Micarea
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''Micarea'' is a
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 ...
of
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 ...
in the family
Pilocarpaceae The Pilocarpaceae are a family of crustose lichens in the order Lecanorales. The species of this family have a cosmopolitan distribution and have been found in a variety of climatic regions. Pilocarpaceae was circumscribed by Alexander Zahlbruck ...
. The widely distributed genus contains 126 species and new species are described actively. Species in the genus are
crustose lichen Crustose lichens are lichens that form a crust which strongly adheres to the Substrate (biology), substrate (soil, rock, tree bark, etc.), making separation from the substrate impossible without destruction. The basic structure of crustose lichen ...
s and 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 ...
.


Taxonomy

''Micarea'' was
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 ...
by
Elias Magnus Fries Elias Magnus Fries (15 August 1794 – 8 February 1878) was a Swedish mycologist and botanist. Career Fries was born at Femsjö (Hylte Municipality), Småland, the son of the pastor there. He attended school in Växjö. He acquired ...
in his 1825 work ''Systema Orbis Vegetabilis''. There have been some
taxonomic 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. ...
disputes about the genus. Nowadays, the genus is accepted, although it is
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 ...
and in need of further research. '' Micarea prasina'' is the
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specimen ...
of the genus.


Reproduction

Lichenized fungi have developed diverse reproduction strategies. The microlichen genus ''Micarea'' is an excellent model for studying the effects of reproductive traits and environmental factors on
speciation Speciation is the evolutionary process by which populations evolve to become distinct species. The biologist Orator F. Cook coined the term in 1906 for cladogenesis, the splitting of lineages, as opposed to anagenesis, phyletic evolution within ...
because it shows intricate variation in substrate requirements and reproduction modes. Certain ''Micarea'' species are generalists and able to grow on various substrata, while some are specialized and live in strict
microhabitat In ecology, the term habitat summarises the array of resources, physical and biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species habitat can be seen as the physical ...
s. Some of the ''Micarea'' species are predominately sexual, while some frequently lack sexual structures but bear numerous
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 ...
where asexual
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 produced. The actual roles of the three types of conidia present are not thoroughly understood, but mesoconidia are likely asexual
propagules In biology, a propagule is any material that functions in propagating an organism to the next stage in its life cycle, such as by dispersal. The propagule is usually distinct in form from the parent organism. Propagules are produced by organisms ...
based on, for example, the observation that many of the species are frequently found with only mesopycnidia and no
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 ...
. Recent
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
analyses together with ancestral state reconstruction among ''Micarea'' species showed that the shift in reproduction mode has evolved independently several times within the group and that facultative and obligate lignicoles are
sister species 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 ...
. This supports the assumption that the ancestor of these species was a facultative lignicole. It is presumed that a shift in substrate requirement from bark to wood leads to differentiation in reproduction mode and becomes a driver of speciation in ''Micarea'' microlichens. The case of ''Micarea'' is the first observation that among lichenized fungi reproduction mode is connected to substrate requirement. It is also the first example where such an association is demonstrated to lead to lichen speciation. The main hypothesis behind this phenomenon is that obligate species on dead wood need to colonize new suitable substrata relatively fast and asexual reproduction is a more effective strategy for successful colonisation.


