Diploschistaceae
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The Graphidaceae are a
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
lichen A lichen ( , ) is a hybrid colony (biology), colony of algae or cyanobacteria living symbiotically among hypha, filaments of multiple fungus species, along with yeasts and bacteria embedded in the cortex or "skin", in a mutualism (biology), m ...
-forming
fungi A fungus (: fungi , , , or ; or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and mold (fungus), molds, as well as the more familiar mushrooms. These organisms are classified as one ...
in the order
Graphidales Graphidales is an order (biology), order of lichen-forming fungi in the class Lecanoromycetes. It contains 6 family (biology), families, about 81 genus, genera and about 2,228 species. Family Graphidaceae are the largest crustose family within G ...
. The family contains nearly a hundred
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 ...
and more than 2000 species. Although 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 ...
, most Graphidaceae species occur in tropical regions, and typically grow on bark.


Taxonomy

Graphidaceae was originally proposed by French botanist
Barthélemy Charles Joseph Dumortier Barthélemy Charles Joseph Dumortier (; 3 April 1797 – 9 July 1878) was a Belgians, Belgian who conducted a parallel career of botanist and Member of Parliament and is the first discoverer of biological cell division. Over the course of his lif ...
in 1822 (as "Graphineae"). '' Graphis'', '' Opegrapha'', and ''
Arthonia ''Arthonia'' is a genus of lichens in the family Arthoniaceae. It was circumscribed by Swedish botanist Erik Acharius in 1806. It is a genus of thin crustose lichen of widely varying forms, commonly called comma lichens.Field Guide to California ...
'' were included in the new family. In 2002, the German lichenologist
Bettina Staiger Bettina Staiger (born 1968) is a German lichenologist known for her systematic studies of tropical lichen families, particularly the Graphidaceae. Born in Stuttgart, she completed her doctoral research at the University of Regensburg under Kl ...
revised the Graphidaceae in a
monograph A monograph is generally a long-form work on one (usually scholarly) subject, or one aspect of a subject, typically created by a single author or artist (or, sometimes, by two or more authors). Traditionally it is in written form and published a ...
, proposing a new classification of genera that was widely accepted until
molecular phylogenetic Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...
studies led to a further reorganization of the family. Two subfamilies are recognized in the Graphidaceae: *Fissurinoideae *Graphidoideae Subfamily Redonographoideae, proposed by Lücking and colleagues in 2013, has since been promoted to familial status (as the monogeneric family
Redonographaceae ''Redonographa'' is a genus of lichen-forming fungi in the monogeneric family Redonographaceae. It has five species. Taxonomy In 2013, lichenologists Robert Lücking, Anders Tehler, and Helge Thorsten Lumbsch proposed the new family (biology ...
).


Synonymy

The family Thelotremataceae was placed in
synonymy 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 ...
with Graphidaceae in 2008, after molecular phylogenetic analysis showed that the families formed several lineages within one strongly supported
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 ...
lineage. This study also showed that many genera in Graphidaceae and Thelotremataceae were
paraphyletic Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...
or
polyphyletic A polyphyletic group is an assemblage that includes organisms with mixed evolutionary origin but does not include their most recent common ancestor. The term is often applied to groups that share similar features known as Homoplasy, homoplasies ...
. In 2018, Kraichak and colleagues, using a "temporal phylogenetic" approach to identify temporal bands for specific
taxonomic rank In biology, taxonomic rank (which some authors prefer to call nomenclatural rank because ranking is part of nomenclature rather than taxonomy proper, according to some definitions of these terms) is the relative or absolute level of a group of or ...
s, proposed removing ''
Fissurina ''Fissurina'' is a genus of lichenized fungi in the family Graphidaceae. It has about 160 species, most of which are found in tropical regions. Taxonomy The genus was circumscription (taxonomy), circumscribed by the French botanist Antoine Lau ...
'' from the Graphidaceae and instead as placing it as the type genus of Fissurinaceae, a family originally proposed by Brendan P. Hodkinson in 2012. They similarly suggested recognizing Diploschistaceae and Thelotremataceae as independent families. This taxonomic proposal was rejected by
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 ...
in a critical 2019 review of the temporal method for the classification of lichen-forming fungi, using these specific examples to highlight several drawbacks of this approach. He noted "there is substantial loss of information content in this alternative classification at the second most important rank, the level of family, and such a revised classification is practically meaningless. It makes more sense to apply such essentially phylogenetic classifications, with less information content, to infraranks, such as subfamily or tribe".


