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Developments in Plant Genetics and Breeding 1A ISOZYMES IN PLANT GENETICS AND BREEDING, PART A edited by S. D. Tanksley and T. J. Orton 1983x + 516 pp. 1B ISOZYMES IN PLANT GENETICS AND BREEDING, PART Β edited by S. D. Tanksley and T. J. Orton 1983viii + 472 pp. 2A CHROMOSOME ENGINEERING IN PLANTS: GENETICS, BREEDING, EVOLUTION, PART A edited by P. K. Gupta and T. Tsuchiya 1991 656 pp. 2B CHROMOSOME ENGINEERING IN PLANTS: GENETICS, BREEDING, EVOLUTION, PART Β edited by T. Tsuchiya and P. K. Gupta 1991 in preparation 3 GENETICS OF SCOTS PINE edited by M. Giertych and Cs. Mâtyés 1991 280 pp. Developments in Plant Genetics and Breeding, 3 Genetics of Scots Pine Edited by Maciej Giertych Institute of Dendrology, Polish Academy of Sciences, Kornik, Poland Co-editor Csaba Maty as Department of Plant Sciences, Sopron University, Sopron, Hungary ELSEVIER Amsterdam-Oxford-New York-Tokyo 1991 Joint edition published by Elsevier Science Publishers, Amsterdam, The Netherlands and Akadémiai Kiado, Budapest, Hungary Distribution of this book is being handled by the following publishers: for the U.S.A. and Canada Elsevier Science Publishing Company, Inc. 655 Avenue of the Americas New York, N.Y. 10010 for the East European countries. Democratic People's Republic of Korea, People's Republic of Mongolia, Republic of Cuba, Socialist Republic of Vietnam Kultura Foreign Trading Company, P. 0. Box 149, 1389 Budapest, Hungary for all remaining areas Elsevier Science Publishers Sara Burgerhartstraat 25 P.O. Box 211, 1000 AE Amsterdam, The Netherlands Library of Congress Cataloging-in-Publication Data Giertych, Maciej. Genetics of Scots pine / editor, Maciej Giertych : co-editor, Csaba Mâtyâs. p. cm. — (Developments in plant genetics and breeding; 3) Includes bibliographical references and index. ISBN 0-444-98724-X 1. Scots pine—Genetics. 2. Scots pine—Breeding. I. Mâtyâs, Csaba. II. Title. III. Series. SD397.P614G54 1991 634.9751 —dc20 91 -6309 CIP ISBN 0-444-98724-X © M. Giertych and Cs. Mâtyâs 1991 All rights reserved. No part of this publication may be reproduced, stored in a retrieval system or transmitted in any form or by any means, mechanical, photocopying, recording, or otherwise without the prior written permission of the copyright owner. Printed in Hungary by Akadémiai Kiado és Nyomda Vâllalat, Budapest. Preface In April 1986 Dr. Csaba Mâtyâs from Hungary, chairman of the IUFRO Working Party S2.03.05 Breeding Scots pine, wrote, asking me to act as editor in an international project aimed at compiling literature on the subject of Scots pine genetics. Since the species is the most important one in Eurasia it was considered necessary to involve authors from as many countries as possible, including Eastern Europe and the USSR where the species dominates all forestry endeavour. Since Scots pine represents the highest proportion of forest cover in Poland (72 per cent), it seemed natural that there should be an editor from Poland. I agreed tentatively and started to discuss the format with prospective authors. During the 18th IUFRO World Congress in Ljubljana, Yugoslavia, in September 1986, at a business meeting of the Working Party, we discussed the details of the project and agreed on the contents of the book. With minor changes the list of chapters of this book is as originally planned. My nomination as editor of the volume was confirmed. It was agreed that publication be arranged through Akadémiai Kiado, the Publishing House of the Hungarian Academy of Sciences. In a letter dated 5th March 1987 from the publisher I was formally asked to organise and edit the monograph on 'Genetics of Scots pine'. I accepted. With help of other members of the Working Party an intensive search for willing and competent authors was started through correspondence. On 4th May 1987 I sent out a circular letter asking prospective authors to write different chapters and also asking them to help with the editing. It was my belief that the best reviewers were to be found among the authors of this monograph, all specialists in Scots pine genetics. Thus I have asked each author to act also as a reviewer for a chapter of the volume. On 27th July 1987 I was able to announce that 19 authors had agreed to my terms and that I expected the agreement of the