Teloschistaceae
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The Teloschistaceae are a large
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 ...
of mostly
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 ...
belonging to the
class Class, Classes, or The Class may refer to: Common uses not otherwise categorized * Class (biology), a taxonomic rank * Class (knowledge representation), a collection of individuals or objects * Class (philosophy), an analytical concept used d ...
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. The group is ...
in the division
Ascomycota Ascomycota is a phylum of the kingdom Fungi that, together with the Basidiomycota, forms the subkingdom Dikarya. Its members are commonly known as the sac fungi or ascomycetes. It is the largest phylum of Fungi, with over 64,000 species. The def ...
. The family has a
cosmopolitan distribution In biogeography, a cosmopolitan distribution is the range of a taxon that extends across most or all of the surface of the Earth, in appropriate habitats; most cosmopolitan species are known to be highly adaptable to a range of climatic and en ...
, although its members occur predominantly in
temperate In geography, the temperate climates of Earth occur in the middle latitudes (approximately 23.5° to 66.5° N/S of the Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ran ...
regions. Most members are lichens that either live on rock or on bark, but about 40 species are
lichenicolous This glossary of mycology is a list of definitions of terms and concepts relevant to mycology, the study of fungi. Terms in common with other fields, if repeated here, generally focus on their mycology-specific meaning. Related terms can be found ...
meaning they are non-lichenised fungi that live on other lichens. Many members of the Teloschistaceae are readily identifiable by their vibrant orange to yellow hue, a result of their frequent
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 ...
content. The presence of these anthraquinone
pigments A pigment is a powder used to add or alter color or change visual appearance. Pigments are completely or nearly solubility, insoluble and reactivity (chemistry), chemically unreactive in water or another medium; in contrast, dyes are colored sub ...
, which confer protection from
ultraviolet Ultraviolet radiation, also known as simply UV, is electromagnetic radiation of wavelengths of 10–400 nanometers, shorter than that of visible light, but longer than X-rays. UV radiation is present in sunlight and constitutes about 10% of ...
light, enabled this group to expand from shaded forest habitats to harsher environmental conditions of sunny and
arid Aridity is the condition of geographical regions which make up approximately 43% of total global available land area, characterized by low annual precipitation, increased temperatures, and limited water availability.Perez-Aguilar, L. Y., Plata ...
ecosystems during the
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the more recent of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''cre ...
. Teloschistaceae lichens typically have one of a few physical
growth forms 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 ...
. Depending on the species, 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 ...
(the main body of the lichen) is either leaf-like (
foliose A foliose lichen is a lichen with flat, leaf-like , which are generally not firmly bonded to the substrate on which it grows. It is one of the three most common lichen growth forms, growth forms of lichens. It typically has distinct upper and lo ...
), bushy or shrub-like (
fruticose A fruticose lichen is a form of lichen fungi that is characterized by a coral-like shrubby or bushy lichen growth forms, growth structure. It is formed from a symbiotic relationship of a photobiont such as green algae or less commonly cyanobacteri ...
), or crust-like (
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 ...
). These lichens typically partner with a
photosynthetic Photosynthesis ( ) is a Biological system, system of biological processes by which Photoautotrophism, photosynthetic organisms, such as most plants, algae, and cyanobacteria, convert light energy, typically from sunlight, into the chemical ener ...
companion (a ) from the green algal 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 ...
''. Teloschistaceae members are also characterised by their (the
fruiting bodies The sporocarp (also known as fruiting body, fruit body or fruitbody) of fungi is a multicellular structure on which spore-producing structures, such as basidia or asci, are borne. The fruitbody is part of the sexual phase of a fungal life cyc ...
where sexual reproduction occurs), which generally have a well-defined . In the Teloschistaceae, the tip of the
ascus An ascus (; : asci) is the sexual spore-bearing cell produced in ascomycete fungi. Each ascus usually contains eight ascospores (or octad), produced by meiosis followed, in most species, by a mitotic cell division. However, asci in some gen ...
, the structure that produces spores, characteristically turns blue when stained with
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 ...
. The
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 are released through a longitudinal slit in the ascus tip, a unique trait common to this family of lichens. The family, first formally proposed in 1898, was extensively revised in 2013, including the creation or resurrection of 31
genera Genus (; : genera ) is a taxonomic rank above species and below family as used in the biological classification of living and fossil organisms as well as viruses. In binomial nomenclature, the genus name forms the first part of the binomial s ...
. Three subfamilies – Caloplacoideae, Teloschistoideae, and Xanthorioideae – are recognised. Since 2013, several dozen new genera have been added to the family, but there has been some debate about these additions. Ongoing
DNA studies Deoxyribonucleic acid (; DNA) is a polymer composed of two polynucleotide chains that coil around each other to form a double helix. The polymer carries genetic instructions for the development, functioning, growth and reproduction of al ...
are helping to provide clearer insights into how the different groups within this family are related. The family contains more than 800 species in around 120 genera. Three species from the Teloschistaceae have been globally assessed for
conservation status The conservation status of a group of organisms (for instance, a species) indicates whether the group still exists and how likely the group is to become extinct in the near future. Many factors are taken into account when assessing conservation ...
and others, such as the rare New Zealand species ''
Caloplaca allanii ''Caloplaca'' is a lichen genus comprising a number of distinct species. Members of the genus are commonly called firedot lichen, jewel lichen.Field Guide to California Lichens, Stephen Sharnoff, Yale University Press, 2014, gold lichens, "ora ...
'', appear on regional lists. The full diversity of this family remains underexplored in vast regions like South America and China. Regarding human interactions and applications, although lacking any major economic impact, several rock-dwelling Teloschistaceae species are known to damage
marble Marble is a metamorphic rock consisting of carbonate minerals (most commonly calcite (CaCO3) or Dolomite (mineral), dolomite (CaMg(CO3)2) that have recrystallized under the influence of heat and pressure. It has a crystalline texture, and is ty ...
surfaces, and others are used in some
traditional medicine Traditional medicine (also known as indigenous medicine or folk medicine) refers to the knowledge, skills, and practices rooted in the cultural beliefs of various societies, especially Indigenous groups, used for maintaining health and treatin ...
s. One member, '' Rusavskia elegans'', is used in research as a
model organism A model organism is a non-human species that is extensively studied to understand particular biological phenomena, with the expectation that discoveries made in the model organism will provide insight into the workings of other organisms. Mo ...
to investigate resilience against the harsh conditions of outer space.


Systematics


Historical taxonomy

The first members of the present-day Teloschistaceae to be
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 ...
were the common sunburst lichen (''
Xanthoria parietina ''Xanthoria parietina'' is a common and widespread lichen-forming fungus in the family Teloschistaceae. Commonly known as the yellow wall lichen, common orange lichen, or maritime sunburst lichen, this foliose lichen, leafy lichen is known f ...
'') and the gold-eye lichen ('' Teloschistes chrysophthalmus''). These were two of several dozen lichen species described by the Swedish taxonomist
Carl Linnaeus Carl Linnaeus (23 May 1707 – 10 January 1778), also known after ennoblement in 1761 as Carl von Linné,#Blunt, Blunt (2004), p. 171. was a Swedish biologist and physician who formalised binomial nomenclature, the modern system of naming o ...
, the former in his influential 1753 treatise , and the latter in his 1771 work . In his 1852 work (), the lichenologist
Abramo Bartolommeo Massalongo Abramo Bartolommeo Massalongo (13 May 1824 – 25 May 1860) was an Italian paleobotanist and lichenologist. He was born in Tregnago in the Province of Verona and took a great interest in botany as a young man. Massalongo joined the faculty of med ...
attempted to classify what he called "blasteniospore lichens". This term referenced species, diverse in
growth forms 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 ...
and appearance, united by the distinct spores now attributed to the family Teloschistaceae. These are spores that are divided into two compartments () separated by a central
septum In biology, a septum (Latin language, Latin for ''something that encloses''; septa) is a wall, dividing a Body cavity, cavity or structure into smaller ones. A cavity or structure divided in this way may be referred to as septate. Examples Hum ...
with a small hole. Although Massalongo's efforts to arrange these
taxa In biology, a taxon (back-formation from ''taxonomy''; : taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular name and ...
into more natural genera were largely ignored by subsequent researchers, several of his proposed genera were resurrected for use 16 decades later, such as '' Blastenia'', '' Gyalolechia'', '' Pyrenodesmia'', and '' Xanthocarpia''. 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 ...
Teloschistaceae was formally
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 ...
by the lichenologist
Alexander Zahlbruckner Alexander Zahlbruckner (31 May 1860, Svätý Jur – 1938, Vienna) was an Austrian- Hungarian botanist who specialized in the study of lichens. Johann Babtist Zahlbruckner, an earlier Austrian botanist, was his grandfather. From 1878 to 1883 ...
in 1898. In his initial version, he grouped together
foliose A foliose lichen is a lichen with flat, leaf-like , which are generally not firmly bonded to the substrate on which it grows. It is one of the three most common lichen growth forms, growth forms of lichens. It typically has distinct upper and lo ...
and
fruticose A fruticose lichen is a form of lichen fungi that is characterized by a coral-like shrubby or bushy lichen growth forms, growth structure. It is formed from a symbiotic relationship of a photobiont such as green algae or less commonly cyanobacteri ...
taxa having polarilocular (i.e. two-locule) or four-locule , including the genera '' Xanthoria'', '' Teloschistes'', and ''Lethariopsis''. At that time, the growth form of the lichen was often used in classical lichen taxonomy to segregate groups of species into families, and so in a subsequent (1926) publication, Zahlbruckner introduced the family Caloplacaceae to contain crustose lichens with polarilocular ascospores; this family included the genera ''
Caloplaca ''Caloplaca'' is a lichen genus comprising a number of distinct species. Members of the genus are commonly called firedot lichen, jewel lichen.Field Guide to California Lichens, Stephen Sharnoff, Yale University Press, 2014, gold lichens, "ora ...
'', ''Blastenia'', ''Bombyliospora'', and ''Protoblastenia''. The distinctness of the family Caloplacaceae was largely rejected by other authors, and it is now a historical
synonym A synonym is a word, morpheme, or phrase that means precisely or nearly the same as another word, morpheme, or phrase in a given language. For example, in the English language, the words ''begin'', ''start'', ''commence'', and ''initiate'' are a ...
of Teloschistaceae. In another older classification, crustose genera were grouped together in the family Blasteniaceae or the Placodiaceae. In 1971,
Carroll William Dodge Carroll William Dodge (January 20, 1895 – July 21, 1988) was an American mycologist and lichenologist. His major fields of study included human and mammalian parasitic fungi, lichen-associated fungi, and fungi forming subterranean sporophore ...
proposed the family Xanthoriaceae to contain ''Xanthodactylon'', ''Xanthopeltis'', and ''Xanthoria'', but it was not
validly published In botanical nomenclature, a validly published name is a name that meets the requirements in the ''International Code of Nomenclature for algae, fungi, and plants'' (''ICN'') for valid publication. Valid publication of a name represents the minim ...
. In the 20th century, particularly with the widespread use of
electron microscopy An electron microscope is a microscope that uses a beam of electrons as a source of illumination. It uses electron optics that are analogous to the glass lenses of an optical light microscope to control the electron beam, for instance focusing i ...
, the details of ascus structure became important considerations in the taxonomy of lichen-forming fungi. Studies on several Teloschistaceae species have noted the consistent presence of a cap-like zone at the tip of the ascus that shows a strong reaction 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 ...
, characteristic of
amyloid Amyloids are aggregates of proteins characterised by a fibrillar morphology of typically 7–13 nm in diameter, a β-sheet secondary structure (known as cross-β) and ability to be stained by particular dyes, such as Congo red. In the human ...
substances. Using advanced
transmission electron microscopy Transmission electron microscopy (TEM) is a microscopy technique in which a beam of electrons is transmitted through a specimen to form an image. The specimen is most often an ultrathin section less than 100 nm thick or a suspension on a g ...
, Rosmarie Honegger confirmed a unique type of
ascus An ascus (; : asci) is the sexual spore-bearing cell produced in ascomycete fungi. Each ascus usually contains eight ascospores (or octad), produced by meiosis followed, in most species, by a mitotic cell division. However, asci in some gen ...
in Teloschistaceae, later named the ''Teloschistes''-type. This ascus is distinguished by a special outer layer that reacts to certain stains and lacks the typical structures seen at the tip, opening in an unusual pattern during spore release. The presence of this ascus type was later used as a diagnostic for the family Teloschistaceae following an
ultrastructural Ultrastructure (or ultra-structure) is the architecture of cells and biomaterials that is visible at higher magnifications than found on a standard optical light microscope. This traditionally meant the resolution and magnification range of a c ...
study that corroborated Honegger's work. In 1989
Ingvar Kärnefelt Jan Eric Ingvar Kärnefelt (born 1944) is a Swedish lichenologist. Early life and education Kärnefelt was born in Gothenburg, Sweden in 1944. His initial goal in his higher-level studies at University of Cologne in 1966–1967 was to become a ...
revised the family, accepting ten genera, and this served as the main taxonomic classification for the family until the
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, ...
era. In one of the last classifications of the family before the widespread use and implementation of molecular techniques, the ''Outline of the Ascomycota'' accepted 12 genera in Teloschistaceae in 2006: ''Caloplaca'', ''Cephalophysis'', ''Fulgensia'', ''Huea'', ''Ioplaca'', ''Josefpoeltia'', ''Seirophora'', ''Teloschistes'', ''Xanthodactylon'', ''Xanthomendoza'', ''Xanthopeltis'', and ''Xanthoria''. The family continues to undergo significant changes. For example, in 2020, of all fungal families, Teloschistaceae had the fourth-highest number of new fungal names (a total of 128), including 8 genera, 48 new species and infraspecific taxa, and 72
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 ...
s.


