Hostname: page-component-586b7cd67f-tf8b9 Total loading time: 0 Render date: 2024-11-24T12:52:14.390Z Has data issue: false hasContentIssue false

The taxonomy of the genus Graphis sensu Staiger (Ascomycota: Ostropales: Graphidaceae)

Published online by Cambridge University Press:  02 July 2009

Robert LÜCKING
Affiliation:
Botany Department, The Field Museum, 1400 South Lake Shore Drive, Chicago, Illinois 60605-2496, USA. Email: [email protected]

Abstract

The species-level taxonomy of the genus Graphis sensu Staiger was revised using numerical taxonomy and cladistic techniques. In its current circumscription, Graphis includes more than 300 species and is characterized by lirellae with well-developed, carbonized labia usually concealing the disc and usually hyaline, I+ blue-violet, transversely septate to muriform ascospores. The study analyzed the importance of morphological, anatomical, and chemical characters for taxonomic purposes in the genus, both at species level and at the level of species groups and lineages. A phenotype-based cladistic analysis, combined with a novel method to estimate the degree of character state homoplasies prior to a cladogram, as well as a species ordination using non-metric multidimensional scaling (NMS), was performed to assess the delimitation of potentially natural groups within the genus. Contrary to previous views, the most important character complex characterizing potentially natural groups in the genus Graphis is lirellae morphology, whereas labia striation, excipulum carbonization, ascospore features, and secondary chemistry, vary greatly even in closely related species. Based on these findings, Wirth and Hale's concept of ‘sporomorphs’ is emended to generally characterize morphologically similar species that differ in a single character, such as labia striation (‘labiomorph’), excipulum carbonization (‘excipulomorph’), hymenium inspersion (‘inspersomorph’), ascospore size and septation (‘sporomorph’) and secondary chemistry (‘chemomorph’).

Type
Research Article
Copyright
Copyright © British Lichen Society 2009

Access options

Get access to the full version of this content by using one of the access options below. (Log in options will check for institutional or personal access. Content may require purchase if you do not have access.)

References

Adawadkar, B. & Makhija, U. (2006) New species and new records of Graphis from India: transseptate species with completely carbonized exciples and norstictic acid. Mycotaxon 96: 5160.Google Scholar
Adawadkar, B. & Makhija, U. (2007) New species and new records of Graphis from India with partially carbonized exciples and transseptate ascospores. Mycotaxon 99: 303326.Google Scholar
Aptroot, A., Lücking, R., Sipman, H. J. M., Umaña, L. & Chaves, J. L. (2008) Pyrenocarpous lichens with bitunicate asci. A first assessment of the lichen biodiversity inventory in Costa Rica. Bibliotheca Lichenologica 97: 1162.Google Scholar
Archer, A. W. (1999) The lichen genera Graphis and Graphina (Graphidaceae) in Australia 1. Species based on Australian type specimens. Telopea 8: 273295.Google Scholar
Archer, A. W. (2000) The lichen genera Phaeographis and Phaeographina (Graphidaceae) in Australia. 1. Species based on Australian type specimens. Telopea 8: 461475.CrossRefGoogle Scholar
