PHYLOC;ENETIC ANALY SIS OF BREEDING-SYSTEM EVOLUTION IN EETEFtOSTYLOUS MONOCOTYLEDONS SEAN W. GRAHAM A THESIS SUBMTED IN CONFORMITY WlTH REQUTREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY IN THE UNZ\rERSm OF TORONTO National Library Bibliothëque nationale I*m of Canada du Canada Acquisitions and Acquisitions et Bibliographie Services services bibliographiques 395 Wellington Street 395, me Wellington Ottawa ON K 1A ON4 OttawaON K1AON4 Canada Canada Your hie Votre relenmce our nie Naire reterence The author has granted a non- L'auteur a accordé une licence non exclusive licence allowing the exclusive permettant à la National Library of Canada to Bibliothèque nationale du Canada de reproduce, loan, distribute or sel1 reproduire, prêter, distribuer ou copies of this thesis in microfonn, vendre des copies de cette thèse sous paper or electronic formats. la forme de microfiche/nlm, de reproduction sur papier ou sur format électronique. The author retains ownership of the L'auteur conserve la propriété du copyright in this thesis. Neither the droit d'auteur qui protège cette thèse. thesis nor substantid extracts from it Ni la thèse ni des extraits substantiels may be printed or otherwise de celle-ci ne doivent être imprimés reproduced without the author's ou autrement reproduits sans son permission. autorisation. Canada PI-IYLOGE~JE~ICANALYSIS OF BREEDING-SYSTEMEVOL~ON IN HE?EROS?rtOUS MONOCOTYLEDONS Sean W. Graham, Department of Botany, University of Toronto The evolutionary histories of various stylar conditions (heierostyly, enantiostyly, stylar dirnorphism and monomorphism) were examined in two groups of rnonocotyledons, Pontederiaceae and Narcissus, using several soures of phylogenetic information and severai different weighting schemes for reconstructions of c haracter evolution. The monophyly of Pontederiaceae is strongly supported by phylogenetic evidence from rbcL, but its local position in the monocotyledons is unclear. Several tests were used to assas the congruence of four phylogenetic data sets for the farnily: two chloroplast genes (ndhF and rbcL);a restriction-site survey of the plastome; and a revised morphology-based data set. Despite different "rules" of evolution for the three chloroplast sources of data, these data sets provided highly congruent. well resolved and weU supported phylogenetic estimates of the fàmily's history. nie morphological data provided a poorly supported phylognetic estimate that showed both congruence and incongruence with the chloroplast evidence. A cornbined analysis of ndhF and rbcL genes from taxa in five superorders of monocotyledons added support to the view that three families containhg enantiostylous taxa, Commelinaceae, Haemodoraceae and Philydraceae. are the nearest Living relatives of Pontederiaceae. Several lines of evidence indicate that despite the evolutionary distance between Pontedenaceae and these families, they contain usefui historicai information conceming the position of the family's root Trees derived €rom the combined chloroplast evidence were used to reconstruct breeding-system evolution in Pontederiaceae. Modentely weighted optimization schemes indicate that tristyly arose once only in the family. while predominantiy-seIfmg, florally- monomorphic Lineages of Eichhomia arose on multiple occasions. Self-incompatibility arose after the ongin of the floral trimorphisrn, a scenario incompatible with some evolutionary models. Enantiostyly arose twice, with its occurrence in Heterunthera (sensu lato) possibly homologous wi th that in related families. Reconstructions of breeding-system evolution in Narcissur are im peded b y the partial lack of resolution of an n&F-based tree of the genus and the probable role played by undetected hybridization events in the genus' phylogenetic history. A preliminary reconstruction of breeding-system evolution in Narcissus suggests that stigma-height dimorphism may have arisen on multiple occasions, and that tristyly in Narcissur triandrus may have evolved diredy from floral monomorphism. It is not very easy to sum up in a few short sentences the debt 1 owe to so many for making the last seven years a frequently wonderfd the, and for making the les-than-wonderful times more than bearable. .. To Spencer, for taking me on and providing a wonderful problem and professional training, for misting me enough to do this my own way, and for frequent jolts of psychic energy. Without Brian Morton this whole thing may not have finally got OHthe grand. Joshua Kohn shed in the prolonged birthing agony of the Evoiution paper that 1 forced myself ont0 so naïvely, frorn which 1 leamed so much about the art of phylogenetics and the science of diplomacy. 