Non-Vocal Communication in the Atlantic Spotted Dolphin
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The University of Southern Mississippi NON-VOCAL COMMUNICATION IN THE ATLANTIC SPOTTED DOLPHIN (STENELLA FRONTALIS) AND THE INDO-PACIFIC BOTTLENOSE DOLPHIN (TURSIOPS ADUNCUS) by Robin Duncan Paulos A Thesis Submitted to the College of Education and Psychology of The University of Southern Mississippi in Partial Fulfillment of the Requirements for the Degree of Master of Arts Approved: ____________________________________ Director ____________________________________ ____________________________________ ____________________________________ ____________________________________ Dean, College of Education and Psychology May 2004 ABSTRACT NON-VOCAL COMMUNICATION IN THE ATLANTIC SPOTTED DOLPHIN (STENELLA FRONTALIS) AND THE INDO-PACIFIC BOTTLENOSE DOLPHIN (TURSIOPS ADUNCUS) by Robin Duncan Paulos May 2004 The goal of this project was to increase our understanding of non-vocal communication in the social lives of two species of dolphins, the Atlantic spotted dolphin (Stenella frontalis) and the Indo-Pacific bottlenose dolphin (Tursiops aduncus). The non- vocal behaviors produced by dolphins in various social contexts and the reactions of other dolphins to these behaviors were described and analyzed. Although no specific patterns of behavior (defined as sequences comprised of greater than two behaviors) were found in either species based on lag sequential analysis, significant associations between target events and non-vocal behaviors were identified. Several behaviors were significantly associated with three “target events” (depart, join, and contact) in both study groups including contact behaviors. Spotted dolphins were more likely to touch another individual after joining than before departing (76.47% vs. 28.57%). The Indo- Pacific bottlenose dolphins were equally likely to touch another after joining or before departing (44.44% vs. 57.14%). However, there was no significant difference between species in the use of these behavioral combinations. Both species engaged in reciprocated contact in only two behavioral contexts (mingle and play). The spotted ii dolphins, however, use a wider variety of contact behaviors than do the Indo-Pacific bottlenose dolphins. Serendipitously, a behavior previously undocumented in the literature was observed in both species of dolphin during data analyses for this study. This behavior was labeled an “oscillating swim”. This swim behavior is defined as a dolphin moving forward while its entire body is involved in a rolling-type movement. The oscillating swim was documented in five social contexts (general social, travel, forage, play, and inquisitive) for both species, but the use of this behavior varied with age (χ2 = 24.241; df = 3; p<.001) as well as by sex (χ2 = 3.98; df = 1; p<.05). iii DEDICATION This thesis is respectfully and lovingly dedicated to Terry and Hope Paulos, and Jennie and Robert Duncan. Your unending support, indefatigable enthusiasm, and constant encouragement kept me going when the light became dim. Thank you for being such wonderful role models and tireless cheerleaders. iv ACKNOWLEDGMENTS This work could not have been completed without the advice and support provided by a number of people. I would first like to thank the Chair of my committee, Stan Kuczaj. Without his generosity, guidance, and artful prodding this project might still be in the planning stage. I would also like to thank him for his flexibility and commitment in making it possible for me to spend time in the field. Sincere thanks are also extended to my other committee members, Tammy Greer and Sheree Watson, for their sage advice and comments throughout this process. Special thanks go to committee member, Kathleen Dudzinski, not only for her counsel, comments, and encouragement, but for her willingness to disseminate and share data used in this project. Without her data and her support, this project could not have even commenced. I also thank her for providing opportunities for me to gain invaluable field experience. Finally, I wish to acknowledge with gratitude the technical assistance provided by Andrew Wright and Peter Sugarman in the creation of macros used in the analyses in this project (without which I would still be calculating Z-scores). v TABLE OF CONTENTS ABSTRACT………………………………………….……………………………ii DEDICATION…………………………………………………………………….iv ACKNOWLEDGEMENTS………………………………………………………..v LIST OF ILLUSTRATIONS…………………………………………………….viii LIST OF TABLES………………………………………………………………….x LIST OF ABBREVIATIONS………………………………………………..