Archer, A. W. (2001 a) New taxa and new reports in the lichen family Graphidaceae (Ascomycotina) from Australia. Mycotaxon 80: 367374.Google Scholar
Archer, A. W. (2001 b) The lichen genera Phaeographis and Phaeographina (Graphidaceae) in Australia 2. Phaeographina – new reports and new species. Telopea 9: 329344.CrossRefGoogle Scholar
Archer, A. W. (2001 c) The lichen genera Phaeographis and Phaeographina (Graphidaceae) in Australia 3. Phaeographis – new reports and new species. Telopea 9: 663677.CrossRefGoogle Scholar
Archer, A. W. (2001 d) The lichen genus Graphis (Graphidaceae) in Australia. Australian Systematic Botany 14: 245271.CrossRefGoogle Scholar
Archer, A. W. (2001 e) The lichen genus Graphina (Graphidaceae) in Australia: new reports and new species. Mycotaxon 77: 153180.Google Scholar
Archer, A. W. (2003 a) Graphina streblocarpa, Graphina subserpentina (Graphidaceae, lichenised Ascomycota) and their synonyms. Mycotaxon 86: 3136.Google Scholar
Archer, A. W. (2003 b) New species in the lichen family Graphidaceae (Ascomycota) from Australia and the Solomon Islands. Mycotaxon 88: 143148.Google Scholar
Archer, A. W. (2005) Australian species in the genus Diorygma (Graphidaceae). Australasian Lichenology 56: 1112.Google Scholar
Archer, A. W. (2006) The lichen family Graphidaceae in Australia. Bibliotheca Lichenologica 94: 1191.Google Scholar
Archer, A. W. (2007) Additional lichen records from Australia 63. Australasian Lichenology 61: 67.Google Scholar
Awasthi, D. D. (1991) A key to the microlichens of India, Nepal and Sri Lanka. Bibliotheca Lichenologica 40: 1340.Google Scholar
Awasthi, D. D. & Joshi, M. (1979) The lichen genera Helminthocarpon, Cyclographa, and Cyclographina (gen. nov.). Norwegian Journal of Botany 26: 165177.Google Scholar
Awasthi, D. D. & Singh, K. P. (1975) Observations on some graphidaceous lichen taxa. Phyta 1: 3440.Google Scholar
Clements, F. E. (1909) The Genera of Fungi. Minneapolis.CrossRefGoogle Scholar
Crisp, M. D. & Chandler, G. T. (1996) Paraphyletic species. Telopea 6: 813844.CrossRefGoogle Scholar
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.CrossRefGoogle ScholarPubMed
Culberson, C. F. & Kristinsson, H. (1970) A standardized method for the identification of lichen products. Journal of Chromatography 46: 8593.CrossRefGoogle Scholar
Dodge, C. W. (1966) New lichens from Chile. Nova Hedwigia 12: 307352.Google Scholar
Dress, A. W., Flamm, C., Fritzsch, G., Grünewald, S., Kruspe, M., Prohaska, S. J. & Stadler, P. F. (2008) Noisy: identification of problematic columns in multiple sequence alignments. Algorithms for Molecular Biology 3: 7 (online: doi:10.1186/1748-7188-3-7).CrossRefGoogle ScholarPubMed
Frisch, A., Kalb, K. & Grube, M. (2006) Contributions towards a new systematics of the lichen family Thelotremataceae. Bibliotheca Lichenologica 92: 1556.Google Scholar
Funk, D. A. & Omland, K. E. (2003) Species-level paraphyly and polyphyly: frequency, causes, and consequences, with insights from animal mitochondrial DNA. Annual Review of Ecology, Evolution, and Systematics 34: 397423.CrossRefGoogle Scholar
Hafellner, J. (1984) Studien in Richtung einer naturlicheren Gliederung der Sammelfamilien Lecanoraceae und Lecideaceae. Beiheft zur Nova Hedwigia 79: 241371.Google Scholar
