Micarea is a genus of lichenized fungi in the family Pilocarpaceae.[2] The widely distributed genus contains 126 species[3] and new species are described actively. Species in the genus are crustose lichens and their photobiont (the non-fungal organism) is a single-celled green alga.[4]
Taxonomy
Micarea was circumscribed by Elias Magnus Fries in his 1825 work Systema Orbis Vegetabilis.[5] There have been some taxonomic disputes about the genus.[6] Nowadays, the genus is accepted, although it is paraphyletic and in need of further research.[4] Micarea prasina is the type species of the genus.[7][8]
Reproduction
Lichenized fungi have developed diverse reproduction strategies. The microlichen genus Micarea is an excellent model for studying the effects of reproductive traits and environmental factors on speciation because it shows intricate variation in substrate requirements and reproduction modes. Certain Micarea species are generalists and able to grow on various substrata, while some are specialized and live in strict microhabitats. Some of the Micarea species are predominately sexual, while some frequently lack sexual structures but bear numerous pycnidia where asexual conidia are produced. The actual roles of the three types of conidia present are not thoroughly understood, but mesoconidia are likely asexual propagules based on, for example, the observation that many of the species are frequently found with only mesopycnidia and no apothecia. Recent phylogenetic analyses together with ancestral state reconstruction among Micarea species showed that the shift in reproduction mode has evolved independently several times within the group and that facultative and obligate lignicoles are sister species.[9] This supports the assumption that the ancestor of these species was a facultative lignicole. It is presumed that a shift in substrate requirement from bark to wood leads to differentiation in reproduction mode and becomes a driver of speciation in Micarea microlichens. The case of Micarea is the first observation that among lichenized fungi reproduction mode is connected to substrate requirement.[9] It is also the first example where such an association is demonstrated to lead to lichen speciation. The main hypothesis behind this phenomenon is that obligate species on dead wood need to colonize new suitable substrata relatively fast and asexual reproduction is a more effective strategy for successful colonisation.[9]
Species
As of September 2021, Species Fungorum accepts 126 species of Micarea,[3] although this tally does not yet include some recently described taxa, such as four species from montane cloud forests of Kenya.[10]
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Micarea adnata Coppins (1983)[11]
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Micarea aeruginoprasina van den Boom, Guzow-Krzemińska, A.M.Brand & Sérusiaux (2019)[12]
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Micarea alabastrites (Nyl.) Coppins (1982)
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Micarea alectorialica M.Brand, van den Boom & Sérus. (2014)[13]
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Micarea amplissima van den Boom & Etayo (2023)[14]
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Micarea argopsinosa P.M.McCarthy & Elix (2016)[15]
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Micarea assimilata (Nyl.) Coppins (1983)[11]
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Micarea austroternaria Coppins & Kantvilas (1990)[16] – Australia
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Micarea azorica van den Boom, Guzow-Krzemińska, A.M.Brand & Sérusiaux (2019)[12]
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Micarea bacidiella (Vain.) Vězda (1976)
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Micarea bebourensis M.Brand, van den Boom & Sérus. (2014)[13]
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Micarea borbonica M.Brand, van den Boom & Sérus. (2014)[13]
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Micarea boryana M.Brand, van den Boom & Sérus. (2014)[13]
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Micarea botryoides (Nyl.) Coppins (1980)
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Micarea byssacea (Th.Fr.) Czarnota, Guzow-Krzem. & Coppins (2010)
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Micarea canariensis van den Boom (2013)[17]
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Micarea capitata M.Svenss. & G.Thor (2011)[18]
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Micarea ceracea Coppins & Kantvilas (2019)[19] – Australasia
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Micarea cilaosensis M.Brand, van den Boom & Sérus. (2014)[13]
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Micarea cinerea (Schaer.) Hedl. (1892)
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Micarea cinereopallida Coppins & Kantvilas (2019)[19] – Australasia; South America
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Micarea contexta Hedl. (1892)[20]
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Micarea coppinsii Tønsberg (1992)[21]
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Micarea corallothallina M.Cáceres, D.A.Mota & Aptroot (2013)
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Micarea coreana Lőkös, S.Y.Kondr. & Hur (2013)
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Micarea curvata Coppins (1983)[11]
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Micarea czarnotae Launis, van den Boom, Sérus. & Myllys (2019)[22]
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Micarea deminuta Coppins (1995)[23]
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Micarea denigrata (Fr.) Hedl. (1892)
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Micarea doliiformis (Coppins & P.James) Coppins & Sérus. (2010)