Etymology

As is standard practice in
botanical nomenclature Botanical nomenclature is the formal, scientific naming of plants. It is related to, but distinct from taxonomy (biology), taxonomy. Plant taxonomy is concerned with grouping and classifying plants; Botany, botanical nomenclature then provides na ...
, the name ''Teloschistaceae'' is based on the name of the
type genus In biological taxonomy, the type genus (''genus typica'') is the genus which defines a biological family and the root of the family name. Zoological nomenclature According to the International Code of Zoological Nomenclature, "The name-bearin ...
, '' Teloschistes'', with the ending indicating the
rank A rank is a position in a hierarchy. It can be formally recognized—for example, cardinal, chief executive officer, general, professor—or unofficial. People Formal ranks * Academic rank * Corporate title * Diplomatic rank * Hierarchy ...
of family. The genus name, assigned by the Norwegian botanist Johannes M. Norman in 1852, comprises two
Ancient Greek Ancient Greek (, ; ) includes the forms of the Greek language used in ancient Greece and the classical antiquity, ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Greek ...
words: (), meaning , , or ; and (), meaning , , or . It refers to the split ends of the thallus branches that are characteristic of that genus.


Subfamilial and ordinal classification

Teloschistaceae is divided into three recognised subfamilies: Xanthorioideae, Caloplacoideae, and Teloschistoideae. In 2015, researchers proposed a fourth subfamily, Brownlielloideae, which was later shown by genetic studies to be a grouping based on mixed or misinterpreted data rather than a distinct lineage. Further analysis placed what was thought to be Brownlielloideae within the already established Teloschistoideae, suggesting the proposed subfamily was not a separate branch of the family tree. DNA evidence also dispersed members of the informally introduced subfamily Ikaerioideae across the three acknowledged subfamilies, primarily within Teloschistoideae. Despite this, Sergey Kondratyuk and colleagues continue to use Brownlielloideae and Ikaerioideae in their publications, assigning nine genera to the former and two to the latter. The well-supported subfamilies (Xanthorioideae, Caloplacoideae, and Teloschistoideae) encompass a range of growth formscrustose, foliose, and fruticosedemonstrating the diverse evolutionary paths within the family. These groups are genetically distinct, each subfamily showing unique patterns in their nuclear large ribosomal subunit RNA sequences. *Caloplacoideae **Type genus: ''
Caloplaca ''Caloplaca'' is a lichen genus comprising a number of distinct species. Members of the genus are commonly called firedot lichen, jewel lichen.Field Guide to California Lichens, Stephen Sharnoff, Yale University Press, 2014, gold lichens, "ora ...
''. Proposed by Ester Gaya and colleagues in 2012 and
validly published In botanical nomenclature, a validly published name is a name that meets the requirements in the ''International Code of Nomenclature for algae, fungi, and plants'' (''ICN'') for valid publication. Valid publication of a name represents the minim ...
in 2020, Caloplacoideae consists mostly of crustose lichens with a wide geographical spread and produces a range of unique chemical compounds. *Teloschistoideae **Type genus: '' Teloschistes''. Initially proposed in 2013 and validly published with a full diagnosis in 2020, this subfamily is predominantly found in the Southern Hemisphere. *Xanthorioideae **Type genus: '' Xanthoria''. Named by Gaya and colleagues in 2012 and formally validated in 2020, Xanthorioideae species are primarily distributed in the Northern Hemisphere. The order
Teloschistales The Teloschistales are an order of mostly lichen-forming fungi belonging to the class Lecanoromycetes in the division Ascomycota. According to one 2008 estimate, the order contains 5 families, 66 genera, and 1954 species. The predominant photob ...
was first proposed by David Hawksworth and Eriksson in 1986, with a single family (Teloschistaceae); other families were added later. In the 1990s, several authors recognised the Teloschistales as a suborder within the
Lecanorales The Lecanorales are an order of mostly lichen-forming fungi belonging to the class Lecanoromycetes in the division Ascomycota. The order contains 26 families, 269 genera, and 5695 species. Families Suborder Lecanorineae * Biatorellacea ...
; as a suborder it was named ''Teloschistineae''. Following the appearance of preliminary molecular studies, the Teloschistaceae was classified by some within the order Lecanorales, although others maintained the Teloschistales as a valid order. A large-scale, multigene phylogenetic study of the class
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. The group is ...
published in 2014 corroborated the ordinal status of the Teloschistales, and showed it comprises two clades: Letrouitineae (containing Brigantiaeaceae and Letrouitiaceae) and its sister clade, Teloschistineae (containing Teloschistaceae and Megalosporaceae). The suborder Teloschistineae was formally proposed by Ester Gaya and François Lutzoni in 2016.


Molecular phylogenetics

Historically, classification of taxa within the family relied on physical characteristics such as growth form, the nature of the outer layer of the lichen (the ), and spore type. Studies using modern
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 ...
have shown that
phenotypic In genetics, the phenotype () is the set of observable characteristics or traits of an organism. The term covers the organism's morphology (physical form and structure), its developmental processes, its biochemical and physiological propert ...
characteristics () are not always reliable markers of phylogenetic relationships, and classifying species based on these characters has occasionally led to inaccurate interpretations of their evolutionary history. Advanced DNA analysis techniques have allowed scientists to identify and differentiate
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 ...
, which, though visually indistinguishable, are genetically distinct. This approach has unveiled distinct species within previously thought homogeneous groups, like the genus ''
Caloplaca ''Caloplaca'' is a lichen genus comprising a number of distinct species. Members of the genus are commonly called firedot lichen, jewel lichen.Field Guide to California Lichens, Stephen Sharnoff, Yale University Press, 2014, gold lichens, "ora ...
'', by uncovering their unique genetic markers. Although Teloschistaceae is now well represented in
GenBank The GenBank sequence database is an open access, annotated collection of all publicly available nucleotide sequences and their protein translations. It is produced and maintained by the National Center for Biotechnology Information (NCBI; a par ...
, with thousands of
DNA sequences A nucleic acid sequence is a succession of bases within the nucleotides forming alleles within a DNA (using GACT) or RNA (GACU) molecule. This succession is denoted by a series of a set of five different letters that indicate the order of the ...
, the early molecular studies were limited by having too few examples of each species to draw definitive conclusions. With the increasing availability of genetic sequences, researchers began to gain a better understanding of the family's phylogeny. One significant finding from molecular data is that the traditional morphological methods had mistakenly grouped different species together. For example, the genus ''Caloplaca'' was once thought to be descended from a single lineage (i.e.
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 ...
), but is now understood to have been composed of multiple, unrelated groups (polyphyletic). This insight has prompted numerous proposals to redefine the genus into smaller, monophyletic groups; but such taxonomic changes have sometimes met with resistance due to the vast number of species reclassifications they would entail. According to the lichenologist
Robert Lücking Robert Lücking (born 1964) is a German lichenologist, known for his extensive research on foliicolous lichens (lichens that live on leaves) and his significant contributions to the taxonomy, ecology, and biodiversity of fungi and lichens. He e ...
, families like Teloschistaceae, which have undergone several changes in genus classification through various studies, require phylogenetic consolidation through extensive multi-locus analysis, incorporating all available data and employing rigorous analytical methods. This strategy, akin to approaches taken with families such as
Collemataceae The Collemataceae are a family of mostly lichen-forming fungi in the order Peltigerales. The family contains twelve genera and about 325 species. The family has a widespread distribution. Taxonomy The family was circumscribed by Jonathan Carl Z ...
,
Graphidaceae The Graphidaceae are a family of lichen-forming fungi in the order Graphidales. The family contains nearly a hundred genera and more than 2000 species. Although the family has a cosmopolitan distribution, most Graphidaceae species occur in tropic ...
,
Pannariaceae The Pannariaceae are a family of lichens in the order Peltigerales (suborder Collematineae). Species from this family have a widespread distribution, but are especially prevalent in southern temperate In geography, the temperate clima ...
, and
Parmeliaceae The Parmeliaceae is a large and diverse family of Lecanoromycetes. With over 2700 species in 71 genera, it is the largest family of lichen-forming fungi. The most speciose genera in the family are the well-known groups: '' Xanthoparmelia'' ( 82 ...
, is essential for accurately revising the taxonomic classification of this diverse and widespread group of lichens. Molecular evidence has also helped to map the family's relationships within the class Lecanoromycetes. A 2018 study identified the Megalosporaceae as the Teloschistaceae's closest relative.


