View source for Biology:Verrucariaceae

{{Short description|Family of mostly lichenised fungi}}
{{Automatic taxobox
| image = Verrucaria nigrescens 080408.jpg
| image_caption = ''[[Biology:Verrucaria nigrescens|Verrucaria nigrescens]]''
| taxon = Verrucariaceae
| authority = Zenker (1827)
| type_genus = ''[[Biology:Verrucaria|Verrucaria]]''
| type_genus_authority = Shrad. (1794)
| subdivision_ranks = Genera
| subdivision = See text
| synonyms_ref = {{sfn|Cannon|Kirk|2007|pages=438–456}}<ref name="Pérez-Ortega et al. 2010"/>
| synonyms = {{collapsible list|bullets=on
|Bagliettoaceae {{small|Servít (1954)}}
|Dermatocarpaceae {{small|Stizenb. (1862)}}
|Endocarpaceae {{small|[[Biology:Antoine Laurent Apollinaire Fée|Fée]] ex Zenker (1828)}}
|Endopyreniaceae {{small|Zahlbr. (1898)}}
|Glomerillaceae {{small|Norman (1868)}}
|Mastodiaceae {{small|Zahlbr. (1908)}}
|Pyrenothamniaceae {{small|Zahlbr. (1898)}}
|Staurotheleaceae {{small|Servít (1954)}}
|Thelidiaceae {{small|Walt.Watson (1929)}}
}}
}}

'''Verrucariaceae''' is a family of [[Biology:Lichen|lichen]]s and a few non-lichenised [[Biology:Fungi|fungi]] in the order [[Biology:Verrucariales|Verrucariales]]. The lichens have a wide variety of [[Biology:Thallus|thallus]] forms, from crustose (crust-like) to foliose (bushy) and squamulose (scaly). Most of them grow on land, some in freshwater and a few in the sea. Many are free-living but there are some species that are [[Earth:Parasitism|parasites]] on other lichens, while one marine species always lives together with a leafy [[Biology:Green algae|green alga]].

Several characteristics of the [[Biology:Spore|spore]]-bearing structures, the ascomata, define the family, including their [[Biology:Ascocarp#Perithecia|perithecioid]] form–more or less spherical or flask-shaped, with a single opening and otherwise completely enclosed by a wall. Squamulose members of the Verrucariaceae with simple [[Biology:Ascospore|ascospore]]s (lacking partitions called [[Biology:Septum|septa]]), and without [[Biology:Algae|algae]] in the [[Biology:Hymenium|spore-bearing region]] are known as {{lichengloss|catapyrenioid}} lichens; there are more than 80 of these species. The family has several dozen lichenicolous (lichen-dwelling) examples, including a few genera that contain solely lichenicolous members. An unusually diverse variety of photobiont partners have been recorded, mostly [[Biology:Green algae|green algae]], but also [[Biology:Brown algae|brown algae]] and [[Biology:Yellow-green algae|yellow-green algae]].

The family, [[Philosophy:Circumscription (taxonomy)|circumscribed]] nearly two centuries ago, now includes 56 [[Biology:Genus|genera]] and about one thousand species, and is the third-largest family of lichen-forming fungi. Most [[Biology:Biodiversity|diversity]] occurs in [[Earth:Temperate climate|temperate climate]]s of the Northern Hemisphere. Rocks and soil are the most common [[Biology:Substrate|substrates]] for the Verrucariaceae, with growth on wood, bark, and leaves less common. Several species are components of [[Biology:Biological soil crust|biological soil crust]]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 of the thallus, and on the [[Biology:Septum|septation]] of the spores. [[Biology:Molecular phylogenetics|Molecular phylogenetics]] research conducted in the past couple of decades has helped to clarify the phylogenetic framework of the family, but many genera remain poorly investigated.

==Systematics==

===Taxonomic history===
{{Multiple image
| align = right
| direction = vertical
| width = 250
| image1 = Dermatocarpon americanum 2929567.jpg
| caption1 = ''[[Biology:Dermatocarpon americanum|Dermatocarpon americanum]]'' is a foliose member of the Verrucariaceae ...
| image2 = Endocarpon pulvinatum (43009591755).jpg
| caption2= ... while ''[[Biology:Staurothele pulvinata|Staurothele pulvinata]]'' is squamulose.
}}

A system of classification for the Verrucariaceae was first suggested by German botanist Franz Gerhard Eschweiler in 1824. In his scheme, [[Biology:Taxon|taxa]] were distributed between two [[Biology:Taxonomic rank|taxonomic rank]]s intended as names of orders ("[[Biology:Taxonomic rank#Ranks in botany|cohors]]"), the Dermatocarpeae and the Verrucariae, based on thallus structure. The Dermatocarpeae contained squamulose (scaly) and foliose (leafy) species, while the Verrucariae had the crustose species. Although several genera were included in each cohor,<ref name="Eschweiler 1824"/> 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 ''[[Biology:Dermatocarpon|Dermatocarpon]]'' and ''[[Biology:Endocarpon|Endocarpon]]'' in the Dermatocarpeae, and ''[[Biology:Verrucaria|Verrucaria]]'' in the Verrucariae. Because Eschweiler published these taxa as "cohors", they do not meet the requirements of [[Biology:Validly published name|valid publication]] according to nomenclatural rules, and the [[Biology:Author citation (botany)|authorship]] of the family cannot be attributed to him.<ref name="Mycobank: Verrucariaceae"/> The family was validly published by Jonathan Carl Zenker three years later in 1827 (as Verrucariae), with ''[[Biology:Verrucaria|Verrucaria]]'' assigned as the [[Biology:Type genus|type genus]].<ref name="Goebel & Kunze 1827"/> 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.<ref name="Gueidan et al. 2007"/>

Nearly a century later, Alexander Zahlbruckner's publication ''Catalogus Lichenum Universalis'' (1922)<ref name="Zahlbruckner 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.<ref name="Gueidan et al. 2007"/> Around this time, German lichenologist Georg Hermann Zschacke contributed the first extensive [[Monograph|monograph]]ic series on the family in a set of publications from 1913 to 1927.<ref name="Zschacke refbundle"/> His classification scheme was similar to that of Zahlbruckner.<ref name="Gueidan et al. 2007"/>

