ARCTIC AND ALPINE MYCOLOGY II ENVIRONMENTAL SCIENCE RESEARCH Editorial Board Alexander Hollaender Council for Research Planning in Biological Sciences, Inc. Washington, D.C. Ronald F. Probstein Massachusetts Institute of Technology Cambridge, Massachusetts Bruce L. Welch Environmental Biomedicine Research, Inc. and The Johns Hopkins University School of Medicine Baltimore, Maryland Recent Volumes in this Series Volume 27 -SHORT-TERM BIOASSA YS IN THE ANALYSIS OF COMPLEX ENVIRONMENTAL MIXTURES III Edited by Michael D. Waters, Shah beg S. Sandhu, Joellen Lewtas, Larry Claxton, Neil Chernoff, and Stephen Nesnow Volume 28-UTILIZATION OF MAMMALIAN SPECIFIC LOCUS STUDIES IN HAZARD EVALUATION AND ESTIMATION OF GENETIC RISK Edited by Frederick J. de Serres and William Sheridan Volume 29-APPLICATION OF BIOLOGICAL MARKERS TO CARCINOGEN TESTING Edited by Harry A. Milman and Stewart Sell Volume 3D-INDIVIDUAL SUSCEPTIBILITY TO GENOTOXIC AGENTS IN THE HUMAN POPULATION Edited by Frederick J. de Serres and Ronald W. Pero Volume 31-MUTATION, CANCER, AND MALFORMATION Edited by Ernest H. Y. Chu and Walderico M. Generoso Volume 32-SHORT-TERM BIOASSAYS IN THE ANALYSIS OF COMPLEX ENVIRONMENTAL MIXTURES IV Edited by Michael D. Waters, Shahbeg S. Sandhu, Joellen Lewtas, Larry Claxton, Gary Strauss, and Stephen Nesnow Volume 33-ACID RAIN: Economic Assessment Edited by Paulette Mandelbaum Volume 34-ARCTIC AND ALPINE MYCOLOGY II Edited by Gary A. Laursen, Joseph F. Ammirati, and Scott A. Redhead A Continuation Order Plan is available for this series. A continuation order will bring delivery of each new volume immediately upon publication. Volumes are billed only upon actual shipment. For further information please contact the publisher. ARCTIC AND ALPINE MYCOLOGY II Edited by Gary A. Laursen School of Agricultllre and Land Hesollrces Management University of Alaslw Fairbanks, Alaska Joseph F. Ammirati Department of Botany University of Washington Seattle, Washington and Scott A. Redhead Biosystenwtics Research Centre Ottawa, Ontario, Canada Springer Science+Business Media, LLC Library of Congress Cataloging in Publication Data International Symposium on Arcto-Alpine Mycology (2nd: 1984: Fetan, Switzerland) Arctic and alpine mycology II. (Environmental science research; v. 34) "Proceedings of the Second International Symposium on Arctic and Alpine Mycology, held August 26-September 2, 1984, in Fetan, Switzerland" - T.p. verso. Bibliography: p. Includes index. 1. Fungi - Arctic regions - Congresses. 2. Mycology - Congresses. 3. Alpine flora Congresses. 1. Laursen, Gary A. II. Ammirati, )oseph F. III. Redhead, Scott A. IV. Ti tie. V. Title: Arctic and alpine mycology. 2. Vi. Series. QK615.157 1984 589.2'0998 87-7814 ISBN 978-1-4757-1941-3 ISBN 978-1-4757-1939-0 (eBook) DOI 10.1007/978-1-4757-1939-0 Proceedings of the Second International Symposium on Arctic ancl Alpine Mycology, held August 26-September 2, 1984, in Fetan, Switzerland © 1987 Springer Science+Business Media New York Originally published by Plenum Press, New York in 1987. AII rights reserved No part of this book may be reproduced, stored in a retricval systcm, or transmittcd in any form or by any means, electronic, mechanical, photocopying, microfilming, recording, or otherwise, without written permission horn the Publisher PREFACE During the summer of 1980, the First International symposium on Arctic and Alpine Mycology (ISAM-I) was held at the then extant Naval Arctic Research Laboratory near Barrow, Alaska, U.S.A., well within the Arctic Circle (Laursen and Ammirati, Arctic and Alpine Mycology. The First International symposium on Arcto-Alpine Mycology. Univ. Wash. Press, 1982). The facility is currently owned and operated by the Utkeagvik Inupiat community and is named the National Academic and Research Laboratory, thus retaining its acronym NARL. Twenty-five scientists participated in that historic first meeting. Their interests in the fungi spanned a vast geographic area of cold