Splitting Or Synonymizing – Genus Concept and Taxonomy Exemplified by the Parmeliaceae in the Nordic Region
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Supplementary Material for Nelson, P. R., B. Mccune & D. K. Swanson
Supplementary material for Nelson, P. R., B. McCune & D. K. Swanson. 2015. Lichen traits and species as indicators of vegetation and environment. The Bryologist 118(3): XX–XX. Supplementary Table S2. Trait matrix (alphabetical by species). “1” indicates a species possesses that trait. cladoniiform Filamentous Squamulose Cyano Erect Appressed 3D s branched Tripartite Fruticose Terricole Epiphyte Lignicole Saxicole p soredia lobules Foliose Simple foliose Green rawlin isida foliose Lichen Species Subspecies richly Only g Alectoria ochroleuca 1 0 0 0 1 0 0 0 1 0 0 0 0 1 0 0 0 0 Allantoparmelia almquistii 1 0 0 1 0 0 0 1 0 0 0 0 1 0 0 0 0 0 Allantoparmelia alpicola 1 0 0 1 0 0 0 1 0 0 0 0 1 0 0 0 0 0 Allocetraria madreporiformis 1 0 0 0 1 0 0 0 1 1 0 0 0 0 0 0 0 0 Anaptychia bryorum 1 0 0 0 1 0 0 1 0 0 0 0 1 0 0 0 0 0 Arctoparmelia centrifuga 1 0 0 1 0 0 0 1 0 0 0 0 1 0 0 0 0 0 Arctoparmelia incurva 1 0 0 1 0 0 0 1 0 0 0 0 1 0 0 1 0 0 Arctoparmelia separata 1 0 0 1 0 0 0 1 0 0 0 0 1 0 0 0 0 0 Arctoparmelia subcentrifuga 1 0 0 1 0 0 0 1 0 0 0 0 1 0 0 0 0 0 Asahinea chrysantha 1 0 0 1 0 0 0 1 0 0 0 0 0 0 1 0 0 0 Baeomyces carneus 1 0 0 0 1 0 0 0 1 1 0 0 0 0 0 0 0 0 Baeomyces placophyllus 1 0 0 0 1 0 0 0 1 1 0 0 0 0 0 0 0 0 Baeomyces rufus 1 0 0 0 1 0 0 0 1 1 0 0 0 0 0 0 0 0 Blennothallia crispa 0 1 0 0 1 0 0 1 0 0 0 0 0 0 1 0 1 0 Brodoa oroarctica 1 0 0 1 0 0 0 1 0 0 0 0 0 0 1 0 0 0 Bryocaulon divergens 1 0 0 0 1 0 0 0 1 0 0 0 0 1 0 0 0 0 Bryoria capillaris 1 0 0 0 0 1 0 0 1 0 0 0 0 1 0 0 0 0 Bryoria chalybeiformis 1 0 0 0 0 1 0 0 1 0 0 0 0 -
Phylogeny of the Cetrarioid Core (Parmeliaceae) Based on Five
The Lichenologist 41(5): 489–511 (2009) © 2009 British Lichen Society doi:10.1017/S0024282909990090 Printed in the United Kingdom Phylogeny of the cetrarioid core (Parmeliaceae) based on five genetic markers Arne THELL, Filip HÖGNABBA, John A. ELIX, Tassilo FEUERER, Ingvar KÄRNEFELT, Leena MYLLYS, Tiina RANDLANE, Andres SAAG, Soili STENROOS, Teuvo AHTI and Mark R. D. SEAWARD Abstract: 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, -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. -
Diversity and Distribution of Lichen-Associated Fungi in the Ny-Ålesund Region (Svalbard, High Arctic) As Revealed by 454 Pyrosequencing
www.nature.com/scientificreports OPEN Diversity and distribution of lichen- associated fungi in the Ny-Ålesund Region (Svalbard, High Arctic) as Received: 31 March 2015 Accepted: 20 August 2015 revealed by 454 pyrosequencing Published: 14 October 2015 Tao Zhang1, Xin-Li Wei2, Yu-Qin Zhang1, Hong-Yu Liu1 & Li-Yan Yu1 This study assessed the diversity and distribution of fungal communities associated with seven lichen species in the Ny-Ålesund Region (Svalbard, High Arctic) using Roche 454 pyrosequencing with fungal-specific primers targeting the internal transcribed spacer (ITS) region of the ribosomal rRNA gene. Lichen-associated fungal communities showed high diversity, with a total of 42,259 reads belonging to 370 operational taxonomic units (OTUs) being found. Of these OTUs, 294 belonged to Ascomycota, 54 to Basidiomycota, 2 to Zygomycota, and 20 to unknown fungi. Leotiomycetes, Dothideomycetes, and Eurotiomycetes were the major classes, whereas the dominant orders were Helotiales, Capnodiales, and