Historically, there were three main [[Biology:Morphology|morphological]] criteria used to separate genera in the Verrucariaceae: spore [[Biology:Septum|septation]] (the degree of partitioning by septa); the structure of the [[Biology:Thallus|thallus]]; and the presence or absence of [[Biology:Algae|algae]] in the spore-bearing tissue, the [[Biology:Hymenium|hymenium]]. However, even before the advent of [[Biology:Molecular phylogenetics|molecular phylogenetics]], the use of these {{lichengloss|characters}} have been contentious, as several authors have considered them artificial, and not representative of true phylogenetic relationships.<ref name="Gueidan et al. 2009"/> As one example, the use of the degree of spore septation as a major character to [[Philosophy:Circumscription (taxonomy)|circumscribe]] genera was shown to be problematic when it was demonstrated that in some instances, spore septation is variable with a single species.<ref name="Orange 1991"/> In 1953, Czech lichenologist Miroslav Servít proposed a new system of classification for the Verrucariaceae, based largely on characteristics of the {{lichengloss|involucrellum}}—the upper, often pigmented, part of the fruiting body covering the {{lichengloss|perithecia}}.<ref name="Servít 1953"/> 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".<ref name="Halda 2003"/> 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.<ref name="Gueidan et al. 2009"/>

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.<ref name="Orange & Chhetri 2022"/> Many morphological [[Biology:Phenotypic trait|traits]] are symplesiomorphic or [[Biology:Homoplasy|homoplastic]], and are not suitable as distinguishing characters to define genera.<ref name="Prieto et al. 2012"/> Studies have demonstrated several examples of cryptic species–genetically distinct lichens that are difficult or impossible to distinguish by morphology alone–in the genera ''[[Biology:Hydropunctaria|Hydropunctaria]]'',<ref name="Orange 2012"/> ''[[Biology:Sporodictyon|Sporodictyon]]'',<ref name="Savić & Tibell 2009"/> and ''Verrucaria''.<ref name="Thüs et al. 2015"/><ref name="Orange 2020"/> 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.<ref name="Orange & Chhetri 2022"/>

===Molecular phylogenetics===
{{cladogram|title=
|caption=[[Biology:Cladogram|Cladogram]] (highly simplified from the original) showing the [[Biology:Phylogeny|phylogeny]] of the main genera in family Verrucariaceae.<ref name="Gueidan et al. 2009"/>
|cladogram={{clade| style=font-size:85%;line-height:75%;width:240px; 
    |1={{clade
       |1={{clade
          |1=''[[Biology:Placocarpus|Placocarpus]]''
          |2=''[[Biology:Verrucula|Verrucula]]''
          }}
       |2={{clade
          |1=''[[Biology:Wahlenbergiella|Wahlenbergiella]]''
          |2={{clade
             |1={{clade
                |1={{clade
                   |1={{clade
                      |1=''[[Biology:Bagliettoa|Bagliettoa]]''
                      |2=''[[Biology:Parabagliettoa|Parabagliettoa]]''
                      }}
                   |2={{clade
                      |1={{clade
                         |1={{clade
                            |1={{clade
                               |1=''[[Biology:Placidium|Placidium]]''
                               |2=''[[Biology:Heteroplacidium|Heteroplacidium]]''
                               }}
                            |2=''[[Biology:Dermatocarpon|Dermatocarpon]]''
                            }}
                         |2={{clade
                            |1=''[[Biology:Endocarpon|Endocarpon]]'', ''[[Biology:Neocatapyrenium|Neocatapyrenium]]''
                            |2={{clade
                               |1={{clade
                                  |1={{clade
                                     |1=''[[Biology:Polyblastia|Polyblastia]]''
                                     |2=''[[Biology:Thelidium|Thelidium]]''
                                     }}
                                  |2={{clade
                                     |1={{clade
                                        |1=''[[Biology:Atla (lichen)|Atla]]''
                                        |2=''[[Biology:Sporodictyon|Sporodictyon]]''
                                        }}
                                     |2={{clade
                                        |1=''[[Biology:Henrica|Henrica]]''
                                        |2=''[[Biology:Verrucaria|Verrucaria]]''
                                        }}
                                     }}
                                  }}
                               |2=''[[Biology:Trimmatothele|Trimmatothele]]''
                               }}
                            }}
                         }}
                      |2={{clade
                         |1=''[[Biology:Placopyrenium|Placopyrenium]]''
                         |2=''[[Biology:Verruculopsis|Verruculopsis]]''
                         }}
                      }}
                   }}
                |2=''[[Biology:Hydropunctaria|Hydropunctaria]]''
                }}
             |2={{clade
                |1={{clade
                   |1={{clade
                      |1=''[[Biology:Catapyrenium|Catapyrenium]]''
                      |2=''[[Biology:Placidiopsis|Placidiopsis]]''
                      }}
                   |2=''[[Biology:Staurothele|Staurothele]]''
                   }}
                |2=''Staurothele'' group
                }}
             }}
          }}
       }}
}}
}}

The first molecular phylogenetic studies involving the Verrucariaceae, published between 2001 and 2006, were used to show the higher-level relationships in the [[Biology:Eurotiomycetes|Eurotiomycetes]]. This research showed that Verrucariales has a [[Biology:Sister group|sister]] relationship to the [[Biology:Chaetothyriales|Chaetothyriales]], an [[Biology:Order|order]] of non-lichenised fungi.<ref name="Lutzoni et al. 2001"/><ref name="Lumbsch et al. 2002"/><ref name="Lumbsch et al. 2004"/><ref name="Lutzoni et al. 2004"/><ref name="Lumbsch et al. 2005"/><ref name="Geiser et al. 2006"/><ref name="Spatafora et al. 2006"/> 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—of mixed evolutionary origins. In their analysis, they identified 4 major [[Biology:Lineage (evolution)|lineages]] in the family, including ten [[Biology:Monophyletic|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 [[Biology:Combinatio nova|new combinations]]. [[Biology:Ancestral reconstruction|Ancestral state reconstruction]] analysis suggests that the [[Biology:Most recent common ancestor|most recent common ancestor]] of the Verrucariaceae was probably crustose, had a weakly differentiated upper {{lichengloss|cortex}}, a [[Biology:Hymenium|hymenium]] free of algae, and {{lichengloss|simple}} ascospores (i.e., without [[Biology:Septum|septa]]).<ref name="Gueidan et al. 2007"/><ref name="Gueidan et al. 2009"/> The first lichen-forming fungus to have its [[Biology:Genome#Sequencing and mapping|genome sequenced]] was the mycobiont of ''Endocarpon pusillum'', the type species of ''[[Biology:Endocarpon|Endocarpon]]'' and a member of the Verrucariaceae.<ref name="Wang et al. 2014"/>

