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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. -
Lichenicolous Biota (Nos 201–230)
ZOBODAT - www.zobodat.at Zoologisch-Botanische Datenbank/Zoological-Botanical Database Digitale Literatur/Digital Literature Zeitschrift/Journal: Fritschiana Jahr/Year: 2015 Band/Volume: 80 Autor(en)/Author(s): Hafellner Josef Artikel/Article: Lichenicolous Biota (Nos 201-230) 21-41 - 21 - Lichenicolous Biota (Nos 201–230) Josef HAFELLNER* HAFELLNER Josef 2015: Lichenicolous Biota (Nos 201–230). – Frit- schiana (Graz) 80: 21–41. - ISSN 1024-0306. Abstract: The 9th fascicle (30 numbers) of the exsiccata 'Lichenicolous Biota' is published. The issue contains ma- terial of 20 non-lichenized fungal taxa (14 teleomorphs of ascomycetes, 4 anamorphic states of ascomycetes, 2 an- amorphic states of basidiomycetes) and 9 lichenized as- comycetes, including paratype material of Dimelaena li- chenicola K.Knudsen et al. (no 223), Miriquidica invadens Hafellner et al. (no 226, 227), and Stigmidium xantho- parmeliarum Hafellner (no 210). Furthermore, collections of the type species of the following genera are distributed: Illosporiopsis (I. christiansenii), Illosporium (I. carneum), Marchandiomyces (M. corallinus), Marchandiobasidium (M. aurantiacum, sub Erythricium aurantiacum), Micro- calicium (M. disseminatum), Nigropuncta (N. rugulosa), Paralecanographa (P. grumulosa), Phaeopyxis (P. punc- tum), Placocarpus (P. schaereri), Rhagadostoma (R. li- chenicola), and Stigmidium (S. schaereri). *Institut für Pflanzenwissenschaften, NAWI Graz, Karl-Franzens-Universität, Holteigasse 6, 8010 Graz, AUSTRIA e-mail: [email protected] Introduction The exsiccata 'Lichenicolous Biota' is continued with fascicle 9, containing 30 numbers. The exsiccata covers all lichenicolous biota, i.e., it is open not only to non- lichenized and lichenized fungi, but also to myxomycetes, bacteria, and even animals, whenever they cause a characteristic symptom on their host (e.g. discoloration or galls). -
A New Species of Aspicilia (Megasporaceae), with a New Lichenicolous Sagediopsis (Adelococcaceae), from the Falkland Islands
The Lichenologist (2021), 53, 307–315 doi:10.1017/S0024282921000244 Standard Paper A new species of Aspicilia (Megasporaceae), with a new lichenicolous Sagediopsis (Adelococcaceae), from the Falkland Islands Alan M. Fryday1 , Timothy B. Wheeler2 and Javier Etayo3 1Department of Plant Biology, Michigan State University, East Lansing, Michigan 48824, USA; 2Division of Biological Sciences, University of Montana, Missoula, Montana 59801, USA and 3Navarro Villoslada 16, 3° dcha., 31003 Pamplona, Navarra, Spain Abstract The new species Aspicilia malvinae is described from the Falkland Islands. It is the first species of Megasporaceae to be discovered on the islands and only the seventh to be reported from South America. It is distinguished from other species of Aspicilia by the unusual secondary metabolite chemistry (hypostictic acid) and molecular sequence data. The collections of the new species support two lichenicolous fungi: Endococcus propinquus s. lat., which is new to the Falkland Islands, and a new species of Sagediopsis with small perithecia and 3-septate ascospores c. 18–20 × 4–5 μm, which is described here as S. epimalvinae. A total of 60 new DNA sequences