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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. -
The Lichens' Microbiota, Still a Mystery?
fmicb-12-623839 March 24, 2021 Time: 15:25 # 1 REVIEW published: 30 March 2021 doi: 10.3389/fmicb.2021.623839 The Lichens’ Microbiota, Still a Mystery? Maria Grimm1*, Martin Grube2, Ulf Schiefelbein3, Daniela Zühlke1, Jörg Bernhardt1 and Katharina Riedel1 1 Institute of Microbiology, University Greifswald, Greifswald, Germany, 2 Institute of Plant Sciences, Karl-Franzens-University Graz, Graz, Austria, 3 Botanical Garden, University of Rostock, Rostock, Germany Lichens represent self-supporting symbioses, which occur in a wide range of terrestrial habitats and which contribute significantly to mineral cycling and energy flow at a global scale. Lichens usually grow much slower than higher plants. Nevertheless, lichens can contribute substantially to biomass production. This review focuses on the lichen symbiosis in general and especially on the model species Lobaria pulmonaria L. Hoffm., which is a large foliose lichen that occurs worldwide on tree trunks in undisturbed forests with long ecological continuity. In comparison to many other lichens, L. pulmonaria is less tolerant to desiccation and highly sensitive to air pollution. The name- giving mycobiont (belonging to the Ascomycota), provides a protective layer covering a layer of the green-algal photobiont (Dictyochloropsis reticulata) and interspersed cyanobacterial cell clusters (Nostoc spec.). Recently performed metaproteome analyses Edited by: confirm the partition of functions in lichen partnerships. The ample functional diversity Nathalie Connil, Université de Rouen, France of the mycobiont contrasts the predominant function of the photobiont in production Reviewed by: (and secretion) of energy-rich carbohydrates, and the cyanobiont’s contribution by Dirk Benndorf, nitrogen fixation. In addition, high throughput and state-of-the-art metagenomics and Otto von Guericke University community fingerprinting, metatranscriptomics, and MS-based metaproteomics identify Magdeburg, Germany Guilherme Lanzi Sassaki, the bacterial community present on L. -
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. -
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. -
<I>Cyanodermella Asteris</I> Sp. Nov. (<I>Ostropales</I>)
MYCOTAXON ISSN (print) 0093-4666 (online) 2154-8889 Mycotaxon, Ltd. ©2017 January–March 2017—Volume 132, pp. 107–123 http://dx.doi.org/10.5248/132.107 Cyanodermella asteris sp. nov. (Ostropales) from the inflorescence axis of Aster tataricus Linda Jahn1,*, Thomas Schafhauser2, Stefan Pan2, Tilmann Weber2,7, Wolfgang Wohlleben2, David Fewer3, Kaarina Sivonen3, Liane Flor4, Karl-Heinz van Pée4, Thibault Caradec5, Philippe Jacques5,8, Mieke M.E. Huijbers6,9, Willem J.H. van Berkel6 & Jutta Ludwig-Müller1,* 1 Institut für Botanik, Technische Universität Dresden, 01062 Dresden, Germany 2 Mikrobiologie und Biotechnologie, Interfakultäres Institut für Mikrobiologie und Infektionsmedizin, Eberhard Karls Universität Tübingen, Auf der Morgenstelle 28, 72076 Tübingen, Germany 3 Microbiology and Biotechnology Division, Dept. of Food and Environmental Sciences, University of Helsinki, Viikinkaari 9, FIN-00014, Helsinki, Finland 4 Allgemeine Biochemie, Technische Universität Dresden, 01069 Dresden, Germany 5 Laboratoire ProBioGEM, Université Lille1- Sciences et Technologies, Villeneuve d’Ascq, France 6 Laboratory of Biochemistry, Wageningen University, Dreijenlaan 3, 6703 HA Wageningen, The Netherlands 7 moved to: Novo Nordisk Foundation Center for Biosustainability, Technical University of Denmark, Kemitorvet Bygning 220, 2800 Kgs. Lyngby, Denmark 8 moved to: Gembloux Agro-Bio Tech, Université de Liege, Passage des Déportés 2, 5030 Gembloux, Belgium 9 moved to: Department of Biotechnology, Technical University Delft, Van der Maasweg 9, 2629 HZ Delft, The Netherlands *Correspondence to: [email protected], [email protected] Abstract—An endophytic fungus isolated from the inflorescence axis ofAster tataricus is proposed as a new species. Phylogenetic analyses based on sequences from the ribosomal DNA cluster (the ITS1+5.8S+ITS2, 18S, and 28S regions) and the RPB2 gene revealed a relationship between the unknown fungus and the Stictidaceae lineage of the Ostropales. -
Studies of the Laboulbeniomycetes: Diversity, Evolution, and Patterns of Speciation
