New Records of Lichenized and Lichenicolous Fungi in Scandinavia
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DISSERTAÇÃO Lidiane Alves Dos Santos.Pdf
UNIVERSIDADE FEDERAL DE PERNAMBUCO CENTRO DE BIOCIÊNCIAS DEPARTAMENTO DE MICOLOGIA PROGRAMA DE PÓS-GRADUAÇÃO EM BIOLOGIA DE FUNGOS LIDIANE ALVES DOS SANTOS RELAÇÕES FILOGENÉTICAS DOS GÊNEROS LECANORA ACH. E NEOPROTOPARMELIA GARIMA SINGH, LUMBSCH & I. SCHIMITT (LECANORALES, ASCOMYCOTA LIQUENIZADOS) Recife 2019 LIDIANE ALVES DOS SANTOS RELAÇÕES FILOGENÉTICAS DOS GÊNEROS LECANORA ACH. E NEOPROTOPARMELIA GARIMA SINGH, LUMBSCH & I. SCHIMITT (LECANORALES, ASCOMYCOTA LIQUENIZADOS) Dissertação apresentada ao Programa de Pós- Graduação em Biologia de Fungos do Departamento de Micologia do Centro de Biociências da Universidade Federal de Pernambuco, como parte dos requisitos para a obtenção do título de Mestre em Biologia de Fungos. Área de Concentração: Micologia Básica Orientadora: Profª. Dra. Marcela Eugenia da Silva Caceres. Coorientador: Dr. Robert Lücking. Recife 2019 Catalogação na fonte Elaine C Barroso (CRB4/1728) Santos, Lidiane Alves dos Relações filogenéticas dos gêneros Lecanora Ach. e Neoprotoparmelia Garima Singh, Lumbsch & I. Schimitt (Lecanorales, Ascomycota Liquenizados) / Lidiane Alves dos Santos- 2019. 52 folhas: il., fig., tab. Orientadora: Marcela Eugênia da Silva Cáceres Coorientador: Robert Lücking Dissertação (mestrado) – Universidade Federal de Pernambuco. Centro de Biociências. Programa de Pós-Graduação em Biologia de Fungos. Recife, 2019. Inclui referências 1. Fungos liquenizados 2. Filogenia 3. Metabólitos secundários I. Cáceres, Marcela Eugênia da Silva (orient.) II. Lücking, Robert (coorient.) III. Título 579.5 CDD (22.ed.) UFPE/CB-2019-312 LIDIANE ALVES DOS SANTOS RELAÇÕES FILOGENÉTICAS DOS GÊNEROS LECANORA ACH. E NEOPROTOPARMELIA GARIMA SINGH, LUMBSCH & I. SCHIMITT (LECANORALES, ASCOMYCOTA LIQUENIZADOS) Dissertação apresentada ao Programa de Pós- Graduação em Biologia de Fungos do Departamento de Micologia do Centro de Biociências da Universidade Federal de Pernambuco, como parte dos requisitos para a obtenção do título de Mestre em Biologia de Fungos. -
Lichen Functional Trait Variation Along an East-West Climatic Gradient in Oregon and Among Habitats in Katmai National Park, Alaska
AN ABSTRACT OF THE THESIS OF Kaleigh Spickerman for the degree of Master of Science in Botany and Plant Pathology presented on June 11, 2015 Title: Lichen Functional Trait Variation Along an East-West Climatic Gradient in Oregon and Among Habitats in Katmai National Park, Alaska Abstract approved: ______________________________________________________ Bruce McCune Functional traits of vascular plants have been an important component of ecological studies for a number of years; however, in more recent times vascular plant ecologists have begun to formalize a set of key traits and universal system of trait measurement. Many recent studies hypothesize global generality of trait patterns, which would allow for comparison among ecosystems and biomes and provide a foundation for general rules and theories, the so-called “Holy Grail” of ecology. However, the majority of these studies focus on functional trait patterns of vascular plants, with a minority examining the patterns of cryptograms such as lichens. Lichens are an important component of many ecosystems due to their contributions to biodiversity and their key ecosystem services, such as contributions to mineral and hydrological cycles and ecosystem food webs. Lichens are also of special interest because of their reliance on atmospheric deposition for nutrients and water, which makes them particularly sensitive to air pollution. Therefore, they are often used as bioindicators of air pollution, climate change, and general ecosystem health. This thesis examines the functional trait patterns of lichens in two contrasting regions with fundamentally different kinds of data. To better understand the patterns of lichen functional traits, we examined reproductive, morphological, and chemical trait variation along precipitation and temperature gradients in Oregon. -