Description

The photosynthetic partner () of Graphidaceae fungi is typically
green algae 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 ...
from the genus '' Trentepohlia'', or very rarely ''
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
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 ...
is typically
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 ...
, which means it forms a crust-like structure on the it grows on. The form of the ascomata of Graphidaceae are usually (cup-shaped), (narrow, and slit-like) or (round or oval-shaped structure and embedded in the substrate), but can be rarely (with a powdery mass of ascospores and formed by the disintegration of the asci). They are usually (with both a and a ), but are sometimes (without a thallus margin) or (surrounded by a pale thalline margin). The , which is the hyphae and tissue that between the asci, consists of usually unbranched, sometimes , rarely branched and anastomosing paraphyses. In some lineages, the paraphyses may be apically (covered with small spines or projections at the tips), and there may be lateral in some lineages. The asci are , which means they have a ring structure protruding into the and with an apical . They are non-
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 ...
or amyloid in some lineages, such as in the genus ''
Diorygma ''Diorygma'' is a genus of lichen-forming fungi in the family Graphidaceae. The genus was circumscribed by Franz Gerhard Eschweiler in 1824. Species of the genus are widely distributed in tropical and subtropical regions of the world. These l ...
''. The asci are to oblong or in shape. The ascospores of Graphidaceae number primarily 8 per ascus but can often be reduced to 2–4 or 1 per ascus. They are transversely
septate In biology, a septum (Latin for ''something that encloses''; septa) is a wall, dividing a cavity or structure into smaller ones. A cavity or structure divided in this way may be referred to as septate. Examples Human anatomy * Interatrial se ...
to (elongated and divided into multiple compartments by transverse and longitudinal septa), usually
ellipsoid An ellipsoid is a surface that can be obtained from a sphere by deforming it by means of directional Scaling (geometry), scalings, or more generally, of an affine transformation. An ellipsoid is a quadric surface;  that is, a Surface (mathemat ...
to oblong, and often have forming and lens-shaped to rounded lumina. This particular set of features is known as "graphidoid". The spores can be
hyaline A hyaline substance is one with a glassy appearance. The word is derived from , and . Histopathology Hyaline cartilage is named after its glassy appearance on fresh gross pathology. On light microscopy of H&E stained slides, the extracellula ...
(colourless) to (dark) brown, and the endospore is often amyloid. Graphidaceae also have
pycnidia A pycnidium (plural pycnidia) is an asexual fruiting body produced by mitosporic fungi, for instance in the order Sphaeropsidales ( Deuteromycota, Coelomycetes) or order Pleosporales (Ascomycota, Dothideomycetes). It is often spherical or inve ...
for producing
conidia A conidium ( ; : conidia), sometimes termed an asexual chlamydospore or chlamydoconidium (: chlamydoconidia), is an asexual, non- motile spore of a fungus. The word ''conidium'' comes from the Ancient Greek word for dust, ('). They are also ...
, which are non-septate, usually oblong, and hyaline.


Distribution and ecology

The vast majority of Graphidaceae species are restricted to the
tropics The tropics are the regions of Earth surrounding the equator, where the sun may shine directly overhead. This contrasts with the temperate or polar regions of Earth, where the Sun can never be directly overhead. This is because of Earth's ax ...
. Most Graphidaceae species are
epiphytic An epiphyte is a plant or plant-like organism that grows on the surface of another plant and derives its moisture and nutrients from the air, rain, water (in marine environments) or from debris accumulating around it. The plants on which epiphyt ...
(i.e. they grow only on plants). Forty-two species are known from the
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 ...
, where they are among the most diverse of the crustose lichens there. Mexico is thought to be a
biodiversity Biodiversity is the variability of life, life on Earth. It can be measured on various levels. There is for example genetic variability, species diversity, ecosystem diversity and Phylogenetics, phylogenetic diversity. Diversity is not distribut ...
hotspot of undiscovered Graphidaceae species, with about 430 species predicted to occur in tropical regions, compared to less than 200 recorded in the entire country.