Studies of the Laboulbeniomycetes: Diversity, Evolution, and Patterns of Speciation The Harvard community has made this article openly available. Please share how this access benefits you. Your story matters Citable link http://nrs.harvard.edu/urn-3:HUL.InstRepos:40049989 Terms of Use This article was downloaded from Harvard University’s DASH repository, and is made available under the terms and conditions applicable to Other Posted Material, as set forth at http:// nrs.harvard.edu/urn-3:HUL.InstRepos:dash.current.terms-of- use#LAA ! STUDIES OF THE LABOULBENIOMYCETES: DIVERSITY, EVOLUTION, AND PATTERNS OF SPECIATION A dissertation presented by DANNY HAELEWATERS to THE DEPARTMENT OF ORGANISMIC AND EVOLUTIONARY BIOLOGY in partial fulfillment of the requirements for the degree of Doctor of Philosophy in the subject of Biology HARVARD UNIVERSITY Cambridge, Massachusetts April 2018 ! ! © 2018 – Danny Haelewaters All rights reserved. ! ! Dissertation Advisor: Professor Donald H. Pfister Danny Haelewaters STUDIES OF THE LABOULBENIOMYCETES: DIVERSITY, EVOLUTION, AND PATTERNS OF SPECIATION ABSTRACT CHAPTER 1: Laboulbeniales is one of the most morphologically and ecologically distinct orders of Ascomycota. These microscopic fungi are characterized by an ectoparasitic lifestyle on arthropods, determinate growth, lack of asexual state, high species richness and intractability to culture. DNA extraction and PCR amplification have proven difficult for multiple reasons. DNA isolation techniques and commercially available kits are tested enabling efficient and rapid genetic analysis of Laboulbeniales fungi. Success rates for the different techniques on different taxa are presented and discussed in the light of difficulties with micromanipulation, preservation techniques and negative results. CHAPTER 2: The class Laboulbeniomycetes comprises biotrophic parasites associated with arthropods and fungi. -
Umbilicariaceae Phylogeny TAXON 66 (6) • December 2017: 1282–1303
Davydov & al. • Umbilicariaceae phylogeny TAXON 66 (6) • December 2017: 1282–1303 Umbilicariaceae (lichenized Ascomycota) – Trait evolution and a new generic concept Evgeny A. Davydov,1 Derek Peršoh2 & Gerhard Rambold3 1 Altai State University, Lenin Ave. 61, Barnaul, 656049 Russia 2 Ruhr-Universität Bochum, AG Geobotanik, Gebäude ND 03/170, Universitätsstraße 150, 44801 Bochum, Germany 3 University of Bayreuth, Plant Systematics, Mycology Dept., Universitätsstraße 30, NW I, 95445 Bayreuth, Germany Author for correspondence: Evgeny A. Davydov, [email protected] ORCID EAD, http://orcid.org/0000-0002-2316-8506; DP, http://orcid.org/0000-0001-5561-0189 DOI https://doi.org/10.12705/666.2 Abstract To reconstruct hypotheses on the evolution of Umbilicariaceae, 644 sequences from three independent DNA regions were used, 433 of which were newly produced. The study includes a representative fraction (presumably about 80%) of the known species diversity of the Umbilicariaceae s.str. and is based on the phylograms obtained using maximum likelihood and a Bayesian phylogenetic inference framework. The analyses resulted in the recognition of eight well-supported clades, delimited by a combination of morphological and chemical features. None of the previous classifications within Umbilicariaceae s.str. were supported by the phylogenetic analyses. The distribution of the diagnostic morphological and chemical traits against the molecular phylogenetic topology revealed the following patterns of evolution: (1) Rhizinomorphs were gained at least four times independently and are lacking in most clades grouping in the proximity of Lasallia. (2) Asexual reproductive structures, i.e., thalloconidia and lichenized dispersal units, appear more or less mutually exclusive, being restricted to different clades. -
Distribution of Lichens on Few Ancient Monuments of Sonitpur District, Assam, North East India
Vol. 8(11), pp. 291-296, November 2016 DOI: 10.5897/IJBC2016.0971 Article Number: 1912C0161158 International Journal of Biodiversity ISSN 2141-243X Copyright © 2016 and Conservation Author(s) retain the copyright of this article http://www.academicjournals.org/IJBC Full Length Research Paper Distribution of Lichens on few ancient monuments of Sonitpur district, Assam, North East India Manas Pratim Choudhury1*, Mandira Sarma2, Sanjeeva Nayaka3 and Dilip Kumar Upreti3 1Department of Botany, Gauhati University, Guwahati-781001, Assam, India. 2Department of Botany, Cotton College, Guwahati-781001, Assam, India. 3Lichenology Laboratory, Plant Biodiversity and Conservation Biology Division, National Botanical Research Institute, CSIR, Lucknow 226001, U.P., India. Received 21 March, 2016; Accepted 15 September, 2016 This preliminary investigation describes the diversity and distribution of lichens from the different archaeological monuments of Sonitpur district, located in the Eastern Himalayan region of India, which is a part of Indo-Burma biodiversity hotspot. It is a land of natural beauty and rich cultural heritage and it is well reflected in the number of archaeological sites and monuments of ancient time at this region. Enumerations of 38 species of lichen belonging to 21 genera and 15 families have been reported from the archaeological monuments of Sonitpur district. 