Description

In general, Teloschistaceae members are known for their vibrant colours, spanning a spectrum of yellow, orange, and red hues, attributed to
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 ...
pigments. This group of lichens has a broad range of physical formsfrom thin, encrusting (
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 ...
) to leaf-like (
foliose A foliose lichen is a lichen with flat, leaf-like , which are generally not firmly bonded to the substrate on which it grows. It is one of the three most common lichen growth forms, growth forms of lichens. It typically has distinct upper and lo ...
) or even bushy (
fruticose A fruticose lichen is a form of lichen fungi that is characterized by a coral-like shrubby or bushy lichen growth forms, growth structure. It is formed from a symbiotic relationship of a photobiont such as green algae or less commonly cyanobacteri ...
) formations. Although it is an atypical growth form for the Teloschistaceae, members of genus '' Ioplaca'' are somewhat , meaning they have a somewhat circular, leafy thallus attached to the
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 ...
at a single point. Teloschistaceae lichens have a symbiotic relationship with a , generally a member of 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 ( ...
l 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 ...
''. The lichen's reproductive structures, or
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 ...
, are usually brightly coloured, and typically in the form of an apotheciuma wide, open, saucer-shaped or cup-shaped fruit body. In most species, these ascomata have a form, in which the apothecial is surrounded by a pale rim of tissue known as a . Fewer Teloschistaceae species have or forms, in which the apothecial disc lacks a thalline margin. Reproductive
propagule 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 ...
s, such as
isidia An isidium (plural: isidia) is a tiny, wart- or finger-like outgrowth on the thallus surface of certain lichen species. It is one of two principal types of vegetative reproduction, vegetative reproductive structures in lichens, the other being ...
and
soredia Soredia are common reproduction, reproductive structures of lichens. Lichens asexual reproduction, reproduce asexually by employing simple fragmentation and production of soredia and isidia. Soredia are powdery propagules composed of fungus, fung ...
, can be found in select species. The presence and distribution of crystalline deposits serve as important diagnostic features in Teloschistaceae taxonomy. While members of this family commonly produce orange anthraquinone crystals that are soluble in potassium hydroxide, some species also accumulate distinct
calcium oxalate Calcium oxalate (in archaic terminology, oxalate of lime) is a calcium salt of oxalic acid with the chemical formula or . It forms hydrates , where ''n'' varies from 1 to 3. Anhydrous and all hydrated forms are colorless or white. The monohydr ...
crystals, particularly in their
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 ...
. These oxalate crystals are especially prevalent in genera such as '' Wetmoreana'', where they can form either as a distinct layer between the algal and medullary layers or throughout the entire medulla. The pattern and abundance of these crystals can help distinguish between similar species. For example, consistent differences in crystal distribution help separate ''
Wetmoreana ochraceofulva ''Wetmoreana'' is a genus of lichen-forming fungi in the family Teloschistaceae. It comprises 15 formally described species, one subspecies, and three undescribed species of crustose or squamulose that are predominantly saxicolous (rock-dwelling) ...
'' from '' W. brouardii'', and '' Variospora flavescens'' from '' V. aurantia''. In some species, such as ''
Wetmoreana texana ''Wetmoreana'' is a genus of lichen-forming fungi in the family Teloschistaceae. It comprises 15 formally described species, one subspecies, and three undescribed species of crustose or squamulose that are predominantly saxicolous (rock-dwelling) ...
'', the abundance of medullary crystals varies with environmental conditions, being more prevalent in specimens exposed to strong sunlight. While surface crystals () may be influenced by environmental factors, the presence of internal crystalline deposits often represents a stable taxonomic characteristic. The ascomata encase , cylindrical formations that commonly contain between four and sixteen , with eight being the most prevalent count. These asci are characterised by a well-developed J+ layer
amyloid Amyloids are aggregates of proteins characterised by a fibrillar morphology of typically 7–13 nm in diameter, a β-sheet secondary structure (known as cross-β) and ability to be stained by particular dyes, such as Congo red. In the human ...
cap; the term "J+" refers to the positive
staining Staining is a technique used to enhance contrast in samples, generally at the Microscope, microscopic level. Stains and dyes are frequently used in histology (microscopic study of biological tissue (biology), tissues), in cytology (microscopic ...
reaction of the ascus tip 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 ...
, specifically when it turns blue or dark blue in the presence of iodine-based solutions like
Melzer's reagent Melzer's reagent (also known as Melzer's iodine reagent, Melzer's solution or informally as Melzer's) is a chemical reagent used by mycologists to assist with the identification of fungi, and by phytopathologists for fungi that are plant pathogens ...
or
Lugol's iodine Lugol's iodine, also known as aqueous iodine and strong iodine solution, is a solution of potassium iodide with iodine in water. It is a medication and disinfectant used for a number of purposes. Taken by mouth it is used to treat thyrotoxic ...
. The internal structure of the ascus is rudimentary, relative to the more complex apical structures in other related families. Teloschistaceae ascospores, typically
translucent In the field of optics, transparency (also called pellucidity or diaphaneity) is the physical property of allowing light to pass through the material without appreciable light scattering by particles, scattering of light. On a macroscopic scale ...
, usually have one to three septa (internal partitions) with a prominent central septum connected by a canal to the spores' internal cavities, or . Although the presence of a two-chambered (polarilocular) structure in these ascospores is generally indicative of the Teloschistaceae, the spores lack other distinctive features that could be useful as defining taxonomic characteristics. Historically, polarilocular ascospores were regarded as a hallmark of the Teloschistaceae. The incorporation of genera such as ''Apatoplaca'', ''Cephalophysis'', ''Fulgensia'', and ''Xanthopeltis'', which have either non-septate or simply septate spores, has necessitated a reassessment of what fundamentally characterises this group. A distinctive feature of Teloschistaceae is the presence of the gelatinous
paraphyses Paraphyses are erect sterile filament-like support structures occurring among the reproductive apparatuses of fungi, ferns, bryophytes and some thallophytes. The singular form of the word is paraphysis. In certain fungi, they are part of the f ...
(filament-like support structures in the reproductive apparatus), with either unbranched or slightly branched structures culminating in bulbous ends. Within this family,
asexual reproduction Asexual reproduction is a type of reproduction that does not involve the fusion of gametes or change in the number of chromosomes. The offspring that arise by asexual reproduction from either unicellular or multicellular organisms inherit the f ...
leads to the creation of -type
conidiomata Conidiomata (singular: Conidioma) are blister-like fruiting structures produced by a specific type of fungus called a coelomycete. They are formed as a means of dispersing asexual spores call conidia, which they accomplish by creating the blister- ...
(i.e., small, flask-shaped fruiting bodies), producing translucent asexual spores () that are either (rod-shaped) or (double-spindle shaped). The tissue composition of the thallus and apothecia is characterised by a loosely structure, meaning the constituent fungal hyphae are oriented in different directions.


Photobionts

In lichens, photobionts are the
photosynthetic Photosynthesis ( ) is a Biological system, system of biological processes by which Photoautotrophism, photosynthetic organisms, such as most plants, algae, and cyanobacteria, convert light energy, typically from sunlight, into the chemical ener ...
organisms that collaborate with fungal partners to enable the unique lichen symbiosis. Members of the Teloschistaceae associate with green algal photobionts (i.e. resembling or belonging to the green algal 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 ...
''). An early study investigating the
ultrastructure Ultrastructure (or ultra-structure) is the architecture of cells and biomaterials that is visible at higher magnifications than found on a standard optical light microscope. This traditionally meant the resolution and magnification range of a c ...
of the interaction between the fungus and alga in several Teloschistaceae species showed that, in most cases, the cells were merely in close proximity to one another, with only a few instances of fungal cells invading the algal cells. The widespread ''Xanthoria parietina''
species complex 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 ...
has been identified to associate with various trebouxioid species, including '' Asterochloris italiana'', '' Trebouxia arboricola'', and '' T. decolorans''. Within the order Teloschistales, unlike the Teloschistaceae, species in the families Letrouitiaceae and Megalosporaceae primarily partner with the green algal genus ''
Dictyochloropsis ''Dictyochloropsis'' is a genus of Unicellular organism, unicellular Green algae, green alga of the phylum Chlorophyta. This genus consists of Free-swimming, free-living algae which have a reticulate (net-like) chloroplast that varies slightly in ...
''. Due to their resilience to
desiccation Desiccation is the state of extreme dryness, or the process of extreme drying. A desiccant is a hygroscopic (attracts and holds water) substance that induces or sustains such a state in its local vicinity in a moderately sealed container. The ...
, ''Trebouxia'' species serve as the main photobionts for lichen-forming fungi found in extreme environments such as the Antarctic, Arctic, alpine regions, and deserts, where lichens face continual exposure to intense dryness and temperature shifts. Studies of photobionts in the Teloschistaceae, including foliose genera (''Xanthoria'', ''Xanthomendoza'') and a fruticose genus (''Teloschistes''), demonstrate a consistent association with specific ''Trebouxia'' clades. This finding suggests a genus-level specificity, with only select ''Trebouxia'' subclades forming symbiotic relationships. This specificity is not absolute and may vary with habitat: lichens in extreme climates have been observed to associate with a broader range of photobionts.


