Gomphillaceae
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The Gomphillaceae 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 ...
. Species in this family are found mostly in
tropical 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 ...
regions. The family underwent a major
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 ...
-led reorganisation in 2023, in which 17 genera were recognised and formally established as new or reinstated.


Taxonomy

The Gomphillaceae has undergone significant taxonomic revision as
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, ...
analysis techniques have improved scientists' understanding of relationships between species. As of 2024, the family includes approximately 440 accepted species, of which 422 are lichenised (form a symbiotic relationship with algae) and 18 are lichenicolous or fungicolous (grow on other lichens or fungi). The family is particularly diverse in tropical regions, where many species grow on leaves in wet forests. However, some members, particularly in the genus '' Gyalideopsis'', can also be found growing on other surfaces and extend into temperate regions. Recent
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 ...
studies (analysis of DNA sequences to determine evolutionary relationships) have led to a major reorganisation of the family's classification. The research identified 46 distinct genera, including 19 newly recognised lineages. This represents a significant expansion from the previous understanding of the family's diversity. A challenge in classifying members of this family is that only about 27% of known species have had their DNA sequenced. To address this, researchers have used a technique called "phylogenetic binning", which uses physical characteristics to predict where unsequenced species fit within the
phylogenetic tree A phylogenetic tree or phylogeny is a graphical representation which shows the evolutionary history between a set of species or taxa during a specific time.Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA. In ...
based on their similarities to sequenced relatives. The actual number of species in the family may be significantly higher than currently recognised, possibly exceeding 800 species. This is because detailed examination often reveals that what appears to be a single species may actually be several distinct but closely related species (known as
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 ...
). Recent (2024) sequencing work has filled one of the most persistent gaps in the family tree, with a two-gene phylogeny for the enigmatic ''Paratricharia paradoxa''. The analysis showed that '' Paratricharia'' is
sister A sister is a woman or a girl who shares parents or a parent with another individual; a female sibling. The male counterpart is a brother. Although the term typically refers to a familial relationship, it is sometimes used endearingly to ref ...
to '' Caleniopsis'', with the pair forming the earliest branch of a broadened ''Aulaxina''
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
that also contains '' Aulaxina'' and '' Aulaxinella''. This placement corroborates earlier morphology-based predictions and removes ''Paratricharia'' from any close relationship with '' Tricharia'' in the strict sense or the independently evolved "black-setae" lineages '' Microxyphiomyces'' and '' Santricharia''. Because the study emphasised apothecial architecture as a better indicator of deep relationships than the presence of sterile black bristles, several "setae-bearing" genera are now recognised as the result of
convergent evolution Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last comm ...
rather than shared ancestry. The same paper, together with the suite of genus-level descriptions produced in 2023, confirms that the ''Aulaxina'' clade represents one of at least five major lineages inside Gomphillaceae that have diversified chiefly in tropical leaf-dwelling habitats. These findings dovetail with a 2025 global phylogeny that sampled more than 500 terminals and predicted a total family richness well above 800 species once cryptic diversity is fully resolved. Taken together, the influx of molecular data and the continued use of phenotype-based "binning" for the many still-unsequenced species are progressively stabilising generic concepts, moving the family towards a classification that reflects both genetic relationships and diagnostic morphology.


