Tuckermannopsis Ciliaris
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''Tuckermannopsis ciliaris'' is a species of
corticolous This glossary of mycology is a list of definitions of terms and concepts relevant to mycology, the study of fungi. Terms in common with other fields, if repeated here, generally focus on their mycology-specific meaning. Related terms can be found ...
(bark-dwelling),
foliose lichen A foliose lichen is a lichen with flat, leaf-like , which are generally not firmly bonded to the substrate on which it grows. It is one of the three most common growth forms of lichens. It typically has distinct upper and lower surfaces, each o ...
in the family
Parmeliaceae The Parmeliaceae is a large and diverse family of Lecanoromycetes. With over 2700 species in 71 genera, it is the largest family of lichen-forming fungi. The most speciose genera in the family are the well-known groups: '' Xanthoparmelia'' ( 82 ...
. It was first described by
Erik Acharius Erik Acharius (10 October 1757 – 14 August 1819) was a Swedish botanist who pioneered the Taxonomy (biology), taxonomy of lichens and is known as the "father of lichenology". Acharius was famously the last pupil of Carl Linnaeus. Life Ac ...
in 1810, initially classified in the genus ''
Cetraria ''Cetraria'' is a genus of fruticose lichens that associate with green algae as photobionts. Most species are found at high latitudes, occurring on sand or heath, and are characterised by their "strap-like" form with spiny edges. The lobes can ...
''. The species was later reclassified into the genera ''
Nephromopsis ''Nephromopsis'' is a genus of lichenized fungi within the Parmeliaceae The Parmeliaceae is a large and diverse family of Lecanoromycetes. With over 2700 species in 71 genera, it is the largest family of lichen-forming fungi. The most speci ...
'' and then ''
Tuckermannopsis ''Tuckermannopsis'' is a genus of foliose lichens in the family Parmeliaceae. Taxonomy The genus was circumscribed in 1933 by Hungarian lichenologist Vilmos Kőfaragó-Gyelnik. The genus name ''Tuckermannopsis'' honours Edward Tuckerman (1817†...
'', with some researchers proposing that certain genera, including ''Tuckermannopsis'', should be merged into ''Nephromopsis''. However, this suggestion was disputed, and both names, ''Tuckermannopsis ciliaris'' and ''Nephromopsis ciliaris'', are used in recent literature to refer to this species. ''Tuckermannopsis ciliaris'' is widely distributed across
temperate In geography, the temperate climates of Earth occur in the middle latitudes (approximately 23.5° to 66.5° N/S of the Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ran ...
regions of North America and the northern
Old World The "Old World" () is a term for Afro-Eurasia coined by Europeans after 1493, when they became aware of the existence of the Americas. It is used to contrast the continents of Africa, Europe, and Asia in the Eastern Hemisphere, previously ...
, found on
birch A birch is a thin-leaved deciduous hardwood tree of the genus ''Betula'' (), in the family Betulaceae, which also includes alders, hazels, and hornbeams. It is closely related to the beech- oak family Fagaceae. The genus ''Betula'' contains 3 ...
es and
coniferous Conifers () are a group of conifer cone, cone-bearing Spermatophyte, seed plants, a subset of gymnosperms. Scientifically, they make up the phylum, division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a sin ...
trees such as
pine A pine is any conifer tree or shrub in the genus ''Pinus'' () of the family Pinaceae. ''Pinus'' is the sole genus in the subfamily Pinoideae. ''World Flora Online'' accepts 134 species-rank taxa (119 species and 15 nothospecies) of pines as cu ...
s, with varying frequencies and habitats across different regions. The
thallus Thallus (: thalli), from Latinized Greek (), meaning "a green shoot" or "twig", is the vegetative tissue of some organisms in diverse groups such as algae, fungi, some liverworts, lichens, and the Myxogastria. A thallus usually names the entir ...
grows in irregular shapes or rosettes up to in diameter, with olive-green to greenish-brown coloration and elongated featuring black, hair-like along their edges. Both
apothecia An ascocarp, or ascoma (: ascomata), is the fruiting body ( sporocarp) of an ascomycete phylum fungus. It consists of very tightly interwoven hyphae and millions of embedded asci, each of which typically contains four to eight ascospores. As ...
(sexual fruiting bodies) and
pycnidia A pycnidium (plural pycnidia) is an asexual fruiting body produced by mitosporic fungi, for instance in the order Sphaeropsidales ( Deuteromycota, Coelomycetes) or order Pleosporales (Ascomycota, Dothideomycetes). It is often spherical or inve ...
(asexual fruiting bodies) also occur along the edges of the lobes. The chemical properties of ''T. ciliaris'' include the presence of olivetoric and physodic acids in the
medulla Medulla (Latin for "marrow") or medullary may refer to: Science * Medulla oblongata, a part of the brain stem * Renal medulla, a part of the kidney * Adrenal medulla, a part of the adrenal gland * Medulla of ovary, a stroma in the center of the ...
. Chemical spot tests for these
lichen product Lichen products, also known as lichen substances, are organic compounds produced by a lichen. Specifically, they are secondary metabolites. Lichen products are represented in several different chemical classes, including terpenoids, orcinol deri ...
s can help distinguish ''T. ciliaris'' from several similar species.


