Arthoniaceae
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The Arthoniaceae are a
family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
of
lichen A lichen ( , ) is a hybrid colony (biology), colony of algae or cyanobacteria living symbiotically among hypha, filaments of multiple fungus species, along with yeasts and bacteria embedded in the cortex or "skin", in a mutualism (biology), m ...
ized,
lichenicolous 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 ...
and
saprobic Saprotrophic nutrition or lysotrophic nutrition is a process of chemoheterotrophic extracellular digestion involved in the processing of decayed (dead or waste) organic matter. It occurs in saprotrophs, and is most often associated with fungi ...
fungi A fungus (: fungi , , , or ; or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and mold (fungus), molds, as well as the more familiar mushrooms. These organisms are classified as one ...
in the order
Arthoniales The Arthoniales is the second largest order of mainly crustose lichens, but fruticose lichens are present as well. The order contains around 1500 species, while the largest order with lichenized fungi, the Lecanorales, contains more than 14000 s ...
.Encyclopedia of Life, consulted at september the seventeenth 2013
/ref> The Arthoniaceae is the largest family of
Arthoniales The Arthoniales is the second largest order of mainly crustose lichens, but fruticose lichens are present as well. The order contains around 1500 species, while the largest order with lichenized fungi, the Lecanorales, contains more than 14000 s ...
, with around 800 species. Most species in Arthoniaceae belong in ''
Arthonia ''Arthonia'' is a genus of lichens in the family Arthoniaceae. It was circumscribed by Swedish botanist Erik Acharius in 1806. It is a genus of thin crustose lichen of widely varying forms, commonly called comma lichens.Field Guide to California ...
'' which is the largest genus with 500 species. The second and third largest genus is ''
Arthothelium ''Arthothelium'' is a genus of lichen-forming fungi in the family Arthoniaceae. Description ''Arthothelium'' is a genus of crustose lichens, which can either be (embedded within the ) or superficial (growing on the surface). The thallus may s ...
'' with 80 species, and ''
Cryptothecia ''Cryptothecia'' is a genus of white to greenish crustose lichens that grow on bark, wood, or leaves, in tropical or subtropical areas worldwide. It has a conspicuous prothallus that develops around its periphery which can be bright red in some s ...
'' with 60 species.


Taxonomy

The family was
circumscribed In geometry, a circumscribed circle for a set of points is a circle passing through each of them. Such a circle is said to ''circumscribe'' the points or a polygon formed from them; such a polygon is said to be ''inscribed'' in the circle. * Circum ...
by
Heinrich Gottlieb Ludwig Reichenbach Heinrich Gottlieb Ludwig Reichenbach (8 January 1793 – 17 March 1879) was a German botanist, ornithologist and illustrator. It was he who first requested Leopold Blaschka to make a set of glass marine invertebrate models for scientific educatio ...
in 1841. ''Arthonia'' is the
type genus In biological taxonomy, the type genus (''genus typica'') is the genus which defines a biological family and the root of the family name. Zoological nomenclature According to the International Code of Zoological Nomenclature, "The name-bearin ...
of Arthoniaceae, and it is known to be a
polyphyletic A polyphyletic group is an assemblage that includes organisms with mixed evolutionary origin but does not include their most recent common ancestor. The term is often applied to groups that share similar features known as Homoplasy, homoplasies ...
and
paraphyletic Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...
genus. The process of splitting ''Arthonia'' into
monophyletic In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent co ...
groups is an ongoing process. In order to make ''Arthonia'' monophyletic, several genera have been described or resurrected.


