''Calvitimela'' is a

lichen A lichen ( , ) is a composite organism that arises from algae or cyanobacteria living among filaments of multiple fungi species in a mutualistic relationship.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 ...

Tephromelataceae The Tephromelataceae are a family of lichenized fungi in the order Lecanorales. The family was circumscribed by Austrian lichenologist Josef Hafellner in 1984. Tephromelataceae comprises the genera '' Tephromela'', '' Calvitimela'', '' Mycoblastu ...

(

Lecanorales The Lecanorales are an order of mostly lichen-forming fungi belonging to the class Lecanoromycetes in the division Ascomycota. The order contains 26 families, 269 genera, and 5695 species. Families * Aphanopsidaceae * Biatorellaceae * Brigan ...

Lecanoromycetes Lecanoromycetes is the largest class of lichenized fungi. It belongs to the subphylum Pezizomycotina in the phylum Ascomycota. The asci (spore-bearing cells) of the Lecanoromycetes most often release spores by rostrate dehiscence. Genera of unc ...

). Members of the

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 ...

are

crustose lichens Crustose lichens are lichens that form a crust which strongly adheres to the substrate (soil, rock, tree bark, etc.), making separation from the substrate impossible without destruction. The basic structure of crustose lichens consists of a cort ...

with green photobionts and lecideine or lecanorine

apothecia 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 mo ...

. The species in ''Calvitimela'' have lecideine

, are saxicolous and are primarily found in

alpine Alpine may refer to any mountainous region. It may also refer to: Places Europe * Alps, a European mountain range ** Alpine states, which overlap with the European range Australia * Alpine, New South Wales, a Northern Village * Alpine National Pa ...

arctic The Arctic ( or ) is a polar regions of Earth, polar region located at the northernmost part of Earth. The Arctic consists of the Arctic Ocean, adjacent seas, and parts of Canada (Yukon, Northwest Territories, Nunavut), Danish Realm (Greenla ...

regions.

Species

''Calvitimela'' currently includes ten species: *'' Calvitimela aglaea'' (Sommerf.) Hafellner *'' Calvitimela armeniaca'' (DC.) Hafellner (type species) *'' Calvitimela austrochilensis'' Fryday *'' Calvitimela cuprea'' Haugan & Timdal *'' Calvitimela livida'' Haugan & Timdal *'' Calvitimela melaleuca'' (Sommerf.) R. Sant *'' Calvitimela perlata'' (Haugan & Timdal) R. Sant *'' Calvitimela septentrionalis'' (Hertel & Rambold) McCune *'' Calvitimela talayana'' (Haugan & Timdal) M.P. Andreev *'' Calvitimela uniseptata'' G. Thor

Taxonomy

The taxonomic history of ''Calvitimela'' is long and relatively complex. In the early days, the species now belonging to ''Calvitimela'' were placed in the classical genus ''

Lecidea ''Lecidea'' is a genus of crustose lichens with a carbon black ring or outer margin (exciple) around the fruiting body disc (apothecium), usually (or always) found growing on (saxicolous) or in (endolithic) rock.Field Guide to California Lichens, ...

'' Ach., e.g. by Fries (1874; as ''Lecidea'' strips ''L. armeniacae''). and Magnusson (1931; as “''Lecidea armeniaca''- und ''elata''-Gruppe”) In the 1980s, several taxonomists revised large groups of lichens 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 ...

'' and ''

''. First, the

generic Generic or generics may refer to: In business * Generic term, a common name used for a range or class of similar things not protected by trademark * Generic brand, a brand for a product that does not have an associated brand or trademark, other ...

name '' Tephromela'' was resurrected, and the new

monotypic In biology, a monotypic taxon is a taxonomic group (taxon) that contains only one immediately subordinate taxon. A monotypic species is one that does not include subspecies or smaller, infraspecific taxa. In the case of genera, the term "unispec ...

described. Then, a group from ''Lecidea'' was moved into the genus ''Tephromela'' M. Choisy. This led to species with both lecanorine and lecideine

being present in the same genus. Finally, the genus ''Calvitimela'' was erected for species with lecideine apothecia previously placed in ''Tephromela''.

consists of the '' Tephromela'', ''Calvitimela'', '' Mycoblastus'' Norman and '' Violella'' T. Sprib, and together these four genera constitute a well-supported monophyletic group.