Chaetothyriales. Interestingly, most fungal OTUs were closely related to fungi from various habitats (e.g., soil, rock, plant tissues) in the Arctic, Antarctic and alpine regions, which suggests that living in association with lichen thalli may be a transient stage of life cycle for these fungi and that long-distance dispersal may be important to the fungi in the Arctic. In addition, host-related factors shaped the lichen-associated fungal communities in this region. Taken together, these results suggest that lichens thalli act as reservoirs of diverse fungi from various niches, which may improve our understanding of fungal evolution and ecology in the Arctic. The Arctic is one of the most pristine regions of the planet, and its environment exhibits extreme condi- tions (e.g., low temperature, strong winds, permafrost, and long periods of darkness and light) and offers unique opportunities to explore extremophiles. -
The Puzzle of Lichen Symbiosis
Digital Comprehensive Summaries of Uppsala Dissertations from the Faculty of Science and Technology 1503 The puzzle of lichen symbiosis Pieces from Thamnolia IOANA ONUT, -BRÄNNSTRÖM ACTA UNIVERSITATIS UPSALIENSIS ISSN 1651-6214 ISBN 978-91-554-9887-0 UPPSALA urn:nbn:se:uu:diva-319639 2017 Dissertation presented at Uppsala University to be publicly examined in Lindhalsalen, EBC, Norbyvägen 14, Uppsala, Thursday, 1 June 2017 at 09:15 for the degree of Doctor of Philosophy. The examination will be conducted in English. Faculty examiner: Associate Professor Anne Pringle (University of Wisconsin-Madison, Department of Botany). Abstract Onuț-Brännström, I. 2017. The puzzle of lichen symbiosis. Pieces from Thamnolia. Digital Comprehensive Summaries of Uppsala Dissertations from the Faculty of Science and Technology 1503. 62 pp. Uppsala: Acta Universitatis Upsaliensis. ISBN 978-91-554-9887-0. Symbiosis brought important evolutionary novelties to life on Earth. Lichens, the symbiotic entities formed by fungi, photosynthetic organisms and bacteria, represent an example of a successful adaptation in surviving hostile environments. Yet many aspects of the lichen symbiosis remain unexplored. This thesis aims at bringing insights into lichen biology and the importance of symbiosis in adaptation. I am using as model system a successful colonizer of tundra and alpine environments, the worm lichens Thamnolia, which seem to only reproduce vegetatively through symbiotic propagules. When the genetic architecture of the mating locus of the symbiotic fungal partner was analyzed with genomic and transcriptomic data, a sexual self-incompatible life style was revealed. However, a screen of the mating types ratios across natural populations detected only one of the mating types, suggesting that Thamnolia has no potential for sexual reproduction because of lack of mating partners. -
New Or Interesting Lichens and Lichenicolous Fungi from Belgium, Luxembourg and Northern France
New or interesting lichens and lichenicolous fungi from Belgium, Luxembourg and northern France. X Emmanuël SÉRUSIAUX1, Paul DIEDERICH2, Damien ERTZ3, Maarten BRAND4 & Pieter VAN DEN BOOM5 1 Plant Taxonomy and Conservation Biology Unit, University of Liège, Sart Tilman B22, B-4000 Liège, Belgique ([email protected]) 2 Musée national d’histoire naturelle, 25 rue Munster, L-2160 Luxembourg, Luxembourg ([email protected]) 3 Jardin Botanique National de Belgique, Domaine de Bouchout, B-1860 Meise, Belgium ([email protected]) 4 