The Families and Genera of Vascular Plants
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THE FAMILIES AND GENERA OF VASCULAR PLANTS Edited by K. Kubitzki Volumes published in this series Volume I Pteridophytes and Gymnosperms Edited by K.U. Kramer and P.S. Green (1990) Date of publication: 28.9.1990 Volume II Flowering Plants. Dicotyledons. Magnoliid, Hamamelid and Caryophyllid Families Edited by K. Kubitzki, J.G. Rohwer, and V. Bittrich (1993) Date of publication: 28.7.1993 Volume III Flowering Plants. Monocotyledons: Lilianae (except Orchidaceae) Edited by K. Kubitzki (1998) Date of publication: 27.8.1998 Volume IV Flowering Plants. Monocotyledons: Alismatanae and Commelinanae (except Gramineae) Edited by K. Kubitzki (1998) Date of publication: 27.8.1998 Volume V Flowering Plants. Dicotyledons: Malvales, Capparales and Non-betalain Caryophyllales Edited by K. Kubitzki and C. Bayer (2003) Date of publication: 12.9.2002 Volume VI Flowering Plants. Dicotyledons: Celastrales, Oxalidales, Rosales, Cornales, Ericales Edited by K. Kubitzki (2004) Date of publication: 21.1.2004 Volume VII Flowering Plants. Dicotyledons: Lamiales (except Acanthaceae including Avicenniaceae) Edited by J.W. Kadereit (2004) Date of publication: 13.4.2004 Volume VIII Flowering Plants. Eudicots: Asterales Edited by J.W. Kadereit and C. Jeffrey (2007) Volume IX Flowering Plants. Eudicots: Berberidopsidales, Buxales, Crossosomatales, Fabales p.p., Geraniales, Gunnerales, Myrtales p.p., Proteales, Saxifragales, Vitales, Zygophyllales, Clusiaceae Alliance, Passifloraceae Alliance, Dilleniaceae, Huaceae, Picramniaceae, Sabiaceae Edited by K. Kubitzki (2007) The Families and Genera of Vascular Plants Edited by K. Kubitzki Flowering Plants · Eudicots IX Berberidopsidales, Buxales, Crossosomatales, Fabales p.p., Geraniales, Gunnerales, Myrtales p.p., Proteales, Saxifragales, Vitales, Zygophyllales, Clusiaceae Alliance, Passifloraceae Alliance, Dilleniaceae, Huaceae, Picramniaceae, Sabiaceae Volume Editor : K. Kubitzki in Collaboration with C. Bayer and P.F. Stevens With 174 Figures 123 Professor Dr. Klaus Kubitzki Universität Hamburg Biozentrum Klein-Flottbek und Botanischer Garten Ohnhorststraße 18 22609 Hamburg Germany Library of Congress Control Number: 2006928744 ISBN-10 3-540-32214-0 Springer Berlin Heidelberg New York ISBN-13 978-3-540-32214-6 Springer Berlin Heidelberg New York This work is subject to copyright. All rights are reserved, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfilm or in any other way, and storage in data banks. Duplication of this publication or parts thereof is permitted only under the provisions of the German Copyright Law of September 9, 1965, in its current version, and permissions for use must always be obtained from Springer-Verlag. Violations are liable for prosecution under the German Copyright Law. Springer is a part of Springer Science+Business Media springer.com © Springer-Verlag Berlin Heidelberg 2007 The use of general descriptive names, registered names, trademarks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. Cover design: WMXDesign, Heidelberg, Germany Typesetting and production: LE-TEX Jelonek, Schmidt & Vöckler GbR, Leipzig, Germany Printed on acid-free paper 31/3150/YL – 5 4 3 2 1 0 Preface The present volume contains treatments of various eudicot orders which all are strongly supported in molecular analyses. A first group comprises Proteales, Buxales and the enigmatic Sabiaceae which, together with Ranunculales and Trochodendrales, treated earlier in Vol. II of this series, represent the basal grade of early-diverging eudicots. Al- though all of them have clearly tri-orate pollen, making them eudicots, they otherwise lack the strict eudicot floral organisation, particularly with regard to flower merosity, phyllotaxis and perianth structure. The same is true for the order Gunnerales which, however, according to findings of molecular systematics, forms part of the strongly sup- ported core eudicots. All these orders to some extent bridge the morphological gap be- tween basal angiosperms and typical core eudicots. Plesiomorphic floral traits, although less pronounced, are also found among the isolated but clearly core eudicotyledonous Saxifragales, treated in this volume, and particularly in the early-diverging woody fam- ilies of this order. Some of these latter families were included in Vol. II, which followed a classification used prior to the discovery of the modern concept of Saxifragales. The exact interrelationships among the early-diverging eudicot orders still remain largely unresolved. This is even more true for many of the core eudicot orders included in the present