family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Idea ...

fungi A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately from ...

Verrucariales Verrucariales is an order of ascomycetous fungi within the subclass Chaetothyriomycetidae of the class Eurotiomycetes. Although most of the Verrucariales are lichenised, the family Sarcopyreniaceae consists of 11 species of lichenicolous (lichen ...

thallus Thallus (plural: 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. Many of these organisms ...

crustose Crustose is a 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 substrates at all points. ''Crustose'' is found on rocks ...

foliose Foliose lichen is one of the morphological classes of lichens, which are complex organisms that arise from the symbiotic relationship between fungi and a photosynthetic partner, typically algae. This partnership allows lichen to live in diverse ...

squamulose A squamulose lichen is a lichen that is composed of small, often overlapping "scales" called squamules. If they are raised from the substrate and appear leafy, the lichen may appear to be a foliose lichen, but the underside does not have a "ski ...

ascomata An ascocarp, or ascoma (), 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. Ascocarps are mos ...

ascospore An ascus (; ) is the sexual spore-bearing cell produced in ascomycete fungi. Each ascus usually contains eight ascospores (or octad), produced by meiosis followed, in most species, by a mitotic cell division. However, asci in some genera or ...

septa The Southeastern Pennsylvania Transportation Authority (SEPTA) is a regional public transportation authority that operates bus, rapid transit, commuter rail, light rail, and electric trolleybus services for nearly 4 million people in five c ...

lichenicolous A lichenicolous fungus is a parasitic fungus that only lives on lichen as the host. A lichenicolous fungus is not the same as the fungus that is the component of the lichen, which is known as a lichenized fungus. They are most commonly specific to ...

photobiont A lichen ( , ) is a composite organism that arises from algae or cyanobacteria living among filaments of multiple fungi species in a mutualistic relationship.green algae, but also

brown algae Brown algae (singular: alga), comprising the class Phaeophyceae, are a large group of multicellular algae, including many seaweeds located in colder waters within the Northern Hemisphere. Brown algae are the major seaweeds of the temperate and p ...

and yellow-green algae. The family, circumscribed nearly two centuries ago, now includes 56 genera and about one thousand species, and is the third-largest family of lichen-forming fungi. Most

diversity Diversity, diversify, or diverse may refer to: Business *Diversity (business), the inclusion of people of different identities (ethnicity, gender, age) in the workforce *Diversity marketing, marketing communication targeting diverse customers * ...

occurs in

temperate climate In geography, the temperate climates of Earth occur in the middle latitudes (23.5° to 66.5° N/S of Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ranges throughout ...

s of the Northern Hemisphere. Rocks and

soil Soil, also commonly referred to as earth or dirt Dirt is an unclean matter, especially when in contact with a person's clothes, skin, or possessions. In such cases, they are said to become dirty. Common types of dirt include: * Debri ...

are the most common substrates for the Verrucariaceae, with growth on

wood Wood is a porous and fibrous structural tissue found in the stems and roots of trees and other woody plants. It is an organic materiala natural composite of cellulose fibers that are strong in tension and embedded in a matrix of lignin ...

, bark, and leaves less common. Several species are components of

biological soil crust Biological soil crusts are communities of living organisms on the soil surface in arid and semi-arid ecosystems. They are found throughout the world with varying species composition and cover depending on topography, soil characteristics, climate, ...

s and contribute to the formation and stabilisation of soil. Some semi-aquatic lichens occur in this family, including about two dozen species of marine lichens. Traditionally, Verrucariaceae species have been grouped into genera based largely on the

growth form Plant life-form schemes constitute a way of classifying plants alternatively to the ordinary species-genus-family scientific classification. In colloquial speech, plants may be classified as trees, shrubs, herbs (forbs and graminoids), etc. The sc ...

of the thallus, and on the septation of the spores.

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 ...

research conducted in the past couple of decades has helped to clarify the

phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...

framework of the family, but many genera remain poorly investigated.

Systematics

Taxonomic history

A system of classification for the Verrucariaceae was first suggested by German botanist Franz Gerhard Eschweiler in 1824. In his scheme,

taxa In biology, a taxon (back-formation from ''taxonomy''; plural 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 nam ...

were distributed between two

taxonomic rank In biological classification, taxonomic rank is the relative level of a group of organisms (a taxon) in an ancestral or hereditary hierarchy. A common system consists of species, genus, family, order, class, phylum, kingdom, domain. While ...

s intended as names of orders ("

cohors A cohort (from the Latin ''cohors'', plural ''cohortes'', see wikt:cohors for full inflection table) was a standard tactical military unit of a Roman legion. Although the standard size changed with time and situation, it was generally compose ...

