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Kinematics and Hydrodynamics of Cephalopod Turning Performance in Routine Swimming and Predatory Attacks

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Kinematics and Hydrodynamics of Cephalopod Turning Performance in Routine Swimming and Predatory Attacks Old Dominion University ODU Digital Commons Biological Sciences Theses & Dissertations Biological Sciences Fall 2015 Kinematics and Hydrodynamics of Cephalopod Turning Performance in Routine Swimming and Predatory Attacks Rachel A. Jastrebsky Old Dominion University, [email protected] Follow this and additional works at: https://digitalcommons.odu.edu/biology_etds Part of the Biomechanics Commons, Marine Biology Commons, and the Physiology Commons Recommended Citation Jastrebsky, Rachel A.. "Kinematics and Hydrodynamics of Cephalopod Turning Performance in Routine Swimming and Predatory Attacks" (2015). Doctor of Philosophy (PhD), Dissertation, Biological Sciences, Old Dominion University, DOI: 10.25777/tcn0-pm67 https://digitalcommons.odu.edu/biology_etds/4 This Dissertation is brought to you for free and open access by the Biological Sciences at ODU Digital Commons. It has been accepted for inclusion in Biological Sciences Theses & Dissertations by an authorized administrator of ODU Digital Commons. For more information, please contact [email protected]. KINEMATICS AND HYDRODYNAMICS OF CEPHALOPOD TURNING PERFORMANCE IN ROUTINE SWIMMING AND PREDATORY ATTACKS by Rachel Anne Jastrebsky B.S. December 2008, University of Rhode Island A Dissertation Submitted to the Faculty of Old Dominion University in Partial Fulfillment of the Requirements for the Degree of DOCTOR OF PHILOSOPHY ECOLOGICAL SCIENCES OLD DOMINION UNIVERSITY December 2015 Approved By: _______________________________ Ian Bartol (Director) _______________________________ Paul Krueger (Member) _______________________________ Michael Vecchione (Member) _______________________________ Mark Butler (Member) _______________________________ Daniel Dauer (Member) _______________________________ Daniel Barshis (Member) ABSTRACT KINEMATICS AND HYDRODYNAMICS OF CEPHALOPOD TURNING PERFORMANCE IN ROUTINE SWIMMING AND PREDATORY ATTACKS Rachel Jastrebsky Old Dominion University, 2015 Director: Dr. Ian Bartol Steady rectilinear swimming has received considerable attention in aquatic animal locomotion studies. Unsteady swimming movements, however, represent a large portion of many aquatic animals’ locomotive repertoire and have not been examined extensively. This study incorporates kinematic analyses of routine turning performance of brief squid Lolliguncula brevis and dwarf cuttlefish Sepia bandensis (Chapter 2), 3D velocimetry techniques to examine hydrodynamic turning performance of L. brevis (Chapter 3) and kinematic analyses of turning performance of L. brevis during predatory attacks on shrimp and fish prey (Chapter 4). Both L. brevis and S. bandensis demonstrated high maneuverability, having the lowest measures of length-specific turning radii reported to date for any aquatic taxa. Lolliguncula brevis was more agile than S. bandensis, i.e., L. brevis exhibited higher angular velocities during turning. In L. brevis, jet flows were the principle driver of angular velocity. Asymmetric fin motions played a reduced role in turning, and arm wrapping increased turning performance to varying degrees depending on the species. Flow patterns and relative torque contributions from the fins and jet varied with the speed of oncoming flow and orientation of the squid. Four turning categories were identified: (1) short tail-first turns, (2) long tail-first turns, (3) vertically oriented turns and (4) arms-first turns. The jet generally contributed more to turning torque than the fins in short tail-first, long tail-first and vertical turns. However, the fins produced a wider repertoire of flows, including isolated vortex rings, linked vortices and regions of elongated tubular vorticity, and were more important than the jet for turning torque generation during arms-first turns. Both the jet and fins produced torque contributing to roll and pitch, but the relative importance of these flows differed by turning category, with jet roll/pitch stabilization being critical for short tail-first turns and fin roll/pitch stabilization being integral to arms-first and vertical turns. Squid attack sequences involved three phases: (1) approach, (2) strike and (3) recoil. Lolliguncula brevis employed different attack strategies for fish and shrimp and turning performance played a significant role during predatory encounters. The squid exhibited high agility during the approach for both prey types. However, positioning, maneuverability and synchronized fin motions were more important for attacks on shrimp than fish. For attacks on fish, squid favored maximizing linear attack speeds over high maneuverability. Squid controlled their translational velocity and tentacle