Hale, M. E. Jr. (1974) Morden-Smithsonian Expedition to Dominica: the lichens (Thelotremataceae). Smithsonian Contributions to Botany 16: 146.CrossRefGoogle Scholar
Hale, M. E. Jr. (1978) A revision of the lichen family Thelotremataceae in Panama. Smithsonian Contributions to Botany 38: 160.CrossRefGoogle Scholar
Hale, M. E. Jr. (1980) Generic delimitation in the lichen family Thelotremataceae. Mycotaxon 11: 130138.Google Scholar
Hale, M. E. Jr. (1981) A revision of the lichen family Thelotremataceae in Sri Lanka. Bulletin of the British Museum (Natural History), Botany Series 8: 227332.Google Scholar
Harris, R. C. (1989) A sketch of the family Pyrenulaceae (Melanommatales) in eastern North America. Memoirs of the New York Botanical Garden 49: 74107.Google Scholar
Harris, R. C. (1990) Some Florida Lichens. Bronx, N.Y.: Published by the Author.Google Scholar
Harris, R. C. (1995) More Florida Lichens. Including the 10ø Tour of the Pyrenolichens. Bronx, N.Y.: Published by the Author.Google Scholar
Hawksworth, D. L. (1976) Lichen chemotaxonomy. In Lichenology: Progress and Problems (Brown, D. H., Hawksworth, D. L. & Bailey, R. H., eds): 139184. London: Academic Press.Google Scholar
Henssen, A. & Jahns, H. M. (1974) Lichenes. Stuttgart: Georg Thieme.Google Scholar
Herrera-Campos, M. A., Huhndorf, S. & Lücking, R. (2005) The foliicolous lichen flora of Mexico IV: a new foliicolous species of Pyrenothrix (Chaetothyriales: Pyrenothrichaceae). Mycologia 97: 356361.CrossRefGoogle Scholar
Kalb, K. & Staiger, B. (2000) Dyplolabia Massalongo. Monographie einer vergessenen Flechtengattung. Hoppea 61: 409422.Google Scholar
Kalb, K., Staiger, B. & Elix, J. A. (2004) A monograph of the lichen genus Diorygma – a first attempt. Symbolae Botanicae Upsalienses 34(1): 133181.Google Scholar
Kirk, P. M., Cannon, P. F., Minter, D. W. & Stalpers, J. A. (2008) Ainsworth & Bisby's Dictionary of the Fungi. 10th edition. Wallingford, Oxon: CAB International.CrossRefGoogle Scholar
Lücking, R. (2007) Kalbographa: Monografie einer unerkannten Flechtengattung. Bibliotheca Lichenologica 96: 185192.Google Scholar
Lücking, R. (2008) Foliicolous lichenized fungi. Flora Neotropopica Monograph 103: 1867.Google Scholar
Lücking, R. & Rivas Plata, E. (2008) Clave y guía ilustrada para géneros de Graphidaceae. Glalia 1: 141.Google Scholar
Lücking, R., Sérusiaux, E. & Vězda, A. (2005) Phylogeny and systematics of the lichen family Gomphillaceae (Ostropales) inferred from cladistic analysis of phenotype data. Lichenologist 37: 123170.CrossRefGoogle Scholar
Lücking, R., Kalb, K., Staiger, B. & McNeill, J. (2007) Proposal to conserve the name Phaeographis, with a conserved type, against Creographa, Ectographis, Flegographa, Hymenodecton, Platygramma, and Pyrographa (Ascomycota: Ostropales: Graphidaceae), along with notes on the names Graphina and Phaeographina. Taxon 56: 12961299.CrossRefGoogle Scholar
Lücking, R., Chaves, J. L., Sipman, H. J. M., Umaña, L. & Aptroot, A. (2008 a) A first assessment of the Ticolichen biodiversity inventory in Costa Rica: the genus Graphis, with notes on the genus Hemithecium (Ascomycota: Ostropales: Graphidaceae). Fieldiana (Botany), New Series 46: 1131.CrossRefGoogle Scholar
