Skip to main content Accessibility help

Phylogeny of the cetrarioid core (Parmeliaceae) based on five genetic markers

  • Arne THELL (a1), Filip HÖGNABBA (a2), John A. ELIX (a3), Tassilo FEUERER (a4), Ingvar KÄRNEFELT (a1), Leena MYLLYS (a2), Tiina RANDLANE (a5), Andres SAAG (a5), Soili STENROOS (a2), Teuvo AHTI (a2) and Mark R. D. SEAWARD (a6)...


Fourteen genera belong to a monophyletic core of cetrarioid lichens, Ahtiana, Allocetraria, Arctocetraria, Cetraria, Cetrariella, Cetreliopsis, Flavocetraria, Kaernefeltia, Masonhalea, Nephromopsis, Tuckermanella, Tuckermannopsis, Usnocetraria and Vulpicida. A total of 71 samples representing 65 species (of 90 worldwide) and all type species of the genera are included in phylogentic analyses based on a complete ITS matrix and incomplete sets of group I intron, β-tubulin, GAPDH and mtSSU sequences. Eleven of the species included in the study are analysed phylogenetically for the first time, and of the 178 sequences, 67 are newly constructed. Two phylogenetic trees, one based solely on the complete ITS-matrix and a second based on total information, are similar, but not entirely identical. About half of the species are gathered in a strongly supported clade composed of the genera Allocetraria, Cetraria s. str., Cetrariella and Vulpicida. Arctocetraria, Cetreliopsis, Kaernefeltia and Tuckermanella are monophyletic genera, whereas Cetraria, Flavocetraria and Tuckermannopsis are polyphyletic. The taxonomy in current use is compared with the phylogenetic results, and future, probable or potential adjustments to the phylogeny are discussed. The single non-DNA character with a strong correlation to phylogeny based on DNA-sequences is conidial shape. The secondary chemistry of the poorly known species Cetraria annae is analyzed for the first time; the cortex contains usnic acid and atranorin, whereas isonephrosterinic, nephrosterinic, lichesterinic, protolichesterinic and squamatic acids occur in the medulla. Notes on the anatomy of Cetraria annae and Flavocetraria minuscula are also provided.



