SYSTEMATIC ANATOMY OF THE DICOTYLEDONS A HANDBOOK FOR LABORATORIES OF \PURE AND APPLIED BOTANY BY DR. HANS SOLEREDER PROFESSOR OF BOTANY IN THE UNIVERSITY OF ERLANG.N 'tRANSLATm BY L. A. BOODLE, F.L.S. ROYAL BOTANIC GARDENS, KEW AND F. E. FRITSCH, D.Se., PH.D., F.L.S. ASSISTANT PROFESS01t. OF BOTANY, UNIVERSI'!"Y OF LONDON, UNIVERSITY COLl.EGII: AND LECTUR.ER IN BO'rANY, EAST LONDON COLLEGE, UNIVERSITY OF LOND(\N REVJ6ED BY D. H. SCOTT, M.A., LL.D" Pn.D., F.R.S. LATE IIONORARY KEEPER ~OF~TlU!.11?DRi'tLL LABORATORY ROYAL BOTANIC GARDENS KEW 1 VOL. I INTRODUCTION, POLYPETALAE, GAMOPETALAE WITH 153 FIGURES IN THE TEXT AJAY BOOK SERVICE NEW DELHI INDIA Published by: AJ A Y BOOK SERVICE (Publishers &: Distributors) 704 Chandni Mahal, Daryaganj, New Delhi-II 0 002 FIrst Printed 1908 Indian Reprint 1986 Printed at: SU5HIl PRINTERS At New Sachdeva Press, A-6/ 1, Mayapuri, Phase-I, NewDelhi-11 0064 TO PROF. DR. L. RADLKOFER SE1NEM HOCHVEREHRTEN LSHR.ER UND VORSTANDE THIS BOOK IS DEDICATED BY THE AUTHOR EDITORIAL PREFACE PROFESSOR SOLEREDER'S prefaces, to the original German edition and to the present English translation, explain so clearly the objecf and 5(",.-0£ his book that .it is unnecessary to enter on any further detailed exposition of its putpose. The work is universally recognized among Botanists as the standard authority on the subject of, the anatomy of plants as applied to systematic purposes; though limited to the Dicotyledons, Professor Solereder's book must serve as the type of all treatises dealing with the stf\1c,ture of other clas~ from a similar point of view. Covering, as it does, with marvellous coI\lplete ness and exactitude, the most extensive and important division of the Vegetable Kingdo~ the' Systematic Anatomy' forms an indispensable work ot reference to all engaged in systematic studies, and serves at the same t.i,m e as by far the best source from which students may gain an idea of the objects and methods of this young and vigorous branch of Botany. Until recently, the study of anatomical structure in relation to taxonomy has been but little pursued in this country, though a beginning was made y many yeal'$ ago, by some of our greatestsystematists. The present t~anslation will, it is hoped, much facilitate the introduction of modern anatomical methods . into the daily work of the Herbarium and the Economic Museum. One 01 _.. th~ advantages of the study of systematic anatomy is that it brings together the work of the laboratory botanist and the taxonomist, each of whom has hitherto often remained too much isolated in his own department. The 'case for the anatomical method is put by the author with singulCLf judgement and moderation, as will be seen from the Introduction and the Concluding Remarks. The latter form a very full and valuable summary of the various anatomical characters and their value in classification; they may be read with interest and advantage by all students of botany, wh¥ther their bent be systematic or otherwise. It may be pointed ot1t that the Engll~h edition has heen brought thoroughly up to date, both the Addenda and the Concluding Remarks having received from the author's hand all the emendations which will be embodied in thel new German edition. vi PREFACE The work of translating so extensive a treatise, embracing such a vast amount of minute and accurate detail, has naturally been extremely arduous; and the translators, Mr. Boodle and Dr. Fritsch, are to be congratulated on the successful completion of their labours. The translation, first begun by Mr. Boodle, was carried out jointly by him and his colleague up to the end of the Natural Orders; the work of translating the Addenda and Concluding Remarks tell to Dr. Fritsch alone. My own part has consisted in th~ .revision of the proofs throughout, and in the necessary consultations with the translators on all questions that arose in the course of·their work. I have aimed at securing a sufficient uniformity of style throughout the book, and have satisfied myself that the transhition faithfully conveys the author's meaning. The rendering of technical terms has in some cases presented a certain amount of difficulty; it is ~roposed to add a glossary of the less familiar terms, with their German originals, at the end of the second volume. ~e additional references to the literature, which came in too late to be appended to the lists under each Natural Order in the Addenda, will be found collech.d at the end of the book. D. H. SCOTT. March 31, '908. PREFACE BY THE AUTHOR TO THE ENGLISH EDITION THE present English edition of the • Systematische Anatomie der Dico tyledonen: originally published in the year 1899 by F. Enke in Stuttgart, contains the translation of the Introduction and of the Anatomical Characters of the Dicotyledonous Orders (pp. 1-903 of the German edition). This, the principal part of the book, is followed by an appendix, compiled at the request of my English colleagues, which summarizes, under the respectiv(.. Orders, the results of recent researches. The book terminates with the • Concluding Remarks,' which give a survey of. the anatomical characters of the vegetative organs and of the distribution of these characters among the Dicotyledons, and constitute a revised treatment of the • Schlussbemerkungert" of the original German edition (pp. 904-79). As regards the arrangement of the subject-matter and the termmology of the species, exactly the same plan has been followed in the Addenda as in the principal part of the book (see the preface to the German edition)~ The Orders are dealt with in th~ sequence of Bentham and Hooker's system. In the