……xi CHAPTER I. INTRODUCTION………………………………………………….1 Non-vocal Communication in Humans Communication in Non-human Animals Communication in Dolphins II. THE PRESENT STUDY………………………………………….23 III. METHODS………………………………………………………..25 The Sample Animal Variables Behavioral Context Behavior Codes Video Data Collection Analysis IV. RESULTS………………………………………………………...36 Sequences of Behavior Behavioral Associations Adventitious Finding: Oscillating Swim vi V. DISCUSSION……………………………………………….…….61 Sequences of Behavior Behavioral Associations Adventitious Finding: Oscillating Swim VI. CONCLUSION…………………………………………………...76 Limitations and Future Study APPENDIXES…………………………………………………………………….79 REFERENCES………………………………………………………………..…115 vii LIST OF ILLUSTRATIONS Figure 1. Geographic location of video data acquisition for Stenella frontalis……….…..26 2. Geographic location of video data acquisition for Tursiops aduncus………..…27 3. Diagram illustrating ‘skeleton’ transitions (solid lines) and intervening transitions (dotted lines) between behaviors C, B, A, and target event………………….…35 4. Three-behavior sequence depicting pattern that occurred more than once.........39 5. Number of total occurrences of contact-depart and the number of occurrences in which the dolphin is both the initiator of contact and the ‘departer’ in both study groups…………………………………………………………….………43 6. Number of total occurrences of contact-depart in the Bahamas study group and the number of occurrences in which the dolphin is both the initiator of contact and the ‘departer’ for each age class……………………………….…..43 7. Number of total occurrences of contact-depart in the Japan study group and the number of occurrences in which the dolphin is both the initiator of contact and the ‘departer’ for each age class…………………………………..44 8. Number of depart occurrences by behavioral context that were preceded by a contact behavior in the Bahamas study group……………………...…….46 9. Number of depart occurrences by behavioral context that were preceded by a contact behavior in the Japan study group………………………………..46 10. Number of total occurrences of join-contact and the number of occurrences in which the dolphin is both the initiator of contact and the ‘joiner’……….....49 11. Total number of identified dolphins in the Bahamas study group by age class and the number of dolphins that were both the joiner and initiator of contact………………………………………………………………………….50 12. Total number of identified dolphins in the Japan study group by age class and the number of dolphins that were both the joiner and initiator of contact……………………………………………………………………….…51 viii 13. Total number of join events in the Bahamas study group by bbc and the number of times that contact followed those join events…………………..53 14. Total number of join events in the Japan study group by bbc and the number of times that contact followed those join events……………..…....53 15. Total number of identified dolphins and number observed performing oscillating swim by sex and site……………………………………………..…58 16. Total number of identified dolphins and number of dolphins observed performing the oscillating swim in the Bahamas study group by age………….59 17. Total number of identified dolphins and number of dolphins observed performing the oscillating swim in the Japan study group by age……………..59 ix LIST OF TABLES Table 1. Spot Class Designations and Associated Color Pattern Changes in Atlantic Spotted Dolphins…………………………………………………….………...29 2. Number of Seconds by Site, Year, and Broad Behavioral Context …………..36 3. Number of Target Events for Atlantic Spotted Dolphins in the Bahamas and Indo-pacific Bottlenose Dolphins in Japan by Broad Behavioral Context…....37 4. Proportion of Animals in Each Age Group by Year…………………………..38 5. Number of Dolphins Included in the Sequential Analyses by Site and Age Group……………………………………………………………………..40 6. List of Significant Behaviors (Z> 3.291) Before (lags -1, -2, or -3) the Target Event depart…………………………………………………………....42 7. Percentage of depart Occurrences that were Preceded by a Contact Behavior……………………………………………………………………….45 8. Results for the Pairwise Comparisons (for contact-depart) using Holm’s Sequential Bonferroni Method……………………………………..…47 9. List of Significant (Z> 3.291) Behaviors After (lags +1, +2, or +3) the Target Event join…………………………………………………………..….48 10. Percentage of join Occurrences that were Followed by a Contact Behavior………………………………………………………………….…...52 11. Results for the Pair-wise Comparisons (for join-contact) using Holm’s Sequential Bonferroni Method……………………………………………......54 12. List of Significant (Z> 3.291) Behaviors that Occurred before (lags -1, -2, or -3) and After (lags +1, +2, or +3) the Target Event contact…..56 13. Results for the Age Pair-wise Comparisons using Holm’s Sequential Bonferroni Method………………………………………………………….…60 x LIST OF ABBREVIATIONS bbc broad behavioral context DCP dolphin communication project MA /I F Mystic Aquarium/ Institute for Exploration MIK Mikura Iruka Kenkyukai sag significantly occurring behavior xi CHAPTER I INTRODUCTION Communication is an integral part of daily life