Description

The family Arthoniaceae consists of lichens that are either lichenized or lichenicolous (living on other lichens), and sometimes lose their lichenized nature. The main body of these lichens, known as 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
crustose Crustose is a Habit (biology), habit of some types of algae and lichens in which the organism grows tightly appressed to a substrate, forming a biological layer. ''Crustose'' adheres very closely to the Substrate (biology), substrates at all poin ...
, meaning it forms a crust-like appearance on the surface it inhabits. In lichen-forming species of Arthoniaceae, the primary
photosynthetic Photosynthesis ( ) is a Biological system, system of biological processes by which Photoautotrophism, photosynthetic organisms, such as most plants, algae, and cyanobacteria, convert light energy, typically from sunlight, into the chemical ener ...
partner (the ), is usually a
green alga The green algae (: green alga) are a group of chlorophyll-containing autotrophic eukaryotes consisting of the phylum Prasinodermophyta and its unnamed sister group that contains the Chlorophyta and Charophyta/ Streptophyta. The land plants ( ...
from the genus '' Trentepohlia''. Occasionally, they associate with other green algae, like '' Chrosonothrix''. The form of the reproductive structures (
ascomata An ascocarp, or ascoma (: ascomata), is the fruiting body (sporocarp (fungi), sporocarp) of an ascomycete phylum fungus. It consists of very tightly interwoven hyphae and millions of embedded ascus, asci, each of which typically contains four to ...
) varies widely within this family. They can be (open and -like), (elongated and furrowed), or scattered. Some genera, like ''
Tylophoron ''Tylophoron'' is a genus of lichen-forming fungi in the family Arthoniaceae. It comprises seven species of crustose lichens, most of which occur in tropical regions. Taxonomy The genus was circumscribed in 1862 by the Finnish lichenologist Wil ...
'', have a unique type of ascomata called . Typically, these structures lack distinct margins made of their own tissue () or host tissue (). The internal structure of the ascomata, known as the , consists of branched and interconnected filaments called ''
paraphyses Paraphyses are erect sterile filament-like support structures occurring among the reproductive apparatuses of fungi, ferns, bryophytes and some thallophytes. The singular form of the word is paraphysis. In certain fungi, they are part of the f ...
''. In some genera, such as ''
Cryptothecia ''Cryptothecia'' is a genus of white to greenish crustose lichens that grow on bark, wood, or leaves, in tropical or subtropical areas worldwide. It has a conspicuous prothallus that develops around its periphery which can be bright red in some s ...
'' and '' Stirtonia'', the hamathecium lacks a gelatinous matrix, while in others, like ''
Arthothelium ''Arthothelium'' is a genus of lichen-forming fungi in the family Arthoniaceae. Description ''Arthothelium'' is a genus of crustose lichens, which can either be (embedded within the ) or superficial (growing on the surface). The thallus may s ...
'', it is densely packed, resembling a different type of fungal reproductive structure known as ascolocular ascomata. These structures can react to
iodine Iodine is a chemical element; it has symbol I and atomic number 53. The heaviest of the stable halogens, it exists at standard conditions as a semi-lustrous, non-metallic solid that melts to form a deep violet liquid at , and boils to a vi ...
(
amyloid Amyloids are aggregates of proteins characterised by a fibrillar morphology of typically 7–13 nm in diameter, a β-sheet secondary structure (known as cross-β) and ability to be stained by particular dyes, such as Congo red. In the human ...
reaction) in various ways: not at all (non-amyloid), partially (hemiamyloid), or fully (amyloid). The asci, which are the spore-producing cells, are typically , meaning they have a double wall that splits during spore release. They come in various shapes, from club-shaped () to sac-like (saccate) or even spherical (), and contain a specialized cap structure (apical ) and an eye-like feature (ocular chamber). The asci may react to iodine in different ways: non-amyloid, hemiamyloid, or amyloid. The spores produced by the asci, the
ascospore In fungi, an ascospore is the sexual spore formed inside an ascus—the sac-like cell that defines the division Ascomycota, the largest and most diverse Division (botany), division of fungi. After two parental cell nucleus, nuclei fuse, the ascu ...
s, usually number eight per ascus but can sometimes be fewer, ranging from two to four or even one. These spores are segmented (transversely
septate In biology, a septum (Latin for ''something that encloses''; septa) is a wall, dividing a cavity or structure into smaller ones. A cavity or structure divided in this way may be referred to as septate. Examples Human anatomy * Interatrial se ...
) or have multiple compartments (), with shapes ranging from
ellipsoid An ellipsoid is a surface that can be obtained from a sphere by deforming it by means of directional Scaling (geometry), scalings, or more generally, of an affine transformation. An ellipsoid is a quadric surface;  that is, a Surface (mathemat ...
to clavate. Often, one end of the spore is larger (macrocephalic), or in muriform spores, there is a single large cell. The spores are typically transparent (
hyaline A hyaline substance is one with a glassy appearance. The word is derived from , and . Histopathology Hyaline cartilage is named after its glassy appearance on fresh gross pathology. On light microscopy of H&E stained slides, the extracellula ...
) or brown and do not react to iodine. Arthoniaceae also produce asexual spores in structures 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 ...
, and less commonly in . The
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) can vary in shape from simple, non-segmented forms to those with transverse segments, and can be oblong, rod-like (), needle-like (-), or sickle-shaped (), and are typically hyaline. The family produces a diverse range of
secondary metabolite Secondary metabolites, also called ''specialised metabolites'', ''secondary products'', or ''natural products'', are organic compounds produced by any lifeform, e.g. bacteria, archaea, fungi, animals, or plants, which are not directly involved ...
s (
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), including
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,
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 ...
s,
anthraquinone Anthraquinone, also called anthracenedione or dioxoanthracene, is an aromatic hydrocarbon, aromatic organic compound with formula . Several isomers exist but these terms usually refer to 9,10-anthraquinone (IUPAC: 9,10-dioxoanthracene) wherein th ...
s, and derivatives of
pulvinic acid Pulvinic acids are natural chemical pigments found in some lichens, derived biosynthetically from the aromatic amino acids phenylalanine and tyrosine, via dimerization and oxidative ring-cleavage of arylpyruvic acids, a process that also produces ...
. Some species also contain
xanthone Xanthone is an organic compound with the molecular formula C13H8O2. It is a white solid. In 1939, xanthone was introduced as an insecticide and it currently finds uses as ovicide for codling moth eggs and as a larvicide. Xanthone is also use ...
s.


