An Ecomorphological Analysis of the Feeding Mechanism in the Cownose Ray (Rhinoptera Bonasus, Mitchill 1815) Matthew A
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Florida State University Libraries Electronic Theses, Treatises and Dissertations The Graduate School 2012 An Ecomorphological Analysis of the Feeding Mechanism in the Cownose Ray (Rhinoptera Bonasus, Mitchill 1815) Matthew A. Kolmann Follow this and additional works at the FSU Digital Library. For more information, please contact [email protected] THE FLORIDA STATE UNIVERSITY DEPARTMENT OF BIOLOGICAL SCIENCE AN ECOMORPHOLOGICAL ANALYSIS OF THE FEEDING MECHANISM IN THE COWNOSE RAY (RHINOPTERA BONASUS, MITCHILL 1815) By MATTHEW A. KOLMANN A Thesis submitted to the Department of Biological Science, College of Arts & Sciences in partial fulfillment of the requirements for the degree of Master of Science Degree Awarded: Fall Semester, 2012 Matthew Kolmann defended this thesis on July 27, 2012. The members of the supervisory committee were: R. Dean Grubbs Professor Directing Thesis Gregory M. Erickson Committee Member Scott J. Steppan Committee Member Daniel R. Huber Committee Member The Graduate School has verified and approved the above-named committee members, and certifies that the thesis has been approved in accordance with university requirements. ii For my parents, Charles F. and Marcia Kolmann And My grandparents, Virginia and Charles A. Kolmann iii ACKNOWLEDGEMENTS I begin this section not knowing who to start with. If such a thing can be said, I would say that I have been extremely fortunate to know some figures of the utmost character throughout my time at FSU. Several figures stand out in my academic “upbringing,” notably Dan Huber and Paul Gignac. Dan kept me out of trouble when I was just starting as a young, upstart student at University of Tampa. His advice, kindness, and help through the good times and the bad times have given me a big brother to look up to. After leaving University of Tampa, Paul Gignac came to be the interim mentor I needed, yet another big brother to look up to. He always encouraged my exploration of new ideas and the advancement of a more critical way of thinking, about science and life too. Dean Grubbs, for whatever reason, has dealt with me for three years and I’m continually amazed by his patience and encouragement. Dean’s impressive knowledge of natural history and his continuous curiosity has been a real inspiration to me. My education in fishing, good music, and hard work are also some side effects of spending so much time on long road trips with Dean. I consider Dean to be my utmost mentor and friend. Greg Erickson has been my dogged, practical, analytical mentor since my start at FSU. Greg has really showed me what to do to become a successful researcher and has shown me to be just as critical of my own work as I can be of others. He’s reminded me that this isn’t just supposed to be just fun, but that I have a responsibility to myself and others to produce quality science. To all of my advisors, Greg, Dan, and Dean – my sincerest thanks for guiding me and accepting me into your labs. Scott Steppan propelled me along a collision course with phylogenetic methods, which is to become my new area of interest. His reading group discussions always encouraged questions and comments, no matter how simple or mundane those comments may be. A notable shout-out to Bonnie Garcia is needed, as she has been the big sister, or mother hen, who has pecked me back into line time and time again. Bonnie has looked out for me, pulled strings, beaten away the critics, and done her best to keep me happy. Neil Aschliman notably, and his brothers in arms, have gotten me in more trouble than I can imagine, but have always removed me from the thick of it. My interest in phylogenetics and expanding my evolutionary thinking is in large part due to Neil. My coauthor, Mason Dean has been an outstanding source of encouragement and opportunity. Thanks Mason for helping me with all the brainstorming and for going to batoid (pun intended) for me again and again. It’s great to be excited about research, and Mason provided a great outlet for discussing batoids and science, most times elegantly mixed together. Robert (Bob) Walsh has helped me do the most frustrating and largely the most important part of this thesis. His patience with a wide-eyed biologist amongst a room full of expensive and potentially lethal machinery has been invaluable. Special thanks goes to the faculty and staff at the marine lab, namely Felicia Coleman, Chris Koenig, Bobby Henderson, Mark Daniels, Dan Overlin, Dennis Tinsley, Kathy Houck, Miranda Johnson, and Chris Stallings for making the FSUCML such a special place. Many folks have helped with stingray specimen collection, namely Mollie Taylor, Chris Bedore, Kristene Parsons, CJ Sweetman, Matt Ajemian, Gregg