{{Short description|Family of lichen-forming fungi}} {{Use dmy dates|date=April 2024}} {{Automatic taxobox | image = Fuscidea cyathoides 139905995.jpg | image_caption =''Fuscidea cyathoides'' | taxon = Fuscideaceae | authority = Hafellner (1984) | type_genus = ''Fuscidea'' | type_genus_authority = V.Wirth & Vězda (1972) | subdivision_ranks = Genera | subdivision = ''Albemarlea''<br> ''Fuscidea''<br> ''Maronea''<br> ''Maronora'' }}
'''Fuscideaceae''' is a family of fungi that form symbiotic relationships with algae to create lichens. These lichens typically have a crust-like appearance and are found worldwide, though they are most common in temperate regions. The family includes four genera and about 45 species, which primarily grow on tree bark, rocks, or occasionally on wood or leaves. Fuscideaceae lichens are characterised by their reproductive structures, cup-like formations called apothecia, which can vary in colour from red to dark brown or black. The family has undergone several changes in its classification over the years, with recent genetic studies placing it within the order Umbilicariales. Fuscideaceae lichens produce various chemical compounds (lichen products), some of which are unique to this family, and these chemicals are often used to help identify different species.
==Systematics== ===Taxonomy=== The family was circumscribed by the lichenologist Josef Hafellner in 1984. He included the type genus, ''Fuscidea'', and ''Maronea'' in his original conception of the family. He considered ''Maronea'' as sort of a version of ''Fuscidea'' with {{lichengloss|lecanorine}} apothecia and multi-spored asci.<ref name="Hafellner 1984"/> In his ''Outline of the Ascomycota'' series (2006), Ove Eriksson included the genus ''Hueidea'' in Fuscideaceae based on its ''Fuscidea''-type asci as the diagnostic character.<ref name="Eriksson 2006"/> Both ''Loxospora'' and ''Sarrameana'' had been proposed for inclusion in the Fuscideaceae,<ref name="Nash 2008"/> but they are now both placed in a new family, the Sarrameanaceae.<ref name="Wijayawardene et al. 2022"/> Ulrik Søchting and colleagues demonstrated with molecular data that Hueidea belongs in the Teloschistaceae (subfamily Caloplacoideae) and is congeneric with the Antarctic genus ''Huea''; the Nomenclature Committee for Fungi has therefore recommended rejection of ''Huea'',<ref name="May & Lendemer 2023"/> leaving ''Hueidea'' in Teloschistaceae and outside Fuscideaceae. The proposal is pending vote by the Nomenclature Committee and, if accepted, will be forwarded to the next International Botanical Congress.<ref name="Søchting et al. 2024"/>
In a proposal by Heidi Lie Andersen and Tor Tønsberg, published in late 2023, there has been a call to conserve the name ''Fuscidea'' over ''Maronea'', to addressing a taxonomic and nomenclatural issue in the Fuscideaceae. This suggestion stems from multiple molecular studies, particularly those by Anya Bylin et al. (2007),<ref name="Bylin et al. 2007"/> Jolanta Miadlikowska et al. (2014),<ref name="Miadlikowska et al. 2014"/> and research by Martina Zahradnikova (2017),<ref name="Zahradníková 2017"/> which demonstrated that ''Maronea'' is paraphyletic to ''Fuscidea'', with ''Maronea'' species nested within a larger monophyletic ''Fuscidea'' clade. The genus ''Maronea'', described by Abramo Bartolommeo Massalongo in 1856, consists of about 12 species, with a more restricted distribution compared to ''Fuscidea'', which comprises over 40 widely distributed species. To preserve nomenclatural stability and avoid the extensive renaming that transferring all ''Fuscidea'' species to ''Maronea'' or splitting ''Fuscidea'' into several smaller genera would entail, the authors advocate for conserving the name ''Fuscidea'' against ''Maronea''. This approach aims to minimise name changes, maintaining a monophyletic group within Fuscideaceae and supporting nomenclatural consistency.<ref name="Andersen & Tønsberg 2023"/>