Taxonomy

The lichen was
formally described A species description is a formal scientific description of a newly encountered species, typically articulated through a scientific publication. Its purpose is to provide a clear description of a new species of organism and explain how it differ ...
in 1810 by the Swedish lichenologist
Erik Acharius Erik Acharius (10 October 1757 – 14 August 1819) was a Swedish botanist who pioneered the Taxonomy (biology), taxonomy of lichens and is known as the "father of lichenology". Acharius was famously the last pupil of Carl Linnaeus. Life Ac ...
, who initially classified it in ''
Cetraria ''Cetraria'' is a genus of fruticose lichens that associate with green algae as photobionts. Most species are found at high latitudes, occurring on sand or heath, and are characterised by their "strap-like" form with spiny edges. The lobes can ...
'', the namesake genus of the lichens. Cetrarioid lichens are characterised by erect, foliose thalli, and apothecia and pycnidia on the margins of the lobes. In his original description, Acharius summarised the main characteristics of the new species, noting that the
thallus Thallus (: thalli), from Latinized Greek (), meaning "a green shoot" or "twig", is the vegetative tissue of some organisms in diverse groups such as algae, fungi, some liverworts, lichens, and the Myxogastria. A thallus usually names the entir ...
is pale brownish with a whitish, - underside. He emphasised two key diagnostic features that remain important today: the crisped, ciliate with thick black or along their margins, and the thallus texture that becomes more reticulated-lacunose with age. The
apothecia An ascocarp, or ascoma (: ascomata), is the fruiting body ( sporocarp) of an ascomycete phylum fungus. It consists of very tightly interwoven hyphae and millions of embedded asci, each of which typically contains four to eight ascospores. As ...
are slightly elevated and dark brownish, with a raised, margin. These morphological characteristics, particularly the distinctive ciliate lobes, helped distinguish it as a new species. ''Cetraria ciliaris'' resembles the
variety Variety may refer to: Arts and entertainment Entertainment formats * Variety (radio) * Variety show, in theater and television Films * ''Variety'' (1925 film), a German silent film directed by Ewald Andre Dupont * ''Variety'' (1935 film), ...
''ulophylla'' of ''
Cetraria sepincola ''Cetraria sepincola'', the chestnut wrinkle-lichen, is a species of foliose lichen in the family Parmeliaceae. It forms compact, cushion-like colonies typically high, with a yellowish-green to dark brown upper surface and lighter underside. T ...
'' but was much larger and more solid. In 1899, Auguste-Marie Hue proposed reclassifying the species in genus ''
Nephromopsis ''Nephromopsis'' is a genus of lichenized fungi within the Parmeliaceae The Parmeliaceae is a large and diverse family of Lecanoromycetes. With over 2700 species in 71 genera, it is the largest family of lichen-forming fungi. The most speci ...
''. Christian August Frege thought it was more appropriate in the genus ''Platysma'', and moved it there in an 1812 publication.
Vilmos Kőfaragó-Gyelnik Vilmos Kőfaragó-Gyelnik (March 30, 1906 – March 15, 1945) was a Hungarian botanist and lichenologist. Prior to earning his PhD in 1929 from Budapest University, he spent a year in Cairo to help organize a botanical museum. In 1930 he started ...
