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Bibliotheca Botanica 161 ISSN 0067-7892 Bibliotheca Botanica Bibliotheca Botanica, Volume 161, 2012 Original Contributions to Botany, Founded in 1886 V

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Bibliotheca Botanica 161 ISSN 0067-7892 Bibliotheca Botanica Bibliotheca Botanica, Volume 161, 2012 Original Contributions to Botany, Founded in 1886 V Bibliotheca Botanica 161 ISSN 0067-7892 Bibliotheca Botanica Bibliotheca Botanica, Volume 161, 2012 Original Contributions to Botany, founded in 1886 V. M. Dörken, The evolutionary relevance of vegetative long-shoot / short-shoot differentiation in gymnospermous tree species Edited by H. W. Lack, Berlin, P. Leins, Heidelberg and S. Porembski, Rostock This well illustrated monograph treats the correla- sequently, the evolutionary pathway to the long- M. Dörken – Gymnospermous tree species V. Volume 161 tion of deciduousness (shedding leaves at a cer- shoot / short-shoot differentiation in gymnospermous tain season) and long-shoot / short-shoot differen- tree species must be different from the evolutionary tiation focused on gymnospermous tree species. traits in angiospermous trees. It is shown that some The vast majority of gymnosperms are evergreen evergreen gymnosperms can be regarded as de- Veit Martin Dörken and within this group deciduousness has generally rived from deciduous ancestors while others still re- been regarded as a derived feature. Comparative present the primitive deciduous condition. The evolutionary relevance of studies of angiospermous tree species indicate that This monograph consists of two parts: In the first the vegetative long-shoot / short-shoot differentiation part several gymnospermous and angiospermous vegetative long-shoot / short-shoot differentiation correlates well with deciduousness. The total leaf tree species have been investigated morphologi- area of an entire short-shoot equals the leaf area of cally, anatomically and physiologically. In the sec- in gymnospermous tree species lamina of a single long-shoot leaf. So the lamina of ond part those data were mapped on palaeobotanic a long-shoot leaf is replaced by the total leaf area of and palaeogeographic data in order to test the initial an entire short-shoot in the following vegetation pe- hypothesis that deciduousness has been more fre- riod. This simple correlation is not observed in any quent among gymnospermous tree species in the of the studied gymnosperms except Ginkgo. Con- past than the recent diversity can reflect. This book is of interest for all botanists and researchers on angiospermous and gymnospermous trees. www.schweizerbart.de ISBN 978-3-510-48032-6 ISSN 0067-7892 E Schweizerbart Science Publishers BIBLIOTHECA BOTANICA, VOLUME 161 BIBLIOTHECA BOTANICA Original Contributions to Botany Edited by H. W. Lack, Berlin, P. Leins, Heidelberg and S. Porembski, Rostock Volume 161 Veit Martin Dörken The evolutionary relevance of vegetative long-shoot/ short-shoot differentiation in gymnospermous tree species with 41 fi gures and 12 tables Schweizerbart Science Publishers Stuttgart 2012 The evolutionary relevance of vegetative long-shoot/short- shoot differentiation in gymnospermous tree species by Veit Martin Dörken with 41 fi gures and 12 tables Schweizerbart Science Publishers Stuttgart 2012 Veit Martin Dörken: The evolutionary relevance of vegetative long-shoot/short-shoot differentiation in gymnospermous tree species Author’s address: Dr. Veit Martin Dörken Department of Biology, University Konstanz Universitätsstraße 10, 78464 Konstanz, Germany [email protected] Front cover: Metasequoia glyptostroboides and Larix decidua ISBN 978-3-510-48032-6 ISSN 0067-7892 Information on this title: www.schweizerbart.com/9783510480326 © 2012 E. Schweizerbart’sche Verlagsbuchhandlung (Nägele u. Obermiller), Stuttgart, Germany 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, electronic, mechanical photocopying, recording, or otherwise, without the prior written permission of E. Schweizerbart’sche Verlagsbuchhandlung, Stuttgart Publisher: E. Schweizerbart’sche Verlagsbuchhandlung (Nägele u. Obermiller), Johannesstraße 3A, 70176 Stuttgart, Germany [email protected] www.schweizerbart.de ∞ Printed on permanent paper conforming to ISO9706-1994 Typesetting: Satzpunkt Ursula Ewert GmbH, Bayreuth Printed in Germany by Tutte Druckerei GmbH, Salzweg bei Passau Contents 1 Contents Abstract . 