Table Of ContentDevelopments 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:
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for all remaining areas
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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