6 species are new records for this state. Members of Physciaceae showed higher dominance with 11 species followed by Teloschistaceae with 4 species and Lecanoraceae and Verrucariaceae each containing 3 species, respectively. Cryptothecia subnidulans is the most widespread species, reported from 7 study sites. Key words: Diversity, Eastern Himalayan, heritage, hot spot, species. INTRODUCTION Lichens are a stable self-supporting association of a can withstand xeric conditions which other plants find mycobiont and a phycobiont in which the mycobiont is the unfavourable for their growth and survival. -
An Evolving Phylogenetically Based Taxonomy of Lichens and Allied Fungi
Opuscula Philolichenum, 11: 4-10. 2012. *pdf available online 3January2012 via (http://sweetgum.nybg.org/philolichenum/) An evolving phylogenetically based taxonomy of lichens and allied fungi 1 BRENDAN P. HODKINSON ABSTRACT. – A taxonomic scheme for lichens and allied fungi that synthesizes scientific knowledge from a variety of sources is presented. The system put forth here is intended both (1) to provide a skeletal outline of the lichens and allied fungi that can be used as a provisional filing and databasing scheme by lichen herbarium/data managers and (2) to announce the online presence of an official taxonomy that will define the scope of the newly formed International Committee for the Nomenclature of Lichens and Allied Fungi (ICNLAF). The online version of the taxonomy presented here will continue to evolve along with our understanding of the organisms. Additionally, the subfamily Fissurinoideae Rivas Plata, Lücking and Lumbsch is elevated to the rank of family as Fissurinaceae. KEYWORDS. – higher-level taxonomy, lichen-forming fungi, lichenized fungi, phylogeny INTRODUCTION Traditionally, lichen herbaria have been arranged alphabetically, a scheme that stands in stark contrast to the phylogenetic scheme used by nearly all vascular plant herbaria. The justification typically given for this practice is that lichen taxonomy is too unstable to establish a reasonable system of classification. However, recent leaps forward in our understanding of the higher-level classification of fungi, driven primarily by the NSF-funded Assembling the Fungal Tree of Life (AFToL) project (Lutzoni et al. 2004), have caused the taxonomy of lichen-forming and allied fungi to increase significantly in stability. This is especially true within the class Lecanoromycetes, the main group of lichen-forming fungi (Miadlikowska et al. -
Preliminary Classification of Leotiomycetes
Mycosphere 10(1): 310–489 (2019) www.mycosphere.org ISSN 2077 7019 Article Doi 10.5943/mycosphere/10/1/7 Preliminary classification of Leotiomycetes Ekanayaka AH1,2, Hyde KD1,2, Gentekaki E2,3, McKenzie EHC4, Zhao Q1,*, Bulgakov TS5, Camporesi E6,7 1Key Laboratory for Plant Diversity and Biogeography of East Asia, Kunming Institute of Botany, Chinese Academy of Sciences, Kunming 650201, Yunnan, China 2Center of Excellence in Fungal Research, Mae Fah Luang University, Chiang Rai, 57100, Thailand 3School of Science, Mae Fah Luang University, Chiang Rai, 57100, Thailand 4Landcare Research Manaaki Whenua, Private Bag 92170, Auckland, New Zealand 5Russian Research Institute of Floriculture and Subtropical Crops, 2/28 Yana Fabritsiusa Street, Sochi 354002, Krasnodar region, Russia 6A.M.B. Gruppo Micologico Forlivese “Antonio Cicognani”, Via Roma 18, Forlì, Italy. 7A.M.B. Circolo Micologico “Giovanni Carini”, C.P. 314 Brescia, Italy. Ekanayaka AH, Hyde KD, Gentekaki E, McKenzie EHC, Zhao Q, Bulgakov TS, Camporesi E 2019 – Preliminary classification of Leotiomycetes. Mycosphere 10(1), 310–489, Doi 10.5943/mycosphere/10/1/7 Abstract Leotiomycetes is regarded as the inoperculate class of discomycetes within the phylum Ascomycota. Taxa are mainly characterized by asci with a simple pore blueing in Melzer’s reagent, although some taxa have lost this character. The monophyly of this class has been verified in several recent molecular studies. However, circumscription of the orders, families and generic level delimitation are still unsettled. This paper provides a modified backbone tree for the class Leotiomycetes based on phylogenetic analysis of combined ITS, LSU, SSU, TEF, and RPB2 loci. In the phylogenetic analysis, Leotiomycetes separates into 19 clades, which can be recognized as orders and order-level clades. -
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.