Chemistry

The predominant chemical compounds found in the Teloschistaceae are pigments known as
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 ...
s. These substances, which are deposited in the upper of the lichen, have
photoprotective Photoprotection is the biochemical process that helps organisms cope with molecular damage caused by sunlight. Plants and other oxygenic phototrophs have developed a suite of photoprotective mechanisms to prevent photoinhibition and oxidative stre ...
properties, as they can absorb
ultraviolet Ultraviolet radiation, also known as simply UV, is electromagnetic radiation of wavelengths of 10–400 nanometers, shorter than that of visible light, but longer than X-rays. UV radiation is present in sunlight and constitutes about 10% of ...
(UV) and blue light. The development 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, particularly anthraquinones, has been essential in the evolutionary adaptation of the family to diverse environments, enabling their transition from shaded, plant-based habitats to sun-exposed,
arid Aridity is the condition of geographical regions which make up approximately 43% of total global available land area, characterized by low annual precipitation, increased temperatures, and limited water availability.Perez-Aguilar, L. Y., Plata ...
regions and contributing significantly to the success of the familial lineage. A 2023 study used
comparative genomics Comparative genomics is a branch of biological research that examines genome sequences across a spectrum of species, spanning from humans and mice to a diverse array of organisms from bacteria to chimpanzees. This large-scale holistic approach c ...
to identify a metabolic gene cluster involved in anthraquinone metabolism and shared uniquely across the Teloschistales. Phylogenetic analyses of fungal
polyketide synthase Polyketide synthases (PKSs) are a family of multi- domain enzymes or enzyme complexes that produce polyketides, a large class of secondary metabolites, in bacteria, fungi, plants, and a few animal lineages. The biosyntheses of polyketides share ...
s (PKSs) reveal a consistent grouping of genes, hinting at a shared ancestral trait for anthraquinone biosynthesis within the subphylum
Pezizomycotina Pezizomycotina is the largest subdivision of Ascomycota, containing the filamentous ascomycetes and most lichenized fungi. It is more or less synonymous with the older taxon Euascomycota. These fungi reproduce by fission rather than budding. Thi ...
. Although the genetic machinery (like the PKSs) involved in anthraquinone biosynthesis in Teloschistales and some non-lichenised fungi is conserved and shows similarities, the specific arrangement of the involved enzymes seems to be a distinguishing feature in the Teloschistales' approach to anthraquinone biosynthesis. The identification of an ABC transporter gene in the pigment gene cluster suggests a mechanism for how lichens accumulate substantial amounts of potentially toxic anthraquinone crystals in their thallus and reproductive structures. Between 1897 and 1906, the mycologist
Friedrich Wilhelm Zopf Friedrich (or Friederich) Wilhelm Zopf (12 December 1846 – 24 June 1909) was a well-known German botanist and mycologist. He dedicated to his whole life with fungal biology, particularly in classification of fungi and dye production in fungi an ...
and the chemist Oswald Hesse conducted a series of early chemical studies on members of the Teloschistaceae, leading to the extraction of the reddish pigment
parietin Parietin is the predominant cortical pigment of lichens in the genus '' Caloplaca'', a secondary product of the lichen '' Xanthoria parietina'', and a pigment found in the roots of curled dock (''Rumex crispus''). It has an orange-yellow color ...
from selected species. Parietin is an
antioxidant Antioxidants are Chemical compound, compounds that inhibit Redox, oxidation, a chemical reaction that can produce Radical (chemistry), free radicals. Autoxidation leads to degradation of organic compounds, including living matter. Antioxidants ...
molecule that is produced in greater amounts in lichen thalli that are exposed to excess nitrogen. In a 1970 publication, Johan Santesson surveyed 230 ''Caloplaca'' species for anthraquinones as part of a
phytochemical Phytochemicals are naturally-occurring chemicals present in or extracted from plants. Some phytochemicals are nutrients for the plant, while others are metabolites produced to enhance plant survivability and reproduction. The fields of ext ...
study of the Teloschistaceae, and concluded that the studied species could be arranged according to their anthraquinone content in thirteen chemical groups. In 1997, Ulrik Søchting analysed secondary metabolites from species of ''Caloplaca'', ''Teloschistes'', and ''Xanthoria'' to look for chemical patterns of consistent combinations and proportions of lichen products. He identified two (characteristic sets of chemical compounds) with parietin,
emodin Emodin (6-methyl-1,3,8-trihydroxyanthraquinone) is an organic compound. Classified as an anthraquinone, it can be isolated from rhubarb, buckthorn, and Japanese knotweed ('' Reynoutria japonica'' syn. ''Polygonum cuspidatum''). Emodin is part ...
, fallacinol, fallacinal, and parietinic acid as the main substances. Parietin acts as a UV-light filter to provide optimal light intensities for the photobionts that are resident in the internal . Studies show that in ''Xanthoria parietina'', the more light the lichen is exposed to, the higher the concentration of parietin. In the Teloschistaceae, parietin may also serve a defensive role. In the
Negev The Negev ( ; ) or Naqab (), is a desert and semidesert region of southern Israel. The region's largest city and administrative capital is Beersheba (pop. ), in the north. At its southern end is the Gulf of Aqaba and the resort town, resort city ...
desert, the parietin-containing Teloschistaceae species '' Elenkiniana ehrenbergii'' and '' Seirophora lacunosa'' are avoided by
grazing In agriculture, grazing is a method of animal husbandry whereby domestic livestock are allowed outdoors to free range (roam around) and consume wild vegetations in order to feed conversion ratio, convert the otherwise indigestible (by human diges ...
snails, but they frequently consume lichens like '' Diploicia canescens'' and '' Buellia subalbula'' (both in family
Caliciaceae The Caliciaceae are a family (taxonomy), family of mostly lichen-forming fungus, fungi belonging to the class (taxonomy), class Lecanoromycetes in the division (mycology), division Ascomycota. Although the family has had its classification chan ...
), which lack parietin. In their large-scale phylogenetic analysis of the Teloschistaceae, the lichenologists Ulf Arup, Ulrik Søchting, and Patrik Frödén analysed about 4000 specimens of members of the family using
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 ...
, and identified more than 100 secondary metabolites, mostly anthraquinones. They noted that in the large majority of cases, the distribution of lichen products was more or less constant within species. In some instances, the secondary chemistry is important at higher taxonomic levels (i.e.,
rank A rank is a position in a hierarchy. It can be formally recognized—for example, cardinal, chief executive officer, general, professor—or unofficial. People Formal ranks * Academic rank * Corporate title * Diplomatic rank * Hierarchy ...
s higher than species). For example, the genus ''Catenaria'' is characterised by the presence of 7-chlorocatenarin, a secondary metabolite previously unknown in lichens. The substance
usnic acid Usnic acid is a naturally occurring dibenzofuran derivative found in several lichen species with the formula C18H16O7. It was first isolated by German scientist W. Knop in 1844 and first synthesized between 1933 and 1937 by Frank H. Curd and Al ...
characterises the genus '' Usnochroma'', and 5-chloroemodin occurs in all but one species of ''
Shackletonia ''Shackletonia'' is a genus of lichen-forming fungi in the family Teloschistaceae. The species in the ''Shackletonia'' genus are lichenicolous and muscicolous, and known from Antarctica and southern Patagonia (Arup et al. 2013). The genus was ...
''. The secondary chemistry of the Caloplacoideae is the most diverse amongst the three Teloschistaceae subfamilies, as it contains both
chlorinated In chemistry, halogenation is a chemical reaction which introduces one or more halogens into a chemical compound. Halide-containing compounds are pervasive, making this type of transformation important, e.g. in the production of polymers, drugs. ...
anthraquinones and
depsidone Depsidones (+ " depside" + "one") are chemical compounds that are sometimes found as secondary metabolites in lichens. They are esters that are both depsides and cyclic ethers. An example is norstictic acid Norstictic acid is a depsidone produ ...
s. Although most Teloschistaceae lichens produce anthraquinone pigments in shades ranging from yellow to orange to red, the genera '' Apatoplaca'' and '' Cephalophysis'' lack these anthraquinones. Similarly, the genus '' Pyrenodesmia'' encompasses species where anthraquinones are absent and replaced by substances such as or ; these insoluble lichen pigments may confer UV-protective ability similar to anthraquinones. Taxa of the closely related genera '' Kuettlingeria'' and ''
Sanguineodiscus ''Sanguineodiscus'' is a genus of lichen-forming fungi in the family Teloschistaceae. It is characterized by its deep red apothecial , found predominantly in the Mediterranean basin, Central Asia, Northern Africa, and Europe. This genus comprise ...
'' have anthraquinones in their apothecia and Sedifolia-grey in their thalli. The species '' Kuettlingeria neotaurica'' features apothecia of two colour variants: orange-red (with anthraquinones) and grey (with Sedifolia-grey). The absence of anthraquinones is not a
synapomorphic In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to hav ...
character, but appears independently in unrelated lineages of Teloschistaceae; as such, it is a phylogenetically unreliable .


Adaptive radiation

Adaptive radiation In evolutionary biology, adaptive radiation is a process in which organisms diversify rapidly from an ancestral species into a multitude of new forms, particularly when a change in the environment makes new resources available, alters biotic int ...
in the Teloschistaceae has been studied to understand the key
phenotypic In genetics, the phenotype () is the set of observable characteristics or traits of an organism. The term covers the organism's morphology (physical form and structure), its developmental processes, its biochemical and physiological propert ...
changes leading to their diversification. This diversification is believed to be connected to the spread of anthraquinone pigments in their thallus. Initially, these pigments were thought to have appeared during the Teloschistaceae's first
divergence In vector calculus, divergence is a vector operator that operates on a vector field, producing a scalar field giving the rate that the vector field alters the volume in an infinitesimal neighborhood of each point. (In 2D this "volume" refers to ...
, with a more widespread occurrence developing later. The distribution of anthraquinones in Teloschistaceae lichens varies, from being dispersed across the organism's surface to localised regions. Analysis suggests that the family's lineage witnessed a loss and subsequent return of these pigments over time, considering their presence in the thallus and apothecia as the
ancestral state In phylogenetics, a primitive (or ancestral) character, trait, or feature of a lineage or taxon is one that is inherited from the common ancestor of a clade (or clade group) and has undergone little change since. Conversely, a trait that appears ...
. Ecologically, these organisms transitioned from inhabiting shaded bark environments to colonising sunlit, rocky areas during their diversification. The analysis of phenotypic traits and diversification rates shows that anthraquinones in the thallus and greater sun exposure have contributed to an acceleration of diversification. On the contrary, living in shaded environments or having a crustose-continuous (smooth, non-scaly) growth form hindered diversification. The choice of substrate, be it rock or bark, did not have a pronounced impact on diversification rates. This adaptive radiation within the Teloschistaceae is estimated to have initiated around 100 million years ago, specifically during the
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the more recent of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''cre ...
period. Factors like climatic shifts, continental separations, and the emergence of
flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (). The term angiosperm is derived from the Ancient Greek, Greek words (; 'container, vessel') and (; 'seed'), meaning that the seeds are enclosed with ...
s are theorised to have influenced the adaptive landscape. Such factors might have promoted the development of light-protective anthraquinones, enabling Teloschistaceae to colonise exposed environments. The diversification of anthraquinone
gene In biology, the word gene has two meanings. The Mendelian gene is a basic unit of heredity. The molecular gene is a sequence of nucleotides in DNA that is transcribed to produce a functional RNA. There are two types of molecular genes: protei ...
s in their evolution is primarily due to gene reshuffling, which has given rise to novel biosynthetic
enzyme An enzyme () is a protein that acts as a biological catalyst by accelerating chemical reactions. The molecules upon which enzymes may act are called substrate (chemistry), substrates, and the enzyme converts the substrates into different mol ...
pathways and
gene cluster A gene cluster is a group of two or more genes found within an organism's DNA that encode similar peptide, polypeptides or proteins which collectively share a generalized function and are often located within a few thousand base pairs of each othe ...
s.


