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Twelve Million Years of Bird History: a Specimen-Based Approach to Reconstructing the Late Neogene Bird Communities of California ______

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Twelve Million Years of Bird History: a Specimen-Based Approach to Reconstructing the Late Neogene Bird Communities of California ______ TWELVE MILLION YEARS OF BIRD HISTORY: A SPECIMEN-BASED APPROACH TO RECONSTRUCTING THE LATE NEOGENE BIRD COMMUNITIES OF CALIFORNIA ____________________________________ A Thesis Presented to the Faculty of California State University, Fullerton ____________________________________ In Partial Fulfillment of the Requirements for the Degree Master of Science in Geological Sciences ____________________________________ By Peter Alfred Kloess Thesis Committee Approval: Dr. James Parham, Chair Dr. Nicole Bonuso, Department of Geological Sciences Dr. Adam Woods, Department of Geological Sciences Fall, 2015 ABSTRACT California has experienced significant climate change from the onset of the Miocene (~23Ma) to the present. The impact of past climate change is often recorded in the fossil record, and can be revealed by studying how ecological communities change through time. Fossil seabirds are the ideal taxon for studying faunal responses to environmental changes because they are numerous in collections, easily identified from fragmentary remains, and since modern seabirds respond quickly to immediate changes in their environments, we expect the fossil record of seabirds to faithfully represent past environments. The first study to look at fossil seabird diversity through the Tertiary of the North Pacific relied entirely on literature records to describe the appearance and disappearance of seabird species and correlated these patterns to geologic and climatic events. My thesis utilizes an empirical, specimen-based approach to accurately describe the seabird response to climate and tectonic change during ~12 million years of coastal California’s geologic history (middle Miocene to early Pliocene). The foundation of my dataset is a previously unstudied collection of 305 bird specimens from the John D. Cooper Center for Archaeology and Paleontology, representing a relatively complete sequence of strata (Topanga Group, Monterey Formation-equivalent, and Capistrano Formation). Representing the middle Miocene to early Pliocene of Orange County, these strata form the basis for delineating chronostratigraphic bins used for studying the ii diversity of the fossil seabird community of California. Comparison of this new collection with 378 seabird specimens from three other institutions across California provides a more complete and detailed view of the seabird community during this period. Diversity, taxonomic richness, and relative abundance were examined using quantitative statistical methods to understand the change in seabird populations over time and depth of deposition. Using these statistical techniques, my data show a clear increase of pan- alcid abundance and decline of sulid abundance that is coincident with global climatic and tectonic changes as well as enhanced nutrient upwelling. Upwelling through this time also accounts for morphological changes in salmon and speciation in marine mammals. In the future, the specimen-based methodology used here can be applied to contemporaneous taxa, such as marine mammals, to quantitatively analyze diversity and relative abundance during the late Neogene and further explore the relationship between physical drivers and faunal change. iii TABLE OF CONTENTS ABSTRACT ................................................................................................................... ii LIST OF TABLES ......................................................................................................... vi LIST OF FIGURES ....................................................................................................... vii ACKNOWLEDGMENTS ............................................................................................. ix Chapter 1. INTRODUCTION ................................................................................................ 1 2. TECTONICS AND STRATIGRAPHIC UNITS OF CALIFORNIA FROM THE MIDDLE MIOCENE TO EARLY PLIOCENE ................................................... 4 Neogene Plate Tectonics of Southern California .................................................. 5 Seabird-Bearing, Marine Rock Formations of California .................................... 7 Middle Miocene Rock Units (Time Bin 1) .................................................... 7 Late Miocene Rock Units (Time Bin 2) ........................................................ 13 Mio-Pliocene Rock Units (Time Bin 3) ......................................................... 17 3. FOSSIL SEABIRDS FROM THE MIDDLE MIOCENE TO EARLY PLIOCENE OF CALIFORNIA ............................................................................ 26 Nomenclature ........................................................................................................ 28 Collections ............................................................................................................ 28 Orange County Paleontology Collection ....................................................... 28 Museum Collections Across California ......................................................... 29 Identification .................................................................................................. 30 Seabird Clades Examined in this Study ................................................................ 34 Pan-Alcidae (Charadriiformes) ...................................................................... 34 Mancallinae .................................................................................................... 34 Procellariidae (Procellariiformes) .................................................................. 35 Sulidae (Suliformes) ...................................................................................... 35 Other families present .................................................................................... 35 Seabirds Through Time ........................................................................................ 36 Time Bin 1 (14.7 to 17.0 Ma) ........................................................................ 36 Time Bin 2 (10.0 to 14.7 Ma) ........................................................................ 37 Time Bin 3 (4.9 to 10.0 Ma) .......................................................................... 38 iv 4. STATISTICAL ANALYSIS OF CALIFORNIA’S FOSSIL SEABIRD COMMUNITY ..................................................................................................... 40 Analyses ................................................................................................................ 41 Diversity ......................................................................................................... 41 Richness ......................................................................................................... 42 Abundance ..................................................................................................... 43 Multivariate Techniques ................................................................................ 44 Results ................................................................................................................... 45 Diversity ......................................................................................................... 45 Richness ......................................................................................................... 46 Abundance ..................................................................................................... 49 Multivariate Techniques ................................................................................ 53 5. PHYSICAL DRIVERS OF SEABIRD COMMUNITY EVOLUTION DURING THE LATE NEOGENE ........................................................................................ 60 Potential Taphonomic Bias from Osteosclerosis within Pan-Alcidae .................. 62 Pacific Circulation Patterns and Plate Tectonics .................................................. 65 The Monterey Formation-equivalent of Orange County ...................................... 67 Response of Contemporaneous Non-Avian Taxa ................................................. 68 Changes in Diversity During The Pliocene/Pleistocene ....................................... 68 Conclusions ........................................................................................................... 71 APPENDICES ............................................................................................................... 72 A. Fossil Seabird Species in Middle Miocene to Early Pliocene California Strata .............................................................................................................. 72 B. Extant Bird Specimens Used for Comparison and Identification .................. 75 C. Identified Seabird Material from middle Miocene to Early Pliocene California Strata ............................................................................................. 78 REFERENCES .............................................................................................................. 91 v LIST OF TABLES Table Page 1. Taxa listed as seabirds ......................................................................................... 27 2. Counts of identified fossil bird specimens from middle Miocene to early Pliocene strata across California, organized by repository .................................. 30 3. Counts of identified fossil bird specimens organized by anatomical element ..... 32 4. Richness and evenness metrics of fossil bird specimens from middle Miocene to early Pliocene
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