===Etymology===
As is standard practice in [[Biology:Botanical nomenclature|botanical nomenclature]],<ref name="Hawksworth 1974"/> the name Verrucariaceae is based on the name of the type genus, ''Verrucaria'', with the ending ''-aceae'' indicating the [[Biology:Taxonomic rank|rank]] of family. The genus name is derived from the [[Social:Latin|Latin]] word ''verruca'' (meaning "wart") and the suffix ''-aria'' (meaning "belonging to" or "possession").<ref name="Ulloa & Aguirre-Acosta 2020"/>

===Synonymy===
Some genera now classified in the Verrucariaceae were considered by past authors to be distinctive enough to warrant inclusion in their own family. These historical family names are considered [[Biology:Synonym (taxonomy)|synonymous]] with Verrucariaceae:{{sfn|Cannon|Kirk|2007|pages=438–456}}<ref name="Pérez-Ortega et al. 2010"/>
*Endocarpaceae {{small|Fée ex Zenker (1828)}} – type genus: ''[[Biology:Endocarpon|Endocarpon]]''
*Dermatocarpaceae {{small|Stizenb. (1862)}}<ref name="Stizenberger 1862"/> – type genus: ''[[Biology:Dermatocarpon|Dermatocarpon]]''
*Glomerillaceae {{small|Norman (1868)}}<ref name="Norman 1868"/> – type genus: ''[[Biology:Glomerilla|Glomerilla]]''
*Endopyreniaceae {{small|Zahlbr. (1898)}}<ref name="Engler 1898"/> – type genus: ''Endocarpon''<ref name="IF: Endopyreniaceae"/>
*Mastodiaceae {{small|Zahlbr. (1908)}}<ref name="Zahlbruckner 1907"/> – type genus: ''[[Biology:Mastodia|Mastodia]]''
*Pyrenothamniaceae {{small|Zahlbr. (1898)}}<ref name="Engler 1898"/> – type genus: ''Pyrenothamnia'' {{small|Tuck. (1883)}}, now synonymised with ''Endocarpon''<ref name="IF: Pyrenothamnia"/>
*Thelidiaceae {{small|Walt.Watson (1929)}}<ref name="Watson 1929"/> – type genus: ''[[Biology:Thelidium|Thelidium]]''
*Bagliettoaceae {{small|Servít (1954)}}<ref name="Servít 1954"/> – type genus: ''[[Biology:Bagliettoa|Bagliettoa]]''
*Staurotheleaceae {{small|Servít (1954)}}<ref name="Servít 1954"/> – type genus: ''[[Biology:Staurothele|Staurothele]]''

==Description==
{{multiple image
 | align = left
 | width = 200
 | image1 = Normandina pulchella 157856252.jpg
 | caption1 = 
 | image2 = Flakea papillata 122726.jpg
 | alt2 = A trout
 | caption2 =
 | footer = ''Normandina pulchella'' (left) and ''[[Biology:Flakea papillata|Flakea papillata]]'' (right) are two Verrucariaceae species with unusual thallus morphologies.
}}
Species of Verrucariaceae occur in a wide variety of [[Biology:Morphology|morphologies]], including crustose, foliose, and squamulose forms.{{sfn|Cannon|Kirk|2007|pages=373–374}} The size of the lichen body, or [[Biology:Thallus|thallus]], usually ranges from a few millimetres to about {{convert|10|cm|in|abbr=on|0}} in diameter.<ref name="Gueidan et al. 2007"/> Typical thallus colours are greenish, brownish, grey, or black.{{sfn|Cannon|Kirk|2007|pages=373–374}} The [[Biology:Medulla (lichenology)|medulla]] is the internal layer of fungal [[Biology:Hypha|hypha]]e below the [[Biology:Cortex (botany)|cortex]] and the algal layer; three types occur in the Verrucariaceae. The {{lichengloss|prosoplectenchyma|prosoplectenchymatous}} type has loosely interlaced hyphae with elongated cells, the {{lichengloss|paraplectenchyma|paraplectenchymatous}} type comprises tightly arranged and rounded cells, and the mixed-type medulla has both rounded and elongated cells.<ref name="Zhang et al. 2022"/>

The ascomata are in the form of a [[Biology:Ascocarp#Perithecium|perithecium]], and these structures are often {{gli|clypeate}}—meaning they have a shield-like [[Biology:Stroma (tissue)|stromatic]] growth, made of both fungal hyphae and [[Biology:Host|host]] tissue, around the [[Biology:Ostiole|ostiole]] (opening). In these instances the involucrellum is typically well developed. The {{lichengloss|hamathecium}} is a term for all kinds of hyphae or other tissues between [[Biology:Ascus|asci]] (the spore-bearing cells). In the Verrucariaceae, the hamathecium is {{lichengloss|evanescent}} (disappearing at maturity), and usually [[Biology:Amyloid (mycology)|hemiamyloid]] (rarely amyloid).<ref name="Frey 2016"/> The perithecia have short {{lichengloss|pseudoparaphyses}};<ref name="Frey 2016"/> these vertical, filament-like support structures are similar to [[Biology:Paraphyses|paraphyses]] but grow downwards in the perithecial cavity before ascus formation.<ref name="Ulloa 2012"/> In two genera, ''[[Biology:Endocarpon|Endocarpon]]'' and ''[[Biology:Staurothele|Staurothele]]'', algal cells occur in the hamathecium.<ref name="Frey 2016"/> Asci are usually bitunicate, meaning they have two functional ascal wall layers.{{sfn|Cannon|Kirk|2007|pages=373–374}} The release of spores from the ascus, or [[Biology:Dehiscence (botany)|dehiscence]], has been shown in some species to involve the gelification of the outer wall apex.<ref name="Grube 1999"/> The [[Biology:Ascospore|ascospore]]s usually number eight per [[Biology:Ascus|ascus]], are variably [[Biology:Septum|septate]], and often have a gelatinous covering.{{sfn|Cannon|Kirk|2007|pages=373–374}}

