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Evolutionary Biology of the Parasitic Angiosperm Arceut.Hobzem Amerjcanum (Viscaceae) As Determined by Population Genetic Anacysis and Infectivi'iy Experiments

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Evolutionary Biology of the Parasitic Angiosperm Arceut.Hobzem Amerjcanum (Viscaceae) As Determined by Population Genetic Anacysis and Infectivi'iy Experiments EVOLUTIONARYBIOLOGY OF THE PARASITIC~JGIOSPERM ARCEUTHOBIUMAME'G~NUM (VISCACEAE:) AS DETERMINED BY POPULATIONGENETIC ANALYSIS AND INFECTIVITYEXPERMENTS Cheryl AM Jerome Department of Botany A thesis submitted to the Faculty of Graduate Studies in partial fùlfillrnent of the requirements for the degree of Doctor of Philosophy at the University of Manitoba National Library ûiilioîhèque nationale I+B of,,, du Canada Acquisitions and Acquisitions et Bibliographic Services services bibliographiques 395 Welington Street 395. rue Weiiinglon Ottawa ON KIA ON4 OttawaON KlAW Carada CaMda The author has granteci 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 sell reproduire, prêter, distribuer ou copies of this thesis in microform, vendre des copies de cette thèse sous paper or electronic formats. la fonne de microfiche/fïh, 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 substantial extracts ffom 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. THE UNIVERSITY OF MANITOBA COPYRIGHT PERMISSION EVOLUTIONARY BIOLOGY OF THE PARASITIC ANGIOSPERM ARCEUT.HOBZEM AMERJCANUM (VISCACEAE) AS DETERMINED BY POPULATION GENETIC ANACYSIS AND INFECTIVI'IY EXPERIMENTS BY CEERYL ANN JEROME A Thesis/Practicum submitted to the Faculty of Graduate Studies of The University of Manitoba in partial fulfillment of the requirement of the degree of DOCTOR OF PHaOSOPHY CHERYL ANN JEROME O 2001 Permission bas been gmnted to the Library of the University of Manitoba to lend or sel1 copies of this thesis/practicum, to the National Library of Canada to microfilm this thesis and to lend or sel1 copies of the film, and to University Microfilms Inc. to publish an abstract of this tbesis/practicum. This reproduction or copy of tbis tbesis bas been made available by authority of the copyright owner solely for the purpose of private study and research, and may only be reproduced and copied as permitted by copyright laws or with express written authorization from the copyright owner. TABLE OF CONTENTS ... ABSTRACT ....................................................................................... ~ii ACKNO WLEDGEMENTS. ....................................................................... -v .. LIST OF TABLES ..................................................................................vu LIST OF FIGURES ...... .............................................................................xi CHAPTERS 1. INTRODUCTION. LITERATUREREVIE w AND THESISGOALS.. ....................-1 1.1 Introduction.. ................................................................... 1 * 1.2 Literature Review .................... ... .................................... 3 1.3 Thesis Goals .................................................................. -19 2 . AFLP MARKERANALYSIS REVEALS EVIDENCEFOR THREEGENETIC RACES OF THE PARASITICANGIOSPERM. ARCEUTHOBIUAIAMERICANUM .................. -22 2.1 Introduction.................................................................... 22 2.2 Materials and Methods ..................................................... -26 2.3 Results........................................................................... 32 2.4 Discussion...................................................................... 37 2.5 Conclusions.................................................................... 53 3 . COMPARATIVEPOPULATION STRUCTURE AND GENETICDIVERSLN OF . A RCEUTHOBIUMAMERICANUM AND 17s PINUSSPP . HOSTS........................... 71 3 .1 Introduction ................................................................... 71 3.2 Materials and Methods ............................... ......... ............ -74 3.3 Results ......................................................................... 81 3.4 Discussion ............................ .. .................................... A37 3 -5 ConcIusions ................................................................... 98 4 . INFECTIVITY OF THE PARASITIC PLANT. ARCEUTHOBIUMAMERICANUM ON PINUS BANKSIANA. PINUSCONTORTA. AND THEIR HYBRlDS ......................... 115 4.1 Introduction ................................................................. 115 4.2 Materials and Methods ....................................................119 4.3 Results ....................................................................... 