H. Thorsten Lumbsch VP, Science & Education the Field Museum 1400
H. Thorsten Lumbsch VP, Science & Education The Field Museum 1400 S. Lake Shore Drive Chicago, Illinois 60605 USA Tel: 1-312-665-7881 E-mail: [email protected] Research interests Evolution and Systematics of Fungi Biogeography and Diversification Rates of Fungi Species delimitation Diversity of lichen-forming fungi Professional Experience Since 2017 Vice President, Science & Education, The Field Museum, Chicago. USA 2014-2017 Director, Integrative Research Center, Science & Education, The Field Museum, Chicago, USA. Since 2014 Curator, Integrative Research Center, Science & Education, The Field Museum, Chicago, USA. 2013-2014 Associate Director, Integrative Research Center, Science & Education, The Field Museum, Chicago, USA. 2009-2013 Chair, Dept. of Botany, The Field Museum, Chicago, USA. Since 2011 MacArthur Associate Curator, Dept. of Botany, The Field Museum, Chicago, USA. 2006-2014 Associate Curator, Dept. of Botany, The Field Museum, Chicago, USA. 2005-2009 Head of Cryptogams, Dept. of Botany, The Field Museum, Chicago, USA. Since 2004 Member, Committee on Evolutionary Biology, University of Chicago. Courses: BIOS 430 Evolution (UIC), BIOS 23410 Complex Interactions: Coevolution, Parasites, Mutualists, and Cheaters (U of C) Reading group: Phylogenetic methods. 2003-2006 Assistant Curator, Dept. of Botany, The Field Museum, Chicago, USA. 1998-2003 Privatdozent (Assistant Professor), Botanical Institute, University – GHS - Essen. Lectures: General Botany, Evolution of lower plants, Photosynthesis, Courses: Cryptogams, Biology -
Different Diversification Histories in Tropical and Temperate Lineages in the Ascomycete Subfamily Protoparmelioideae (Parmeliaceae)
A peer-reviewed open-access journal MycoKeys 36: 1–19 Different(2018) diversification histories in tropical and temperate lineages... 1 doi: 10.3897/mycokeys.36.22548 RESEARCH ARTICLE MycoKeys http://mycokeys.pensoft.net Launched to accelerate biodiversity research Different diversification histories in tropical and temperate lineages in the ascomycete subfamily Protoparmelioideae (Parmeliaceae) Garima Singh1, Francesco Dal Grande1, Jan Schnitzler2, Markus Pfenninger1, Imke Schmitt1,3 1 Senckenberg Biodiversity and Climate Research Centre (SBiK-F), Frankfurt am Main, Germany 2 Department of Molecular Evolution and Plant Systematics, Institute of Biology, Leipzig University, Germany 3 Department of Biological Sciences, Institute of Ecology, Evolution and Diversity, Goethe Universität Frankfurt am Main, Germany Corresponding author: Garima Singh ([email protected]) Academic editor: P. Divakar | Received 27 November 2018 | Accepted 19 June 2018 | Published 2 July 2018 Citation: Singh G, Grande FD, Schnitzler J, Pfenninger M, Schmitt I (2018) Different diversification histories in tropical and temperate lineages in the ascomycete subfamily Protoparmelioideae (Parmeliaceae). MycoKeys 36: 1–19. https://doi.org/10.3897/mycokeys.36.22548 Abstract Background: Environment and geographic processes affect species’ distributions as well as evolutionary processes, such as clade diversification. Estimating the time of origin and diversification of organisms helps us understand how climate fluctuations in the past might have influenced the diversification and present distribution of species. Complementing divergence dating with character evolution could indicate how key innovations have facilitated the diversification of species. Methods: We estimated the divergence times within the newly recognised subfamily Protoparmel- ioideae (Ascomycota) using a multilocus dataset to assess the temporal context of diversification events. We reconstructed ancestral habitats and substrate using a species tree generated in *Beast. -