There have been several publications investigating the anatomy and [[Biology:Ontogeny|development]] of the perithecia in Verrucariaceae species.<ref name="Doppelbauer 1959"/><ref name="Janex-Favre 1970"/><ref name="Janex-Favre 1975"/><ref name="Wagner 1987"/> 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).<ref name="Gueidan et al. 2007"/>
[[File:Catapyrenium boccanum 208899.jpg|thumb|''[[Biology:Catapyrenium boccanum|Catapyrenium boccanum]]'', shown here growing in a hole in a calcareous rock, is a catapyrenioid lichen.]]
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 ''[[Biology:Catapyrenium|Catapyrenium]]''. These species were later divided into several genera based on the structure of their pycnidia; the validity of this generic splitting was confirmed with phylogenetic analyses. The so-called "catapyrenioid" lichens include members of ''[[Biology:Anthracocarpon|Anthracocarpon]]'', ''[[Biology:Catapyrenium|Catapyrenium]]'', ''[[Biology:Heteroplacidium|Heteroplacidium]]'', ''[[Biology:Involucropyrenium|Involucropyrenium]]'', ''[[Biology:Neocatapyrenium|Neocatapyrenium]]'', ''[[Biology:Placidiopsis|Placidiopsis]]'', ''[[Biology:Placidium|Placidium]]'', and ''[[Biology:Scleropyrenium|Scleropyrenium]]'', and, as of 2010, numbered 81 species.<ref name="Breuss 2010"/>

The [[Biology:Morphology|morphological]] variety of the thallus in the Verrucariaceae includes some rarely encountered forms. For example, the thallus of ''[[Biology:Flakea papillata|Flakea papillata]]'' consists of what has been described as "minute 'lawns' of small leaflets",<ref name="Muggia et al. 2010"/> in which the algae are arranged in a net-like structure of adjacent cells, one cell layer thick. In ''[[Biology:Psoroglaena stigonemoides|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 [[Biology:Basidiolichen|basidiolichen]] species ''[[Biology:Lichenomphalia hudsoniana|Lichenomphalia hudsoniana]]'', or in other instances transform into soralia, somewhat resembling the leprose genus ''[[Biology:Lepraria|Lepraria]]''.<ref name="Muggia et al. 2010"/>

==Photobionts==

[[File:Mastodia y Papúa.jpg|thumb|left|upright=0.7|The marine species ''Mastodia tessellata'' (dark colour) is the only lichen known to form a symbiotic association with a foliose green alga.]]

There are several green algal genera that are known to become photobiont partners with Verrucariaceae 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.<ref name="Thüs et al. 2011"/><ref name="Sanders & Masumoto 2021"/> Known green algal associates include members of the class [[Biology:Trebouxiophyceae|Trebouxiophyceae]] (genera ''Asterochloris'', ''[[Biology:Auxenochlorella|Auxenochlorella]]'', ''[[Biology:Chlorella|Chlorella]]'', ''Chloroidium'', ''Diplosphaeria'', ''[[Biology:Elliptochloris|Elliptochloris]]'', ''[[Biology:Myrmecia (alga)|Myrmecia]]'', ''Pseudostichococcus'', ''[[Biology:Trebouxia|Trebouxia]]'', and ''[[Biology:Trochiscia|Trochiscia]]''), class [[Biology:Chlorophyceae|Chlorophyceae]] (genus ''[[Biology:Coccobotrys (alga)|Coccobotrys]]''), and class [[Biology:Ulvophyceae|Ulvophyceae]] (genus ''Dilabiphilum'').<ref name="Thüs et al. 2011"/><ref name="Frey 2016"/><ref name="Sanders & Masumoto 2021"/> Research continues to reveal new symbiont relationships, such as the newly described (2022) green algal genus and species, ''Rindifilum ramosum'' (family [[Biology:Ctenocladaceae|Ctenocladaceae]], order [[Biology:Ulvales|Ulvales]]), which associates with some members of ''Verrucaria''.<ref name="Malavasi et al. 2022"/>

One study of photobiont diversity in the Verrucariceae suggests that genera with simple crustose thalli (examples include ''[[Biology:Bagliettoa|Bagliettoa]]'', ''[[Biology:Hydropunctaria|Hydropunctaria]]'', and ''[[Biology:Wahlenbergiella|Wahlenbergiella]]'',) use a high diversity and unusual selection of photobionts, whereas the genera with more complex thalli (such as ''[[Biology:Placidium|Placidium]]'' and ''[[Biology:Dermatocarpon|Dermatocarpon]]'') tend to have lower photobiont diversity.<ref name="Thüs et al. 2011"/> [[Biology:Brown algae|Brown algae]] are far less frequent photobiont partners for the Verrucariaceae.<ref name="Frey 2016"/> An example is the [[Earth:Intertidal zone|intertidal]] marine lichen formed by the fungus ''[[Biology:Wahlenbergiella tavaresiae|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.<ref name="Sanders et al. 2004"/> The Verrucariaceae is the only mostly lichen-forming family to have photobiont associations with [[Biology:Yellow-green algae|yellow-green algae]] (class Xanthophyceae, genus ''[[Biology:Heterococcus|Heterococcus]]''). This genus was shown to be a photobiont partner in the thalli of three unrelated species of Verrucariaceae: ''[[Biology:Hydropunctaria rheitrophila|Hydropunctaria rheitrophila]]'', ''[[Biology:Verrucaria funckii|Verrucaria funckii]]'', and ''V.&nbsp;hydrela''.<ref name="Thüs et al. 2011"/> In 2022, a crustose, seashore-inhabiting member of the Verrucariaceae was reported as a mycobiont partner for the free-living macroscopic genus ''[[Biology:Urospora (alga)|Urospora]]'', the first time that genus, or even any in the order [[Biology:Ulotrichales|Ulotrichales]], has been recorded as a photobiont.<ref name="Černajová et al. 2022"/>

