Behavioural and Physiological Studies of Fighting in the Velvet Swimming Crab, Necora Puber (L.) (Brachyura, Portunidae)
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Behavioural and Physiological Studies of Fighting in the Velvet Swimming Crab, Necora puber (L.) (Brachyura, Portunidae) by Kathleen Elaine Thorpe B.Sc. (Leicester) Department of Zoology University of Glasgow A thesis submitted for the degree of Doctor of Philosophy at the Faculty of Science at the University of Glasgow © K. E. Thorpe August 1994 ProQuest Number: 13834084 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 13834084 Published by ProQuest LLC(2019). 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 'ft-M 113 ^ i P -I Irsssssr UNIVERSITY C( I I { library _ Dedicated to the memory of my Dad, Ian Grenville Thorpe. THAT'S THE WHOLE PROBLEM WITH SCIENCE. 'fCU'VE GOT A BUNCH OF EMPIRICISTS TRT1NG TO DESCRIBE THINGS OF UNIMAGINABLE WONDER. ACKNOWLEDGEMENTS First of all, sincere thanks and great respect are due to my supervisors, Felicity Huntingford and Alan Taylor, for their help, advice and enthusiastic encouragement throughout the course of this work. Special thanks go to Felicity for constructive criticism, discussion and guidance during the preparation of this thesis, and also for a welcomed source of pocket money. Thanks go to all the people who have helped me in anyway, especially Cathy McLagan and June Freel, who answered my questions and cries for help with patience; Alan McGregor and Graham Adam for making bits and pieces for me and assisting my own efforts in the workshop; Roddy Macdonald and John Laurie for looking after my beasties; and, Liz Denton for Apple Mac advice and showing me the joys of the scanner. I am very grateful to Cathy McLagan, Brenda McPherson and Davy Watson for action above and beyond the call of duty. To my chums in Glasgow - Kathleen, Alan, Jayne and Isabel (my fellow water babies), Kate, Alwyn, Catherine, Fiona, Richard, June, Neil, Alan, Roddy, Sunil and Andy. You made it fun! Many thanks go to the members of Kirkintilloch Thistle B.S.- A.C., especially Eddy, Gavin, Grant, Greg, Henry, Jeanette, Kevin and Sammy, for good crack on diving trips and for buying me numerous pints when I was broke. Drinks are on me! A big, big thank you goes to Kathleen for being a true friend. For managing to share a flat with me for three years, Chinese takeaways, cinema outings, for being a fellow shopaholic, cheering me up when I was down and innumerable drinking sessions. Thank you for allowing me to use your Apple Mac. Finally, words are inadequate to express my thanks to my Mum and my sister, Irene, for their love and support in all my ventures. Special thanks to Mum for additional financial support for me and Dewy. Many thanks to Catherine Bettinson for helping out in the early days. Thanks to Catherine & Rob Bettinson, & Uncle Jimmy & Uncle Louie for keeping me stocked with finest Lincolnshire produce. The work described in this thesis was funded by a studentship from the Natural Environment Research Council, to whom I am grateful. DECLARATION I declare that this thesis is my own composition and that the research described herein was performed entirely by myself except where expressly stated. K. Elaine Thorpe August 1994 Table of Contents Summary vi Chapter 1 General Introduction 1 1.1 Why fight? 2 1.2 Effects of agonistic behaviour 2 1.3 Agonistic behaviour and game theory 3 1.4 The velvet swimming crab, Necora puber (L.) (Crustacea, Decapoda, Brachyura, Portunidae) 5 Chapter 2 Relative size and agonistic behaviour in the female velvet swimming crab, Necora puber 9 2.1 Introduction 10 2.1.1 Agonistic displays of the Crustacea 10 2.1.2 Application of game theory to agonistic behaviour 11 2.1.3 Effect of relative size on agonistic behaviour in Crustacea 12 2.1.4 Aims 14 2.2 Materials & Methods 15 2.2.1 Experimental animals 15 2.2.2 Observation of agonistic behaviour 15 2.2.3 Agonistic behaviour of female Necora puber 16 2.2.4 Definition of initiation and duration of interactions 18 2.2.5 Content of agonistic interactions 19 2.2.6 Statistical analyses 21 2.3 Results 22 2.3.1 Effect of relative size on initiation and outcome 22 2.3.2 Contests won by small crabs 22 2.3.3 Evaluation of contest costs 22 2.3.4 Behaviour of winners and losers 25 2.4 Discussion 29 i Chapter 3 Escalation and communication