Circinaria Digitata
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''Circinaria digitata'' is a species of
fruticose lichen A fruticose lichen is a form of lichen fungi that is characterized by a coral-like shrubby or bushy growth structure. It is formed from a symbiotic relationship of a photobiont such as green algae or less commonly cyanobacteria and one, two or m ...
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 ...
. This small, soil-dwelling species is a vagrant lichen, and grows as loose, finger-like tufts that roll freely across the ground. It is restricted to high-elevation valleys of the
Tian Shan The Tian Shan, also known as the Tengri Tagh or Tengir-Too, meaning the "Mountains of God/Heaven", is a large system of mountain ranges in Central Asia. The highest peak is Jengish Chokusu at high and located in Kyrgyzstan. Its lowest point is ...
mountain range in
Kyrgyzstan Kyrgyzstan, officially the Kyrgyz Republic, is a landlocked country in Central Asia lying in the Tian Shan and Pamir Mountains, Pamir mountain ranges. Bishkek is the Capital city, capital and List of cities in Kyrgyzstan, largest city. Kyrgyz ...
, where it forms part of the so-called "manna" lichens – vagrant species whose
thalli 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 ...
are not anchored to a
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 ...
. The species was originally described in 2010 as ''Aspicilia digitata'' and transferred to the genus ''
Circinaria ''Circinaria'' is a genus of lichen-forming fungi in the family Megasporaceae. It was circumscribed by Johann Heinrich Friedrich Link in 1809. Species , Species Fungorum ''Index Fungorum'' is an international project to index all formal na ...
'' after
molecular A molecule is a group of two or more atoms that are held together by attractive forces known as chemical bonds; depending on context, the term may or may not include ions that satisfy this criterion. In quantum physics, organic chemistry, ...
work clarified relationships within the Megasporaceae.


Taxonomy

''Circinaria digitata'' was first
formally described A species description is a formal scientific description of a newly encountered species, typically articulated through a scientific publication. Its purpose is to provide a clear description of a new species of organism and explain how it differ ...
in 2010 as ''Aspicilia digitata'' by Mohammad Sohrabi and Birgit Litterski, based on vagrant thalli collected at 2,900 m on gravel terraces above the Jal-Jyr River in the central
Tian Shan The Tian Shan, also known as the Tengri Tagh or Tengir-Too, meaning the "Mountains of God/Heaven", is a large system of mountain ranges in Central Asia. The highest peak is Jengish Chokusu at high and located in Kyrgyzstan. Its lowest point is ...
. The authors contrasted its narrowly , intricately branched and the distinctive black spots at the lobe tips with the broader, more robust branches of the superficially similar '' A. fruticulosa''. In a comprehensive revision of the "manna lichens" the following year, Sohrabi transferred the species to the resurrected genus ''Circinaria'', publishing the
new combination In Taxonomy (biology), biological taxonomy, a combinatio nova (abbreviated comb. nov. or n. comb.) refers to the formal renaming of an organism's scientific name when it is transferred to a different genus, reclassified within a different specie ...
''Circinaria digitata''. That same paper synonymised the vagrant genera ''Agrestia'', ''Chlorangium'' and ''Sphaerothallia'' under a broadly circumscribed ''Circinaria'', leaving no unattached ("vagrant") species in ''Aspicilia''. The original
holotype A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...
(H 5185-A) is housed in the
Finnish Museum of Natural History The Finnish Museum of Natural History (, ), established in 1988, is a research institution under the University of Helsinki in Finland, based in Helsinki, Finland. It is a natural history museum responsible for the national botanical, zoological, ...
, with an isotype (duplicate) preserved in
Greifswald Greifswald (), officially the University and Hanseatic City of Greifswald (, Low German: ''Griepswoold'') is the fourth-largest city in the German state of Mecklenburg-Western Pomerania after Rostock, Schwerin and Neubrandenburg. In 2021 it surpa ...
(GFW). Multilocus phylogenetic analyses (nrITS, nrLSU and mtSSU rDNA) show that ''C. digitata'' belongs to a strongly supported "sphaerothallioid"
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 contains other vagrant or erratic members of the Megasporaceae; within this group it clusters with ''C. elmorei'' and ''C. hispida'' rather than with the morphologically alike ''C. fruticulosa''. The study further demonstrates that several diagnostic features traditionally relied upon for generic delimitation—such as the presence of a two-layered cortex—have evolved more than once inside ''Circinaria'' and therefore lack
phylogenetic In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...
significance. Together these results support treating ''C. digitata'' as a specialised, vagrant offshoot within a morphologically heterogeneous but
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 ...
''Circinaria''.


