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Information to Users INFORMATION TO USERS This manuscript has been reproduced fix>m the microfilm master. UMI films the text directly fi'om the original or copy submitted. Thus, some thesis and dissertation copies are in typewriter face, while others may be fi'om any type of computer printer. The quality of this reproduction is dependent upon the quality of the copy subm itted. Broken or indistinct print, colored or poor quality illustrations and photographs, print bleedthrough, substandard margins, and improper alignment can adversely affect reproduction. In the unlikely event that the author did not send UMI 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 comer and continuing fi'om 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 quality 6” x 9” black and white photographic prints are available for any photographs or illustrations spearing in this copy for an additional charge. Contact UM I directly to order. UMI A Bell & Howell Infonnation ConqKtny 300 North Zeeb Road, Ann Arbor MI 48106-1346 USA 313/761-4700 800/521-0600 UNIVERSITY OF OKLAHOMA GRADUATE COLLEGE COMMUNITY ECOLOGY OF WATER-FILLED TREE HOLES IN PANAMA A Dissertation SUBMITTED TO THE GRADUATE FACULTY in partial fulfillment of the requirements for the degree of Doctor of Philosophy By STEPHEN P. YANOVIAK Norman, Oklahoma 1999 ÜMX Number: 9925606 UMI Microform 9925606 Copyright 1999, by UMI Company. All rights reserved. This microform edition is protected against unauthorized copying under Title 17, United States Code. UMI 300 North Zeeb Road Ann Arbor, MI 48103 by STEPHEN P. YANOVIAK 1999 All Rights Reserved. COMMUNITY ECOLOGY OF WATER-FILLED TREE HOLES IN PANAMA A Dissertation APPROVED FOR THE DEPARTMENT OF ZOOLOGY BY Dr. Ola M. Fincke (major professor) Dr. Victor H, Hut . Michael f. John J. SS^arla DrCarj^uVaughn ACKNOWLEDGEMENTS I would not have completed this work without the support of my major professor. Dr. Ola M. Fincke. Ola helped me in more ways than I can list, but I especially thank her for introducing me to the neotropics, for always being forthright and honest, for encouraging me when I needed it most, and for balancing constructive criticism with positive reinforcement. Her thoughtful advice enhanced my personal and professional growth, and will guide me well into the future. I also greatly appreciate the support provided by my committee members, Drs. Victor H. Hutchison, Michael E. Kaspari, Caryn C. Vaughn, and John J. Skvarla. Dr. Hutchison quickly and thoroughly reviewed my work, and offered excellent advice regarding matters of professionalism. Dr. Kaspari improved my understanding of ecological theory, provided funding during my Anal year in Norman, and allowed me to work a flexible schedule. This research required me to leave Norman for many months at a time. I am indebted to Caroline Tawes and Karen Voltura, who forwarded my mail, paid my bills, and handled my academic and personal responsibilities while I was away. Their help and patience substantially reduced the stress of my travels. During my periods of residence in Norman, Curtis Creighton, Michael Fuller and Michael D. Weiser served as sounding boards for research ideas. I appreciate their fliendship, advice, and criticisms very much. In Panama, S. Joseph Wright, Egbert G. Leigh, Jr., Phyllis D. Coley, E. Allan Herre, William Wcislo, Julie Denslow, Annette Aiello, Diane DeSteven, Walter IV Carson, and Wayne Sousa offered a wealth of advice and thoughtful discussions of ideas. L t Col. Nelson Powers and coworkers at Gorgas Laboratory provided reference materials and assistance with mosquito identifications. While observing monkeys for their projects, Christina Campbell and Betsy Mitchell noted the locations of canopy tree holes. Their help saved me a large amount of time and effort, and is greatly appreciated. Christina Campbell, Jack McFarland, Kirsten Montague and Sabine Stuntz cared for my insect cultures and monitored various aspects of ongoing projects during my absences from Panam a. Alejandro Almanza volunteered his time to assist me in the field, helped improve my knowledge of Spanish, and taught me much about Panamanian culture. Gerhard Zotz introduced me to the forest canopy, John Barone and Justine Evans taught me the basics of getting up there on my own, and Deedra McCleam loaned or gave me several pieces of climbing equipment. Their expert advice helped make my canopy research safe and enjoyable. I thank Betsy Arnold, Phyllis D. Coley, Jonathan Gonzales, and Charles Handley for assisting me with tree species identifications. Harley