dominated habitats in both the northern and southern hemispheres that included four continents (N. and S. America, Eurasia,and Antarctica), nine countries, and numerous islands ranging from Greenland to Jan Mayen in the Svalbard group. ISAM-I helped to develop ongoing interests and initiate others. This is what ISAM-I founders hoped would happen. Asa result, the organizing committee for ISAM-II was formed. Its mandate was to: involve a maximum of one third new participants in future ISAM meetings: divide the responsibility for organizing future meetings at sites located in areas of interest to research thrusts in Arctic and alpine environments: keep the number of participants small enough to ensure manageability, taking full advantage of field collecting opportunities with minimal complications and cost. ISAM-II was held in eastern Switzerland in the Unter-Engadin at the Hochalpines Tochter-Institut at Fetan/Ftan near the swiss National Park, from August 26 to September 2, 1984. The National Park was a favorite collecting area of a swiss pioneer in alpine mycology, Jules Favre. Twenty-eight participants, all pictured in Figure 1, represented twelve countries as follows: Austria (1); Canada (1); Denmark (3); Finland (2); France (2); Germany (1); Japan (1); Norway (3); Scotland (2); Sweden (2); Switzerland (7); and the U. S. A. (3). They visited a variety of alpine sites, including type localities, in a carefully planned and executed itinerary of field trips organized by Prof. E. Muller and Dr. E. Horak and their students. The group was fortunate enough to have good weather for the duration of the symposium, as the following week the entire area became snowed under with an early storm. Evening sessions were devoted to the presentation of papers and examination of specimens which often led to animated discussions between seasoned older researchers and younger scientists, between North Americans and Europeans, and between taxonomists and ecologists. There was an important balance between senior, middle-level, younger and graduate-level scientists, insuring continued growth, an influx of new research ideas, and an infusion of individuals researching mycological problems in Arctic and alpine habitats. v Growing popularity in Artic and alpine mycology requires that organizers of future ISAM meetings restrict the number in forays to about 25-30 individuals. This is mainly due to the difficulties involved in supporting and transporting large groups to remote sites. Holding formal symposia in conjunction with other international meetings in more central metropolitan areas may be necessary to facilitate the interaction of a larger group. This could be coupled with a 7-10 day foray to a'remote site with a smaller group of participants. This problem needs resolving if groups of more than 30 participants are going to meet at future symposium gatherings. The establishment of Arctic and Alpine Mycology Mycology as a branch of study was addressed at ISAM-II. A preamble for "Arranging the ISAM" was drafted by M. Lange (Copenhagen) and adopted at the ISAM-II business meeting. It sets guidelines for organizing future ISAM gatherings in a meaningful 1 way and allows for the development of all aspects of Arctic and alpine mycology. The preamble is as follows: 1. Invitations for the next assembly are considered and determined at each ISAM. The session should accept one invitation and indicate a second priority. 2. A representative of the inviting nation is selected as President for the next ISAM period. He/she has the responsibility for making all arrangements. 3. The ISAM is open only by invitation. Persons invited shall constitute a qualified group with broad expertise in arcto-alpine mycology. No past ISAM members may claim the privilage of an invitation. 4. It shall be the right of members of the last ISAM to suggest participants to the next assembly. 