APPLICATIONS OF QUANTITATIVE METHODS AND CHAOS THEORY IN ICHNOLOGY FOR ANALYSIS OF INVERTEBRATE BEHAVIOR AND EVOLUTION by James Richard Woodson Lehane A dissertation submitted to the faculty of The University of Utah in partial fulfillment of the requirements for the degree of Doctor of Philosophy in Geology Department of Geology and Geophysics The University of Utah August 2014 Copyright © James Richard Woodson Lehane 2014 All Rights Reserved The University of Utah Graduate School STATEMENT OF DISSERTATION APPROVAL The dissertation of James Richard Woodson Lehane has been approved by the following supervisory committee members: Allan A. Ekdale , Chair May 5th, 2014 Date Approved Randall B. Irmis , Member June 6th, 2014 Date Approved Marjorie A. Chan , Member May 5th, 2014 Date Approved Elena A. Cherkaev , Member June 12th, 2014 Date Approved Leif Tapanila , Member June 6th, 2014 Date Approved and by John M. Bartley , Chair/Dean of the Department/College/School of Geology and Geophysics and by David B. Kieda, Dean of The Graduate School. ABSTRACT Trace fossils are the result of animal behaviors, such as burrowing and feeding, recorded in the rock record. Previous research has been mainly on the systematic description of trace fossils and their paleoenvironmental implications, not how animal behaviors have evolved. This study analyzes behavioral evolution using the quantification of a group of trace fossils, termed graphoglyptids. Graphoglyptids are deep marine trace fossils, typically found preserved as casts on the bottom of turbidite beds. The analytical techniques performed on the graphoglyptids include calculating fractal dimension, branching angles, and tortuosity, among other analyses, for each individual trace fossil and were performed on over 400 trace fossils, ranging from the Cambrian to the modem. These techniques were used to determine various behavioral activities of the trace makers, including feeding and behavioral evolution. Graphoglyptids have been previously identified as representing mining, grazing, farming, and/or trapping. By comparing graphoglyptids to known mining burrows and grazing trails, using fractal analysis, it was possible to rule out mining and grazing behaviors for graphoglyptids. To determine between farming and trapping, a review of all known trapping burrows was required. The hypothesis that graphoglyptids were trappers was based entirely on the hypothesized feeding behaviors of the worm Pciraonis. Close examination of Paraonis burrows indicated that the burrows are not traps. This means that, since Paraonis does not trap prey, graphoglyptids should not be considered traps either. Therefore, graphoglyptids likely represent farming behavior. This study also shows that previous interpretations of graphoglyptid behavioral evolution was far too simple. The results of the morphological analyses indicate that major changes to the behavioral evolution occurred during the Late Cretaceous and the Early Eocene. Previous hypotheses about Late Cretaceous evolutionary influences were validated. However there were additional influences like the Paleocene-Eocene Thermal Maximum that were not overly emphasized before. Finally, of the many theories about the driving force of evolution, chaos theory has often been overlooked. Chaos theory is a powerful tool, such that, by knowing the similarities between chaos theory and evolutionary theory, it may be possible to map out how environmental changes could shift the evolution of a species. iv This dissertation is dedicated to my lovely daughter, who has taught me the valuable lesson that no hardship is too tough, that you can’t press on through it. “For our own species, evolution occurs mostly through our behavior. We innovate new behavior to adapt.” —Michael Crichton, The Lost World (1995) TABLE OF CONTENTS ABSTRACT............................................................................................................. .....iii LIST OF TABLES........................................................................................................x ACKNOWLEDGEMENTS.................................................................................... .....xi Chapters 1. INTRODUCTION AND BACKGROUND................................................... .....1 1.1 Overview..................................................................................................... 1 1.2 Graphoglyptids...................................................................................... .....4 1.3 Sampling..................................................................................................... 11 1.4 Significance of research....................................................................... ..... 20 1.5 Summary of following chapters................................................................22 2. FRACTAL ANALYSIS OF GRAPHOGLYPTID TRACE FOSSILS...................................................................................................27 2.1 Abstract....................................................................................................... 27 2.2 Introduction........................................................................................... ..... 28 2.3 Methods and materials...............................................................................32 2.4 Results.........................................................................................................46 2.5 Discussion............................................................................................. ..... 47 2.6 Conclusion............................................................................................ .....51 3. PITFALLS, TRAPS, AND WEBS IN ICHNOLOGY: TRACES AND TRACE FOSSILS OF AN UNDERSTUDIED BEHAVIORAL STRATEGY............................................................................................................52 3.1 Abstract.......................................................................................................52 3.2 Introducti on........................................................................................... .....53 3.3 Modem trapping traces and their fossil equivalents.......................... .....54 3.4 Possible ancient traps.................................................................................71 3.5 Discussion............................................................................................. .....77 3.6 Conclusion............................................................................................ .....79 4. ANALYTICAL TOOLS FOR QUANTIFYING THE MORPHOLOGY OF INVERTEBRATE TRACE FOSSILS...................................................... .....80 4.1 Abstract.......................................................................................................80 4.2 Introducti on........................................................................................... .....81 4.3 Methodology: Starting out................................................................... .....81 4.4 Morphology dependent methods...............................................................87 4.5 Morphology independent methods...........................................................101 4.6 Materials.....................................................................................................109 4.7 Discussion............................................................................................. .....I ll 5. BEHAVIORAL EVOLUTION OF BENTHIC ORGANISMS REFLECTED IN THE GEOLOGIC RECORD OF GRAPHOGLYPTID TRACE FOSSILS .............................................................119 5.1 Abstract.......................................................................................................119 5.2 Introducti on........................................................................................... .....120 5.3 Basis of analyses........................................................................................122 5.4 Ichnotaxonomy and topology groups.................................................. .....123 5.5 Materials.....................................................................................................134 5.6 Methodology......................................................................................... .....135 5.7 Results.........................................................................................................141 5.8 Discussion............................................................................................. .....163 5.9 Conclusion............................................................................................ .....177 6. EVOLUTION IN CHAOS: THEORETICAL APPROACH OF CHAOS THEORY AS A GUIDING PRINCIPLE FOR UNDERSTANDING PATTERNS IN BIOLOGICAL EVOLUTION.............................................. .....180 6.1 Abstract.......................................................................................................180 6.2 Introducti on........................................................................................... .....181 6.3 Chaos theory......................................................................................... .....182 6.4 Chaos theory in evolution.................................................................... .....185 6.5 Morphospace......................................................................................... .....188 6.6 B ehavi oral evoluti on..................................................................................191 6.7 Discussion............................................................................................