''Carbonicola'' is a small

genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...

lichen A lichen ( , ) is a composite organism that arises from algae or cyanobacteria living among filaments of multiple fungi species in a mutualistic relationship.fungi A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately from ...

. It is the sole genus in the monogeneric

family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...

Carbonicolaceae. The genus, which collectively has an almost

cosmopolitan distribution In biogeography, cosmopolitan distribution is the term for the range of a taxon that extends across all or most of the world in appropriate habitats. Such a taxon, usually a species, is said to exhibit cosmopolitanism or cosmopolitism. The ext ...

, contains three

squamulose lichen A squamulose lichen is a lichen that is composed of small, often overlapping "scales" called squamules. If they are raised from the substrate and appear leafy, the lichen may appear to be a foliose lichen, but the underside does not have a "ski ...

s that prefer to grow on burned wood in

temperate In geography, the temperate climates of Earth occur in the middle latitudes (23.5° to 66.5° N/S of Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ranges throughout t ...

areas of the world.

Taxonomy

Both the genus and the family were

circumscribed In geometry, the circumscribed circle or circumcircle of a polygon is a circle that passes through all the vertices of the polygon. The center of this circle is called the circumcenter and its radius is called the circumradius. Not every polyg ...

in 2013 by Mika Bendiksby and Ernst Timdal. In a reassessment of the genus ''Hypocenomyce'', they found that the genus–as then circumscribed–was polyphyletic, dividable into seven

clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...

s belonging to different genera, families,

orders Order, ORDER or Orders may refer to: * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of d ...

and even subclasses. ''Carbonicola'' includes the three species formerly included in the ''Hypocenomyce anthracophila'' group. Using

molecular phylogenetic Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...

analysis, genus ''Carbonicola'' was shown outside to lie outside the Lecanoraceae, in a

sister group In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and t ...

position to a clade containing the Cladoniaceae and Stereocaulaceae. The generic name ''Carbonicola'' combines the

Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...

roots ''carbo'' ("charcoal") and -''cola'' ("dweller") to refer to burned wood, its preferred substrate. In 2017, Divakar and colleagues proposed to subsume the Carbonicolaceae into the

Lecanoraceae The Lecanoraceae are a family of lichenized fungi in the order Lecanorales. Species of this family have a widespread distribution. Taxonomy Lecanoraceae was circumscribed by German lichenologist Gustav Wilhelm Körber in 1855. Genera According ...

, because in their analysis, genus ''Carbonicola'' was found to be nested in ''

Lecanora ''Lecanora'' is a genus of lichen commonly called rim lichens.Field Guide to California Lichens, Stephen Sharnoff, Yale University Press, 2014, Lichens in the genus ''Squamarina'' are also called rim lichens. Members of the genus have roughly ci ...

'' in the broad sense. This synonymy was not accepted in a review published soon afterwards, as the phylogenetic conclusions contradicted the findings of Bendiksby and Timdal, making such a decision premature.

Description

''Carbonicola'' species have a

crustose Crustose is a 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 substrates at all points. ''Crustose'' is found on rocks and ...

squamulose A squamulose lichen is a lichen that is composed of small, often overlapping "scales" called squamules. If they are raised from the substrate and appear leafy, the lichen may appear to be a foliose lichen, but the underside does not have a "skin ...

thallus. Their

photobiont A lichen ( , ) is a composite organism that arises from algae or cyanobacteria living among filaments of multiple fungi species in a mutualistic relationship.green alga The green algae (singular: green alga) are a group consisting of the Prasinodermophyta and its unnamed sister which contains the Chlorophyta and Charophyta/Streptophyta. The land plants (Embryophytes) have emerged deep in the Charophyte alga as ...

e. The ascomata are in the form of an

apothecium An ascocarp, or ascoma (), 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. Ascocarps are ...

, biatorine, with reduced margin. The hamathecium (i.e.,

hypha A hypha (; ) is a long, branching, filamentous structure of a fungus, oomycete, or actinobacterium. In most fungi, hyphae are the main mode of vegetative growth, and are collectively called a mycelium. Structure A hypha consists of one or ...

e between the

asci ASCI or Asci may refer to: * Advertising Standards Council of India * Asci, the plural of ascus, in fungal anatomy * Accelerated Strategic Computing Initiative * American Society for Clinical Investigation * Argus Sour Crude Index * Association of ...

) is amyloid, and comprises unbranched

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 fe ...

. The asci are semifissitunicate, meaning that the two walls of the ascus do not completely separate during the discharge of the

ascospore An ascus (; ) is the sexual spore-bearing cell produced in ascomycete fungi. Each ascus usually contains eight ascospores (or octad), produced by meiosis followed, in most species, by a mitotic cell division. However, asci in some genera or s ...

. Asci, club-shaped (clavate), and have a weakly amyloid apical tholus that contains a darker amyloid, tubular structure. Ascospores number eight per ascus, and lack

septa The Southeastern Pennsylvania Transportation Authority (SEPTA) is a regional public transportation authority that operates bus, rapid transit, commuter rail, light rail, and electric trolleybus services for nearly 4 million people in five coun ...

, are

ellipsoid An ellipsoid is a surface that may be obtained from a sphere by deforming it by means of directional scalings, or more generally, of an affine transformation. An ellipsoid is a quadric surface; that is, a surface that may be defined as the ...

hyaline A hyaline substance is one with a glassy appearance. The word is derived from el, ὑάλινος, translit=hyálinos, lit=transparent, and el, ὕαλος, translit=hýalos, lit=crystal, glass, label=none. Histopathology Hyaline cartilage is ...

, and non-amyloid. The conidiomata are in the form of pycnidia; the

conidia A conidium ( ; ), sometimes termed an asexual chlamydospore or chlamydoconidium (), is an asexual, non-motile spore of a fungus. The word ''conidium'' comes from the Ancient Greek word for dust, ('). They are also called mitospores due to the ...

are non-septate, thread-shaped (filiform), and hyaline.

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 produced ...

s are the predominant secondary chemicals found in ''Carbonicola''. These include colensoic acid, 4-O- methylphysodic acid and related compounds, which are found in all species, and fumarprotocetraric and protocetraric acid, which are found in ''C. anthracophila''. The diagnostic characters of Carbonicolaceae differentiate it from its sister taxa, the Cladoniaceae and the Stereocaulaceae. Unlike those families, which usually form

fruticose A fruticose lichen is a form of lichen fungi that is characterized by a coral-like shrubby or bushy growth structure. It is formed from a symbiotic relationship of a photobiont such as green algae or less commonly cyanobacteria and one, two or mor ...

secondary thalli (a feature absent in the Carbonicolaceae), Carbonicolaceae species form purely crustose to squamulose thalli that are dark brown with a thick, shiny upper cortex.

Habitat and distribution

Typically

epiphytic An epiphyte is an organism that grows on the surface of a plant and derives its moisture and nutrients from the air, rain, water (in marine environments) or from debris accumulating around it. The plants on which epiphytes grow are called phoroph ...

on burnt wood or bark, ''Carbonicola'' has a sub

, and is found in

regions.

Species

*'' Carbonicola anthracophila'' *'' Carbonicola foveata'' *'' Carbonicola myrmecina''

References

{{Taxonbar, from1=Q64755645, from2=Q21229378 Lecanorales Lichen genera Lecanorales genera Taxa described in 2013