Skip to main content

Morphological, chemical and species delimitation analyses provide new taxonomic insights into two groups of Rinodina

  • Philipp RESL (a1), Helmut MAYRHOFER (a1), Stephen R. CLAYDEN (a2), Toby SPRIBILLE (a3), Göran THOR (a4), Tor TØNSBERG (a5) and John W. SHEARD (a6)...
  • Please note a correction has been issued for this article.

The genus Rinodina (Physciaceae), with approximately 300 species, has been subject to few phylogenetic studies. Consequently taxonomic hypotheses in Rinodina are largely reliant on phenotypic data, while hypotheses incorporating DNA dependent methods remain to be tested. Here we investigate Rinodina degeliana/R. subparieta and the Rinodina mniaraea group, which previously have not been subjected to comprehensive molecular and phenotypic studies. We conducted detailed morphological, anatomical, chemical, molecular phylogenetic and species delimitation studies including 24 newly sequenced specimens. We propose that Rinodina degeliana and R. subparieta are conspecific and that chemical morphs within the R. mniaraea group should be recognized as distinct species. We also propose the placement of the recently described genus Oxnerella in Physciaceae.

  • View HTML
    • Send article to Kindle

      To send this article to your Kindle, first ensure is added to your Approved Personal Document E-mail List under your Personal Document Settings on the Manage Your Content and Devices page of your Amazon account. Then enter the ‘name’ part of your Kindle email address below. Find out more about sending to your Kindle. Find out more about sending to your Kindle.

      Note you can select to send to either the or variations. ‘’ emails are free but can only be sent to your device when it is connected to wi-fi. ‘’ emails can be delivered even when you are not connected to wi-fi, but note that service fees apply.

      Find out more about the Kindle Personal Document Service.

      Morphological, chemical and species delimitation analyses provide new taxonomic insights into two groups of Rinodina
      Available formats
      Send article to Dropbox

      To send this article to your Dropbox account, please select one or more formats and confirm that you agree to abide by our usage policies. If this is the first time you use this feature, you will be asked to authorise Cambridge Core to connect with your <service> account. Find out more about sending content to Dropbox.

      Morphological, chemical and species delimitation analyses provide new taxonomic insights into two groups of Rinodina
      Available formats
      Send article to Google Drive

      To send this article to your Google Drive account, please select one or more formats and confirm that you agree to abide by our usage policies. If this is the first time you use this feature, you will be asked to authorise Cambridge Core to connect with your <service> account. Find out more about sending content to Google Drive.