extension velocity during the strike, and demonstrated considerable rotational control during the recoil phase despite prey escape attempts. This study represents the most comprehensive quantitative turning performance study of cephalopods to date and demonstrates that the unique body architecture of these taxa provides exceptional advantages for maneuvering in the marine environment. iv Copyright, 2015, by Rachel Anne Jastrebsky and Ian Bartol, All Rights Reserved v This dissertation is dedicated to all of the people in my life that have helped me to develop and grow into the person I am today. vi ACKNOWLEDGEMENTS There are many people who have contributed to the completion of this dissertation. This process especially would not have been possible without all of the support, input, patience and copious amounts of red ink from my major advisor. Dr. Ian Bartol -- thank you for not only conducting research with me and helping me to put together the words to bring that research to the world, but also for being a friend and mentor. I would like to thank my dissertation committee for their collaboration, support, patience and guidance. Dr. Paul Krueger deserves special recognition for all of his assistance with code, as well as for patiently answering my many fluid mechanics questions. My lab mate Carly -- chatting with you across our desk partition for the last five years has been an absolute pleasure. I will miss our trawling trips together and thank you for all of your help, tips and advice. My lab mates Kristy and Maggie -- you made this fun and enjoyable and I am glad to have made great, lasting friendships with you in the process. My family and friends -- thank you for the unending love and support; it has not gone unnoticed, and this work would have not been possible without it. Lastly, but certainly not least, thank you to my husband, Brian, for trawling trips, presentation run throughs, listening to me “geek out” over my results, lab construction checks, de-stressing workouts together, and all of the many many other ways that you have supported me through this process. “This way of living that we once took for granted isn’t necessarily a ‘natural’ process at all. It’s not like water flowing down to the sea, not like aging. It takes effort, determination, conviction. But mostly it takes will. It takes a conscious decision to follow one difficult uphill path, and then the will to stay with it and not waver, to not give up.” ~John L. Parker Jr. in Again to Carthage vii NOMENCLATURE DML dorsal mantle length COR center of rotation L total body length R radius of the center of rotation (R/L)mean mean length specific turning radius (R/L)min minimum length specific turning radius using a 90% cut-off (R/L)amin absolute minimum length specific turning radius using a 90% cut-off ωavg mean angular velocity ωmax maximum angular velocity ωamax absolute maximum angular velocity Max D maximum distance in cm between the COR at any two instances during the turn ϴv ventral angle between the arms and mantle ϴvmin minimum ventral angle between the arms and mantle ϴvmean mean ventral angle between the arms and mantle ϴlam lateral angle between the arms and mantle ϴlmh lateral angle between the mantle and the horizontal ϴtotal total angular displacement Ty net torque acting about the yaw axis Txz torque acting about the roll and pitch axes I hydrodynamic impulse viii IA angular impulse T period of flow generation Aavg mean acceleration Apeak peak acceleration Vavg mean swimming velocity Vpeak peak swimming velocity ix TABLE OF CONTENTS Page LIST OF TABLES ...............................................................................................................x LIST OF FIGURES ........................................................................................................... xi Chapter 1. INTRODUCTION ...................................................................................................1 2. TURNING PERFORMANCE IN SQUID AND CUTTLEFISH: UNIQUE DUAL MODE MUSCULAR HYDROSTAT SYSTEMS ...............................................18 INTRODUCTION .................................................................................................18 METHODS ............................................................................................................22 RESULTS ..............................................................................................................29 DISCUSSION ........................................................................................................38 3. EXAMINATION OF TURNING HYDRODYNAMICS OF BRIEF SQUID LOLLIGUNCULA BREVIS USING VOLUMETRIC (3D) VELOCIMETRY ....47 INTRODUCTION .................................................................................................47 METHODS ............................................................................................................49
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