One Hundred New Species of Lichenized Fungi: a Signature of Undiscovered Global Diversity
Phytotaxa 18: 1–127 (2011) ISSN 1179-3155 (print edition) www.mapress.com/phytotaxa/ Monograph PHYTOTAXA Copyright © 2011 Magnolia Press ISSN 1179-3163 (online edition) PHYTOTAXA 18 One hundred new species of lichenized fungi: a signature of undiscovered global diversity H. THORSTEN LUMBSCH1*, TEUVO AHTI2, SUSANNE ALTERMANN3, GUILLERMO AMO DE PAZ4, ANDRÉ APTROOT5, ULF ARUP6, ALEJANDRINA BÁRCENAS PEÑA7, PAULINA A. BAWINGAN8, MICHEL N. BENATTI9, LUISA BETANCOURT10, CURTIS R. BJÖRK11, KANSRI BOONPRAGOB12, MAARTEN BRAND13, FRANK BUNGARTZ14, MARCELA E. S. CÁCERES15, MEHTMET CANDAN16, JOSÉ LUIS CHAVES17, PHILIPPE CLERC18, RALPH COMMON19, BRIAN J. COPPINS20, ANA CRESPO4, MANUELA DAL-FORNO21, PRADEEP K. DIVAKAR4, MELIZAR V. DUYA22, JOHN A. ELIX23, ARVE ELVEBAKK24, JOHNATHON D. FANKHAUSER25, EDIT FARKAS26, LIDIA ITATÍ FERRARO27, EBERHARD FISCHER28, DAVID J. GALLOWAY29, ESTER GAYA30, MIREIA GIRALT31, TREVOR GOWARD32, MARTIN GRUBE33, JOSEF HAFELLNER33, JESÚS E. HERNÁNDEZ M.34, MARÍA DE LOS ANGELES HERRERA CAMPOS7, KLAUS KALB35, INGVAR KÄRNEFELT6, GINTARAS KANTVILAS36, DOROTHEE KILLMANN28, PAUL KIRIKA37, KERRY KNUDSEN38, HARALD KOMPOSCH39, SERGEY KONDRATYUK40, JAMES D. LAWREY21, ARMIN MANGOLD41, MARCELO P. MARCELLI9, BRUCE MCCUNE42, MARIA INES MESSUTI43, ANDREA MICHLIG27, RICARDO MIRANDA GONZÁLEZ7, BIBIANA MONCADA10, ALIFERETI NAIKATINI44, MATTHEW P. NELSEN1, 45, DAG O. ØVSTEDAL46, ZDENEK PALICE47, KHWANRUAN PAPONG48, SITTIPORN PARNMEN12, SERGIO PÉREZ-ORTEGA4, CHRISTIAN PRINTZEN49, VÍCTOR J. RICO4, EIMY RIVAS PLATA1, 50, JAVIER ROBAYO51, DANIA ROSABAL52, ULRIKE RUPRECHT53, NORIS SALAZAR ALLEN54, LEOPOLDO SANCHO4, LUCIANA SANTOS DE JESUS15, TAMIRES SANTOS VIEIRA15, MATTHIAS SCHULTZ55, MARK R. D. SEAWARD56, EMMANUËL SÉRUSIAUX57, IMKE SCHMITT58, HARRIE J. M. SIPMAN59, MOHAMMAD SOHRABI 2, 60, ULRIK SØCHTING61, MAJBRIT ZEUTHEN SØGAARD61, LAURENS B. SPARRIUS62, ADRIANO SPIELMANN63, TOBY SPRIBILLE33, JUTARAT SUTJARITTURAKAN64, ACHRA THAMMATHAWORN65, ARNE THELL6, GÖRAN THOR66, HOLGER THÜS67, EINAR TIMDAL68, CAMILLE TRUONG18, ROMAN TÜRK69, LOENGRIN UMAÑA TENORIO17, DALIP K. -
Lichens and Associated Fungi from Glacier Bay National Park, Alaska
The Lichenologist (2020), 52,61–181 doi:10.1017/S0024282920000079 Standard Paper Lichens and associated fungi from Glacier Bay National Park, Alaska Toby Spribille1,2,3 , Alan M. Fryday4 , Sergio Pérez-Ortega5 , Måns Svensson6, Tor Tønsberg7, Stefan Ekman6 , Håkon Holien8,9, Philipp Resl10 , Kevin Schneider11, Edith Stabentheiner2, Holger Thüs12,13 , Jan Vondrák14,15 and Lewis Sharman16 1Department of Biological Sciences, CW405, University of Alberta, Edmonton, Alberta T6G 2R3, Canada; 2Department of Plant Sciences, Institute of Biology, University of Graz, NAWI Graz, Holteigasse 6, 8010 Graz, Austria; 3Division of Biological Sciences, University of Montana, 32 Campus Drive, Missoula, Montana 59812, USA; 4Herbarium, Department of Plant Biology, Michigan State University, East Lansing, Michigan 48824, USA; 5Real Jardín Botánico (CSIC), Departamento de Micología, Calle Claudio Moyano 1, E-28014 Madrid, Spain; 6Museum of Evolution, Uppsala University, Norbyvägen 16, SE-75236 Uppsala, Sweden; 7Department of Natural History, University Museum of Bergen Allégt. 41, P.O. Box 7800, N-5020 Bergen, Norway; 8Faculty of Bioscience and Aquaculture, Nord University, Box 2501, NO-7729 Steinkjer, Norway; 9NTNU University Museum, Norwegian University of Science and Technology, NO-7491 Trondheim, Norway; 10Faculty of Biology, Department I, Systematic Botany and Mycology, University of Munich (LMU), Menzinger Straße 67, 80638 München, Germany; 11Institute of Biodiversity, Animal Health and Comparative Medicine, College of Medical, Veterinary and Life Sciences, University of Glasgow, Glasgow G12 8QQ, UK; 12Botany Department, State Museum of Natural History Stuttgart, Rosenstein 1, 70191 Stuttgart, Germany; 13Natural History Museum, Cromwell Road, London SW7 5BD, UK; 14Institute of Botany of the Czech Academy of Sciences, Zámek 1, 252 43 Průhonice, Czech Republic; 15Department of Botany, Faculty of Science, University of South Bohemia, Branišovská 1760, CZ-370 05 České Budějovice, Czech Republic and 16Glacier Bay National Park & Preserve, P.O. -