      Morphological, chemical and species delimitation analyses provide new taxonomic insights into two groups of Rinodina
      Available formats
This is an Open Access article, distributed under the terms of the Creative Commons Attribution licence (, which permits unrestricted re-use, distribution, and reproduction in any medium, provided the original work is properly cited.
Corresponding author
Hide All
Altermann, S., Leavitt, S. D., Goward, T., Nelsen, M. P. & Lumbsch, H. T. (2014) How do you solve a problem like Letharia? A new look at cryptic species in lichen-forming fungi using Bayesian clustering and SNPs from multilocus sequence data. PLoS ONE 9: e9755614.
Arnold, F. (1870) Lichenologische Fragmente. X. Flora (Regensburg) 53: 465488.
Baloch, E. & Grube, M. (2009) Pronounced genetic diversity in tropical epiphyllous lichen fungi. Molecular Ecology 18: 21852197.
Blackwell, M. (2011) The fungi: 1, 2, 3 ... 5.1 million species? American Journal of Botany 98: 426438.
Bouckaert, R., Heled, J., Kühnert, D., Vaughan, T., Wu, C.-H., Xie, D., Suchard, M. A., Rambaut, A. & Drummond, A. J. (2014) BEAST 2: a software platform for Bayesian evolutionary analysis. PLoS Computational Biology 10 (4): e10035376.
Brodo, I. M. & Lendemer, J. C. (2012) On the perplexing variability of reproductive modes in the genus Ochrolechia: notes on O. africana and O. arborea in eastern North America. Opuscula Philolichenum 11: 120134.
Coppins, B. J. (1983) A new corticolous sorediate Rinodina from Swedish Lappland. Lichenologist 15: 146150.
Crespo, A. & Lumbsch, H. T. (2010) Cryptic species in lichen-forming fungi. IMA Fungus 1: 167170.
Crespo, A., Blanco, O., Llimona, X. & Ferencová, Z. L. (2004) Coscinocladium, an overlooked endemic and monotypic Mediterranean lichen genus of Physciaceae, reinstated by molecular phylogenetic analysis. Taxon 53: 405414.
Culberson, C. F. (1972) Improved conditions and new data for the identification of lichen products by a standardized thin layer chromatographic method. Journal of Chromatography 72: 113125.
Culberson, C. F. & Kristinsson, H. (1970) A standardized method for the identification of lichen products. Journal of Chromatography 46: 8593.
Ertl, R. (2000) Die Flechtengattung Rinodina in Zentralasien unter besonderer Berücksichtigung der muscicolen Arten. Diploma thesis, University of Graz.
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.
Giavarini, V., James, P. W. & Purvis, O. W. (2009) Rinodina (Ach.) Gray (1821). In The Lichens of Great Britain and Ireland (C. W. Smith, A. Aptroot, B. J. Coppins, A. Fletcher, O. L. Gilbert, P. W. James & P. A. Wolseley, eds): 812825. London: British Lichen Society.
Grube, M. & Arup, U. (2001) Molecular and morphological evolution in the Physciaceae (Lecanorales, lichenized Ascomycotina), with special emphasis on the genus Rinodina . Lichenologist 33: 6372.
Grube, M. & Kroken, S. (2000) Molecular approaches and the concept of species and species complexes in lichenized fungi. Mycological Research 104: 12841294.
Hall, T. A. (1999) BioEdit: a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nucleic Acids Symposium Series 41: 9598.
Helms, G., Friedl, T. & Rambold, G. (2003) Phylogenetic relationships of the Physciaceae inferred from rDNA sequence data and selected phenotypic characters. Mycologia 95: 10781099.
Hennig, C. (2013) fpc: flexible procedures for clustering. R package version 2.1-5. Available at:
Hibbett, D. S. & Taylor, J. W. (2013) Fungal systematics: is a new age of enlightenment at hand? Nature Reviews Microbiology 11: 129133.
