Phoebus (lichen)
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''Phoebus hydrophobius'' is the single
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 ...
species in the fungal
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 ...
''Phoebus'', which is placed in the
family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
Lecanographaceae ''Lecanographaceae'' is a family of mostly lichens (also some lichenicolous fungi) in the order Arthoniales. The family was circumscribed in 2014, prompted by a molecular phylogenetic-based restructuring of the Arthoniales. Taxonomy The fami ...
(order
Arthoniales The Arthoniales is the second largest order of mainly crustose lichens, but fruticose lichens are present as well. The order contains around 1500 species, while the largest order with lichenized fungi, the Lecanorales, contains more than 14000 s ...
). This distinctive lichen was discovered in 2007 in the Ozark region of the central United States and later found in
limestone Limestone is a type of carbonate rock, carbonate sedimentary rock which is the main source of the material Lime (material), lime. It is composed mostly of the minerals calcite and aragonite, which are different Polymorphism (materials science) ...
areas of northeastern Mexico. It forms orange-brown crusty patches on sun-exposed rock faces and can be identified by its intense violet-black color reaction when treated with
potassium hydroxide Potassium hydroxide is an inorganic compound with the formula K OH, and is commonly called caustic potash. Along with sodium hydroxide (NaOH), KOH is a prototypical strong base. It has many industrial and niche applications, most of which utili ...
solution, a standard chemical spot test.


Taxonomy

Genus ''Phoebus'' contains only one species, ''Phoebus hydrophobius'', found in the
Ozarks The Ozarks, also known as the Ozark Mountains, Ozark Highlands or Ozark Plateau, is a physiographic region in the U.S. states of Missouri, Arkansas, and Oklahoma, as well as a small area in the southeastern corner of Kansas. The Ozarks cover ...
of the central United States, and described as new to science in 2007. When erecting the genus, Harris and Ladd compared its , intensely K-positive thallus with that of certain orange members of the
Roccellaceae The Roccellaceae are a family (biology), family of mostly lichen-forming fungi in the order Arthoniales, established by the French botanist François Fulgis Chevallier in 1826. Species in the family exhibit various lichen growth forms, growth ...
and provisionally placed the new genus in that family. Subsequent check-lists that lacked DNA evidence transferred ''Phoebus'' to "Arthoniales genera ''
incertae sedis or is a term used for a taxonomy (biology), taxonomic group where its broader relationships are unknown or undefined. Alternatively, such groups are frequently referred to as "enigmatic taxa". In the system of open nomenclature, uncertainty ...
''", reflecting uncertainty over its deeper relationships; as of 2025, it is still classified in this way by
Species Fungorum ''Index Fungorum'' is an international project to index all formal names (Binomial nomenclature, scientific names) in the fungus Kingdom (biology), kingdom. As of 2015, the project is based at the Royal Botanic Gardens, Kew, one of three partn ...
. A three- locus phylogeny (nuLSU + mtSSU + RPB2) published in 2020 resolved the species within Lecanographaceae, as an isolated lineage that is
sister A sister is a woman or a girl who shares parents or a parent with another individual; a female sibling. The male counterpart is a brother. Although the term typically refers to a familial relationship, it is sometimes used endearingly to ref ...
to a
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 ...
containing ''Phacographa'' spp., two "''Opegrapha''" species and '' Simonyella variegata''. This placement makes ''Phoebus'' the eighth confirmed genus in the family and shows that its placodioid
growth form Plant life-form schemes constitute a way of classifying plants alternatively to the ordinary species-genus-family scientific classification. In colloquial speech, plants may be classified as trees, shrubs, herbs (forbs and graminoids), etc. The sci ...
evolved independently of the superficially similar genus '' Roccellina''. The study also extended the known range of ''P. hydrophobius'' to six
xerophytic A xerophyte () is a species of plant that has adaptations to survive in an environment with little liquid water. Examples of xerophytes include cactus, cacti, pineapple and some gymnosperm plants. The morphology (biology), morphology and physiology ...
limestone localities in the
Sierra Madre Oriental The Sierra Madre Oriental () is a mountain range in northeastern Mexico. The Sierra Madre Oriental is part of the American Cordillera, a chain of mountain ranges (cordillera) that consists of an almost continuous sequence of mountain ranges that ...
of
Coahuila Coahuila, formally Coahuila de Zaragoza, officially the Free and Sovereign State of Coahuila de Zaragoza, is one of the 31 states of Mexico. The largest city and State Capital is the city of Saltillo; the second largest is Torreón and the thi ...
and
Nuevo León Nuevo León, officially the Free and Sovereign State of Nuevo León, is a Administrative divisions of Mexico, state in northeastern Mexico. The state borders the Mexican states of Tamaulipas, Coahuila, Zacatecas, and San Luis Potosí, San Luis ...
, Mexico, demonstrating that the species is not restricted to the Ozarks.


