{{Short description|Genus of lichen}} {{Use British English|date=January 2024}} {{Use dmy dates|date=January 2024}} {{Automatic taxobox | image = Schaereria fuscocinerea 50-Digulleville,Jardeheu 2015-05-30 01.jpg | image_caption = ''Schaereria fuscocinerea'' | display_parents = 3 | grandparent_authority = Lumbsch & Leavitt (2018) | parent_authority = M.Choisy ex Hafellner (1984) | taxon = Schaereria | authority = Körb. (1855) | type_species = ''Schaereria lugubris'' | type_species_authority = (A.Massal.) Körb. (1855) | synonyms_ref = <ref name="Species Fungorum synonymy"/> | synonyms = *''Hafellnera'' {{small|Houmeau & Cl.Roux (1984)}} *''Ropalospora'' {{small|A.Massal. (1860)}} | subdivision_ranks = | subdivisions = }}
'''''Schaereria''''' is a genus of lichen-forming fungi. It is the sole genus in the family '''Schaereriaceae''', which itself is the only family in the '''Schaereriales''', an order in the subclass Ostropomycetidae of the class Lecanoromycetes.<ref name="Wijayawardene et al. 2020"/> Most ''Schaereria'' species are crustose lichens that live on rocks. ''Schaereria'' was first proposed by Gustav Wilhelm Körber in 1855 and was later taken up by other lichenologists despite periods of disuse.
Distinctive characteristics of species in the ''Schaereria'' genus include a crustose to squamulose thallus, ascomata (fruiting bodies) with a blackened ring and a blackish {{lichengloss|disk}}, and asci (spore-bearing cells) that lack {{lichengloss|tholus}} (a thickened part of the inner wall near the tip). The secondary chemistry of the genus produces specific substances, including the pigment known as {{lichengloss|Cinereorufa-green}} in ''Schaereria cinereorufa''. The genus, having a cosmopolitan distribution, primarily favours cold to cool climates of the Northern Hemisphere. However, certain species have been identified in both polar regions and others in warmer locales like Queensland (Australia). ''Schaereria'' species often grow on siliceous rocks, with a few variants being epiphytic (plant-dwelling) or even lichen-dwelling. Various lichenicolous fungi and lichens have been observed to grow on ''Schaereria'' species.
==Systematics== ===Historical taxonomy=== Genus ''Schaereria'' was named and circumscribed by the German lichenologist Gustav Wilhelm Körber in 1855, with ''Schaereria lugubris'' assigned as the type, and at the time, only species.<ref name="Körber 1855"/> According to Hannes Hertel and L. Zürn, it is clear that he was actually referring to ''Schaereria cinereorufa'' {{au|(Schaer.) Th.Fr.}}, but he mistakenly believed he had ''Ropalospora lugubris'' (formerly ''Lecidea lugubris'') on hand. For this reason, Körber's name is not available for the genus.<ref name="Hertel & Zürn 1986"/> Despite this, ''Schaereria'' was accepted a few years later (in 1860) by Theodor Magnus Fries, who used it in his work on Arctic, European, and Greenlandic lichens.<ref name="Fries 1860"/> It subsequently fell into disuse for more than a century after William Nylander placed it in synonymy with ''Lecidea''.<ref name="Lunke et al. 1996"/> Josef Poelt and Antonín Vězda resurrected ''Schaereria'' in 1977, and included ''S. cinereorufa''.<ref name="Poelt & Vězda 1977"/> The genus name honours the Swiss pastor and lichenologist Ludwig Emanuel Schaerer.<ref name="Hertel 2012"/>