during agonistic behaviour in Necora puber 33 3.1 Introduction 34 3.1.1 Escalation in animal fights 34 3.1.2 Explaining and predicting the occurrence of escalation - the use of game theory 34 3.1.3 Assessment of fighting ability and resource value 36 3.1.4 Behavioural mechanisms underlying escalation 38 3.1.5 Communication during aggressive encounters 39 3.1.6 Information availability 41 3.1.7 Resolution of fights 42 3.1.8 Describing and analysing sequences of behaviour 42 3.1.9 Communication in Crustacea 45 3.1.10 Aims 46 3.2 Materials & Methods 47 3.2.1 Experimental design 47 3.2.2 Statistical analyses 47 3.3 Results 50 3.3.1 Males 50 3.3.1.1 Inter-individual transitions: winners 50 3.3.1.2 Inter-individual transitions: losers 55 3.3.1.3 Comparison of inter-individual (winner & loser) transition matrices 62 3.3.1.4 Intra-individual transitions: winners 64 3.3.1.5 Intra-individual transitions: losers 69 3.3.1.6 Comparison of intra-individual (winner & loser) transition matrices 74 3.3.2 Females 76 3.3.2.1 Inter-individual transitions: winners 76 3.3.2.2 Inter-individual transitions: losers 82 3.3.2.3 Comparison of inter-individual (winner & loser) transition matrices 88 3.3.2.4 Intra-individual transitions: winners 90 3.3.2.5 Intra-individual transitions: losers 95 3.3.2.6 Comparison of intra-individual (winner & loser) transition matrices 100 ii 3.3.3 Comparison of males and females 100 3.4 Discussion 102 Chapter 4 Metabolic consequences of agonistic behaviour and exercise in male Necora puber 108 4.1 Introduction 109 4.1.1 Costs of agonistic behaviour 109 4.1.2 Measurement of the physiological costs associated with behaviours 109 4.1.3 Physiological consequences of various activities in non-crustacean species 110 4.1.3.1 Anuran amphibians 110 4.1.3.2 Birds 112 4.1.4 Crustacean metabolism 113 4.1.5 Responses of brachyuran crabs to exercise 116 4.1.6 Physiological consequences of fighting in Necora puber 117 4.1.7 Aims 118 4.2 Materials & Methods 119 4.2.1 Experimental animals 119 4.2.2 Experimental development 119 4.2.3 Experimental design 119 4.2.3.1 Metabolic consequences of agonistic behaviour 119 4.2.3.2 Metabolic consequences of exercise 120 4.2.4 Preparation of haemolymph and tissue samples using perchloric acid (PCA) extraction 120 4.2.4.1 Haemolymph samples 120 4.2.4.2 Tissue samples 121 4.2.5 Enzymatic analyses 121 4.2.5.1 L-lactate determination 121 4.2.5.2 D-glucose determination 122 4.2.5.3 Glycogen determination 123 4.2.6 Statistical analyses 124 4.2.6.1 Metabolic consequences of agonistic behaviour 124 iii 4.2.6.2 Metabolic consequences of exercise 124 4.3 Results 126 4.3.1 Comparison of the metabolic consequences of agonistic behaviour, exercise and low activity 126 4.3.1.1 Haemolymph metabolites 126 4.3.1.2 Tissue metabolites 126 4.3.1.3 Relationship between metabolic and behavioural variables 129 4.3.2 Metabolic events during recovery from exercise 129 4.3.2.1 Haemolymph metabolites 129 4.3.2.2 Tissue metabolites 133 4.4 Discussion 138 Chapter 5 Constraints imposed by agonistic behaviour in male Necora puber 143 5.1 Introduction 144 5.1.1 Effect of aggression on reef-building corals 144 5.1.2 Behavioural constraints of reproduction in crickets - calling versus courtship gifts in males 145 5.1.3 Metabolic constraints on spider web relocation 146 5.1.4 Metabolic constraints on behaviour in decapods 147 5.1.5 Aims 148 5.2 Materials & Methods 149 5.2.1 Experimental animals 149 5.2.2 Experimental design 149 5.2.3 Statistical analyses 150 5.3 Results 151 5.3.1 Immediate re-fights 151 5.3.2 Delayed re-fights 151 5.4 Discussion 156 Chapter 6 General Discussion 158 iv References 165 v SUMMARY The velvet swiming crab, Necora puber (L.), is a marine brachyuran commonly found in shallow, rocky, sublittoral areas on the Atlantic coasts of Europe. In Britain, the commercial importance of N. puber has increased over the last fifteen years, following the collapse of the Spanish fishery for this species, and the crab now forms the basis of an important fishery based mainly off the west coast of Scotland. N. puber has traditionally been regarded as "aggressive", due mainly to its rapid defensive response of striking and grasping humans who attempt to handle them, but relatively little was known of their intra-specific agonistic behaviour until the work of Smith (1990; Smith & Taylor 1993; Smith et al.