"), the Dermatocarpeae and the Verrucariae, based on thallus structure. The Dermatocarpeae contained

squamulose A squamulose lichen is a lichen that is composed of small, often overlapping "scales" called squamules. If they are raised from the substrate and appear leafy, the lichen may appear to be a foliose lichen, but the underside does not have a "ski ...

(scaly) and

foliose Foliose lichen is one of the morphological classes of lichens, which are complex organisms that arise from the symbiotic relationship between fungi and a photosynthetic partner, typically algae. This partnership allows lichen to live in diverse ...

(leafy) species, while the Verrucariae had the crustose species. Although several genera were included in each cohor, most of these are now known to be not closely related and are classified in other families; the only currently applicable genera from Eschweiler's list are '' Dermatocarpon'' and '' Endocarpon'' in the Dermatocarpeae, and '' Verrucaria'' in the Verrucariae. Because Eschweiler published these taxa as "cohors", they do not meet the requirements of valid publication according to nomenclatural rules, and the

authorship An author is the writer of a book, article, play, mostly written work. A broader definition of the word "author" states: "''An author is "the person who originated or gave existence to anything" and whose authorship determines responsibility f ...

of the family cannot be attributed to him. The family was validly published by

Jonathan Carl Zenker Jonathan Carl (or Karl) Zenker (1 March 1799 – 6 November 1837) was a German naturalist born in Remda-Teichel, Sundremda, located in the Saalfeld-Rudolstadt district. Originally a student of theology at the University of Jena, he subsequentl ...

three years later in 1827 (as Verrucariae), with '' Verrucaria'' assigned as the

type genus In biological taxonomy, the type genus 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-bearing type of a nominal ...

. In Zenker's proposed classification, the family was divided into ''Cryolichenes'' (''Verrucariae''), which contained ''Verrucaria'' and other unrelated crustose genera, and ''Phyllolichenes'' (''Endocarpa''), with genus ''Endocarpon'' representing the squamulose and foliose taxa. Nearly a century later, Alexander Zahlbruckner's publication ''Catalogus Lichenum Universalis'' (1922) became an influential work for lichen classification. He divided the order Verrucariales into two families, Dermatocarpaceae and Verrucariaceae, the latter of which was divided into 13 genera, 9 of which remain in the family. Around this time, German lichenologist Georg Hermann Zschacke contributed the first extensive monographic series on the family in a set of publications from 1913 to 1927. His classification scheme was similar to that of Zahlbruckner. Historically, there were three main morphological criteria used to separate genera in the Verrucariaceae: spore septation (the degree of partitioning by septa); the structure of the

thallus Thallus (plural: 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. Many of these organisms ...

; and the presence or absence of algae in the spore-bearing tissue, the hymenium. However, even before the advent of

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 ...

, the use of these have been contentious, as several authors have considered them artificial, and not representative of true

phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...

relationships. As one example, the use of the degree of spore septation as a major character to

circumscribe In geometry, the circumscribed circle or circumcircle of a polygon is a circle that passes through all the vertices of the polygon. The center of this circle is called the circumcenter and its radius is called the circumradius. Not every polyg ...

genera was shown to be problematic when it was demonstrated that in some instances, spore septation is variable with a single species. In 1953, Czech lichenologist Miroslav Servít proposed a new system of classification for the Verrucariaceae, based largely on characteristics of the —the upper, often pigmented, part of the fruiting body covering the . Josef Halda summarised Servít's contribution: "Servít's studies contributed to complicacy of the taxonomy of the whole order Verrucariales. The huge amount of newly described taxa (families, genera, species, varieties or forms) and new combinations almost brought an end to good orientation necessary for new researchers. Any of his studies is not really a revision since it contains only a further list of new taxa and their combinations within the many genera and family framework. Servít neither returned to his newly described taxa nor he revised them in his monographs". Servít's classification was not widely adopted by future authors, and according to Cécile Gueidan and colleagues, it was the lack of clear morphological characteristics in the Verrucariaceae that hindered future proposals for changes in classification. Because of the relatively simple morphology of most of the crustose Verrucariaceae (especially in the type genus ''Verrucaria''), and the fact that this morphology is often variable depending on environmental conditions, identification and delimitation of the species in this family is difficult. Many morphological traits are symplesiomorphic or

homoplastic Homoplasy, in biology and phylogenetics, is the term used to describe a Phenotypic trait, feature that has been gained or lost independently in separate lineages over the course of evolution. This is different from Homology (biology), homology, wh ...