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Micarea elachista (Körb.) Coppins & R.Sant. (1983)[11]
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Micarea eucalypti P.M.McCarthy & Elix (2016)
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Micarea eximia Hedl. (1892)
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Micarea fallax Launis & Myllys (2019)[24]
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Micarea farinosa Coppins & Aptroot (2008)
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Micarea fennica Launis & Myllys (2019)[25]
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Micarea flagellispora Coppins & Kantvilas (1990)[16] – Australasia
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Micarea flavoleprosa Launis, Malíček & Sérus. (2019)[24]
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Micarea globulosella (Nyl.) Coppins (1983)[11]
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Micarea granuloblastidiata van den Boom (2017)[26] – Panama
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Micarea hedlundii Coppins (1983)[11]
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Micarea herbarum M.Brand, Coppins, Sérus. & van den Boom (2017)[27] – Europe
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Micarea humilis P.M.McCarthy & Elix (2018)[28] – Australia
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Micarea hyalinoxanthonica M.Brand, van den Boom & Sérus. (2014)[13]
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Micarea hylocomii Poelt & Döbbeler (1975)[29]
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Micarea hypoviolascens Czarnota & Coppins (2005)[30]
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Micarea incrassata Hedl. (1892)
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Micarea inquinans Tul.) Coppins (1992)
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Micarea intersociella (Stirt.) Coppins (1994)
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Micarea isabellina Coppins & Kantvilas (1990)[16] – Australia
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Micarea isidioprasina van den Boom, Guzow-Krzemińska, Sérusiaux, M.Kukwa (2019)[12]
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Micarea isidiosa M.Brand, van den Boom & Sérus. (2014)[13]
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Micarea kartana Kantvilas & Coppins (2018)[31] – Australia
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Micarea kemmleri Brackel (2016)[32]
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Micarea laeta Launis & Myllys (2019)[22]
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Micarea lapillicola (Vain.) Coppins & Muhr (1997)
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Micarea leprosula (Th.Fr.) Coppins & A.Fletcher (1975)
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Micarea levicula (Nyl.) Coppins (2001)
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Micarea lignaria (Ach.) Hedl. (1892)
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Micarea lithinella (Nyl.) Hedl. (1892)
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Micarea magellanica (Müll.Arg.) Fryday (2004)
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Micarea marginata Coppins & Muhr (1988)[33]
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Micarea melaena (Nyl.) Hedl. (1892)
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Micarea melaenida (Nyl.) Coppins (1983)[11]
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Micarea melanobola (Nyl.) Coppins (1983)[11]
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Micarea melanoprasina M.Brand, van den Boom & Sérus. (2014)[13]
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Micarea meridionalis van den Boom, M. Brand, Coppins & Sérus. (2017)[27] – Europe
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Micarea microareolata Launis, Pykälä & Myllys (2019)[22]
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Micarea micrococca (Körb.) Gams ex Coppins (2002)
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Micarea micromelaena Kantvilas & Coppins (2019)[19] – Australasia
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Micarea microsorediata van den Boom, Guzow-Krzemińska, Sérusiaux, M.Kukwa (2019)[12]
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Micarea minuta van den Boom, Guzow-Krzem. & Kukwa (2020)[34] – western Europe
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Micarea misella (Nyl.) Hedl. (1892)
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Micarea mutabilis Coppins & Kantvilas (1990)[16] – Australia
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Micarea myriocarpa V.Wirth & Vězda ex Coppins (1983)[11]
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Micarea neostipitata Coppins & P.F.May (2001)[35] – eastern North America
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Micarea nigella Coppins (1983)[11]
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Micarea nigra van den Boom, Guzow-Krzemińska, A.M.Brand & Sérusiaux (2019)[12]
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Micarea nigrata (Müll.Arg.) Kalb (1981)
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Micarea nitschkeana (J.Lahm ex Rabenh.) Harm. (1899)
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Micarea nowakii Czarnota & Coppins (2007)
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Micarea olivacea Coppins (1983)[11]
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Micarea oreina Kantvilas & Coppins (2019)[19] – Australasia
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Micarea pallida Coppins & Kantvilas (2019)[19] – Australasia
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Micarea pannarica Fryday (2004)[36]