In 1989, David Leslie Hawksworth and John Charles David put forth a proposal to conserve the name ''Schaereria'' (as defined by Theodor Magnus Fries in 1874) for a group of species with ''S. cinereorufa'' as the type species, instead of using ''Schaereria'' as defined by Körber. The reason for this proposal is that ''S. cinereorufa'' is a well-known species under the name ''Schaereria'', while Körber applied the name ''S. lugubris'' to a different species that should have been named ''S. cinereorufa''. By conserving the name ''Schaereria'' as suggested, it avoids the need to create a new name for the same group of lichens and prevents confusion with another genus called ''Ropalospora''. By accepting this proposal, both ''Ropalospora'' and ''Schaereria'' could continue to be used with their current meanings, avoiding any unnecessary changes.<ref name="Hawksworth & David 1989"/> In the subsequent recommendations of the Committee for Fungi and Lichens, they agreed that it was desirable to conserve the genus with one of the original specimens used by the author, noting that this specimen was erroneously labelled ''Schaereria lugubris'', but clearly belongs to ''S. cinereorufa''.<ref name="Gams 1993"/>
''Schaereria'' is one of several dozen genera whose species were previously included in the large genus ''Lecidea''.<ref name="Hertel & Rambold 1985"/> However, ''Lecidea'' has a different ascus structure than ''Schaereria''. The family Schaereriaceae was first proposed by French lichenologist Maurice Choisy in 1949, but he did not publish the name validly as it did not meet the criteria for publication as determined by the International Code of Nomenclature for algae, fungi, and plants.<ref name="Index Fungorum: Schaereriaceae"/> Josef Hafellner published the family name Schaereriaceae validly in 1984.<ref name="Hafellner 1984"/>
===Classification=== Hafellner noted some similarities in the characteristics of the hymenium between the Schaereriaceae and the order Pezizales,<ref name="Hafellner 1984"/> and the family was included there in the 1985 version of the ''Outline of the Ascomycota''.<ref name="Eriksson & Hawksworth 1985"/> This classification was later shown to be inappropriate as the Schaereriaceae do not have operculate asci{{emdash}}a characteristic of the Pezizales. After this the family was associated with either the Teloschistineae (a suborder of the Teloschistales), the Agyriineae (a suborder of the Lecanorales),<ref name="Lunke et al. 1996"/> or placed in the order Sarrameanales.<ref name="Frey 2016"/> ''Schaereria'' has also been included and excluded from the family Lecideaceae several times.<ref name="Schmull et al. 2011"/> Molecular studies that have included ''Schaereria'' species have shown that it occupies a relatively isolated phylogenetic position.<ref name="Wedin et al. 2005"/><ref name="Schmull et al. 2011"/><ref name="Miadlikowska et al. 2014"/>
In 2018, H. Thorsten Lumbsch and Steven Leavitt proposed the new order Schaereriales to contain the family, segregating the order Schaereriales from Sarrameanales. They considered the Schaereriales to have "unresolved relationships" in the parent taxon, Ostropomycetidae. In their analysis, which used a "temporal" method to group together comparable ordinal and family ranks, Schaereriales has a sister group relationship with the order Sarrameanales. Both Schaereriales and Sarrameanales form a clade that is sister to the Baeomycetales.<ref name="Kraichak et al. 2018a"/><ref name="Kraichak et al. 2018b"/> In a critical review of the temporal method for lichen classification, Robert Lücking found flaws in their analysis and rejected the proposed split, instead retaining both Sarrameanaceae and Schaereriaceae in the Sarrameanales.<ref name="Lücking 2019"/>