, and are not suitable as distinguishing characters to define genera. Studies have demonstrated several examples of

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 ...

–genetically distinct lichens that are difficult or impossible to distinguish by morphology alone–in the genera '' Hydropunctaria'', '' Sporodictyon'', and ''Verrucaria''. These difficulties are reflected in the number of species of uncertain status described by previous lichenologists: of the 84 Verrucariaceae species described by Zahlbruckner, about 37 are currently accepted as valid species; Zschacke described 133 species, of which about 50 are now accepted; of Servít's 424 described species, only about 60 are accepted.

Molecular phylogenetics

The first molecular phylogenetic studies involving the Verrucariaceae, published between 2001 and 2006, were used to show the higher-level relationships in the

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 tha ...

. This research showed that Verrucariales has a

sister A sister is a woman or a girl who shares one or more parents with another individual; a female sibling. The male counterpart is a brother. Although the term typically refers to a family, familial relationship, it is sometimes used endearingly to r ...

relationship to the

Chaetothyriales The Chaetothyriales are an order of ascomycetous fungi in the class Eurotiomycetes and within the subclass Chaetothyriomycetidae. The order was circumscribed in 1987 by mycologist Margaret Elizabeth Barr-Bigelow. Families and genera , Species ...

, an order of non-lichenised fungi. In 2007 and 2009 publications, Cécile Gueidan and colleagues used molecular data from 83 Verrucariaceae taxa to demonstrate that many of the morphologically-defined genera were

polyphyletic A polyphyletic group is an assemblage of organisms or other evolving elements that is of mixed evolutionary origin. The term is often applied to groups that share similar features known as homoplasies, which are explained as a result of conver ...

—of mixed evolutionary origins. In their analysis, they identified 4 major lineages in the family, including ten monophyletic subgroups. They proposed several taxonomic changes to more closely align the morphology-based classification with the molecular phylogeny, including the new genera ''Parabagliettoa'', ''Hydropunctaria'', and ''Wahlenbergiella'' and several new combinations. Ancestral state reconstruction analysis suggests that the

most recent common ancestor In biology and genetic genealogy, the most recent common ancestor (MRCA), also known as the last common ancestor (LCA) or concestor, of a set of organisms is the most recent individual from which all the organisms of the set are descended. The ...

of the Verrucariaceae was probably crustose, had a weakly differentiated upper , a hymenium free of algae, and ascospores (i.e., without

septa The Southeastern Pennsylvania Transportation Authority (SEPTA) is a regional public transportation authority that operates bus, rapid transit, commuter rail, light rail, and electric trolleybus services for nearly 4 million people in five c ...

). The first lichen-forming fungus to have its genome sequenced was the mycobiont of '' Endocarpon pusillum'', the type species of '' Endocarpon'' and a member of the Verrucariaceae.

Etymology

As is standard practice in botanical nomenclature, the name Verrucariaceae is based on the name of the type genus, ''Verrucaria'', with the ending ''-aceae'' indicating the

rank Rank is the relative position, value, worth, complexity, power, importance, authority, level, etc. of a person or object within a ranking, such as: Level or position in a hierarchical organization * Academic rank * Diplomatic rank * Hierarchy * ...

of family. The genus name is derived from the

Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...

word ''verruca'' (meaning "wart") and the suffix ''-aria'' (meaning "belonging to" or "possession").

Synonymy

Some genera now classified in the Verrucariaceae were considered by past authors to be unique enough to warrant inclusion in their own family. These historical family names are considered synonymous with Verrucariaceae: *Endocarpaceae – type genus: '' Endocarpon'' *Dermatocarpaceae – type genus: '' Dermatocarpon'' *Glomerillaceae – type genus: '' Glomerilla'' *Endopyreniaceae – type genus: ''Endocarpon'' *Mastodiaceae – type genus: '' Mastodia'' *Pyrenothamniaceae – type genus: ''Pyrenothamnia'' , now synonymised with ''Endocarpon'' *Thelidiaceae – type genus: '' Thelidium'' *Bagliettoaceae – type genus: '' Bagliettoa'' *Staurotheleaceae – type genus: '' Staurothele''

Description

Species of Verrucariaceae occur in a wide variety of morphologies, including

crustose Crustose is a 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 substrates at all points. ''Crustose'' is found on rocks ...

