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Rediscovery of pycnidia in Thamnolia vermicularis: implications for chemotype occurrence and distribution

Published online by Cambridge University Press:  07 May 2013

Janice M. LORD
Department of Botany, University of Otago, PO Box 56, Dunedin 9054, New Zealand. Email:
Allison KNIGHT
Department of Botany, University of Otago, PO Box 56, Dunedin 9054, New Zealand. Email:
Department of Botany, University of Otago, PO Box 56, Dunedin 9054, New Zealand. Email:
Department of Botany, University of Otago, PO Box 56, Dunedin 9054, New Zealand. Email:
William M. MALCOLM
PO Box 320, Nelson 7040, New Zealand
Department of Botany, University of Otago, PO Box 56, Dunedin 9054, New Zealand. Email:


Thamnolia vermicularis is a globally widespread arctic-alpine lichen with two distinct chemotypes. The intermixing of these chemotypes at the local scale and lack of genetic variation between them is puzzling given the apparent absence of specialized reproductive structures. Apothecia and conidiomata were originally reported for Thamnolia in the second half of the 19th century, but putative apothecia proved to be parasitic in origin. This study presents conclusive microscopic evidence for the existence of pycnidial conidiomata containing conidia on thalli of Thamnolia vermicularis from geographically widespread locations. Sequences of the internal transcribed spacer (ITS) region of ribosomal DNA from excised pycnidial conidiomata matched that of thallus tips from individual Thamnolia thalli. Our examination of the historic literature found that the occurrence of pycnidial conidiomata was never disproved, but this information was lost from the post-1920s English-language literature. Our rediscovery of pycnidial conidiomata indicates that Thamnolia vermicularis possesses a reproductive strategy with the potential for long-distance dispersal of multitudes of mitotically produced conidia, providing a plausible explanation for the cosmopolitan distribution of the species and the low levels of genetic variation among populations. Reversible processes, for example alternative splicing, warrant consideration as explanations for chemotype intermixing.

Copyright © British Lichen Society 2013

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Abbayes, H. des (1951) Traite de Lichenologie. Paris: Paul Lechevalier.Google Scholar
Anders, J. (1928) Die Strauch- und Laubflechten Mitteleuropas. Anleitung zum Bestimmen der in Mitteleuropa vorkommenden Strauch- und Laubflechten. Jena: Fischer.Google Scholar
Armaleo, D., Zhang, Y. & Cheung, S. (2008) Light might regulate divergently depside and depsidone accumulation in the lichen Parmotrema hypotropum by affecting thallus temperature and water potential. Mycologia 100: 565576.CrossRefGoogle ScholarPubMed
Arnold, F. (1874) Lichenologische Fragmente XVI. Flora (Regensburg) 57: 97110. Google Scholar
Asahina, Y. (1937) Lichenologische Notizen IX. Journal of Japanese Botany 13: 315321.Google Scholar
Bagley, S. J. & Orlovich, D. A. (2004) Genet size and distribution of Amanita muscaria in a suburban park, Dunedin, New Zealand. New Zealand Journal of Botany 42: 939947.CrossRefGoogle Scholar
Brodo, I. M., Duran Sharnoff, S. & Sharnoff, S. (2001) Lichens of North America. New Haven & London: Yale University Press.Google Scholar
Cassie, D. M. & Piercey-Normore, M. D. (2008) Dispersal in a sterile lichen-forming fungus, Thamnolia subuliformis (Ascomycotina: Icmadophilaceae). Botany 86: 751762.CrossRefGoogle Scholar
