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Open Thesis Currano Final.Pdf The Pennsylvania State University The Graduate School Department of Geosciences VARIATIONS IN INSECT HERBIVORY ON ANGIOSPERM LEAVES THROUGH THE LATE PALEOCENE AND EARLY EOCENE IN THE BIGHORN BASIN, WYOMING, USA A Dissertation in Geosciences by Ellen Diane Currano © 2008 Ellen D. Currano Submitted in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy August 2008 The dissertation of Ellen D. Currano was reviewed and approved* by the following: Peter Wilf Associate Professor of Geosciences John T. Ryan, Jr., Faculty Fellow Dissertation Advisor Chair of Committee Russell W. Graham Director of the Earth and Mineral Sciences Museum Associate Professor of Geosciences Conrad C. Labandeira Curator of Paleoentomology, Smithsonian Institution Chairman of the Department of Paleobiology, Smithsonian Institution Special Member Lee Ann Newsom Associate Professor of Anthropology Member Scientist of the Penn State Institutes of the Environment Mark E. Patzkowsky Associate Professor of Geosciences Scott L. Wing Curator of Paleobotany, Smithsonian Institution Special Member Katherine H. Freeman Associate Department Head of Graduate Programs Professor of Geosciences *Signatures are on file in the Graduate School ii ABSTRACT Climate, terrestrial biodiversity, and distributions of organisms all underwent significant changes across the Paleocene-Eocene boundary (55.8 million years ago, Ma). However, the effects of these changes on interactions among organisms have been little studied. Here, I compile a detailed record of insect herbivory on angiosperm leaves for the Bighorn Basin of Wyoming and investigate the causes of variation in insect herbivory. I test whether the changes in temperature, atmospheric carbon dioxide, and floral diversity observed across the Paleocene-Eocene boundary correlate with changes in insect damage frequency, diversity, and composition. Because these correlations cannot be recognized without regional, high-resolution studies, this thesis makes a major contribution to the ecological understanding of disturbance and biotic response. Insect damage censuses were conducted at nine stratigraphic levels ranging in age from 59 to 52.5 Ma. A total of 9071 fossil angiosperm leaves belonging to 107 species were examined for the presence or absence of 71 DTs. Damage frequency, diversity, and composition were analyzed on the bulk floras and individual host species. Chapter 1 focuses on insect herbivory during the Paleocene-Eocene Thermal Maximum (PETM). The abrupt global warming and increase in atmospheric CO2 during the PETM make it the best geologic analog for modern anthropogenic warming. Chapter 2 examines small-scale spatial variability in insect damage along two early Eocene carbonaceous shale beds. I test whether spatial variability within a bed exceeds differences between beds. Chapter 3 extends the study interval through the early Eocene Cool Period and into the Eocene Thermal Maximum, when temperatures cool and then warm to a sustained Cenozoic maximum. Temporal trends in insect damage are generally greater than intra-bed variation, which is primarily due to differing floral composition. The Bighorn Basin dataset shows a very strong positive correlation between damage diversity and temperature. Damage diversity increases as temperature increases through the late Paleocene, peaks in the PETM, decreases during the early Eocene cooling, and then increases again during the warming to the sustained Eocene Thermal Maximum. Temperature probably affects insect herbivory by allowing diverse insect populations from lower latitudes to migrate northwards and by influencing insect metabolism and population density. iii TABLE OF CONTENTS Commonly Used Abbreviations................................................................................ vi List of Tables............................................................................................................vii List of Figures...........................................................................................................viii Acknowledgements...................................................................................................x Introduction.............................................................................................................. 1 Geologic and Paleontologic Setting............................................................. 2 Insect Damage on Angiosperm Leaves........................................................4 Research Objectives and Hypotheses..........................................................6 Methodology: Insect Damage Censuses......................................................8 Summary...................................................................................................... 9 Chapter 1: Sharply Increased Insect Herbivory during the Paleocene-Eocene Thermal Maximum..................................................................................................................11 Abstract........................................................................................................ 12 Introduction...................................................................................................12 Results and Discussion................................................................................ 13 Methods Data Collection................................................................................. 16 Quantitative Analyses of Insect Damage......................................... 16 Estimation of Leaf Mass per Area.................................................... 17 Figure Captions............................................................................................ 20 Supplementary Text Age Calibration for the Sites............................................................. 26 Paleotemperature Estimate for the Tiffanian.................................... 26 Information on Photographed Specimens........................................ 28 Chapter 2: Patchiness and Long-Term Change in Early Eocene Insect Feeding Damage Abstract........................................................................................................ 30 Introduction.................................................................................................. 30 Geologic Setting........................................................................................... 32 Methods....................................................................................................... 33 Results Damage Frequency.......................................................................... 37 Damage Diversity............................................................................. 38 Damage Composition....................................................................... 39 Early Eocene Plant-Insect Interactions.............................................41 Discussion.................................................................................................... 42 Conclusions.................................................................................................. 44 Chapter 3: A Quantitative Analysis of Insect Feeding on Angiosperm Leaves through the Late Paleocene and Early Eocene in the Bighorn Basin, Wyoming Introduction.................................................................................................. 55 Geologic Background................................................................................... 56 Paleotemperature Record............................................................................ 57 iv Methodology Insect Damage Censuses................................................................ 58 Statistical Analyses Damage Frequency.............................................................. 60 Number of Damage Types....................................................60 Damage Composition........................................................... 62 Time Series Analysis............................................................ 63 Leaf Mass per Area Analyses...........................................................64 Plant Composition and Diversity.................................................................. 65 Insect Damage through Time Damage Frequency.......................................................................... 69 DTL................................................................................................... 70 DTO.................................................................................................. 71 Damage on Individual Host Families................................................ 72 Damage Composition....................................................................... 73 Changes in MA from Skeleton Coast to 15 Mile Creek................................. 75 The Effects of Floral Diversity on Insect Damage........................................ 76 The Effects of Temperature on Insect Herbivory......................................... 78 Comparing Damage Frequency, DTL, and DTO.......................................... 80 Moving Beyond the Bighorn Basin: A Brief Look at Insect Damage in the Western Interior during the Early Paleogene Bulk Floras........................................................................................81 Individual Species.............................................................................83 Tracing Insect Damage on Plant Lineages...................................... 83 Floral Diversity, Temperature, and Insect Damage.........................
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