,

foliose Foliose lichen is one of the morphological classes of lichens, which are complex organisms that arise from the symbiotic relationship between fungi and a photosynthetic partner, typically algae. This partnership allows lichen to live in diverse ...

, and

squamulose A squamulose lichen is a lichen that is composed of small, often overlapping "scales" called squamules. If they are raised from the substrate and appear leafy, the lichen may appear to be a foliose lichen, but the underside does not have a "ski ...

forms. The size of the lichen body, or

thallus Thallus (plural: 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. Many of these organisms ...

, usually ranges from a few millimetres to about in diameter. Typical thallus colours are greenish, brownish, grey, or black. The

medulla Medulla or Medullary may refer to: Science * Medulla oblongata, a part of the brain stem * Renal medulla, a part of the kidney * Adrenal medulla, a part of the adrenal gland * Medulla of ovary, a stroma in the center of the ovary * Medulla of t ...

is the internal layer of fungal hyphae below the

cortex Cortex or cortical may refer to: Biology * Cortex (anatomy), the outermost layer of an organ ** Cerebral cortex, the outer layer of the vertebrate cerebrum, part of which is the ''forebrain'' *** Motor cortex, the regions of the cerebral cortex i ...

and the algal layer; three types occur in the Verrucariaceae. The type has loosely interlaced hyphae with elongated cells, the type comprises tightly arranged and rounded cells, and the mixed-type medulla has both rounded and elongated cells. The

ascomata An ascocarp, or ascoma (), 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. Ascocarps are mos ...

are in the form of a

perithecium An ascocarp, or ascoma (), 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. Ascocarps are mos ...

, and these structures are often —meaning they have a shield-like stromatic growth, made of both fungal hyphae and host tissue, around the ostiole (opening). In these instances the involucrellum is typically well developed. The is a term for all kinds of hyphae or other tissues between asci (the spore-bearing cells). In the Verrucariaceae, the hamathecium is (disappearing at maturity), and usually hemiamyloid (rarely amyloid). The perithecia have short ; these vertical, filament-like support structures are similar to paraphyses but grow downwards in the perithecial cavity before ascus formation. In two genera, '' Endocarpon'' and '' Staurothele'', algal cells occur in the hamathecium. Asci are usually bitunicate, meaning they have two functional ascal wall layers. The release of spores from the ascus, or dehiscence, has been shown in some species to involve the gelification of the outer wall apex. The

ascospore An ascus (; ) is the sexual spore-bearing cell produced in ascomycete fungi. Each ascus usually contains eight ascospores (or octad), produced by meiosis followed, in most species, by a mitotic cell division. However, asci in some genera or ...

s usually number eight per ascus, are variably

septate In biology, a septum (Latin for ''something that encloses''; plural 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 * Interatr ...

, and often have a gelatinous covering. There have been several publications investigating the anatomy and

development Development or developing may refer to: Arts *Development hell, when a project is stuck in development *Filmmaking, development phase, including finance and budgeting *Development (music), the process thematic material is reshaped * Photograph ...

of the perithecia in Verrucariaceae species. Despite this, it is not clear whether the development of ascomata in the family should be considered ascohymenial (formation originates from special hyphal branches that are usually enveloped by the hyphae that will form the ascocarp) or ascolocular (formation originates from a small stroma, and the asci develop in cavities whose wall consists of stromal tissue). Catapyrenium boccanum 208899

Members of the Verrucariaceae that are squamulose, have simple ascospores (without any septa), and lack algae in the hymenium were historically classified in the genus '' Catapyrenium''. These species were later divided into several genera based on the structure of their

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 inverse ...