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 ''[[Biology:Pelvetia|Pelvetia]]'' and ''[[Biology:Ascophyllum|Ascophyllum]]'', and the green alga ''[[Biology:Prasiola crispa|Prasiola crispa]]''.<ref name="Thüs et al. 2011"/> 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.<ref name="Garrido-Benavent et al. 2017"/> Because the fungus grows independently as [[Biology:Mycelium|mycelium]] in close contact with the algae, the interaction between ''Ascophyllum'' and verrucariacean fungi is not technically considered a lichen symbiosis.<ref name="Hawksworth 1988"/> This type of biological partnership has been referred to as an "intimate, mutualistic association",<ref name="Sanders & Masumoto 2021"/> or a [[Biology:Mycophycobiosis|mycophycobiosis]];<ref name="Hawksworth 1988"/> more generally, associated species with this type of loose symbiosis have been called "borderline lichens".<ref name="Kohlmeyer et al. 2004"/>

==Habitat, distribution, and ecology==

[[File:Coastal vegetation zones at St Malo.jpg|thumb|''Hydropunctaria maura'' forming a stripe of black lichen (a vegetation zone) along the high-tide line in Saint-Malo, northwestern France]]

Collectively, Verrucariaceae has a [[Biology:Cosmopolitan distribution|cosmopolitan distribution]], although the majority of its species occur in [[Earth:Temperate climate|temperate climate]]s. Rocks and soil are the most common [[Biology:Substrate|substrates]], with growth on wood and bark less common.{{sfn|Cannon|Kirk|2007|pages=373–374}} Among the rock-dwelling species, there are both epiliths (those that grow on the surface) and endoliths (those that grow "within" the rocks, i.e., under and around the rock crystals). [[Earth:Calcareous|Calcareous]] rocks are the most common substrate for the rock-dwellers, but some, especially those in aquatic or semi-aquatic habitats, grow on [[Earth:Siliceous rock|siliceous rock]]s.<ref name="Gueidan et al. 2007"/> Some species grow on mosses,<ref name="Döbbeler & Triebel 1985"/><ref name="Döbbeler 1997"/> and some grow on leaves, including all species of genus ''[[Biology:Phylloblastia|Phylloblastia]]''.<ref name="Lee & Hur 2022"/> Several Verrucariaceae species are components of [[Biology:Biological soil crust|biological soil crust]]s and contribute to the formation and stabilisation of soil, particularly in somewhat [[Earth:Arid|arid]] areas; examples include members of ''[[Biology:Endocarpon|Endocarpon]]'' and ''[[Biology:Catapyrenium|Catapyrenium]]''.<ref name="Belnap et al. 2001"/> Although most species are terrestrial, there are some semi-aquatic species, with representatives from both fresh and salt water.<ref name="Frey 2016"/> As of 2015, there were 26 known marine Verrucariaceae, distributed amongst the genera ''[[Biology:Hydropunctaria|Hydropunctaria]]'' (6 species), ''[[Biology:Mastodia|Mastodia]]'' (1 sp.), ''[[Biology:Verrucaria|Verrucaria]]'' (16 spp.), and ''[[Biology:Wahlenbergiella|Wahlenbergiella]]'' (3 spp.).<ref name="Jones et al. 2015"/> Several of these marine lichens have been investigated preliminarily to determine their suitability for use as [[Earth:Bioindicator|bioindicator]]s of coastal [[Earth:Water pollution|water pollution]].<ref name="Nokes et al. 2022"/>

[[File:Verrucula arnoldaria Jymm.jpg|thumb|right|''[[Biology:Verrucula arnoldaria|Verrucula arnoldaria]]'' (darkened parts of thallus) growing parasitically on the crustose lichen ''Calogaya arnoldii'']]

Several Verrucariaceae genera consist entirely of lichenicolous (lichen-dwelling) fungi: ''[[Biology:Bellemerella|Bellemerella]]'', ''[[Biology:Clauzadella|Clauzadella]]'', ''[[Biology:Haleomyces|Haleomyces]]'', ''[[Biology:Halospora|Halospora]]'', ''[[Biology:Norrlinia|Norrlinia]]'', ''[[Biology:Phaeospora|Phaeospora]]'', ''[[Biology:Telogalla|Telogalla]]''. Lichenicolous fungi occur in other genera to a lesser extent. Some genera, including ''[[Biology:Heteroplacidium|Heteroplacidium]]'', ''[[Biology:Placidiopsis|Placidiopsis]]'', ''[[Biology:Placocarpus|Placocarpus]]'', ''[[Biology:Placopyrenium|Placopyrenium]]'', ''[[Biology:Thelidium|Thelidium]]'', ''[[Biology:Verrucaria|Verrucaria]]'', and ''[[Biology:Verruculopsis|Verruculopsis]]'' contain lichenicolous lichens.<ref name="Diederich et al. 2018"/>

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.<ref name="Orange & Chhetri 2022"/>

==Uses==
There are no species in the Verrucariaceae that have any major economic significance.{{sfn|Cannon|Kirk|2007|pages=373–374}} The only lichen in the family that has been recorded as being used in [[Medicine:Traditional medicine|traditional medicine]] is ''[[Biology:Dermatocarpon miniatum|Dermatocarpon miniatum]]''. In [[Medicine:Traditional Chinese medicine|Traditional Chinese medicine]], the lichen, known as "white stone ear" ({{lang|und-Hani|皮果衣}}, pí guǒ yī), is used in treatments for high blood pressure, as a [[Chemistry:Diuretic|diuretic]], for expelling [[Biology:Parasite|parasite]]s, for malnutrition in children, for [[Medicine:Dysentery|dysentery]], to improve digestion, and for abdominal distension.<ref name="Crawford 2019"/>