Choisy, M. (1955) Sur le Thamnolia vermicularis et la systematique de ce genre. Bulletin Mensuel de la Société Linnéenne de Lyon 24: 245248.CrossRefGoogle Scholar
Crombie, J. M. (1894) A Monograph of Lichens Found in Britain: Being a Descriptive Catalogue of the Species of the British Museum. Part I. London: British Museum. Google Scholar
Culberson, C. F. (1963) The lichen genus Thamnolia . Brittonia 15: 140144.CrossRefGoogle Scholar
Culberson, C. F. & Elix, J. A. (1989) Lichen substances. In Methods in Plant Biochemistry. Vol. 1. Plant Phenolics (Dey, P. M. & Harborne, J. B., eds): 509535. London: Academic Press.Google Scholar
Dobson, F. S. (2011) Lichens: An Illustrated Guide to the British and Irish Species, Sixth Edition. Slough: The Richmond Publishing Co. Google Scholar
Dodge, C. W. (1973) Lichen Flora of the Antarctic Continent and Adjacent Islands. Canaan, New Hampshire: Phoenix Publishing Co. Google Scholar
Drummond, A. J., Ashton, B., Buxton, S., Cheung, M., Cooper, A., Heled, J., Kearse, M., Moir, R., Stones-Havas, S., Sturrock, S., et al. (2011) Geneious v. 5.5. Available from Google Scholar
Elix, J. A. (1996) Biochemistry and secondary metabolites. In Lichen Biology (Nash, T. H. III, ed.): 154180. Cambridge: Cambridge University Press.Google Scholar
Elix, J. A. & Gaul, K. L. (1986) The interconversion of the lichen depsides para- and meta-scrobiculin, and the biosynthetic implications. Australian Journal of Chemistry 39: 613624.CrossRefGoogle Scholar
Elix, J. A., Jenie, U. A. & Parker, J. L. (1987) A novel synthesis of the lichen depsidones divaronic acid and stenosporonic acid, and the biosynthetic implications. Australian Journal of Chemistry 40: 14511464.CrossRefGoogle Scholar
Filson, R. B. (1972) Studies in Australasian lichens II. The alpine lichen Thamnolia vermicularis (Sw.) Schaer. in Australia. Muelleria 2: 180187.CrossRefGoogle Scholar
Fletcher, A. (1975) Key for the identification of British marine and maritime lichens I. Siliceous rocky shore species. Lichenologist 7: 152.CrossRefGoogle Scholar
Fries, T. M. (1860) Lichenes Arctoi Europae Groenlandiaeque Hactenus Cogniti. Upsaliae: C. A. Leffler. Google Scholar
Fries, T. M. (1867) Lichenes Spitsbergenses. Kongliga Svenska Vetenskaps-Akademiens Handlingar 7: 153. Google Scholar
Galloway, D. J. (1985) Flora of New Zealand Lichens. Wellington: Government Printer.Google Scholar
Galloway, D. J. (1991) Phytogeography of Southern Hemisphere lichens. In Quantitative Approaches to Phytogeography (Nimis, P. L. & Crovello, T. J., eds): 233262. Dordrecht: Kluwer Academic Publishers.CrossRefGoogle Scholar
Galloway, D. J. (1996) Lichen biogeography. In Lichen Biology (Nash, T. H. III, ed.): 199216. Cambridge: Cambridge University Press.Google Scholar
Galloway, D. J. (2007) Flora of New Zealand Lichens, Revised Second Edition: Including Lichen-forming and Lichenicolous Fungi. Lincoln, New Zealand: Manaaki Whenua Press.Google Scholar
Galloway, D. J. & Aptroot, A. (1995) Bipolar lichens: a review. Cryptogamic Botany 5: 184191.Google Scholar
Gardes, M. & Bruns, T. D. (1993) ITS primers with enhanced specificity for basidiomycetes – application to the identification of mycorrhizae and rusts. Molecular Ecology 2: 113118.CrossRefGoogle Scholar
Gargas, A., DePriest, P. T. & Taylor, J. W. (1995) Positions of multiple insertions in SSU rDNA of lichen-forming fungi. Molecular Biology and Evolution 12: 208218.Google ScholarPubMed