Klipperwerf 5, NL-2317 DX Leiden, the Netherlands ([email protected]) 5 Arafura 16, NL-5691 JA Son, the Netherlands ([email protected]) Sérusiaux, E., P. Diederich, D. Ertz, M. Brand & P. van den Boom, 2006. New or interesting lichens and lichenicolous fungi from Belgium, Luxembourg and northern France. X. Bul- letin de la Société des naturalistes luxembourgeois 107 : 63-74. Abstract. Review of recent literature and studies on large and mainly recent collections of lichens and lichenicolous fungi led to the addition of 35 taxa to the flora of Belgium, Lux- embourg and northern France: Abrothallus buellianus, Absconditella delutula, Acarospora glaucocarpa var. conspersa, Anema nummularium, Anisomeridium ranunculosporum, Artho- nia epiphyscia, A. punctella, Bacidia adastra, Brodoa atrofusca, Caloplaca britannica, Cer- cidospora macrospora, Chaenotheca laevigata, Collemopsidium foveolatum, C. sublitorale, Coppinsia minutissima, Cyphelium inquinans, Involucropyrenium squamulosum, Lecania fructigena, Lecanora conferta, L. pannonica, L. xanthostoma, Lecidea variegatula, Mica- rea micrococca, Micarea subviridescens, M. vulpinaris, Opegrapha prosodea, Parmotrema stuppeum, Placynthium stenophyllum var. isidiatum, Porpidia striata, Pyrenidium actinellum, Thelopsis rubella, Toninia physaroides, Tremella coppinsii, Tubeufia heterodermiae, Verru- caria acrotella and Vezdaea stipitata. -
Phylogenetic Diversity of the Lichenized Algal Genus Trebouxia (Trebouxiophyceae, Chlorophyta): a New Lineage and Novel Insights
applyparastyle “fig//caption/p[1]” parastyle “FigCapt” Botanical Journal of the Linnean Society, 2020, XX, 1–9. With 3 figures. Downloaded from https://academic.oup.com/botlinnean/article-abstract/doi/10.1093/botlinnean/boaa050/5873858 by National and University Library of Iceland user on 05 August 2020 Phylogenetic diversity of the lichenized algal genus Trebouxia (Trebouxiophyceae, Chlorophyta): a new lineage and novel insights from fungal-algal association Keywords=Keywords=Keywords_First=Keywords patterns of Icelandic cetrarioid lichens (Parmeliaceae, HeadA=HeadB=HeadA=HeadB/HeadA Ascomycota) HeadB=HeadC=HeadB=HeadC/HeadB HeadC=HeadD=HeadC=HeadD/HeadC MAONIAN XU1, , HUGO DE BOER2, ELIN SOFFIA OLAFSDOTTIR1, Extract3=HeadA=Extract1=HeadA SESSELJA OMARSDOTTIR1 and STARRI HEIDMARSSON3,*, REV_HeadA=REV_HeadB=REV_HeadA=REV_HeadB/HeadA REV_HeadB=REV_HeadC=REV_HeadB=REV_HeadC/HeadB 1Faculty of Pharmaceutical Sciences, University of Iceland, Hagi, Hofsvallagata 53, IS-107 Reykjavik, REV_HeadC=REV_HeadD=REV_HeadC=REV_HeadD/HeadC Iceland 2Natural History Museum, University of Oslo, Sars’ gate 1, NO-0562 Oslo, Norway REV_Extract3=REV_HeadA=REV_Extract1=REV_HeadA 3Icelandic Institute of Natural History, Akureyri Division, IS-600 Akureyri, Iceland BOR_HeadA=BOR_HeadB=BOR_HeadA=BOR_HeadB/HeadA BOR_HeadB=BOR_HeadC=BOR_HeadB=BOR_HeadC/HeadB Received 3 June 2019; revised 20 March 2020; accepted for publication 11 June 2020 BOR_HeadC=BOR_HeadD=BOR_HeadC=BOR_HeadD/HeadC BOR_Extract3=BOR_HeadA=BOR_Extract1=BOR_HeadA EDI_HeadA=EDI_HeadB=EDI_HeadA=EDI_HeadB/HeadA -
A New Species of Allocetraria (Parmeliaceae, Ascomycota) in China
