The genus Cheiromycina is one of the few genera of lichenized hyphomycetes for which no sexual reproductive stages have been observed. The genus includes species from boreal to temperate regions of the Northern Hemisphere where it is found growing on bark or wood. Congeners in Cheiromycina are characterized by a noncorticate thallus, nearly immersed in the substrate and presenting powdery unpigmented sporodochia, and containing chlorococcoid photobionts. The relationships of members of Cheiromycina with other fungi are not known. Here we inferred the phylogenetic placement of Cheiromycina using three loci (nuSSU, nuLSU, and mtSSU) representing C. flabelliformis, the type species for the genus, C. petri, and C. reimeri. Our results revealed that the genus Cheiromycina is found within the family Malmideaceae (Lecanorales) where members formed a monophyletic clade sister to the genera Savoronala and Malmidea. This phylogenetic placement and the relationships of Cheiromycina with other lichenized hyphomycetous taxa are here discussed.

b Institute of Environmental Sciences CML Leiden University Einsteinweg 2 2333 CC Leiden The Netherlands

c Department of Natural History University Museum University of Bergen Allegt 41 7800 N 5020 Bergen Norway

Department of Life Sciences University of Trieste via Giorgieri 10 34127 Trieste Italy

Department of Plant Taxonomy and Nature Conservation University of Gdańsk Wita Stwosza 59 80 308 Gdańsk Poland

e Institute of Botany Academy of Science of Czech Republic Zámek 1 CZ 25243 Průhonice Czech Republic

f Department of Botany Faculty of Natural Sciences Charles University Benátská 2 CZ 12801 Praha Czech Republic

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Altschul SF, Gish W, Miller W, Myers EW, Lipman DJ.. 1990. Basic local alignment search tool. The Journal of Molecular Biology 215:403–410. PubMed

An K-D, Degawa Y, Fujihara E, Mikawa T, Ohkuma M, Okada G.. 2012. Molecular phylogenetic analyses based on the nuclear rRNA genes and the intron-exon structures of the nuSSU rRNA gene in PubMed

Aptroot A, Schiefelbein U.. 2003. Additional species of PubMed

Aptroot A, Sipman HJM.. 2011.

Berbee ML, Taylor JW.. 1993. Ascomycete relationships: dating the origin of asexual lineages with 18S ribosomal RNA gene sequence data In: Reynolds DR, Taylor JW, eds. The fungal holomorph: mitotic, meiotic, and pleomorphic speciation in fungal systematics. Wallingford, UK: CAB International; p. 68–77.

MES Cáceres, Aptroot A.. 2016. First inventory of lichens from the Brazilian Amazon in Amapá State. The Bryologist 119:250–265.

MES Cáceres, Aptroot A, Oliveira Mendoca C, LA dos Santos, Lücking R.. 2017.

Cai L, Guo XY, Hyde KD.. 2008. Morphological and molecular characterization of a new anamorpjhic genus PubMed PMC

Crous PW, Groenewald JZ, Mansilla JP, Hunter GC, Wingfield MJ.. 2004. Phylogenetic reassessment of PubMed PMC

Crous PW, Kang JC, Braun U.. 2001. A phylogenetic redefinition of anamorph genera in

Crous PW, Wingfield MJ, Mansilla JP, Alfenas AC, Groenewald JZ.. 2006. Phylogenetic reassessment of PubMed PMC

Cubero OF, Crespo A, Fatehi J, Bridge PD.. 1999. DNA extraction and PCR amplification method suitable for fresh, herbarium stored and lichenized fungi. Plant Systematics and Evolution 217:243–249.

Darriba D, Taboada GL, Doallo R, Posada D.. 2012. jModelTest 2: more models, new heuristics and parallel computing. Natural Methods 9: article 772. PubMed PMC

Dettman JR, Jacobson DJ, Taylor JW.. 2003. A multilocus genealogical approach to phylogenetic species recognition in the model eukaryote PubMed

Diederich P. 2011. Lichenicolous hyphomycete genera In: Seifert K, Morgan-Jones G, Gams W, Kendrick B, eds. The genera of Hyphomycetes. CBS Biodiversity Series 9. Utrecht, The Netherlands: CBS-KNAW Fungal Biodiversity Center p. 917–920.

Diederich P. 2013. A redescription of

Diederich P, Coppins BJ.. 2009.

Diederich P, Ertz D, Lawrey JD, Sikaroodi M, Untereiner WA.. 2013. Molecular data place the hyphomycetous lichenicolous genus

Diederich P, Palice Z, Ertz D.. 2008.

Diederich P, Scheidegger C.. 1996.

Earland-Bennett PM, Hawksworth DL.. 2005. The first lichen-forming species of

Ekman S. 1996. The corticolous and lignicolous species of

Ekman S, Tønsberg T.. 2002. Most species of

Elix JA, Tønsberg T.. 2004. Notes on the chemistry of some lichens, including four species of

Ertz D, Bungartz F, Diederich P, Tibell L.. 2011. Molecular and morphological data place

Ertz D, Fischer E, Killmann D, Razafindrahaja T, Sérusiaux E.. 2013.