Gierl, C. & Kalb, K. (1993) Die Flechtengattung Dibaeis. Eine Übersicht über die rosafrüchtigen Arten von Baeomyces sens. lat. nebst Anmerkungen zu Phyllobaeis gen. nov. Herzogia 9: 593645.CrossRefGoogle Scholar
Glück, H. (1899) Entwurf zu einer vergleichenden Morphologie der Flechtenspermogonien. Verhandlungen des Naturhistorisch-Medizinischen Vereins zu Heidelberg N.F. 6: 81216. Google Scholar
Golubkova, N. S., Savicz, V. P. & Trass, H. H. (1978) Handbook of the Lichens of the USSR. 5. Cladoniaceae Acarosporaceae. Leningrad: Nauka.Google Scholar
Grube, M. & Kantvilas, G. (2006) Siphula represents a remarkable case of morphological convergence in sterile lichens. Lichenologist 38: 241249.CrossRefGoogle Scholar
Grube, M., Gargas, A. & DePriest, P. T. (1996) A small insertion in the SSU rDNA of the lichen fungus Arthonia lapidicola is a degenerate group-I intron. Current Genetics 29: 582586.CrossRefGoogle ScholarPubMed
Hale, M. E. Jr. (1979) How to Know the Lichens. 2nd edn. Dubuque, Iowa: Brown.Google Scholar
Hawksworth, D. L. (1980) Notes on British lichenicolous fungi: III. Notes from the Royal Botanic Garden Edinburgh 38: 165183.Google Scholar
Hawksworth, D. L. (1988) Conidiomata, conidiogenesis, and conidia. In CRC Handbook of Lichenology, Volume I (Galun, M., ed.): 181193. Boca Raton: CRC Press, Inc. Google Scholar
Henssen, A. & Jahns, H. M. (1974) Lichenes. Eine Einführung in die Flechtenkunde. Stuttgart: Thieme.Google Scholar
Honegger, R. & Scherrer, S. (2008) Sexual reproduction in lichen-forming ascomycetes. In Lichen Biology. 2nd edn. (Nash, T. H. III, ed.): 94103. Cambridge: Cambridge University Press.CrossRefGoogle Scholar
Ihlen, P. G. (1995) The lichenicolous fungi on Thamnolia vermicularis in Norway. Graphis Scripta 7: 1724.Google Scholar
Jahns, H. M. (1988) The lichen thallus. In CRC Handbook of Lichenology. Volume I (Galun, M., ed.): 95146. Boca Raton: CRC Press, Inc. Google Scholar
Jahns, H. M. & Schuster, G. (1981) Morphogenetische untersuchungen an Cetraria islandica . Beitrage zur Biologie der Pflanzen 55: 427473.Google Scholar
Jatta, A. (1911) Lichenes. Flora Italica Cryptogama, Firenze. Pars III, Fasc. 4: 461694. Google Scholar
Kantvilas, G. (1998) Studies on the lichen genus Siphula in Tasmania II. The S. decumbens group. Herzogia 13: 119138.CrossRefGoogle Scholar
Kantvilas, G. (2002) Studies on the lichen genus Siphula Fr. Bibliotheca Lichenologica 82: 3753.Google Scholar
Kärnefelt, E. I. & Thell, A. (1995) Genotypical variation and reproduction in natural populations of Thamnolia . Bibliotheca Lichenologica 58: 213243.Google Scholar
Keissler, K. v. (1960) Usneaceae. In Rabenhorst's Kryptogamen-Flora von Deutschland, Österreich und der Schweiz. 2. Aufl., Band 9, 5. Abt., 4. Teil. Leipzig: Akademische Verlagsgesellschaft.Google Scholar
Koerber, G. W. (1855) Systema Lichenum Germaniae (I–XXXIV). Die Flechten Deutschlands Mikroscopisch Geprüft, Kritisch Gesichtet, Charakteristisch Beschrieben und Systematisch Geordnet. Breslau: Verlag Trevendt & Granier. Google Scholar
Krempelhuber, A. von (1869) Geschichte und Litteratur der Lichenologie von den Ältesten Zeiten bis zum Schlusse des Jahres 1865, i–iv. München: Selbstverlag. Google Scholar
Krog, H., Østhagen, H. & Tønsberg, T. (1994) Lavflora. Norske busk- og bladlav. Oslo: Universitetsforlaget.Google Scholar
Lambley, P. W. & Purvis, O. W. (2009) Thamnolia. In The Lichens of Great Britain and Ireland (Smith, C. W., Aptroot, A., Coppins, B. J., Fletcher, A., Gilbert, O. L., James, P. W. & Wolseley, P. A., eds): 877. London: British Lichen Society.Google Scholar