Hide All
Arup, U., Ekman, S., Grube, M., Mattsson, J.-E. & Wedin, M. (2007) The sister group relation of Parmeliaceae (Lecanorales, Ascomycota). Mycologia 99: 4249.
Blanco, O., Crespo, A., Divakar, P. K., Elix, J. A., Hawksworth, D. L. & Lumbsch, H. T. (2004 a) A molecular phylogeny and a new classification of parmelioid lichens containing Xanthoparmelia-type lichenan (Ascomycota: Lecanorales). Taxon 53: 959975.
Blanco, O., Crespo, A., Divakar, P. K., Esslinger, T. L., Hawksworth, D. L. & Lumbsch, H. T. (2004 b) Melanelixia and Melanohalea, two new genera segregated from Melanelia (Parmeliaceae) based on molecular data. Mycological Research 108: 873884.
Blanco, O., Crespo, A., Divakar, P. K., Elix, J. A. & Lumbsch, H. T. (2005) Molecular phylogeny of parmotremoid lichens (Ascomycota, Parmeliaceae). Mycologia 97: 150159.
Blanco, O., Crespo, A., Ree, R. H. & Lumbsch, H. T. (2006) Major clades of parmelioid lichens (Parmeliaceae, Ascomycota) and the evolution of their morphological and chemical diversity. Molecular Phylogenetics and Evolution 39: 5269.
Crespo, A., Lumbsch, H. T., Mattsson, J.-E., Blanco, O., Divakar, P. K., Articus, K., Wiklund, E., Bawingan, P. A. & Wedin, M. (2007) Testing morphology-based hypotheses of phylogenetic relationships in Parmeliaceae (Ascomycota) using three ribosomal markers and the nuclear RPB1 gene. Molecular Phylogenetics and Evolution 44: 812824.
Divakar, P. K., Crespo, A., Blanco, O. & Lumbsch, H. T. (2006) Phylogenetic significance of morphological characters in the tropical Hypotrachyna clade of parmelioid lichens (Parmeliaceae, Ascomycota). Molecular Phylogenetics and Evolution 40: 448458.
Divakar, P. K., Molina, M. C., Lumbsch, H. T. & Crespo, A. (2005) Parmelia barrenoae, a new lichen species related to Parmelia sulcata (Parmeliaceae) based on molecular and morphological data. Lichenologist 37: 3746.
Egan, R. S. (1987) A fifth checklist of the lichen-forming, lichenicolous and allied fungi of the continental United States and Canada. Bryologist 90: 77173.
Ekman, S. (2001) Molecular phylogeny of the Bacidiaceae (Lecanorales, lichenized Ascomycota). Mycological Research 105: 783797.
Elix, J. A. (1993) Progress in the generic delimitation of Parmelia sensu lato lichens (Ascomycotina: Parmeliaceae). Bryologist 96: 359383.
Elix, J. A., Tønsberg, T. & Wardlaw, J. H. (2004) The structure of friesiic acid, a novel lichen substance from Hypocenomyce friesii. Bibliotheca Lichenologica 88: 103109.
Esslinger, T. L. (2003) Tuckermanella, a new cetrarioid genus in western North America. Mycotaxon 85: 135141.
Gardes, M. & Bruns, T. D. (1993) ITS primers with enhanced specificity for basidiomycetes. Application to the identification mycorrhizae and rusts. Molecular Ecology 2: 113118.
Goward, T. (1985) Ahtiana, a new lichen genus in the Parmeliaceae. Bryologist 88: 367371.
Hafellner, J. (1984) Studien in Richtung einer naturlicheren Gliederung der Sammelfamilien Lecanoraceae und Lecideaceae. Nova Hedwigia 79: 241371. Vaduz: J. Cramer.
Henssen, A. & Jahns, H. M. (1974) Lichenes. Stuttgart: G. Thieme.
Kärnefelt, I. (1977 a) Masonhalea, a new lichen genus in the Parmeliaceae. Botaniska Notiser 130: 101107.
Kärnefelt, I. (1977 b) Three new species of brown fruticose Cetraria. Botaniska Notiser 130: 125129.
Kärnefelt, I. (1979) The brown fruticose species of Cetraria. Opera Botanica 46: 1150.
Kärnefelt, I. (1986) The genera Bryocaulon, Coelocaulon and Cornicularia and formerly associated taxa. Opera Botanica 86: 190.
Kärnefelt, I., Mattsson, J.-E. & Thell, A. (1992) Evolution and phylogeny of cetrarioid lichens. Plant Systematics and Evolution 183: 113160.
Kärnefelt, I., Mattsson, J.-E. & Thell, A. (1993) The lichen genera Arctocetraria, Cetraria and Cetrariella (Parmeliaceae) and their presumed evolutionary affinities. Bryologist 96: 394404.
Kärnefelt, I. & Thell, A. (1992) The evaluation of characters in lichenized families, exemplified with the alectorioid and some parmelioid genera. Plant Systematics and Evolution 180: 181204.
Kärnefelt, I. & Thell, A. (1993) Chemical evolution in cetrarioid lichens. Bibliotheca Lichenologica 53: 115127.
Kärnefelt, I. & Thell, A. (1994) Sexual reproductive characters vs. morphological characters in lichen genera. Journal of the Hattori Botanical Laboratory 76: 207219.
Kärnefelt, I. & Thell, A. (1996) A new classification for the Dactylina/Dufourea complex. Nova Hedwigia 62: 487511.