==Description== thumb|Closeup of {{lichengloss|lecideine}} ascomata of ''Schaereria cinereorufa'' Several characteristics unite taxa in the order Schaereriales. These include: a thallus that is crustose to squamulose; a trebouxioid photobiont partner (spherical unicellular green algae); ascomata being lecideine (having an apothecium which lacks algae and lacks an {{lichengloss|amphithecium}}); apothecia that are hemiangiocarpous (meaning they open before the spores are mature); and a cup-shaped {{lichengloss|excipulum}}. Microscopic characteristics of the Schaereriales include asci of the ''Schaereria''-type (lacking a {{lichengloss|tholus}}–the thickened inner part of an ascus tip), and ascospores that are hyaline, thin-walled, and one-celled (i.e., lacking septa).<ref name="Kraichak et al. 2018a"/> There are eight spores per ascus. The {{lichengloss|hamathecium}} contains non-amyloid, non-gelatinous, unbranched {{lichengloss|paraphyses}}.<ref name="Frey 2016"/> The secondary chemistry of this group includes depsides and unknown compounds.<ref name="Frey 2016"/> ''Schaereria cinereorufa'' produces a green- to turquoise-coloured insoluble lichen pigment known as {{lichengloss|Cinereorufa-green}}.<ref name="Meyer & Printzen 2000"/> The identification of species within ''Schaereria'' is primarily based on specific traits: the chemical composition and form of the thallus, the colouration of the apothecia, and the distinctive shape and organization of ascospores within the ascus.<ref name="Kantvilas 2023"/>
==Anatomy and development== The anatomy and development of several ''Schaereria'' species have been studied in some detail. The genus displays a varied morphology, encompassing both crustose and squamulose lichens. For instance, ''Schaereria cinereorufa'' is characterised by a squamulose to {{lichengloss|bullate}} (blistered) thallus, often found on siliceous rocks, with {{lichengloss|sessile}} or marginally located apothecia. In contrast, ''S. corticola'' has a sparsely developed, endo- to episubstral thallus with soredia and sessile apothecia. ''Schaereria fuscocinerea'', somewhat intermediate, features an {{lichengloss|areolate}} thallus on siliceous rocks, and its fruiting bodies are either {{lichengloss|immersed}} in or between {{lichengloss|areolae}}.<ref name="Lunke et al. 1996"/>
The anatomy within the genus is more consistent. In ''S. cinereorufa'' and ''S. fuscocinerea'', the thallus is covered by an {{lichengloss|epinecral layer}} (a layer of dead fungal hyphae), and no distinct cortex is formed. However, the upper parts of the hyphae in the thallus are pigmented brownish. In contrast, ''S. corticola'' lacks a distinct upper layer, primarily consisting of soredia.<ref name="Lunke et al. 1996"/>
The development of ''Schaereria'' ascomata is somewhat variable depending on species. In ''S. cinereorufa'', the ascomatal {{lichengloss|primordium}} begins as groups of ascogonia forming a more or less spherical cluster, followed by coiled {{lichengloss|ascogonia}} with {{lichengloss|trichogynes}}. The primordium eventually rises to the thallus's upper surface, with the upper part forming the hymenium. In ''S. corticola'', spherical primordia with ascogonia are observed, becoming pigmented and forming the cup-shaped {{lichengloss|excipulum}}. True paraphyses replace the {{lichengloss|paraphysoid|paraphysoidal}} network in the {{lichengloss|hamathecium}}. In ''S. fuscocinerea'', ascogonia with trichogynes appear in the {{lichengloss|algal layer}}, with subsequent growth of paraphysoids, followed by the differentiation of ascogenous hyphae and true paraphyses.<ref name="Lunke et al. 1996"/>
==Habitat, distribution, and ecology== Collectively, the genus has a cosmopolitan distribution.<ref name="Frey 2016"/> The majority of ''Schaereria'' species inhabit cold to cool climates within the Northern Hemisphere. ''S. fuscocinerea'', however, is an exception, as it can be found in both polar regions. Additionally, ''S. xerophila'' thrives in the lowland regions of Queensland. ''Schaereria'' species are typically found growing on rocky surfaces composed of siliceous materials. In contrast, ''S. corticola'' is bark-dwelling, ''S. dolodes'' is both bark- and wood-dwelling,<ref name="Schmull et al. 2011"/> ''S. corticola'' has an epiphytic lifestyle, while ''S. parasemella'' has a lichenicolous (lichen-dwelling) habit.<ref name="Kantvilas 1999"/>