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Micarea parasitica van den Boom & Etayo (2023)[14]
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Micarea paratropa (Nyl.) Alstrup (1994)
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Micarea parva Coppins (1995)[23]
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Micarea pauli Guzow-Krzemińska, A.Łubek & M.Kukwa (2019)[12]
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Micarea peliocarpa (Anzi) Coppins & R.Sant. (1979)
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Micarea poliocheila (Vain.) Aptroot (2002)
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Micarea polycarpella (Erichsen) Coppins & Palice (1995)
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Micarea prasina Fr. (1825)
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Micarea prasinastra Coppins & Kantvilas (2019)[19] – Australasia
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Micarea prasinella (Jatta) I.M.Lamb (1954)
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Micarea pseudocoppinsii M.Brand, van den Boom & Sérus. (2014)[13]
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Micarea pseudolignaria M.Brand, van den Boom & Sérus. (2014)[13]
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Micarea pseudomarginata Coppins (1989)
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Micarea pseudomicrococca Launis & Myllys (2019)[22]
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Micarea pseudotsugae van den Boom, Guzow-Krzem. & Kukwa (2020)[34] – western Europe
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Micarea pumila Kantelinen & Myllys (2021)[10] – Kenya
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Micarea pusilla Launis, Malíček & Myllys (2019)
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Micarea pycnidiophora Coppins & P.James (1979)
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Micarea rubiginosa Coppins & Kantvilas (2019)[19] – Australasia; South America
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Micarea sandyana Kantvilas (2019)[19] – Australasia
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Micarea saxicola Coppins & Kantvilas (2019)[19] – Australasia
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Micarea senecionis van den Boom (2010)
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Micarea sipmanii Sérus. & Coppins (2009)[37]
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Micarea soralifera Guzow-Krzem., Czarnota, Łubek & Kukwa (2016)
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Micarea squamulosa Aptroot, Lücking & M.Cáceres (2019)
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Micarea stellaris Kantelinen & Myllys (2021)[10] – Kenya
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Micarea stereocaulorum van den Boom & Etayo (2017)[27]
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Micarea stipitata Coppins & P.James (1979)[38]
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Micarea subalpina Coppins & T.Sprib. (2004)[39]
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Micarea subcinerea M.Brand & van den Boom (2004)
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Micarea subconfusa (Nyl.) Alstrup (1994)
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Micarea subgranulans (Vain.) Aptroot (2002)
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Micarea sublithinella M.Brand, van den Boom & Sérus. (2014)[13]
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Micarea submilliaria (Nyl.) Coppins (1994)
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Micarea subnigrata (Nyl.) Coppins & H.Kilias (1981)
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Micarea subternaria (Vain.) Aptroot (2002)
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Micarea subviridescens (Nyl.) Hedl. (1892)
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Micarea synotheoides (Nyl.) Coppins (1982)
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Micarea taitensis Kantelinen & Myllys (2021)[10] – Kenya
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Micarea takamakae M.Brand, van den Boom & Sérus. (2014)[13]
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Micarea tenuispora M.Brand, van den Boom & Sérus. (2014)[13]
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Micarea termitophila Aptroot & M.Cáceres (2014)
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Micarea ternaria (Nyl.) Vězda (1970)
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Micarea tomentosa Czarnota & Coppins (2007)[40]
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Micarea tubaeformis Coppins & Kantvilas (2019)[19] – Australasia
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Micarea turfosa (A.Massal.) Du Rietz (1923)
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Micarea usneae van den Boom & Ertz (2014)[41] – Madeira
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Micarea versicolor Kantelinen, Hyvärinen & Myllys (2021)[10] – Kenya
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Micarea viridiatra Coppins (1985)[42] – Europe
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Micarea viridicapitata Córd.-Cháv., Aptroot & R.-E.Pérez (2014)[43] – Mexico
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Micarea viridileprosa Coppins & van den Boom (2001)[44] – western Europe
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Micarea vulpinaris (Nyl.) Muhr (1987)
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Micarea xanthonica Coppins & Tønsberg (2001)[45] – Europe; North America
References
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^ "Synonymy: Micarea Fr., Syst. orb. veg. (Lundae) 1: 256 (1825)". Species Fungorum. Retrieved 12 May 2021.
