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Sex-Specific Aggressive Decision-Making in the African Cichlid Melanochromis Auratus

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Sex-Specific Aggressive Decision-Making in the African Cichlid Melanochromis Auratus SEX-SPECIFIC AGGRESSIVE DECISION-MAKING IN THE AFRICAN CICHLID MELANOCHROMIS AURATUS Kamela De Stamey 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 August 2014 Committee: Moira van Staaden, Advisor Daniel Wiegmann Sheryl Coombs © 2014 Enter your First and Last Name All Rights Reserved iii ABSTRACT Moira van Staaden, Advisor Effective fighting strategies are essential to successfully navigate competitive social interactions. Probing the fighting ability of opponents requires that individuals employ assessment behaviors so that appropriate decisions about fighting strategies can be made. Inherent properties, such as sex and body size, have the potential to influence tactical fighting choices. African cichlids are well known for their hyper-aggressive nature and make ideal models for probing the underlying factors that impact decision-making during aggressive encounters. Here, an ethogram was constructed comprising seventeen behaviors to probe the sex- and size- related differences in the fighting decisions of same-sex pairs of Melanochromis auratus, a highly territorial Malawian cichlid. A modified Mirror Image Stimulation (MMIS) test was developed that utilized mirrors with curved surfaces to query sex-dependent strategies based on altered apparent opponent size. Differential behavior based on the sex of the fish was also observed in staged encounters of size-matched dyads. Males showed little progressive assessment behavior and instead engaged in immediate and intense fighting, whereas females exhibit longer latencies to engage opponents and prolonged assessment phases. The sexes also exhibited distinct, but different, size-dependent strategies. During MMIS, males bit and displayed at higher rates towards larger mirror-image opponents, while female responses were more circumspect. Comparisons of MMIS and size-matched encounters indicate that live opponents elicit more aggression from larger males than do flat mirror images. We conclude that male M. auratus do not conform to expectations based on typical progressive opponent assessment, but rather escalate to full-contact, high-intensity attacking very rapidly. iv I dedicate this to my parents Kevin and Debi Stamey for the unfailing love and support throughout my academic career. v ACKNOWLEDGMENTS Firstly, I would like to thank my graduate advisor Dr. Moira van Staaden for her continual support, advice, and constructive feedback throughout the entire course of this project. I would also like to thank Robert Huber for his insightful assessment and critical evaluation at every stage. In addition, his substantial statistical and technical contribution, specifically with the development of tracking software, was an invaluable part of my study. I would also like to mention special gratitude to all the members of the van Staaden (Sarah Jenevieve Jackson and Jessica LaHurd) and Huber (Udita Datta and Rohan Bhimani) labs for their feedback, manuscript review and, most importantly, for their continual support and encouragement. Thanks are also extended to my committee members, Dan Wiegmann and Sheryl Coombs, for their critique and feedback of my experimental design, notably for Dr. Wiegmanns’ expert statistical advise. Special thanks to Joe Coleman for the extensive time he took from his own thesis work to assist me in writing code for my statistical analysis. I thank Steve Queen for constructing all experimental arenas and the BGSU Stats Department for the free statistical support service they offer graduate students. Lastly, I thank my family and friends for their loving support, practical advice, and unconditional encouragement, without which this work would not have been possible. vi TABLE OF CONTENTS Page INTRODUCTION………………………………………………………………………..... 1 MATERIALS AND METHODS ........................................................................................... 5 Test subjects and Housing ......................................................................................... 5 Behavioral Analysis and Score .................................................................................. 5 Aggressive Dyadic Interactions ................................................................................. 6 Experimental Arena ....................................................................................... 6 Testing Procedures ........................................................................................ 6 Modified Mirror Image Stimulation ......................................................................... 7 Experimental Arena ....................................................................................... 7 Testing Procedures ........................................................................................ 7 Statistical Analysis ..................................................................................................... 8 Size-matched Dyads ...................................................................................... 8 Modified Mirror Image Stimulation ............................................................. 9 Comparisons of Flat Mirror Images to Size-matched Opponents ................ 10 RESULTS ………………………. ........................................................................................ 11 Behavioral Observations………………………………………………………………..11 Sex-Specific Differences in Size-matched Dyadic Contests ………………………… 11 Size-Dependent Differences in Fighting Behavior .................................................... 13 Altering Opponent Size.............................................................................................. 13 Comparing Responses to Mirror Images and Live Opponents ................................. 14 Body Size Alters Fighting Towards Flat Mirror Images ........................................... 15 vii DISCUSSION ............................................................................................................ 16 Sex-Specific Fighting Strategies ................................................................................ 16 Size-Specific Fighting Strategies ............................................................................... 17 Sequential Assessment is Sex-Dependent ................................................................. 18 Modifying Opponent Size with MMIS Alters Decision-Making .............................. 18 Differential Response Based on Opponent Type ....................................................... 21 Conclusions ............................................................................................................ 22 REFERENCES……………………………………………………………………………… 23 APPENDIX …………………………………………………………… ............................... 48 viii LIST OF TABLES Table Page 1 Ethogram .................................................................................................................... 34 2 Replicated Goodness of Fit (G-statistic) for differences in the frequency of Attack, Assessment, and Non-aggressive behaviors for males and females .......................... 35 3 Behavioral frequency. A summary of the overall frequencies for each type of behavior (N = 17) exhibited by males and females during size-matched dyads ............................ 35 4 Principle Components Analysis ................................................................................. 37 5 Male Transition Matrix .............................................................................................. 38 6 Partial correlation matrix that shows behavioral associations to Body Size in size- matched dyads ............................................................................................................ 40 7 Partial correlation matrix that shows associations between Body Size and the frequency of individual fighting behaviors in size-matched dyads ............................................ 41 8 MMIS. Behavioral Frequency ................................................................................... 42 9 MMIS: ANOVA ........................................................................................................ 43 10 Test Effects of the ANOVA (Table 9) ....................................................................... 43 11 MMIS: MANOVA ..................................................................................................... 43 12 Replicated Goodness of Fit (G-statistic) for opponent type ...................................... 45 13 Partial correlation matrix that shows behavioral associations to Body Size in MMIS trials for males and females ............................................................................................... 46 14 Partial correlation matrix that shows associations between Body Size and the frequency of individual fighting behaviors in (a) females and (b) males facing flat mirror images 47 ix LIST OF FIGURES Figure Page 1 Oblique view of experimental arena for MMIS ......................................................... 33 2 Percentage of Behaviors: Size-matched dyads ......................................................... 36 3 Flow chart of male Transition Matrix ........................................................................ 39 4 S-Shape Display exhibited by female M. auratus ...................................................... 43 5 Canonical Centroid Plot ............................................................................................ 44 6 Percentage of Behaviors
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