Epibionts on Brachiopods from the Devonian Dundee Formation of Ohio
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EPIBIONTS ON BRACHIOPODS FROM THE DEVONIAN DUNDEE FORMATION OF OHIO Rituparna Bose A Thesis Submitted to the Graduate College of Bowling Green State University in partial fulfillment of the requirements for the degree of MASTER OF SCIENCE December 2006 Committee: Margaret M. Yacobucci, Advisor Richard D. Hoare Don C. Steinker © 2006 Rituparna Bose All Rights Reserved iii ABSTRACT Dr. Margaret M. Yacobucci, Advisor Epibionts seem to be more common in siliciclastic units than in carbonate units. To evaluate this difference, the paleontology of the Middle Devonian Dundee Formation has been explored. A total of 245 brachiopod specimens were collected from a fossiliferous horizon of the Dundee Formation exposed at Whitehouse Quarry and identified to the generic level. Brachiopod genera identified were Strophodonta, Rhipidomella, Rhynchotrema, Atrypa, and Mucrospirifer. All the brachiopod shells were examined under a stereomicroscope for evidence of epibionts, and preferred host taxa were determined. Epibionts are absent on all the brachiopod shells except some Rhipidomella shells. Further examination of these Rhipidomella shells under 100x magnification showed evidence of biotic interactions in 21 out of 48 specimens. Large boreholes were produced by worm borers, scars were left on a few specimens by worms, branching grooves were the traces of soft-bodied ctenostome bryozoans, and sheet-like encrustation was produced by an indeterminate group of bryozoans. Ctenostome bryozoans had a commensal relationship with their host while a few worms had a parasitic relation with the host. While one might expect encrustation on hardgrounds within this carbonate unit, field work has determined that much of the Dundee Formation was extensively bioturbated, implying a soft substrate. It may be that bioturbation mixed shells down into the substrate before epibionts could attach. iv DEDICATION With the completion of my Master’s thesis, I would like to express my regard to my dear parents and my family members, and my love to my dear friend Arnab for his moral support and his encouragement in completing my thesis. (Dedicated to my close ones) v ACKNOWLEDGMENTS I acknowledge my irrepayable debt to my advisor, Dr. Margaret M. Yacobucci, for her suggestions, great ideas, and guidance. I am indebted to her for her help and encouragement to explore the paleontology of a less well-known unit. I sincerely thank her for she urged me to keep up with the challenges as at times it was really very disappointing for not finding the desired results for my research. She was very informative and that made me keep going with my research and focus entirely on my goals. I also thank my research committee members, Dr. Don C. Steinker and Dr. Richard Hoare, for being a support for my research. I especially want to thank Dr. Hoare for providing me with a lot of information on the host-epibiont relationship of the overlying Silica Formation. I am also grateful to him for his suggestions for a better result on my research. I also thank my fellow graduate students Christopher Wright and David VanDeVelde for their assistance in fieldwork. I specially acknowledge Chris for providing stratigraphic information about the Dundee, including the measured section that is shown in Figure 6. I would also like to take the opportunity to thank Dr. Sheila J. Roberts and all the faculty and staff members of BGSU for their decision to select me for graduate studies in this department and for funding me for my Master’s program. And last, but definitely not least, I would also like to thank my friend Tumpa from the Computer Science department of BGSU for helping me in Microsoft Word related questions. At the end, a silent salute to Bowling Green State University from one of its many infatuated admirers. vi TABLE OF CONTENTS INTRODUCTION ................................................................................................................. 1 PREVIOUS WORK......................................................................................................... 1 Epibionts .................................................................................................................... 1 Paleontological Studies of Epibionts ........................................................................ 6 RESEARCH OBJECTIVES .................................................................................................. 10 GEOLOGICAL BACKGROUND......................................................................................... 11 PALEOENVIRONMENTAL INTERPRETATION OF THE DUNDEE FORMATION (WHITEHOUSE QUARRY)........................................................................................... 18 PREVIOUS PALEONTOLOGICAL FAUNAL LISTS FROM THE DUNDEE FORMATION ................................................................................................................. 20 TRACE FOSSILS IN THE DUNDEE FORMATION ................................................... 21 METHODS ............................................................................................................................ 26 RESULTS .............................................................................................................................. 28 TAPHONOMY OF SHELLS .......................................................................................... 28 TRACE TYPES ............................................................................................................... 30 Sheet-like Spotty Encrustation................................................................................... 30 Branching Grooves .................................................................................................... 30 Scars........................................................................................................................... 30 Large Boreholes......................................................................................................... 30 NATURE AND POSITION OF TRACES ON RHIPIDOMELLA SHELLS ................ 32 Microstructures.......................................................................................................... 32 Sheet-like Spotty Encrustations................................................................................. 38 vii Branching Grooves ............................................................................................................................................................ 39 Scars.................................................................................................................................. 50 Large Boreholes......................................................................................................... 57 Mixed Traces........................................................................................................... 57 ENCRUSTATION ON NON-BRACHIOPOD HOSTS ................................................. 58 IDENTIFICATION OF TRACE-MAKERS ................................................................... 61 DISCUSSION........................................................................................................................ 63 EPIZOAN-HOST RELATIONSHIPS ............................................................................. 63 Sheet-like Spotty Encrustations................................................................................. 63 Branching Grooves .................................................................................................... 63 Scars.................................................................................................................................. 66 Large Borehole .......................................................................................................... 67 Trace-Marks on Non-Brachiopod Hosts ....................................................................................................................................... 67 FOUR POSSIBLE CAUSES FOR RARE ENCRUSTATION IN THE DUNDEE FORMATION ................................................................................................................... 68 CONCLUSION...................................................................................................................... 70 REFERENCES ...................................................................................................................... 72 viii LIST OF FIGURES/TABLES Figures Page 1 Examples of epibionts…………………………………………………………...2-3 2 Epibionts on Paraspirifer bownockeri from the Middle Devonian of Ohio (from Sparks et al., 1980)………………………………………………………….6 3 Geologic map of southeastern Michigan and northwestern Ohio, showing the narrow belt of Dundee Formation along the Lucas County Monocline…..……....12 4 Regional geologic structures of Ohio and adjacent states…………………….......13 5 Conodont-based correlation of the Middle Devonian strata in the Lake Erie region (modified from Sparling, 1988)…………………………….......15 6 Stratigraphic section of Whitehouse Quarry (modified from Wright, 2006)…......17 7 A fossiliferous horizon of the Dundee Formation in the Whitehouse Quarry, Lucas County, Ohio……………………………………………………………….21 8 T-shaped branching network of burrows in the Dundee Formation identified as Thalassinoides ……………………………………………………….22 9 Y-shaped branching burrows in the Dundee Formation identified as Thalassinoides…..................................................................................................22 10 Criss-cross pattern of the trace fossils in in units 10 to