transferred the
taxon In biology, a taxon (back-formation from ''taxonomy''; : taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular name and ...
to the genus ''
Tuckermannopsis ''Tuckermannopsis'' is a genus of foliose lichens in the family Parmeliaceae. Taxonomy The genus was circumscribed in 1933 by Hungarian lichenologist Vilmos Kőfaragó-Gyelnik. The genus name ''Tuckermannopsis'' honours Edward Tuckerman (1817†...
'' in 1933. In 1963,
Mason Hale Mason Ellsworth Hale, Jr. (September 23, 1929 – April 23, 1990) was one of the most prolific American lichenologists of the 20th century. Many of his scholarly articles focused on the taxonomy of the family Parmeliaceae. Hale was one of the fi ...
identified three chemical strains within the ''Cetraria ciliaris'' group in North America, distinguished by their production of alectoronic acid, olivetoric acid, and protolichesteric acid. This study emphasised the importance of chemical characteristics in distinguishing morphologically identical species and laid the groundwork for further chemical taxonomy in this group of lichens. Building on this, a study by
William William is a masculine given name of Germanic languages, Germanic origin. It became popular in England after the Norman Conquest, Norman conquest in 1066,All Things William"Meaning & Origin of the Name"/ref> and remained so throughout the Middle ...
and Chicita Culberson in 1967 on the ''Cetraria ciliaris''
species complex In biology, a species complex is a group of closely related organisms that are so similar in appearance and other features that the boundaries between them are often unclear. The taxa in the complex may be able to hybridize readily with each oth ...
identified several species within the group, each distinguished by its unique chemical composition and geographic distribution. The study revealed that ''Cetraria ciliaris'' produces olivetoric and physodic acids and is found in eastern North America and Finland. The species is morphologically similar to '' Cetraria halei'', which produces alectoronic acid and has a broader geographic distribution, including the Old World from Finland through the Soviet Union to Japan. Despite the morphological similarities, these species are chemically distinct and occupy slightly different ecological niches. A key aspect of the study was the finding that environmental factors do not significantly influence the chemical composition of these lichens. Instead, the chemical profiles are genetically determined, underscoring the importance of chemical characteristics in lichen taxonomy. Using a
phylogenetic In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...
approach that incorporated estimates of when different groups evolved over time, researchers proposed that certain genera, including ''Tuckermannopsis'', should be merged with ''
Nephromopsis ''Nephromopsis'' is a genus of lichenized fungi within the Parmeliaceae The Parmeliaceae is a large and diverse family of Lecanoromycetes. With over 2700 species in 71 genera, it is the largest family of lichen-forming fungi. The most speci ...
''. However, this suggestion was later disputed, and the change in classification has not been universally accepted, particularly by some North American lichenologists. Consequently, both names, ''Tuckermannopsis ciliaris'' and ''Nephromopsis ciliaris'', have been used in recent literature to refer to this species.


