Graphidales Genera
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Graphidales is an
order Order, ORDER or Orders may refer to: * A socio-political or established or existing order, e.g. World order, Ancien Regime, Pax Britannica * Categorization, the process in which ideas and objects are recognized, differentiated, and understood ...
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 class Lecanoromycetes. It contains 6
families Family (from ) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictability, structure, and safety as ...
, about 81
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 about 2,228 species. Family Graphidaceae are the largest
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 ...
family within Graphidales order comprising more than 2000 species, which are widely distributed in tropical and subtropical regions of the world.


History

The Graphidales were introduced in a 1884 publication by Frigyes Ákos Hazslinszky in Magyar Birodalom Zuzmó-Flórája on page 216 as family Graphideae. In 1907, they were established as an
order Order, ORDER or Orders may refer to: * A socio-political or established or existing order, e.g. World order, Ancien Regime, Pax Britannica * Categorization, the process in which ideas and objects are recognized, differentiated, and understood ...
by American botanist Bessey (1845–1915), When the order was introduced, it contained just two families, the Graphidaceae and Thelotremataceae who were both mainly tropical based and each family had about 800–1000 species. Sherwood in 1977 proposed to maintain a distinction between the Graphidales with mostly lichenised members and the Ostropales which included mostly non-lichenised fungi, based on different spore septation types. Molecular data by Winka et al. in 1998, supported a close relationship between the two groups of species. The Graphidales were then included in the Ostropales order (Lecanoromycetes) for a long time (Staiger 2002; Kalb et al. 2004; Hibbett et al. 2007; Lumbsch et al. 2007; Kirk et al. 2008; Baloch et al. 2010; Rivas Plata and Lumbsch 2011; Rivas Plata et al. 2012; Lumbsch et al. 2014; Lücking et al. 2017; Wijayawardene et al. 2018). In 2004, the phylogenetic relationships of class Lecanoromycetes were examined by using
mitochondrial A mitochondrion () is an organelle found in the cells of most eukaryotes, such as animals, plants and fungi. Mitochondria have a double membrane structure and use aerobic respiration to generate adenosine triphosphate (ATP), which is used ...
small subunit
ribosomal DNA The ribosomal DNA (rDNA) consists of a group of ribosomal RNA encoding genes and related regulatory elements, and is widespread in similar configuration in all domains of life. The ribosomal DNA encodes the non-coding ribosomal RNA, integral struc ...
sequencing which found that orders Graphidales and Ostropales were
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 ...
. Using molecular data (partial DNA sequencing) in 2012, it was also shown that Graphidaceae and Thelotremataceae were non-monophyletic and consequently Thelotremataceae was included in Graphidaceae as a synonym. Graphidaceae also included subfamilies Fissurinoideae and Graphidoideae. However, Kraichak et al. in 2018, ranked Graphidales as a separate order based on a temporal approach, and accepted five families; Diploschistaceae, Fissurinaceae, Gomphillaceae, Graphidaceae and Thelotremataceae. Wijayawardene et al. 2020 agreed but also added family Redonographaceae to the order, Other authors have agreed on the use of reinstated order of Graphidales.


Description

Most species in the order are
lichens 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 ...
which have a
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 ...
(vegetative tissue) which is
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 ...
and ascocarps (fruiting body) which are apothecioid (cup-shaped).Michael Allaby (Editor) The Graphidaceae are mostly
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 ...
lichens with trentepohlioid
photobiont A lichen ( , ) is a hybrid colony of algae or cyanobacteria living symbiotically among filaments of multiple fungus species, along with yeasts and bacteria embedded in the cortex or "skin", in a mutualistic relationship.
(i.e., filamentous, multicellular
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 genus '' Trentepohlia''), and graphidoid, distoseptate (forming a layer)
ascospores 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 of fungi. After two parental nuclei fuse, the ascus undergoes meiosis (halving of ...
. Family Gomphillaceae was originally based on a single species, '' Gomphillus calycioides'' (Watson, 1929), which is an unusual taxon growing over
bryophytes Bryophytes () are a group of land plants ( embryophytes), sometimes treated as a taxonomic division referred to as Bryophyta '' sensu lato'', that contains three groups of non-vascular land plants: the liverworts, hornworts, and mosses. In t ...
. It includes a common asexual fungus '' Lawreya glyphidiphila'' ( Teratosphaeriaceae family) which is described as growing on lichenized fungi '' Glyphis scyphulifera'' (Graphidaceae family).