==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.<ref name="Wijayawardene et al. 2022"/> Following the genus name is the [[Biology:Author citation (botany)|taxonomic authority]] (those who first circumscribed the genus; standardised author abbreviations are used), year of publication, and the estimated number of species.<ref name="Wijayawardene et al. 2022"/> Other recent estimates of the number of Verrucariaceae taxa include: 46 genera and 931 species (2008),<ref name="Kirk et al. 2008"/> 48–54 genera and 850–870 species (2016),<ref name="Frey 2016"/> and 43 genera and 943 species (2017).<ref name="Lücking et al. 2017"/> The GBIF accept over 60+ genera and nearly a thousand species.<ref>{{cite web |title=Verrucariaceae |url=https://www.gbif.org/species/4841 |website=www.gbif.org |access-date=29 August 2022 |language=en}}</ref> {{As of|2022|September}}, Species Fungorum (in the [[Biology:Catalogue of Life|Catalogue of Life]]), accepts 56 genera and 575 species (including 8 [[Biology:Variety (botany)|varieties]] and 6 [[Biology:Form (botany)|forms]]) in the Verrucariaceae. The largest of these is the type genus, ''Verrucaria'', at 146 accepted species.<ref name="CoL-Species Fungorum"/>{{#tag:ref|The Verrucariaceae species accepted by Species Fungorum are those for which they have expressed a taxonomic opinion, and does not necessarily represent the total number of species in the family.|group=note}} Verrucariaceae is the third-largest family of lichen-forming fungi, following the [[Biology:Parmeliaceae|Parmeliaceae]] and the [[Biology:Graphidaceae|Graphidaceae]].<ref name="Lücking et al. 2017"/>
{{Multiple image|perrow = 2|total_width=450
| image1 = Bagliettoa marmorea 66981341.jpg
| image2 = Streamside Stippleback (4504253631).jpg
| image3 = Verrucaria compacta - Flickr - pellaea (2).jpg
| image4 = Verrucaria maura Kergulan 2008-01-29 13.jpg
| footer = Clockwise from upper left: ''[[Biology:Bagliettoa marmorea|Bagliettoa marmorea]]'' (crustose, endolithic); ''[[Biology:Dermatocarpon luridum|Dermatocarpon luridum]]'' (foliose/umbilicate); ''[[Biology:Heteroplacidium compactum|Heteroplacidium compactum]]'' (crustose/[[Biology:Areolate|areolate]]); and ''[[Biology:Hydropunctaria maura|Hydropunctaria maura]]'' (crustose, epilithic)}}
{{Multiple image|perrow = 2|total_width=450
| image1 = Tree Stipplescale (4752735810).jpg
| image2 = Placocarpus schaereri Jymm.jpg
| image3 = Staurothele elenkinii - Flickr - pellaea (2).jpg
| image4 = Wahlenbergiella mucosa Jymm.jpg
| footer = Clockwise from upper left: ''[[Biology:Placidium arboreum|Placidium arboreum]]'' (squamulose, corticolous); ''[[Biology:Placocarpus schaereri|Placocarpus schaereri]]'' (crustose/areolate, epilithic); ''[[Biology:Staurothele elenkinii|Staurothele elenkinii]]'' (crustose, epilithic); and ''[[Biology:Wahlenbergiella mucosa|Wahlenbergiella mucosa]]'' (crustose, epilithic)}}