Kärnefelt, I. & Thell, A. (2001) Delimitation of the genus Tuckermannopsis Gyeln. (Ascomycotina, Parmeliaceae) based on morphology and DNA sequences. Bibliotheca Lichenologica 78: 193209.
Kärnefelt, I., Thell, A., Randlane, T. & Saag, A. (1994) The genus Flavocetraria Kärnefelt & Thell (Parmeliaceae, Ascomycotina) and its affinities. Acta Botanica Fennica 150: 7986.
Kurokawa, S. & Lai, M. J. (1991) Allocetraria, a new lichen genus in the Parmeliaceae. Bulletin of the National Science Museum (Tokyo), Series B (Botany) 17: 5965.
Lai, M. J. 1980. Studies on the cetrarioid lichens in Parmeliaceae of East Asia (I). Quarterly Journal of the Taiwan Museum 33: 215229.
Lai, M. J. & Elix, J. A. (2002) A new species of Cetreliopsis (Ascomycotina, Parmeliaceae) from Thailand. Mycotaxon 84: 355360.
Lai, M. J., Qian, Z. G. & Xu, L. (2007) Synopsis of the cetrarioid lichen genera and species (Parmeliaceae, lichenized Ascomycotina) in China. Journal of the National Taiwan Museum 60: 4561.
Larget, B. & Simon, D. L. (1999) Markov chain Monte Carlo algorithms for the Bayesian analysis of Phylogenetic trees. Molecular Biology and Evolution 16: 750759.
Lumbsch, H. T., Schmitt, I., Mangold, A. & Wedin, M. (2007) Ascus types are phylogenetically misleading in Trapeliaceae and Agyriaceae (Ostropomycetidae, Ascomycota). Mycological Research 111: 11331141.
Mattsson, J.-E. (1993) A monograph of the genus Vulpicida (Parmeliaceae, Ascomycetes). Opera Botanica 119: 161.
Mattsson, J.-E. & Lai, M. J. (1993) Vulpicida, a new genus in Parmeliaceae (lichenized ascomycetes). Mycotaxon 46: 425428.
Müller [Argoviensis], J. (1891) Lichenologische Beiträge 35. Flora (Regensburg) 74: 371382.
Myllys, L., Lohtander, K. & Tehler, A. (2001) β-tubulin, ITS and group I intron sequences challenge the species pair Physcia aipolia and P. caesia. Mycologia 93: 335343.
Myllys, L., Stenroos, S. & Thell, A. (2002) New genes for phylogenetic studies of lichenized fungi: glyceraldehyde-3-phosphate dehydrogenase and β-tubulin genes. Lichenologist 34: 237246.
Obermayer, W. (2008) Fotografische Dokumentation einer ungewöhnlich reich fruchtenden Aufsammlung von Cetraria islandica (L.) Ach. [mit einem historischen Abriss zur Darstellung fertiler Thalli, Anmerkungen zur Gestalt der Pycnosporen und einigen Notizen zum Gebrauch des ‘Kramperltees’]. Mitteilungen des naturwissenschaftlichen Vereins für Steiermark 138: 113158.
Ohmura, Y. (2002) Phylogenetic evaluation of infrageneric groups of the genus Usnea based on ITS regions in rDNA. Journal of the Hattori Botanical Laboratory 92: 231243.
Ohmura, Y. & Kanda, H. (2004) Taxonomic status of section Neuropogon in the genus Usnea elucidated by morphological comparison and ITS rDNA sequences. Lichenologist 36: 217225.
Oxner, A. N. (1933) Species Lichenum novae ex Asia. Journal du Cycle Botanique de l'Academie des Sciences d'Ukraine 2: 167172.
Prado, R. del, Ferencová, Z., Armas-Crespo, V., Amo de Paz, G., Cubas, P. & Crespo, A. (2007) The arachiform vacuolar body, an overlooked shared character in the ascospores of a large monophyletic group within Parmeliaceae (Xanthoparmelia clade, Lecanorales). Mycological Research 91: 455467.
Randlane, T. & Saag, A. (1998) Synopsis of the genus Nephromopsis (fam. Parmeliaceae, lichenized Ascomycota). Cryptogamie, Bryologie-Lichénologie 19: 175191.
Randlane, T. & Saag, A. (2000) Revision of the second updated world list of cetrarioid lichens. January 17, 2002.
Randlane, T. & Saag, A. (2003) Taxonomic notes on some cetrarioid lichens. Mycotaxon 87: 479487.
Randlane, T. & Saag, A. (2004) Distribution patterns of some primary and secondary cetrarioid species. Acta Universitatis Upsaliensis Symbolae Botanicae Upsalienses 34: 359376.
Randlane, T. & Saag, A. (2005) Distribution patterns of primary and secondary species in the genus Vulpicida. Folia Cryptogamica Estonica 41: 8996.
Randlane, T., Saag, A. & Obermayer, W. (2001) Cetrarioid lichens containing usnic acid from the Tibetan area. Mycotaxon 80: 389425.
Randlane, T., Saag, A. & Thell, A. (1997) A second updated world list of cetrarioid lichens. Bryologist 100: 109122.
Randlane, T., Thell, A. & Saag, A. (1995) New data about the genera Cetrariopsis, Cetreliopsis and Nephromopsis (fam. Parmeliaceae, lichenized Ascomycotina). Cryptogamie, Bryologie-Lichénologie 16: 3560.
Rico, V. J., van den Boom, P. P. G. & Barrasa, J. M. (2005) Morphology, chemistry and distribution of Melanelia sorediella (Parmeliaceae) and similar species in the Iberian Peninsula. Lichenologist 37: 199215.