Poulakis, Dana Bethea, Lisa Hollensead, the FWC Eastpoint Lab, Bob Fisher, and others. iv Another round of folks have helped with shellfish collection, namely Emily Field, Robyn Zerebecki, Johnny Sheridan, David Kimbro, Hanna Garland & Mary Balthrop. I need to sincerely thank Paul (again), Aki Watanabe, David Kay, and Mariah Pfleger for their assistance in and around the lab(s). I can’t forget the help that the Motta Lab and their associated AES and FISH minions have given me. Finally, I am in debt to K. James Soda, Dennis Slice, and Lisa Whitenack for their tireless help regarding statistics and programming. Mollie Taylor has kept me on the ball with her encouragement and there hasn’t been anyone else who has convinced me of my own worth more than she has. v TABLE OF CONTENTS List of Tables ................................................................................................................................ vii List of Figures .............................................................................................................................. viii Abstract ........................................................................................................................................ xiii INTRODUCTION………………………………………………………………………………... 1 1. A REVIEW OF THE CEPHALIC MUSCULATURE IN MYLIOBATIFORM STINGRAYS: FUNCTIONAL AND EVOLUTIONARY CONCEPTS………………… ...5 1.1 Methodology……………………………………………………............................6 1.2 Results……..……………………………………………………............................8 1.3 Discussion………………………………………………………..........................19 1.4 Conclusion….……………………………………………………........................25 2. FEEDING BIOMECHANICS OF THE COWNOSE RAY, RHINOPTERA BONASUS, OVER ONTOGENY…………………………….. ........................................37 2.1 Methodology……………………………………………………..........................39 2.2 Results……..……………………………………………………..........................44 2.3 Discussion………………………………………………………..........................46 2.4 Conclusion….……………………………………………………........................49 REFERENCES.………………………………………………………….………………..……...64 BIOGRAPHICAL SKETCH……………………………………………………...…………….. 76 vi LIST OF TABLES 1.1 List of relevant batoid muscle abbreviations .........................................................................28 1.2 Collected muscle synonyms from associated literature. n/a: this muscle was not referenced in the study, dashes (-): author’s term matches the proposed terminology for that muscle, (?): author is unclear with regards to which taxa these terms apply to. ...............................................36 2.1 Results of RMA regression scaling analyses to determine the predictive ability of cranial morphometrics and body size with respect to bite force in Rhinoptera bonasus (log Y = b log x +log a). 95% confidence intervals obtained from RMA regression of log-transformed variables versus log-transformed disk width. PBL: prebranchial length, HW: head width, HH: head height, IS: inter-spiracle distance, IO: inter-orbital distance, MBF: medial bite force, LBF: lateral bite force, DW: disk width. ..................................................................................................................54 2.2 Results of RMA regression scaling analyses of morphometric and biomechanical variables with respect to disk width (cm) in the feeding apparatus of Rhinoptera bonasus (log Y = b log x +log a). P-values obtained from permutation test of the regression, 95% confidence intervals obtained from RMA regression of log-transformed variables versus log-transformed disk width. AMMe: adductor mandibulae medialis, SB: suborbitalis, AMLa: adductor mandibulae lateralis, AMMa: adductor mandibulae major, AMD: adductor mandibulae deep, AMLi: adductor mandibulae lingualis, Res. In-Lever: resultant in-lever, Med: medial, Lat: lateral, BF: bite force, MBF: medial bite force, LBF: lateral bite force, CSA: cross-sectional area. ................................55 2.3 Descriptive statistics of morphological variables and percent contribution of each jaw adductor to bite-force generation in Rhinoptera bonasus. Theoretical bite-force values assume biting using a Class 3 lever system at the medial bite point. Cross-sectional areas (CSA) for the adductor mandibulae major (AMMa) are measures of physiological CSA, other adductors are morphological CSAs. AMMe: adductor mandibulae medialis, SB: suborbitalis, AMLa: adductor mandibulae lateralis, AMMa: adductor mandibulae major, AMD: adductor mandibulae deep, AMLi: adductor mandibulae lingualis, Res. In-Lever: resultant in-lever, Med: medial, Lat: lateral, BF: bite force, MBF: medial bite force, LBF: lateral bite force, CSA: cross-sectional area. .......................................................................................................................59 2.4 Measured forces from electrostimulation trials and analysis