Description

''Tuckermannopsis ciliaris'' is a
foliose lichen A foliose lichen is a lichen with flat, leaf-like , which are generally not firmly bonded to the substrate on which it grows. It is one of the three most common growth forms of lichens. It typically has distinct upper and lower surfaces, each o ...
that typically forms loosely attached, irregular to
rosette Rosette is the French diminutive of ''rose''. It may refer to: Flower shaped designs * Rosette (award), a mark awarded by an organisation * Rosette (design), a small flower design *hence, various flower-shaped or rotational symmetric forms: ** R ...
-shaped growths, reaching up to about in diameter. Its colour ranges from olive-green to greenish-brown. The comprising the
thallus Thallus (: thalli), from Latinized Greek (), meaning "a green shoot" or "twig", is the vegetative tissue of some organisms in diverse groups such as algae, fungi, some liverworts, lichens, and the Myxogastria. A thallus usually names the entir ...
(the main body of the lichen) are elongated and range from 1 to 4 mm in width, with a slightly wrinkled surface. These lobes often have scattered, long, black, hair-like structures called along their edges; the cilia are (unbranched), and measure up to 7 mm long. The upper surface of the thallus varies in color from pale to dark greenish-brown, and it lacks
pseudocyphella Pseudocyphellae (singular ''pseudocyphella'') are structures in lichens that appear as tiny pores on the outer surface (the cortex) of the lichen. They are caused when there is a break in the cortex of the lichen, and the medullary hyphae extend ...
e, which are tiny pores sometimes found in lichens. The
medulla Medulla (Latin for "marrow") or medullary may refer to: Science * Medulla oblongata, a part of the brain stem * Renal medulla, a part of the kidney * Adrenal medulla, a part of the adrenal gland * Medulla of ovary, a stroma in the center of the ...
is white. The lower surface of the thallus ranges in color from brown to blackish and has a somewhat wrinkled texture. There are simple brown
rhizine In lichens, rhizines are multicellular root-like structures arising mainly from the lower surface. A lichen with rhizines is termed rhizinate, while a lichen lacking rhizines is termed erhizinate. Rhizines serve only to anchor the lichen to their s ...
s (small, rootlike structures) in the central part of the thallus that measure 1.5–3 long. The reproductive structures of the lichen, the
apothecia An ascocarp, or ascoma (: ascomata), is the fruiting body ( sporocarp) of an ascomycete phylum fungus. It consists of very tightly interwoven hyphae and millions of embedded asci, each of which typically contains four to eight ascospores. As ...
, are found along the edges and sides of the lobes and can grow up to 3 mm in diameter. These apothecia produce spores that are roughly spherical and measure 4 to 5 Î¼m in both length and width. Additionally, ''Tuckermannopsis ciliaris'' has small, black, protruding structures along the margins called
pycnidia A pycnidium (plural pycnidia) is an asexual fruiting body produced by mitosporic fungi, for instance in the order Sphaeropsidales ( Deuteromycota, Coelomycetes) or order Pleosporales (Ascomycota, Dothideomycetes). It is often spherical or inve ...
, which produce
conidia A conidium ( ; : conidia), sometimes termed an asexual chlamydospore or chlamydoconidium (: chlamydoconidia), is an asexual, non- motile spore of a fungus. The word ''conidium'' comes from the Ancient Greek word for dust, ('). They are also ...
(asexual spores). The conidia are dumbbell-shaped and measure approximately 5 Î¼m in length and 1 Î¼m in width.


Chemistry

The expected results of standard chemical spot tests for ''Tuckermannopsis ciliaris'' are K−, C−, KC+ (reddish), and P− in the medulla; all spot tests are negative on the . The species produces olivetoric and physodic acids in the medulla and atranorin in the cortex. The joint occurrence of olivetoric acid and its corresponding
depsidone Depsidones (+ " depside" + "one") are chemical compounds that are sometimes found as secondary metabolites in lichens. They are esters that are both depsides and cyclic ethers. An example is norstictic acid Norstictic acid is a depsidone produ ...
, physodic acid, was the first known example of such a relationship, supporting the theory of the origin of depsidones from
depside A depside is a type of polyphenolic compound composed of two or more monocyclic aromatic units linked by an ester group. Depsides are most often found in lichens, but have also been isolated from higher plants, including species of the Ericaceae, ...
s.