Distribution

They are mainly found in warmer regions living on bark. Genera in the order of Graphidales has been found worldwide, from North America (including
Florida Florida ( ; ) is a U.S. state, state in the Southeastern United States, Southeastern region of the United States. It borders the Gulf of Mexico to the west, Alabama to the northwest, Georgia (U.S. state), Georgia to the north, the Atlantic ...
,), South America (including
Venezuela Venezuela, officially the Bolivarian Republic of Venezuela, is a country on the northern coast of South America, consisting of a continental landmass and many Federal Dependencies of Venezuela, islands and islets in the Caribbean Sea. It com ...
, Costa Rica, and
Guianas The Guianas, also spelled Guyanas or Guayanas, are a geographical region in north-eastern South America. Strictly, the term refers to the three Guianas: Guyana, Suriname, and French Guiana, formerly British Guiana, British, Surinam (Dutch colo ...
,), Africa (including
Kenya Kenya, officially the Republic of Kenya, is a country located in East Africa. With an estimated population of more than 52.4 million as of mid-2024, Kenya is the 27th-most-populous country in the world and the 7th most populous in Africa. ...
, and South Africa,), Asia (including China,
Vietnam Vietnam, officially the Socialist Republic of Vietnam (SRV), is a country at the eastern edge of mainland Southeast Asia, with an area of about and a population of over 100 million, making it the world's List of countries and depende ...
,
Sri Lanka Sri Lanka, officially the Democratic Socialist Republic of Sri Lanka, also known historically as Ceylon, is an island country in South Asia. It lies in the Indian Ocean, southwest of the Bay of Bengal, separated from the Indian subcontinent, ...
, India, and
Thailand Thailand, officially the Kingdom of Thailand and historically known as Siam (the official name until 1939), is a country in Southeast Asia on the Mainland Southeast Asia, Indochinese Peninsula. With a population of almost 66 million, it spa ...
,) Australia, and also New Zealand. Species of family Gomphillaceae are found in north-eastern Brazil, Mexico, Guatemala, Costa Rica,
Panama Panama, officially the Republic of Panama, is a country in Latin America at the southern end of Central America, bordering South America. It is bordered by Costa Rica to the west, Colombia to the southeast, the Caribbean Sea to the north, and ...
and Cuba.


Families and genera

This is a list of the families and genera contained within the Graphidales, based on a 2020 review and summary of ascomycete classification. Following the taxon name is the
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 (for genera) the number or estimated number of species:


Diploschistaceae (1905)

*'' Acanthothecis'' (ca. 60) *'' Acanthotrema'' (6) *'' Aggregatorygma'' (2) *'' Ampliotrema'' (17) *''
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) *'' Austrotrema'' (3) *'' Borinquenotrema'' (1) *'' Byssotrema'' (1) *''
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. ...
'' (28) *'' Compositrema'' (4) *'' Corticorygma'' (1) *'' Diploschistes'' (33) *'' Fibrillithecis'' (15) *'' Gintarasia'' (8) *'' Glaucotrema'' (5) *'' Gyrotrema'' (6) *'' Heiomasia'' (5) *'' Melanotopelia'' (4) *'' Melanotrema'' (12) *'' Myriochapsa'' (3) *'' Myriotrema'' (55) *'' Nadvornikia'' (5) *'' Nitidochapsa'' (5) *''
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) ...
'' (ca. 400) *''
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) *'' Pseudoramonia'' (4) *'' Redingeria'' (9) *'' Reimnitzia'' (1) *'' Rhabdodiscus'' (36) *'' Sanguinotrema'' (1) *'' Schizotrema'' (8) *'' Stegobolus'' (16) *'' Topeliopsis'' (20) *'' Wirthiotrema'' (5) *'' Xalocoa'' (1)


Fissurinaceae (2012)

*'' Clandestinotrema'' (17) *'' Cruentotrema'' (7) *'' Dyplolabia'' (5) *'' Enigmotrema'' (1) *''
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 ...
'' (ca. 155) *'' Pycnotrema'' (2)