; the validity of this generic splitting was confirmed with phylogenetic analyses. The so-called "catapyrenioid" lichens include members of '' Anthracocarpon'', '' Catapyrenium'', '' Heteroplacidium'', '' Involucropyrenium'', '' Neocatapyrenium'', '' Placidiopsis'', '' Placidium'', and '' Scleropyrenium'', and, as of 2010, numbered 81 species. The morphological variety of the thallus in the Verrucariaceae includes some rarely encountered forms. For example, the thallus of '' Flakea papillata'' consists of what has been described as "minute 'lawns' of small leaflets", in which the algae are arranged in a net-like structure of adjacent cells, one cell layer thick. In '' Psoroglaena stigonemoides'', the thallus consists of tiny coral-like branches of algal threads that are covered by a layer of pimpled (''papillate'') fungal hyphae. In the sterile lichen '' Normandina pulchella'', the thallus comprises small bluish-green squamules. These squamules have edges can extend upward to form structures similar in appearance to the

basidiolichen Basidiolichens are lichenized members of the Basidiomycota, a much smaller group of lichens than the far more common ascolichens in the Ascomycota. In arctic, alpine, and temperate forests, the most common basidiolichens are in the agaric genus ...

species '' Lichenomphalia hudsoniana'', or in other instances transform into soralia, somewhat resembling the leprose genus '' Lepraria''.

Photobionts

There are several

green alga The green algae (singular: green alga) are a group consisting of the Prasinodermophyta and its unnamed sister which contains the Chlorophyta and Charophyta/Streptophyta. The land plants (Embryophytes) have emerged deep in the Charophyte alga as ...

l genera that are known to become

photobiont A lichen ( , ) is a composite organism that arises from algae or cyanobacteria living among filaments of multiple fungi species in a mutualistic relationship.mycobionts. The high photobiont diversity observed in this family is rather unusual, as most higher-level taxonomic groups of lichen-forming fungi tend to associate preferentially with only one or two groups of algae. Known green algal associates include members of the class

Trebouxiophyceae The Trebouxiophyceae are a class of green algae, in the division Chlorophyta. Their circumscription within the green algae is not well established due to the need for more genetic studies at higher levels within the group. Genera without interve ...

(genera '' Asterochloris'', '' Auxenochlorella'', ''

Chlorella ''Chlorella'' is a genus of about thirteen species of single-celled green algae belonging to the division Chlorophyta. The cells are spherical in shape, about 2 to 10 μm in diameter, and are without flagella. Their chloroplasts contain the ...

'', '' Chloroidium'', '' Diplosphaeria'', ''

Elliptochloris ''Elliptochloris'' is a genus of green algae in the order Prasiolales Prasiolales is an order of green algae in the class Trebouxiophyceae.See the NCBI The National Center for Biotechnology Information (NCBI) is part of the United St ...

'', ''

Myrmecia Myrmecia can refer to: * ''Myrmecia'' (alga), genus of algae associated with lichens * ''Myrmecia'' (ant), genus of ants called bulldog ants * Myrmecia (skin), a kind of deep wart on the human hands or feet See also * '' Copromorpha myrmecias'' ...

'', '' Pseudostichococcus'', ''

Trebouxia ''Trebouxia'' is a unicellular green alga.Silverside, A. J. (2009). Retrieved from http://www.bioref.lastdragon.org/Chlorophyta/''Trebouxia''.html It is a photosynthetic organism that can exist in almost all habitats found in polar, tropical, a ...

'', and '' Trochiscia''), class Chlorophyceae (genus '' Coccobotrys''), and class

Ulvophyceae The Ulvophyceae or ulvophytes are a class of green algae, distinguished mainly on the basis of ultrastructural morphology, life cycle and molecular phylogenetic data.Graham LE, Graham JM, Wilcox LW (2009) ''Algae''. 2nd Edition. Benjamin Cumming ...

(genus '' Dilabiphilum''). Research continues to reveal new symbiont relationships, such as the newly described (2022) green algal genus and species, '' Rindifilum ramosum'' (family

Ctenocladaceae Ctenocladaceae is a family of green algae in the order Ulvales Ulvales is an order of green alga The green algae (singular: green alga) are a group consisting of the Prasinodermophyta and its unnamed sister which contains the Chlorophyta ...

, order

Ulvales Ulvales is an order of green alga The green algae (singular: green alga) are a group consisting of the Prasinodermophyta and its unnamed sister which contains the Chlorophyta and Charophyta/Streptophyta. The land plants (Embryophytes) have e ...

), which associates with some members of ''Verrucaria''. One study of photobiont diversity in the Verrucariceae suggests that genera with simple crustose thalli (examples include '' Bagliettoa'', '' Hydropunctaria'', and '' Wahlenbergiella'',) use a high diversity and unusual selection of photobionts, whereas the genera with more complex thalli (such as '' Placidium'' and '' Dermatocarpon'') tend to have lower photobiont diversity.