Genera

The classification and number of species within the Teloschistaceae have evolved significantly over time. Historical estimates have varied, with 10 genera and 47 species recognised in 2001, increasing to 12 genera and 644 species by 2008. The new classification proposed by Arup and colleagues in their 2013 taxonomy article recognised 39 genera, including 31 that were newly described or resurrected. By 2016, there were between 51 and 53 genera and about 700 species; 65 genera and 755 species in 2017; and 71 genera and about 840 species by 2022. In the same year, Kondratyuk and colleagues enumerated all members of the Teloschistaceae with publicly available DNA sequences, and confirmed 590 species across 115 genera. ,
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 ...
(in the
Catalogue of Life The Catalogue of Life (CoL) is an online database that provides an index of known species of animals, plants, fungi, and microorganisms. It was created in 2001 as a partnership between the global Species 2000 and the American Integrated Taxono ...
), accepts 117 genera and 805 species in the Teloschistaceae. The largest genus is ''Caloplaca'', at 173 accepted species. In terms of
species diversity Species diversity is the number of different species that are represented in a given community (a dataset). The effective number of species refers to the number of equally abundant species needed to obtain the same mean proportional species abundan ...
, Teloschistaceae stood as the sixth-largest lichen-forming fungal family by 2017, following
Parmeliaceae The Parmeliaceae is a large and diverse family of Lecanoromycetes. With over 2700 species in 71 genera, it is the largest family of lichen-forming fungi. The most speciose genera in the family are the well-known groups: '' Xanthoparmelia'' ( 82 ...
,
Graphidaceae The Graphidaceae are a family of lichen-forming fungi in the order Graphidales. The family contains nearly a hundred genera and more than 2000 species. Although the family has a cosmopolitan distribution, most Graphidaceae species occur in tropic ...
,
Verrucariaceae Verrucariaceae is a family of lichens and a few non-lichenised fungi in the order Verrucariales. The lichens have a wide variety of thallus forms, from crustose (crust-like) to foliose (bushy) and squamulose (scaly). Most of them grow on lan ...
,
Ramalinaceae The Ramalinaceae are a family of lichen-forming fungi in the order Lecanorales. First proposed by Carl Adolph Agardh in 1821, the family now comprises 63 genera and about 750 species. Ramalinaceae lichens exhibit diverse growth forms, includin ...
, and
Lecanoraceae The Lecanoraceae are a family 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 c ...
. Each genus is paired with its
taxonomic authority In biology, taxonomy () is the scientific study of naming, defining ( circumscribing) and classifying groups of biological organisms based on shared characteristics. Organisms are grouped into taxa (singular: taxon), and these groups are given ...
, denoting the first describers using standardised author abbreviations, the publication year, and the number of species. Genera are organised by subfamily:


Caloplacoideae

*'' Apatoplaca'' – 1 sp. *'' Blastenia'' – 11 spp. *'' Bryoplaca'' – 3 spp. *''
Caloplaca ''Caloplaca'' is a lichen genus comprising a number of distinct species. Members of the genus are commonly called firedot lichen, jewel lichen.Field Guide to California Lichens, Stephen Sharnoff, Yale University Press, 2014, gold lichens, "ora ...
'' 351 spp. *'' Cephalophysis'' – 1 sp. *'' Eilifdahlia'' – 2 spp. *''
Elenkiniana ''Elenkiniana'' is a genus of lichen-forming fungi in the family Teloschistaceae. It has three species, all of which occur in Eurasia. Taxonomy The genus was circumscribed in 2014 by the lichenologists Sergey Kondratyuk, Ingvar Kärnefelt, Joh ...
'' – 3 spp. *'' Fauriea'' – 7 spp. *'' Franwilsia'' – 3 spp. *''
Fulgensia ''Fulgensia'' is a fungal genus in the family Teloschistaceae. The genus was circumscribed by Abramo Bartolommeo Massalongo and Giuseppe De Notaris in 1853, with ''Fulgensia vulgaris'' assigned as the type species. Although the genus has had se ...
'' – 1 sp. *'' Gintarasiella'' – 1 sp. *'' Gyalolechia'' – 40 spp. *'' Hanstrassia'' – 2 spp. *'' Hueidea'' – 6 spp. *'' Huneckia'' – 4 spp. *'' Ioplaca'' – 2 spp. *'' Jasonhuria'' – 1 sp. *'' Klauderuiella'' – 3 spp. *'' Kuettlingeria'' – 15 spp. *'' Lacrima'' – 4 spp. *'' Laundonia'' – 2 spp. *'' Lendemeriella'' – 9 spp. *''
Leproplaca ''Leproplaca'' is a genus of lichen-forming fungi in the family Teloschistaceae. Its exhibits a leprose growth form, where the thallus consists primarily of asexual propagules called soredia Soredia are common reproduction, reproductive struct ...
'' – 7 spp. *''
Loekoesia ''Loekoesia'' is a genus of lichen-forming fungi in the family Teloschistaceae. It contains three species of saxicolous (rock-dwelling), crustose lichens. Collectively, the genus occurs in South Korea, Mauritius, and the United States. The genu ...
'' – 3 spp. *'' Marchantiana'' – 7 spp. *'' Mikhtomia'' – 4 spp. *'' Neoplaca'' – 1 sp. *'' Obscuroplaca'' – 3 spp. *'' Oceanoplaca'' – 6 spp. *'' Olegblumia'' – 1 sp. *''
Opeltia ''Opeltia'' is a genus of lichen-forming fungi in the family Teloschistaceae. It has four species of corticolous (bark-dwelling), crustose lichens. Taxonomy The genus was circumscribed in 2017 by lichenologists Sergey Kondratyuk and Laszlo L ...
'' – 4 spp. *''
Oxneriopsis ''Oxneriopsis'' is a genus of lichen-forming fungi in the family Teloschistaceae. It has four species of corticolous (bark-dwelling), crustose lichens. Taxonomy The genus was circumscribed in 2017 by the lichenologists Sergey Kondratyuk, Dalip ...
'' – 4 spp. *'' Pisutiella'' – 6 spp. *'' Pyrenodesmia'' – 6 spp. *'' Rufoplaca'' – 10 spp. *''
Sanguineodiscus ''Sanguineodiscus'' is a genus of lichen-forming fungi in the family Teloschistaceae. It is characterized by its deep red apothecial , found predominantly in the Mediterranean basin, Central Asia, Northern Africa, and Europe. This genus comprise ...
'' – 4 spp. *''
Seirophora ''Seirophora'' is a genus of lichen-forming fungi in the family Teloschistaceae. It has eight species. The genus was circumscribed by German lichenologist Josef Poelt in 1983, with '' Seirophora magara'' assigned as the type species. Several s ...
'' – 8 spp. *'' Sucioplaca'' – 1 sp. *'' Taedigera'' – 6 spp. *'' Upretia'' – 2 spp. *'' Usnochroma'' – 2 spp. *''
Variospora ''Variospora'' is a genus of crustose lichens, belonging to the family Teloschistaceae The Teloschistaceae are a large family (biology), family of mostly lichen-forming fungi belonging to the class (taxonomy), class Lecanoromycetes in the d ...
'' – 16 spp. *'' Xanthaptychia'' – 3 spp. *''
Yoshimuria ''Yoshimuria'' is a genus of lichen-forming fungi in the family Teloschistaceae. It has four species of crustose lichens. Taxonomy The genus was circumscribed in 2014 by a group of lichenologists, who assigned '' Yoshimuria spodoplaca'' as th ...
'' – 4 spp.


Teloschistoideae

*'' Aridoplaca'' – 1 sp. *'' Brownliella'' – 2 spp. *'' Catenarina'' – 3 spp. *'' Cinnabaria'' – 1 sp. *''
Elixjohnia ''Elixjohnia'' is a genus of lichen-forming fungi in the family Teloschistaceae. It has four species of saxicolous (rock-dwelling), crustose lichens that occur in Australasia. Taxonomy ''Elixjohnia'' was circumscribed in 2017 by Sergey Kondraty ...
'' – 4 spp. *'' Filsoniana'' – 9 spp. *'' Follmannia'' – 2 spp. *''
Haloplaca ''Haloplaca'' is a genus of lichen-forming fungi in the subfamily Teloschistaceae of the family Teloschistaceae. It contains three species of crustose lichens. The genus was circumscribed by Ulf Arup and colleagues in 2013, with ''Haloplaca bri ...
'' – 3 spp. *'' Harusavskia'' – 1 sp. *'' Hosseusiella'' – 3 spp. *''
Ikaeria ''Ikaeria'' is a genus of two species of crustose lichens in the family Teloschistaceae. Both species grow on twig bark of shrubs and trees. It was circumscribed in 2017 by lichenologists Sergey Kondratyuk, Dalip Kumar Upreti, and Jae-Seoun Hur, ...
'' – 2 spp. *'' Iqbalia'' – 1 sp. *'' Josefpoeltia'' – 3 spp. *''
Kaernefia ''Kaernefia'' is a genus of crustose lichens in the family Teloschistaceae. It has three species, found in Australia or South Africa. Taxonomy ''Kaernefia'' was circumscribed in 2013 by Sergey Kondratyuk, John Elix, Arne Thell, and Jae-Seoun Hu ...
'' – 3 spp. *'' Lazarenkoiopsis'' – 1 sp. *'' Loekoeslaszloa'' – 2 spp. *'' Neobrownliella'' – 5 spp. *'' Nevilleiella'' – 2 spp. *'' Niorma'' – 5 spp. *'' Raesaeneniana'' – 1 sp. *'' Rehmanniella'' – 5 spp. *'' Scutaria'' – 1 sp. *''
Sirenophila ''Sirenophila'' is a genus of crustose lichens in the subfamily Teloschistoideae of the family Teloschistaceae. It has five species with an Australasian distribution. Taxonomy The genus was circumscribed in 2013 by Ulrik Søchting, Ulf Arup, and ...
'' – 4 spp. *'' Stellarangia'' – 3 spp. *'' Streimanniella'' – 4 spp. *'' Tarasginia'' – 2 spp. *'' Tassiloa'' – 2 spp. *'' Tayloriellina'' – 2 spp. *'' Teloschistes'' – ca. 24 spp. *'' Teloschistopsis'' – 3 spp. *'' Thelliana'' – 1 sp. *''
Villophora ''Villophora'' is a genus of lichen-forming fungi in the subfamily Teloschistoideae of the family Teloschistaceae. Eight species are accepted in the genus. First circumscribed in 2013 and confirmed by a 2021 DNA study, the genus forms a well-s ...
'' – 9 spp. *'' Wetmoreana'' – 2 spp. *'' Wilketalia'' – 1 sp.