descriptions of the individual Orders supplementary details to the' review of the anatomical features,' the structure of the leaf,' the structure of the f f axis • and in some cases also the' structure of the root • succeed one another. In the sections devoted to the structure of the leaf and the structure of the axis the anatomical characters are discussed in the same serial order as in the principal part of the book. A considerable amount of trouble has been taken in completing the references to literature; papers, of which I was acquainted with the title only or of which only abstracts were available, are again placed in square brackets. Readers who are conversant with the subject will also find new observations, which have not yet been published elsewhere, intro duced from time to time. In concluding, I should like above all to express the pleasure which the translation of my book has afforded me, for it carries with it the hope that the anatomical method will find new adherents, who are so necessary for its further elaboration and obligatory introduction into all systematic researches (monographs, questions of affinity, establishment of new genera and species). At the same time I wish to tender my heartiest thanks to the translators for the great labour expended on the preparation of this work, as well as to the Delegates of the Clarendon Press for their invariable readiness to meet my wishes, and,-last not least,-to all the colleagues who ha\'e assisted me in my work by the forwarding of separate copies or of material for investigation. H. SOLEREDER. ERLANGItN, Odoher, 1907. PREFACE TO THE GERMAN EDITION 'THE next hundred years will be devoted to the anatomical methotl; I these significant words of an illustrious Botanist are directly respQnsible for 1 the production of this book. Repeated earlier attempts had bee'll made to employ the charac!ers afforded by the anatomy of the vegetatiVe and' reproductive organs in conjunction with external features. for syst~matic y purposes and for a long time such characters had been of service to those y engaged in applied science as a means of determining vegetable products. Both De Candolle and Endlicher had found anatomical characters useful In the division of the Phanerogams into the two great groups the Monocotyledons and Dicotyledons. It is only in recent times, however, that certain well-known investigators. and foremost among them Radlkofer have succeeded in y demonstrating in a convincing manner the scientific 'value of anatomical characters for systematic purposes, and in founding an anatomical method. With the help of this they have been able to solve systematic problems which otherwise could scarcely have been elucidated, and to throw an essentially new ]~ht on scientifio .questions relating to the delimitation of species, genera and Orders, as well as to natural affinities. Considering these brilliant results it is not surprising that small succe$S attended the opposition of systematists of the old school who were averse to the un J accustomed use of the microscope in this department of Botany, and feared lest they should have. to resign their favourite science to their younger, initiated colleagues; and it is not to be wondered at that, the new me~od gained a sound footing. On the other hand, it is not unnatural that a number of Botanists of the new school expected too much from the anatomical jllethod,and not only looked for a revival of systematic Botany on a surer foundation, but also for an entirely new Natural System. Now that nearly a decade and a half have elapsed since Radlkofer's memorable speech was delivered, and hundreds of papers have been published affording numerous data on the val ue of anatomical characters considered generally or within the limits of certain Orders, the time appears to have come for summarizing the results attained, and, with the aid of the data which have been established, for forming a decision as to how far our expectations of the anatomical method have been justified. Having been trained in Radlkofer's school of Bo{any, my own scientific researches, as well as a study of the literature of the subject and the share in the direction of Radlkofer'.s laboratory entrusted to me, imbued me with a keen interest in the development of the new line of systematic aootomical investigation-first elevated by Radlkofer to the rank of a scientific method. A(a consequence I have for many years realized the need for a summary of 1 R~(l)kofeTI Ueber die Melb. in d. bot. Syst. insbes., d. anat. Methode. Festre(le, Milnich, J883. PREFACE TO THE GERMAN EDITION rx the results acquired, and have desired to undertake the work. I therefore gladly accepted the publisher's invitation to write a book on this subject. It cannQt be expected that the value of this work will rise to the level of the high aims of the anatomical method, not\\'ithstanding the great amount of labour expended by me on the study of the various treatises (which sprang up like mUshrooms!)