Brown algae Brown algae (singular: alga), comprising the class Phaeophyceae, are a large group of multicellular algae, including many seaweeds located in colder waters within the Northern Hemisphere. Brown algae are the major seaweeds of the temperate and p ...

are far less frequent photobiont partners for the Verrucariaceae. An example is the

intertidal The intertidal zone, also known as the foreshore, is the area above water level at low tide and underwater at high tide (in other words, the area within the tidal range). This area can include several types of habitats with various species ...

marine lichen formed by the fungus '' Wahlenbergiella tavaresiae'' and the brown alga '' Petroderma maculiforme''. The free-living alga produces filaments that grow and branch upwards; when lichenised and surrounded by mycobiont tissue, they grow downwards. The Verrucariaceae is the only mostly lichen-forming family to have photobiont associations with yellow-green algae (class Xanthophyceae, genus '' Heterococcus''). This genus was shown to be a photobiont partner in the thalli of three unrelated species of Verrucariaceae: '' Hydropunctaria rheitrophila'', '' Verrucaria funckii'', and '' V. hydrela''. In 2022, a crustose, seashore-inhabiting Verrucariaceae was reported as a mycobiont partner for the free-living macroscopic genus '' Urospora'', the first time that genus, or even any in the order

Ulotrichales Ulotrichales is an order of green algae in the class Ulvophyceae.See the NCBIbr>webpage on Ulotrichales Data extracted from the Genera unplaced to family (''incertae sedis''): *'' Trichosarcina'' Families * '' Binucleariaceae'' * '' Collin ...

, has been recorded as a photobiont. The nature of the interaction between mycobiont and photobiont sometimes differs from the typical lichen arrangement in which the photobiont is embedded in a fungal tissue created by the mycobiont. In the Verrucariaceae, some lichens have thalloid algae, where the mycobiont grows within the independently formed algal thallus. Examples include the brown algae ''

Pelvetia ''Pelvetia canaliculata'', the channelled wrack, is a very common brown alga (Phaeophyceae) found on the rocks of the upper shores of Europe. It is the only species remaining in the monotypic genus ''Pelvetia''. In 1999, the other members of this ...

'' and ''

Ascophyllum ''Ascophyllum nodosum'' is a large, common cold water seaweed or brown alga (Phaeophyceae) in the family Fucaceae, being the only species in the genus ''Ascophyllum''. It is a seaweed that only grows in the northern Atlantic Ocean, also known in ...

'', and the green alga '' Prasiola crispa''. This last species is the photobiont partner of '' Mastodia tessellata'', the type species of genus ''Mastodia''; this is the only known lichen symbiosis involving a foliose green alga. Because the fungus grows independently as

mycelium Mycelium (plural mycelia) is a root-like structure of a fungus consisting of a mass of branching, thread-like hyphae. Fungal colonies composed of mycelium are found in and on soil and many other substrates. A typical single spore germinates ...

in close contact with the algae, the interaction between ''Ascophyllum'' and verrucariacean fungi is not technically considered a lichen symbiosis. This type of biological partnership has been referred to as an "intimate, mutualistic association", or a mycophycobiosis; more generally, associated species with this type of loose symbiosis have been called "borderline lichens".

Habitat, distribution, and ecology

Collectively, Verrucariaceae has a cosmopolitan distribution, although the majority of its species occur in

temperate climate In geography, the temperate climates of Earth occur in the middle latitudes (23.5° to 66.5° N/S of Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ranges throughout ...

s. Rocks and

soil Soil, also commonly referred to as earth or dirt Dirt is an unclean matter, especially when in contact with a person's clothes, skin, or possessions. In such cases, they are said to become dirty. Common types of dirt include: * Debri ...

are the most common substrates, with growth on

wood Wood is a porous and fibrous structural tissue found in the stems and roots of trees and other woody plants. It is an organic materiala natural composite of cellulose fibers that are strong in tension and embedded in a matrix of lignin ...

and bark less common. Among the rock-dwelling species, there are both endoliths (those that grow on the surface) and epiliths (those that grow "within" the rocks, i.e., under and around the rock crystals).