5. The President will draw up a list of potential participants from nations participating in arcto-alpine research in mycology, keeping in mind geographical distribution. He/she shall consult with previous presidents on matters of principle or of major importance. 6. Definite invitations shall be issued not later than nine months before the ISAM assembly. 7. If the President must cancel the arrangement, then he/she shall inform the two previous Presidents before defintive action is taken. He/she shall inform a representative of the nation that extended the second priority. The country in question shall then appoint a President who shall assume the above responsibilities. 8. ISAM members shall submit a relevant paper to be printed in a special ISAM publication in accordance with decisions and arrangements made by the President. To facilitate this, the President may appoint an editorial committee. The organizational structure and membership, albeit small, exists. Previously unresolved and long range objectives may be developed for adoption. These and other professional activities have resulted in our making positive strides toward meeting and correcting criticisms of the ISAM-I volumne. vi A third ISAM is scheduled to take place in 1988 at Svalbard (Norway). The site for the meeting is positioned above 780 N latitude. The following committees have been established: Executive Committee: Sigmund Sivertsen (Norway-Trondheim) President Egon Horak (Switzerland-Zurich) Past President Gary Laursen (U. S. A.-Alaska) Secretary organizing committee (Norway): Sigmund Sivertsen, Chairperson Gro Gulden Trond Schumacher ACKNOWLEDGEMENTS No meeting of this magnitude is ever conducted without support given from several sources. From Vice Chancellor for Research and Advanced Study at the University of Alaska came a travel grant: from the Mikrobiologisches Institut and the Geobotanisches Institut, Eidegenossische Technische Hochschule, ZUrich, came all logistical support and arrangements, and from President H. Ursprung's office came financial support; the Swiss Army provided vehicles for local foray transportation; the Hochalpines Tochter-Institut and its summer staff provided gracious living accommodations; the Swiss National Park provided collecting permits for several forays into various sections of the park; and Mr. G. A. Mulligan, Director of the Biosystematics Research Centre, Ottawa, provided the resources necessary to prepare and index all manuscripts in a camera-ready format. G. A. Laursen J. F. Ammirati S. A. Redhead vii CONTENTS Introduct ion •.••....•........•....•••...•••••••••..••••••••••••••••• 1 BIOGEOGRAPHY On the Origin and Ecology of Alpine Plant Parasitic Fungi ..•...•..•. 3 E. Muller and J. A. Magnuson Larger Arctic-Alpine Fungi in Scotland 17 R. Watling ECOLOGY/PHYSIOLOGY Ecophysiological Studies on Alpine Macromycetes: Saprophytic Clitocybe and Mycorrhizal Hebeloma associated with Dryas octopetala 47 J. C. Debaud sociology and Ecology of Larger Fungi in the Subarctic and Oroarctic Zones in Northwest Finnish Lapland .....•.....•••••••• 61 K. Metsanheimo Endophytic Fungi of Alpine Ericaceae. The Endophytes of Loiseleuria procumbens .........••..•..•.....••.•••••••••....••. 71 o. Petrini TAXONOMY: ASCOMYCETES Three 4-spored Saccobolus species from North East Greenland ••.....•. 79 H. Dissing Ascomycetes Growing on Polytrichum sexangulare ...•.••••••••••.....•. 87 P. Dobbeler Nordic Juncicolous Mycosphaerellae 109 L. Holm and K. Holm New Svalbard Fungi 123 S. Huhtinen ix Phaeosphaeria in the Arctic and Alpine Zones ........................ 153 A. Leuchtmann Sarcoleotia globosa (Sommerf.: Fr.) Korf. Taxonomy Ecology and Distribution....................................... 163 T. Schumacher and S. Sivertsen (see also Muller and Magnuson. Petrini. Watling) TAXONOMY: BASIDIOMYCETES The Genus Galerina on Svalbard 177 G. Gulden Astrosporina in the Alpine Zone of the Swiss National Park (SNP) and Adjacent Regions ................................ 205 E. Horak Agaricaceae. Amanitaceae. Boletaceae. Gomphidiaceae. Paxillaceae and Pluteaceae in Greenland ........................ 