obtained from species of Megasporaceae (mostly Aspicilia) are also introduced. Key words: DNA sequences, Endococcus, Lecanora masafuerensis, lichen, southern South America, southern subpolar region (Accepted 18 March 2021) Introduction Materials and Methods Species of Megasporaceae Lumbsch et al. are surprisingly scarce Morphological methods in the Southern Hemisphere. Whereas 97 species are known Gross morphology was examined under a dissecting microscope from North America (Esslinger 2019), 104 from Russia and apothecial characteristics by light microscopy (compound (Urbanavichus 2010), 40 from Svalbard (Øvstedal et al. -
Die Gattung Aspicilia, Ihre Ableitungen Nebst
Acta Botánica Malacitana, 16(1): 133-140 Malaga, 1991 DIE GAUG ASPICILIA, IE AEIUGE ES EMEKUGE OE CYOECAOIE ASCOCAOGAISAIO EI AEE GENERA E LECANORALES (ASCOMYCEES ICEISAI Josef HAFELLNER SUMMARY: Some evolutionary lines are shown within Aspicilia coll, and the taxonomical and nomenclatural consequences for species commonly classified in Sphaerothallia Nees are discussed. The generic rank (Lobothallia (Clauzade & Roux) Hafellner) is proposed for the Aspicilia radiosa group and the following new combinations are introduced: Lobothallia alphoplaca (Wahlenb. in Ach.)Haf., Lobothallia melanaspis (Ach.)Haf., Lobothallia praeradiosa (Nyl.)Haf. and Lobothallia radiosa (Hoffm.)Haf. Key words: Lichenized Ascomycetes, Aspicilia, Lobothallia, taxonomy. ZUSAMMENFASSUNG: Innerhalb der Gattung Aspicilia coll. werden einige Evolutionslinien aufgezeigt und taxonomische wie nomenklatorische Konsequenzen für gewtihnlich als Sphaerothallia Nees bezeichnete Arten werden diskutiert. Lobothallia (Clauzade & Roux) Hafellner wird in den Gattungsrang erhoben und folgende neue Kombinationen werden vorgeschlagen: Lobythallia alphoplaca (Wahlenb. in Ach) Haf., Lobothallia melanaspis (Ach) Haf., Lobothallia praeradiosa (NYL.) Haf. und Lobothallia radiosa (Hoffm.)Haf. Schltisselwórter: Licheniscerte Ascomyceten, Aspiclia, Lobothallia, Taxonomie. EINLEITUNG Der schon von Massalongo (1852) beschriebenen Gattung Aspicilia war emn wechselvolles Schicksal beschieden. Ober lange Zeit in Lecanora eingeschlossen und oft in dieser als Subgenus bewertet (z.B. Magnusson 1939, Poelt 1958, Eigler 1969), hat sich erst in jtingerer Zeit die Erkenntnis allgemein durchgesetzt, daB Aspicilia mit Lecanora nicht ndher verwandt ist (Poelt 1974, Roux 1977, Hawksworth & al. 1980, Santesson 1984, Clauzade & Roux 1984, 1987, Hafellner 1984, Esnault 1985), obwohl einige Autoren schon früh die Selbstdndigkeit betont hatten (z.B. Kürber 1855, Hue 1910, Choisy 1929). Poelt (1974) hat sogar die neue Familie Aspiciliaceae vorgeschlagen, urn die taxonomische Distanz zwischen Aspicilia und Lecanora augenfdllig zu machen. -
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. -
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. -
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. -
The Fungi of Slapton Ley National Nature Reserve and Environs