1have ken exceptionally lucky in the lab cornpanions 1 have had over the years - Elizabeth Bush, Domenica Manicacci, Majid Ghassemian, Taline Sarkissian, Holly Trewhitt, Mara Kerry, Brendon Larson, LinIey Jesson, Peter Toppings, Brian Husband, Chris Eckert, Pam Diggle and Mitch Cruzan. In particuiar, Fanny Stxumas, Bill Cole, Anne Worïey, Angela Baker, Andrea Case, Brian Morton, and John Panneil provided friendship and support well above and beyond die cdof duty. David McKnight provided crucial encouragement early in my acadernic training and Richard Abbott more recently. Inside and outside the depanment, Tim Gray, Connie Soros, Sue and Emma Dexter, Kristjan Vitols, Rodger Evans. Petra Comelly, Zum Gadallah, Roberta Fulthorpe, Mara Kerry, Luba Skambara, Steven Schultz, Ben Teh, John Maxwell, Sandra hnde,Kate Frego, Rolfe Vinebrooke, Christine Karnpny, Fereshteh Hashemi, Usha Goel, Val& Zobens, Nick Provart, Elizabeth Tillier, Bob Latta, Rebecca Dotterer, Esther Levesque, Brenda Kosmer and James Scott supplied endess ta&, food. hin, and countless hours of procrastination. 1 rhank Tamw Mamourian and Valerie Anderson for their caring professionaüsrn, and Fernanda and the night- tirne crew for keeping me Company into the wee houn. Various faculty members provided support, help and advice dong the way, but Jim Eckenwalder, Jim Anderson, Linda Kohn, Tim Dickinson, Tarnmy Sage and Nancy Dengler deserve particular thanks for ths. Dick Olrnstead and Mike Donoghue provided invaluable suppon and mentorship from afar. Thahto Ken Sytsma and bdaKohn for working so hard as my external and interna1 examiners. Mike Clegg and Jeff Palmer lundly pennitted me to use their labs at criticai points early on in my degree. Jennifer Richards, Mike Simpson and Charles Hom supplied usehl advice on several papers. Many people wittingly or unwittingly supplied me with data and plant maienal over the years, but Linda Prince, John Blanchard, Mike Simpson. Tom Givnish, Alan Meemw and Mark Chase deseme particular thanks for going out of their way to do so. Many individuals and institutions provided assistance on a less personal basis, but not less valuable for that. David Swofford, Wayne and David Maddison and Joe Felsenstein wrote the software withwt which my data would be a meaningless collection of letters and numbers. Gerard Zurawski and the DNAX Institute kindiy provided the rbcL primers that allowed me to start. The taxpayers of Canada and Ontario providecl the financial support that aîiowed me to do my work The Connaught commitîee provided the fmcial support that aiiowed me to corne to Canada CUEW 2 and the University of Toronto provided exceIlent reasons for hypothemiia on mid-Febmary picket hes. 1 thank Alan Cross and the staff of CFNY for helping to keep me sane in an otherwîse IoneIy molecular lab. In Angela Baker, Jacquie Be&, Tracy Solomon and Andrea Case 1 have made wondefil fnends. Thanks for ail the 1ark.s and japes. I thank Christina Heidom and Sue Dexter for "bean"great fnends, for support in times of ned, and for endless hours of hilarity. 1 thank Wendy Untereiner for her friendship, for aii the chocolate, and for helping to make it go. I thank Lisa Law for her love and friendship, Rmnnhh. 1 thank Philip Aw&a for his friendshp, for his inteilectual companionship, for sharing hiç epic love of life and most especially for laughing at my jokes. 1 thank Catherine Glass and Catriona Gordon for king the most complete and utter pair of bints 1 am ever likely to know . Thank you for showing me how . 1 thank Cary List and Peter Au for their fkiendship and for king the ones. 1thank my mum and dad, Joan and Jimmy Graham, and my sibs, Paula, Martyn and Peter, for their love and unwavering support across the great divide. This thesis is dedicated to Elsie and Eric McDuff, and to my gmdparents, Wannah and Sammy Nelson. ".. ..w hat ya thinking? What am 1 singing? A Song of seeds, The food of love, Eat the music!" Frorn "The Red Shoes," Kate Bush, 1993. CONTENTS Page LIST OF TABLES xiv LIST OF FIGURES xvi Mo&ls for the evolutionary origin and breakhm of heterostyly Objectives of the thesis ÇHAFTER 2: PHYLOGENEIIC SYSTEMATXCS OF PONTEDERIACEAE: IMPLICATIONS FOR BREEDING-SYSTEM EVOLUTlON 21 INTRODUCTION 22 Sup rafamilia1 Systernatics 23 A. Morplwlogical evidence 23 B. Molecular evidence 27 lnrrafamilial Systernatics 34 A. Morpho bg icai evidence 34 B. Molecuiar evidence 37 Breeding-System Evolution in Pontederiaceae 39 A. Origin and evofutionof the tristylous syndrome 39 B. Eflects of the seljing syndrome on phylogenetic reconstruction 49 C. Evolution and ahptive signiftcance of enmriostyly 5 1 CONCLUSION 58 CH- PHYLOGENETIC CONGRUENCE AND DISCORDANCE AMONG ONE MORPHOLOGICAL AND THREE MOLECULAR DATA SECS FROM PONTEDERLACEALE INTRODUCTION MATERIALS AND METHODS Analyses Measwes of tree resolrrtion and support Memures of characrer incongrme