Teuvoa Alpina
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''Teuvoa'' is a
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 ...
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 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 ...
in the family
Megasporaceae Megasporaceae are a family of fungi belonging to the order Pertusariales. Taxa are lichenized with green algae, and grow on rocks, often in maritime climates close to fresh water. Phylogenetic In biology, phylogenetics () is the study of the ...
. It was first classified by lichenologists Mohammad Sohrabi and Steven Leavitt in 2013, with ''
Teuvoa uxoris ''Teuvoa'' is a genus of lichen-forming fungi in the family Megasporaceae. It was first classified by lichenologists Mohammad Sohrabi and Steven Leavitt in 2013, with ''Teuvoa uxoris'' assigned as the type species. This genus was delineated from ...
'' assigned as the
type species In International_Code_of_Zoological_Nomenclature, 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 spe ...
. This genus was delineated from the larger genus, ''
Aspicilia ''Aspicilia'' (sunken disk lichen) is a genus of mostly crustose lichen, crustose areolate lichen, areolate lichens that grow on rock. Most members have black apothecia discs that are slightly immersed in the areolas, hence the List of common na ...
'', following a
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 which revealed that the ''Aspicilia uxoris''
species group 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 ...
constituted a distinct lineage in the Megasporaceae. Initially containing three species, two additional species native to China were added in 2018. ''Teuvoa'' is characterised by its small and , and the absence of
secondary metabolites Secondary metabolites, also called ''specialised metabolites'', ''secondary products'', or ''natural products'', are organic compounds produced by any lifeform, e.g. bacteria, archaea, fungi, animals, or plants, which are not directly involved ...
. The physical characteristics of ''Teuvoa'' include 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 ...
thallus, forming a distinct or patchy pattern, which can range from white to grey in colour with a dull surface. The innermost layer, or
medulla Medulla (Latin for "marrow") or medullary may refer to: Science * Medulla oblongata, a part of the brain stem * Renal medulla, a part of the kidney * Adrenal medulla, a part of the adrenal gland * Medulla of ovary, a stroma in the center of the ...
, is white. The genus's , a photosynthesising partner, generally belongs to the green alga genus ''
Trebouxia ''Trebouxia'' is a unicellular green alga. It is a photosynthetic organism that can exist in almost all habitats found in polar, tropical, and temperate regions.Erokhina, L. G., Shatilovich, A. V., Kaminskaya, O. P., & Gilichinskii, D. A. (2004 ...
'' or similar genera. Reproduction in ''Teuvoa'' occurs through the formation of , cup-like structures that bear spore-producing asci. ''Teuvoa'' is distinct from related genera such as ''Aspicilia'' and '' Lobothallia'' due to a variety of features including size,
secondary metabolite Secondary metabolites, also called ''specialised metabolites'', ''secondary products'', or ''natural products'', are organic compounds produced by any lifeform, e.g. bacteria, archaea, fungi, animals, or plants, which are not directly involved ...
s, and ecological amplitude.


