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Phylogenetic and Biogeographic Relationships of Cheirodendron

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Phylogenetic and Biogeographic Relationships of Cheirodendron PHYLOGENETIC AND BIOGEOGRAPHIC RELATIONSHIPS OF CHEIRODENDRON NUTT. EX. SEEMAN (ARALIACEAE) A THESIS SUBMITTED TO THE GRADUATE DIVISON OF THE UNIVERSITY OF HAWAIʻI AT MĀNOA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE IN BOTANY AUGUST 2014 By Chelsea Osaki Thesis Committee: Clifford Morden, Chairperson Donald Drake David Lorence ACKNOWLEDGEMENTS I would like to acknowledge the following people whom I am grateful for in helping me to complete this project: Dr. Clifford Morden for his patience and kindness with the project and throughout my time as a graduate student; Dr. Donald Drake for his insightful comments on my thesis; and Dr. David Lorence for his knowledge of the Marquesas Islands. I would also like to thank Joel Lau, whose ideas about the evolution and speciation of Cheirodendron have served as the foundations of this project. To Mitsuko Yorkston, who has taught me every lab technique I know. Thank you to Dr. Anthony Mitchell for his comments and suggestions for primers and Dr. Ines Schönberger from Allan Herbarium (CHR) for providing outgroup samples. There were many people who have been involved in field work, collecting specimen, sorting herbarium samples, performing DNA extractions, PCRs, and analyses, whose help I greatly appreciate: Dr. Gregory Plunkett, Dr. Pei-Luen Lu, Dr. Richard Pender, Nipuni Sirimalwatta, Adam Williams, Seana Walsh, Dylan Morden, Wendy Kishida, Steve Perlman, Dr. Timothy Gallaher, Vianca Cao, Jesse Adams, Peter Wiggin, Bao Ying Chen, April Cascasan, Dylan Davis, Jacy Miyaki, Gavin Osaki, Erin Fujimoto, George Akau, Matthew Campbell, Isaiah Smith, Dr. Daniel Rubinoff, Dr. Michael Thomas, and Robert Tamayo. Finally, I would like to thank the Botany Department at the University of Hawaiʻi at Mānoa for providing me with opportunities to do my research; National Tropical Botanical Garden for excellent hospitality during my stay on Kauaʻi; and finally, the Kōkeʻe Resource Conservation Program for providing me with the opportunity to learn about conservation, native Hawaiian plants and the beautiful island of Kauaʻi. ii TABLE OF CONTENTS ACKNOWLEDGEMENTS ...................................................................................................... ii LIST OF TABLES .................................................................................................................... v LIST OF FIGURES ................................................................................................................. vi CHAPTER 1. LITERATURE REVIEW AND THESIS PROPOSAL ................................... 1 INTRODUCTION ................................................................................................................. 1 Family Araliaceae ............................................................................................................. 2 Taxonomic history of the genus Cheirodendron................................................................ 3 Traditional uses ................................................................................................................. 5 PROPOSED RESEARCH AND HYPOTHESES ................................................................ 6 MATERIALS AND METHODS .......................................................................................... 7 Taxon sampling, outgroup sampling and DNA extraction ................................................ 7 DNA sequencing and analysis ........................................................................................... 8 Preliminary data ................................................................................................................ 9 Future directions ............................................................................................................... 9 CHAPTER 2. PHYLOGENETIC AND BIOGEOGRPAHIC RELATIONSHIPS OF CHEIRODENDRON NUTT. EX. SEEM. (ARALIACEAE) ................................................. 13 ABSTRACT ........................................................................................................................ 13 INTRODUCTION ............................................................................................................... 14 MATERIALS AND METHODS ........................................................................................ 19 iii Sampling and DNA extraction ......................................................................................... 19 DNA amplification and DNA sequencing ........................................................................ 20 Primer screening and analysis ........................................................................................ 