Description

The family Gomphillaceae is characterized by its crust-like
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 ...
, which is the outer layer of the lichen body. These lichens contain photobionts, which are symbiotic algae that help the lichen to produce food through
photosynthesis Photosynthesis ( ) is a system of biological processes by which photosynthetic organisms, such as most plants, algae, and cyanobacteria, convert light energy, typically from sunlight, into the chemical energy necessary to fuel their metabo ...
. The reproductive structures of Gomphillaceae lichens, known as
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 ...
, can be apothecioid or lirellate in form. Apothecia occur in a few forms, including or to sometimes . The internal structure of
hymenium The hymenium is the tissue layer on the hymenophore of a fungal fruiting body where the cells develop into basidia or asci, which produce spores. In some species all of the cells of the hymenium develop into basidia or asci, while in oth ...
(the fertile, spore-producing part of the lichen), called the , consists primarily of branched and interwoven . The asci, or spore-producing sacs, are and feature a distinctive apical and ring structure. They can be , oblong, or in shape and do not exhibit any
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 ...
properties. These asci typically produce eight
ascospore In fungi, an ascospore is the sexual spore formed inside an ascus—the sac-like cell that defines the division Ascomycota, the largest and most diverse Division (botany), division of fungi. After two parental cell nucleus, nuclei fuse, the ascu ...
s, although some may produce fewer, ranging from one to four spores. The ascospores are
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 ...
(transparent), and have thin walls and distinct . They can be
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 ...
or oblong in shape, and their internal divisions (
septa SEPTA, the Southeastern Pennsylvania Transportation Authority, is a regional public transportation authority that operates bus, rapid transit, commuter rail, light rail, and electric trolleybus services for nearly four million people througho ...
) can be transverse or . Gomphillaceae lichens also produce , which are asexual reproductive structures. These are mostly , and the conidia are formed as branched
hypha A hypha (; ) is a long, branching, filamentous structure of a fungus, oomycete, or actinobacterium. In most fungi, hyphae are the main mode of vegetative growth, and are collectively called a mycelium. Structure A hypha consists of one o ...
e, called diahyphae, within gelatinous masses. The conidia are typically septate, often taking on a or bead-like appearance, and are also hyaline. In terms of secondary chemistry, Gomphillaceae lichens generally lack any notable substances.


Genera

According to a recent (2024) compilation of fungal classification, the Gomphillaceae comprise 44 genera and about 340 species. The following list indicates the genus name, 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 the number of species: * '' Actinoplaca'' – 2 spp. * '' Adelphomyces'' – 3 spp. * ''
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 spp. * '' Aplanocalenia'' – 1 sp. * '' Aptrootidea'' – 6 spp. * '' Arthotheliopsis'' – 5 spp. * ''
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 ...
'' – 12 spp. * '' Aulaxina'' – 4 spp. * '' Aulaxinella'' – 3 spp. * '' Batistomyces'' – 2 spp. * '' Bezerroplaca'' – 5 spp. * '' Calenia'' – 20 spp. * '' Caleniella'' – 2 spp. * '' Caleniopsis'' – 2 spp. * '' Cladosterigma'' – 2 spp. * '' Corticifraga'' – 9 spp. * ''
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 ...
'' – 5 spp. * ''
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' ...
'' – 20 spp. * '' Ferraroa'' – 1 sp. * ''
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 ...
'' – 4 spp. * '' Gyalectidium'' – 40 spp. * ''
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 ...
'' – 30 spp. * '' Gyalideopsis'' – 50 spp. * ''
Hippocrepidea ''Hippocrepidea'' is a genus of fungi in the family Gomphillaceae. This is a monotypic In biology, a monotypic taxon is a taxonomic group (taxon) that contains only one immediately subordinate taxon. A monotypic species is one that does not ...
'' – 1 sp. * '' Jamesiella'' – 4 spp. * '' Lithogyalideopsis'' – 4 spp. * '' Microxyphiomyces'' – 15 spp. * '' Monocalenia'' – 1 sp. * '' Paragyalideopsis'' – 4 spp. * '' Paratricharia'' – 1 spp. * '' Phyllogyalidea'' – 2 spp. * '' Psathyromyces'' – 4 spp. * '' Pseudocalenia'' – 1 sp. * '' Psorotheciopsis'' – 7 spp. * '' Rolueckia'' – 3 spp. * '' Roselviria'' – 2 spp. * '' Santricharia'' – 1 sp. * '' Sipmanidea'' – 2 spp. * '' Spinomyces'' – 9 spp. * '' Sporocybomyces'' – 4 spp. * '' Taitaia'' – 1 spp. * '' Tricharia'' – ca. 30 spp. * '' Verruciplaca'' – 2 spp. * '' Vezdamyces'' – 2 spp.


References

{{Taxonbar, from=Q3110397 Gomphillaceae Lichen families Lecanoromycetes families Taxa described in 1929 Taxa named by Walter Watson (lichenologist)