This page intentionally left blank Flowers on the Tree of Life Genetic and molecular studies have recently come to dominate botanical research at the expense of more traditional morphological approaches. This broad intro- duction to modern flower systematics demonstrates the great potential that floral morphology has to complement molecular data in phylogenetic and evolutionary investigations. Contributions from experts in floral morphology and evolution take the reader through examples of how flowers have diversified in a large variety of lineages of extant and fossil flowering plants. They explore angiosperm origins and the early evolution of flowers and analyse the significance of morphological characters for phylogenetic reconstructions on the tree of life. The importance of integrating morphology into modern botanical research is highlighted through case studies exploring specific plant groups where morpho- logical investigations are having a major impact. Examples include the clarification of phylogenetic relationships and an understanding of the significance and evolu- tion of specific floral characters, such as pollination mechanisms and stamen and carpel numbers. Livia Wanntorp currently works as a researcher at the Swedish Museum of Natural History in Stockholm, where she leads several projects involving flower morphology and systematics of many different groups of flowering plants. She is Associate Editor of the journal Plant Systematics and Evolution and President of the Swedish Systematics Association. Louis P. Ronse De Craene is director of the M.Sc. course on the Biodiversity and Taxonomy of Plants at the Royal Botanic Garden Edinburgh. His research interests include the morphology and evolution of flowers and encompass a broad range of angiosperm families. He is the author of Floral Diagrams: An Aid to Understanding Flower Morphology and Evolution (Cambridge University Press, 2010). The Systematics Association Special Volume Series series editor David J. Gower Department of Zoology, The Natural History Museum, London, UK The Systematics Association promotes all aspects of systematic biology by organizing conferences and workshops on key themes in systematics, running annual lecture series, publishing books and a newsletter, and awarding grants in support of systematics research. Membership of the Association is open globally to professionals and amateurs with an interest in any branch of biology, including palaeobiology. Members are entitled to attend conferences at discounted rates, to apply for grants and to receive the newsletter and mailed information; they also receive a generous discount on the purchase of all volumes produced by the Association. The first of the Systematics Association’s publications The New Systematics (1940) was a classic work edited by its then-president Sir Julian Huxley. Since then, more than 70 volumes have been published, often in rapidly expanding areas of science where a modern synthesis is required. The Association encourages researchers to organize symposia that result in multiauthored volumes. In 1997 the Association organized the first of its international Biennial Conferences. This and subsequent Biennial Conferences, which are designed to provide for systematists of all kinds, included themed symposia that resulted in further publications. The Association also publishes volumes that are not specifically linked to meetings, and encourages new publications (including textbooks) in a broad range of systematics topics. More information about the Systematics Association and its publications can be found at our website: www.systass.org Previous Systematics Association publications are listed after the index for this volume. Systematics Association Special Volumes published by Cambridge University Press: 78. Climate Change, Ecology and Systematics Trevor Hodkinson, Michael Jones, Stephen Waldren and John Parnell 79. Biogeography of Microscopic Organisms: Is Everything Small Everywhere? Diego Fontaneto T S A S he ystematics ssociation pecial V 80 olume Flowers on the Tree of Life Edited by Livia Wanntorp Department of Phanerogamic Botany, Swedish Museum of Natural History, Stockholm, Sweden Louis P. Ronse De Craene Royal Botanic Garden Edinburgh, UK THE Systematics ASSOCIATION cambridge university press Cambridge, New York, Melbourne, Madrid, Cape Town, Singapore, São Paulo, Delhi, Tokyo, Mexico City Cambridge University Press The Edinburgh Building, Cambridge CB2 8RU, UK Published in the United States of America by Cambridge University Press, New York www.cambridge.org Information on this title: www.cambridge.org/9780521765992 © The Systematics Association 2011 This publication is in copyright. Subject to statutory exception and to the provisions of relevant collective licensing agreements, no reproduction of any part may take place without the written permission of Cambridge University Press. First published 2011 Printed in the United Kingdom at the University Press, Cambridge A catalogue record for this publication is available from the British Library Library of Congress Cataloguing in Publication data Flowers on the tree of life / [edited by] Livia Wanntorp, Louis P. Ronse De Craene. p. cm. – (Systematics Association special volume series) Includes bibliographical references and indexes. ISBN 978-0-521-76599-2 1. Flowers – Morphology. 