Taxonomy

The genus was
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 lichenologists Mohammad Sohrabi and Steven Leavitt with ''
Teuvoa uxoris ''Teuvoa'' is a genus of lichen-forming fungi in the family Megasporaceae. It was first classified by lichenologists Mohammad Sohrabi and Steven Leavitt in 2013, with ''Teuvoa uxoris'' assigned as the type species. This genus was delineated from ...
'' assigned as the
type species In International_Code_of_Zoological_Nomenclature, 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 spe ...
. ''Teuvoa'' was segregated from the large genus ''
Aspicilia ''Aspicilia'' (sunken disk lichen) is a genus of mostly crustose lichen, crustose areolate lichen, areolate lichens that grow on rock. Most members have black apothecia discs that are slightly immersed in the areolas, hence the List of common na ...
'' (family
Megasporaceae Megasporaceae are a family of fungi belonging to the order Pertusariales. Taxa are lichenized with green algae, and grow on rocks, often in maritime climates close to fresh water. Phylogenetic In biology, phylogenetics () is the study of the ...
) after
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 revealed that the ''Aspicilia uxoris''
species group 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 ...
formed a
monophyletic In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent co ...
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 ...
that represented an independent lineage within the Megasporaceae. Three species were initially placed in the genus; two additional species from China were added in 2018. ''Teuvoa'' is distinguished from ''Aspicilia'' by its small
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 ...
s and
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 ...
(the latter measuring 5–8 
μm The micrometre (Commonwealth English as used by the International Bureau of Weights and Measures; SI symbol: μm) or micrometer (American English), also commonly known by the non-SI term micron, is a unit of length in the International System ...
), and the lack of
secondary metabolite Secondary metabolites, also called ''specialised metabolites'', ''secondary products'', or ''natural products'', are organic compounds produced by any lifeform, e.g. bacteria, archaea, fungi, animals, or plants, which are not directly involved ...
s. The genus name honours Finnish lichenologist
Teuvo Ahti Teuvo "Ted" Tapio Ahti (born 14 June 1934) is a Finnish botanist and lichenologist who has made significant contributions to the taxonomy and biogeography of lichens. Known particularly for his work on the lichen family Cladoniaceae, he has h ...
, "one of the prominent lichen taxonomists of the 20th century".


Description

The crustose
thallus Thallus (: 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. A thallus usually names the entir ...
, or body, of ''Teuvoa'', forms a distinctly or patchy pattern. It can appear white to grey in colour and possesses a dull surface. Unlike some lichens, ''Teuvoa'' lacks (specialised structures for
nitrogen-fixing Nitrogen fixation is a chemical process by which molecular dinitrogen () is converted into ammonia (). It occurs both biologically and abiological nitrogen fixation, abiologically in chemical industry, chemical industries. Biological nitrogen ...
cyanobacteria) and (small pores on the lichen surface). Beneath the surface, ''Teuvoa'' has a single-layered comprising tissue, a particular type of arrangement of fungal
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. Its
medulla Medulla (Latin for "marrow") or medullary may refer to: Science * Medulla oblongata, a part of the brain stem * Renal medulla, a part of the kidney * Adrenal medulla, a part of the adrenal gland * Medulla of ovary, a stroma in the center of the ...
, or innermost layer, is white in colour and does not turn blue when exposed to
