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A Systematic Revision of Gomphandra (Stemonuraceae)

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A Systematic Revision of Gomphandra (Stemonuraceae) A dissertation presented to the faculty of the College of Arts and Sciences of Ohio University In partial fulfillment of the requirements for the degree Doctor of Philosophy Melanie Schori June 2010 © 2010 Melanie Schori. All Rights Reserved. 2 This dissertation titled A Systematic Revision of Gomphandra (Stemonuraceae) by MELANIE SCHORI has been approved for the Department of Environmental and Plant Biology and the College of Arts and Sciences by Philip D. Cantino Professor of Environmental and Plant Biology Harvey E. Ballard, Jr. Associate Professor of Environmental and Plant Biology Benjamin M. Ogles Dean, College of Arts and Sciences 3 ABSTRACT SCHORI, MELANIE, Ph.D., June 2010, Plant Biology A Systematic Revision of Gomphandra (Stemonuraceae) (480 pp.) Directors of Dissertation: Philip D. Cantino and Harvey E. Ballard, Jr. The genus Gomphandra Wall. ex Lindl. was studied and revised to more accurately reflect the diversity of the group. Floral morphology and fruit anatomy were compared to other genera of Stemonuraceae to assess reproductive diversity within the family and identify potential synapomorphies. Seven genera of Stemonuraceae possess what appears to be a unique fruit type: a laterally compressed drupe with fleshy appendage that aids in dispersal. Fruit of nine genera were sectioned, and all exhibit sclerified longitudinal bundles in the inner mesocarp; one bundle, which appears bright in cross-section under polarized light, may be a synapomorphy for the family. Seed coats are vascularized with a chalazal/post-chalazal bundle (7 genera) or a pachychalazal bundle (2 genera). Molecular data were sampled from four gene regions (rbcL, trnL intron, trnL-F spacer, matK) in 10 species of three genera (Gomphandra, Grisollea, Stemonurus), for a total of 24 new sequences. Preliminary evidence suggests Grisollea and Codiocarpus (both with a laterally compressed fruit) are more closely related to each other than to Stemonurus or Gomphandra, which have uncompressed fruits. Members of Stemonuraceae may have a molecular synapomorphy in a 100 base pair region of TA repeats in the trnL-F spacer that is apparently not shared with any genera outside the family. Gomphandra has two moderately supported clades, one with the synapomorphy of terminal (vs. axillary) inflorescences. The revision of the genus recognizes 62 taxa: 56 4 species, three subspecies, and three varieties. Diversity is highest in the Philippines (17 taxa), followed by Borneo and Papua New Guinea (10 taxa each), and Sumatra (9 taxa). Twenty-nine new taxa are described, three species are resurrected, one species is reduced in status, and five taxa are synonymized. Problems with typification and nomenclature are corrected for 24 species of Gomphandra and six species in four other genera of Stemonuraceae. Approved: _____________________________________________________________ Philip D. Cantino Professor of Environmental and Plant Biology Approved: _____________________________________________________________ Harvey E. Ballard, Jr. Associate Professor of Environmental and Plant Biology 5 ACKNOWLEDGMENTS This dissertation would not have been possible without the help of many individuals and organizations. I apologize to anyone I may have inadvertently omitted. The librarians at Harvard University Herbaria, especially Judy Warnement, provided invaluable help, as did Peter Fraissinet at Cornell University’s Bailey Hortorium. Curators and staff at 29 herbaria sent loans, checked specimens, hosted me as a visitor, and photographed specimens (A/GH, BH, BISH, BM, BRI, C, CAHUP, CAL, CANB, E, F, G, K, L, LBC, MEL, MICH, MO, MU, NY, P, PNH, PUH, TAIF, TNM, U, UC, US, WRSL). I would especially like to thank Linda Hollenberg (US), Emily Wood and Walter Kittredge (A/GH), Laura Pearce and Timothy Utteridge (K), Gerard Thijsse (L), Jim Solomon (MO), and Peter Lowry (P). Alice Schori databased hundreds of specimens at Harvard for me, showing true maternal devotion. Several people provided translations - Maria Teresa Bonsatti (Italian), Tineke Blokzijl (Dutch), David Tees (French and Japanese), Angelika Stammler (German), and Yunjing Wang, Leonardo Co, and Yi-Ting Wang (Chinese). Neil Bernstein provided assistance with Latin translations and patiently proof-read my Latin descriptions and diagnoses. I received research funding from a variety of sources. Ohio University’s Graduate Student Senate awarded me two original work grants and one travel grant. The American Society of Plant Taxonomists provided support for a research trip to Kew, Leiden, and Paris, and paid for conference attendance at its 2009 meeting in Snowbird, Utah. Ohio University’s Student Enhancement Award