Ertz D, Lawrey JD, Common RS, Diederich P.. 2014. Molecular data resolve a new order of Arthoniomycetes sister to the primarily lichenized Arthoniales and composed of black yeasts, lichenicolous and rock-inhabiting species. Fungal Diversity 66:113–137.

Ertz D, Tehler A.. 2011. The phylogeny of Arthoniales (Pezizomycotina) inferred from nucLSU and RPB2 sequences. Fungal Diversity 49:47–71.

Etayo J. 1995. Two new species of lichenicolous fungi from the Pyrenees. Nova Hedwigia 61:189–197.

Fehrer J, Š Bayerová, Orange A.. 2008. Large genetic divergence of new, morphologically similar species of sterile lichens from Europe ( PubMed

Flakus A, Elix JA, Rodriguez P, Kukwa M.. 2011. New species and records of

Frisch A, Thor G, Sheil D.. 2014. Four new Arthoniomycetes from Bwindi Impenetrable National Park, Uganda—supported by molecular data. Nova Hedwigia 98:295–312.

Fryday AM. 2001. The lichen vegetation associated with areas of late snow-lie in the Scottish Highlands. The Lichenologist 33:121–150.

Fryday AM, Schmitt I, Perez-Ortega S.. 2017. The genus

Gargas A, Taylor JW.. 1992. Polymerase chain reaction (PCR) primers for amplifying, sequencing nuclear 18S rDNA from lichenized fungi. Mycologia 84:589–592.

Hafellner J. 2012. Lichenicolous Biota (Nos 101–120). Fritschiana 74:1–17.

Hall TA. 1999. BioEdit: a user friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nucleic Acid Symposia Series 41:95–98.

Harris RC, Ladd D.. 2007. New taxa of lichens and lichenicolous fungi from the Ozark Ecoregion. Opuscula Philolichenum 4:57–68.

Hawksworth DL. 1979. The lichenicolous hyphomycetes. Bulletin of the British Museum (Natural History), Botany 6:183–300.

Hawksworth DL, Coppins BJ, James PW.. 1979.

Hawksworth DL, Poelt J.. 1986. Five additional genera of conidial lichen-forming fungi from Europe. Plant Systematics and Evolution 154:195–211.

Hawksworth DL, Poelt J.. 1990. A second lichen-forming species of

Heuchert B, Braun U.. 2006. On some dematiaceous lichenicolous hyphomycetes. Herzogia 19:11–21.Huelsenbeck JP, Ronquist F. 2003. MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics 19:1572e1574. PubMed

Hughes SJ. 1953. Conidiophores, conidia and classification. Canadian Journal of Botany 31:577–657.

Hughes SJ. 1958. Revisiones Hyphomycetum aliquot cum appendice de nominibus rejiciendis. Canadian Journal of Botany 36:727–836.

Huhndorf SM, Fernández FA.. 2005. Teleomorph-anamorph connections:

Kalb K, Rivas Plata E, Lücking R, Lumbsch HT.. 2011. The phylogenetic position of

Kukwa M. 2006. The lichen genus

Lendemer JC. 2013. A monograph of the crustose members of the genus

Lendemer JC, Harris RC.. 2004. A checklist of the lichens collected on the 28th A. Leroy Andrews Foray. Evansia 21:88–100.

Lendemer JC, Harris RC.. 2015.

Lendemer JC, Hodkinson BP.. 2012.

Liu ZY, Yao YJ, Liang ZQ, Liu AY, Pegler DN, Chase MW.. 2001. Molecular evidence for the anamorph—teleomorph connection in

Lizel M, Crous PW, Groenewald JZE, Gams W, Summerbell C.. 2003. PubMed

Lücking R, Hodkinson BP, Leavitt SD.. 2017. The 2016 classification of lichenized fungi in the Ascomycota and Basidiomycota—approaching one thousand genera. The Bryologist 119:361–416.

Mason-Gamer RJ, Kellogg EA.. 1996. Testing for phylogenetic conflict among molecular data sets in the tribe Triticeae (Gramineae). Systematic Biology 45:524–545.

Mercado Sierra A, Calduch M, Gené J, Guarro J, Delgado G.. 2003. PubMed

Mercado Sierra A, Mena Portales J.. 1988. Nuevos o raros hifomicetes de Cuba. B. Un nuevo género sobre

Miadlikowska J, Kauff F, Högnabba F, Oliver JC, Molnár K, Fraker E, Otálora MA.. 2014. A multigene phylogenetic synthesis for the class Lecanoromycetes (Ascomycota): 1307 fungi representing 1139 infrageneric taxa, 317 genera and 66 families. Molecular Phylogenetics and Evolution 79:132–168. PubMed PMC

Morris EF, Finley DE.. 1967. Two new genera of stilbellaceous fungi. The American Midland Naturalist 77:200–204.