===Classification===
Since its proposal in 1984, the classification of family Fuscidaceae has been a matter of some debate. Hafellner originally thought the family to be more closely related to the Teloschistaceae than to the Lecideaceae or Lecanoraceae, and he classified the family in the suborder Teloschistineae of the order Teloschistales. He described the similarities in the ascus structure between the Teloschistaceae and Fuscideaceae as "unmistakable".<ref name="Hafellner 1984"/> Miadlikowska and colleagues (2006) discovered that the resemblance in ascus structure between the Fuscideaceae and Teloschistales is due to homoplasy.<ref name="Miadlikowska et al. 2006"/>
In some preliminary molecular work, Lutzoni and colleagues concluded that the family was associated with the Ostropomycetidae.<ref name="Lutzoni et al. 2004"/> Reeb and colleagues (2004) identified the Umbilicariaceae as a closely related group to the Fuscideaceae, suggesting the establishment of a new order named Umbilicariales to encompass both groups.<ref name="Reeb et al. 2004"/> Wedin and others (2005) acknowledged a potential relationship between the Fuscideaceae and Umbilicariaceae, though with limited evidence.<ref name="Wedin et al. 2005"/> In 2006, Miadlikowska and her team recommended placing the Fuscideaceae-Ophioparmaceae-Umbilicariaceae grouping into a distinct order, Umbilicariales, under the class Lecanoromycetidae, yet this classification was not officially adopted.<ref name="Miadlikowska et al. 2014"/> Bylin and associates (2007) observed Fuscideacea as a close relative to Umbilicariales, despite a lack of strong evidence.<ref name="Bylin et al. 2007"/> The Ropalosporaceae were reclassified with a provisional placement within the Umbilicariales due to an unclear relationship. Contrarily, Bendiksby and Timdal (2013) contested the inclusion of Fuscideaceae within the Umbilicariales despite noting similarities in the ascus structure between ''Umbilicaria'' and ''Fuscidea'', characterised by an amyloid inner and outer layer separated by a nonamyloid layer.<ref name="Bendiksby & Timdal 2013"/> Molecular work by Miadlikowska and colleagues in 2014 suggested that the family occupied an outlying clade within the order Umbilicariales, which includes the Ophioparmaceae and Ropalosporaceae as well as the Umbilicariaceae itself.<ref name="Miadlikowska et al. 2014"/> More recent multigene phylogenies (nrITS, nrLSU, mtSSU, RPB2) place Fuscideaceae firmly within Umbilicariales, sister to Umbilicariaceae + Ophioparmaceae, corroborating earlier suggestions based on ascus architecture. The current "Outline of Fungi" (2024) recognises Fuscideaceae as the sole family in Umbilicariales lacking foliose thalli.<ref name="2024 Outline"/>
==Description==
Fuscideaceae lichens have a crust-like body (thallus), which often presents as a collection of tightly packed, cracked segments ({{lichengloss|areolate}}). It is not uncommon for these lichens to have a darker outline or base ({{lichengloss|prothallus}}), and some species may develop clusters of reproductive structures (soralia) on their surface. The photosynthetic partner within these lichens is typically {{lichengloss|chlorococcoid}}—a form of green algae.<ref name="Cannon et al. 2024"/>