Distribution

The species in Arthoniaceae have a worldwide distribution, but are especially prevalent in
tropical The tropics are the regions of Earth surrounding the equator, where the sun may shine directly overhead. This contrasts with the temperate or polar regions of Earth, where the Sun can never be directly overhead. This is because of Earth's ax ...
areas with a
Mediterranean climate A Mediterranean climate ( ), also called a dry summer climate, described by Köppen and Trewartha as ''Cs'', is a temperate climate type that occurs in the lower mid-latitudes (normally 30 to 44 north and south latitude). Such climates typic ...
. They are known from arctic to tropical latitudes, as well as variating altitudes from sea level to alpine regions, distributed in both humid forests and dry habitats.


Ecology

Collectively, the family have a highly variable ecology with
lichen A lichen ( , ) is a hybrid colony (biology), colony of algae or cyanobacteria living symbiotically among hypha, filaments of multiple fungus species, along with yeasts and bacteria embedded in the cortex or "skin", in a mutualism (biology), m ...
ized,
lichenicolous 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 ...
and
saprobic Saprotrophic nutrition or lysotrophic nutrition is a process of chemoheterotrophic extracellular digestion involved in the processing of decayed (dead or waste) organic matter. It occurs in saprotrophs, and is most often associated with fungi ...
fungi A fungus (: fungi , , , or ; or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and mold (fungus), molds, as well as the more familiar mushrooms. These organisms are classified as one ...
. The majority of species are lichenized with a
photobiont A lichen ( , ) is a hybrid colony of algae or cyanobacteria living symbiotically among filaments of multiple fungus species, along with yeasts and bacteria embedded in the cortex or "skin", in a mutualistic relationship.
from
Trentepohliaceae Trentepohliaceae are a family (biology), family of green algae, and the only family in the order Trentepohliales. It is a member of the Ulvophyceaen green algae. The family is characterized by traits like net-like chloroplasts without pyrenoids, ...
and a few species in ''Arthonia'' are lichenized with a photobiont from Chlorococcaleae. They grow on leaves, bark, bryophytes, and rocks. Other species are lichenicolous (growing on other lichens), and a few species are known to be saprobic.