235 H. Knudsen and T. Borgen Agaricales de la Zone Alpine. Genus Cortinarius Fr .• Subgenus Telamonia (Fr.) Loud. Part III ....................... 255 D. Lamoure Arctic Gasteromycetes. The Genus Bovista in Greenland and Svalbard ......................................... 261 M. Lange Hygrophoraceae from Arctic and Alpine Tundra in Alaska .............. 273 G. A. Laursen. J. F. Ammirati and D. F. Farr Higher Fungi in Tundra and Subalpine Tundra from the Yukon Territory and Alaska ..................................... 287 o. K. Miller. Jr. Fungi (Agaricales. Russulales) from the Alpine Zone of Yellowstone National Park and the Beartooth Mountains with Special Emphasis on Cortinarius ........................... 299 M. Moser and K. H. McKnight Lichenized Agarics: Taxonomic and Nomenclatural Riddles ............. 319 S. A. Redhead and Th. W. Kuyper (see also Debaud. Huhtinen. Metsanheimo. Watling) Contributors 349 Part icipants Photograph ............................................. 351 Index ............................................................... 353 x INTRODUCTION Research topics from the following papers which constitute the second International Symposium on Arctic and Alpine Mycology (ISAM) are largely biased in favour of taxonomy, perhaps because of the pioneering work being conducted in the field of Arctic and alpine mycology. The present volumne contains a collection of 20 papers dealing with fungi from Arctic, Arctic-like, or alpine habitats (c.f. Watling's paper); eight of which are devoted wholly or inpart to the Ascomycotina and eleven that are dedicated to investigations mainly on the Basidiomycotina. Abstracts for most these works were published separately (Laursen and Ammirati, Univ. Alaska, Agric. Exp. Stn., Misc. Pub. 84-2, 1984). Leading papers of these proceedings are two biogeographically oriented papers which give overviews on the historically linked origins of alpine fungi, illucidated by examples of parasitic ascomycetes from the Alps (Muller and Magnuson), and contemporary climatic factors influencing the distribution of Arctic-alpine fungi, with relatively low altitude areas of Scotland serving as an unusual example (watling). These are followed by three papers detailing specific factors related to the ecological niches and fruiting periods of mycorrhizal and saprophytic agarics in alpine Dryas mounds, subarctic birch and spruce forests, and heaths, and endophytic ascomycetes in Lo1seleur1a (Debaud, Metsanheimo, Petrini). Finally there are two sections largely devoted to taxonomy of first the Ascomycetes, and secondly the Basidiomycetes. Two of these address problems in the large fleshy terrestrial or coprophilous discomycetes in the genera Saccobolus and Sarcoleot1a (Schumacher and Sivertsen), another documents microascomycetes specifically parasitizing a characteristic Arctic-alpine moss, Polytr1chum sexangulare (Dobbeler), and a fourth summarizes data on Nordic juncicolous species in the parasitic/saprophytic genus Mycosphaerella (Holm and Holm). A key to the Arctic and alpine Phaeosphaer1a is given in another (Leuchtmann). Finally, this section leads into the Basidiomycete section with a paper covering a variety of Ascomycetes and Basidiomycetes from Svalbard (Huhtinen). In the Basidiomycete section one contribution is an in depth examination of the historical facts concerning the study of the omphaloid, mainly Arctic and alpine basidiolichens, and the repercussions of these studies with novel interpretations (Redhead and Kuyper). Two are studies on mycorrhizal species in Astrospor1na/Inocybe; one historically linked to the studies and collecting sites of J. Favre (Horak), the other an exploration of new territory (Miller). Two contributions concentrate on the genus Cortinar1us - the subgenus Telamonia from European alpine sites (Lamoure), and notable taxa from alpine sites in western North
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