Genera

According to 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 ...
, there are 94 genera and more than 2100 species in Graphidaceae. In terms of number of species, Graphidaceae is the second-largest family of lichen-forming fungi, after the
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 ...
(2765 species) and ahead of the
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 ...
(943 species). The following list gives the genus name, 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 ...
, year of publication, and the number of species: *''
Acanthothecis ''Acanthothecis'' is a genus of lichen-forming fungi in the family Graphidaceae. The genus was circumscribed by Frederick Edward Clements in 1909. These lichens form pale grey-brown to olive-green crusts on tree bark and are characterized by ...
'' – 5 spp. *'' Acanthotrema'' – 1 sp. *'' Aggregatorygma'' – 3 spp. *''
Allographa ''Allographa'' is a genus of script lichens in the family Graphidaceae. It has nearly 200 species. Formally circumscription (taxonomy), circumscribed in 1824 by François Fulgis Chevallier, ''Allographa'' was formerly included in ''Graphis (liche ...
'' – 183 spp. *'' Amazonotrema'' – 1 sp. *'' Ampliotrema'' – 1 sp. *'' Anomalographis'' – 2 spp. *''
Anomomorpha ''Anomomorpha'' is a genus of lichens in the family Graphidaceae. The genus, described in 1891, has a pantropical distribution. These lichens form pale grey to dull olive crusts on tree bark that often develop low wart-like bumps covered with ...
'' – 8 spp. *''
Asteristion ''Asteristion'' is a genus of lichen-forming fungi in the family Graphidaceae. It has seven species. Previously considered a synonym (taxonomy), synonym of either ''Phaeotrema'' or ''Thelotrema'', molecular phylogenetics, molecular evidence led ...
'' – 7 spp. *'' Astrochapsa'' – 29 spp. *''
Austrotrema ''Austrotrema'' is a genus of lichen-forming fungi in the family Graphidaceae. It is primarily found in the Australian-Southeast Asian region. The genus is characterized by small, pore-like with a double margin, faintly amyloid ascospores, and ...
'' – 3 spp. *'' Borinquenotrema'' – 1 sp. *'' Byssotrema'' – 1 sp. *''
Carbacanthographis ''Carbacanthographis'' is a genus of corticolous lichen, corticolous (bark-dwelling) lichens in the family Graphidaceae. The genus was circumscription (taxonomy), circumscribed by the German lichenologists Bettina Staiger and Klaus Kalb in 2002. ...
'' – 22 spp. *'' Chapsa'' – 51 spp. *'' Chroodiscus'' – 17 spp. *'' Clandestinotrema'' – 17 spp. *'' Compositrema'' – 4 spp. *'' Corticorygma'' – 1 sp. *'' Creographa'' – 1 spp. *'' Cruentotrema'' – 6 spp. *'' Crutarndina'' – 1 sp. *'' Cryptoschizotrema'' – 2 spp. *'' Cyclographina'' – 8 spp. *'' Diaphorographis'' – 2 spp. *''
Diorygma ''Diorygma'' is a genus of lichen-forming fungi in the family Graphidaceae. The genus was circumscribed by Franz Gerhard Eschweiler in 1824. Species of the genus are widely distributed in tropical and subtropical regions of the world. These l ...
'' – 74 spp. *'' Diploschistes'' – 25 spp. *'' Dyplolabia'' – 5 spp. *'' Enigmotrema'' – 1 sp. *'' Fibrillithecis'' – 15 spp. *''
Fissurina ''Fissurina'' is a genus of lichenized fungi in the family Graphidaceae. It has about 160 species, most of which are found in tropical regions. Taxonomy The genus was circumscription (taxonomy), circumscribed by the French botanist Antoine Lau ...
'' – 117 spp. *'' Gintarasia'' – 7 spp. *'' Glaucotrema'' – 5 spp. *'' Glyphis'' – 7 spp. *'' Graphina'' – 25 spp. *'' Graphis'' – ca. 275 spp. *'' Gyrotrema'' – 6 spp. *'' Halegrapha'' – 9 spp. *'' Heiomasia'' – 3 spp. *''
Hemithecium ''Hemithecium'' is a genus of lichen-forming fungi in the family Graphidaceae. The genus was circumscribed by Vittore Benedetto Antonio Trevisan de Saint-Léon in 1853. Species , Species Fungorum (in the Catalogue of Life The Catalogue of ...
'' – ca. 50 spp. *'' Jocatoa'' – 1 sp. *''
Kalbographa ''Kalbographa'' is a genus of lichen-forming fungi in the family Graphidaceae. These lichens form thin, grey-white to pale yellow-olive crusts on tree bark and are distinguished by their striking bright orange to brick-red fruiting bodies that ...
'' – 5 spp. *'' Leiorreuma'' – 18 spp. *'' Leptotrema'' – 14 spp. *'' Leucodecton'' – 32 spp. *'' Malmographina'' – 1 sp. *'' Mangoldia'' – 4 spp. *'' Medusulina'' *'' Melanotopelia'' – 4 spp. *'' Melanotrema'' – 12 spp. *'' Myriochapsa'' – 3 spp. *'' Myriotrema'' – 76 spp. *'' Nadvornikia'' – 5 spp. *'' Nitidochapsa'' – 5 spp. *''
