''Carbonicola'' is a small

genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...

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 ...

-forming

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 ...

. It is the sole genus in the monogeneric

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 ...

Carbonicolaceae. The genus, which collectively has an almost

cosmopolitan distribution In biogeography, a cosmopolitan distribution is the range of a taxon that extends across most or all of the surface of the Earth, in appropriate habitats; most cosmopolitan species are known to be highly adaptable to a range of climatic and en ...

, contains three

squamulose lichen A squamulose lichen is a lichen that is composed of small, often overlapping "scales" called . 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" (cortex), ...

s that prefer to grow on burned wood in

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 ...

areas of the world.

Taxonomy

Both the genus and the family were

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 ...

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 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 ...

, dividable into seven

clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...

s belonging to different genera, families,

orders Order, ORDER or Orders may refer to: * A socio-political or established or existing order, e.g. World order, Ancien Regime, Pax Britannica * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * H ...

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

group A group is a number of persons or things that are located, gathered, or classed together. Groups of people * Cultural group, a group whose members share the same cultural identity * Ethnic group, a group whose members share the same ethnic iden ...

. 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 ...

position to a clade containing the

Cladoniaceae The Cladoniaceae are a family of lichen-forming fungi in the order Lecanorales, comprising about 560 species distributed amongst 18 genera. This family is one of the largest among lichen-forming fungi and is globally distributed, from Arctic t ...

and

Stereocaulaceae The Stereocaulaceae are a family of lichen-forming fungi in the order Lecanorales. It contains five genera. Species of this family are widely distributed in temperate boreal and austral regions. Genera *'' Hertelidea'' – 6 spp. *''Lepraria ...

. The generic name ''Carbonicola'' combines the

Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...

roots ''carbo'' ("charcoal") and -''cola'' ("dweller") to refer to burned wood, its preferred

substrate Substrate may refer to: Physical layers *Substrate (biology), the natural environment in which an organism lives, or the surface or medium on which an organism grows or is attached ** Substrate (aquatic environment), the earthy material that exi ...

. In 2017, Divakar and colleagues proposed to subsume the Carbonicolaceae into the

Lecanoraceae The Lecanoraceae are a family 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 c ...

, 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 c ...

'' 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 (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 ...

squamulose A squamulose lichen is a lichen that is composed of small, often overlapping "scales" called . 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" (cortex) ...

thallus. Their

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.

partner is chlorococcoid–spherical, single-celled

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 ( ...

e. The

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 ...

are in the form of an

apothecium 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. Ascoc ...

, 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 o ...

e between the asci) is

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 ...

, 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 f ...

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

. 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 SEPTA, the Southeastern Pennsylvania Transportation Authority, is a regional public transportation authority that operates bus, rapid transit, commuter rail, light rail, and electric trolleybus services for nearly four million people througho ...

, are

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 ...

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 ...

, and non-amyloid. The

conidiomata Conidiomata (singular: Conidioma) are blister-like fruiting structures produced by a specific type of fungus called a coelomycete. They are formed as a means of dispersing asexual spores call conidia, which they accomplish by creating the blister- ...

are in the form of

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 ...

; 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 ...

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 produ ...

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 lichen growth forms, growth structure. It is formed from a symbiotic relationship of a photobiont such as green algae or less commonly cyanobacteri ...

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 Cortex or cortical may refer to: Biology * Cortex (anatomy), the outermost layer of an organ ** Cerebral cortex, the outer layer of the vertebrate cerebrum, part of which is the ''forebrain'' *** Motor cortex, the regions of the cerebral cortex i ...

Habitat and distribution

Typically

epiphytic An epiphyte is a plant or plant-like organism that grows on the surface of another 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 epiphyt ...

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 Taxa named by Einar Timdal