Calvitimela Armeniaca
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''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,
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 ...
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 ...
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. The species in ''Calvitimela'' have lecideine apothecia, 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 ...
to
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 ''Calvitimela'' is a lichen genus in the family Tephromelataceae ( Lecanorales, Lecanoromycetes). Members of the family Tephromelataceae are crustose lichens with green photobionts and lecideine or lecanorine apothecia. The species in ''Calviti ...
'' Fryday *'' Calvitimela cuprea'' Haugan & Timdal *'' Calvitimela livida'' Haugan & Timdal *'' Calvitimela melaleuca'' (Sommerf.) R. Sant *''
Calvitimela perlata ''Calvitimela'' is a lichen genus in the family Tephromelataceae ( Lecanorales, Lecanoromycetes). Members of the family Tephromelataceae are crustose lichens with green photobionts and lecideine or lecanorine apothecia. The species in ''Calviti ...
'' (Haugan & Timdal) R. Sant *''
Calvitimela septentrionalis ''Calvitimela'' is a lichen genus in the family Tephromelataceae ( Lecanorales, Lecanoromycetes). Members of the family Tephromelataceae are crustose lichens with green photobionts and lecideine or lecanorine apothecia. The species in ''Calviti ...
'' (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 ''
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, ...
''. 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 ''Tephromela'' is a genus of lichens in the family Tephromelataceae. There are about 25 species in this widespread genus. Species *'' Tephromela alectoronica'' *'' Tephromela antarctica'' *'' Tephromela arafurensis'' *''Tephromela atra ''T ...
'' 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 ...
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 ...
described. Then, a group from ''Lecidea'' was moved into the genus ''Tephromela'' M. Choisy. This led to species with both lecanorine and lecideine apothecia being present in the same genus. Finally, the genus ''Calvitimela'' was erected for species with lecideine apothecia previously placed in ''Tephromela''.
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 ...
consists of the ''
Tephromela ''Tephromela'' is a genus of lichens in the family Tephromelataceae. There are about 25 species in this widespread genus. Species *'' Tephromela alectoronica'' *'' Tephromela antarctica'' *'' Tephromela arafurensis'' *''Tephromela atra ''T ...
'', ''Calvitimela'', ''
Mycoblastus ''Mycoblastus'' is a genus of crustose lichens in the family Tephromelataceae. Members of the genus are commonly called blood lichens. Taxonomy The genus was circumscribed in 1852 by Johannes Musaeus Norman, who selected the widespread '' Mycob ...
'' Norman and ''
Violella ''Violella'' is a genus of two species of crustose lichens in the family Tephromelataceae. The genus is characterized by its brownish inner ascospore walls, brilliant violet hymenial pigment (called Fucatus-violet), and thallus chemistry. The ty ...
'' T. Sprib, and together these four genera constitute a well-supported monophyletic group.


Morphology, anatomy, and chemistry

The species of ''Calvitimela'' 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 ...
. 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 apothecia lecideine. The apothecia are convex, black and shiny. Apothecia 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 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 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. 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. ...
of
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 ...
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. In ''Calvitimela'' there are some species that are morphologically identical, but are genetically distinct and have different chemotypes. In the ''C. melaleuca''– complex 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 ''Tephromela'' is a genus of lichens in the family Tephromelataceae. There are about 25 species in this widespread genus. Species *'' Tephromela alectoronica'' *'' Tephromela antarctica'' *'' Tephromela arafurensis'' *''Tephromela atra ''T ...
'' and ''
Violella ''Violella'' is a genus of two species of crustose lichens in the family Tephromelataceae. The genus is characterized by its brownish inner ascospore walls, brilliant violet hymenial pigment (called Fucatus-violet), and thallus chemistry. The ty ...
'' 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