Calcareous Calcareous () is an adjective meaning "mostly or partly composed of calcium carbonate", in other words, containing lime or being chalky. The term is used in a wide variety of scientific disciplines. In zoology ''Calcareous'' is used as an ad ...

rocks are the most common substrate for the rock-dwellers, but some, especially those in aquatic or semi-aquatic habitats, grow on

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 ...

s. Some species grow on mosses, and some grow on leaves, including all species of genus '' Phylloblastia''. Several Verrucariaceae species are components of

biological soil crust Biological soil crusts are communities of living organisms on the soil surface in arid and semi-arid ecosystems. They are found throughout the world with varying species composition and cover depending on topography, soil characteristics, climate, ...

s and contribute to the formation and stabilisation of soil, particularly in somewhat

arid A region is arid when it severely lacks available water, to the extent of hindering or preventing the growth and development of plant and animal life. Regions with arid climates tend to lack vegetation and are called xeric or desertic. Most ...

areas; examples include members of '' Endocarpon'' and '' Catapyrenium''. Although most species are terrestrial, there are some semi-aquatic species, with representatives from both fresh and salt water. As of 2015, there were 26 known marine Verrucariaceae, distributed amongst the genera '' Hydropunctaria'' (6 species), '' Mastodia'' (1 sp.), '' Verrucaria'' (16 spp.), and '' Wahlenbergiella'' (3 spp.). Several of these marine lichens have been investigated preliminarily to determine their suitability for use as

bioindicator A bioindicator is any species (an indicator species) or group of species whose function, population, or status can reveal the qualitative status of the environment. The most common indicator species are animals. For example, copepods and other sma ...

s of coastal

water pollution Water pollution (or aquatic pollution) is the contamination of water bodies, usually as a result of human activities, so that it negatively affects its uses. Water bodies include lakes, rivers, oceans, aquifers, reservoirs and groundwater. ...

. Several Verrucariaceae genera consist entirely of

lichenicolous A lichenicolous fungus is a parasitic fungus that only lives on lichen as the host. A lichenicolous fungus is not the same as the fungus that is the component of the lichen, which is known as a lichenized fungus. They are most commonly specific to ...

(lichen-dwelling) fungi: '' Bellemerella'', '' Clauzadella'', '' Haleomyces'', '' Halospora'', '' Norrlinia'', '' Phaeospora'', '' Telogalla''. Lichenicolous fungi occur in other genera to a lesser extent. Some genera, including '' Heteroplacidium'', '' Placidiopsis'', '' Placocarpus'', '' Placopyrenium'', '' Thelidium'', '' Verrucaria'', and '' Verruculopsis'' contain lichenicolous lichens. Crustose Verrucariaceae species are often under-represented in local or national checklists (i.e., lists of all species in a certain area) due to their typical habitat on rocks and their inconspicuousness. For example, with the exception of Japan (77 species on a 2018 checklist), the number of Verrucariaceae species in East Asia is poorly known.

Uses

There are no species in the Verrucariaceae that have any major economic significance. The only lichen in the family that has been recorded as being used in

traditional medicine Traditional medicine (also known as indigenous medicine or folk medicine) comprises medical aspects of traditional knowledge that developed over generations within the folk beliefs of various societies, including indigenous peoples, before the ...

is '' Dermatocarpon miniatum''. In

Traditional Chinese medicine Traditional Chinese medicine (TCM) is an alternative medical practice drawn from traditional medicine in China. It has been described as "fraught with pseudoscience", with the majority of its treatments having no logical mechanism of acti ...

, the lichen, known as "white stone ear" (, pí guǒ yī), is used in treatments for high blood pressure, as a

diuretic A diuretic () is any substance that promotes diuresis, the increased production of urine. This includes forced diuresis. A diuretic tablet is sometimes colloquially called a water tablet. There are several categories of diuretics. All diuretics i ...

, for expelling

parasite Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson has ...

s, for malnutrition in children, for

dysentery Dysentery (UK pronunciation: , US: ), historically known as the bloody flux, is a type of gastroenteritis that results in bloody diarrhea. Other symptoms may include fever, abdominal pain, and a feeling of incomplete defecation. Complications ...

, to improve digestion, and for

abdominal distension Abdominal distension occurs when substances, such as air (gas) or fluid, accumulate in the abdomen causing its expansion. It is typically a symptom of an underlying disease or dysfunction in the body, rather than an illness in its own right. Pe ...

.

Genera

These are the genera that are in the Verrucariaceae (including estimated number of species in each genus, totalling 1017 species), according to a 2021 review of fungal classification. Following the genus 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 ...