James, T. Y., Kauff, F., Schoch, C. L., Matheny, P. B., Hofstetter, V., Cox, C. J., Celio, G., Gueidan, C., Fraker, E., Miadlikowska, J. et al. (2006) Reconstructing the early evolution of fungi using a six-gene phylogeny. Nature 443: 818822.
Kaschik, M. (2006) Taxonomic studies on saxicolous species of the genus Rinodina (lichenized Ascomycetes, Physciaceae) in the Southern Hemisphere with emphasis in Australia and New Zealand. Bibliotheca Lichenologica 93: 1162.
Katoh, K. & Standley, D. M. (2013) MAFFT multiple sequence alignment software version 7: improvements in performance and usability. Molecular Biology and Evolution 30: 772780.
Kauff, F. & Lutzoni, F. (2002) Phylogeny of the Gyalectales and Ostropales (Ascomycota, Fungi): among and within order relationships based on nuclear ribosomal RNA small and large subunits. Molecular Phylogenetics and Evolution 25: 138156.
Kondratyuk, S. Y., Lőkös, L., Kim, J. A., Joeng, M.-H., Kondratiuk, A., Ou, S.-O. & Hur, J.-S. (2014 a) Kashiwadia gen. nov. (Physciaceae, lichen-forming Ascomycota), proved by phylogenetic analysis of eastern Asian Physciaceae . Acta Botanica Hungarica 56 (3–4): 369378.
Kondratyuk, S. Y., Lőkös, L., Kim, J., Kondratiuk, A., Jeong, M. H., Zarei-Darki, B. & Hur, J. S. (2014 b) Oxnerella safavidiorum gen. et spec. nov. (Lecanoromycetidae, Ascomycota) from Iran (Asia) proved by phylogenetic analysis. Acta Botanica Hungarica 56 (3–4): 379398.
Lanfear, R., Calcott, B., Ho, S. Y. W. & Guindon, S. (2012) PartitionFinder: combined selection of partitioning schemes and substitution models for phylogenetic analyses. Molecular Biology and Evolution 29: 16951701.
Leavitt, S. D., Johnson, L. & St. Clair, L. L. (2011 a) Species delimitation and evolution in morphologically and chemically diverse communities of the lichen-forming genus Xanthoparmelia (Parmeliaceae, Ascomycota) in western North America. American Journal of Botany 98: 175188.
Leavitt, S. D., Johnson, L. A., Goward, T. & St. Clair, L. L. (2011 b) Species delimitation in taxonomically difficult lichen-forming fungi: an example from morphologically and chemically diverse Xanthoparmelia (Parmeliaceae) in North America. Molecular Phylogenetics and Evolution 60: 317332.
Leavitt, S. D., Fernández-Mendoza, F., Pérez-Ortega, S., Sohrabi, M., Divakar, P., Lumbsch, H. T. & St. Clair, L. L. (2013 a) DNA barcode identification of lichen-forming fungal species in the Rhizoplaca melanophthalma species-complex (Lecanorales, Lecanoraceae), including five new species. MycoKeys 7: 122.
Leavitt, S. D., Esslinger, T. L., Nelsen, M. P. & Lumbsch, H. T. (2013 b) Further species diversity in Neotropical Oropogon (Lecanoromycetes: Parmeliaceae) in Central America. Lichenologist 45: 553564.
Lendemer, J. C. (2012) A tale of two species: molecular data reveal the chemotypes of Lepraria normandinoides (Stereocaulaceae) to be two sympatric species. Journal of the Torrey Botanical Society 139: 118130.
Lendemer, J. C., Tripp, E. A. & Sheard, J. (2014) A review of Rinodina (Physciaceae) in Great Smoky Mountains National Park highlights the growing significance of this “island of biodiversity” in eastern North America. Bryologist 117: 259281.
Lohtander, K., Källersjö, M., Moberg, R. & Tehler, A. (2000) The family Physciaceae in Fennoscandia: phylogeny inferred from ITS sequences. Mycologia 92: 728735.
Lohtander, K., Myllys, L., Källersjö, M., Moberg, R., Stenroos, S. & Tehler, A. (2009) New entities in Physcia aipoliaP. caesia group (Physciaceae, Ascomycetes): an analysis based on mtSSU, ITS, group I intron and betatubulin sequences. Annales Botanici Fennici 46: 4353.
