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Grasses S YSTEMATICS AND E VOLUTION S W. L. J J E ( ) URREY ACOBS AND OY VERETT EDITORS National Library of Australia Cataloguing-in-Publication entry International Symposium on Grass Systematics and Evolution (3rd : 1998). Grasses : systematics and evolution. Bibliography. Includes index. ISBN 0 643 06438 9 (hardback). ISBN 0 643 06393 5 (set). ISBN 0 643 09010 X (eBook). 1. Grasses – Congresses. I. Jacobs, S.W.L. (Surrey W. L.) II. Everett, Joy. III. Title. 584.9 © CSIRO 2000 This book is available from: CSIRO PUBLISHING PO Box 1139 (150 Oxford Street) Collingwood VIC 3066 Australia Tel: (03) 9662 7666 Int: +(613) 9662 7666 Fax: (03) 9662 7555 Int: +(613) 9662 7555 Email: [email protected] http://www.publish.csiro.au Every effort has been made to trace the original source of copyright material contained in this book. On behalf of the authors, the publisher would be grateful to hear from any copyright owners it has not been able to contact and apologises for any errors or omissions. Printed in Australia by Brown Prior Anderson C ONTENTS GENERAL A Phylogeny of the Grass Family (Poaceae), as Inferred from Eight Character Sets 3 The Grass Phylogeny Working Group (GPWG) A Systematic View of the Development of Vascular Systems in Culms and Inflorescences of Grasses 8 Thompson Demetrio Pizzolato The Grass Inflorescence 29 C. Vegetti and Ana M. Anton BAMBOOS Phylogeny and Classification of the Bambusoideae (Poaceae) 35 Weiping Zhang and Lynn G. Clark Woody Bamboos (Gramineae-Bambusoideae) of Madagascar 43 Soejatmi Dransfield Preliminary Studies on Taxonomy and Biosystematics of the AA Genome Oryza Species (Poaceae) 51 Bao-Rong Lu, Ma. Elizabeth B. Naredo, Amita B. Juliano, and Michael T. Jackson POOIDS Phylogenetic Structure in Poaceae Subfamily Pooideae as Inferred from Molecular and Morphological Characters: Misclassification versus Reticulation 61 Robert J. Soreng, and Jerrold I. Davis Relationships within the Stipoid Grasses (Gramineae) 75 S.W.L. Jacobs, Joy Everett, Mary E. Barkworth and Cathy Hsiao Preliminary Views on the Tribe Meliceae (Gramineae: Pooideae) 83 Teresa Mejia-Saulés and Frank A. Bisby The Species of Bromus (Poaceae: Bromeae) in South America 89 Ana M. Planchuelo and Paul M. Peterson Phylogenetic Analysis of the Triticeae Using the Starch Synthase Gene, and a Preliminary Analysis of some North American Elymus species 102 Roberta J. Mason-Gamer and Elizabeth A. Kellogg Changing Perceptions of the Triticeae 110 Mary E. Barkworth III PANICOIDS A Cladistic Analysis of the Paniceae: a Preliminary Approach 123 Fernando O. Zuloaga, Osvaldo Morrone and Liliana M. Giussani Phylogeny of the Subfamily Panicoideae with Emphasis on the Tribe Paniceae: Evidence from the trnL-F cpDNA Region 136 Rosalba Gómez-Martínez and Alastair Culham Austral South American Species of Eriochloa 141 Mirta O. Arriaga Molecular and Morphological Evolution in the Andropogoneae 149 Elizabeth A. Kellogg The Relation of Space and Geography to Cladogenic Events in Agenium and Homozeugos (Poaceae: Andropogoneae) in South America and Africa 159 Gerald F. Guala Andropogoneae Systematics and Generic Limits in Sorghum 167 Russell E. Spangler CHLORIDOIDS Phylogenetic Relationships in Subfamily Chloridoideae (Poaceae) Based on matK Sequences: A Preliminary Assessment 173 K. W. Hilu and L. A. Alice On the Classification of the Chloridoideae: Results from Morphological and Leaf Anatomical Data Analyses 180 An Van den Borre and Leslie Watson Phylogenetic Relationships of the Genus Sporobolus (Poaceae: Eragrostideae) Based on Nuclear Ribosomal DNA ITS Sequences 184 Juan-Javier Ortiz-Diaz and Alastair Culham Phylogenetics of Bouteloua and Relatives (Gramineae: Chloridoideae): Cladistic Parsimony Analysis of Internal Transcribed Spacer (nrDNA) and trnL-F (cpDNA) Sequences 189 J. Travis Columbus, Michael S. Kinney, Maria Elena Siqueiros Delgado, and J. Mark Porter Systematics of the Muhlenbergiinae (Chloridoideae: Eragrostideae) 195 Paul M. Peterson A Phylogeny of Triodieae (Poaceae: Chloridoideae) based on the ITS Region of nrDNA: Testing Conflict between Anatomical and Inflorescence Characters 213 J.G. Mant, R.J. Bayer, M.D. Crisp, and J.W.H. Trueman ARUNDINOIDS The Danthonieae: Generic Composition and Relationships 221 Nigel P. Barker, Cynthia M. Morton, and H. Peter Linder Biogeography of the Danthonieae 231 H.P. Linder and N.P. Barker IV PHYSIOLOGY/ECOLOGY Contributions of Prolamin Size Diversity and Structure to the Systematics of the Poaceae 241 K.W. Hilu Ecological Significance of South-West African Grass Leaf Phytoliths: A Climatic Response of Vegetation Biomes to Modern Aridification Trends 248 Christian Mulder and Roger P. Ellis Effects of Elevated Atmospheric [CO ] in Panicum Species of Different 2 Photosynthetic Modes (Poaceae: Panicoideae) 259 Claudia Tipping and David R. Murray Ecophysiological Investigations of the Distribution of Poaceae and Restionaceae in the Cape Floristic Region, South Africa 267 T. L. Bell, W. D. Stock and H. P. Linder BREEDING SYSTEMS Adaptive Plasticity in Reproduction and Reproductive Systems of Grasses 281 James A. Quinn Dioecism in Grasses in Argentina 287 H.E. Connor, Ana M. Anton and Marta E. Astegiano Apomixis and Amphimixis Comparative Biogeography: a Study in Poa (Poaceae) 294 Robert J. Soreng Comparative Reproductive Biology of the Vulnerable and Common Grasses in Bothriochloa and Dichanthium 307 Ping Yu, N. Prakash and R. D. B. Whalley Apomixis in Monocotyledons 316 R. Czapik BIOGEOGRAPHY Chorology of Grasses – a Review 325 T.A. Cope Grasses in North America: A Geographic Perspective 331 Mary E. Barkworth and Kathleen M. Capels Tropical