Lücking, R., Del Prado, R., Lumbsch, H. T., Will-Wolf, S., Aptroot, A., Sipman, H. J. M., Umaña, L., and Chaves, J. L. (2008 b) Phylogenetic patterns of morphological and chemical characters and reproductive mode in the Heterodermia obscurata group in Costa Rica (Ascomycota, Physciaceae). Systematics and Biodiversity 6: 3141.CrossRefGoogle Scholar
Lücking, R., Archer, A. W. & Aptroot, A. (2009) A world-wide key to the genus Graphis (Ostropales: Graphidaceae). Lichenologist 41: 363452.CrossRefGoogle Scholar
Lumbsch, H. T. (1998) The use of metabolic data in lichenology at the species and subspecific levels. Lichenologist 30: 357367.CrossRefGoogle Scholar
Lumbsch, H. T. (2002) Analysis of phenolic products in lichens for identification and taxonomy. In Protocols in Lichenology. Culturing, Biochemistry, Ecophysiology and Use in Biomonitoring (Kranner, I., Beckett, R. P. & Varma, A. K., eds): 281295. Berlin, Heidelberg: Springer.Google Scholar
Lumbsch, H. T., Mangold, A., Martín, M. P. & Elix, J. A. (2008 a) Species recognition and phylogeny of Thelotrema species in Australia (Ostropales, Ascomycota). Australian Systematic Botany 21:217227.CrossRefGoogle Scholar
Lumbsch, H. T., Nelsen, M. P. & Lücking, R. (2008 b) The phylogenetic position of Haematommataceae (Lecanorales, Ascomycota), with notes on secondary chemistry and species delimitation. Nova Hedwigia 86: 105114.CrossRefGoogle Scholar
Maddison, W. P. & Maddison, D. R. (2008) Mesquite: a Modular System for Evolutionary Analysis. Version 2.5. http://mesquiteproject.org.Google Scholar
Makhija, U. & Adawadkar, B. (2005 a) Some additions to the Graphidaceae in the Andaman Islands, India. Mycotaxon 91: 347352.Google Scholar
Makhija, U. & Adawadkar, B. (2005 b) Some new species of Graphis (lichenised Ascomycota) from the Andaman and Nicobar Islands of India. Mycotaxon 91: 369379.Google Scholar
Makhija, U., Dube, A., Adawadkar, B. & Chitale, G. (2005) Five trans-septate species of Hemithecium from India. Mycotaxon 93: 365372.Google Scholar
Mangold, A. (2008) Taxonomic studies on members of thelotrematoid Ostropales (lichenized Ascomycota) in Australia. Ph.D. Thesis, Universität Duisburg-Essen.Google Scholar
Mangold, A., Martín, M. P., Lücking, R. & Lumbsch, H. T. (2008) Molecular phylogeny places Thelotremataceae within Graphidaceae (Ascomycota: Ostropales). Taxon 57: 476486.Google Scholar
Mangold, A., Elix, J. A. & Lumbsch, H. T. (2009) Thelotremataceae. Flora of Australia (in press).Google Scholar
McCune, B. & Grace, J. B. (2002). Analysis of Ecological Communities. Gleneden Beach, Oregon: MjM Software Design.Google Scholar
McCune, B. & Mefford, M. J. (1999) PC-ORD. Multivariate Analysis of Ecological Data. Version 4. Gleneden Beach, Oregon: MjM Software Design.Google Scholar
McDonald, T., Miądlikowska, J. & Lutzoni, F. (2003) The lichen genus Sticta in the Great Smoky Mountains: a phylogenetic study of morphologial, chemical, and molecular data. Bryologist 106: 6179.CrossRefGoogle Scholar
Miądlikowska, J. & Lutzoni, F. (2000) Phylogenetic revision of the genus Peltigera (lichen-forming Ascomycota) based on morphological, chemical, and large subunit nuclear ribosomal DNA data. International Journal of Plant Science 161: 925958.CrossRefGoogle Scholar
Müller Argoviensis, J. (1880) Lichenologische Beiträge X. Flora 63: 1724, 4045.Google Scholar
Müller Argoviensis, J. (1882) Lichenologische Beiträge XV. Flora 65: 291306, 313322, 326337, 381386, 397402.Google Scholar