volume, i.e. Vitales, Crossosomatales, Geraniales, Zygophyllales and Myrtales, and also for two of the families not assigned to order, i.e. Huaceae and Picramniaceae. A possible relationship between Dilleniaceae and woody Caryophyllales, as suggested by recent studies, opens an interesting new perspective on the evolution of this family. Included in this volume are two subclades of the vast Malpighiales. These are Passi- floraceae with two satellite families, and Clusiaceae/Hypericaceae with Podostemaceae, which recently have been identified as very close relatives. My deep thanks go to all authors of this volume, who have provided such highly in- teresting and scholarly contributions, and to all those who have freely shared additional information and/or have commented on earlier drafts of the contributions. These include B.G. Briggs, T. Clifford, G. Jordan, B. Makinson, P. Olde, R. Barker, J.A. Doyle, P.J. Rudall and C.A. Furness (Proteaceae); H. Manitz (Aphanopetalaceae); A.E. Orchard (Halor- agaceae); P.H. Linder and M. Weigend (Geraniales); A. Bernhard and W.J.J.O. de Wilde (Passifloraceae); the late M. Ricardi S. (Malesherbiaceae); I. Jäger-Zürn and T. Philbrick (Podostemaceae); and S. Renner, P.G. Wilson and J. Schönenberger (Myrtales). I am also grateful to M.L. Matthews and P.K. Endress for showing me their papers prior to publication elsewhere. Mark C. Chase is thanked for always making available the newest results of his pathbreaking studies. The copyright holders of the illustrations included in this volume are thanked for their generous permission to use their valuable material. As always, it is a pleasure to acknowledge the agreeable collaboration with the staff of Springer-Verlag, who kindly responded to all requests I had in connection with the production of this volume, and to thank Monique T. Delafontaine for her meticulous copy editing of the manuscript. Hamburg, September 2006 K. Kubitzki Contents Introduction to the Groups Treated in this Volume K. Kubitzki ................................. 1 IntroductiontoBerberidopsidales ................................. 1 IntroductiontoBuxales .......................................... 2 IntroductiontotheClusiaceaeAlliance(Malpighiales) ................. 3 IntroductiontoCrossosomatales ................................... 3 IntroductiontoFabales ........................................... 5 IntroductiontoGeraniales ........................................ 5 IntroductiontoGunnerales ....................................... 7 IntroductiontoMyrtales ......................................... 7 Introduction to the Passifloraceae Alliance (“Passiflorales” = Malpighiales) .................................... 12 IntroductiontoProteales ......................................... 12 IntroductiontoSaxifragales ....................................... 15 IntroductiontoVitales ........................................... 18 IntroductiontoZygophyllales ..................................... 19 FamiliesUnassignedtoOrder ..................................... 20 General References ............................................ 21 Aextoxicaceae K. Kubitzki ................................. 23 Alzateaceae S.A. Graham ................................ 26 Aphanopetalaceae K. Kubitzki ................................. 29 Aphloiaceae K. Kubitzki ................................. 31 Berberidopsidaceae K. Kubitzki ................................. 33 Bonnetiaceae A.L. Weitzman, K. Kubitzki and P.F.Stevens ..... 36 Buxaceae E. Köhler ................................... 40 Clusiaceae-Guttiferae P.F. Stevens ................................. 48 Combretaceae C.A. Stace .................................. 67 Crassulaceae J. Thiede and U. Eggli ........................ 83 Crossosomataceae V. S o s a ..................................... 119 Crypteroniaceae S.S. Renner ................................. 123 Daphniphyllaceae K. Kubitzki ................................. 127 VIII Contents Didymelaceae E. Köhler ................................... 129 Dilleniaceae J.W. Horn ................................... 132 Geissolomataceae F. Forest .................................... 155 Geraniaceae F. Albers and J.J.A. Van der Walt .............. 157 Grossulariaceae M. Weigend ................................. 168 Gunneraceae H.P. Wilkinson and L. Wanntorp .............. 177 Haloragaceae K. Kubitzki ................................. 184 Huaceae C. Bayer .................................... 191 Hypericaceae P.F. Stevens ................................. 194 Iteaceae K. Kubitzki ................................. 202 Ixerbaceae J.V. Schneider ............................... 205 Krameriaceae B.B. Simpson ................................ 208 Ledocarpaceae M.