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A Molecular Taxonomic Treatment of the Neotropical Genera

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An Intrageneric and Intraspecific Study of Morphological and Genetic Variation in the Neotropical Compsoneura and Virola (Myristicaceae) by Royce Allan David Steeves A Thesis Presented to The University of Guelph In partial fulfillment of requirements for the degree of Doctor of Philosophy in Botany Guelph, Ontario, Canada © Royce Steeves, August, 2011 ABSTRACT AN INTRAGENERIC AND INTRASPECIFIC STUDY OF MORPHOLOGICAL AND GENETIC VARIATION IN THE NEOTROPICAL COMPSONEURA AND VIROLA (MYRISTICACEAE) Royce Allan David Steeves Advisor: University of Guelph, 2011 Dr. Steven G. Newmaster The Myristicaceae, or nutmeg family, consists of 21 genera and about 500 species of dioecious canopy to sub canopy trees that are distributed worldwide in tropical rainforests. The Myristicaceae are of considerable ecological and ethnobotanical significance as they are important food for many animals and are harvested by humans for timber, spices, dart/arrow poison, medicine, and a hallucinogenic snuff employed in medico-religious ceremonies. Despite the importance of the Myristicaceae throughout the wet tropics, our taxonomic knowledge of these trees is primarily based on the last revision of the five neotropical genera completed in 1937. The objective of this thesis was to perform a molecular and morphological study of the neotropical genera Compsoneura and Virola. To this end, I generated phylogenetic hypotheses, surveyed morphological and genetic diversity of focal species, and tested the ability of DNA barcodes to distinguish species of wild nutmegs. Morphological and molecular analyses of Compsoneura. indicate a deep divergence between two monophyletic clades corresponding to informal sections Hadrocarpa and Compsoneura. Although 23 loci were tested for DNA variability, only the trnH-psbA intergenic spacer contained enough variation to delimit 11 of 13 species sequenced. A morphological and molecular investigation of Compsoneura capitellata showed little discrete morphological variation among populations but significant genetic structure among populations. Phylogenetic analysis of Virola also revealed a deep molecular divergence between two clades having numerous contrasting morphologies. In contrast to Compsoneura, the trnH-psbA intergenic spacer failed to differentiate the majority of Virola species tested. An infraspecific morphological and molecular study of V. sebifera and V. loretensis showed that each of these species contains morphologically and ecologically discrete sympatric morphotypes that likely represent new species. In total, this investigation found 5 provisional new species from fewer than 600 collections at biological stations in Ecuador and Peru where these new species were among the most abundant trees in the forest. This suggests that much diversity likely remains to be described in the Myristicaceae and other tropical plant families. iv ACKNOWLEDGEMENTS This thesis wouldn’t have been possible without the help of many people. Firstly, I’d like to thank my supervisor, Steve Newmaster, for taking me on 5 years ago as a new masters student, pardon the pun. Your patience and infectious botanical curiosity has been an inspiration to me. Aron Fazekas, thanks for your unrelenting patience with me in the lab and for always having time for those “quick” questions that turn into a lenghty discussion. As for jungle-man John Janovec, thanks for all the great times in the field, your help in getting permits, access to BRIT’s plant and human resources and our late- night Compsoneura campfire discussions. I’m also very grateful to all the members of my committee Dr’s Rob Hanner, Usher Posluszny, and David Galbraith for your valuable input and guidance throughout these years. Carol-Anne Lacroix and Ragupathy Sabrumanyam; thanks for sharing your taxonomic and ethnobotanical expertise over the years. I’d also like to thank the following labmates for their input, thoughtfull discussions, help in procrastinating, and most importantly their friendship: Heather Cole, Jillian Bainard, John Gerrath, Troy Mcmullin, Kevin Berg, Kevin Burgess, Jose Maloles, Evan Lusty and Brian Lacey as well as all the other folks that have worked in the herbarium over the years. An extra thanks to Jose and Brian for their help with measuring specimens and preparing figures. This work couldn’t have been completed without all the dedicated field botanists that I’ve had the pleasure of working with. Derek Piper and Andrea Reid: thanks for your patience with me in the tropics. I know I’m not the easiest person to get along with in the tropical heat while climbing a tree and trying to get the pruning poles together. I also got v to work with some great parataxonomists in my 4 months in the field: Danilo Simba Larco (Ecaudor), Wilson Quizhpe (Ecuador), Angel Balarezo (Peru-BRIT), Pedro Centeno (Peru-BRIT), Benjamin Chambi (Peru-BRIT). Thank you all for the many laughs in the field, your unspeakable tree-climbing/pruning abilities and all your patience with a neotropcial neophytologist like myself. Funding for my research and travel were provided in part byby NSERC, Amazon Conservation Association, and the Latournelle travel grant. David Neil, Renan Valega, Keri McNew, and Tiana Franklin made this research possible by coordinating collection and export permits. I am deeply indebted to innumerable people that have helped me troubleshoot DNA extraction, PCR, sequencing and other molecular methods over the past few years; namely I’d like to thank the Husband and Crease labs for sharing equipment and Jeff and Angela at the Genomics Facility for their expertise. I’d also like to take the opportunity to my fellow graduate students for their companionship and numerous beer-fueled discussions at the grad lounge. To my girlfriend friend Sarah Baldwin: thank for your love and patience through this whole process, thanks for our long-winded plant genetic discussions and thanks for being you. Last but not least I’d like to thank my parents, Bob and Diane, for their unrelenting love and support throughout my schooling. Our innumerable forays into the woods and streams of New Brunswick to go hunting, fishing, trapping, berry and fiddlehead picking have spawned a curiosity in nature that I hope will only grow stronger. This thesis is dedicated to you. vi Table of Contents ACKNOWLEDGEMENTS ............................................................................................ iv LIST OF TABLES ........................................................................................................... ix LIST OF FIGURES .......................................................................................................... x GENERAL INTRODUCTION........................................................................................ 1 Taxonomy ...................................................................................................................... 1 Ethnotaxonomy .......................................................................................................... 1 DNA Barcoding.......................................................................................................... 4 Phenetic versus Cladistic Analyses in Systematics................................................... 6 Human and geologic history of Central and South America.................................... 9 Myristicaceae............................................................................................................... 12 General Morphology and Cytology ......................................................................... 12 Ecology ..................................................................................................................... 14 Ethnobotany ............................................................................................................. 15 Myristicaceae Systematics........................................................................................ 17 Biogeography of the Myristicaceae ......................................................................... 19 Challenges to Molecular work with the Myristicaceae........................................... 20 Why work with the Myristicaceae?.......................................................................... 21 Chapter 1 ......................................................................................................................... 33 A PHYLOGENETIC AND MORPHOLIGICAL ANALYSIS OF COMPSONEURA ........................................................................................................................................... 33 Abstract........................................................................................................................ 33 Introduction................................................................................................................. 34 Materials and Methods............................................................................................... 39 Taxon sampling and outgroup selection ............................................................... 39 Results .......................................................................................................................... 44 DNA sequencing and alignment ............................................................................ 44 Phylogenetic Analyses............................................................................................. 45 Morphology ............................................................................................................
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