Composition and Specialization of the Lichen Functional Traits in a Primeval Forest—Does Ecosystem Organization Level Matter?
Article Composition and Specialization of the Lichen Functional Traits in a Primeval Forest—Does Ecosystem Organization Level Matter? Anna Łubek 1,*, Martin Kukwa 2 , Bogdan Jaroszewicz 3 and Patryk Czortek 3 1 Division of Environmental Biology, Institute of Biology, The Jan Kochanowski University in Kielce, Uniwersytecka 7, PL-25-406 Kielce, Poland 2 Department of Plant Taxonomy and Nature Conservation, Faculty of Biology, University of Gda´nsk, Wita Stwosza 59, PL-80-308 Gda´nsk,Poland; [email protected] 3 Białowieza˙ Geobotanical Station, Faculty of Biology, University of Warsaw, Sportowa 19, PL-17-230 Białowieza,˙ Poland; [email protected] (B.J.); [email protected] (P.C.) * Correspondence: [email protected] Abstract: Current trends emphasize the importance of the examination of the functional composition of lichens, which may provide information on the species realized niche diversity and community assembly processes, thus enabling one to understand the specific adaptations of lichens and their interaction with the environment. We analyzed the distribution and specialization of diverse mor- phological, anatomical and chemical (lichen secondary metabolites) traits in lichen communities in a close-to-natural forest of lowland Europe. We considered these traits in relation to three levels of forest ecosystem organization: forest communities, phorophyte species and substrates, in order to recognize the specialization of functional traits to different levels of the forest complexity. Traits related to the sexual reproduction of mycobionts (i.e., ascomata types: lecanoroid apothecia, lecideoid Citation: Łubek, A.; Kukwa, M.; apothecia, arthonioid apothecia, lirellate apothecia, stalked apothecia and perithecia) and asexual Jaroszewicz, B.; Czortek, P. -
<I> Lecanoromycetes</I> of Lichenicolous Fungi Associated With
Persoonia 39, 2017: 91–117 ISSN (Online) 1878-9080 www.ingentaconnect.com/content/nhn/pimj RESEARCH ARTICLE https://doi.org/10.3767/persoonia.2017.39.05 Phylogenetic placement within Lecanoromycetes of lichenicolous fungi associated with Cladonia and some other genera R. Pino-Bodas1,2, M.P. Zhurbenko3, S. Stenroos1 Key words Abstract Though most of the lichenicolous fungi belong to the Ascomycetes, their phylogenetic placement based on molecular data is lacking for numerous species. In this study the phylogenetic placement of 19 species of cladoniicolous species lichenicolous fungi was determined using four loci (LSU rDNA, SSU rDNA, ITS rDNA and mtSSU). The phylogenetic Pilocarpaceae analyses revealed that the studied lichenicolous fungi are widespread across the phylogeny of Lecanoromycetes. Protothelenellaceae One species is placed in Acarosporales, Sarcogyne sphaerospora; five species in Dactylosporaceae, Dactylo Scutula cladoniicola spora ahtii, D. deminuta, D. glaucoides, D. parasitica and Dactylospora sp.; four species belong to Lecanorales, Stictidaceae Lichenosticta alcicorniaria, Epicladonia simplex, E. stenospora and Scutula epiblastematica. The genus Epicladonia Stictis cladoniae is polyphyletic and the type E. sandstedei belongs to Leotiomycetes. Phaeopyxis punctum and Bachmanniomyces uncialicola form a well supported clade in the Ostropomycetidae. Epigloea soleiformis is related to Arthrorhaphis and Anzina. Four species are placed in Ostropales, Corticifraga peltigerae, Cryptodiscus epicladonia, C. galaninae and C. cladoniicola -