The Lichenologist 47(1): 31–34 (2015) 6 British Lichen Society, 2015 doi:10.1017/S0024282914000528 A new species of Allocetraria (Parmeliaceae, Ascomycota) in China Rui-Fang WANG, Xin-Li WEI and Jiang-Chun WEI Abstract: Allocetraria yunnanensis R. F. Wang, X. L. Wei & J. C. Wei is described as a new species from the Yunnan Province of China, and is characterized by having a shiny upper surface, strongly wrinkled lower surface, and marginal pseudocyphellae present on the lower side in the form of a white continuous line or spot. The phylogenetic analysis based on nrDNA ITS sequences suggests that the new species is related to A. sinensis X. Q. Gao. Key words: Allocetraria yunnanensis, lichen, taxonomy Accepted for publication 26 June 2014 Introduction genus, as all ten species have been reported there (Kurokawa & Lai 1991; Thell et al. The lichenized genus Allocetraria Kurok. & 1995; Randlane et al. 2001; Wang et al. M. J. Lai was described in 1991, with a new 2014). During our taxonomic study of Allo- species A. isidiigera Kurok. & M. J. Lai, and cetraria, a new species was found. two new combinations: A. ambigua (C. Bab.) Kurok. & M. J. Lai and A. stracheyi (C. Bab.) Kurok. & M. J. Lai (Kurokawa & Lai 1991). The main distribution area of Allocetraria Materials and Methods species was reported to be in the Himalayas, A dissecting microscope (ZEISS Stemi SV11) and com- including China, India, and Nepal. pound microscope (ZEISS Axioskop 2 plus) were used Allocetraria is characterized by dichoto- to study the morphology and anatomy of the specimens. Colour test reagents [10% aqueous KOH, saturated mously or subdichotomously branched lobes aqueous Ca(OCl)2, and concentrated alcoholic p- and a foliose to suberect or erect thallus with phenylenediamine] and thin-layer chromatography sparse rhizines, angular to sublinear pseudo- (TLC, solvent system C) were used for the detection cyphellae, palisade plectenchymatous upper of lichen substances (Culberson & Kristinsson 1970; Culberson 1972). -
1307 Fungi Representing 1139 Infrageneric Taxa, 317 Genera and 66 Families ⇑ Jolanta Miadlikowska A, , Frank Kauff B,1, Filip Högnabba C, Jeffrey C
Molecular Phylogenetics and Evolution 79 (2014) 132–168 Contents lists available at ScienceDirect Molecular Phylogenetics and Evolution journal homepage: www.elsevier.com/locate/ympev A multigene phylogenetic synthesis for the class Lecanoromycetes (Ascomycota): 1307 fungi representing 1139 infrageneric taxa, 317 genera and 66 families ⇑ Jolanta Miadlikowska a, , Frank Kauff b,1, Filip Högnabba c, Jeffrey C. Oliver d,2, Katalin Molnár a,3, Emily Fraker a,4, Ester Gaya a,5, Josef Hafellner e, Valérie Hofstetter a,6, Cécile Gueidan a,7, Mónica A.G. Otálora a,8, Brendan Hodkinson a,9, Martin Kukwa f, Robert Lücking g, Curtis Björk h, Harrie J.M. Sipman i, Ana Rosa Burgaz j, Arne Thell k, Alfredo Passo l, Leena Myllys c, Trevor Goward h, Samantha Fernández-Brime m, Geir Hestmark n, James Lendemer o, H. Thorsten Lumbsch g, Michaela Schmull p, Conrad L. Schoch q, Emmanuël Sérusiaux r, David R. Maddison s, A. Elizabeth Arnold t, François Lutzoni a,10, Soili Stenroos c,10 a Department of Biology, Duke University, Durham, NC 27708-0338, USA b FB Biologie, Molecular Phylogenetics, 13/276, TU Kaiserslautern, Postfach 3049, 67653 Kaiserslautern, Germany c Botanical Museum, Finnish Museum of Natural History, FI-00014 University of Helsinki, Finland d Department of Ecology and Evolutionary Biology, Yale University, 358 ESC, 21 Sachem Street, New Haven, CT 06511, USA e Institut für Botanik, Karl-Franzens-Universität, Holteigasse 6, A-8010 Graz, Austria f Department of Plant Taxonomy and Nature Conservation, University of Gdan´sk, ul. Wita Stwosza 59, 80-308 Gdan´sk, Poland g Science and Education, The Field Museum, 1400 S. -