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^ Wijayawardene, Nalin; Hyde, Kevin; Al-Ani, Laith Khalil Tawfeeq; Somayeh, Dolatabadi; Stadler, Marc; Haelewaters, Danny; et al. (2020). "Outline of Fungi and fungus-like taxa". Mycosphere. 11: 1060–1456. doi:10.5943/mycosphere/11/1/8.
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^ a b Andersen, Heidi L.; Ekman, Stefan (2005). "Disintegration of the Micareaceae (lichenized Ascomycota): a molecular phylogeny based on mitochondrial rDNA sequences". Mycological Research. 109 (1): 21–30. doi:10.1017/S0953756204001625.
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^ Fries, Elias M. (1825). Systema Orbis Vegetabilis (in Latin). Vol. 1. Lund: Typographia Academica. p. 256.
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^ Jørgensen, Per M. (1984). "Coppins, B. J. 1983. A taxonomic study of the lichen genus Micarea in Europe. – Bull". Nordic Journal of Botany. 4 (4): 544. doi:10.1111/j.1756-1051.1984.tb02061.x.
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^ Coppins, B.J. (1989). "(945) Proposal to conserve Micarea Fries (Dec. 1825) against Micarea Fries (pre-May 1825) (Fungi)". Taxon. 38 (3): 499–501. doi:10.2307/1222305. JSTOR 1222305.
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^ "Index Fungorum - Names Record - Micarea". Index Fungorum. Retrieved 12 May 2021.
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^ a b c Kantelinen, Annina; Prinzen, Christian; Poczai, Péter; Myllys, Leena (2022). "Lichen speciation is sparked by a substrate requirement shift and reproduction mode differentiation". Scientific Reports. 12: 11048. doi:10.1038/s41598-022-14970-9.
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^ a b c d e Kantelinen, Annina; Hyvärinen, Marko T.; Kirika, Paul; Myllys, Leena (2021). "Four new Micarea species from the montane cloud forests of Taita Hills, Kenya". The Lichenologist. 53 (1): 81–94. doi:10.1017/S0024282920000511. hdl:10138/329202.
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^ a b c d e f Guzow-Krzemińska, Beata; Sérusiaux, Emmanuël; van den Boom, Pieter P.G.; Brand, A. Maarten; Launis, Annina; Łubek, Anna; Kukwa, Martin (2019). "Understanding the evolution of phenotypical characters in the Micarea prasina group (Pilocarpaceae) and descriptions of six new species within the group". MycoKeys. 57: 1–30. doi:10.3897/mycokeys.57.33267.
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^ a b c d e f g h i j k l m Brand, A. Maarten; van Den Boom, Pieter P. G.; Sérusiaux, Emmanuël (2014). "Unveiling a surprising diversity in the lichen genus Micarea (Pilocarpaceae) in Réunion (Mascarenes archipelago, Indian Ocean)". The Lichenologist. 46 (3): 413–439. doi:10.1017/S0024282913000911.
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^ a b van den Boom, Pieter; Etayo, Javier; de Silanes, M. Eugenia López (2023). "Notes on lichenicolous Micarea species in Spain and Macaronesia, with the descriptions of two new species". Nova Acta Científica Compostelana. doi:10.15304/nacc.id8595. ISSN 2340-0021.
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^ Hedlund, T. (1892). "Kritische Bemerkungen über einige Arten der. Flechtengattungen Lecanora (Ach.), Lecidea (Ach.) und Micarea (Fr.)". Bihang til Kongliga Svenska Vetenskaps-Akademiens Handlingar (in German). 18 (3): 83, 96.
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^ a b Coppins, B.J. (1995). "Two new, diminutive Micarea species from Western Europe". Bibliotheca Lichenologica. 58: 57–62.