Genera

,
Species Fungorum ''Index Fungorum'' is an international project to index all formal names (Binomial nomenclature, scientific names) in the fungus Kingdom (biology), kingdom. As of 2015, the project is based at the Royal Botanic Gardens, Kew, one of three partn ...
accepts 25 genera and 392 species in the family Arthoniaceae. This is a list of the genera in the Arthoniaceae based on a 2020 review and summary of fungal
classification Classification is the activity of assigning objects to some pre-existing classes or categories. This is distinct from the task of establishing the classes themselves (for example through cluster analysis). Examples include diagnostic tests, identif ...
by Wijayawardene and colleagues. Following the genus name is the
taxonomic authority In biology, taxonomy () is the scientific study of naming, defining ( circumscribing) and classifying groups of biological organisms based on shared characteristics. Organisms are grouped into taxa (singular: taxon), and these groups are given ...
(those who first
circumscribed In geometry, a circumscribed circle for a set of points is a circle passing through each of them. Such a circle is said to ''circumscribe'' the points or a polygon formed from them; such a polygon is said to be ''inscribed'' in the circle. * Circum ...
the genus; standardized author abbreviations are used), year of publication, and the number of species: *'' Amazonomyces'' – 2 spp. *''
Arthonia ''Arthonia'' is a genus of lichens in the family Arthoniaceae. It was circumscribed by Swedish botanist Erik Acharius in 1806. It is a genus of thin crustose lichen of widely varying forms, commonly called comma lichens.Field Guide to California ...
'' – ca. 50 + ca. 300 orphaned spp. *''
Arthothelium ''Arthothelium'' is a genus of lichen-forming fungi in the family Arthoniaceae. Description ''Arthothelium'' is a genus of crustose lichens, which can either be (embedded within the ) or superficial (growing on the surface). The thallus may s ...
'' – 10 + ca. 100 orphaned spp. *'' Briancoppinsia'' – 1 sp. *'' Coniangium'' – 4 spp. *'' Coniarthonia'' – 12 spp. *'' Coniocarpon'' – 6 spp. *'' Crypthonia'' – 16 spp. *'' Cryptophaea'' – 1 spp. *''
Cryptothecia ''Cryptothecia'' is a genus of white to greenish crustose lichens that grow on bark, wood, or leaves, in tropical or subtropical areas worldwide. It has a conspicuous prothallus that develops around its periphery which can be bright red in some s ...
'' – ca. 65 *'' Diarthonis'' – 1 sp. *'' Eremothecella'' – 8 spp. *'' Glomerulophoron'' – 2 spp. *'' Helicobolomyces'' – 2 spp. *'' Herpothallon'' – ca. 50 *'' Inoderma'' – 4 spp. *'' Leprantha'' – 1 sp. *'' Myriostigma'' – 7 spp. *'' Naevia'' – 2 spp. *'' Pachnolepia'' – 2 spp. *'' Reichlingia'' – 4 spp. *'' Snippocia'' – 1 sp. *'' Sporodophoron'' – 4 spp. *'' Staurospora'' – 1 sp. *'' Stirtonia'' – ca. 25 spp. *''
Tylophoron ''Tylophoron'' is a genus of lichen-forming fungi in the family Arthoniaceae. It comprises seven species of crustose lichens, most of which occur in tropical regions. Taxonomy The genus was circumscribed in 1862 by the Finnish lichenologist Wil ...
'' – 11 spp.


References

{{Taxonbar, from=Q2864921 Lichen families Ascomycota families Taxa named by Ludwig Reichenbach Taxa described in 1841