*''[[Biology:Agonimia|Agonimia]]'' {{small|Zahlbr. (1909)}}<ref name="Zahlbruckner 1909"/> – ca. 20 spp.
*''[[Biology:Anthracocarpon|Anthracocarpon]]'' {{small|Breuss (1996)}}<ref name="Breuss 1996"/> – 1 sp.
*''[[Biology:Atla (lichen)|Atla]]'' {{small|S.Savić & Tibell (2008)}}<ref name="Savić & Tibell 2008"/> – 10 spp.
*''[[Biology:Awasthiella|Awasthiella]]'' {{small|Kr.P.Singh (1980)}}<ref name="Singh 1980"/> – 1 sp.
*''[[Biology:Bagliettoa|Bagliettoa]]'' {{small|A.Massal. (1853)}}<ref name="Massalongo 1853"/> – 17 spp.
*''[[Biology:Bellemerella|Bellemerella]]'' {{small|Nav.-Ros. & Cl.Roux (1997)}}<ref name="Navarro-Rosinés & Roux 1997"/> – 4 spp.
*''[[Biology:Catapyrenium|Catapyrenium]]'' {{small|Flot. (1850)}}<ref name="Flotow 1850"/> – 6 spp.
*''[[Biology:Clauzadella|Clauzadella]]'' {{small|Nav.-Ros. & Cl.Roux (1996)}}<ref name="Navarro-Rosinés & Roux 1996"/> – 1 sp.
*''[[Biology:Clavascidium|Clavascidium]]'' {{small|Breuss (1996)}}<ref name="Breuss 1996"/> – 9 spp.
*''[[Biology:Dermatocarpon|Dermatocarpon]]'' {{small|Eschw. (1824)}}<ref name="Eschweiler 1824"/> – 20 spp.
*''[[Biology:Diederimyces|Diederimyces]]'' {{small|Etayo (1995)}}<ref name="Etayo 1995"/> – 2 spp.
*''[[Biology:Endocarpon|Endocarpon]]'' {{small|Hedw. (1789)}}<ref name="Hedwig 1789"/> – ca. 75 spp.
*''[[Biology:Flakea|Flakea]]'' {{small|O.E.Erikss. (1992)}}<ref name="Eriksson 1992"/> – 1 sp.
*''[[Biology:Glomerilla|Glomerilla]]'' {{small|Norman (1869)}}<ref name="Norman 1868"/> – 1 sp.
*''[[Biology:Haleomyces|Haleomyces]]'' {{small|D.Hawksw. & Essl. (1993)}}<ref name="Hawksworth & Esslinger 1993"/> – 1 sp.
*''[[Biology:Halospora|Halospora]]'' {{small|(Zschacke) Tomas. & Cif. (1952)}}<ref name="Ciferri & Tomaselli 1952"/> – 4 spp.
*''[[Biology:Henrica|Henrica]]'' {{small|B.de Lesd. (1921)}}<ref name="Bouly de Lesdain 1921"/> – 4 spp.
*''[[Biology:Heterocarpon|Heterocarpon]]'' {{small|Müll.Arg. (1885)}}<ref name="Müller 1885"/> – 1 sp.
*''[[Biology:Heteroplacidium|Heteroplacidium]]'' {{small|Breuss (1996)}}<ref name="Breuss 1996"/> – 12 spp.
*''[[Biology:Hydropunctaria|Hydropunctaria]]'' {{small|C.Keller, Gueidan & Thüs (2009)}}<ref name="Gueidan et al. 2009"/> – 8 spp.
*''[[Biology:Involucropyrenium|Involucropyrenium]]'' {{small|Breuss (1996)}}<ref name="Breuss 1996"/> – 9 spp.
*''[[Biology:Mastodia|Mastodia]]'' {{small|Hook.f. & Harv. (1847)}}<ref name="Hooker 1844"/> – 5 spp.
*''[[Biology:Moriola|Moriola]]'' {{small|Norman (1872)}}<ref name="Norman 1872"/> – ca. 15 spp.
*''[[Biology:Muellerella|Muellerella]]'' {{small|Hepp ex Müll.Arg. (1862)}}<ref name="Müller 1862"/> – 9 spp.
*''[[Biology:Neocatapyrenium|Neocatapyrenium]]'' {{small|H.Harada (1993)}}<ref name="Harada 1993"/> – 5 spp.
*''[[Biology:Nesothele|Nesothele]]'' {{small|Orange (2022)}}<ref name="Orange & Chhetri 2022"/> – 5 spp.
*''[[Biology:Normandina|Normandina]]'' {{small|Nyl. (1855)}}<ref name="Nylander 1855"/> – 3 spp.
*''[[Biology:Norrlinia|Norrlinia]]'' {{small|Theiss. & Syd. (1918)}}<ref name="Theissen & Sydow 1918"/> – 2 spp.
*''[[Biology:Parabagliettoa|Parabagliettoa]]'' {{small|Gueidan & Cl.Roux (2009)}}<ref name="Gueidan et al. 2009"/> – 3 spp.
*''[[Biology:Phaeospora|Phaeospora]]'' {{small|Hepp ex Stein (1879)}}<ref name="Stein 1879"/> – 14 spp.
*''[[Biology:Phylloblastia|Phylloblastia]]'' {{small|Vain. (1921)}}<ref name="Vainio 1921"/> – 12 spp.
*''[[Biology:Placidiopsis|Placidiopsis]]'' {{small|Beltr. (1858)}}<ref name="Beltramini 1858"/> – 20 spp.
*''[[Biology:Placidium|Placidium]]'' {{small|A.Massal. (1855)}}<ref name="Massalongo 1855b"/> – 28 spp.
*''[[Biology:Placocarpus|Placocarpus]]'' {{small|Trevis. (1860)}}<ref name="Trevisan 1860"/> – 5 spp.
*''[[Biology:Placopyrenium|Placopyrenium]]'' {{small|Breuss (1987)}}<ref name="Nimis & Poelt 1987"/> – 22 spp.
*''[[Biology:Placothelium|Placothelium]]'' {{small|Müll.Arg. (1893)}}<ref name="Müller 1893"/> – 1 sp.
*''[[Biology:Plurisperma|Plurisperma]]'' {{small|Sivan. (1970)}}<ref name="Sivanesan 1970"/> – 1 sp.
*''[[Biology:Polyblastia|Polyblastia]]'' {{small|A.Massal. (1852)}}<ref name="Massalongo 1852"/> – ca. 40 + ca. 50 orphaned{{#tag:ref|"Orphaned" species are those that have been named and [[Biology:Species description|formally described]], but have not been updated and reassessed following a revision of the genus they are in.|group=note}}
*''[[Biology:Psoroglaena|Psoroglaena]]'' {{small|Müll.Arg. (1891)}}<ref name="Müller 1891"/> – 17 spp.
*''[[Biology:Rhabdopsora|Rhabdopsora]]'' {{small|Müll.Arg. (1888)}}<ref name="Müller 1888"/> – 2 spp.
*''[[Biology:Scleropyrenium|Scleropyrenium]]'' {{small|H.Harada (1993)}}<ref name="Harada 1993"/> – 2 spp.
*''[[Biology:Servitia|Servitia]]'' {{small|M.S.Christ. & Alstrup (2001)}}<ref name="Alstrup & Hansen 2001"/> – 1 sp.
*''[[Biology:Spheconisca|Spheconisca]]'' {{small|(Norman) Norman (1876)}}<ref name="Norman 1876"/> – ca. 20 spp.
*''[[Biology:Sporodictyon|Sporodictyon]]'' {{small|A.Massal. (1852)}}<ref name="Massalongo 1852b"/> – 5 spp.
*''[[Biology:Staurothele|Staurothele]]'' {{small|Norman (1852)}}<ref name="Norman 1852"/> – ca. 40 spp.
*''[[Biology:Telogalla|Telogalla]]'' {{small|Nik.Hoffm. & Hafellner (2000)}}<ref name="Hoffmann & Hafellner 2000"/> – 2 spp.
*''[[Biology:Thelidiopsis|Thelidiopsis]]'' {{small|Vain. (1921)}}<ref name="Vainio 1921"/> – 4 spp.
*''[[Biology:Thelidium|Thelidium]]'' {{small|A.Massal. (1855)}}<ref name="Massalongo 1855"/> – ca. 50 + ca. 50 orphaned
*''[[Biology:Trimmatothele|Trimmatothele]]'' {{small|Norman ex Zahlbr. (1903)}}<ref name="Engler & Prantl 1903"/> – 3 spp.
*''Turgidosculum'' {{small|Kohlm. & E.Kohlm. (1972)}}<ref name="Kohlmeyer & Kohlmeyer 1972"/> – 1 sp.
*''[[Biology:Verrucaria|Verrucaria]]'' {{small|Schrad. (1794)}}<ref name="Schrader 1794"/> – ca. 300 spp.
*''[[Biology:Verrucula|Verrucula]]'' {{small|J.Steiner (1896)}}<ref name="Steiner 1896"/> – 22 spp.
*''[[Biology:Verruculopsis|Verruculopsis]]'' {{small|Gueidan, Nav.-Ros. & Cl.Roux (2007)}}<ref name="Navarro-Rosinés et al. 2007"/> – ca. 10 spp.
*''[[Biology:Wahlenbergiella|Wahlenbergiella]]'' {{small|Gueidan & Thüs (2009)}}<ref name="Gueidan et al. 2009"/> – 3 spp.
*''[[Biology:Willeya|Willeya]]'' {{small|Müll.Arg. (1883)}}<ref name="Müller 1883"/> – 12 spp.