provided funds for travel, equipment, and lab expenses. The Ohio Center for Ecology and Evolutionary Studies (OCEES) paid for two quarters of tuition and stipend in support of my research. Ohio University’s Clippinger Fellowship allowed me to concentrate on research without teaching responsibilities for one academic year. A Fulbright grant permitted me to travel to the Philippines for nine months to conduct field research. I would like to thank Elizabeth Clodfelter and the staff at the Philippine-American Educational Foundation for their unwavering support during the Fulbright. I cannot possibly individually thank all the people in the Philippines who helped me, but I can acknowledge those who contributed the most to my success. Mary Ann 6 Cajano was my travel companion during field work, and one could not find a better person with whom to visit remote leech-infested mountains. Julie Barcelona took me under her wing at PNH and helped me connect with other Filipino botanists. Leonardo Co provided amazing botanical assistance. Dr. Edwino Fernando mentored me during my time at the College of Forestry, University of the Philippines – Los Baños. The Quimado family, especially Grace and Marilyn, opened their home to me, as did Rhodora and Nestor Aldemita. Aurora and Gene Hettel and the IT staff at the International Rice Research Institute provided cultural, travel, and technical assistance. The following people and organizations were also invaluably helpful: Dr. Domingo Madulid, Luisto Evangelista (PNH); Dr. Antonio Manila, Anson Tagtag (Department of Environment and Natural Resources); DENR Regional Executive Directors, PENR and CENR officers, and mayors who gave permission to collect and provided collection permits; Rey Luna and Bert Almonte (National Commission on Indigenous Peoples); Andy Pascual, Artaban (Buboy) Famenia, Pompeo Sanchez, and all the other DENR foresters; Nonito Antoque and the other men and women who acted as local guides. Maraming salamat po! Drs. David Tank, Kobinah Abdul-Salim, Vicki Funk, and Félix Forest provided assistance with molecular systematics research. Dr. Wolfgang Stuppy mailed a tungsten carbide microtome blade from the United Kingdom so I could section fruit and then helped me interpret my results. Dr. Timothy Utteridge and Dr. Peter Lowry have served as collaborators on manuscripts and worked with me to make sense of difficult specimens. Drs. James Reveal, Richard Brummitt, and Kanchi Gandhi provided nomenclatural assistance. I need to thank the people at Ohio University who helped me with my dissertation. Dr. Philip Cantino served as my primary advisor, meticulously edited everything I wrote, and supported my efforts tirelessly for six years. Dr. Harvey Ballard coadvised me, provided encouragement when I needed it most, and told me he had no doubt I would finish my dissertation on time. I could not have asked for a better pair of advisors. My other committee members, Drs. Gar Rothwell, Morgan Vis, and Kelly Johnson, have supervised my tenure as a graduate student and read most, if not all, of my dissertation, which is a lot to ask from anyone. Vijay Nadella and Tara Killen at the Ohio 7 University Genomics Facility helped me finally get useable sequence data. Martha Bishop helped me find obscure but essential equipment for my fruit study. Dr. Robert Hikida taught me how to use several kinds of microtomes and allowed me to use his lab. Noorliza Warner produced several hundred annotation slips for me. Dr. Brian McCarthy provided advice on statistical analyses. Connie Pollard found an answer to every administrative question I had, gave me two cats (who provided fur therapy), and was always willing to listen to my frustrations and triumphs, no matter how busy she was (and she’s always busy). Finally, I need to thank Dr. David Tees, who started out translating French for me and ended up as my husband. He called every night while I was in the Philippines, never complained when I worked late, and provided the cheerful loving support every woman should be lucky enough to have from her partner. 8 TABLE OF CONTENTS Page Abstract 3 Acknowledgments 5 List of Tables 10 List of Figures 11 Chapter 1: Introduction 14 Systematics of Gomphandra 14 Stemonuraceae 16 Biogeography 17 Objectives 18 Herbarium and Field Work 18 Taxonomic Concepts 19 Overview of Results 19 Dissemination 20 Chapter 2: Stemonuraceae 21 Introduction 21 Morphology and Anatomy 22 Ecology and Reproductive Biology 30 Economic Importance 32 Relationships among the Genera 32 Family Description 36 Key to the Genera 37 Generic Descriptions 38 Chapter 3: Systematics and Phylogeny of Stemonuraceae 44 Introduction 44 Previous Studies 44 Materials and Methods 46 Results 49 Parsimony Analyses 52 Preliminary Conclusions 55 Chapter 4: Characters, Species Concepts, and Morphometrics 57 Characteristics
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