The reproductive structures of these lichens, known as ascomata, take the form of {{lichengloss|apothecia}}, which are cup-like fungal structures. These can be either partially embedded in the thallus or sitting atop it (ranging from more or less {{lichengloss|immersed}} to {{lichengloss|sessile}}), and may or may not have a noticeable rim ({{lichengloss|thalline margin}}). The colour of the apothecia's {{lichengloss|disc}} varies from red to dark brown or black. Inside these structures, the supportive tissue ({{lichengloss|hamathecium}}) consists of sparsely to richly branched filaments (paraphyses), which often have coloured tips.<ref name="Cannon et al. 2024"/> [[File:Fuscidea recensa-8.jpg|thumb|upright=0.7|right|Kidney bean-shaped spore of ''Fuscidea recensa'']] The spore-producing cells (asci) feature a distinctive staining pattern when treated with certain chemical dyes (K/I+ dark blue for the external and internal layers, surrounded by a thick K/I+ pale blue gelatinous cap, but with a K/I– tube at the centre, or {{lichengloss|tholus}}). Fuscideaceae asci typically release eight or more spores, which are generally non-segmented (aseptate) or rarely divided once (1-septate), and range in colour from clear to pale brown. The shape of these spores can vary, with some resembling beans or showing a pinched middle section.<ref name="Cannon et al. 2024"/> The conidiomata are in the form of pycnidia; the conidia are non-septate, {{lichengloss|fusiform}} to ellipsoid in shape, and hyaline.<ref name="Frey 2016"/>
==Chemistry==
Fuscideaceae species typically have either depsides, depsidones, or no lichen products at all.<ref name="Frey 2016"/> Other secondary chemical compounds found in Fuscideaceae include anthraquinones, benzyl esters, higher aliphatic acids, and usnic acid.<ref name="Tønsberg 1992"/><ref name="Ekman 1993"/><ref name="Kantvilas 2004"/> Specific compounds identified from various members of the family include the orcinol depsides divaricatic acid, nordivaricatic acid, and imbricaric acid; the benzyl esters alectorialic acid and barbatolic acid.<ref name="Tønsberg 1992"/>
==Habitat and distribution==
Collectively, the Fuscideaceae has a cosmopolitan distribution, although species are predominantly found in temperate locales.<ref name="Frey 2016"/> The most extensive genus within this group, ''Fuscidea'', contains over 40 species that are primarily found in the Northern Hemisphere, although they also occur in the Southern Hemisphere.<ref name="Andersen & Tønsberg 2023"/> Genus ''Albemarlea'' contains a single species that is found exclusively in North Carolina's Albemarle-Pamlico Peninsula.<ref name="Lendemer et al. 2016"/>
Fuscideaceae species usually grow on bark, on rocks, or more rarely, on wood or leaves.<ref name="Frey 2016"/>
==Genera==
* ''Albemarlea'' {{au|Lendemer & R.C.Harris (2016)}}<ref name="Lendemer et al. 2016"/> – 1 sp. :Tentative placement in the Fuscideaceae. Its single species has a crust-like thallus, ''Fuscidea''-type asci, and green coccoid photobiont, but lacks any lichen products.<ref name="Lendemer et al. 2016"/> * ''Fuscidea'' {{au|V.Wirth & Vězda (1972)}}<ref name="Wirth & Vězda 1972"/> – 40+ spp.<ref name="Andersen & Tønsberg 2023"/> * ''Maronea'' {{au|A.Massal. (1856)}} – 12 spp.<ref name="Andersen & Tønsberg 2023"/> * ''Maronora'' {{au|Kalb & Aptroot (2018)}}<ref name="Kalb & Aptroot 2018"/> – 1 sp.