Ocellularia ''Ocellularia'' is a genus of lichens in the family Graphidaceae. The genus was circumscribed by Georg Friedrich Wilhelm Meyer Georg may refer to: * Georg (film), ''Georg'' (film), 1997 *Georg (musical), Estonian musical * Georg (given name) ...
'' – 343 spp. *'' Pallidogramme'' – 13 spp. *'' Paratopeliopsis'' – 1 sp. *'' Phaeographina'' – 11 spp. *''
Phaeographis ''Phaeographis'' is a genus of lichen-forming fungi in the family Graphidaceae. It has an estimated 180 species. Taxonomy The genus ''Phaeographis'' was circumscribed by the Swiss lichenologist Johannes Müller Argoviensis in 1882. However, its ...
'' – ca. 180 spp. *''
Phaeographopsis ''Phaeographopsis'' is a genus of script lichens in the family Graphidaceae. It has three species. Taxonomy The genus was circumscription (taxonomy), circumscribed by the Dutch lichenologist Harrie Sipman in 1997, with ''Phaeographopsis indica, ...
'' – 3 spp. *''
Phaeotrema ''Phaeotrema'' is a genus of lichen-forming fungi in the family Graphidaceae. The genus was circumscribed in 1887 by Johannes Müller Argoviensis, with '' P. subfarinosum'' assigned as the type species. Species , Species Fungorum (in the C ...
'' – 19 spp. *''
Platygramme ''Platygramme'' is a genus of lichen-forming fungi in the family Graphidaceae consisting of about 30 species. The genus was circumscribed by Antoine Laurent Apollinaire Fée in 1874. Several features characterise the genus ''Platygramme'': the ...
'' – 30 spp. *''
Platythecium ''Platythecium'' is a genus of lichen-forming fungi in the family Graphidaceae. It comprises 25 species. Described in 2002 by the German lichenologist Bettina Staiger, these bark-dwelling lichens are found in humid, shaded forests across trop ...
'' – 27 spp. *''
Pliariona ''Pliariona'' is a fungal genus in the family Graphidaceae. It comprises the single species ''Pliariona montagnei'', a corticolous (bark-dwelling) script lichen widely distributed across various tropical and subtropical regions. Taxonomy ''Pl ...
'' – 1 sp. *'' Polistroma'' – 1 sp. *'' Pseudochapsa'' – 18 spp. *'' Pseudoramonia'' – 4 spp. *'' Pseudotopeliopsis'' – 4 spp. *'' Pycnotrema'' – 2 spp. *'' Redingeria'' – 8 spp. *'' Reimnitzia'' – 1 sp. *'' Rhabdodiscus'' – 36 spp. *'' Sanguinotrema'' – 1 sp. *''
Sarcographa ''Sarcographa'' is a genus of lichen-forming fungi in the family Graphidaceae. Established in 1825 by the French botanist Antoine Laurent Apollinaire Fée, the genus contains 22 species that are recognised by their distinctive star-shaped coloni ...
'' – 37 spp. *'' Sarcographina'' – 6 spp. *''
Schistophoron ''Schistophoron'' is a genus of lichen-forming fungi in the family Graphidaceae. It comprises five species. Established in 1876 by the Scottish naturalist James Stirton, these unusual lichens grow exclusively on the undersides of living leaves ...
'' – 5 spp. *''
Schistostoma ''Schistostoma'' is a genus of fly, flies in the family Dolichopodidae, subfamily Microphorinae. Species At least 50 extant species have been described in the genus, with 16 from the Palaearctic realm, two from the Oriental realm, four from the ...
'' – 1 sp. *'' Schizotrema'' – 7 spp. *'' Stegobolus'' – 16 spp. *''
Thalloloma ''Thalloloma'' is a genus of lichen-forming fungi in the family Graphidaceae. Established in 1853 by the Italian botanist Vittore Trevisan de Saint-Léon, the genus comprises 20 species distinguished by their narrow script-like slits immersed ...
'' – 20 spp. *''
Thecaria ''Thecaria'' is a genus of lichen-forming fungi in the family 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 fam ...
'' – 4 spp. *'' Thecographa'' – 3 spp. *''
Thelotrema ''Thelotrema'' is a genus of lichen-forming fungi in the family Graphidaceae, the family to which all taxa in the former Thelotremataceae now belong. Members of the genus ''Thelotrema'' are commonly called barnacle lichens. Description ''Thelo ...
'' – 165 spp. *'' Topeliopsis'' – 20 spp. *'' Tremotylium'' – 4 spp. *''
Wirthiotrema ''Wirthiotrema'' is a genus of lichen-forming fungi in the family Graphidaceae. The genus was circumscribed in 2010 by Eimy Rivas Plata, Klaus Kalb, Andreas Frisch, and H. Thorsten Lumbsch, with '' Wirthiotrema glaucopallens'' assigned as the t ...
'' – 5 spp. *'' Xalocoa'' – 1 sp.


References

{{Authority control * Lecanoromycetes families Taxa described in 1822 Taxa named by Barthélemy Charles Joseph Dumortier Lichen families