Saag, A., Randlane, T., Thell, A. & Obermayer, W. (2002) Phylogenetic analysis of cetrarioid lichens with globose ascospores. Proceedings of the Estonian Academy of Sciences, Biology, Ecology 51: 103123.
Swofford, D. L. (1998) PAUP: Phylogenetic Analysis Using Parsimony, Version 4.0b. Sunderland, Mass.: Sinauer Associates.
Thell, A. (1995 a) A new position of the Cetraria commixta group in Melanelia (Ascomycotina, Parmeliaceae). Nova Hedwigia 60: 407422.
Thell, A. (1995 b) Pycnoconidial types and their presence in cetrarioid lichens (Ascomycotina, Parmeliaceae). Cryptogamie, Bryologie-Lichénologie 16: 247256.
Thell, A. (1998) Phylogenetic relationships of some cetrarioid species in British Columbia with notes on Tuckermannopsis. Folia Cryptogamica Estonica 32: 113122.
Thell, A. (1999) Group I intron versus ITS sequences in phylogeny of cetrarioid lichens. Lichenologist 31: 441449.
Thell, A., Feuerer, T., Elix, J. A. & Kärnefelt, I. (2006) A contribution to the phylogeny and taxonomy of Xanthoparmelia (Ascomycota, Parmeliaceae). Journal of the Hattori Botanical Laboratory 100: 797807.
Thell, A., Feuerer, T., Kärnefelt, I., Myllys, L. & Stenroos, S. (2004) Monophyletic groups within the Parmeliaceae identified by ITS rDNA, betatubulin and GAPDH sequences. Mycological Progress 3: 297314.
Thell, A. & Goward, T. (1996) The new cetrarioid genus Kaernefeltia and related groups in the Parmeliaceae (lichenized Ascomycotina). Bryologist 99: 125136.
Thell, A., Goward, T., Randlane, T., Kärnefelt, I. & Saag, A. (1995 a) A revision of the North American lichen genus Ahtiana (Parmeliaceae). Bryologist 98: 596605.
Thell, A., Mattsson, J.-E. & Kärnefelt, I. (1995 c) Lecanoralean ascus types in the lichenized families Alectoriaceae and Parmeliaceae. Cryptogamic Botany 5: 120127.
Thell, A. & Miao, V. (1998) Phylogenetic analysis of ITS and group I intron sequences from European and North American samples of cetrarioid lichens. Annales Botanici Fennici 35: 275286.
Thell, A., Randlane, T., Kärnefelt, I., Gao, X.-Q. & Saag, A. (1995 b) The lichen genus Allocetraria (Ascomycotina, Parmeliaceae). In Flechten Follmann. Contributions to Lichenology in Honour of Gerhard Follmann (Daniels, J. A., Schulz, M. & Peine, J., eds): 353370. Cologne: University of Cologne.
Thell, A., Randlane, T., Saag, A. & Kärnefelt, I. (2005) A new circumscription of the lichen genus Nephromopsis (Parmeliaceae, lichenized Ascomycetes). Mycological Progress 4: 303316.
Thell, A., Søchting, U., Kärnefelt, I., Elix, J. A. & Sancho, L. G. (2007) Phylogeny of Himantormia – an Antarctic genus in the Parmeliaceae (lichenized ascomycetes). Bibliotheca Lichenologica 95: 95541.
Thell, A., Stenroos, S., Feuerer, T., Kärnefelt, I., Myllys, L. & Hyvönen, J. (2002) Phylogeny of cetrarioid lichens (Parmeliaceae) inferred from ITS and β-tubulin sequences, morphology, anatomy and secondary chemistry. Mycological Progress 1: 335354.
Thell, A., Stenroos, S. & Myllys, L. (2000) A DNA-study of the Cetraria aculeata and C. islandica groups. Folia Cryptogamica Estonica 36: 95106.
van den Boom, P. P. G. & Sipman, H. J. M. (1994) Cetraria obtusata comb. et stat nov., an overlooked lichen species from the central Alps. Lichenologist 26: 105112.
White, T. J., Burns, T., Lee, S. & Taylor, J. W. (1991) 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.
Wirtz, N., Printzen, C., Sancho, L. G. & Lumbsch, H. T. (2006) The phylogeny and classification of Neuropogon and Usnea (Parmeliaceae, Ascomycota) revisited. Taxon 55: 367376.
Zhurbenko, M. P., Laursen, G. A. & Walker, D. A. (2005) New and rare lichenicolous fungi and lichens from the North American Arctic. Mycotaxon 92: 201212.
Zoller, S., Scheidegger, C. & Sperisen, C. (1999) PCR-primers for the amplification of mitochondrial small subunit ribosomal DNA of lichen-forming ascomycetes. Lichenologist 31: 511516.


Phylogeny of the cetrarioid core (Parmeliaceae) based on five genetic markers

  • Arne THELL (a1), Filip HÖGNABBA (a2), John A. ELIX (a3), Tassilo FEUERER (a4), Ingvar KÄRNEFELT (a1), Leena MYLLYS (a2), Tiina RANDLANE (a5), Andres SAAG (a5), Soili STENROOS (a2), Teuvo AHTI (a2) and Mark R. D. SEAWARD (a6)...


Full text views

Total number of HTML views: 0
Total number of PDF views: 0 *
Loading metrics...

Abstract views

Total abstract views: 0 *
Loading metrics...

* Views captured on Cambridge Core between <date>. This data will be updated every 24 hours.

Usage data cannot currently be displayed