2. Flowers – Evolution. 3. Angiosperms – Morphology. 4. Angiosperms – Evolution. I. Wanntorp, Livia. II. Ronse Decraene, L. P. (Louis Philippe) QK653.F59 2011 581.3′8–dc23 2011019851 ISBN 978-0-521-76599-2 Hardback Cambridge University Press has no responsibility for the persistence or accuracy of URLs for external or third-party internet websites referred to in this publication, and does not guarantee that any content on such websites is, or will remain, accurate or appropriate. v Contents List of contributors page vii Acknowledgements ix 1 Introduction: Establishing the state of the art – the role of morphology in plant systematics 1 Louis P. Ronse De Craene and Livia Wanntorp 2 Spatial separation and developmental divergence of male and female reproductive units in gymnosperms, and their relevance to the origin of the angiosperm flower 8 Richard M. Bateman, Jason Hilton and Paula J. Rudall 3 New flowers of Laurales from the Early Cretaceous (Early to Middle Albian) of eastern North America 49 Maria von Balthazar, Peter R. Crane, Kaj Raunsgaard Pedersen and Else Marie Friis 4 Tracing the early evolutionary diversification of the angiosperm flower 88 James A. Doyle and Peter K. Endress 5 Changing views of flower evolution and new questions 120 Peter K. Endress 6 Centrifugal stamens in a modern phylogenetic context: was Corner right? 142 Paula J. Rudall 7 Evolution of the palm androecium as revealed by character mapping on a supertree 156 Sophie Nadot, Julie Sannier, Anders Barfod and William J. Baker 8 Comparative floral structure and development of Nitrariaceae (Sapindales) and systematic implications 181 Julien B. Bachelier, Peter K. Endress and Louis P. Ronse De Craene 9 Multiplications of floral organs in flowers: a case study in Conostegia (Melastomataceae, Myrtales) 218 Livia Wanntorp, Carmen Puglisi, Darin Penneys and Louis P. Ronse De Craene vi contents 10 Ontogenetic and phylogenetic diversification in Marantaceae 236 Alexandra C. Ley and Regine Claßen-Bockhoff 11 Floral ontogeny of Acacia celastrifolia: an enigmatic mimosoid legume with pronounced polyandry and multiple carpels 256 Gerhard Prenner 12 Floral development of Napoleonaea (Lecythidaceae), a deceptively complex flower 279 Louis P. Ronse De Craene Taxon index 296 Subject index 305 Colour plate section appears between pages 214 and 215. vii Contributors Julien B. Bachelier Institute of Systematic Botany, University of Zurich, Switzerland William J. Baker Royal Botanic Gardens, Kew, UK Maria von Balthazar Department of Systematic and Evolutionary Botany, University of Vienna, Austria Anders Barfod Department of Biological Sciences, Aarhus University, Denmark Richard M. Bateman School of Geography, Earth and Environmental Sciences, University of Birmingham, UK Regine Claßen-Bockhoff Institut für Spezielle Botanik und Botanischer Garten, University Mainz, Germany Peter R. Crane School of Forestry and Environmental Studies, Yale University, New Haven, CT, USA James A. Doyle Department of Evolution and Ecology, University of California, Davis, CA, USA Peter K. Endress Institute of Systematic Botany, University of Zurich, Switzerland Else Marie Friis Department of Palaeobotany, Swedish Museum of Natural History, Stockholm, Sweden Jason Hilton School of Geography, Earth and Environmental Sciences, University of Birmingham, UK Alexandra C. Ley Department of Evolutionary Biology and Ecology, Université Libre de Bruxelles, Belgium Sophie Nadot Laboratoire Ecologie, Systématique et Evolution, Université Paris- Sud, Orsay, France Kaj Raunsgaard Pedersen Department of Geology, University of Aarhus, Denmark Darin Penneys Department of Botany, California Academy of Sciences, San Francisco, CA, USA Gerhard Prenner Jodrell Laboratory, Royal Botanic Gardens, Kew, UK Carmen Puglisi Royal Botanic Garden Edinburgh, UK viii list of contributors Louis P. Ronse De Craene Royal Botanic Garden Edinburgh, UK Paula J. Rudall Jodrell Laboratory, Royal Botanic Gardens, Kew, UK Julie Sannier Laboratoire Ecologie, Systématique et Evolution, Université Paris- Sud, Orsay, France Livia Wanntorp Department of Phanerogamic Botany, Swedish Museum of Natural History, Stockholm, Sweden