Müller Argoviensis, J. (1887) Lichenologische Beiträge XXV. Flora 70: 5664, 7480.Google Scholar
Müller Argoviensis, J. (1891) Lichenes Brisbanenses a cl. F. M. Bailey, Gouvernment Botanist, prope Brisbane (Queensland) in Australia orientali lecti. Nuovo Giornale Botanico Italiano 23: 385404.Google Scholar
Myllys, L., Stenroos, S., Thell, A. & Ahti, T. (2003) Phylogeny of bipolar Cladonia arbuscula and Cladonia mitis (Lecanorales, Euascomycetes). Molecular Phylogenetics and Evolution 27: 5869.CrossRefGoogle ScholarPubMed
Nagarkar, M. B. & Patwardhan, P. G. (1982) Notes on some lichens from North East India IV: Genus Graphis. Biovigyanam 8: 125131.Google Scholar
Nakanishi, M. (1966) Taxonomical studies on the family Graphidaceae of Japan. Journal of Science of the Hiroshima University, Series B, Division 2 (Botany) 11: 51126.Google Scholar
Nakanishi, M., Kashiwadani, H. & Moon, K. H. (2003 a) New species of Graphis, Phaeographina and Sarcographa (Ascomycotina, Graphidaceae) from Vanuatu. Bulletin of the National Science Museum (Tokyo), Series B (Botany) 28: 107111.Google Scholar
Nakanishi, M., Kashiwadani, H. & Moon, K. H. (2003 b) Taxonomical notes on Japanese Graphidaceae (Ascomycotina), including some new combinations. Bulletin of the National Science Museum (Tokyo), Series B (Botany) 29: 8390.Google Scholar
Nylander, W. (1891) Sertum Lichenaeae Tropicae e Labuan et Singapore. Paris: Schmidt.Google Scholar
Patwardhan, P. G. & Kulkarni, C. R. (1976) Some additions to the lichen flora of India. IV: Graphis and Graphina (family Graphidaceae). Biovigyanam 2: 123132.Google Scholar
Patwardhan, P. G. & Kulkarni, C. R. (1977) Some additions to the lichen flora of India. V. Genera Phaeographis and Phaeographina (family: Graphidaceae). Current Science 46: 720721.Google Scholar
Patwardhan, P. G. & Kulkarni, C. R. (1979 a) Some new taxa of the family Graphidaceae from Western Ghats, southwestern India. Norwegian Journal of Botany 26: 4552.Google Scholar
Patwardhan, P. G. & Kulkarni, C. R. (1979 b) The lichen genus Phaeographina (family Graphidaceae) in the Western Ghats, Southwestern India. Indian Journal of Botany 2: 132143.Google Scholar
Patwardhan, P. G. & Kulkarni, C. R. (1979 c) Three new species of Graphina of the Graphina multistriata complex. Biovigyanam 5: 58.Google Scholar
Patwardhan, P. G. & Nagarkar, M. B. (1979) Notes on some lichens from North East India I: Family Graphidaceae. Biovigyanam 5: 131138.Google Scholar
Poe, S. & Wiens, J. J. (2000) Character selection and the methodology of morphological phylogenetics. In Phylogenetic Analysis of Morphological Data (Wiens, J. J., ed.): 2036. Washington, D.C.: Smithsonian Institution Press.Google Scholar
Redinger, K. (1933) Die Graphidineen der ersten Regnell'schen Expedition nach Brasilien 1892–95. I. Glyphis, Medusulina und Sarcographa. Arkiv för Botanik 25A(13): 120.Google Scholar
Redinger, K. (1934) Die Graphidineen der ersten Regnell'schen Expedition nach Brasilien 1892–94 II. Graphina und Phaeographina. Arkiv för Botanik 26A(1): 1105.Google Scholar
Redinger, K. (1935) Die Graphidineen der ersten Regnell'schen Expedition nach Brasilien 1892–94. III. Graphis und Phaeographis, nebst einem Nachtrage zu Graphina. Arkiv för Botanik 27A(3): 1103.Google Scholar
