THE FAMILIES AND GENERA OF VASCULAR

Edited by K. Kubitzki

For further volumes see list at the end of the book and: http://www.springer.com/series/1306 . The Families and Genera of Vascular Plants

Edited by K. Kubitzki

Flowering Plants  Aquifoliales, Boraginales, Bruniales, XIV , Escalloniales, Garryales, Paracryphiales, (except Convolvulaceae), , , Vahliaceae Volume Editors: Joachim W. Kadereit  Volker Bittrich

With 76 Figures Editors Joachim W. Kadereit Volker Bittrich Johannes Gutenberg Campinas Universit€at Mainz Brazil Mainz Germany Series Editor

Prof. Dr. Klaus Kubitzki Universit€at Hamburg Biozentrum Klein-Flottbek und Botanischer Garten 22609 Hamburg Germany

ISBN 978-3-319-28532-0 ISBN 978-3-319-28534-4 (eBook) DOI 10.1007/978-3-319-28534-4

Library of Congress Control Number: 2016937409

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Printed on acid-free paper

This Springer imprint is published by Springer Nature The registered company is Springer International Publishing AG Switzerland Preface

It is with great pleasure that I present volume XIV of the Families and Genera of Vascular Plants, a further step forward in the treatment of asterid (sympetalous) families of the eudicots. Recent advances in molecular systematics have necessitated an extensive rearrangement of the families, including the lumping, splitting or revival of well-known family concepts or, in some cases, the creation of completely new ones. To cite one example, the position of the Aquifoliaceae has long been unclear—in previous classifications the family has peregrinated through at least eight orders, whereas now it is placed in a new order at the base of the , the Aquifoliales, which also comprise a few other presumably related families. The Icacinaceae, which always have been a problematic group for taxonomists, can now be presented in a substantially revised circumscription, although their closest relatives still remain unknown. It is worth remembering that early naturalists, who based their classifica- tions on visible traits and geographical patterns, achieved classifications that have proven useful for integrating findings from both non-molecular and molecular data. Thus, the five families of the Dipsacales as treated in this volume are today confirmed as a monophylum—already in the 19th century, they had been recognised as a natural group but, despite much effort, their interrelationships still are not understood in all details. It is with pride that we can include in this volume the monumental treatments of the two very large families Solanaceae and Boraginaceae, resulting from year- (if not life-) long involvement of their two main authors accompanied by numerous co- workers dealing with specific aspects, so that their treatments rest on broad factual bases. These include fully revised classifications of the two families as well as a wealth of biological data. Boraginaceae, which are predominantly herbaceous, comprise a wide range of growth forms such as rhizomatous herbs and other geophytes, and their flowers, fruits and seeds are diverse in terms of size, structure and functional adapta- tions. Solanaceae exploit habitats worldwide from coastal areas to high-montane regions, preferably in the tropics. Thus, their considerable variation of life-forms and reproductive structures is no surprise; almost every known pollination syndrome has been uncovered in this family and, in addition to nectar and pollen, pollinator attractants include curious substances such as liquid perfume and oil. I am stressing all these biological traits included into the family treatments in this series from the very beginning, following the conviction that they are important for an understanding of the life history of plants. Very often, groups of closely related species differ in a single trait, their theme of diversification, while other traits may remain unchanged. Understanding adaptive variation may help understand life history evolution. I would like to acknowledge the hard work and commitment of all authors and the volume editors; I am deeply indebted for their scholarly contributions and careful

v vi Preface editorial work. I am also grateful to the copyright holders of the illustrations published under their responsibility. The artist Bobbi Angell, New York, is thanked for the generosity with which she authorized the use of her artwork. Finally, I have great pleasure in thanking the copy editor of the present volume, Dr. Monique Delafontaine, for her dedicated editorial work. I also wish to acknowledge the very pleasant collaboration with Dr. Andrea Schlitzberger from Springer Verlag and with Mr. V. Anand from Spi Technologies India Pvt Limited.