Species

,
Species Fungorum ''Index Fungorum'' is an international project to index all formal names (scientific names) in the fungus kingdom. the project is based at the Royal Botanic Gardens, Kew, one of three partners along with Landcare Research and the Institute of Mi ...
accepts 126 species of ''Micarea'', although this tally does not yet include some recently described taxa, such as four species from montane
cloud forest A cloud forest, also called a water forest, primas forest, or tropical montane cloud forest (TMCF), is a generally tropical or subtropical, evergreen, montane, moist forest characterized by a persistent, frequent or seasonal low-level cloud c ...
s of Kenya. *'' Micarea adnata'' *'' Micarea aeruginoprasina'' *'' Micarea alabastrites'' *'' Micarea alectorialica'' *'' Micarea argopsinosa'' *'' Micarea assimilata'' *'' Micarea austroternaria'' – Australia *'' Micarea azorica'' *'' Micarea bacidiella'' *'' Micarea bebourensis'' *'' Micarea borbonica'' *'' Micarea boryana'' *'' Micarea botryoides'' *'' Micarea byssacea'' *'' Micarea canariensis'' *'' Micarea capitata'' *'' Micarea ceracea'' – Australasia *'' Micarea cilaosensis'' *'' Micarea cinerea'' *'' Micarea cinereopallida'' – Australasia; South America *'' Micarea contexta'' *'' Micarea coppinsii'' *'' Micarea corallothallina'' *'' Micarea coreana'' *'' Micarea curvata'' *'' Micarea czarnotae'' *'' Micarea deminuta'' *'' Micarea denigrata'' *'' Micarea doliiformis'' *'' Micarea elachista'' *'' Micarea eucalypti'' *'' Micarea eximia'' *'' Micarea fallax'' *'' Micarea farinosa'' *'' Micarea fennica'' *'' Micarea flagellispora'' – Australasia *'' Micarea flavoleprosa'' *'' Micarea globulosella'' *'' Micarea granuloblastidiata'' – Panama *'' Micarea hedlundii'' *'' Micarea herbarum'' – Europe *'' Micarea humilis'' – Australia *'' Micarea hyalinoxanthonica'' *'' Micarea hylocomii'' *'' Micarea hypoviolascens'' *'' Micarea incrassata'' *'' Micarea inquinans'' *'' Micarea intersociella'' *'' Micarea isabellina'' – Australia *'' Micarea isidioprasina'' *'' Micarea isidiosa'' *'' Micarea kartana'' – Australia *'' Micarea kemmleri'' *'' Micarea laeta'' *'' Micarea lapillicola'' *'' Micarea leprosula'' *'' Micarea levicula'' *'' Micarea lignaria'' *'' Micarea lithinella'' *'' Micarea magellanica'' *'' Micarea marginata'' *'' Micarea melaena'' *'' Micarea melaenida'' *'' Micarea melanobola'' *'' Micarea melanoprasina'' *'' Micarea meridionalis'' – Europe *'' Micarea microareolata'' *'' Micarea micrococca'' *'' Micarea micromelaena'' – Australasia *'' Micarea microsorediata'' *'' Micarea minuta'' – western Europe *'' Micarea misella'' *'' Micarea mutabilis'' – Australia *'' Micarea myriocarpa'' *'' Micarea neostipitata'' – eastern North America *'' Micarea nigella'' *'' Micarea nigra'' *'' Micarea nigrata'' *'' Micarea nitschkeana'' *'' Micarea nowakii'' *'' Micarea olivacea'' *'' Micarea oreina'' – Australasia *'' Micarea pallida'' – Australasia *'' Micarea pannarica'' *'' Micarea paratropa'' *'' Micarea parva'' *'' Micarea pauli'' *'' Micarea peliocarpa'' *'' Micarea poliocheila'' *'' Micarea polycarpella'' *'' Micarea prasina'' *'' Micarea prasinastra'' – Australasia *'' Micarea prasinella'' *'' Micarea pseudocoppinsii'' *'' Micarea pseudolignaria'' *'' Micarea pseudomarginata'' *'' Micarea pseudomicrococca'' *'' Micarea pseudotsugae'' – western Europe *'' Micarea pumila'' – Kenya *'' Micarea pusilla'' *'' Micarea pycnidiophora'' *'' Micarea rubiginosa'' – Australasia; South America *'' Micarea sandyana'' – Australasia *'' Micarea saxicola'' – Australasia *'' Micarea senecionis'' *'' Micarea sipmanii'' *'' Micarea soralifera'' *'' Micarea squamulosa'' *'' Micarea stellaris'' – Kenya *'' Micarea stereocaulorum'' *'' Micarea stipitata'' *'' Micarea subalpina'' *'' Micarea subcinerea'' *'' Micarea subconfusa'' *'' Micarea subgranulans'' *'' Micarea sublithinella'' *'' Micarea submilliaria'' *'' Micarea subnigrata'' *'' Micarea subternaria'' *'' Micarea subviridescens'' *'' Micarea synotheoides'' *'' Micarea taitensis'' – Kenya *'' Micarea takamakae'' *'' Micarea tenuispora'' *'' Micarea termitophila'' *'' Micarea ternaria'' *'' Micarea tomentosa'' *'' Micarea tubaeformis'' – Australasia *'' Micarea turfosa'' *'' Micarea usneae'' – Madeira *'' Micarea versicolor'' – Kenya *'' Micarea viridiatra'' – Europe *'' Micarea viridicapitata'' – Mexico *'' Micarea viridileprosa'' – western Europe *'' Micarea vulpinaris'' *'' Micarea xanthonica'' – Europe; North America


References

{{Taxonbar, from=Q6827942 Pilocarpaceae Lichen genera Lecanorales genera Taxa named by Elias Magnus Fries Taxa described in 1825