Larkin, M. A., Blackshields, G., Brown, N. P., Chenna, R., McGettigan, P. A., McWilliam, H., Valentin, F., Wallace, I. M., Wilm, A., Lopez, R., et al. (2007) Clustal W and Clustal X version 2.0. Bioinformatics 23: 29472948.CrossRefGoogle ScholarPubMed
Leighton, W. A. (1879) The Lichen-Flora of Great Britain, Ireland and the Channel Islands, 3rd edn. Shrewsbury: published by author. CrossRefGoogle Scholar
Lindau, G. (1923) Die Flechten. 2. Aufl. Kryptogamenflora für Anfänger, III. Berlin: Springer. CrossRefGoogle Scholar
Lindsay, W. L. (1859) Memoir on the Spermogones and Pycnides of Filamentous, Fruticulose and Foliaceous Lichens. Transactions of the Royal Society of Edinburgh 22: 101303. CrossRefGoogle Scholar
Lücking, R. & Santesson, R. (2002) On the identity of Pyrenotrichum ‘atrocyaneum’, P. ‘mirum’ and P. ‘podosphaera’, Campylidia of lichenized Ascomycota (Lecanorales: Ectolechiaceae). Bryologist 105: 5762.CrossRefGoogle Scholar
Ludwig, L. [R.] (2011) Marginal soralia and conidiomata in Icmadophila splachnirima (Icmadophilaceae) from southern New Zealand. Australasian Lichenology 68: 411.Google Scholar
Malcolm, N. & Malcolm, W. M. (2000) New Zealand Lichens. Nelson, New Zealand: Micro-Optics Press.Google Scholar
Massalongo, A. B. (1856) De Thamnolia enere lichenum nodum rite definito breve commentorium. Flora 39: 231235. Google Scholar
Migula, W. (1929) Kryptogamenflora von Deutschland, Österreich und der Schweiz, Band IV: Flechten, 1. Teil. Berlin-Lichterfelde: Hugo Bermühler Verlag. Google Scholar
Minks, A. (1874) Thamnolia vermicularis, eine Monographie. Flora 57: 337361. Google Scholar
Motyka, J. (1960) Zmienność Thamnolia vermicularis (Sw.) Schaer. — De variabilitate Thamnoliae vermicularis (Sw.) Schaer. Fragmenta Floristica et Geobotanica 6: 627635.Google Scholar
Mudd, W. (1865) A Monograph of British Cladoniae, Illustrated with Dried Specimens of Eighty Species and Varieties. Cambridge: privately printed.Google Scholar
Müller Argoviensis, J. (1862) Principes de classification des lichens et énumeration des lichens de Genève. Memoires de la Société de physique et d'histoire naturelle de Genève 16: 343433. Google Scholar
Nelsen, M. P. & Gargas, A. (2009 a) Assessing clonality and chemotype monophyly in Thamnolia (Icmadophilaceae). Bryologist 112: 4253.CrossRefGoogle Scholar
Nelsen, M. P. & Gargas, A. (2009 b) Symbiont flexibility in Thamnolia vermicularis (Pertusariales: Icmadophilaceae). Bryologist 112: 404417.CrossRefGoogle Scholar
Nilsen, T. W. & Graveley, B. R. (2010) Expansion of the eukaryotic proteome by alternative splicing. Nature 463: 457463.CrossRefGoogle ScholarPubMed
Nylander, J. A. A., Wilgenbusch, J. C., Warren, D. L. & Swofford, D. L. (2007) AWTY (are we there yet?): a system for graphical exploration of MCMC convergence in Bayesian phylogenetics. Bioinformatics Applications Note 24: 581583.Google Scholar
Nylander, W. (1856) Animadversiones adhuc quaedam lichenographicae. Flora (Regensburg) 39: 577579. Google Scholar
Nylander, W. (1857) Prodromus lichenographicae Galliae et Algeriae. Actes de la Société linnéenne de Bordeaux 21: 249267. Google Scholar
Nylander, W. (1860) Synopsis Methodica Lichenum Omnium Hucusque Cognitorum Praemissa Introductione Lingua Gallica Tractata. Paris: L. Martinet. Google Scholar
Nylander, W. (1861) Lichenes Scandinaviae (sive Prodromus Lichenographiae Scandinaviae). Notiser ur Sällskapets pro Fauna et Flora Fennica Förhandlingar 5: 1313. Google Scholar