Xanthorioideae

*'' Amundsenia'' – 2 spp. *'' Athallia'' – 17 spp. *'' Austroplaca'' – 10 spp. *'' Calogaya'' – 19 spp. *''
Cerothallia ''Cerothallia'' is a genus of crustose lichens in the family Teloschistaceae. It has four species, all of which occur in the Southern Hemisphere. The genus was circumscribed in 2013 by Ulf Arup, Patrik Frödén, and Ulrik Søchting, with '' Cerot ...
'' – 4 spp. *'' Charcotiana'' – 1 sp. *'' Coppinsiella'' – 3 spp. *'' Dijigiella'' – 2 spp. *'' Dufourea'' – 25 spp. *'' Erichansenia'' – 3 spp. *''
Flavoplaca ''Flavoplaca'' is a genus of crust-like or scaly lichens in the family Teloschistaceae. It has about 30 species with a mostly Northern Hemisphere distribution. Taxonomy The genus was circumscribed in 2013 by Ulf Arup, Patrik Frödén and Ulr ...
'' – 28 spp. *'' Fominiella'' – 2 spp. *'' Gallowayella'' – 15 spp. *'' Golubkovia'' – 1 sp. *'' Gondwania'' – 4 spp. *'' Honeggeria'' – 1 sp. *'' Huriella'' – 5 spp. *'' Igneoplaca'' – 1 sp. *''
Jackelixia ''Dufourea'' is a genus of mostly foliose lichen species in the subfamily Xanthorioideae of the family Teloschistaceae. Species in the genus are found in the Southern Hemisphere. Taxonomy The genus was originally circumscription (taxonomy), circ ...
'' *'' Jesmurraya'' – 1 sp. *'' Langeottia'' – 2 spp. *'' Lazarenkoella'' – 2 spp. *'' Martinjahnsia'' – 1 sp. *'' Massjukiella'' – 8 spp. *'' Orientophila'' – 15 spp. *'' Ovealmbornia'' – 3 spp. *'' Oxneria'' – 4 spp. *'' Pachypeltis'' – 7 spp. *'' Parvoplaca'' – 6 spp. *''
Polycauliona ''Polycauliona'' is a genus of lichen-forming fungi in the family Teloschistaceae. Species *'' Polycauliona antarctica'' *'' Polycauliona ascendens'' *'' Polycauliona austrogeorgica'' *'' Polycauliona bolacina'' *'' Polycauliona brattiae'' ...
'' – 18 spp. *'' Rusavskia'' – 19 spp. *''
Scythioria ''Scythioria'' is a small genus of lichen-forming fungi in the family Teloschistaceae. It comprises three species. Species in this genus are found primarily in the Palaearctic realm, occurring in Eurasia and North Africa. They typically grow on ...
'' – 3 spp. *'' Seawardiella'' – 2 spp. *''
Shackletonia ''Shackletonia'' is a genus of lichen-forming fungi in the family Teloschistaceae. The species in the ''Shackletonia'' genus are lichenicolous and muscicolous, and known from Antarctica and southern Patagonia (Arup et al. 2013). The genus was ...
'' – 5 spp. *'' Solitaria'' – 1 sp. *''
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. ...
'' – 15 spp. *'' Teuvoahtiana'' – 3 spp. *''
Tomnashia ''Tomnashia'' is a genus of lichen-forming fungi in the family Teloschistaceae. It has four species of saxicolous (rock-dwelling), crustose lichens that occur in southwestern North America. Taxonomy ''Tomnashia'' was circumscribed in 2017 by ...
'' – 4 spp. *'' Transdrakea'' – 2 spp. *'' Verrucoplaca'' – 1 sp. *'' Xanthocarpia'' – 12 spp. *'' Xanthodactylon'' – 2 spp. *'' Xanthokarrooa'' – 2 spp. *'' Xanthomendoza'' – 20 spp. *'' Xanthopeltis'' – 1 sp. *'' Xanthoria'' – 10 spp. *'' Zeroviella'' – 8 spp.


Invalid names

Some of the genera proposed during the restructuring of the family have since been shown to be nomenclaturally illegitimate or unavailable for use. For example: *''Andina'' has been replaced with '' Wilketalia''. *''Phaeoplaca'' has been replaced with '' Obscuroplaca''. *''Tayloriella'' has been replaced with '' Tayloriellina''.