The genus ''Orphniospora'' {{au|Körb. (1874)}} was previously included in the family, but molecular analysis published in 2017 has since excluded it,<ref name="Zahradníková 2017"/> and its phylogenetic affinity is considered uncertain.<ref name="Cannon et al. 2024"/> ''Lettauia'' is another genus that was included in the Fuscideaceae (sometimes tentatively),<ref name="Frey 2016"/> but it has since been synonymised with ''Cryptodiscus'' (family Stictidaceae).<ref name="IF: Lettauia"/> ''Maronea'' is kept in the Fuscideaceae pending the nomenclatural decision regarding its name.<ref name="Andersen & Tønsberg 2023"/>
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|last447=Taylor |first447=P.W.J. |last448=Tennakoon |first448=D.S. |last449=Thakshila |first449=S.A.D. |last450=Thambugala |first450=K.M. |last451=Thamodini |first451=G.K. |last452=Thilanga |first452=D. |last453=Thines |first453=M. |last454=Tiago |first454=P.V. |last455=Tian |first455=X.G. |last456=Tian |first456=W.H. |last457=Tibpromma |first457=S. |last458=Tkalčec |first458=Z. |last459=Tokarev |first459=Y.S. |last460=Tomšovský |first460=M. |last461=Torruella |first461=G. |last462=Tsurykau |first462=A. |last463=Udayanga |first463=D. |last464=Ulukapı |first464=M. |last465=Untereiner |first465=W.A. |last466=Usman |first466=M. |last467=Uzunov |first467=B.A. |last468=Vadthanarat |first468=S. |last469=Valenzuela |first469=R. |last470=Van den Wyngaert |first470=S. |last471=Van Vooren |first471=N. |last472=Velez |first472=P. |last473=Verma |first473=R.K. |last474=Vieira |first474=L.C. |last475=Vieira |first475=W.A.S. |last476=Vinzelj |first476=J.M. |last477=Tang |first477=A.M.C. |last478=Walker |first478=A. |last479=Walker |first479=A.K. |last480=Wang |first480=Q.M. |last481=Wang |first481=Y. |last482=Wang |first482=X.Y. |last483=Wang |first483=Z.Y. |last484=Wannathes |first484=N. |last485=Wartchow |first485=F. |last486=Weerakoon |first486=G. |last487=Wei |first487=D.P. |last488=Wei |first488=X. |last489=White |first489=J.F. |last490=Wijesundara |first490=D.S.A. |last491=Wisitrassameewong |first491=K. |last492=Worobiec |first492=G. |last493=Wu |first493=H.X. |last494=Wu |first494=N. |last495=Xiong |first495=Y.R. |last496=Xu |first496=B. |last497=Xu |first497=J.P. |last498=Xu |first498=R. |last499=Xu |first499=R.F. |last500=Xu |first500=R.J. |last501=Yadav |first501=S. |last502=Yakovchenko |first502=L.S. |last503=Yang |first503=H.D. |last504=Yang |first504=X. |last505=Yang |first505=Y.H. |last506=Yang |first506=Y. |last507=Yang |first507=Y.Y. |last508=Yoshioka |first508=R. |last509=Youssef |first509=N.H. |last510=Yu |first510=F.M. |last511=Yu |first511=Z.F. |last512=Yuan |first512=L.L. |last513=Yuan |first513=Q. |last514=Zabin |first514=D.A. |last515=Zamora |first515=J.C. |last516=Zapata |first516=C.V. |last517=Zare |first517=R. |last518=Zeng |first518=M. |last519=Zeng |first519=X.Y. |last520=Zhang |first520=J.F. |last521=Zhang |first521=J.Y. |last522=Zhang |first522=S. |last523=Zhang |first523=X.C. |last524=Zhao |first524=C.L. |last525=Zhao |first525=H. |last526=Zhao |first526=Q. |last527=Zhao |first527=H. |last528=Zhao |first528=H.J. |last529=Zhou |first529=H.M. |last530=Zhu |first530=X.Y. |last531=Zmitrovich |first531=I.V. |last532=Zucconi |first532=L. |last533=Zvyagina |first533=E. |title=The 2024 Outline of Fungi and fungus-like taxa |journal=Mycosphere |volume=15 |issue=1 |pages=5146–6239 [5225] |year=2024 |doi=10.5943/mycosphere/15/1/25 |url=https://www.researchgate.net/publication/385939154|hdl=11584/429245 |hdl-access=free }}</ref>
}}
{{Taxonbar|from=Q5509956}}
Category:Umbilicariales Category:Lecanoromycetes families Category:Lichen families Category:Taxa described in 1984 Category:Taxa named by Josef Hafellner