remain- ing few shortly, which indeed happened. With minor modifications the list of authors and chapter titles was fixed in mid-1987. The writing and reviewing proved longer than expected, so that the manuscript was ready for presentation to the publisher by the end of July 1989. This volume is a truly international effort, prepared in the traditional IUFRO spirit of selfless co-operation. In all, 24 authors from 9 countries are involved. Each chapter was reviewed by two reviewers from two different countries, by myself as the editor of the whole volume, and by Cs. Mâtyâs as associate editor 5 commissioned by the publisher, who was responsible for the final, co-ordinative editing work and the linguistic revision of the chapters. Needless to say a lot of international correspondence was involved and the process proved time-consum- ing. I hope it was worth. I leave this to our readers to judge. I wish to express my sincere gratitude to all authors for their co-operation, for their willingness to submit to the editorial pressure and for their help with the reviewing. Maciej Giertych 6 Contributors Barnéoud, C, AFOCEL, Direction des Recherches, 2 Place Ferber, 69009 Lyon, France Boratynski, Α., Institute of Dendrology, 62-035 Kornik, Poland Borzan, Z., Forestry Faculty, Department of Forest Genetics and Dendrology, P.O.Box 178, 41001 Zagreb, Yugoslavia Chaiupka, W., Institute of Dendrology, 62-035 Kornik, Poland Ericson, B., SLU Department of Forest Yield Research, S-770 73 Garpenberg, Sweden Gerhold, H. D., School of Forest Resources, The Pennsylvania State University, University Park, PA 16802, USA Giertych, M., Institute of Dendrology, 62-035 Kornik, Poland Hattemer, Η. H., Abteilung Forstliche Biometrie, Büsgenweg 5, 3400 Göttingen, FRG van Haverbeke, D. F., USDA Department of Agriculture, Rocky Mountain Forest and Range Experimental Station, University of Nebraska, Lincoln, NE 68583, USA Kosinski, G., Institute of Dendrology, 62-035 Kornik, Poland Koski, V., Department of Forest Genetics, The Finnish Forest Research Institute, P.O. Box 18, SF-01301, Vantaa 30, Finland Lindgren, D., SLU Department of Forest Genetics, S-901 83 Urnea, Sweden Mâtyâs, Cs., Forestry Faculty, Department of Plant Sciences, P.O.Box 132, 9401 Sopron, Hungary Mikola, J., The Foundation for Forest Tree Breeding, Viljatie 4 A 5, SF-00700 Helsinki, Finland 17 Molotkov, P. I., Ukrainian Forest and Melioration Research Institute, Pushkinskaya 86, 310024 Kharkov 24, USSR Monteuuis, 0., AFOCEL, Domaine de l'Etancon, F-77370 Nangis, France Müller-Starck, G., Abteilung Forstgenetik und Forstpflanzenzüchtung, Büsgenweg 2, 3400 Göttingen, FRG Nilsson, J.-E., SLU Department of Forest Genetics, S-90183 Umeä, Sweden Oleksyn, J., Institute of Dendrology, 62-035 Kornik, Poland Patlaj, I. N., Ukrainian Forest and Melioration Research Institute, Pushkinskaya 86, 310024 Kharkov 24, USSR Pihelgas, E., Eesti Pollumajanduse Akadeemia, Tartu, Estonia, USSR Prus-Gtowacki, W., UAM Department of Genetics, Ul. Dabrowskiego 165, 60-594 Poznan, Poland Stâhl, Ε. G., SLU Department of Forest Yield Research, S-7770 73 Garpenberg, Sweden Stephan, Β. R., Institut für Forstgenetik und Forstpflanzenzüchtung, Sieker Landstrasse 2, 2070 Grosshansdorf 2, FRG 18 CHAPTER 1 Range of natural distribution A. Boratynski The range of genus Pinus L. The genus Pinus occupies an extensive area in the Northern Hemisphere, covering primarily the north temperate zone, particularly the boreal forest belt (taiga). In this zone pines, together with species from other genera of the Pinaceae family, such as Abies, Picea and Larix, are the most frequent trees which, depending on the environmental conditions, locally even dominate the natural vegetation. In spite of their abundance, there are not too many pine species in the boreal zone. The majority of species are found in regions situated much further south, in areas linking the temperate and tropical zones, with a particular increase in diversity in the mountains of the southern part of North America, in Central America, as well as in south-eastern Asia (Critchfield and Little, 1966; Mirov, 1967; Little and Critchfield, 1969; Farjon, 1984). In the last-mentioned region are to be found the southernmost natural stands of the genus Pinus (Fig. 1.1 ). Pinus merkusii Jungh et De Vriese crosses the