THE FUNGI OF SLAPTON LEY NATIONAL NATURE RESERVE AND ENVIRONS APRIL 2019 Image © Visit South Devon ASCOMYCOTA Order Family Name Abrothallales Abrothallaceae Abrothallus microspermus CY (IMI 164972 p.p., 296950), DM (IMI 279667, 279668, 362458), N4 (IMI 251260), Wood (IMI 400386), on thalli of Parmelia caperata and P. perlata. Mainly as the anamorph <it Abrothallus parmeliarum C, CY (IMI 164972), DM (IMI 159809, 159865), F1 (IMI 159892), 2, G2, H, I1 (IMI 188770), J2, N4 (IMI 166730), SV, on thalli of Parmelia carporrhizans, P Abrothallus parmotrematis DM, on Parmelia perlata, 1990, D.L. Hawksworth (IMI 400397, as Vouauxiomyces sp.) Abrothallus suecicus DM (IMI 194098); on apothecia of Ramalina fustigiata with st. conid. Phoma ranalinae Nordin; rare. (L2) Abrothallus usneae (as A. parmeliarum p.p.; L2) Acarosporales Acarosporaceae Acarospora fuscata H, on siliceous slabs (L1); CH, 1996, T. Chester. Polysporina simplex CH, 1996, T. Chester. Sarcogyne regularis CH, 1996, T. Chester; N4, on concrete posts; very rare (L1). Trimmatothelopsis B (IMI 152818), on granite memorial (L1) [EXTINCT] smaragdula Acrospermales Acrospermaceae Acrospermum compressum DM (IMI 194111), I1, S (IMI 18286a), on dead Urtica stems (L2); CY, on Urtica dioica stem, 1995, JLT. Acrospermum graminum I1, on Phragmites debris, 1990, M. Marsden (K). Amphisphaeriales Amphisphaeriaceae Beltraniella pirozynskii D1 (IMI 362071a), on Quercus ilex. Ceratosporium fuscescens I1 (IMI 188771c); J1 (IMI 362085), on dead Ulex stems. (L2) Ceriophora palustris F2 (IMI 186857); on dead Carex puniculata leaves. (L2) Lepteutypa cupressi SV (IMI 184280); on dying Thuja leaves. (L2) Monographella cucumerina (IMI 362759), on Myriophyllum spicatum; DM (IMI 192452); isol. ex vole dung. (L2); (IMI 360147, 360148, 361543, 361544, 361546). -
Taxonomie, Phylogénie Et Écogéographie Des Peltigerales De La Réunion (Archipel Des Mascareignes)
Département des sciences et gestion de l’environnement Taxonomie végétale et biologie de la conservation Taxonomie, phylogénie et écogéographie des Peltigerales de la Réunion (archipel des Mascareignes) Mémoire de fin d’études en vue de l’obtention du grade de Master en biologie des organismes et écologie, à finalité approfondie Nicolas MAGAIN Promoteur : Emmanuël SÉRUSIAUX Août 2010 Sticta macrophylla Résumé -Une mission sur le terrain à la Réunion en novembre 2009 a permis de récolter environ 450 échantillons de Peltigerales. Un stage d’un mois a été effectué à l’Université du Connecticut dans le laboratoire du Professeur Bernard Goffinet. Taxonomie -Nous avons réalisé une checklist des Peltigerales de la Réunion présentant 69 espèces, dont 11 nouvelles pour la Réunion. Les genres Collema et Leptogium n’ont pas été étudiés. -Nous avons identifié, provenant de la Réunion, du Rift Albertin et de Madagascar, pas moins de 27 espèces de Sticta. Parmi ces 27 espèces, 14 d’entre elles représentent probablement des nouvelles espèces pour lesquelles nous n’avons trouvé aucun épithète validement publié. Phylogénie -Nous avons réalisé plusieurs études phylogénétiques sur l’ordre des Peltigerales par extraction d’ADN, amplification de trois loci (ITS nucléaire, LSU nucléaire et mtSSU mitochondrial), et analyses en maximum de parcimonie, maximum de vraisemblance et analyse bayesienne. -La famille des Pannariaceae au sens large est monophylétique, tout comme le genre Pannaria sensu stricto, à condition de revoir le nom de certaines espèces. Le genre Psoroma est polyphylétique, tout comme le genre Parmeliella. -Le genre Erioderma est monophylétique, et forme un groupe avec Leioderma et une partie des Degelia. -
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.