Habitat, distribution, and ecology

The family has a worldwide distribution, although members occur predominantly in
temperate In geography, the temperate climates of Earth occur in the middle latitudes (approximately 23.5° to 66.5° N/S of the Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ran ...
regions, and in semi-arid regions of the
subtropics The subtropical zones or subtropics are geographical and climate zones immediately to the north and south of the tropics. Geographically part of the temperate zones of both hemispheres, they cover the middle latitudes from to approximately ...
. Most members either grow on rock (
saxicolous This glossary of mycology is a list of definitions of terms and concepts relevant to mycology, the study of fungi. Terms in common with other fields, if repeated here, generally focus on their mycology-specific meaning. Related terms can be found ...
) or on bark (
corticolous This glossary of mycology is a list of definitions of terms and concepts relevant to mycology, the study of fungi. Terms in common with other fields, if repeated here, generally focus on their mycology-specific meaning. Related terms can be found ...
). As an exception to this general ecological preference, the genus '' Bryoplaca'' contains species that only grow on mosses and
detritus In biology, detritus ( or ) is organic matter made up of the decomposition, decomposing remains of organisms and plants, and also of feces. Detritus usually hosts communities of microorganisms that colonize and decomposition, decompose (Reminera ...
. Some ''
Fulgensia ''Fulgensia'' is a fungal genus in the family Teloschistaceae. The genus was circumscribed by Abramo Bartolommeo Massalongo and Giuseppe De Notaris in 1853, with ''Fulgensia vulgaris'' assigned as the type species. Although the genus has had se ...
'' species grow on soils, particularly those rich in
lime Lime most commonly refers to: * Lime (fruit), a green citrus fruit * Lime (material), inorganic materials containing calcium, usually calcium oxide or calcium hydroxide * Lime (color), a color between yellow and green Lime may also refer to: Bo ...
. Several crustose Teloschistaceae species, typically saxicolous in nature, have been recorded growing on human bone remains recovered at a looted Late
Holocene The Holocene () is the current geologic time scale, geological epoch, beginning approximately 11,700 years ago. It follows the Last Glacial Period, which concluded with the Holocene glacial retreat. The Holocene and the preceding Pleistocene to ...
aboriginal
cairn A cairn is a human-made pile (or stack) of stones raised for a purpose, usually as a marker or as a burial mound. The word ''cairn'' comes from the (plural ). Cairns have been and are used for a broad variety of purposes. In prehistory, t ...
burial site in Argentina. Many species of the family are moderately to strongly nitrophilous, meaning that they prefer habitats rich in
nitrogen Nitrogen is a chemical element; it has Symbol (chemistry), symbol N and atomic number 7. Nitrogen is a Nonmetal (chemistry), nonmetal and the lightest member of pnictogen, group 15 of the periodic table, often called the Pnictogen, pnictogens. ...
, particularly in the form of
nitrate Nitrate is a polyatomic ion with the chemical formula . salt (chemistry), Salts containing this ion are called nitrates. Nitrates are common components of fertilizers and explosives. Almost all inorganic nitrates are solubility, soluble in wa ...
. Adapted to environments with high sunlight exposure, Teloschistaceae lichens show an enhanced ability to fix carbon from the atmosphere, a crucial process for their sustenance and growth. These lichens also have an increased capacity to absorb excess nitrogen, a trait linked to their adaptation to intense light conditions. This dual adaptation – improved carbon fixation and nitrogen absorption – is evident in the family's small foliose and crustose species, which show greater tolerance to elevated nitrogen levels. The absorption of additional nitrogen is thought to be associated with the presence of specific compounds like anthraquinones in these lichens. These compounds not only contribute to their characteristic yellow/orange colouration but may also help protect fungal
cell membrane The cell membrane (also known as the plasma membrane or cytoplasmic membrane, and historically referred to as the plasmalemma) is a biological membrane that separates and protects the interior of a cell from the outside environment (the extr ...
s. ''Xanthoria parietina'' is one example of a widespread lichen that appears to be experiencing an increase in its
range Range may refer to: Geography * Range (geographic), a chain of hills or mountains; a somewhat linear, complex mountainous or hilly area (cordillera, sierra) ** Mountain range, a group of mountains bordered by lowlands * Range, a term used to i ...
due to its ability to tolerate nitrogenous
pollutant A pollutant or novel entity is a substance or energy introduced into the environment that has undesired effect, or adversely affects the usefulness of a resource. These can be both naturally forming (i.e. minerals or extracted compounds like oi ...
s, and its potential ability to displace native lichen species as a result. ''Caloplaca'', ''Fulgensia'', ''Teloschistes'', and ''Xanthoria'' are genera that are characteristic of sun-exposed habitats. In some extreme desert environments, ''Caloplaca'' (in the broad senseincluding lichens historically classified in this genus) may be the only genus present; ''Caloplaca'' and ''Xanthoria'' dominate harsh coastal environments. There are several Teloschistaceae genera that contain
lichenicolous This glossary of mycology is a list of definitions of terms and concepts relevant to mycology, the study of fungi. Terms in common with other fields, if repeated here, generally focus on their mycology-specific meaning. Related terms can be found ...
(lichen-dwelling) species. These non-lichenised fungi originate from subfamily Caloplacoideae: ''
Caloplaca ''Caloplaca'' is a lichen genus comprising a number of distinct species. Members of the genus are commonly called firedot lichen, jewel lichen.Field Guide to California Lichens, Stephen Sharnoff, Yale University Press, 2014, gold lichens, "ora ...
'' (26 spp.), '' Gyalolechia'' (1 sp.), ''
Variospora ''Variospora'' is a genus of crustose lichens, belonging to the family Teloschistaceae The Teloschistaceae are a large family (biology), family of mostly lichen-forming fungi belonging to the class (taxonomy), class Lecanoromycetes in the d ...
'' (1 sp.); from subfamily Teloschistoideae: '' Catenarina'' (1 sp.), ''Sirenophila'' 1; and from subfamily Xanthorioideae: ''
Flavoplaca ''Flavoplaca'' is a genus of crust-like or scaly lichens in the family Teloschistaceae. It has about 30 species with a mostly Northern Hemisphere distribution. Taxonomy The genus was circumscribed in 2013 by Ulf Arup, Patrik Frödén and Ulr ...
'' (4 spp.), '' Pachypeltis'' (1 sp.), and ''
Shackletonia ''Shackletonia'' is a genus of lichen-forming fungi in the family Teloschistaceae. The species in the ''Shackletonia'' genus are lichenicolous and muscicolous, and known from Antarctica and southern Patagonia (Arup et al. 2013). The genus was ...
'' (3 spp.). Lichenicolous species within the Teloschistaceae generally have a broad range of
hosts A host is a person responsible for guests at an event or for providing hospitality during it. Host may also refer to: Places * Host, Pennsylvania, a village in Berks County * Host Island, in the Wilhelm Archipelago, Antarctica People * ...
. Their geographical distribution seems to be influenced not just by the classification of their host lichen, but also by the they grow on. Teloschistaceae has a high diversity in
polar regions The polar regions, also called the frigid geographical zone, zones or polar zones, of Earth are Earth's polar ice caps, the regions of the planet that surround its geographical poles (the North Pole, North and South Poles), lying within the pol ...
and a substantial number of bipolar species, i.e., species occurring in both northern and southern hemispheres but largely absent from intermediate, tropical latitudes. Examples include '' Gallowayella borealis'', '' Austroplaca soropelta'', and ''
Scythioria phlogina ''Scythioria'' is a small genus of lichen-forming fungi in the family Teloschistaceae. It comprises three species. Species in this genus are found primarily in the Palaearctic realm, occurring in Eurasia and North Africa. They typically grow on ...
''. There is a relatively low diversity of crustose Teloschistaceae in
Central Europe Central Europe is a geographical region of Europe between Eastern Europe, Eastern, Southern Europe, Southern, Western Europe, Western and Northern Europe, Northern Europe. Central Europe is known for its cultural diversity; however, countries in ...
. Localised exceptions occur primarily in sunlit locations with either
calcareous Calcareous () is an adjective meaning "mostly or partly composed of calcium carbonate", in other words, containing lime (mineral), lime or being chalky. The term is used in a wide variety of Science, scientific disciplines. In zoology ''Calcare ...
or nutrient-rich
siliceous rock Siliceous rocks are sedimentary rocks that have silica (SiO2) as the principal constituent. The most common siliceous rock is chert; other types include diatomite. They commonly form from silica-secreting organisms such as radiolarians, diatoms, o ...
formations; these habitats are predominant in the
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 P ...
regions of the
Alps The Alps () are some of the highest and most extensive mountain ranges in Europe, stretching approximately across eight Alpine countries (from west to east): Monaco, France, Switzerland, Italy, Liechtenstein, Germany, Austria and Slovenia. ...
and the
Carpathian Mountains The Carpathian Mountains or Carpathians () are a range of mountains forming an arc across Central Europe and Southeast Europe. Roughly long, it is the third-longest European mountain range after the Ural Mountains, Urals at and the Scandinav ...
, as well as in the arid, warm rocky
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 ...
s. Some Teloschistaceae genera have a strong geographic centre of
species richness Species richness is the number of different species represented in an community (ecology), ecological community, landscape or region. Species richness is simply a count of species, and it does not take into account the Abundance (ecology), abunda ...
; examples include ''
Elixjohnia ''Elixjohnia'' is a genus of lichen-forming fungi in the family Teloschistaceae. It has four species of saxicolous (rock-dwelling), crustose lichens that occur in Australasia. Taxonomy ''Elixjohnia'' was circumscribed in 2017 by Sergey Kondraty ...
'' (Australasia), '' Orientophila'' (east Asia), ''
Shackletonia ''Shackletonia'' is a genus of lichen-forming fungi in the family Teloschistaceae. The species in the ''Shackletonia'' genus are lichenicolous and muscicolous, and known from Antarctica and southern Patagonia (Arup et al. 2013). The genus was ...
'' (Antarctic and subantarctic), '' Stellarangia'' (south-western Africa), and '' Xanthoria'' (Mediterranean area). Several studies published since 2014 have enumerated the Teloschistaceae taxa occurring in certain defined geographical areas. These include: *
Altai-Sayan region The Altai-Sayan region is an area of Inner Asia proximate to the Altai Mountains and the Sayan Mountains, near to where Russia, China, Mongolia and Kazakhstan come together. This region is one of the world centers of Temperate climate, temperate pl ...
, 103 species in 31 genera *
Crimea Crimea ( ) is a peninsula in Eastern Europe, on the northern coast of the Black Sea, almost entirely surrounded by the Black Sea and the smaller Sea of Azov. The Isthmus of Perekop connects the peninsula to Kherson Oblast in mainland Ukrain ...
, 119 species *
Dagestan Dagestan ( ; ; ), officially the Republic of Dagestan, is a republic of Russia situated in the North Caucasus of Eastern Europe, along the Caspian Sea. It is located north of the Greater Caucasus, and is a part of the North Caucasian Fede ...
, 85 species *India, 115 species in 36 genera *Italy, about 160 species *
Galápagos Islands The Galápagos Islands () are an archipelago of volcanic islands in the Eastern Pacific, located around the equator, west of the mainland of South America. They form the Galápagos Province of the Republic of Ecuador, with a population of sli ...
, 24 species *Mexico, 142 species in 6 genera *New Zealand, about 100 species *
Russian Far East The Russian Far East ( rus, Дальний Восток России, p=ˈdalʲnʲɪj vɐˈstok rɐˈsʲiɪ) is a region in North Asia. It is the easternmost part of Russia and the Asia, Asian continent, and is coextensive with the Far Easte ...
, 84 species * Ural, 81 species


Species interactions

Teloschistaceae species host a variety of lichenicolous fungi. Some, such as '' Cercidospora caudata'' and '' Stigmidium cerinae'', infect a wide range of hosts within the family. More generally, these
parasitic Parasitism is a close relationship between species, where one organism, the parasite, lives (at least some of the time) on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The ent ...
fungi have a preference for specific species or genera of Teloschistaceae. An example is the relationship between Teloschistaceae lichens and the fungus '' Tremella caloplacae''. Integrative studies combining molecular data and ecological approaches revealed at least six distinct lineages of ''T. caloplacae'', each specialised to a particular host, indicating a complex of closely related species. This diversification of ''T. caloplacae'' appears to have occurred in tandem with the rapid diversification of the Teloschistaceae since the Late Cretaceous period, implying
coevolution In biology, coevolution occurs when two or more species reciprocally affect each other's evolution through the process of natural selection. The term sometimes is used for two traits in the same species affecting each other's evolution, as well a ...
. Further molecular studies have delineated the ''T. caloplacae'' group into a complex of at least nine distinct species. Out of these, five new species were formally described in 2023, each adapted to a single host species or genus within the Teloschistaceae.


Human interactions and uses

Although Teloschistaceae species are not economically significant, their tendency to grow on rock surfaces has been documented to cause damage to marble structures. In some cases, the lichens, the major contributor of which was '' Xanthocarpia feracissima'', penetrated up to into the stone along larger cracks and beneath loose surface crystals, leading to crumbling of the marble surface. ''Caloplaca pseudopoliotera'' and '' C. cupulifera'' are two crustose species responsible for the slow degradation of the
Konark Sun Temple Konark Sun Temple is a Hindu temple, Hindu Surya, Sun temple at Konark about northeast from Puri, Puri city on the coastline in Puri district, Odisha, India.