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. -
Lichens of Alaska's South Coast
United States Department of Agriculture Lichens of Alaska’s South Coast Forest Service R10-RG-190 Alaska Region Reprint April 2014 WHAT IS A LICHEN? Lichens are specialized fungi that “farm” algae as a food source. Unlike molds, mildews, and mushrooms that parasitize or scavenge food from other organisms, the fungus of a lichen cultivates tiny algae and / or blue-green bacteria (called cyanobacteria) within the fabric of interwoven fungal threads that form the body of the lichen (or thallus). The algae and cyanobacteria produce food for themselves and for the fungus by converting carbon dioxide and water into sugars using the sun’s energy (photosynthesis). Thus, a lichen is a combination of two or sometimes three organisms living together. Perhaps the most important contribution of the fungus is to provide a protective habitat for the algae or cyanobacteria. The green or blue-green photosynthetic layer is often visible between two white fungal layers if a piece of lichen thallus is torn off. Most lichen-forming fungi cannot exist without the photosynthetic partner because they have become dependent on them for survival. But in all cases, a fungus looks quite different in the lichenized form compared to its free-living form. HOW DO LICHENS REPRODUCE? Lichens sexually reproduce with fruiting bodies of various shapes and colors that can often look like miniature mushrooms. These are called apothecia (Fig. 1) and contain spores that germinate and Figure 1. Apothecia, fruiting grow into the fungus. Each bodies fungus must find the right photosynthetic partner in order to become a lichen. Lichens reproduce asexually in several ways. -
BLS Bulletin 111 Winter 2012.Pdf
1 BRITISH LICHEN SOCIETY OFFICERS AND CONTACTS 2012 PRESIDENT B.P. Hilton, Beauregard, 5 Alscott Gardens, Alverdiscott, Barnstaple, Devon EX31 3QJ; e-mail [email protected] VICE-PRESIDENT J. Simkin, 41 North Road, Ponteland, Newcastle upon Tyne NE20 9UN, email [email protected] SECRETARY C. Ellis, Royal Botanic Garden, 20A Inverleith Row, Edinburgh EH3 5LR; email [email protected] TREASURER J.F. Skinner, 28 Parkanaur Avenue, Southend-on-Sea, Essex SS1 3HY, email [email protected] ASSISTANT TREASURER AND MEMBERSHIP SECRETARY H. Döring, Mycology Section, Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AB, email [email protected] REGIONAL TREASURER (Americas) J.W. Hinds, 254 Forest Avenue, Orono, Maine 04473-3202, USA; email [email protected]. CHAIR OF THE DATA COMMITTEE D.J. Hill, Yew Tree Cottage, Yew Tree Lane, Compton Martin, Bristol BS40 6JS, email [email protected] MAPPING RECORDER AND ARCHIVIST M.R.D. Seaward, Department of Archaeological, Geographical & Environmental Sciences, University of Bradford, West Yorkshire BD7 1DP, email [email protected] DATA MANAGER J. Simkin, 41 North Road, Ponteland, Newcastle upon Tyne NE20 9UN, email [email protected] SENIOR EDITOR (LICHENOLOGIST) P.D. Crittenden, School of Life Science, The University, Nottingham NG7 2RD, email [email protected] BULLETIN EDITOR P.F. Cannon, CABI and Royal Botanic Gardens Kew; postal address Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AB, email [email protected] CHAIR OF CONSERVATION COMMITTEE & CONSERVATION OFFICER B.W. Edwards, DERC, Library Headquarters, Colliton Park, Dorchester, Dorset DT1 1XJ, email [email protected] CHAIR OF THE EDUCATION AND PROMOTION COMMITTEE: S. -
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.