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'' were included in the new family.
In 2002, the German lichenologist
Bettina Staiger 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).
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'' 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 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''. 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 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 (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''
– 5 spp.
*''
Acanthotrema''
– 1 sp.
*''
Aggregatorygma''
– 3 spp.
*''
Allographa''
– 183 spp.
*''
Amazonotrema''
– 1 sp.
*''
Ampliotrema''
– 1 sp.
*''
Anomalographis'' – 2 spp.
*''
Anomomorpha'' – 8 spp.
*''
Asteristion'' – 7 spp.
*''
Astrochapsa''
– 29 spp.
*''
Austrotrema'' – 3 spp.
*''
Borinquenotrema''
– 1 sp.
*''
Byssotrema''
– 1 sp.
*''
Carbacanthographis''
– 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'' – 74 spp.
*''
Diploschistes''
– 25 spp.
*''
Dyplolabia''
– 5 spp.
*''
Enigmotrema'' – 1 sp.
*''
Fibrillithecis''
– 15 spp.
*''
Fissurina''
– 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''
– 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''
– 343 spp.
*''
Pallidogramme''
– 13 spp.
*''
Paratopeliopsis''
– 1 sp.
*''
Phaeographina''
– 11 spp.
*''
Phaeographis'' – ca. 180 spp.
*''
Phaeographopsis''
– 3 spp.
*''
Phaeotrema''
– 19 spp.
*''
Platygramme''
– 30 spp.
*''
Platythecium''
– 27 spp.
*''
Pliariona''
– 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'' – 37 spp.
*''
Sarcographina'' – 6 spp.
*''
Schistophoron'' – 5 spp.
*''
Schistostoma'' – 1 sp.
*''
Schizotrema''
– 7 spp.
*''
Stegobolus'' – 16 spp.
*''
Thalloloma''
– 20 spp.
*''
Thecaria'' – 4 spp.
*''
Thecographa'' – 3 spp.
*''
Thelotrema'' – 165 spp.
*''
Topeliopsis''
– 20 spp.
*''
Tremotylium''
– 4 spp.
*''
Wirthiotrema''
– 5 spp.
*''
Xalocoa''
– 1 sp.
References
{{Authority control
*
Lecanoromycetes families
Taxa described in 1822
Taxa named by Barthélemy Charles Joseph Dumortier
Lichen families