Gomphillaceae (1984)

*'' Actinoplaca'' (2) *''
Aderkomyces ''Aderkomyces'' is a genus of lichens in the family Gomphillaceae. It was circumscription (taxonomy), circumscribed by Brazilian mycologist Augusto Chaves Batista in 1961. Species , Species Fungorum accepts 28 species of ''Aderkomyces''. *''Ade ...
'' (30) *'' Aplanocalenia'' (1) *'' Arthotheliopsis'' (5) *''
Asterothyrium ''Asterothyrium'' is a genus of foliicolous lichen, leaf-dwelling lichens in the family Gomphillaceae. Species , Species Fungorum (in the Catalogue of Life) accepts 13 species of ''Asterothyrium'': *''Asterothyrium argenteum'' *''Asterothyriu ...
'' (32) *'' Aulaxina'' (14) *'' Calenia'' (30) *'' Caleniopsis'' (2) *'' Cladosterigma'' (1) *'' Corticifraga'' (9) *''
Diploschistella ''Diploschistella'' is a genus of 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 mushro ...
'' (4) *''
Echinoplaca ''Echinoplaca'' is a genus of lichens in the family Gomphillaceae. Species , Species Fungorum (in the Catalogue of Life) accepts 18 species of ''Echinoplaca''. * ''Echinoplaca areolata'' * ''Echinoplaca atromuralis'' * ''Echinoplaca basalis' ...
'' (40) *'' Ferraroa'' (1) *''
Gomphillus ''Gomphillus'' is a genus of lichen-forming fungi in the family Gomphillaceae. Taxonomy The genus was circumscription (taxonomy), circumscribed by the Finnish lichenologist William Nylander (botanist), William Nylander in 1854, with ''Gomphillu ...
'' (6) *'' Gyalectidium'' (52) *''
Gyalidea ''Gyalidea'' is a genus of crustose lichens in the family Gomphillaceae. It has 50 species. Species *''Gyalidea asteriscus'' *''Gyalidea austrocoreana'' *''Gyalidea corticola'' *''Gyalidea diaphana'' *''Gyalidea fritzei'' *''Gyalidea fruct ...
'' (50) *'' Gyalideopsis'' (91) *'' Hippocrepidea'' (1) *'' Jamesiella'' (4) *'' Lithogyalideopsis'' (4) *'' Paragyalideopsis'' (4) *'' Paratricharia'' (1) *'' Phyllogyalidea'' (2) *'' Psorotheciopsis'' (7) *'' Rolueckia'' (3) *'' Taitaia'' (1) *'' Tricharia'' (ca. 30)


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 ...
(1822)

*'' Allographa'' (ca. 185) *'' Anomalographis'' (2) *'' Anomomorpha'' (8) *'' Creographa'' (2) *'' Cryptoschizotrema'' (2) *'' Diaphorographis'' (2) *'' Diorygma'' (77) *'' Flegographa'' (1) *'' Glyphis'' (7) *'' Graphis'' (ca. 275) *'' Halegrapha'' (9) *''
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) *'' Jocatoa'' (1) *'' Kalbographa'' (5) *'' Leiorreuma'' (18) *'' Malmographina'' (1) *'' Mangoldia'' (2) *'' Pallidogramme'' (14) *'' Phaeographis'' (ca. 180) *'' Platygramme'' (30) *'' Platythecium'' (27) *'' Pliariona'' (1) *'' Polistroma'' (1) *'' Pseudochapsa'' (19) *'' Pseudotopeliopsis'' (4) *'' Sarcographa'' (37) *'' Sarcographina'' (6) *'' Schistophoron'' (5) *'' Thalloloma'' (20) *'' Thecaria'' (4) *'' Thecographa'' (3)


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 ...
(2020)

*'' Gymnographopsis'' (3) *'' Redonographa'' (5)


Thelotremataceae (1862)

*'' Astrochapsa'' (29) *'' Chapsa'' (ca. 60) *'' Chroodiscus'' (17) *'' Crutarndina'' (1) *'' Leucodecton'' (31) *'' Paratopeliopsis'' (1) *''
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 ...
'' (includes ''Tremotylium'' ) (165)


References

{{Authority control Graphidales Ascomycota orders Taxa described in 1884