Lücking, R., Dal-Forno, M., Sikaroodi, M., Gillevet, P. M., Bungartz, F., Moncada, B., Yánez-Ayabaca, A., Chaves, J. L., Coca, L. F. & Lawrey, J. D. (2014) A single macrolichen constitutes hundreds of unrecognized species. Proceedings of the National Academy of Sciences of the United States of America 111: 1109111096.
Lumbsch, H. T. & Leavitt, S. D. (2011) Goodbye morphology? A paradigm shift in the delimitation of species in lichenized fungi. Fungal Diversity 50: 5972.
Mayrhofer, H. & Moberg, R. (2002) Rinodina . In Nordic Lichen Flora Volume 2. Physciaceae (T. Ahti, P. M. Jørgensen, H. Kristinsson, R. Moberg, U. Søchting & G. Thor, eds): 4169. Uddevalla: Nordic Lichen Society.
Mayrhofer, H., Denchew, C. M., Stoykov, D. Y. & Nikolova, S. O. (2005) Catalogue of the lichenized and lichenicolous fungi in Bulgaria. Mycologia Balcanica 2: 361.
Miadlikowska, J., Kauff, F., Högnabba, F., Oliver, J. C., Molnar, K., Fraker, E., Gaya, E., Hafellner, J., Hofstetter, V., Gueidan, C. et al. (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: 132168.
Myllys, L., Velmala, S., Holien, H., Halonen, P., Wang, L.-S. & Goward, T. (2011) Phylogeny of the genus Bryoria . Lichenologist 43: 617638.
Nadyeina, O., Grube, M. & Mayrhofer, H. (2010) A contribution to the taxonomy of the genus Rinodina (Physciaceae, lichenized Ascomycotina) using combined ITS and mtSSU rDNA data. Lichenologist 42: 521531.
Nylander, W. (1870) Addenda nova ad lichenographiam Europaeam. Continuatio tertia decima. Flora (Regensburg) 53: 3338.
Nylander, W. (1900) Lichenes Ceylonensis et additamentum ad lichens Japoniae. Opus posthumum. Acta Societatis Scientiarum Fennicae 26: 133.
Olech, M. (1989) Lichens from the Admiralty Bay region, King George Island (South Shetland Islands, Antarctica). Acta Societatis Botanicorum Poloniae 58: 493512.
Ortiz-Álvarez, R., de los Ríos, A., Fernández-Mendoza, F., Torralba-Burrial, A. & Pérez-Ortega, S. (2015) Ecological specialization of two photobiont-specific maritime cyanolichen species of the genus Lichina . PLoS ONE 10: e0132718.
Poelt, J. & Mayrhofer, H. (1979) Studien über Ascosporen-Typen der Flechtengattung Rinodina . Beiheft zur Sydowia 8: 312331.
Pons, J., Barraclough, T., Gomez-Zurita, J., Cardoso, A., Duran, D., Hazell, S., Kamoun, S., Sumlin, W. D. & Vogler, A. P. (2006) Sequence-based species delimitation for the DNA taxonomy of undescribed insects. Systematic Biology 55: 595609.
R Development Core Team (2013) R: A Language and Environment for Statistical Computing, reference index version 3.0.2. R Foundation for Statistical Computing, Vienna, Austria.
Rambaut, A., Suchard, M. A., Xie, D. & Drummond, A. J. (2014) Tracer v1.6. Available at: Accessed 24 August 2015.
Räsänen, V. (1931) Die Flechten Estlands. Mit einer Bestimmungstabelle der wichtigsten Nord- und Mitteleuropäischen Flechtenarten und Varietäten. I. Annales Academiae Scientiarum Fennicae 34 (4): 1163.
Reid, N. M. & Carstens, B. C. (2012) Phylogenetic estimation error can decrease the accuracy of species delimitation: a Bayesian implementation of the general mixed Yule-coalescent model. BMC Evolutionary Biology 12: 196.
Resl, P. (2015) phylo-scripts: python scripts for phylogenetics, v0.1. Available at: doi:10.5281/zenodo.15983
Resl, P., Schneider, K., Westberg, M., Printzen, C., Palice, Z., Thor, G., Fryday, A., Mayrhofer, H. & Spribille, T. (2015) Diagnostics for a troubled backbone: testing topological hypotheses of trapelioid lichenized fungi in a large-scale phylogeny of Ostropomycetidae (Lecanoromycetes). Fungal Diversity 73: 239258.
Ropin, K. & Mayrhofer, H. (1993) Zur Kenntis corticoler Arten der Gattung Rinodina (lichenisierte Ascomyceten) in den Ostalpen und angrenzenden Gebieten. Herzogia 9: 779835.