Morphology, anatomy, and chemistry

The species of ''Calvitimela'' are

. Their

thallus Thallus (plural: 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. Many of these organisms wer ...

are areolate and their

lecideine. The apothecia are convex, black and shiny.

Apothecia 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 mo ...

are rare or entirely absent in some sorediate species e.g. ''C. cuprea'', ''C. livida'' and ''C. talayana''. Ascii are generally of the ''Lecanora''-type, but ''Bacidia''-type ascii are also observed in ''C. aglaea'' and ''C. perlata''. The spores are usually simple and ellipsoid. There is much chemical variation of

secondary metabolites Secondary metabolites, also called specialised metabolites, toxins, secondary products, or natural products, are organic compounds produced by any lifeform, e.g. bacteria, fungi, animals, or plants, which are not directly involved in the nor ...

in the species of ''Calvitimela''. Most prominently observed in the ''C. melaleuca'' – complex, with alectorialic, norstictic, roccellic and psoromic acids occurring in different combinations in the species´ several chemotypes.

Ecology and distribution

All species of ''Calvitimela'' grow on rocks. The species in ''Calvitimela'' reside either on

boulders In geology, a boulder (or rarely bowlder) is a rock fragment with size greater than in diameter. Smaller pieces are called cobbles and pebbles. While a boulder may be small enough to move or roll manually, others are extremely massive. In co ...

of varying size or directly on mountainous walls and are occasionally found on pebbles. A peculiar ecological preference is observed in ''C. cuprea''. This species is more or less morphologically identical to ''C. livida'', but seem to only grow on

copper Copper is a chemical element with the symbol Cu (from la, cuprum) and atomic number 29. It is a soft, malleable, and ductile metal with very high thermal and electrical conductivity. A freshly exposed surface of pure copper has a pinkis ...

rich rocks, and is associated with old copper mine localities. Species in ''Calvitimela'' are predominantly distributed in alpine to arctic regions, and they seem to have a

circumpolar distribution A circumpolar distribution is any range of a taxon that occurs over a wide range of longitudes but only at high latitudes; such a range therefore extends all the way around either the North Pole or the South Pole. Taxa that are also found in isolat ...

. On the other hand, due to lack of sampling in certain regions of the world (e.g. Africa, Asia and South America), the true distribution of ''Calvitimela'' is only partly known. ''C. austrochilensis'' is described from Chile and ''C. uniseptata'' from Antarctica, respectively. These species have not yet been included in any

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 ...

studies. Therefore, whether they belong in ''Calvitimela'' or not, is yet to be confirmed by molecular data.

Phylogeny

Molecular phylogenetics 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 ...

has revolutionized the

taxonomy Taxonomy is the practice and science of categorization or classification. A taxonomy (or taxonomical classification) is a scheme of classification, especially a hierarchical classification, in which things are organized into groups or types. ...

and revealed an extensive amount of

cryptic species 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 ...

diversity Diversity, diversify, or diverse may refer to: Business *Diversity (business), the inclusion of people of different identities (ethnicity, gender, age) in the workforce *Diversity marketing, marketing communication targeting diverse customers * ...

. In ''Calvitimela'' there are some species that are morphologically identical, but are genetically distinct and have different chemotypes. In the ''C. melaleuca''–

complex Complex commonly refers to: * Complexity, the behaviour of a system whose components interact in multiple ways so possible interactions are difficult to describe ** Complex system, a system composed of many components which may interact with each ...

at least two distinct chemical lineages are observed with no currently known morphological correlation. There have been few studies investigating ''Calvitimela'' from a

molecular phylogenetics 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 ...

perspective. Only one

phylogeny A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram or a tree showing the evolutionary relationships among various biological spec ...

focusing exclusively on ''Calvitimela'' has been published. They found ''Calvitimela'', '' Tephromela'' and '' Violella'' to constitute a well-supported

monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...

group. The relationship between the main clades remained partly unresolved, however. No study has yet included all species currently circumscribed to ''Calvitimela'', but the genus is so far indicated to be

paraphyletic In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...

References

{{Taxonbar, from=Q10440768 Lecanorales Lecanorales genera Lichen genera Taxa named by Josef Hafellner Taxa described in 2001