==Notes==
{{Reflist|group=note}}

==References==
{{Reflist|refs=

<ref name="Alstrup & Hansen 2001">{{cite journal |last1=Alstrup |first1=V. |last2=Hansen |first2=E.S. |year=2001 |title=New lichens and lichenicolous fungi from Greenland |journal=Graphis Scripta |volume=12 |issue=2 |pages=41–50}}</ref>

<ref name="Belnap et al. 2001">{{cite book |editor-last1=Belnap |editor-first1=J. |editor-first2=O.L. |editor-last2=Lange |last1=Belnap |first1=J. |last2=Büdel |first2=B. |last3=Lange |first3=O.L. |title=Biological Soil Crusts: Structure, Function, and Management |series=Ecological Studies |volume=150 |chapter=Biological Soil Crusts: Characteristics and Distribution |publisher=Springer Berlin Heidelberg |publication-place=Berlin, Heidelberg |year=2001 |isbn=978-3-540-43757-4 |doi=10.1007/978-3-642-56475-8_1 |pages=3–30}}</ref>

<ref name="Beltramini 1858">{{cite book |last1=Beltramini |first1=Francesco |year=1858 |title=I Licheni Bassanesi Enumerati e Descritti |location=Bassano |publisher=A. Roberti |page=212 |url=https://books.google.com/books?id=1TDbGh0i6_UC&pg=PA212 |language=it}}</ref>

<ref name="Bouly de Lesdain 1921">{{cite journal |last1=Bouly de Lesdain |first1=M. |year=1921 |title=Notes lichénologiques. XVIII |trans-title=Lichenological notes. XVIII |journal=Bulletin de la Société Botanique de France |volume=68 |issue=2 |pages=203–207 |doi=10.1080/00378941.1921.10836763 |language=fr |url=https://www.biodiversitylibrary.org/page/29643215|doi-access=free }}</ref>

<ref name="Breuss 1996">{{cite journal |last1=Breuss |first1=O. |year=1996 |title=Ein verfeinertes Gliederungskonzept für ''Catapyrenium'' (lichenisierte Ascomyceten, Verrucariaceae) |trans-title=A refined outline concept for ''Catapyrenium'' (Lichenized Ascomycetes, Verrucariaceae) |journal=Annalen des Naturhistorischen Museums Wien |volume=98 (Suppl.) |pages=35–50 |language=de}}</ref>

<ref name="Breuss 2010">{{cite journal |last1=Breuss |first1=Othmar |title=An updated world-wide key to the catapyrenioid lichens (Verrucariaceae) |journal=Herzogia |volume=23 |issue=2 |year=2010 |doi=10.13158/heia.23.2.2010.205 |pages=205–216}}</ref>

<ref name="Černajová et al. 2022">{{cite journal |last1=Černajová |first1=Ivana |last2=Schiefelbein |first2=Ulf |last3=Škaloud |first3=Pavel |title=Lichens from the littoral zone host diverse Ulvophycean photobionts |journal=Journal of Phycology |volume=58 |issue=2 |year=2022 |doi=10.1111/jpy.13234 |pages=267–280 |pmid=35032341}}</ref>

<ref name="CoL-Species Fungorum">{{cite web |author=Source dataset. Species Fungorum Plus: Species Fungorum for CoL+ |title=Verrucariaceae |url=https://www.catalogueoflife.org/data/taxon/HT9 |work=Catalog of Life Version: Annual Checklist 2022 |access-date=5 September 2022}}</ref>

<ref name="Ciferri & Tomaselli 1952">{{cite journal |last1=Ciferri |first1=R. |last2=Tomaselli |first2=R. |year=1952 |title=Scissioni di generi di licheni sulla base delle caratteristische del fungo |trans-title=Splits of lichen genera on the basis of the characteristics of the fungus |journal=Archivio Botanico Italiano |volume=28 |pages=1–15 |language=it}}</ref>

<ref name="Crawford 2019">{{cite book |last1=Crawford |first1=Stuart |chapter=Lichens Used in Traditional Medicine |page=81 |title=Lichen Secondary Metabolites. Bioactive Properties and Pharmaceutical Potential |edition=2 |year=2019 |editor-last1=Ranković |editor-first1=Branislav  |publisher=Springer Nature Switzerland AG |location= |isbn=978-3-030-16813-1}}</ref>

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<ref name="Zschacke refbundle">{{Unbulleted list citebundle
      |{{cite journal |last1=Zschacke |first1=Hermann |year=1913 |title=Die mitteleuropäischen Verrucariaceen. I |trans-title=The Central European Verrucariacea. I |journal=Hedwigia |volume=54 |pages=183–198 |language=de |url=https://www.biodiversitylibrary.org/page/420507}}
|{{cite journal |author-mask=9 |last1=Zschacke |first1=Hermann |year=1914 |title=Die mitteleuropäischen Verrucariaceen. II |journal=Hedwigia |volume=55 |pages=286–324 |language=de |url=https://www.biodiversitylibrary.org/page/424428}}
|{{cite journal |author-mask=9 |last1=Zschacke |first1=Hermann |year=1918 |title=Die mitteleuropäischen Verrucariaceen. Nachträge zu I und II. |journal=Hedwigia |volume=60 |pages=1–9 |language=de |url=https://www.biodiversitylibrary.org/page/424638}}
|{{cite journal |author-mask=9 |last1=Zschacke |first1=Hermann |year=1921 |title=Die mitteleuropäischen Verrucariaceen. III |journal=Hedwigia |volume=62 |pages=90–154 |language=de |url=https://www.biodiversitylibrary.org/page/425931}}
|{{cite journal |author-mask=9 |last1=Zschacke |first1=Hermann |year=1924 |title=Die mitteleuropäischen Verrucariaceen. IV |journal=Hedwigia |volume=65 |pages=46–64|language=de}}
|{{cite journal |author-mask=9 |last1=Zschacke |first1=Hermann |year=1927 |title=Die mitteleuropäischen Verrucariaceen. V |journal=Hedwigia |volume=67 |pages=45–85|language=de}}
   }}</ref>

}}

===Cited literature===
*{{cite book |last1=Cannon |first1=Paul F. |last2=Kirk |first2=Paul M. |title=Fungal Families of the World |publisher=CABI |publication-place=Wallingford |year=2007 |isbn=978-0-85199-827-5 |oclc=60741230}}

{{Taxonbar|from=Q3123598}}

[[Category:Verrucariales]]
[[Category:Ascomycota families]]

{{Sourceattribution|Verrucariaceae}}