Description

The
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 ...
forms minute, cushion-like balls about 0.5–1.5 cm across. Each tuft consists of hundreds of cylindrical branchlets that radiate from a narrow basal core. Individual are 0.45–1 mm thick, irregularly divided and often curve back upon themselves, giving the whole thallus a coral-like appearance. Their upper surface is dull dark-green to grey-green, occasionally with pale olive tones. Near the tips the cortex breaks to form tiny white
pseudocyphella Pseudocyphellae (singular ''pseudocyphella'') are structures in lichens that appear as tiny pores on the outer surface (the cortex) of the lichen. They are caused when there is a break in the cortex of the lichen, and the medullary hyphae extend ...
e—microscopic pores that assist
gas exchange Gas exchange is the physical process by which gases move passively by diffusion across a surface. For example, this surface might be the air/water interface of a water body, the surface of a gas bubble in a liquid, a gas-permeable membrane, or a b ...
—and the very apices usually bear a charcoal-black spot interpreted as a sterile
pycnidium 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 ...
(a flask-shaped reproductive cavity). Cross-sections show a two-layered : an outer band of tightly packed, brownish cells (25–35 
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 ...
thick) and an inner zone up to 100 Î¼m thick. A cluster of
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 ( ...
l cells (5–15 Î¼m diameter) lies immediately beneath the cortex, enveloping a white, loosely woven
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 ...
. No apothecia have yet been observed, and standard chemical spot tests as well as thin-layer and
high-performance liquid chromatography High-performance liquid chromatography (HPLC), formerly referred to as high-pressure liquid chromatography, is a technique in analytical chemistry used to separate, identify, and quantify specific components in mixtures. The mixtures can origin ...
analyses failed to detect any
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.


Habitat and distribution

''Circinaria digitata'' is a strictly
terricolous A terricolous lichen is a lichen that grows on the soil as a substrate. Examples include some members of the genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classificati ...
(ground-dwelling) species: its tufts rest on gravelly river terraces or among coarse alpine
steppe In physical geography, a steppe () is an ecoregion characterized by grassland plains without closed forests except near rivers and lakes. Steppe biomes may include: * the montane grasslands and shrublands biome * the tropical and subtropica ...
soils rather than attaching to rock or plant material. Field observations place it between 2900 m and 3100 m in subalpine belts of the Jangy-Jer and Moldo Too ridges, where summer conditions are windy, cool and arid. The thalli are free-moving; wind or water can roll them across the stony surface, a strategy that disperses fragments and allows the lichen to colonise sparsely vegetated ground. At the time of its original publication, the species was known only from two closely spaced localities in central Kyrgyzstan, suggesting a narrow distribution and possible
endemism Endemism is the state of a species being found only in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also foun ...
to the inner Tian-Shan. Its apparent confinement to high-elevation
river valley A valley is an elongated low area often running between hills or mountains and typically containing a river or stream running from one end to the other. Most valleys are formed by erosion of the land surface by rivers or streams over a ...
s contrasts with related vagrant species such as '' C. fruticulosa'' and '' C. hispida'', which occupy drier lowland steppes across Eurasia and North America.


References

{{Taxonbar , from1=Q22108742 , from2=Q17387335 Megasporaceae Lichen species Lichens described in 2010 Lichens of Western Asia