~ on the re-investigation of doubtful statements, and on numerous supplemelltary investigations of my own, which a reader conversant with the subject will find scattered here and there. It was not possible to give the different Orders a completely uniform treatment, since the investigations were not ·a1ways equally thorough; in those cases in which only preliminary statements were available, it was necessary to quote them in some detail, in order to avoid generalizations, and to enable the reader to appreciate the gaps in our present knowledge. The prolonged period during which this work has been in course of pz:.cparation may also form an excuse for such inequalities of treatment as have been retained. As a result of my honest endeavours, r however, I think may say that this book gives a review of the results hitherto obtained, and will thus form a basis for future supplementary investigations, and a useful handbook for those concerned with systematic problems. It will also be of value in applied Botany, inasmuch as the economic Botanist had hitherto no handbook for the determination of raw vegetable products of unknown origin, and also to the physiologist who wishes to obtain information on the anatomy of the plants used in his experiments, and on the distribution of certain anatomi~a1 features which are cOl1necte(l with special functions. As regards the general plan of the book and the arrangement of the 8ubject-matter in the description of the individual Order"), the following points may be mentioned. The Introduction deals chiefly with the value of the anatomical method and of anatomical characters, and touches on the problems connected with this subject. The Introduction is followed by a description of the anatomical features in the individual Dicotyledonous Orders, the serial arrangement of the latter being the same as in Bentham and Hook~ Genera Plantarum. The description of each Order begins with a short review of the anatomical characters, with the help of which it is possible to determine whether a given plant may be regarded as a member of that Order or not. More detfliled statements, forming an amplification of this review, are given in the succeeding sections, which are, as a rule (in the larger Natural Orders), arranged under -the headings-' Structure of the Leaf: and ' Structure of the Axis.' In order to avoid confusion, the different species cited in connexion with the various anatomical features are enumerated, without regard to synonymy, under the names assigned to them by the authors who demonstrated these anatomical characters in the species in question; in those cases in which to the specific name is placed between inverted commas, this is done indicate that I was unacquainted with the name. In the sections on the structure of the leaf and axis the individual anatomical features are taken serially, and certain of them afe made prominent by the use of Clarendon type. In this connexion it is only necessary to point out that characters which are common x PREFACE TO THE GER,l{AN EDITION to the leaf and axis (such as the hairy covering, oxalate of lime and other contents, internal secretory organs, &c.) are in most cases described jointly for both these organs ~ these features are generally dealt with 1-mder the structure of the leaf, since they are more readily .observed in the leaf, and have consequently been examined for the most part with special reference to that organ. Statements on the structure of the root are occa'\ionally appended to the section on the structure of the axis. In general, however, the root is not dealt with in detail, because the existing researches on this subject are not sufficiently comprehensive, owing to the difficulty in procuring material, while the results obtained are not of any special consequence in the diagnosis of an Order. At the end of the description of each Order there will be found a list of the papers published up to 18g8; these fonn the basis of the statements included under the Ox:der, and from them the reader may The obtain further information. arrangement of the refer.ences in these lists of literature is chronological; as a rule the titles of the papers are quoted in an abbreviated form, and .only such as contain nq more than a few isolated data are omitted; papers, of which I was acquainted with the title only, are placed in square brackets. The principal part of the book is followed by the' Concluding Remarks'; these constitute a summary of the existing data in the form of an enumeration of the anatomical characters, whkh are con sidered in relation to their systematic value: the Concluding Remarks together with the Introduction may also serve as a guide to the anatomical method. In the use of this book a knowledge of plant-anatomy is taken for granted. Readers requiring information on this subject may, besides using the ordinary botanical textbooks, consult such works as De Bary's Vergleichende Anatomic der Vegetationsorgane, translated in this series, Haberlandt's Physiologische Pflanzenanatomie, Tschirch's Angewandte Pftanzenanatomie, or Van Tieghern's Traite de Botanique, vol. i. In conduding this preface I wish to express my warmest thanks to all those who have assisted me""in the publication of this book. Above all I am indebted to Professor L. Radlkofer, to whom I dedicate this book with grateful veneration. I am also deeply indebted to the Koniglich Bayerische Akadenu.e der Wissenschaften, who favoured my work with material assistance, and to the Directors of the Konigliche Hof- und Staatsbibliothek in Munich, whose generosity enabled me to make ample use of the valuable books contained in the library; I also wish to express my thanks to Profes50rs Goebel, Urban, Schenck and Vol kens, and to other colleagues, who assisted me either i~ obtaining literature or by granting permission to make use of a large number of illustrations in their published works, and to Herr Dunzinger, who drew a number of the original figures from my preparations. Lastly, I wish to thank the publisher for the care he has bestowed on the prod uction of this book. I trust that it may prove worthy of the yaluable assistance which I have received. THE AUTHOR. MUNL(lI, .hbyut1Yy, J 898.
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