Colville National Forest Non-Vascular Plant Survey and Reference Collection Interagency Special Status/Sensitive Species Program Final Report, December 3, 2009
Colville National Forest Non-vascular Plant Survey and Reference Collection Interagency Special Status/Sensitive Species Program Final Report, December 3, 2009 Kathy Ahlenslager and Amy Cabral Colville National Forest, 765 S. Main, Colville, WA, 509-684-7178, [email protected]. Introduction Through this project the populations of 12 rare nonvascular taxa were revisited at seven locations on the Colville National Forest. The rarity status for each is displayed on Table 1. These taxa were included on the Interagency Special Status/Sensitive Species Program (ISSSSP) High Priority Special Status and Sensitive Species List, Regional Forester’s Special Status Species List, or a Washington Natural Heritage Program (WNHP) list. Bryophytes were identified by Erica Heinlen and lichens by Dr. Katherine Glew. In 2009 23 collections of lichens were identified to the 18 taxa in Appendix A and were gathered from 15 sites, as displayed on Appendix B. In addition, 132 vouchers of mosses were identified to the 74 taxa in Appendix C and were collected from 37 locations as shown on Appendix D. Table 1. Rankings of Rare Nonvascular Taxa Documented from the Colville National Forest. (Note: FSSO = Forest Service Sensitive in Oregon, FSSW = Forest Service Sensitive in Washington, G = Global Ranking, S = Washington State Ranking.) Taxa Status on Regional Status (WNHP ISSSSP High Forester’s 20091 or Nature Priority Special Status Serve 20092) Special Status Species List and Sensitive (2008) Species List (2007) Liverworts Calypogeia sphagnicola First Priority -
No. 20: Further Notes on Species from the Eastern North America
Opuscula Philolichenum, 15: 105-131. 2016. *pdf effectively published online 15November2016 via (http://sweetgum.nybg.org/philolichenum/) Studies in Lichens and Lichenicolous Fungi – No. 20: Further notes on species from the eastern North America JAMES C. LENDEMER1 AND RICHARD C. HARRIS2 ABSTRACT. – Arthothelium lichenale is placed in synonymy with Mycoporum compositum. The morphological distinctions between Collema pustulatum and Leptogium apalachense are discussed and the North American distributions of the species are revised. The distribution of Lecidea roseotincta in North America is extended to include the central and southern Appalachian Mountains. The distribution of Lecidella subviridis is expanded in northeastern North America. Pyrenula reebiae is placed in synonymy with P. leucostoma and both species are illustrated and discussed. Pyrenula shirabeicola is removed from synonymy with P. pseudobufonia and both species are illustrated and discussed. The following taxa are newly reported from North America: Calvitimela cuprea (Canada, Newfoundland & Labrador), Hypotrachyna consimilis (U.S.A., North Carolina), Schismatomma graphidioides (U.S.A., Alabama and New Jersey). KEYWORDS. – Biogeography, Collemataceae, North American Checklist, Lecanoraceae, Lecideaceae, Parmeliaceae, Pyrenulaceae, sterile crust. INTRODUCTION As a result of fieldwork carried out throughout eastern North America, particularly in the southern Appalachian Mountains, we have studied new collections that prompted us to reexamine our understanding of recognized species, their distributions, and ecologies. Routine curation of the herbarium at the New York Botanical Garden led to similar studies, as has the identification of specimens that were newly donated to the herbarium. While such studies typically result in discrete taxonomic or floristic publications, there are often also brief notes that merit publication but are not easily accommodated in a standalone contribution. -