Hamburg Klaus Kubitzki April 2016 List of Contributors

M.C.E. Amaral Depto. de Biologia Vegetal, Instituto de Biologia, Universidade Estadual de Campinas, Campinas, SP, Brazil. [email protected] S. Andrews Kew Gardens, Royal Botanic Gardens, Kew, Richmond, Surrey, UK. [email protected] A. Anton Instituto Multidisciplinario de Biologı´a Vegetal, Co´rdoba, Argentina. [email protected] S. Arroyo Botanischer Garten und Botanisches Museum Berlin-Dahlem, Freie Universit€at Berlin, Berlin, Germany. [email protected] A. Backlund Division of Pharmacognosy, Dept. of Medicinal Chemistry, Uppsala University, BMC - Biomedical Center, Uppsala, Sweden. [email protected] G.E. Barboza Instituto Multidisciplinario de Biologı´a Vegetal, Co´rdoba, Argentina. [email protected] G. Barriera Conservatoire et Jardin botaniques de la Ville de Gene`ve, Chambe´sy/Gene`ve, Suisse. [email protected] G. Bernardello Instituto Multidisciplinario de Biologı´a Vegetal, Co´rdoba, Argentina. [email protected] V. Bittrich Rua Dr. Ma´rio de Nucci 500, 13083-290 Campinas, SP, Brazil. [email protected] C. Carrizo Garcia Instituto Multidisciplinario de Biologı´a Vegetal, Co´rdoba, Argentina. [email protected] R. Claßen-Bockhoff Institut fur€ Spezielle Botanik, Johannes Gutenberg- Universit€at Mainz, Mainz, Germany. [email protected] A.A. Cocucci Instituto Multidisciplinario de Biologı´a Vegetal, Co´rdoba, Argentina. [email protected]

vii viii List of Contributors

J.G. Conran ACEBB & SGC, School of Earth and Environmental Sciences, The University of Adelaide, North Terrace, SA, Australia. [email protected] M.T. Cosa Instituto Multidisciplinario de Biologı´a Vegetal, Co´rdoba, Argentina. [email protected] L.P. Ronse De Craene Royal Botanic Garden Edinburgh, Edinburgh, UK. [email protected] N. Diane Milowstraße 10, 14195 Berlin, Germany. [email protected] W.C. Dickison (deceased) Formerly at University of North Carolina, Chapel Hill, NC, USA. M.O. Dillon Botany Department, The Field Museum, Chicago, IL, USA. mdillon@fieldmusem.org R. Duno Centro de Investigacio´n Cientı´fica de Yucata´n, Merida, Yucatan, Mexico. [email protected] H. Fo€rther Heilmannstr. 5, D-82049 Pullach, Germany. V. Fuentes (deceased) Formerly at Instituto de Investigaciones de Fruticultura Tropical, La Habana, Cuba. M. Gottschling Department Biologie I, Ludwig-Maximilians-Universit€at Munchen,€ Munchen,€ Germany. [email protected] H.H. Hilger Systematische Botanik und Pflanzengeographie, Institut fur€ Biologie, Freie Universit€at Berlin, Berlin, Germany. [email protected] M. Hofmann Formerly at Systematische Botanik und Pflanzengeographie, Institut fur€ Biologie, Freie Universit€at Berlin, Berlin, Germany. U. Hofmann (deceased) Formerly at Systematic Geobotanical Institute, Georg August Universit€at Go€ttingen, 37073 Go€ttingen, Germany. A.T. Hunziker (deceased) Formerly at Instituto Multidisciplinario de Biologı´a Vegetal, Co´rdoba, Argentina. J.W. Kadereit Institut fur€ Spezielle Botanik und Botanischer Garten, Johannes Gutenberg-Universit€at Mainz, Mainz, Germany. [email protected] T.G. Lammers Department of Biology and Microbiology, University of Wisconsin Oshkosh, Oshkosh, WI, USA. [email protected] A. Liston Department of Botany & Plant Pathology, Oregon State University, Corvallis, OR, USA. [email protected] P.-A. Loizeau Conservatoire et Jardin botaniques de la Ville de Gene`ve, Chambe´sy/Gene`ve, Suisse. [email protected] F. Luebert Nees-Institut fur€ Biodiversit€at der Pflanzen, Rheinische Friedrich-Wilhelms-Universit€at, Bonn, Germany. fl[email protected] List of Contributors ix

J. Lundberg Department of Botany, The Swedish Museum of Natural History, Stockholm, Sweden. [email protected] T.D. Macfarlane Western Australian Herbarium, Science and Conservation Division, Department of Parks and Wildlife, Manjimup, WA, Australia. [email protected] V. Mayer Department of Botany and Biodiversity Research, Faculty of Life Sciences, University of Vienna, Vienna, Austria. [email protected] A.E. Moscone Instituto Multidisciplinario de Biologı´a Vegetal, Co´rdoba, Argentina. M.J. Potgieter Department of Biodiversity, University of Limpopo, Polokwane, South Africa. [email protected] A. Romanutti Instituto Multidisciplinario de Biologı´a Vegetal, Co´rdoba, Argentina. [email protected] V. Savolainen Imperial College London, Ascot, Berkshire, UK. [email protected] M. Schori Department of Environmental & Plant Biology, Ohio University, Athens, OH, USA. [email protected] F. Selvi Dipartimento di Scienze delle Produzioni Agroalimentari e dell’Ambiente (DISPAA), Sez. Scienza del Suolo e della Pianta, Laboratori di Botanica, Universita` di Firenze, Firenze, Italy. federico.selvi@unifi.it R. Spichiger Conservatoire et Jardin botaniques de la Ville de Gene`ve, Chambe´sy/Gene`ve, Suisse. [email protected] R. Subils Instituto Multidisciplinario de Biologı´a Vegetal, Co´rdoba, Argentina. M. Thiv Staatliches Museum fur€ Naturkunde Stuttgart, Stuttgart, Germany. [email protected] D.C. Thomas School of Biological Sciences, The University of Hong Kong, Hong Kong, China. [email protected] T.M.A. Utteridge Herbarium, Library, Art and Archives, Royal Botanic Gardens, Kew, Kew, Surrey, UK. [email protected] G.K. Walden Department of Integrative Biology, University of California, Berkeley, CA, USA. [email protected] F. Weberling (deceased) Formerly at Abteilung f. Spezielle Botanik, Universit€at Ulm, Ulm, Germany. x List of Contributors