P. Brown, Jon K. Gelhaus, Pedro Gnaspini, David Penney, Ricardo Pinto da Rocha, John Polhemus, A. Stanley Rand, Paul Spangler, F. Christian Thompson, and Norman E. Woodley identified specimens for me or advised me of resources for the identification of organisms I collected. Many people, especially Sabine Stuntz, Christina Campbell, Hubert Herz, Betsy Arnold, and Bettina Englebrecht, made my time on Barro Colorado Island one of the best and most unforgettable experiences of nqr life. I also extend very special thanks to David Seitz and his family for their friendship and hospitality. Finally, I thank my parents and my siblings for their unconditional support of my interests and goals. My appreciation of their patience and understanding goes far beyond words. Large portions of this research were made possible by funding from the National Academy of Sciences through Sigma Xi, the Explorers Club, the Institute for International Education/Fulbright Foundation, the Smithsonian Institution, the University of Oklahoma, and research assistantships through National Science Foundation grants IBN-9408143 to Ola Fincke and DEB-9524004 to Mike Kaspari. This work is dedicated to two outstanding educators, Drs. G. W. Folkerts and R. C. Timoney. Both made many contributions to my academic and personal growth; Dr. Folkerts introduced me to phytotelmata and Dr. Timoney helped me explore the diversity of knowledge outside of science. VI TABLE OF CONTENTS Abstract ............................................................................................................. viii Chapter 1 The macrofauna of water-filled tree holes on Barro Colorado Island, Panam a ................................................................................... 1 Chapter 2 Community structure in water-filled tree holes of Panama: effects of hole height and siz e ..................................................................... 56 Chapter 3 Distribution and abundance of Microvelia cavicola Polhemus (Heteroptera: Veliidae) on Barro Colorado Island, Panama .... 99 Chapter 4 Effects of leaf litter species on macroinvertebrate community properties and mosquito yield in neotropical tree hole microcosms ........................................................................................ 119 Chapter 5 Trophic interactions in neotropical tree hole microcosms 167 Chapter 6 Predation, nutrients, and community structure in water-filled tree holes of P a n a m a ................................................................................ 194 Appendix A Correlation of community parameters with abiotic factors .... 238 Appendix B Body length-biomass relationships among odonates .................... 239 Appendix C Biomass of selected mosquito sp e c ie s ............................................ 243 Appendix D Size-selective predation by top predators in tree holes .................. 245 Appendix E Species-selective predation by top predators in tree holes 247 vu ABSTRACT Most studies of aquatic community ecology have focused on large systems (e.g., lakes and streams) in temperate biomes. The primary goal of my dissertation research was to understand how abiotic factors, nutrient availability and predation affect macroorganism communier structure in a small aquatic habitat. Water-filled tree holes are phytotelmata formed when rain collects in branch crotches and other wood-lined cavities, and are common in many neotropical forests. Unlike larger systems, the ratio of organism biomass to habitat volume is often great in tree holes, and their small size is amenable to experimental manipulation. In addition, it is possible to mimic this habitat with simple materials (i.e., plastic cups containing leaf litter and water). Natural and artificial tree holes in the forest of Barro Colorado Island (BCI), Panama served as focal systems for my doctoral research. Chapter 1 provides a descriptive foundation for the experimental studies summarized in later chapters. Repeated surveys of natural and artifîcial water-filled tree holes on BCI (1994-1998) revealed 59 species of macroorganisms associated with this habitat. Over half are true flies (Insecta: Diptera), and half of the fly species are mosquitoes (Culicidae). Densities and occurrence frequencies of most taxa were similar between artificial and natural holes. A typical tree hole contains 5 or 6 macroorganism species at any time during the wet season, but up to 14 taxa occurred in some holes. Chapter 2 addresses abiotic factors as determinants of species’ distributions and community structure. Water-filled
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