Grasslands and Savannas 351 Robert B. Shaw Temperate Grasslands of the Southern Hemisphere 356 R.H. Groves Future of Temperate Natural Grasslands in the Northern Hemisphere 361 Arthur W. Bailey Endemic Grass Genera of Maharashtra State, Peninsular India 369 C.B. Salunkhe INDEX V This page intentionally left blank P REFACE The grass family is the single most important family of organisms The Third International Symposium on Grass Systematics and for the survival of mankind. Of the top ten crops we use to sus- Evolution was held in Sydney in the week beginning 28th Sep- tain ourselves (directly or indirectly as stock feed), the first ten tember 1998, concurrently with the Second International Con- are all grasses. An understanding of the relationships and charac- ference on Monocot Systematics. This was the first time that the teristics of grasses has played an important role so far in our his- two areas of Grass and Monocot research had joined together. So tory, and the importance of that role can only increase as we many of the methods and philosophies are now universally struggle to balance exploitation, sustainability and conservation. adopted that it seemed ideal that researchers had the opportunity to attend a wide range of presentations. With increasing interest There is some discussion as to when the ‘first’ international in evolutionary studies, few families can be studied in isolation. meeting on the systematics and evolution of the grasses was held. To this end the first session of each day’s grass program was held A meeting was held in 1959 as a special session of the Ninth jointly with the general monocots session, with the latter part of International Botanical Congress in Montreal. It took 27 years the day devoted to grasses. before the next International Symposium occurred at the Smith- sonian Institution in 1986 and, while the Symposium was adver- The grasses are a family with confusing relationships, with differ- tised as ‘The First …’ the publication did not include that word. ent data sets suggesting different arrangements. The develop- In the period between the Montreal and Smithsonian meetings, ment of techniques to obtain and analyse molecular data has work on molecular evolution, physiology and numerical meth- meant the dawning of a new age in the study of the grasses. It seems we have been given a key to a whole new suite of rooms ods had revolutionised many areas of grass research. The Smith- previously hidden behind the dark curtains of hybridisation, sonian meeting was an inspiration to the 150 participants from polyploidy and reticulate evolution. Numerical and DNA analy- around the world, and the published proceedings, including ses are developing to a point where they are contributing to the many review papers, took its place as a new benchmark and interpretation of relationships in the cloudiest of tribes. Even major reference for further work in all of the tribes. And so it though we have these new data, the role of more traditional seemed essential to hold the next meeting on the grasses without information on breeding systems, genetic control, developmental such a long gap. There was a further little-publicised Conference studies and biogeography is becoming increasingly important as held at the Main Botanical Garden, Moscow in 1994 but inter- we aim to assemble corroborative and/or multi-sourced data national attendance at this conference was very restricted. matrices. The development of statistical methods for testing It was 13 years after the Smithsonian Symposium before the Syd- molecular hypotheses still has some way to go, and corroboration ney commitment to hold the 2nd International Monocot Sym- with independent data sets still has an important role to play. posium coincided with a request from some of the organisers of There were 42 lectures on grass systematics and evolution, 34 of the Smithsonian Symposium for us to host the next Interna- which are published here, along with two others presented in the tional Grass Symposium. It quickly became clear that the most ‘general’ sessions of Monocots II, and the grass presentation on efficient way of doing this, from both a pragmatic and scientific composition of cell walls was combined to make a single presen- point of view, was to hold them jointly. This presented great tation in the first volume. Of the 75 posters at Monocots II, pre- opportunities to form new research associations and generally senters of six of the many high quality contributions presenting have a good time with others interested in similar research sub- research on grasses were invited to elaborate and present them as jects. If all goes well the next grass symposium should obey the papers to be included in this volume. All manuscripts submitted inverse exponential rule and occur approximately 6 years from were refereed. Monocots II, such is the rate of research on this important eco- nomic group. Joy Everett and Surrey Jacobs VII This page intentionally left blank ORGANISING COMMITTEE OF MONOCOTS II ACKNOWLEDGEMENTS Royal Botanic Gardens Sydney: Barbara Briggs, Joy Everett, Our thanks go to our sponsors, whose assistance made the con- Gwen Harden, Alistair Hay, Surrey Jacobs, Peter Weston, Karen ference feasible: Wilson • University of New South Wales University of New England: Jeremy Bruhl • Royal Botanic Gardens Sydney University of New South Wales: Christopher Quinn University of Sydney: Murray Henwood • Orlando Wyndham Wines University of Technology, Sydney: David Morrison • Australian Systematic Botany Society • Roads and Traffic Authority of New South Wales • Swane’s Garden Centre • Ansett Australia Airlines Australian Systematic Botany Society IX