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^ a b Launis, Annina; Malíček, Jiří; Svensson, Måns; Tsurykau, Andrei; Sérusiaux, Emmanuël; Myllys, Leena (2019). "Sharpening species boundaries in the Micarea prasina group, with a new circumscription of the type species M. prasina". Mycologia. 111 (4): 574–592. doi:10.1080/00275514.2019.1603044. hdl:10138/326192.
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^ Launis, Annina; Myllys, Leena (2019). "Micarea fennica, a new lignicolous lichen species from Finland". Phytotaxa. 409 (3): 179–188. doi:10.11646/phytotaxa.409.3.6.
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^ van den Boom, Pieter P.G.; Sipman, Harrie J.M.; Divakar, Pradeep K.; Ertz, Damien (2017). "New or interesting records of lichens and lichenicolous fungi from Panama, with descriptions of ten new species". Sydowia. 69: 47–72. doi:10.12905/0380.sydowia69-2017-0047.
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^ a b c van den Boom, Pieter P.G.; Brand, A. Maarten; Coppins, Brian John; Sérusiaux, Emmanuel (2017). "Two new species in the Micarea prasina group from Western Europe". The Lichenologist. 49 (1): 13–25. doi:10.1017/S0024282916000633.
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^ Poelt, J.; Döbbeler, P. (1975). "Über moosparasitische Arten der Flechtengattung Micarea und Vezdaea". Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie. 96 (1–4): 328–352.
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^ Kantvilas, G. (2018). "Micarea kartana sp. nov. (lichenised Ascomycetes) from Kangaroo Island, South Australia". Swainsona. 31 (4): 55–58.
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^ a b van den Boom, Pieter; Guzow-Krzemińska, Beata; Kukwa, Martin (2020). "Two new Micarea species (Pilocarpaceae) from Western Europe". Plant and Fungal Systematics: 189–199. doi:10.35535/pfsyst-2020-0014.
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^ Coppins, B.J.; May, P.F. (2007). "Micarea neostipitata, a new species with pale stipitate pycnidia from eastern North America". The Lichenologist. 33 (6): 487–490. doi:10.1006/lich.2001.0350.
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^ Fryday, A.M. (2004). "New species and records of lichenized fungi from Campbell Island and the Auckland Islands, New Zealand". Bibliotheca Lichenologica. 88: 127–146.
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^ Sérusiaux, E.; Coppins, B.J. (2009). "Micarea sipmanii, a new species with arbuscular pycnidia from the West Indies". Bibliotheca Lichenologica. 99: 367–372.
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^ Coppins, B.J.; James, P.W. (2007). "New or interesting British lichens IV". The Lichenologist. 11 (2): 139–179. doi:10.1017/S0024282979000190.
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^ Coppins, Brian J.; Spribille, Toby (2004). "Micarea subalpina Coppins & Spribille, a new subalpine species from the Rocky Mountains, USA". The Lichenologist. 36 (2): 97–102. doi:10.1017/S0024282904014070.
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^ Czarnota, P. (2007). "The lichen genus Micarea (Lecanorales, Ascomycota) in Poland". Polish Botanical Studies. 23: 174.
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^ van den Boom, Pieter P.G.; Ertz, Damien (2014). "A new species of Micarea (Pilocarpaceae) from Madeira growing on Usnea". The Lichenologist. 46: 295–301. doi:10.1017/S0024282913000698.
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^ Coppins, B.J. (1985). "A new Micarea from the Scottish Highlands". The Lichenologist. 17 (1): 99–101.
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^ CórdovaÇhávez, Octavio; Aptroot, André; Castillo-Campos, Gonzalo; Cáceres, Marcela E.S; Pérez-Pérez, Rosa Emilia (2014). "Three new lichen species from cloud forest in Veracruz, Mexico". Cryptogamie, Mycologie. 35: 157–162. doi:10.7872/crym.v35.iss2.2014.157.
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^ van den Boom, P.P.G.; Coppins, B.J. (2001). "Micarea viridileprosa sp. nov., an overlooked lichen species from western Europe". The Lichenologist. 33 (2): 87–91. doi:10.1006/lich.2000.0310.
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^ Coppins, B.J.; Tønsberg, T. (2001). "A new xanthone-containing Micarea from northwest Europe and the Pacific Northwest of North America". The Lichenologist. 33 (2): 93–96. doi:10.1006/lich.2000.0311.