M. Weigend Nees-Institut fur€ Biodiversit€at der Pflanzen, Rheinische Friedrich-Wilhelms-Universit€at, Bonn, Germany. [email protected] Q.Y. Xiang Department of Plant and Microbial Biology, North Carolina State University, Raleigh, NC, USA. [email protected] L.-B. Zhang Science & Conservation Division, Missouri Botanical Garden, St. Louis, MO, USA. [email protected] Contents

Introduction to the Orders and Families V. B ITTRICH and J.W. KADEREIT ...... 1 of Uncertain Placement of this Volume Adoxaceae A. BACKLUND and V. BITTRICH ...... 19 Aquifoliaceae P.-A. LOIZEAU,V.SAVOLAINEN, S. ANDREWS,G.BARRIERA, and R. SPICHIGER ...... 31

Aucubaceae Q.-Y. XIANG ...... 37 Boraginaceae M. WEIGEND,F.SELVI, D.C. THOMAS, and H.H. HILGER ...... 41

Bruniaceae R. CLAßEN-BOCKHOFF ...... 103 (with Zabelia incert. sed.) U. HOFMANN and V. BITTRICH ...... 117 M. SCHORI ...... 131 Codonaceae M. WEIGEND and H.H. HILGER ...... 137 Columelliaceae A. BACKLUND ...... 141 (inclusive Triplostegia) V. M AYER ...... 145 Ehretiaceae M. GOTTSCHLING,M.WEIGEND, and H.H. HILGER ...... 165

Eremosynaceae J.G. CONRAN and T.D. MACFARLANE ... 179 Escalloniaceae J. LUNDBERG ...... 185 Eucommiaceae L.-B. ZHANG ...... 193 Garryaceae A. LISTON ...... 197 Heliotropiaceae N. DIANE, H.H. HILGER,H.Fo€RTHER, M. WEIGEND, and F. LUEBERT ...... 203

Helwingiaceae Q.-Y. XIANG ...... 213 Hydroleaceae V. B ITTRICH and M.C.E. AMARAL ... 217 Hydrophyllaceae M. HOFMANN, G.K. WALDEN, H.H. HILGER, and M. WEIGEND ..... 221

Icacinaceae M.J. POTGIETER and R. DUNO ...... 239 Lennoaceae V. B ITTRICH ...... 257

xi xii Contents

Metteniusaceae (incl. Dendrobangia W.C. DICKISON and V. BITTRICH .... 263 incert. sed.) Montiniaceae LOUIS P. R ONSE DE CRAENE ...... 269 Morinaceae U. HOFMANN and V. BITTRICH ...... 275 Paracryphiaceae W.C. DICKISON and J. LUNDBERG ... 281 Phyllonomaceae V. B ITTRICH ...... 287 Polyosmaceae J. LUNDBERG ...... 291 Solanaceae G.E. BARBOZA, A.T. HUNZIKER, G. BERNARDELLO, A.A. COCUCCI, A.E. MOSCONE,C.CARRIZO GARCÍA, V. F UENTES, M.O. DILLON,V.BITTRICH, M.T. COSA,R.SUBILS,A.ROMANUTTI, S. ARROYO, and A. ANTON ...... 295

Sphenocleaceae T.G. LAMMERS ...... 359 Sphenostemonaceae M. THIV ...... 363 Stemonuraceae M.J. POTGIETER,M.SCHORI, and T.M.A. UTTERIDGE ...... 367

Tribelaceae A. BACKLUND ...... 377 Vahliaceae M. THIV ...... 381 Valerianaceae F. WEBERLING and V. BITTRICH ...... 385 Wellstediaceae H.H. HILGER and M. WEIGEND ...... 403 Index ...... 407