iodine Iodine is a chemical element; it has symbol I and atomic number 53. The heaviest of the stable halogens, it exists at standard conditions as a semi-lustrous, non-metallic solid that melts to form a deep violet liquid at , and boils to a vi ...
(I−). The , or photosynthesising partner, in this lichen genus can be from the
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 ( ...
genus ''
Trebouxia ''Trebouxia'' is a unicellular green alga. It is a photosynthetic organism that can exist in almost all habitats found in polar, tropical, and temperate regions.Erokhina, L. G., Shatilovich, A. V., Kaminskaya, O. P., & Gilichinskii, D. A. (2004 ...
'' or similar genera, with the algal cells being more or less spherical in shape. The reproductive structures of ''Teuvoa'', 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 ...
, form , cup-like structures bearing spore-producing asci. The of these structures ranges from black to brown-black and can be flat or occasionally concave or convex. The surrounding the disc is elevated and matches the colour of the thallus. The , or external tissue of the ascomata, is thin and can be stained by iodine (I+). Internally, the , the top layer of the
hymenium The hymenium is the tissue layer on the hymenophore of a fungal fruiting body where the cells develop into basidia or asci, which produce spores. In some species all of the cells of the hymenium develop into basidia or asci, while in oth ...
(spore-bearing tissue), presents a green to olive-brown colour. The subhymenium and (layers beneath the hymenium) are colourless but turn blue when exposed to iodine (I+ blue). The asci resemble those of the genus ''
Aspicilia ''Aspicilia'' (sunken disk lichen) is a genus of mostly crustose lichen, crustose areolate lichen, areolate lichens that grow on rock. Most members have black apothecia discs that are slightly immersed in the areolas, hence the List of common na ...
'', with their wall and not turning blue when iodine is applied (I−), while their outer coat does (I+ blue). The are simple, colourless, and can be spherical to
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 ...
in shape. Additionally, ''Teuvoa'' forms , small flask-shaped structures producing asexual spores or , which are
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 ...
(translucent), simple, and tend to be more or less straight. Species in this genus do not react with common chemical spot tests. Moreover, no
secondary metabolites Secondary metabolites, also called ''specialised metabolites'', ''secondary products'', or ''natural products'', are organic compounds produced by any lifeform, e.g. bacteria, archaea, fungi, animals, or plants, which are not directly involved ...
(
lichen product Lichen products, also known as lichen substances, are organic compounds produced by a lichen. Specifically, they are secondary metabolites. Lichen products are represented in several different chemical classes, including terpenoids, orcinol deri ...
s) that are often important for lichen identification, have been detected in ''Teuvoa''. ''Teuvoa'' is distinct from ''Aspicilia'' due to its smaller ascospore and conidia size, measuring 5–8 μm, and the lack of secondary metabolites It also differs from the genus '' Lobothallia'', in that it does not have , radiating thalli, a subhypothecial , or certain secondary metabolites, and it grows on organic substrates such as bark, wood, and dead plant debris. ''Teuvoa'' also stands apart from ''Aspicilia''
subgenus In biology, a subgenus ( subgenera) is a taxonomic rank directly below genus. In the International Code of Zoological Nomenclature, a subgeneric name can be used independently or included in a species name, in parentheses, placed between the ge ...
''Pachyothallia'' due to its absence of a subhypothecial algal layer, apothecia, certain secondary metabolites, and its different ecological amplitude, growing on organic substrates.