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 ...
of ''Phoebus hydrophobius'' forms an orange-brown crust made up of small, slightly swollen polygonal blocks () that coalesce into rough patches up to about a millimetre thick. When a drop of
potassium hydroxide Potassium hydroxide is an inorganic compound with the formula K OH, and is commonly called caustic potash. Along with sodium hydroxide (NaOH), KOH is a prototypical strong base. It has many industrial and niche applications, most of which utili ...
solution (the standard "K" spot test) is placed on the surface, those blocks turn a deep violet-black, betraying the presence of
anthraquinone Anthraquinone, also called anthracenedione or dioxoanthracene, is an aromatic hydrocarbon, aromatic organic compound with formula . Several isomers exist but these terms usually refer to 9,10-anthraquinone (IUPAC: 9,10-dioxoanthracene) wherein th ...
-like pigments. In section the outer skin () is a firm, layer of tightly interwoven, gelatinised fungal threads; above it sits a thin, jelly-like tinged the same orange as the surface. Unusually, the white medulla—the cottony interior of the lichen—occurs both above and below the green algal band, giving the thallus a double-decker construction. The algal partner is '' Trentepohlia'', a group of orange-pigmented, sausage-shaped cells that typically favour humid, sheltered rock faces. Black, slit-to-irregularly rounded fruiting bodies ( apothecia) erupt on or between the areoles. They start flush with the surface but soon become slightly elevated, with a coarse black rim and, at times, a dusting of orange powder. Inside, a dark brown underlies a 100-
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 ...
-tall spore-bearing layer (
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 ...
). The asci each contain eight
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 arranged in two rows; the spores are colourless at first, surrounded by a gelatinous , divided by three cross-walls, and measure roughly 16–20 × 5–6.5 
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 ...
before maturing to a pale brown and developing a roughened coat. No
asexual Asexual or Asexuals may refer to: *Asexual reproduction **Asexual reproduction in starfish *Asexuality, the lack of sexual attraction to anyone or lack of interest in or desire for sexual activity. **Gray asexuality, the spectrum between asexualit ...
propagules (
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 ...
) have been observed. This combination of an intensely K-positive orange crust, ''Trentepohlia'' photobiont, and opegraphoid apothecia separates ''Phoebus'' from superficially similar orange rock-dwelling lichens such as '' Caloplaca''.


Habitat and distribution

''Phoebus hydrophobius'' was originally known only from
dolomitic Dolomite () is an anhydrous carbonate mineral composed of calcium magnesium carbonate, ideally The term is also used for a sedimentary carbonate rock composed mostly of the mineral dolomite (see Dolomite (rock)). An alternative name sometimes ...
and other
carbonate A carbonate is a salt of carbonic acid, (), characterized by the presence of the carbonate ion, a polyatomic ion with the formula . The word "carbonate" may also refer to a carbonate ester, an organic compound containing the carbonate group ...
bluffs in the
Missouri Missouri (''see #Etymology and pronunciation, pronunciation'') is a U.S. state, state in the Midwestern United States, Midwestern region of the United States. Ranking List of U.S. states and territories by area, 21st in land area, it border ...
–
Arkansas Arkansas ( ) is a landlocked state in the West South Central region of the Southern United States. It borders Missouri to the north, Tennessee and Mississippi to the east, Louisiana to the south, Texas to the southwest, and Oklahoma ...
Ozark Highlands, where the type was collected in 2007. Targeted surveys and molecular work later revealed the species on six additional
limestone Limestone is a type of carbonate rock, carbonate sedimentary rock which is the main source of the material Lime (material), lime. It is composed mostly of the minerals calcite and aragonite, which are different Polymorphism (materials science) ...
outcrops in the Sierra Madre Oriental of north-eastern Mexico (one site in Coahuila, five in Nuevo León), showing that its range bridges the southern United States and northern Mexico rather than being
Ozark The Ozarks, also known as the Ozark Mountains, Ozark Highlands or Ozark Plateau, is a physiographic region in the U.S. states of Missouri, Arkansas, and Oklahoma, as well as a small area in the southeastern corner of Kansas. The Ozarks cov ...
-restricted. Across this expanded range the lichen occupies sun-exposed,
xeric Deserts and xeric shrublands are a biome defined by the World Wide Fund for Nature. Deserts and xeric (Ancient Greek 'dry') shrublands form the largest terrestrial biome, covering 19% of Earth's land surface area. Ecoregions in this habita ...
faces of hard limestone—both vertical canyon walls and sloping pavements—within xerophytic and
piedmont Piedmont ( ; ; ) is one of the 20 regions of Italy, located in the northwest Italy, Northwest of the country. It borders the Liguria region to the south, the Lombardy and Emilia-Romagna regions to the east, and the Aosta Valley region to the ...
scrub vegetation at roughly elevation. It often grows intermixed with other calcicolous crusts such as '' Alyxoria sierramadrensis'' and various '' Caloplaca'', ''
Buellia ''Buellia'' is a genus of mostly lichen-forming fungi in the family Caliciaceae. The fungi are usually part of a crustose lichen. In this case, the lichen species is given the same name as the fungus. But members may also grow as parasites on lic ...
'' and ''
Squamulea ''Squamulea'' is a genus of lichen-forming fungi in the family Teloschistaceae. It has 15 species. The genus was circumscribed in 2013 by Ulf Arup, Ulrik Søchting, and Patrik Frödén, with '' Squamulea subsoluta'' assigned as the type species. ...
'' species, and appears locally common at several Mexican sites. Most Nuevo León populations lie on heavily visited mountains around Monterrey, where urban expansion,
quarry A quarry is a type of open-pit mining, open-pit mine in which dimension stone, rock (geology), rock, construction aggregate, riprap, sand, gravel, or slate is excavated from the ground. The operation of quarries is regulated in some juri ...
ing of limestone, and chronic industrial
air pollution Air pollution is the presence of substances in the Atmosphere of Earth, air that are harmful to humans, other living beings or the environment. Pollutants can be Gas, gases like Ground-level ozone, ozone or nitrogen oxides or small particles li ...
pose potential threats to the species' limited habitat.


References

{{Taxonbar , from1=Q7186699 , from2=Q20963090 Arthoniomycetes Fungi of Mexico Fungi of the United States Fungi described in 2007 Fungus species Taxa named by Richard Clinton Harris