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INFORMATION TO USERS This manuscript has been reproduced from the microfilm master. U M I films the text directly from the original or copy submitted. Thus, some thesis and dissertation copies are in typewriter face, while others may be from any type of computer printer. The quality of this reproduction is dependent upon the quality of the copy submitted. Broken or indistinct print, colored or poor quality illustrations and photographs, print bleedthrough, substandard margins, and improper alignment can adversely afreet reproduction. In the unlikely event that the author did not send UM I a complete manuscript and there are missing pages, these will be noted. Also, if unauthorized copyright material had to be removed, a note will indicate the deletion. Oversize materials (e.g., maps, drawings, charts) are reproduced by sectioning the original, beginning at the upper left-hand corner and continuing from left to right in equal sections with small overlaps. Each original is also photographed in one exposure and is included in reduced form at the back of the book. Photographs included in the original manuscript have been reproduced xerographically in this copy. Higher quality6 " x 9" black and white photographic prints are available for any photographs or illustrations appearing in this copy for an additional charge. Contact U M I directly to order. University Microfilms International A Bell & Howell Information Company 300 North Zeeb Road. Ann Arbor. Ml 48106-1346 USA 313/761-4700 800/521-0600 Order Number 0816213 The application of fossil diatom assemblages in reconstructing mid to late Holocene climate in the neotropics: The paleolimnology of Huarmicocha and Lake Ayauch1, Ecuador Riedinger, Melanie Ann, Ph.D. The Ohio State University, 1993 Copyright ©1903 by Riedinger, Melanie Ann. AH rights reserved. UMI 300 N. Zeeb Rd. Ann Arbor, MI 48106 THE APPLICATION OF FOSSIL DIATOM ASSEMBLAGES IN RECONSTRUCTING MID TO LATE HOLOCENE CLIMATE IN THE NEOTROPICS : THE PALEOLIMNOLOGY OF HUARMICOCHA AND LAKE AYAUCHI, ECUADOR DISSERTATION Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate School of The Ohio State University By Melanie A. Riedinger, B.S., M.S. ***** The Ohio State University 1993 Dissertation Committee: Approved by D.A. Culver R.A. Stein R.E.J Boemer Adviser M. Steinitz-Kannan Department of Zoology Copyright by Melanie Ann Riedinger 1993 Dedicated to Virginia Mary Zeis Riedinger ACKNOWLEDG EMENTS There are many people who have contributed their time and encouragement to this work, and for their support and efforts I am very grateful. I am indebted to Dave Culver, Roy Stein, Miriam Steinitz-Kannan, Ralph Boerner and Tom Hetherington for their assistance in the last phases of this process, for without their help I would not have completed. I would like to thank Miriam Steinitz-Kannan for stimulating my interest in diatom ecology, for her advice, enthusiasm and encouragement throughout all phases of this research, and for graciously welcoming me into her lab. I also would like to thank Paul Colinvaux, my major professor throughout most of this project, for his support, patience and advice. This work has benefited from many stimulating discussions with Paulo de Oliveira on diatom ecology and paleoecology and I am grateful for both his friendship and his interest in this research. I look forward to many years of collaboration with him in the future. , Mike Miller, Mark Bush and Mark Brenner provided many important insights into my data, and I thank them for their time. Ramahurthi Kannan read and re-read many versions of this draft, and I thank him for his iii comments and observations. Throughout this project I have been fortunate to have had the opportunity to work with many bright and promising scientists, who kindly shared their knowledge, provided polite criticism when required, and offered me their friendship. My thanks and appreciation go to Ann Anton, Becky Bixby, Wendy Eisner, David Harwood, Eric Hugo, Vickie Hugo, Jim Lyons-Weiler, Vicki Martin-Kier, Mark Nienaber, Brian Reeder, Fausto Sarmiento, Mark Scheutzow, Mike Stewart, Carol Stevens, Tim Sharon, and Mike Ware. I thank Tom Whitmore for his friendship and continued efforts to help me maintain my sanity through this process. I thank my mother, Virginia Riedinger, and my brothers Michael and Joseph Riedinger for their support, encouragement, love and patience for so many years, and my nieces Lori and Erin for understanding my absence from their lives. For welcoming me into their country and helping me complete my field work, I thank Elena Sarmiento, Eduardo and Ana Christina Asanza, Trudi and Hans Steinitz, Ana Steinitz, Maria Elena Hervas, Montserrat Rios, Doris Ortiz, and the Museo Ecuatoriano de Ciencias Naturales. I also