equator on Sumatra (Indonesia) and goes as far south as 2° S (Stein, 1978), whereas P. caribaea Morelet only reaches 12° Ν in Nicaragua (Central America) (Critchfield and Little, 1966). In the far north some pines spread into the subarctic zone, and at times reach the arctic timberline. On the Eurasian continent P. pumila (Pall.) Regel has its northernmost stands at about 72Ί Ο' Ν along the Lower Lena tributaries in Siberia (Sokolov et al., 1977; Koropachinskij, 1983), and P. sylvestris L. reaches about 70°20' Ν in northern Norway (Hultén, 1950). In America the northern limit of pines is at latitudes about 10° lower than in Europe and Asia (see Fig. 1.1). P. banks/ana Lamb, ranges furthest to the north in America to about 62°30' Ν in the middle part of the Mackenzie river basin, while P. conforta Dougl. occurs up to latitude 62° Ν in the Upper Yukon (Little, 1971 ). Pines also have a large range with respect to altitude, since they grow from the sea level up to the timberline in the mountains. Of all the representatives of the genus, P. montezumae Lamb, occurs at highest elevations, at about 4 000 m in the mountains of Mexico (Mirov, 1967; Farjon, 1984). 19 2 0 Fig. 1.1. Range of the genus Pinus Natural distribution of Pinus sylvestris L Scots pine occurs on a very large area and it has formed many geographical races and varieties (Ruby and Wright 1976). Some of these are relatively easily distin- guishable on the basis of their morphology and frequently have been described in terms of lower taxonomical units (varieties or subspecies) of P. sylvestris (Gaus- sen et al., 1964; Pravdin, 1964). Some occurrences, growing most frequently at the limits of the range, have sometimes been discussed as independent species. In the present account, following the practice of the "Flora Europaea", all of these "small species" are considered only as subspecies or varieties of P. sylvestris s. I. (Gaussen et al., 1964), and the range described below (Fig. 1.2) concerns the species understood in this broad sense. Horizontal distribution The geographical distribution of P. sylvestris is described on the basis of the maps elaborated by Meusel and his co-workers (1965) and by Critchfield and Little (1966), simultaneously taking into account a number of rather insignificant changes and corrections in papers either not considered by the authors mentioned above, or published later (Wu Ch.-L., 1956; Coode and Cullen, 1965; Jasicova, 1966; Eliçin, 1971; Svoboda, 1971; Jalas and Suominen, 1973; Bobrov, 1975, 1978; Fenaroli and Gambi, 1976; Sokolov et al., 1977; Boratynska et al., 1981; Grubov, 1982; Welten and Sutter, 1982; Barbarich et al., 1986; Christensen, 1986). Scots pine is characterized by a very extensive range, most extensive of all species of the genus Pinus, and even of the whole Pinaceae family. The range of Scots pine spreads out longitudinally over a distance of about 14000 km. The westernmost occurrences of P. sylvestris are situated at about 8° W in Spain and the easternmost ones at 141 ° Ε in the Soviet Far East. In north-south direction the range of Scots pine covers a distance of about 2 700 km, with the northernmost occurrences at the latitude of 70°20' Ν in northern Scandinavia and the southern- most ones at about 37° Ν in the Sierra Nevada Mountains in Spain (see Fig. 1.2). In Scandinavia and on the Kola Peninsula P. sylvestris reaches the arctic timberline (Hultén, 1950; Sokolov et al., 1977), but further to the east the northern limit of the species stays south of the arctic timberline, which is locally formed there by Picea obovata Ledeb., Larix sibrica Ledeb. (including L. sukaczewii Dyils), L. gmelini (Rupr.) Kuzeneva and birch populations. The course of the northern range limit of P. sylvestris is mostly determined by climatic conditions; where continentality is considered to be a restrictive factor, characterized by high temperature amplitudes and low precipitation. It was observed that Scots pine does not grow in extremely continental conditions,, where the mean annual temperature amplitude is above 40 °C, the temperature sum of the growing season (above 10 °C) is less than 1 200 °C degree-days and the mean annual precipitation is less than 300 to 350 mm (Polikarpov, 1970, after Koropachinskij, 1983). 21

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