Diorygma
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''Diorygma'' is a
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
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 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 family has a cosmopolitan distribution, most Graphidaceae species occur in tropic ...
. The genus was circumscribed by Franz Gerhard Eschweiler in 1824. Species of the genus are widely distributed 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 ...
and
subtropical The subtropical zones or subtropics are geographical zone, geographical and Köppen climate classification, climate zones immediately to the Northern Hemisphere, north and Southern Hemisphere, south of the tropics. Geographically part of the Ge ...
regions of the world. These lichens form paint-like crusts on bark and rock that range from chalky white to light green, with elongated, pencil-like slits containing their
spore In biology, a spore is a unit of sexual reproduction, sexual (in fungi) or asexual reproduction that may be adapted for biological dispersal, dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores fo ...
s that may flex and branch across the surface. The genus was established in 1824 for tropical script lichens with large, many-celled
spore In biology, a spore is a unit of sexual reproduction, sexual (in fungi) or asexual reproduction that may be adapted for biological dispersal, dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores fo ...
s, but
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, ...
studies in the 2000s and 2010s expanded it significantly by transferring species from other genera and revealing new diversity.


Taxonomy

The name ''Diorygma'' was coined by Franz Gerhard Eschweiler in 1824 for a group of neotropical script lichens 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 family has a cosmopolitan distribution, most Graphidaceae species occur in tropic ...
that have large, many-celled () spores and a thin, naked
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 ...
.
Elias Fries Elias Magnus Fries (15 August 1794 – 8 February 1878) was a Swedish mycologist and botanist. He is sometimes called the "Linnaeus of Mycology". In his works he described and assigned botanical names to hundreds of fungus and lichen sp ...
accepted the genus in 1825—likening it to what is now '' Arthonia''—but made no additional
combinations In mathematics, a combination is a selection of items from a set that has distinct members, such that the order of selection does not matter (unlike permutations). For example, given three fruits, say an apple, an orange and a pear, there are t ...
. Eschweiler himself broadened the concept in 1833 by adding several species that subsequent work has reassigned to ''
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 ...
'' and '' Platythecium''. Towards the end of the nineteenth century,
Johannes Müller Argoviensis Johann Müller (9 May 1828 – 28 January 1896) was a Swiss botanist who was a specialist in lichens. He published under the name Johannes Müller Argoviensis to distinguish himself from other naturalists with similar names. Biography Müller ...
erected the genus '' Graphina'' for -spored taxa, retaining a handful of ''Diorygma'' species within his
sections Section, Sectioning, or Sectioned may refer to: Arts, entertainment and media * Section (music), a complete, but not independent, musical idea * Section (typography), a subdivision, especially of a chapter, in books and documents ** Section sig ...
''Platygraphina'' and ''Platygrammina'', thereby narrowing the scope of Eschweiler's original
taxon 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 ...
. Interest in the group revived in the twentieth century. Dharani Dhar Awasthi and Mamta Joshi, while revising '' Helminthocarpon'' in 1979, recognised that these lirellate lichens formed a discrete lineage, but instead of resurrecting Eschweiler's name they introduced ''Cyclographina''. In a comprehensive
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 ...
of 2002, Bettina Staiger restored ''Diorygma'' for this assemblage and, on the basis of its rather unusual tissue organisation, suggested that it might fall outside the core Graphidaceae. Subsequent
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, ...
analyses have confirmed that ''Diorygma'' does indeed belong within that family and have expanded its limits to include species with either single-spored, muriform asci or with spores divided only by transverse walls.


Description

Species of ''Diorygma'' form a
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 ...
(tightly adherent, paint-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 ...
that ranges in hue from chalky white or cream to light olive-grey or green. The surface is generally dull and may be uneven, slightly wrinkled, or minutely warted; fine cracks are common. Unlike many other script lichens, the thallus lacks special powdery
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 (
soralia Soredia are common reproductive structures of lichens. Lichens reproduce asexually by employing simple fragmentation and production of soredia and isidia. Soredia are powdery propagules composed of fungal hyphae wrapped around cyanobacteria or g ...
or
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 ...
). A true outer "skin" () is poorly developed or absent, while the interior medulla is white and often riddled with conspicuous crystal deposits. The embedded partner alga is '' Trentepohlia'', a filamentous
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 ( ...
that imparts a faint orange tinge when exposed. The sexual fruit-bodies are —elongate, pencil-like slits that may flex and branch across the thallus. Their margins can be thick and swollen or remain almost flush with the surface, and the exposed is usually brown, sometimes dusted with a whitish or yellowish bloom (). Internally, the lateral wall () may be pale or blackened ("") and often shows fine striations produced by repeated regrowth events. The
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 spore-producing layer—is clear and oil-free but stains bluish-violet in
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 ...
at its upper and outer portions. At its roof lies a tangle of gelatinous, net-like
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 ...
that carry nodular tips and brown or colourless . Club-shaped asci have the so-called "''Graphis''-type" apex with a thickened inner cap, and they bear one to eight colourless
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. These spores are oblong with rounded ends, occasionally spindle-shaped, and show transverse walls (sometimes with additional vertical walls to give a brick-like, pattern); they measure roughly 15–250 × 5–60 μm and may turn blue-violet in iodine.
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 ...
is rare and takes place in tiny
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 ...
embedded in wart-like swellings about 0.1 mm across. These structures release minute, sausage-shaped to rod-shaped conidia 3.5–6 × 1–1.5 μm. Chemically, most species contain compounds of the norstictic or
stictic acid Stictic acid is an aromatic organic compound, a product of secondary metabolism in some species of lichens. Stictic acid is the subject of preliminary biomedical research. Stictic acid has cytotoxic and apoptotic effects ''in vitro ''In v ...
families, while others produce protocetraric-series substances—
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 that can aid in species identification through chemical spot tests and
thin-layer chromatography Thin-layer chromatography (TLC) is a chromatography technique that separates components in non-volatile mixtures. It is performed on a TLC plate made up of a non-reactive solid coated with a thin layer of adsorbent material. This is called the sta ...
.