am grateful to John Thieret, Tom Rambo, Elinor Rambo, Jerry Carpenter, Uewellya Hillis, Charlie Reimer, Bettie Palmison, Mike Riley, Paul Thiel, and Sue Longenbacker for their continued interest in my progress. This work would not have been possible without the cooperation of the Ecuadorian government which granted me permits to study within their country. The Department of Zoology at The Ohio State University and the Department of Biological Sciences at Northern Kentucky University provided me with funding, equipment and laboratory space to complete my research, as well as many hours of stimulating conversation. This research was supported by grant # DEB-8202658 from the National Science Foundation to Paul A. Colinvaux, and grant #ATM- 8713211 from the National Science Foundation to Paul A. Colinvaux and Miriam Steinitz-Kannan. Portions of this project were supported by a Sigma Xi Grant-In-Aid of Research Award and a Tinker Foundation Travel Grant awarded to the author. v VITA July 15,1960 .....................................Born - Ft. Thomas, Kentucky 1983 ........................................................ B.S., Northern Kentucky University, Highland Heights, Kentucky 1983-1989 ................................................. Graduate Research Associate and Teaching Associate, Department of Zoology, The Ohio State University, Columbus, Ohio 1986 ........................................................ M.S. Department of Zoology, The Ohio State University, Columbus, Ohio 1989-1991 .............................................. Instructor and Research Associate, Department of Biological Sciences, Northern Kentucky University, Highland Heights, Kentucky 1991-Present ............................................ Visiting Assistant Professor, Department of Biological Sciences, Northern Kentucky University, Highland Heights, Kentucky PUBLICATIONS Steinitz-Kannan, M., M. Nienaber, M. Riedinger, and R. Kannan. The fossil diatoms of Lake Yambo, Ecuador. A 2500 year record of El Nifto events. Bull. Institut Frangais d"Etudes Andines (In Press). vi Colinvaux, P.A., M.B. Bush, K.-b. Liu, P.E. De Oliveira, M. Steinitz-Kannan, M. Riedinger, and M. Miller. 1989. Amazon without refugia: vegetation and climate of the Amazon basin through a glacial cycle. Int. Symp. on Global changes in South America during the Quaternary: Past- Present- Future. Sao Paulo (Brazil). Steinitz-Kannan, M., M. Riedinger and M. Miller. 1987. Lasdiatomeas del Ecuador: II. Diatomeas de la laguna de Limoncocha, Provincia de Napo. Publicaciones del Museo Ecuatoriano de Ciencias Naturales. 5(7):51-77. Steinitz-Kannan, M., M. Nienaber, M. Riedinger, L. Harrel, and M. Miller. 1982. Estudios limnologicos en la Laguna de San Marcos con descripciones de las especies principales de diatomeas. Publicationes del Museo Ecuatoriano de Ciencias Naturales. 3(3): 39-65. FIELDS OF STUDY Major Field: Zoology Studies in Tropical Paleolimnology, Paleoecology, Community Ecology TABLE OF CONTENTS DEDICATION ii ACKNOWLEDGEMENTS iii VITA vi LIST OF TABLES xii LIST OF FIGURES xv LIST OF PLATES xxiv CHAPTER PAGE I. INTRODUCTION 1 FOCUS OF THE STUDY 5 Objectives of this study and chapter sub-division 7 Limitations in paleoecological investigation 11 II. A LATE HOLOCENE RECORD FROM THE ECUADORIAN ANDES 14 INTRODUCTION 14 STUDY SITE 16 METHODS 27 RESULTS AND DISCUSSION 34 Sediment Stratigraphy of Huarmicocha 34 LOI Data 38 The Diatom Percentage Diagram 41 viii Species Richness in Huarmicocha ................. 49 Paleoecological Implications from the Huarmicocha re c o rd .................... .59 Late Holocene Climate in E cuador ................62 III. MID-HOLOCENE LAKE-LEVEL FLUCTUATIONS: A FOSSIL DIATOM RECORD FROM ECUADORIAN AMAZONIA ......................................................................................70 INTRODUCTION....................................................70 STUDY SITE .... .7 4 METHODS...............................................................81 RESULTS AND DISCUSSION.............................. 87 Sediment Stratigraphy of AyaucW ..................87 LOI Data for Ayauchi.................................... 93 Fossil Pigment and Chemistry Data for Ayauchi ...............................................99 Fossil Diatom P ro file .................................... 106 Species Richness in the Ayauchi Record . .,121 The Paleolimnological Record of Ayauchi. ., .125 Paleoecological implications of the Ayauchi diatom re c o rd .................................. .129 IV. CONTEMPORARY AND HISTORICAL RECORDS OF MORPHOLOGICAL VARIATION IN TROPICAL DIATOM POPULATIONS.............................................................. 136 INTRODUCTION.................................................... 136 METHODS...............................................................