Santesson, R. (1952) Foliicolous lichens I. A revision of the taxonomy of the obligately foliicolous, lichenized fungi. Symbolae Botanicae Upsalienses 12(1): 1590.Google Scholar
Sherwood, M. A. (1977) The ostropalean fungi. Mycotaxon 5: 1277.Google Scholar
Sipman, H. J. M. & Harris, R. C. (1989) Lichens. In Tropical Rain Forest Ecosystems Biogeographical and Ecological Studies (Ecosystems of the World 14B). (Lieth, H. & Werger, M. J. A., eds): 303309. Amsterdam: Elsevier.CrossRefGoogle Scholar
Staiger, B. (2002) Die Flechtenfamilie Graphidaceae. Studien in Richtung einer natürlicheren Gliederung. Bibliotheca Lichenologica 85: 1526.Google Scholar
Staiger, B. & Kalb, K. 1999. Acanthothecis and other graphidoid lichens with warty periphysoids or paraphysis-tips. Mycotaxon 73: 69134.Google Scholar
Staiger, B., Kalb, K. & Grube, M. (2006) Phylogeny and phenotypic variation in the lichen family Graphidaceae (Ostropomycetidae, Ascomycota). Mycological Research 110: 765772.CrossRefGoogle ScholarPubMed
Stocker-Wörgötter, E., Elix, J. A. & Grube, M. (2004) Secondary chemistry of lichen-forming fungi: chemosyndromic variation and DNA-analyses of cultures and chemotypes in the Ramalina farinacea complex. Bryologist 107: 152162.CrossRefGoogle Scholar
Swofford, D. L. (2003) PAUP*. Phylogenetic Analysis Using Parsimony (*and Other Methods). Sunderland, Massachussetts: Sinauer Associates.Google Scholar
Tehler, A. & Källersjö, M. (2001) Parmeliopsis ambigua and P. hyperopta (Parmeliaceae): species or chemotypes? Lichenologist 33: 403408.CrossRefGoogle Scholar
Vainio, E. A. (1890) Etude sur la classification et la morphologie des lichens du Brésil, I. Acta Societatis pro Fauna et Flora Fennica 7: VXXIX, 1247.Google Scholar
Vainio, E. A. (1921) Lichenes Insularum Philippinarum III. Acta Societatis Scienciarum Fennicae 15: 1368.Google Scholar
Walker, F. J. & James, P. W. (1980) A revised guide to microchemical techniques for the identification of lichen products. British Lichen Society Bulletin 46 (supplement): 1329.Google Scholar
Watson, W. (1929) The classification of lichens I & II. New Phytologist 28: 136, 85116.CrossRefGoogle Scholar
Wiley, E. O. & Mayden, R. L. (2000) The evolutionary species concept. In Species Concepts and Phylogenetic Systematics. (Wheeler, Q. & Meier, R., eds): 7089. New York: Columbia University Press.Google Scholar
Wiley, E. O., Siegel-Causey, D., Brooks, D. R. & Funk, V. A. (1991) The Compleat Cladist. Lawrence: University of Kansas Museum of Natural History.Google Scholar
Wirth, M. & Hale, M. E. Jr. (1963) The lichen family Graphidaceae in Mexico. Contributions from the U.S. National Herbarium 36: 63119.Google Scholar
Wirth, M., and Hale, M. E. Jr. (1978) Morden-Smithsonian Expedition to Dominica: the lichens (Graphidaceae). Smithsonian Contributions to Botany 40: 164.CrossRefGoogle Scholar
Zahlbruckner, A. (1907) Flechten. Spezieller Teil. In Engler, A. & Prantl, H., Die Natürlichen Pflanzenfamilien 1(1): 49249. Leipzig: Engelmann.Google Scholar
Zahlbruckner, A. (1923) Catalogus Lichenum Universalis III. Leipzig.Google Scholar
Zahlbruckner, A. (1926) Flechten. Spezieller Teil. In Die Natürlichen Pflanzenfamilien 2(8) (Engler, A. & Prantl, H., eds): 61270. Leipzig: Engelmann.Google Scholar
Zander, R. H. (2007) Paraphyly and the species concept, a reply to Ebach et al. Taxon 56: 642644.CrossRefGoogle Scholar