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University Microfilms, a XEROX Company , Ann Arbor, Michigan REVISIONARY and POPULATION STUDIES in the AMERICAN

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University Microfilms, a XEROX Company , Ann Arbor, Michigan REVISIONARY and POPULATION STUDIES in the AMERICAN 71-27,513 LOWDEN, Richard Max, 1943- REV I SI ONARY AND POPULATION STUDIES IN THE AMERICAN AQUATIC PLANT GENUS PONTEDERIA L. The Ohio State University, Ph.D., 1971 Botany University Microfilms, A XEROX Company , Ann Arbor, Michigan REVISIONARY AND POPULATION STUDIES IN THE AMERICAN AQUATIC PLANT GENUS PONTEDERIA L- DISSERTATION Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate School of The Ohio State University By Richard Max Lowden* B,A., M.Sc. The Ohio State University 1971 Approved by Adviser Graduate Program in Botany To my Parents for their continued interest in Education ACKNOWLEDGMENTS I have sincerest admiration for my industrious assistant, Rafaela Joaquin, my wife, for the countless hours she rendered to this research in the field and laboratory. The receipt of a Grant-In-Aid of Research from The Society of the Sigma Xi is acknowledged, which en­ abled me, while carrying out other research, to study populations in British Honduras during December 1969. I am most indebted to the Organization for Tropical Studies, Inc. (OTS) for an introduction to tropical Biology (Costa Rica, summer of 1968) and for granting an OTS Pilot Research Grant F 70-23 (The Ohio State University Research Foundation Project No. 3031-Al) to R. L. Stuckey (Faculty Adviser, Department of Botany, The Ohio State University) and myself (Principal Investigator) for aid in performing the necessary field research in Mexico and Central America during the summer of 1970. To the Graduate Committee of the Department of Botany and the Graduate School Fellowship Committee (The Ohio State University), I am most grateful for the University Dissertation Year Fellowship awarded to me for the completion of this investigation. Appreciation is expressed to Curators of the following herbaria for specimens loaned: Brooklyn Botanic Garden (BKL), Brooklyn, New York National Museum of Canada, National Herbarium (CAN), Ottawa, Ontario; Duke University Herbarium (DUKE), Durham, North Carolina; Field Museum of Natural History, Department of Botany (F), Chicago, Illinois; The Florida State University Herbarium (FSU), Tallahassee, Florida; Arnold Arboretum and Gray Herbarium of Harvard University (A, GH), Cambridge, Massachusetts; The University of Michigan Herbarium (MICH), Ann Arbor, Michigan; New England Botanical Club Herbarium (NEBC), Cambridge, Massachusetts; Missouri Botanical Garden (MO), St. Louis, Missouri; New York Botanical Garden (NY), Bronx, New York; The Ohio State University Herbarium (OS), Columbus and The Franz Theodore Stone Laboratory Herbarium (FTSL), Put-In-Bay, Ohio; The Academy of Natural Sciences of Philadelphia (PH), Philadelphia, Pennsylvania; Smithsonian Institution, Department of Botany (US), Washington, D. C.: and University of Wisconsin Herbarium (WIS), Madison, Wisconsin. Voucher specimens for investigations in Pontederia have been deposited in The Ohio State University Herbarium, Columbus, Ohio. Constructive criticism of the manuscript by J. A. Schmitt, E. D. Rudolph, R. L. Stuckey, and T. F. Stuessy (Dissertation Reading Committee, Department of Botany, The Ohio State University) is grate­ fully appreciated. Thanks are extended to R. M. Giesy, T. J* Johnson (Department of Botany, The Ohio State University) and D* W. Stevenson (Graduate Program in Botany, The Ohio State University), for assistance in photography of plates. iii VITA September 27, 1943 . Born, Columbus, Ohio June 1964 ....... B.A., Botany Department, The Ohio State University, Columbus, Ohio June 1964-June 1966 . Peace Corps Volunteer, United States Peace Corps, taught Biology and Mathematics at Austin High School, Stann Creek Town, British Honduras, Central America December 1967 ..... M.Sc., thesis entitled "A Vascular Flora of Winous Point, Ottawa and Sandusky Counties, Ohio,’' Botany Program, The Ohio State Uni­ versity, Columbus, Ohio September 1966-June 1968 Research Assistant to Professor T. R. Fisher, National Science Foundation Grant, The Ohio State University, Columbus, Ohio July-August 1968 .... Student in the course, Tropical Biology--An Ecological Approach, sponsored by the Organization for Tropical Studies (OTS) in Costa Rica, Central America September 1968-June 1970 Teaching Assistantshlp, Graduate Program in Botany, The Ohio State University, taught Biology 100 during Autumn quarter 1968 and Winter quarter 1969; Botany 410 during Spring and Summer quarters 1969; and Botany 102 during Spring quarter 1970 January 1970 ..... Awarded a Grant-In-Aid of Research from The Society of The Sigma Xi for study of "William A. Schipp's Botanical Explorations in the Stann Creek and Toledo Districts, British Honduras 1929-1935" May 1970 . ..A Pilot Study research grant was awarded by The Organization for Tropical Studies to Associate Professor Ronald L. Stuckey (Faculty Adviser to Mr. Lowden, Academic Faculty of Botany, The Ohio State University) and Mr. Richard M. Lowden (Principal Investigator) iv aiding field research carried out in Central America for this Dissertation June 27, 1970 Married Rafaela Andrea Joaquin of the Dominican Republic June 1970-June 1971 • . Awarded a University Dissertation Fellowship, Graduate School, The Ohio State University, Columbus, Ohio 4 PUBLICATIONS Lowden, Richard M. 1969. A Vascular Flora of Winous Point, Ottawa and Sandusky Counties, Ohio. Ohio J. Sci. 69 (5):257-284. ________ . 