Habitat, ecology, and distribution

The lichen genus ''Teuvoa'' consists of five species, each having a unique habitat preference and geographical distribution. ''Teuvoa junipericola'' is primarily found across the arid continental regions of the
western United States The Western United States (also called the American West, the Western States, the Far West, the Western territories, and the West) is List of regions of the United States, census regions United States Census Bureau. As American settlement i ...
. Its distribution is notably concentrated within the
Colorado Plateau The Colorado Plateau is a physiographic and desert region of the Intermontane Plateaus, roughly centered on the Four Corners region of the Southwestern United States. This plateau covers an area of 336,700 km2 (130,000 mi2) within w ...
and
Great Basin The Great Basin () is the largest area of contiguous endorheic watersheds, those with no outlets to the ocean, in North America. It spans nearly all of Nevada, much of Utah, and portions of California, Idaho, Oregon, Wyoming, and Baja Californi ...
, regions known for their unique ecological characteristics. ''Teuvoa uxoris'' is a species with a Madrean-Thethyan disjunction, a distribution pattern typically found in plants of
Mediterranean The Mediterranean Sea ( ) is a sea connected to the Atlantic Ocean, surrounded by the Mediterranean basin and almost completely enclosed by land: on the east by the Levant in West Asia, on the north by Anatolia in West Asia and Southern ...
and Tethyan regions. Its distribution spans across vast continental Mediterranean zones that include the Mediterranean, Irano-Turanian, and Saharo-Sindian phytogeographical regions, suggesting a possible shared ecology with other lichens and
bryophyte Bryophytes () are a group of embryophyte, land plants (embryophytes), sometimes treated as a taxonomic Division (taxonomy), division referred to as Bryophyta ''Sensu#Common qualifiers, sensu lato'', that contains three groups of non-vascular pla ...
s known to inhabit these areas. In terms of habitat, ''Teuvoa uxoris'' favours coniferous trees and shrubs, including ''
Cedrus atlantica ''Cedrus atlantica'', the Atlas cedar, is a species of tree in the pine family Pinaceae, native to the Rif and Atlas Mountains of Morocco ( Middle Atlas, High Atlas), and to the Tell Atlas in Algeria.Gaussen, H. (1964). Genre ''Cedrus''. Les F ...
'', several ''
Juniperus Junipers are coniferous trees and shrubs in the genus ''Juniperus'' ( ) of the cypress family Cupressaceae. Depending on the taxonomy, between 50 and 67 species of junipers are widely distributed throughout the Northern Hemisphere as far south ...
'' species, and ''
Pinus halepensis ''Pinus halepensis'', commonly known as the Aleppo pine, also known as the Jerusalem pine, is a pine native to the Mediterranean region. It was officially named by the botanist Philip Miller in his 1768 book ''The Gardener's Dictionary''; he pro ...
''. Given these hosts' acidic bark, it is plausible that ''T. uxoris'' may also be found on other conifer species such as ''
Juniperus excelsa ''Juniperus excelsa'', commonly called the Greek juniper (though originating in Iran), is a juniper found throughout the eastern Mediterranean, from northeastern Greece and southern Bulgaria across Turkey to Syria and Lebanon, Jordan, the Caucasu ...
'', '' J. osteosperma'', '' J. polycarpos'', and '' J. sabina''. The geographical range of these
Cupressaceae Cupressaceae or the cypress family is a family of conifers. The family includes 27–30 genera (17 monotypic), which include the junipers and redwoods, with about 130–140 species in total. They are monoecious, subdioecious or (rarely) dioecio ...
is broad and includes the western and eastern Mediterranean, Minor Asia, Central Asia, Pakistan, and India. ''T. uxoris'' seems to prefer open forests in semi-arid regions, away from coastal influences, and in areas with distinct seasonal climate changes. There have been instances where ''T. uxoris'' was discovered on different substrates, indicating its capacity to adapt to varied habitats. ''Teuvoa tibetica'' is native to the Himalaya Range in Tibet. It particularly thrives in high-altitude regions, specifically between , in areas dotted with ''
Kobresia ''Carex'' is a vast genus of over 2,000 species of grass-like plants in the family Cyperaceae, commonly known as sedges (or seg, in older books). Other members of the family Cyperaceae are also called sedges, however those of genus ''Carex'' may ...
'' meadows and rocky slopes. There, it grows on soil. ''Teuvoa saxicola'' is recognised in the
Qinghai Qinghai is an inland Provinces of China, province in Northwestern China. It is the largest provinces of China, province of China (excluding autonomous regions) by area and has the third smallest population. Its capital and largest city is Xin ...
and
Xinjiang Xinjiang,; , SASM/GNC romanization, SASM/GNC: Chinese postal romanization, previously romanized as Sinkiang, officially the Xinjiang Uygur Autonomous Region (XUAR), is an Autonomous regions of China, autonomous region of the China, People' ...
provinces of northwestern China. This species displays a preference for
siliceous Silicon dioxide, also known as silica, is an oxide of silicon with the chemical formula , commonly found in nature as quartz. In many parts of the world, silica is the major constituent of sand. Silica is one of the most complex and abundant ...
rock, thriving in arid and semi-arid habitats, thereby mirroring the conditions found in the habitats of its genus companions. ''Teuvoa alpina'' is also native to northwestern China, with a distribution similar to ''T. saxicola''. However, unlike ''T. saxicola'', ''T. alpina'' favours coniferous wood as a substrate, reflecting the wide range of habitats that''Teuvoa'' species have adapted to within similar geographical regions.


Species

*'' Teuvoa alpina'' *''
Teuvoa junipericola ''Teuvoa'' is a genus of lichen-forming fungi in the family Megasporaceae. It was first classified by lichenologists Mohammad Sohrabi and Steven Leavitt in 2013, with '' Teuvoa uxoris'' assigned as the type species. This genus was delineated fro ...
'' *'' Teuvoa saxicola'' *'' Teuvoa tibetica'' *''
Teuvoa uxoris ''Teuvoa'' is a genus of lichen-forming fungi in the family Megasporaceae. It was first classified by lichenologists Mohammad Sohrabi and Steven Leavitt in 2013, with ''Teuvoa uxoris'' assigned as the type species. This genus was delineated from ...
''


References

{{Taxonbar, from=Q55579544 Megasporaceae Pertusariales genera Lichen genera Taxa described in 2013