Some lichenicolous fungi and lichens have been recorded growing on ''Schaereria'' species. These include ''Endococcus perpusillus'', ''Buellia miriquidica'', ''Buellia uberior'', and ''Halecania parasitica'' (the last three on ''Schaereria fuscocinerea'').<ref name="Diederich et al. 2018"/> A ''Sclerococcum'' species in the ''S. australe''–''S. saxatile'' group has been reported infesting ''Schaereria bullata''.<ref name="Kantvilas 1999"/>
==Species== {{As of|2024|January}}, Species Fungorum (in the Catalogue of Life) accepts 11 species of ''Schaereria''.<ref name="CoL"/> *''Schaereria albomarginata'' {{au|Øvstedal (2009)}}<ref name="Øvstedal & Lewis Smith 2009"/> *''Schaereria argentina'' {{au|Gyeln. (1934)}}<ref name="Gyelnik 1934"/> *''Schaereria australis'' {{au|Kantvilas (2023)}}<ref name="Kantvilas 2023"/> – Australia *''Schaereria brunnea'' {{au|Björk, T.Sprib. & T.B.Wheeler (2009)}}<ref name="Spribille et al. 2009"/> – Canada *''Schaereria bullata'' {{au|Kantvilas (1999)}}<ref name="Kantvilas 1999"/> – Tasmania *''Schaereria cinereorufa'' {{au|(Schaer.) Th.Fr. (1861)}} – Northern Hemisphere *''Schaereria corticola'' {{au|Muhr & Tønsberg (1992)}}<ref name="Tønsberg 1992"/> – temperate Northern Hemisphere *''Schaereria dolodes'' {{au|(Nyl.) Schmull & T.Sprib. (2005)}}<ref name="Schmull & Spribille 2005"/> – western North America; Tasmania *''Schaereria fabispora'' {{au|Hertel & Zürn (1986)}}<ref name="Hertel & Zürn 1986"/> – Norway *''Schaereria fuscocinerea'' {{au|(Nyl.) Clauzade & Cl.Roux (1985)}} – cosmopolitan distribution *''Schaereria gresinonis'' {{au|(B.de Lesd.) Gyeln. (1937)}} *''Schaereria parasemella'' {{au|(Nyl.) Lumbsch (1997)}}<ref name="Lumbsch 1997"/> – Finland *''Schaereria porpidioides'' {{au|Fryday & Common (2001)}}<ref name="Fryday & Common 2001"/> – Falkland Islands *''Schaereria serenior'' {{au|(Vain.) Vitik. (2004)}} – Fennoscandia *''Schaereria xerophila'' {{au|Rambold & H.Mayrhofer (1989)}}<ref name="Rambold 1989"/> – Australia
===Former species=== Several taxa once classified in ''Schaereria'' have since been reduced to synonymy, or transferred to other genera. Examples include: *''Schaereria decipiens'' {{au|(Hedw.) Gyeln. (1937)}} is now known as ''Psora decipiens'';<ref name="IF: Schaereria decipiens"/> similarly, ''Schaereria icterica'' {{au|(Mont.) Gyeln. (1937)}} is now ''Psora icterica''.<ref name="IF: Schaereria icterica"/> *Both ''Schaereria endocyanea'' {{au|(Stirt.) Hertel & Gotth.Schneid. (1980)}} and ''Schaereria tenebrosa'' {{au|(Flot.) Hertel & Poelt 1980)}} are now considered synonymous with ''Schaereria fuscocinerea''.<ref name="IF: Schaereria endocyanea"/><ref name="IF: Schaereria tenebrosa"/> *''Schaereria lugubris'' {{au|(Sommerf.) Körb. (1855)}} is now ''Ropalospora lugubris''.<ref name="IF: Schaereria lugubris"/> *''Schaereria lurida'' {{au|(Dill. ex With.) Gyeln. (1937)}} is now ''Dermatocarpon luridum''.<ref name="IF: Schaereria lurida"/> *''Schaereria ostreata'' {{au|(Hoffm.) Gyeln. (1937)}} is now ''Hypocenomyce scalaris''.<ref name="IF: Schaereria ostreata"/> *''Schaereria pissodes'' {{au|(Stirt.) Clauzade & Cl.Roux (1985)}} is now ''Clauzadeana macula''.<ref name="IF: Schaereria pissodes"/>
==References== {{Reflist|colwidth=30em|refs=
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{{Taxonbar|from=Q10663129}}
Category:Lecanoromycetes Category:Lecanoromycetes genera Category:Lichen genera Category:Taxa described in 1855 Category:Taxa named by Gustav Wilhelm Körber