Ozenda, P. & Clauzade, G. (1970) Les Lichens. Étude Biologique et Flore Illustrée. Paris: Masson & Cie.Google Scholar
Pérez, F. L. (1991) Particle sorting due to off-road vehicle traffic in a high Andean Paramo. Catena 18: 239254.CrossRefGoogle Scholar
Pérez, F. L. (1994) Vagrant cryptograms in a Paramo of the high Venezuelan Andes. Flora 189: 263276.CrossRefGoogle Scholar
Platt, J. L. & Spatafora, J. W. (2000) Evolutionary relationships of nonsexual lichenized fungi: molecular phylogenetic hypotheses for the genera Siphula and Thamnolia from SSU and LSUr DNA. Mycologica 92: 475487.CrossRefGoogle Scholar
Poelt, J. (1963) Bestimmungsschlüssel der höheren Flechten von Europa. Mitteilungen der Botanischen Staatssammlung München 4: 301571. Google Scholar
Poelt, J. (1969) Bestimmungsschlüssel Europäischer Flechten. Berlin, Stuttgart: J. Cramer.Google Scholar
Purvis, O. W. (2000) Lichens. London: Natural History Museum / London, Washington D.C.: Smithsonian Books.Google Scholar
Purvis, O. W., Coppins, B. J., Hawksworth, D. L., James, P. W. & Moore, D. M. (eds) (1992) The Lichen Flora of Great Britain and Ireland. London: Natural History Museum Publications.Google Scholar
Rambold, G., Triebel, D. & Hertel, H. (1993) Icmadophilaceae, a new family in the Leotiales . Bibliotheca Lichenologica 53: 217240.Google Scholar
Räsänen, V. (1932) Thamnolia vermicularis (Sw.) Schaer. mit Apothecien und Sporen gefunden. Annales Botanici Societatis Zoologicae-Botanicae Fennicae “Vanamo” 2: 1113. Google Scholar
Räsänen, V. (1937) Das Rätsel der Flechtengattung Siphula Fr. gelöst. Siphula patagonica Vain. mit Apothecien und Sporen gefunden. Annales Botanici Societatis Zoologicae-Botanicae Fennicae “Vanamo” 8: 38.Google Scholar
Rehm, H. (1896) Rabenhorst's Kryptogamen-Flora, Band 1, Abth. 3, 2. Auflage. Leipzig: Kummer.Google Scholar
Rogers, R. W. (1992) Lichen ecology and biogeography. In Flora of Australia. Volume 54. Lichens – Introduction, Lecanorales 1 (George, A. S., ed.): 3042. Canberra: Australian Government Publishing Service.Google Scholar
Ronquist, F., Teslenko, M., van der Mark, P., Ayres, D. L., Darling, A., Höhna, S., Larget, B., Liu, L., Suchard, M. A. & Huelsenbeck, J. A. (2012) MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology 61: 539542.CrossRefGoogle ScholarPubMed
St. Clair, L. L. (1999) A Color Guidebook to Common Rocky Mountain Lichens. Provo, Utah: M. L. Bean Life Science Museum of Brigham Young University.Google Scholar
Sadowska-Des, A., Otte, J. & Schmitt, I. (2012) Analysis of twelve molecular loci suggests high photobiont and low mycobiont diversity in populations of Lasallia pustulata . In Abstracts of the 7th International Association for Lichenology Symposium, 9–13 January 2012, Bangkok, Thailand, p. 131.Google Scholar
Sato, M. (1965) The mixture ratio of the lichen genus Thamnolia in New Zealand. Bryologist 68: 320324.CrossRefGoogle Scholar
Sato, M. (1968) The mixture ratio of the lichen genus Thamnolia in Tasmania and New Guinea. Journal of Japanese Botany 43: 328334.Google Scholar
Serusiaux, E. (1986) The nature and origin of campylidia in lichenized fungi. Lichenologist 18: 135.CrossRefGoogle Scholar
Sheard, J. W. (1977) Paleogeography, chemistry and taxonomy of the lichenized Ascomycetes Dimelaena and Thamnolia . Bryologist 80: 100118.CrossRefGoogle Scholar
Smith, A. L. (1918) A Monograph of the British Lichens. A descriptive Catalogue of the Species in the Department of Botany, British Museum. 1, Ed 2 London: British Museum. Google Scholar
