The tail flip escape response of the brown shrimp Crangon crangon (L.) in the context of predator-prey interactions Stephen Andrew Arnott This thesis is presented for the degree of Doctor of Philosophy at the University of Glasgow, Institute of Biomedical & Life Sciences, Division of Environmental & Evolutionary Biology. November 1996 ProQuest Number: 11007907 All rights reserved INFORMATION TO ALL USERS The quality of this reproduction is dependent upon the quality of the copy submitted. In the unlikely event that the author did not send a com plete manuscript and there are missing pages, these will be noted. Also, if material had to be removed, a note will indicate the deletion. uest ProQuest 11007907 Published by ProQuest LLC(2018). Copyright of the Dissertation is held by the Author. All rights reserved. This work is protected against unauthorized copying under Title 17, United States C ode Microform Edition © ProQuest LLC. ProQuest LLC. 789 East Eisenhower Parkway P.O. Box 1346 Ann Arbor, Ml 48106- 1346 Iq U! S' ^LASCOw' ©NIVERSIfS’ I h b b a r y I Declaration: I declare that this thesis represents, except where a note is made to the contrary, work carried out by myself. The text was composed by myself. Stephen Andrew Amott. November 1996. ACKNOWLEDGEMENTS There are many people who have helped me in all manner of ways during the period of my PhD, and I am extremely grateful for all of the assistance which they have given me. First and foremost, I would like to thank my supervisors Dr. Douglas Neil and Dr. Alan Ansell for offering me the opportunity to undertake this project, for the excellent guidance they have provided through all stages of my laboratory and field work, and especially for their speedy and productive feed-back during my writing up period. The quality of writing and discussion has been greatly enhanced by their ideas and suggestions. I am grateful to Prof. Felicity Huntingford for the support she has offered, and for the provision of facilities at the University of Glasgow. Considerable technical assistance was provided by Graham Adam, June Freel, Kate Orr, Willie Orr, Nosrat Mirzai and Graham Tobasnick. I also have Dr. Neil Metcalfe and Dr. Willie Fowler to thank for the discussion they have offered on various aspects of my work, and which helped in generating ideas and ways of presenting data. Iain Barber, Phil Caimey, Andrea Fidgett, Mhairi Forbes, Jan Holmes, Cornelia Leibrock, Francesca Marubini, Francis Neat and Murray Roberts have all helped, either by reading through chapters of the thesis, helping with experiments, offering their artistic expertise, or just helping to maintain sanity. A large proportion of the work presented here was conducted at the Dunstaffnage Marine Laboratory near Oban. I would like to thank Dr. Robin Gibson for providing me with facilities in the S.R.P.2 group, and Dr. Bob Batty for the provision of, and assistance with, the high speed video equipment. The field work that was conducted at Tralee Beach would not have been possible without the cheerful labour of Tim Ellis and Simon Morley in particular, but also Alan Gage, Amanda McLean, Lyn Newton, Elvira Poloczanska, Paul Provost and numerous others to whom I can only offer my apologies if they are not included here. I would also like thank Simeon Hill, Lois Nickell and Linda Robb for their assistance, and for arranging space for me at the laboratory. Mike Burrows has helped in numerous aspects of my work, particularly with his computer expertise during analysis of the escape trajectory data. Paolo Domenici also read through an early version of the escape trajectory work, and I am extremely grateful for the thought-provoking and influential discussion which he offered in reply. I have Jim Watson to thank for the computing back-up and assistance provided whilst I was at Dunstaffnage, and John Joyce for the provision of aquarium space. Dr. Stefan Galler and Dr. Karlheiz Hilber of Salzburg University have offered extremely generous support during various stages of my PhD, and I am truly grateful for the selfless assistance which they have provided. This project was funded by NERC grant GT4/92/167 in collaboration with the Dunstaffnage Marine Laboratory, and I would like to thank them for their financial backing. I would also like to thank Dr. Mike Adams, without whom I would probably never have embarked upon this PhD. Lastly, but my no means least, I would like to thank my parents for all the encouragement and support they have provided me with leading up to, as well as during my PhD. Crangon crangon List of Contents i-vii List of Tables viii List of Figures ix-xii Summary xiii-xv List of contents LIST OF CONTENTS Page SUMMARY................................................................................................................................xiii CHAPTER 1: General Introduction 1.1 Interactions between animals, and associated adaptations ............................................. 1 1.2 The biology of Crangon crangon .....................................................................................2 1.3 Defence mechanisms of C. crangon against predators .....................................................8 1.4 Optimal foraging theory, and the effect of escape behaviour ......................................... 10 1.5 Tail flip swimming in Crustaceans .................................................................................... 11 1.6 Optimal evasion strategies ..................................................................................................16 1.7 Aims of the study .................................................................................................................19 CHAPTER 2: High speed video analysis of tail flip swimming in Crangon crangon 2.1 INTRODUCTION................................................................................................................21 2.2 MATERIALS & METHODS.............................................................................................23 2.2.1 Animals ........................................................................................................................... 23 2.2.2 Estimation of the centre of mass ................................................................................... 24 2.2.3 Experimental set-up ....................................................................................................... 24 2.2.4 Experimental protocol ................................................................................................... 25 2.2.5 Analysis of video films ..................................................................................................25 2.2.5.i Analysis of body angle and centre of mass parameters ...................................... 26 2.2.5.ii Measurement of the rotation angle in the shrimp’s pitch plane ........................26 2.2.5.iii Measurement of antennal scale and uropod extension and retraction rates................................................................................................................................ 26 2.2.5.iv Measurement of the velocity of the head and tail fan relative to the centre of mass ............................................................................................................... 26 2.2.5.V Measurement of escape response latencies ............................................................27 2.2.6 Statistical analysis of data ............................................................................................... 27 List of contents Page 2.3 RESULTS..............................................................................................................................27 2.3.1 Description of the tail flip swimming behaviour ...........................................................27 2.3.2 Movements of the head and tail fan during tail flips .....................................................28 2.3.3 Orientation of shrimps whilst tail flipping .................................................................... 29 2.3.3.1 Movements in the shrimp’s roll plane ................................................................... 29 2.3.3.1.a Roll movements during the first tail flip of an escape response ................... 29 2.3.3.1.b Roll movements during subsequent tail flips of the escape response ............................................................................................................................30 2.3.3.11 Movements in the shrimp’s pitch plane ................................................................. 30 2.3.3.11.a Pitch movements during the first tail flip of an escape response ................30 2.3.3.11.b Pitch movements during subsequent tail flips of the escape response ...................................................................................................................... 30 2.3.3.11.c Effects of removing the head and/or the tail fan upon steering in the pitch plane ............................................................................................................ 31 2.3.3.111 Movement in the shrimp’s yaw plane .................................................................. 31 2.3.4 Effect of shrimp length on the duration of tail flips ......................................................31 2.3.5 Effect of shrimp length on body angle measurements ..................................................32 2.3.5.1 Body angle .............................................................................................................