1970. William Ashbrook Kellerman’s Botanical Expeditions to Guatemala (1905-1908). Taxon 19 (l):l9-35. 1970. William A. Schipp's Botanical Explorations in the Stann Creek and Toledo Districts, British Honduras (1929-1935), Taxon 19 (6):831-861. FIELDS OF STUDY Major Field: Botany Studies in Vascular Plant Taxonomy (Plant Systematics). Associate Professor Ronald L. Stuckey Studies of Botanical Collectors and Explorers in Central America. Professor Emanuel D. Rudolph and Associate Professor Ronald L. Stuckey v CONTENTS Page ACKNOWLEDGMENTS............................................. 11 VITA ........................................................ lv TABLES ........................................... vll FIGURES ...................................................... vlil PLATES .......................................................xii PONTEDERIA L. THE GENERIC NAME!: NOMENCLATURE SINCE 1737 .. 1 REVISION OF PONTEDERIA L.: TAXONOMY ......................... 5 Disposition of Excluded Names ............... 197 A GENERIC BASIS: CYTOLOGY .................................... 199 GEOGRAPHICAL DISTRIBUTION OF PONTEDERIA L.: THE AMERICAS . 204 A TRISTYLOUS GENUS WITH AN ISOLATED FLORAL FORM: POPULATION REPRODUCTIVE BIOLOGY .......................................... 211 STATISTICAL ANALYSIS OF PHENOLIC QUALITATIVE FEATURES: CHROMATOGRAPHIC D A T A ........................................ 228 B10SYSTEMATIC ASPECTS: EVOLUTIONARY IMPLICATIONS ............. 245 APPENDIX (TABLE 7 ) .......................................... 251 LITERATURE CITED .............................................. 286 vl TABLES Table Page 1 Representative length measurements of perigones, stamens and styles in Pontederia L. ....... 195 2 Seed germinations in growth chamber conditions, with and without hardened perigone bases ....... 210 3 Indication of floral form frequency in Pontederia L. based on population and herbarium studies . • 222 4 Spot colors of phenolic compounds in Pontederia L. and related genera ................ 237 5 Summary of spot occurrence of phenolic compounds in Pontederia L. and related genera (Table 4) • . ■ . 238 6 A comparison of two dimensional paper chromatographic profiles of Pontederia L. and related genera based on summary of phenolic spot occurrence (Table 7) , 240 7 Spot occurrence of phenolic compounds in Pontederia L. and related genera as exemplified by population study and herbarium specimens examined ...... 251 vii FIGURES Figure Page 1 Numerous blue-purple flowered inflorescence) clasping subtending spathe and flowering stalk leaf of Pontederia cordata L. var. cordata, U.S., Ohio, Ottawa County, V/inous Point"! 3% miles southwest of Port Clinton, off Route 53 at the western end of the Sandusky Bay ................ 22 2 Pontederia cordata L. var. cordata in marsh habitat, U.S., Ohio, Ottawa County, Winous Point, 3% miles southwest of Port Clinton, off Route 53 at the western end of the Sandusky Bay 22 3 Fruiting specimen of Pontederia cordata L. var. cordata, showing cordate flowering stalk leaves and clasping sheath at base of flowering stalk stem shoot, U.S., Ohio, Ottawa County, Winous Point, Lowden 25 (OS) ............... 23 A Fruiting and flowering specimen of Pontederia cordata L. var. cordata, showing sagittate-hastate leaves, U.S., Ohio, Ottawa County, Winous Point, Lowden 25 (OS) .................................................... 25 5 Geographical distribution of Pontederia cordata L. var. cordata in Ontario, Quebec, New Brunswick, Prince Edward Island and Nova Scotia Provinces of Canada . 27 6 Geographical distribution of Pontederia cordata L. var. cordata (dots), var. lancifolia (Muhl.) Torr. (triangles), and var. ovalis (Hart.) Solms (stars) in the United States 29 7 Numerous white flowered inflorescence, clasping sub­ tending spathe and flowering stalk leaf of Pontederia cordata L. var. cordata in British Honduras, C.A., 9 miles from Belize City, along the Northern Highway .................. 31 Pontederia cordata 1.. var. cordata in drainage ditch habitat, British Honduras, C.A., 9 miles from Belize City, along the Northern Highway ........... 31 vili FIGURES (continued) Figure Page 9 Provisional
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