Sommerfelt, S. C. (1826) Supplementum Florae Lapponicae (quam edidit Dr. Georgius Wahlenberg). Christianiae (Oslo): Borgianis et Grondahlianis. CrossRefGoogle Scholar
Stein, B. (1879) Flechten. In Kryptogamen-Flora von Schlesien. Im Namen der Schlesischen (Cohn, F., ed.): 1400. Breslau: Kern.Google Scholar
Stenroos, S., Myllys, L., Thell, A. & Hyvönen, J. (2002) Cladoniaceae, Stereocaulaceae, Icmadophilaceae, and Baeomycetaceae revisited, phylogeny inferred from SSU rDNA sequences. Mycological Progress 1: 267282.CrossRefGoogle Scholar
Stenroos, S., Ahti, T., Lohtander, K. & Myllys, L. (2011) Suomen Jäkäläopas [Lichens of Finland]. Helsinki: Kasvimuseo, Luonnontieteellinen Keskus-Museo.Google Scholar
Stocker-Wörgötter, E. (1998) Culture methods and culture of selected tropical mycobionts and photobionts as exemplified by South American lichens. In Lichenology in Latin America: History, Current Knowledge and Applications (Marcelli, M. P. & Seaward, M. R. D., eds): 143154. Sao Paulo: CETESB – Companhia de Tecnologia de Saneamento Ambiental.Google Scholar
Stocker-Wörgötter, E. (2001) Experimental lichenology and microbiology of lichens: culture experiments, secondary chemistry of cultured mycobionts, resynthesis, and thallus morphogenesis. Bryologist 104: 576581.CrossRefGoogle Scholar
Thomson, J. W. (1984) American Arctic Lichens. 1. The Macrolichens. New York: Columbia University Press.Google Scholar
Tuckerman, E. (1882) A Synopsis of the North American Lichens. Part. I, Comprising the Parmeliacei, Cladoniei and Coenogoniei. Boston: S. F. Cassino. CrossRefGoogle Scholar
Vainio, E. A. (1921) Lichenographia fennica I. Pyrenolichenes iisque proximi Pyrenomycetes et Lichenes imperfecti. Acta Societatis pro Fauna et Flora Fennica 49: 1274. Google Scholar
Vareschi, V. (1970) Flora de los Paramos de Venezuela. Merida: Universidad de los Andes, Ediciones del Rectorado.Google Scholar
Viaud, M., Pasquier, A. & Brygoo, Y. (2000) Diversity of soil fungi studied by PCR-RFLP of ITS. Mycological Research 104: 10271032.CrossRefGoogle Scholar
Vobis, G. (1977) Studies on the germination of lichen conidia. Lichenologist 9: 131136.CrossRefGoogle Scholar
Vobis, G. (1980) Bau und Entwicklung der Flechten-Pycnidien und ihrer Conidien. Bibliotheca Lichenologica 14: 1141.Google Scholar
Vobis, G. & Hawksworth, D. L. (1981) Conidial lichen-forming fungi. In Biology of Conidial Fungi, Volume 1 (Cole, G. T. & Kendrick, B., eds): 245273. London: Academic Press.CrossRefGoogle Scholar
White, T. J., Bruns, T., Lee, S. & Taylor, J. (1990) Amplification and direct sequencing of fungal ribosomal RNA genes for phylogenetics. In PCR Protocols: A Guide to Methods and Applications (Innis, M. A., Gelfand, D. H., Sninsky, J. J. & White, T. J., eds): 315322. San Diego: Academic Press.Google Scholar
Wirth, V. (1980), (1995 a) Flechtenflora. Bestimmung und Ökologische Kennzeichnung der Flechten Südwestdeutschlands und Angrenzender Gebiete. Stuttgart: Verlag Eugen Ulmer. [1995a refers to 2nd edition] Google Scholar
Wirth, V. (1987), (1995 b) Die Flechten Baden-Württembergs, Teil 1 & 2. Stuttgart: Eugen Ulmer GmbH & Co. [1995b refers to 2nd edition] Google Scholar
Zahlbruckner, A. (1907) Ascolichenes. In Die Natürlichen Pflanzenfamilien I. Teil, Abt. 1* (Engler, A. & Prantl, K., eds): 49240. Leipzig: Verlag von Wilhelm Engelmann. Google Scholar
Zhurbenko, M. P. (2012) Lichenicolous fungi growing on Thamnolia, mainly from the Holarctic, with a worldwide key to the known species. Lichenologist 44: 147177.CrossRefGoogle Scholar