Similar species

''Tuckermannopsis ciliaris'' can be easily confused with some other species due to its morphology and presence of cilia. One such species is '' Nephromopsis ahtii'', which shares a similar appearance and the presence of cilia. However, ''Tuckermannopsis ciliaris'' can be distinguished by the presence of atranorin and olivetoric acids as secondary metabolites. Another species that resembles ''Tuckermannopsis ciliaris'' is '' Nephromopsis chlorophylla''. While it has a similar thallus colour and size, it lacks cilia and contains
protolichesterinic acid Protolichesterinic acid is a naturally occurring γ-lactone compound found in various lichen species. Its structure consists of a combination of a lactone ring with a carboxylic acid group and a long aliphatic side chain. First isolated in the ...
in its thallus, which sets it apart from ''Tuckermannopsis ciliaris''. Two other potential lookalikes, '' Tuckermannopsis orbata'' and '' Tuckermannopsis fendleri'', can be distinguished from ''T. ciliaris'' by their negative reaction to
calcium hypochlorite Calcium hypochlorite is an inorganic compound with chemical formula , also written as . It is a white solid, although commercial samples appear yellow. It strongly smells of chlorine, owing to its slow decomposition in moist air. This compound is ...
(C−) in the medulla. '' Tuckermannopsis americana'' is a chemical variant (
chemotype A chemotype (sometimes chemovar) is a chemically distinct entity in a plant or microorganism, with differences in the composition of the secondary metabolites. Minor genetic and epigenetic changes with little or no effect on morphology or anatomy ...
) of ''T. ciliaris'' that produces different substances, alectoronic acid and α-collatolic acid, instead of the olivetoric and physodic acids found in typical ''T. ciliaris''. It is usually treated as a distinct species, although some sources treat the two species as one. ''Tuckermannopsis americana'' is C− in the medulla, but shows a positive UV+ (bluish-white) reaction because of the alectoronic acid.


Habitat and distribution

''Tuckermannopsis ciliaris'' is distributed across a wide range of regions. In North America, it is found from southeastern Canada to the Lake States and south throughout the
Appalachian Mountains The Appalachian Mountains, often called the Appalachians, are a mountain range in eastern to northeastern North America. The term "Appalachian" refers to several different regions associated with the mountain range, and its surrounding terrain ...
and associated foothills. It is generally common in appropriate habitats in northern North America, except for the
Great Plains The Great Plains is a broad expanse of plain, flatland in North America. The region stretches east of the Rocky Mountains, much of it covered in prairie, steppe, and grassland. They are the western part of the Interior Plains, which include th ...
region. In the
Old World The "Old World" () is a term for Afro-Eurasia coined by Europeans after 1493, when they became aware of the existence of the Americas. It is used to contrast the continents of Africa, Europe, and Asia in the Eastern Hemisphere, previously ...
, it is distributed from Finland through Russia to Japan. In Finland, it is considered rare or possibly extinct, and has a
conservation status The conservation status of a group of organisms (for instance, a species) indicates whether the group still exists and how likely the group is to become extinct in the near future. Many factors are taken into account when assessing conservation ...
of "regionally extinct" in the Finnish
red list The International Union for Conservation of Nature (IUCN) Red List of Threatened Species, also known as the IUCN Red List or Red Data Book, founded in 1964, is an inventory of the global conservation status and extinction risk of biological sp ...
. It was reported as new to India in 2022, where it is considered rare. The species typically grows on the trunks and branches of
pine A pine is any conifer tree or shrub in the genus ''Pinus'' () of the family Pinaceae. ''Pinus'' is the sole genus in the subfamily Pinoideae. ''World Flora Online'' accepts 134 species-rank taxa (119 species and 15 nothospecies) of pines as cu ...
s and other
conifer Conifers () are a group of conifer cone, cone-bearing Spermatophyte, seed plants, a subset of gymnosperms. Scientifically, they make up the phylum, division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a sin ...
s. In the Nordic countries, it has only been recorded growing on
birch A birch is a thin-leaved deciduous hardwood tree of the genus ''Betula'' (), in the family Betulaceae, which also includes alders, hazels, and hornbeams. It is closely related to the beech- oak family Fagaceae. The genus ''Betula'' contains 3 ...
bark.


Uses

''Tuckermannopsis ciliaris'' is an auxiliary component of a hair tint that was patented in Korea in 2004.


References

{{Taxonbar, from1=Q10705992 , from2=Q59590859 , from3=Q61722033 , from4=Q61727013 , from5=Q61727777 Parmeliaceae Lichen species Lichens described in 1810 Lichens of Asia Lichens of North America Taxa named by Erik Acharius