23 RESULTS............................................................................................................................ 27 DISCUSSION ..................................................................................................................... 33 Relationships among taxa ................................................................................................ 34 Current taxonomy vs. phylogenetic relationships ........................................................... 37 Biogeographic relationships............................................................................................ 39 Future directions ............................................................................................................. 40 CHAPTER 3. SYNTHESIS- HYPOTHESES REVISITED ................................................. 42 LITERATURE CITED ........................................................................................................... 44 iv LIST OF TABLES Table Page 1.1 Comparison of revisions of Cheirodendron………………………………………...10-11 2.1 Voucher information and locality of specimen used in this study………………….21-22 2.2 List of primers and references………………………………………………………24-26 v LIST OF FIGURES Figure Page 1.1 Infrafamilial phylogenetic relationships of Araliaceae………………………………12 2.1 ETS phylogeny of Cheirodendron…………………………………………………...29 2.2 ITS phylogeny of Cheirodendron………………………………………………….…30 2.3 ndhF-rpl32 phylogeny of Cheirodendron…………………………………………....31 2.4 Combined phylogeny comparing MP, ML, and BI methods………………………..32 vi CHAPTER 1. LITERATURE REVIEW AND THESIS PROPOSAL INTRODUCTION Understanding infrageneneric relationships are crucial in providing a basis in analysis for other studies such as biogeography, ecology, macroevolution, systematics, and conservation biology (Sites & Marshall 2004). One way to determine these relationships is through phylogenetics. Understanding phylogeny, or how species are related to one another, provides a comprehensive interpretation of evolutionary processes, including speciation (Harrison 1998). The study of molecular phylogenetics and phylogenetic trees has provided a direct record of the speciation events that have led to the extant species we see today (Barraclough & Nee 2001). Phylogenetic analysis has been a useful tool in elucidating the microevolutionary and macroevolutionary relationships in a number of lineages, including the family Araliaceae. Progress has been made in resolving the placement of Araliaceae within the order Apiales (Henwood & Hart 2001, Plunkett & Lowry 2001, Plunkett et al. 2004) as well as understanding relationships within and among closely related genera (e.g. Wen & Zimmer 1996, Mitchell & Wagstaff 1997, Costello & Motley 2001, Eibl, Plunkett & Lowry 2001). Although studies have been done on some of the more horticulturally (Hedera L., ivy) or ethnobotanically (Panax L., ginseng) important genera, little attention has been given to other genera within Araliaceae, particularly Cheirodendron Nutt. ex Seem. The phylogenetic relationships among species of Cheirodendron are currently unknown. To date, only a few taxonomical studies have assessed species relationships. However, speciation is not always accompanied by clear morphological differentiation 1 (Kenfack 2011). The importance of phylogenetics in discovering species relationships of Cheirodendron will sort out the taxonomy of the genus. My proposed research aims to uncover species relationships by sequencing various regions of both chloroplast and nuclear DNA. Using a molecular approach, genetic differences will be assessed, rather than morphological characters which can be influenced by the environment and highly plastic. Family Araliaceae The Araliaceae consists of about 50 genera (Liu et al. 2012) and 1500 species that are widely distributed in the tropics and subtropics of Asia, the Pacific Islands, and the Neotropics (Wen et al. 2001, Yi et al. 2004). Most members of Araliaceae are woody with variable leaf morphologies (simple, palmately compound and pinnately compound), but maintain conserved floral characteristics (5-merous flowers with inferior ovaries in a compound umbel) (Wen et al. 2001, Yi et al. 2004, Liu et al. 2012). Some well-known species in Araliaceae include medicinal herbs such as Panax ginseng C. A. Meyer (ginseng), and ornamentals including Hedera helix L. (English ivy) and Schefflera actinophylla (Endl.) Harms (umbrella tree) (Liu et al. 2012). Molecular studies of Araliaceae agree on a phylogeny with multiple polytomies and poorly resolved basal lineages (Figure 1.1) (Mitchell & Wagstaff 1997, Plunkett, Wen & Lowry 2004). Poorly resolved genera that are basal in Araliaceae include Schefflera J. R. Forster & G. Forster, Cheirodendron Nutt. ex. Seem., Raukaua Seem., Cussonia Thunb., Osmoxylon Miq., and Hydrocotyle L. (Plunkett, Wen
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