Traditional medicine

Some Teloschistaceae species have been used in
traditional medicine Traditional medicine (also known as indigenous medicine or folk medicine) refers to the knowledge, skills, and practices rooted in the cultural beliefs of various societies, especially Indigenous groups, used for maintaining health and treatin ...
practices across several cultures. ''
Xanthoria parietina ''Xanthoria parietina'' is a common and widespread lichen-forming fungus in the family Teloschistaceae. Commonly known as the yellow wall lichen, common orange lichen, or maritime sunburst lichen, this foliose lichen, leafy lichen is known f ...
'' has a significant history of use. In Spain, this lichen has been traditionally included in wine-based
decoction Decoction is a method of extraction by boiling herbal or plant material (which may include stems, roots, bark and rhizomes) to dissolve the chemicals of the material. It is the most common preparation method in various herbal medicine systems. D ...
s for
menstrual The menstrual cycle is a series of natural changes in hormone production and the structures of the uterus and ovaries of the female reproductive system that makes pregnancy possible. The ovarian cycle controls the production and release of egg ...
issues, and infused in water as a purported remedy for kidney and tooth ailments. It has also served as an
analgesic An analgesic drug, also called simply an analgesic, antalgic, pain reliever, or painkiller, is any member of the group of drugs used for pain management. Analgesics are conceptually distinct from anesthetics, which temporarily reduce, and in s ...
and a component in a
cold medicine Cold medicines are a group of medications taken individually or in combination as a treatment for the symptoms of the common cold and similar conditions of the upper respiratory tract. The term encompasses a broad array of drugs, including ana ...
. During the
early modern era The early modern period is a historical period that is defined either as part of or as immediately preceding the modern period, with divisions based primarily on the history of Europe and the broader concept of modernity. There is no exact date ...
in Europe, ''Xanthoria parietina'' was commonly boiled in milk as a treatment for
jaundice Jaundice, also known as icterus, is a yellowish or, less frequently, greenish pigmentation of the skin and sclera due to high bilirubin levels. Jaundice in adults is typically a sign indicating the presence of underlying diseases involving ...
, a practice also applied to '' Polycauliona candelaria''. ''X. parietina'' has been used to treat diarrhoea,
dysentery Dysentery ( , ), historically known as the bloody flux, is a type of gastroenteritis that results in bloody diarrhea. Other symptoms may include fever, abdominal pain, and a feeling of incomplete defecation. Complications may include dehyd ...
, bleeding, as a
malaria Malaria is a Mosquito-borne disease, mosquito-borne infectious disease that affects vertebrates and ''Anopheles'' mosquitoes. Human malaria causes Signs and symptoms, symptoms that typically include fever, Fatigue (medical), fatigue, vomitin ...
remedy in the absence of
quinine Quinine is a medication used to treat malaria and babesiosis. This includes the treatment of malaria due to ''Plasmodium falciparum'' that is resistant to chloroquine when artesunate is not available. While sometimes used for nocturnal leg ...
, and for
hepatitis Hepatitis is inflammation of the liver parenchyma, liver tissue. Some people or animals with hepatitis have no symptoms, whereas others develop yellow discoloration of the skin and whites of the eyes (jaundice), Anorexia (symptom), poor appetite ...
. In
Traditional Chinese medicine Traditional Chinese medicine (TCM) is an alternative medicine, alternative medical practice drawn from traditional medicine in China. A large share of its claims are pseudoscientific, with the majority of treatments having no robust evidence ...
, this lichen has been used as an
antibacterial An antibiotic is a type of antimicrobial substance active against bacteria. It is the most important type of antibacterial agent for fighting bacterial infections, and antibiotic medications are widely used in the treatment and prevention ...
. In regional practices, '' Rusavskia elegans'' is used for treating wounds; in
Afghanistan Afghanistan, officially the Islamic Emirate of Afghanistan, is a landlocked country located at the crossroads of Central Asia and South Asia. It is bordered by Pakistan to the Durand Line, east and south, Iran to the Afghanistan–Iran borde ...
, it is applied directly. 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 ...
, the lichen is mixed with butter and used as a remedy for diarrhoea in livestock. ''
Teloschistes flavicans ''Teloschistes flavicans'', also known as the golden hair-lichen, is a lichenized species of fungus in the genus ''Teloschistes'' (meaning "split-ends"; a reference to the plant's finely divided thallus and dense coils which appear almost like Br ...
'' is used in China for its purported properties of "clearing heat" in the lung and liver, and removing toxins. '' Oxneria fallax'' has been incorporated into traditional Tibetan medicinal treatments.


In science

The presence of ''Xanthoria parietina'' in
urban ecosystem In ecology, urban ecosystems are considered a ecosystem functional group within the intensive land-use biome. They are structurally complex ecosystems with highly heterogeneous and dynamic spatial structure that is created and maintained by hum ...
s, even in high pollution areas, offers potential insights for
biomonitoring In analytical chemistry, biomonitoring is the measurement of the body burden of toxic chemical compounds, elements, or their metabolites, in biological substances. Often, these measurements are done in blood and urine. Biomonitoring is performe ...
. Although its tolerance to
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 ...
allows it to persist where other species may not, its presence can be used in conjunction with more pollution-sensitive species to gauge overall air quality. In polluted environments, the health and abundance of ''X. parietina'', relative to other less tolerant species, can provide data on the level and impact of urban pollution. ''Rusavskia elegans'' has been studied in experiments where specimens were exposed to outer space conditions, including extreme temperatures,
ultraviolet Ultraviolet radiation, also known as simply UV, is electromagnetic radiation of wavelengths of 10–400 nanometers, shorter than that of visible light, but longer than X-rays. UV radiation is present in sunlight and constitutes about 10% of ...
radiation, and
ultra-high vacuum Ultra-high vacuum (often spelled ultrahigh in American English, UHV) is the vacuum regime characterised by pressures lower than about . UHV conditions are created by pumping the gas out of a UHV chamber. At these low pressures the mean free path of ...
. The results demonstrated the lichen's substantial capability to endure these conditions.


Conservation

The
conservation status The conservation status of a group of organisms (for instance, a species) indicates whether the group still exists and how likely the group is to become extinct in the near future. Many factors are taken into account when assessing conservation ...
of three Teloschistaceae species has been assessed for the global
IUCN Red List The International Union for Conservation of Nature (IUCN) Red List of Threatened Species, also known as the IUCN Red List or Red Data Book, founded in 1964, is an inventory of the global conservation status and extinction risk of biological ...
. '' Caloplaca rinodinae-albae'' ( vulnerable, 2017) is at risk from
tourism Tourism is travel for pleasure, and the Commerce, commercial activity of providing and supporting such travel. World Tourism Organization, UN Tourism defines tourism more generally, in terms which go "beyond the common perception of tourism as ...
development and increased
erosion Erosion is the action of surface processes (such as Surface runoff, water flow or wind) that removes soil, Rock (geology), rock, or dissolved material from one location on the Earth's crust#Crust, Earth's crust and then sediment transport, tran ...
on
Sardinia Sardinia ( ; ; ) is the Mediterranean islands#By area, second-largest island in the Mediterranean Sea, after Sicily, and one of the Regions of Italy, twenty regions of Italy. It is located west of the Italian Peninsula, north of Tunisia an ...
's coasts. '' Xanthaptychia aurantiaca'' (
Endangered An endangered species is a species that is very likely to become extinct in the near future, either worldwide or in a particular political jurisdiction. Endangered species may be at risk due to factors such as habitat loss, poaching, inv ...
, 2020) faces multiple threats due to
climate change Present-day climate change includes both global warming—the ongoing increase in Global surface temperature, global average temperature—and its wider effects on Earth's climate system. Climate variability and change, Climate change in ...
in the
Canadian Arctic Northern Canada (), colloquially the North or the Territories, is the vast northernmost region of Canada, variously defined by geography and politics. Politically, the term refers to the three territories of Canada: Yukon, Northwest Territories a ...
. These include
habitat loss Habitat destruction (also termed habitat loss or habitat reduction) occurs when a natural habitat is no longer able to support its native species. The organisms once living there have either moved elsewhere, or are dead, leading to a decrease ...
from rapidly eroding coasts, increased
sea ice Sea ice arises as seawater freezes. Because ice is less density, dense than water, it floats on the ocean's surface (as does fresh water ice). Sea ice covers about 7% of the Earth's surface and about 12% of the world's oceans. Much of the world' ...
melt,
saline Saline may refer to: Salt-related * Saline (medicine), a liquid with salt content to match the human body * Saline water, non-medicinal salt water * Saline, a historical term (especially American) for a salt works or saltern Places United States ...
wash from
storm surge A storm surge, storm flood, tidal surge, or storm tide is a coastal flood or tsunami-like phenomenon of rising water commonly associated with low-pressure weather systems, such as cyclones. It is measured as the rise in water level above the ...
s, and
permafrost Permafrost () is soil or underwater sediment which continuously remains below for two years or more; the oldest permafrost has been continuously frozen for around 700,000 years. Whilst the shallowest permafrost has a vertical extent of below ...
melting. Further, the changing climate may allow the advancement of southern vegetation communities and the introduction of
invasive species An invasive species is an introduced species that harms its new environment. Invasive species adversely affect habitats and bioregions, causing ecological, environmental, and/or economic damage. The term can also be used for native spec ...
, potentially exacerbating the impact on this lichen by altering its native habitat. '' Teloschistes peruensis'' (
Critically Endangered An IUCN Red List critically endangered (CR or sometimes CE) species is one that has been categorized by the International Union for Conservation of Nature as facing an extremely high risk of extinction in the wild. As of December 2023, of t ...
, 2021) is at risk due to multiple threats in Peru and Chile, including potential development,
habitat fragmentation Habitat fragmentation describes the emergence of discontinuities (fragmentation) in an organism's preferred environment (habitat), causing population fragmentation and ecosystem decay. Causes of habitat fragmentation include geological proces ...
,
4x4 A four-wheel drive, also called 4×4 ("four by four") or 4WD, is a two-axled vehicle drivetrain capable of providing torque to all of its wheels simultaneously. It may be full-time or on-demand, and is typically linked via a transfer case provi ...
races like the
Dakar Rally The Dakar Rally () or simply "The Dakar" (), formerly known as the Paris–Dakar Rally (), is an annual rally raid organised by the Amaury Sport Organisation (ASO). It is an off-road endurance event traversing terrain much tougher than convent ...
, air pollution, and the impact of livestock like goats and cows, which alter the habitat through grazing and trampling. Other Teloschistaceae members, some with limited geographic distributions, make appearances on regional red lists. For example, the crustose New Zealand endemic ''
Caloplaca allanii ''Caloplaca'' is a lichen genus comprising a number of distinct species. Members of the genus are commonly called firedot lichen, jewel lichen.Field Guide to California Lichens, Stephen Sharnoff, Yale University Press, 2014, gold lichens, "ora ...
'', first documented in 1932, was not collected again until 81 years later. Because of its sparsity and small total area of occupancy, it has been assessed as "Threatened/Nationally Critical" in the
New Zealand Threat Classification System The New Zealand Threat Classification System is used by the Department of Conservation to assess conservation priorities of species in New Zealand. The system was developed because the IUCN Red List, a similar conservation status system, had s ...
. In some large geographical areas, the full extent of the
diversity Diversity, diversify, or diverse may refer to: Business *Diversity (business), the inclusion of people of different identities (ethnicity, gender, age) in the workforce *Diversity marketing, marketing communication targeting diverse customers * ...
of Teloschistaceae taxa is not well known. Examples include South America, where the family has not historically received much attention, and China, where of 2,164 lichen species evaluated for inclusion on its red list, only 49 were members of the Teloschistaceae; 13 of those were listed as
least-concern species A least-concern species is a species that has been evaluated and categorized by the International Union for Conservation of Nature (IUCN) as not being a focus of wildlife conservation because the specific species is still plentiful in the wil ...
, and the other 36 as
data deficient A data deficient (DD) species is one which has been categorized by the International Union for Conservation of Nature (IUCN) as offering insufficient information for a proper assessment of conservation status to be made. This does not necessaril ...
.


Notes


References


Citations


Cited literature

* * * * * * * * * * * * * {{Taxonbar, from=Q2033908 Teloschistales Lichen families Lecanoromycetes families Taxa named by Alexander Zahlbruckner Taxa described in 1898