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The Candelariales are 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
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
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 include subspecies or smaller, infraspecific taxa. In the case of genera, the term "unisp ...
class Candelariomycetes. It contains the families Candelariaceae and Pycnoraceae. The order was circumscribed by Jolanta Miadlikowska, François Lutzoni, and
Helge Thorsten Lumbsch Helge Thorsten Lumbsch (born 1964) is a German-born lichenology, lichenologist living in the United States. His research interests include the phylogeny, taxonomy (biology), taxonomy, and phylogeography of lichen-forming fungi; lichen diversity; ...
as part of a comprehensive
phylogenetic In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...
classification of the kingdom Fungi published in 2007. The class Candelariomycetes was proposed in 2018 by Hermann Voglmayr and Walter Jaklitsch.


Taxonomy

The yellow "candle-wax" lichens now grouped in Candelariomycetes were long treated as part of the large class Lecanoromycetes. A multigene study of lichen-forming fungi published in 2007 first recognised them as a distinct order, Candelariales – comprising the families Candelariaceae and Pycnoraceae – but still left the group inside Lecanoromycetes. Even earlier, Miądlikowska and co-workers had informally coined the name "Candelariomycetidae" (2006) for this lineage, and Lücking and colleagues later formalised that name at subclass rank (2017). Subsequent DNA analyses that combined ribosomal (nSSU, nLSU, mtSSU) and protein-coding (RPB1, RPB2) genes from virtually every recognised class of Leotiomyceta showed that Candelariales never clusters with the lecanoroid core. Instead, it forms an isolated, well-supported branch whose internal topology places Pycnoraceae as the
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 ...
family to Candelariaceae. In the best-scoring maximum-likelihood tree the class received 88 % bootstrap support (79 % under parsimony) and a posterior probability of 1.0, values that molecular systematists regard as strong evidence. Recognising this consistent phylogenetic independence, Voglmayr and Jaklitsch erected the new class Candelariomycetes in 2018 (published 2019), with Candelariales as its
type Type may refer to: Science and technology Computing * Typing, producing text via a keyboard, typewriter, etc. * Data type, collection of values used for computations. * File type * TYPE (DOS command), a command to display contents of a file. * ...
order and ''Candelaria'' as the nomenclatural
type genus In biological taxonomy, the type genus (''genus typica'') is the genus which defines a biological family and the root of the family name. Zoological nomenclature According to the International Code of Zoological Nomenclature, "The name-bearin ...
. Although different analyses place the class variously near
Eurotiomycetes Eurotiomycetes is a large class of ascomycetes with cleistothecial ascocarps within the subphylum Pezizomycotina, currently containing around 3810 species according to the Catalogue of Life. It is the third largest lichenized class, with more th ...
,
Arthoniomycetes Arthoniomycetes are a class of ascomycete fungi. It includes two orders: Arthoniales and Lichenostigmatales. Most of the taxa in these orders are tropical and subtropical lichens. Systematics Phylogenetic analysis supports the monophyly of thi ...
or
Dothideomycetes Dothideomycetes is the largest and most diverse class of ascomycete fungi. It comprises 11 orders 90 families, 1,300 genera and over 19,000 known species. Wijayawardene et al. in 2020 added more orders to the class. Traditionally, most of it ...
, none recover it within Lecanoromycetes when a broad taxon sample is used; the lineage is therefore now treated as one of the younger, small classes of filamentous Ascomycota.


Description

Members of the Candelariomycetes are readily recognised in the field by their vivid lemon- to orange-yellow thalli. The vegetative body (
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 ...
) ranges from a paper-thin crust tightly appressed to the
substrate Substrate may refer to: Physical layers *Substrate (biology), the natural environment in which an organism lives, or the surface or medium on which an organism grows or is attached ** Substrate (aquatic environment), the earthy material that exi ...
to small, scale-like lobes () that may pile up into tiny, leaf-like rosettes. In a few species the thallus is reduced or absent and the fungus grows directly on other lichens. A green, spherical alga (a photobiont) provides
photosynthetic Photosynthesis ( ) is a Biological system, system of biological processes by which Photoautotrophism, photosynthetic organisms, such as most plants, algae, and cyanobacteria, convert light energy, typically from sunlight, into the chemical ener ...
energy, and unlike some other lichen groups no
cyanobacteria Cyanobacteria ( ) are a group of autotrophic gram-negative bacteria that can obtain biological energy via oxygenic photosynthesis. The name "cyanobacteria" () refers to their bluish green (cyan) color, which forms the basis of cyanobacteri ...
l "cephalodia" are formed. The bright colour comes from pulvinic acid pigments in the Candelariaceae or, in the wood-inhabiting Pycnoraceae, from depsidic compounds such as alectorialic acid. Most species colonise bark or
siliceous rock Siliceous rocks are sedimentary rocks that have silica (SiO2) as the principal constituent. The most common siliceous rock is chert; other types include diatomite. They commonly form from silica-secreting organisms such as radiolarians, diatoms, o ...
in well-lit, nutrient-enriched habitats; a few grow on mosses, soil, or old timber. The
cosmopolitan Cosmopolitan may refer to: Internationalism * World citizen, one who eschews traditional geopolitical divisions derived from national citizenship * Cosmopolitanism, the idea that all of humanity belongs to a single moral community * Cosmopolitan ...
species '' Candelaria concolor''—common on street trees and fence rails—illustrates the group's tolerance of urban conditions.
Sexual reproduction Sexual reproduction is a type of reproduction that involves a complex life cycle in which a gamete ( haploid reproductive cells, such as a sperm or egg cell) with a single set of chromosomes combines with another gamete to produce a zygote tha ...
takes place in
apothecia An ascocarp, or ascoma (: ascomata), is the fruiting body ( sporocarp) of an ascomycete phylum fungus. It consists of very tightly interwoven hyphae and millions of embedded asci, each of which typically contains four to eight ascospores. As ...
, the tiny, often disc-shaped fruiting bodies that dot the thallus surface. These apothecia usually carry a narrow rim of thallus tissue ( margin) in Candelariaceae or a blackened rim without algal cells () in Pycnoraceae. Inside, unbranched
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 ...
(slender sterile filaments) are imbued with starch-like (
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 ...
) material that turns blue 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 ...
-based
staining Staining is a technique used to enhance contrast in samples, generally at the Microscope, microscopic level. Stains and dyes are frequently used in histology (microscopic study of biological tissue (biology), tissues), in cytology (microscopic ...
—a useful diagnostic character. The spore sacs ( asci) are club-shaped, also amyloid, and contain eight to as many as sixty-four colourless, smooth
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 ascospores are
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 lemon-shaped and lack cross-walls or show, at most, one to three indistinct
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 ...
. When sexual spores are not being produced, many species form minute flask-shaped
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 ...
that exude colourless, single-celled
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 ...
.


References

{{Taxonbar, from=Q147328 Candelariales Ascomycota orders Lichen orders Taxa described in 2007 Taxa named by Helge Thorsten Lumbsch