3 1 Introduction . 5 2 Materials and methods . 9 2.1 Length-frequence diagrams . 9 2.2 Measurement of leaf area . 12 2.3 Measurement of the chlorophyll content . 12 2.4 Measurement of shoot weight . 12 2.5 Electronic microscopy, paraffi n technique and photography . 12 2.6 Special terms used in this study . 13 3 Results . 15 3.1 Correlation between foliage and shoot system . 15 3.2 Differences in length between long- and short-shoots . 16 3.3 Comparisons of annual investment in shoot biomass between long- and short shoots . 16 3.4 Number of long- and short-shoots in different ages of an individual . 24 3.5 Number of leaves and single-leaf area of long- and short-shoot leaves . 25 3.6 Annual investment in shoot biomass per long-shoot leaf and leaf area . 29 3.7 Annual investment in shoot biomass per short-shoot leaf and leaf area . 29 3.8 Annual investment in shoot biomass per single-leaf and leaf area in taxa without shoot differentiation . 30 3.9 Comparison of annual investment in shoot biomass per 1 cm2 long-shoot leaf area and 1 cm2 short-shoot leaf area . 31 3.10 Comparison of the total leaf area of an entire short-shoot with single long-shoot leaf . 31 3.11 Anatomical comparisons between long and short-shoot leaves . 31 3.12 Chlorophyll content . 31 3.13 Shoot abscission . 31 3.13.1 Shoot abscission in Cupressaceae (Gymnospermae) . 38 3.13.2 Shoot abscission in Larix decidua (Pinaceae, Gymnospermae) . 38 3.13.3 Shoot abscission in Quercus robur (Fagaceae, Angiospermae) . 38 3.14 Anomaliess . 50 3.14.1 Pinus with anomalous formed short-shoot . 50 Pinus sylvestris . 50 Pinus monophylla . 50 3.14.2 Sciadopitys verticillata with anomalous formed cladodes . 56 4 Discussion . 61 4.1 Correlation between shoot differentiation and deciduousness . 61 4.2 Correlation between long- and short-shoots . 61 4.3 Correlation between long- and short-shoot leaves . 63 4.4 Shoot abscission and its evolutionary meaning . 65 4.4.1 Shoot abscission in Quercus robur, Fagaceae (Angiospermae) . 65 4.4.2 Shoot abscission in Larix, Pinaceae (Gymnospermae) . 66 4.4.3 Shoot abscission Cupressaceae (Gymnospermae) . 66 4.5 Evolutionary implications for different groups . 70 4.5.1 Angiospermous trees . 70 4.5.1.1 Cercidiphyllum (Cercidiphyllaceae, Angiospermae) . 70 4.5.1.2 Nothofagus (Nothofagaceae, Angiospermae) . 70 2 Contents 4.5.2 Gymnospermous trees . 72 4.5.2.1 Pinus (Pinaceae, Gymnospermae) . 72 4.5.2.2 Cedrus (Pinaceae, Gymnospermae) . 75 4.5.2.3 Larix (Pinaceae, Gymnospermae) . 76 4.5.2.4 Pseudolarix (Pinaceae, Gymnospermae) . 76 4.5.2.5 Metasequoia (Cupressaceae s.l., Gymnospermae) . 76 4.5.2.6 Sciadopitys verticillata (Sciadopityaceae, Gymnospermae) . 77 4.5.2.7 Ginkgo (Ginkgoaceae, Gymnospermae). 78 5 Conclusions . 81 6 Acknowledgements . 83 7 Bibliography . 85 Index . 91 List of genera and species . 92 Abstract 3 Abstract A vast majority of gymnosperms is evergreen and with- entiation correlates well with deciduousness. The total in this group deciduousness has generally been regard- leaf area of an entire short-shoot has the same leaf area ed as a derived feature. Evergreen taxodioid Cupres- as the lamina of a single long-shoot leaf. So the lamina saceae s.l. like Sequoia and Sequoiadendron, however, of a long-shoot leaf is replaced by the total leaf area of show an abscission structure at the base of short-shoots an entire short-shoot in the next vegetation period. This or short-shoot-like branches similar to the abscission simple correlation does not occur in any of the gymno- structure of deciduous taxa like Taxodium, Glyptostro- sperms studied except Ginkgo. bus, and Metasequoia. Such an abscission structure in It could be shown that some evergreen gymno- evergreen taxa should be better understood.
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