(those who first circumscribed the genus; standardised author abbreviations are used), year of publication, and the estimated number of species. Other recent estimates of the number of Verrucariaceae taxa include: 46 genera and 931 species (2008), 48–54 genera and 850–870 species (2016), and 43 genera and 943 species (2017). The

GBIF The Global Biodiversity Information Facility (GBIF) is an international organisation that focuses on making scientific data on biodiversity available via the Internet using web services. The data are provided by many institutions from around the ...

accept over 60+ genera and nearly 1 thousand species. ,

Species Fungorum ''Index Fungorum'' is an international project to index all formal names (scientific names) in the fungus kingdom. the project is based at the Royal Botanic Gardens, Kew, one of three partners along with Landcare Research and the Institute of M ...

(in the

Catalogue of Life The Catalogue of Life 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 Taxonomic I ...

), accepts 56 genera and 575 species (including 8

varieties Variety may refer to: Arts and entertainment Entertainment formats * Variety (radio) * Variety show, in theater and television Films * ''Variety'' (1925 film), a German silent film directed by Ewald Andre Dupont * ''Variety'' (1935 film), ...

and 6

forms Form is the shape, visual appearance, or configuration of an object. In a wider sense, the form is the way something happens. Form also refers to: *Form (document), a document (printed or electronic) with spaces in which to write or enter data * ...

) in the Verrucariaceae. The largest of these is the type genus, ''Verrucaria'', at 146 accepted species. Verrucariaceae is the third-largest family of lichen-forming fungi, following 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'' ( ...

and the Graphidaceae. *'' Agonimia'' – ca. 20 spp. *'' Anthracocarpon'' – 1 sp. *'' Atla'' – 10 spp. *'' Awasthiella'' – 1 sp. *'' Bagliettoa'' – 17 spp. *'' Bellemerella'' – 4 spp. *'' Catapyrenium'' – 6 spp. *'' Clauzadella'' – 1 sp. *'' Clavascidium'' – 9 spp. *'' Dermatocarpon'' – 20 spp. *''Diederimyces'' – 2 spp. *'' Endocarpon'' – ca. 75 spp. *''Flakea'' – 1 sp. *'' Glomerilla'' – 1 sp. *'' Haleomyces'' – 1 sp. *'' Halospora'' – 4 spp. *''Henrica'' – 4 spp. *''Heterocarpon'' – 1 sp. *'' Heteroplacidium'' – 12 spp. *'' Hydropunctaria'' – 8 spp. *'' Involucropyrenium'' – 9 spp. *'' Mastodia'' – 5 spp. *''Moriola'' – ca. 15 spp. *''Muellerella'' – 9 spp. *'' Neocatapyrenium'' – 5 spp. *''Nesothele'' – 5 spp. *''Normandina'' – 3 spp. *'' Norrlinia'' – 2 spp. *''Parabagliettoa'' – 3 spp. *'' Phaeospora'' – 14 spp. *'' Phylloblastia'' – 12 spp. *'' Placidiopsis'' – 20 spp. *'' Placidium'' – 28 spp. *'' Placocarpus'' – 5 spp. *'' Placopyrenium'' – 22 spp. *''Placothelium'' – 1 sp. *''Plurisperma'' – 1 sp. *''Polyblastia'' – ca. 40 + ca. 50 orphaned *''Psoroglaena'' – 17 spp. *''Rhabdopsora'' – 2 spp. *'' Scleropyrenium'' – 2 spp. *''Servitia'' – 1 sp. *''Spheconisca'' – ca. 20 spp. *'' Sporodictyon'' – 5 spp. *'' Staurothele'' – ca. 40 spp. *'' Telogalla'' – 2 spp. *''Thelidiopsis'' – 4 spp. *'' Thelidium'' – ca. 50 + ca. 50 orphaned *''Trimmatothele'' – 3 spp. *''Turgidosculum'' – 1 sp. *'' Verrucaria'' – List of Verrucaria species, ca. 300 spp. *''Verrucula'' – 22 spp. *'' Verruculopsis'' – ca. 10 spp. *'' Wahlenbergiella'' – 3 spp. *''Willeya'' – 12 spp.

Notes

References

Cited literature

* {{Taxonbar, from=Q3123598 Verrucariales Ascomycota families Taxa named by Jonathan Carl Zenker Taxa described in 1827 Lichen families