Species

,
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 ...
accepts 80 species of ''Diorygma''. *'' Diorygma aeolum'' *'' Diorygma africanum'' *'' Diorygma agumbense'' – India *'' Diorygma alagoense'' – Brazil *'' Diorygma albocinerascens'' – India *'' Diorygma albovirescens'' – India *'' Diorygma angusticarpum'' – Thailand *'' Diorygma antillarum'' *'' Diorygma archeri'' – Vietnam *'' Diorygma australasicum'' *'' Diorygma basinigrum'' *'' Diorygma cameroonense'' – Africa *'' Diorygma chumphonense'' – Thailand *'' Diorygma circumfusum'' *'' Diorygma citri'' – Thailand *'' Diorygma confluens'' *'' Diorygma conprotocetraricum'' – Thailand *'' Diorygma dandeliense'' – India *'' Diorygma dealbatum'' – India *'' Diorygma defectoisidiatum'' – Brazil *'' Diorygma epiglaucum'' *'' Diorygma erythrellum'' *'' Diorygma excipuloconvergentum'' – India *'' Diorygma extensum'' *'' Diorygma fuscopruinosum'' – Thailand *'' Diorygma fuscum'' – China *'' Diorygma gyrosum'' – Brazil *'' Diorygma hieroglyphicellum'' – Thailand *'' Diorygma hieroglyphicum'' *'' Diorygma hololeucum'' *'' Diorygma inaequale'' – India *'' Diorygma incantatum'' – Brazil *'' Diorygma inexpectatum'' – Thailand *'' Diorygma intermedium'' *'' Diorygma isabellinum'' *'' Diorygma isidiolichexanthonicum'' – Brazil *'' Diorygma junghuhnii'' *'' Diorygma karnatakense'' – India *'' Diorygma kurnoolense'' – India *'' Diorygma lichexanthonicum'' – Brazil *'' Diorygma longilirellatum'' – India *'' Diorygma longisporum'' – Brazil *'' Diorygma macgregorii'' *'' Diorygma manipurense'' – India *'' Diorygma megaspermum'' – India *'' Diorygma megasporum'' *'' Diorygma megistosporum'' – India *'' Diorygma microsporum'' *'' Diorygma minisporum'' *'' Diorygma monophorum'' *'' Diorygma nigricans'' – Peru *'' Diorygma norsubmuriforme'' – Brazil *'' Diorygma occultum'' *'' Diorygma panchganiense'' – India *'' Diorygma pauciseptatum'' – Brazil *'' Diorygma poitaei'' *'' Diorygma pruinosum'' *'' Diorygma radiatum'' *'' Diorygma reniforme'' *'' Diorygma roseopruinatum'' – New Caledonia *'' Diorygma rufopruinosum'' *'' Diorygma rufosporum'' – India *'' Diorygma rupicola'' – India *'' Diorygma salazinicum'' – Thailand *'' Diorygma salvadoriense'' *'' Diorygma salxanthonicum'' – Brazil *'' Diorygma saxicola'' – India *'' Diorygma sipmanii'' *'' Diorygma soozanum'' *'' Diorygma sophianum'' – Brazil *'' Diorygma spilotum'' *'' Diorygma sticticum'' *'' Diorygma streimannii'' – Australia *'' Diorygma subalbatum'' – India *'' Diorygma subpruinosum'' – Thailand *'' Diorygma talisense'' *'' Diorygma thailandicum'' – Thailand *'' Diorygma tibellii'' *'' Diorygma tocantinsense'' – Brazil *'' Diorygma toensbergianum'' – Brazil *'' Diorygma tuberculosum'' *'' Diorygma upretii'' *'' Diorygma verrucirimosum'' – India *'' Diorygma wallamanensis'' – Australia *'' Diorygma wilsonianum''


References


Cited literature

* {{Taxonbar, from=Q5279571 Graphidales genera Lichen genera Taxa named by Franz Gerhard Eschweiler Taxa described in 1824