Muggia L, Baloch E, Stabentheiner E, Grube M, Wedin M.. 2011. Photobiont association and genetic diversity of the optionally lichenized fungus PubMed

Muggia L, Fernandez Brime S, Grube M, Wedin M.. 2016. PubMed

Muggia L, Pérez-Ortega S, Fryday A, Spribille T, Grube M.. 2014. Global assessment of genetic variation and phenotypic plasticity in the lichen-forming species

Nelsen MP, Gargas A.. 2008. Phylogenetic distribution and evolution of secondary metabolites in the lichenized fungal genus

Page RDM. 1996. TREEVIEW: an application to display phylogenetic trees on personal computers. Computational Application in Bioscience 12:357–358. PubMed

Pérez-Ortega S, Suija A, de Los Rios A.. 2011. The connection between

Posada D, Crandall KA.. 1998. Modeltest—testing the model of DNA substitution. Bioinformatics 14:817–818. PubMed

Printzen C. 2007. New records of

Rambaut A, Drummond A. Available from: beast.bio.ed.ac.uk/Tracer. 2007. Tracer. [cited 2007 Oct 11]

Réblová M, Seifert KA.. 2004.

Réblová M, Seifert KA.. 2011. Discovery of the teleomorph of the hyphomycete, PubMed PMC

Reeb V, Lutzoni F, Roux C.. 2004. Contribution of RPB2 to multilocus phylogenetic studies of the euascomycetes (Pezizomycotina, Fungi) with special emphasis on the lichen-forming Acarosporaceae and evolution of polyspory. Molecular Phylogenetics and Evolution 32:1036–1060. PubMed

Reynolds DR, Taylor JW.. 1993. The fungal holomorph: mitotic, meiotic and pleomorphic speciation in fungal systematics. Oxon, UK: CAB International.

Ronquist F, Huelsenbeck JP, Van der Mark P.. 2005. MrBayes 3.1 manual. [cited 2005 May 26]. Available from: http://mrbayes.csit.fsu.edu/mb3.1_manual.pdf

Saccardo PA. 1886.

Seifert KA, Kendrick B, Morgan-Jones G, Gams W.. 2011. The genera of Hyphomycetes. Utrecht, The Netherlands: CBS-KNAW Fungal Biodiversity Centre. 997 p.

Seifert KA, Okada G, Rao V.. 1987. The synnematous hyphomycete

Seymour F, Crittenden P, Dyer P.. 2005. Sex in the extremes: lichen-forming fungi. Mycologist 19:51–58.

Shenoy BD, Jeewon R, Hyde KD.. 2007. Impact of DNA sequence-data on the taxonomy of anamorphic fungi. Fungal Diversity 26:1–54.

Smith CW. 2009.

Sodamuk M, Boonpragob K, Mongkolsuk P, Tehler A, Leavitt SD, Lumbsch HT.. 2017.

Stamatakis A, Ludwig T, Meier H.. 2005. RAxML-iii: a fast program for maximum likelihood-based inference of large phylogenetic trees. Bioinformatics 21:456–463. PubMed

Sutton BC. 1985. Notes on some deuteromycete genera with cheiroid or digitate brown conidia. Proceeding of the Indian Academy of Science (Plant Science) 94:229–244.

Sutton BC, Carmaran CC, Romero AI. 1996.

Sutton BC, Muhr L-E.. 1986.

Vilgalys R, Hester M.. 1990. Rapid genetic identification and mapping of enzymatically amplified ribosomal DNA from several PubMed PMC

Vobis G, Hawksworth DL.. 1981. Conidial lichen-forming fungi In: Cole GT, Kandrick WB, eds. The biology of conidial fungi 1. New York: Academic Press; p. 245–273.

Wedin M, Döring H, Gilenstam G.. 2004. Saprotrophy and lichenization as options for the same fungal species on different substrata: environmental plasticity and fungal lifestyles in the

Wedin M, Döring H, Gilenstam G.. 2006. PubMed

White TJ, Burns TD, Lee S, Taylor J.. 1990. Amplification and direct sequencing of fungal ribosomal DNA genes for phylogenies In: Innis MA, Gelfand DH, Snisky JJ, White TJ, eds. PCR protocols, a guide to methods and applications. San Diego, California: Academic Press; p. 315–322.

Zhou S, Stanosz GR.. 2001. Primers for amplification of mtSSU rDNA, and a phylogenetic study of

Zhurbenko MP, Braun U, Heuchert B, Kobzeva AA.. 2015. New lichenicolous hyphomycetes from Eurasia. Herzogia 28:584–598.

Zoller S, Scheidegger C, Sperisen C.. 1999. PCR primers for the amplification of mitochondrial small subunit ribosomal DNA of lichen-forming ascomycetes. The Lichenologist 31:511–516.