125 4.4 Discussion .................................................................. -130 4.5 Conclusions .................... ... ................................... -142 LITERATURE CITED............................................................................. 168 APPENDICES ..................................................................................... -184 ABSTRACT In this study, a multidisciplinary approach incorporating population genetic analysis and infectivity experiments was used to explore the evolutionary biology of Arceuthobium americanum Nutt. ex Engelm. (Viscaceae). This study represents one of the few comprehensive investigations into the evolutionary, geographical, biological, ecologicd, and historical factors influencing a parasitic pIant. Arcezithobium americanum infects three principal hosts and has the most extensive geographic range of any N. Amencan dwarf mistletoe. Based on the lack of apparent morphological and phenologicai dserences between populations of A. americanum, pst researchers have found no evidence for recognizing subspecific taxa. In the present study, molecular analysis using amplified hgment length polymorphism analysis (AFLP) indicated that A. americanum is divided into three distinct genetic races, each associated with a different host taxon in regions of allopatry: (1) Pinus banksiana in - western Canada; (2) Pinus contortu var. murrayanu in the Sierra Nevada and Cascade Mountain ranges in the westem U.S.A.; and (3) Pinus conrorta var. Zatifotia in the westem U.S.A. and Canada. These observations suggested that host identity, geographic isolation, and environmental factors have aü contributed to race formation in A. americanum. Molecular analysis using AFLP indicated that hosts are divided into only two genetic groups: (1) P. banksiana and hybrids; and (2) P. conforta var. ZatifoIia and var. murrayana. This observation suggests that host identity is not the primary factor leading to race formation in A. americanum. Findings fiom infection experiments also question the role played by host identity since there is no indication of host X parasite interactions. Nonetheless, the role of host cmot be completeIy ded out since the genetic races of A. americanum can clearly be defined by this parameter, The lack of fine-scde patterning within A- americanum races was attributed to random dispersal of seeds over long distances by animal vectors, as well as to adaptation of parasite populations to non-geographically patterned host genotypes and local environmental conditions. Historical factors such as glaciations and founder events were also found to impact structuring and genetic diversity in A. arnericanum populations. Despite this lack of fine-scale patterning, the existence of three distinct genetic races of A. americanum provides insight into the evolutionary potential of this taxon. Given sufficient tirne, it is possible that these races will become reproductively isolated, and undergo speciation. ACKNOWLEDGEMENTS There are many people who have contributed to this thesis in distinct ways and 1 would like to take this opportunity to express my appreciation towards them- Firstly, to my major professor, Dr. Bruce Ford (Caricologist), 1 owe many thanks. 1 would like to start by thanking you for accepting me as a Ph.D. student in your lab, despite my king a zoologist by training, Your open-mindedness has allowed me to expand my horizons and comprehend the wonder and beauty of the plant kingdom. 1 appreciate al1 that you have taught me about botany, systematics, and scientific research. Ir has been both an honour and a pleasure to train under your guidance. 1 hope that we will maintain a lifelong professional and personal relationship. To my supewisory committee, Drs, Greg Pemer, Dave Punter, Simon Shamoun and Ray Smith, thank you for your contributions to the development and completion of this research project. In particular, 1 must extend my utrnost gratitude to Dr. Greg Penner (Monsanto) for your enthusiasm and for allowing me to cary out much of my molecular research in your lab. 1 would also like to express my gratitude to Dr. Dave Punter (Chair, Botany Department) for introducing me to the intriguing world of parasitic angiosperms. Thank you also for joining Bruce and me on the field trip to the western U.S.A. in 1999. It was a great expenence that 1 will always remember -- an opportunity to see a diversity of parasitic and non-parasitic plants, as well as nurnerous interesting landscapes including Craters of the Moon (Idaho), the Beartooth Pass (Montana), Zion National Park (Utah), and Crater Lake (Oregon). Thank you also to Dr. Daryl Somee (Agiculture Canada) for allowing me to run ULPgels in your lab and to Dr. Kermit Ritland WC)for statistical advice. Further thanks to several other professors ors. N.
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