A New Species of Protoparmelia (Lichenized Ascomycota) from Australia
Nova Hedwigia 89 3—4 355—360 Stuttgart, November 2009 A new species of Protoparmelia (lichenized Ascomycota) from Australia by John A. Elix1 and Gintaras Kantvilas2 1Research School of Chemistry, Building 33, Australian National University, Canberra, ACT 0200, Australia 2Tasmanian Herbarium, Private Bag 4, Hobart, Tasmania 7001, Australia With 1 figure Elix, J.A. & G. Kantvilas (2009): A new species of Protoparmelia (lichenized Ascomycota) from Australia. - Nova Hedwigia 89: 355–360 Abstract: The new species, Protoparmelia nebulosa Elix & Kantvilas, based on material from Western Australia and New South Wales, is described. It is characterized chiefly by an isidiate thallus, and by the presence of constipatic, isomyelochroic, myelochroic and protoconstipatic acids. The Australasian species, P. pulchra Diederich, Aptroot & Sérus., is recorded from Tasmania for the first time. eschweizerbartxxx author Key words: Australia, fatty acids, lichens, Tasmania, taxonomy. Introduction Protoparmelia M.Choisy is a cosmopolitan genus with approximately 25 species recognized worldwide (Farr & Rossman 2009, Robert et al. 2009). It is characterized by a crustose thallus, typically with a cortex of anticlinal hyphae, a coccal green photobiont, lecanorine apothecia with a meristematic proper excipulum, eight-spored asci approximating to the Lecanora-type (after Hafellner 1984, Aptroot et al. 1997a), with a well-developed amyloid tholus with or without an ocular chamber but always with a distinct, non-amyloid masse axiale, sparsely branched paraphyses, and simple (rarely 1-septate when old), hyaline, narrowly ellipsoidal to oblong ascospores, 7–17 × 2–7 μm. It is superficially similar to Lecanora but separated from that genus by the grey-brown to reddish-brown thallus, the generally smaller, narrower ascospores and straight conidia. -
Effects of Growth Media on the Diversity of Culturable Fungi from Lichens
molecules Article Effects of Growth Media on the Diversity of Culturable Fungi from Lichens Lucia Muggia 1,*,†, Theodora Kopun 2,† and Martin Grube 2 1 Department of Life Sciences, University of Trieste, via Giorgieri 10, 34127 Trieste, Italy 2 Institute of Plant Science, Karl-Franzens University of Graz, Holteigasse 6, 8010 Graz, Austria; [email protected] (T.K.); [email protected] (M.G.) * Correspondence: [email protected] or [email protected]; Tel.: +39-04-0558-8825 † These authors contributed equally to the work. Academic Editor: Joel Boustie Received: 1 March 2017; Accepted: 11 May 2017; Published: 17 May 2017 Abstract: Microscopic and molecular studies suggest that lichen symbioses contain a plethora of associated fungi. These are potential producers of novel bioactive compounds, but strains isolated on standard media usually represent only a minor subset of these fungi. By using various in vitro growth conditions we are able to modulate and extend the fraction of culturable lichen-associated fungi. We observed that the presence of iron, glucose, magnesium and potassium in growth media is essential for the successful isolation of members from different taxonomic groups. According to sequence data, most isolates besides the lichen mycobionts belong to the classes Dothideomycetes and Eurotiomycetes. With our approach we can further explore the hidden fungal diversity in lichens to assist in the search of novel compounds. Keywords: Dothideomycetes; Eurotiomycetes; Leotiomycetes; nuclear ribosomal subunits DNA; nutrients; Sordariomycetes 1. Introduction Lichens are self-sustaining symbiotic associations of specialized fungi (the mycobionts), and green algae or cyanobacteria (the photobionts), which are located extracellularly within a matrix of fungal hyphae and from which the fungi derive carbon nutrition [1].