Schoch, C. L., Seifert, K. A., Huhndorf, S., Robert, V., Spouge, J. L., Levesque, C. A. & Chen, W., Fungal Barcoding Consortium (2012) Nuclear ribosomal internal transcribed spacer (ITS) region as a universal DNA barcode marker for Fungi. Proceedings of the National Academy of Sciences of the United States of America 109: 62416246.
Sheard, J. W. (2010) The Lichen Genus Rinodina (Ach.) Gray (Lecanoromycetidae, Physciaceae ) in North America, North of Mexico. Ottawa: NRC Research Press.
Sheard, J. W., McCune, B. & Tønsberg, T. (2014) A new corticolous species of Rinodina (Physciaceae) and two interesting range extensions for species collected in Katmai National Park, Alaska. Bryologist 117: 253258.
Singh, G., Dal Grande, F., Divakar, P. K., Otte, J., Leavitt, S. D., Szczepanska, K., Crespo, A., Rico, V. J., Aptroot, A., Cáceres, M. E. S. et al. (2015) Coalescent-based species delimitation approach uncovers high cryptic diversity in the cosmopolitan lichen-forming fungal genus Protoparmelia (Lecanorales, Ascomycota). PLoS ONE 10: e0124625.
Spribille, T., Resl, P., Ahti, T., Pérez-Ortega, S., Tønsberg, T., Mayrhofer, H. & Lumbsch, H. T. (2014) Molecular systematics of the wood-inhabiting, lichen-forming genus Xylographa (Baeomycetales, Ostropomycetidae) with eight new species. Symbolae Botanicae Upsalienses 37 (1): 193.
Stamatakis, A. (2014) RAxML version 8: a tool for phylogenetic analysis and post-analysis of large phylogenies. Bioinformatics 30: 13121313.
Strasser, E. A., Hafellner, J., Stešević, D., Geci, F. & Mayrhofer, H. (2015) Lichenized and lichenicolous fungi from the Albanian Alps (Kosovo, Montenegro). Herzogia 28: 520544.
Tehler, A., Irestedt, M., Bungartz, F. & Wedin, M. (2009) Evolution and reproduction modes in the Roccella galapagoensis aggregate (Roccellaceae, Arthoniales). Taxon 58: 438456.
Thell, A., Crespo, A., Divakar, P. K., Kärnefelt, I., Leavitt, S. D., Lumbsch, H. T. & Seaward, M. R. D. (2012) A review of the lichen family Parmeliaceae – history, phylogeny and current taxonomy. Nordic Journal of Botany 30: 641664.
Timpe, E. B. (1990) Untersuchungen an muscicolen, arktisch-alpin verbreiteten Arten der Gattung Rinodina (Physciaceae, Lichenes). Diploma thesis, University of Graz.
Tønsberg, T. (1992) The sorediate and isidiate, corticolous, crustose lichens in Norway. Sommerfeltia 14: 1331.
Velmala, S., Myllys, L., Goward, T., Holien, H. & Halonen, P. (2014) Taxonomy of Bryoria section Implexae (Parmeliaceae, Lecanoromycetes) in North America and Europe, based on chemical, morphological and molecular data. Annales Botanici Fennici 51 (6): 345371.
Wedin, M., Döring, H., Nordin, A. & Tibell, L. (2000) Small subunit rDNA phylogeny shows the lichen families Caliciaceae and Physciaceae (Lecanorales, Ascomycotina) to form a monophyletic group. Canadian Journal of Botany 78: 246254.
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 (M. A. Innis, D. H. Gelfand, J. J. Sninsky & T. J. White, eds): 315322. New York: Academic Press.
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.
Recommend this journal

Email your librarian or administrator to recommend adding this journal to your organisation's collection.

The Lichenologist
  • ISSN: 0024-2829
  • EISSN: 1096-1135
  • URL: /core/journals/lichenologist
Please enter your name
Please enter a valid email address
Who would you like to send this to? *



Full text views

Total number of HTML views: 74
Total number of PDF views: 396 *
Loading metrics...

Abstract views

Total abstract views: 307 *
Loading metrics...

* Views captured on Cambridge Core between 27th September 2016 - 21st May 2018. This data will be updated every 24 hours.