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Vocal Communication of the Brown Hawk Owl Ninox Scutulata in Japan

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Vocal Communication of the Brown Hawk Owl Ninox Scutulata in Japan Vocal Communication of the Brown Hawk Owl Ninox scutulata in Japan A thesis submitted for the degree of PhD in Zoology by Teruyo Oba Royal Holloway and Bedford "New College (University of London) ProQuest Number: 10090135 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 complete 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 10090135 Published by ProQuest LLC(2016). Copyright of the Dissertation is held by the Author. All rights reserved. This work is protected against unauthorized copying under Title 17, United States Code. Microform Edition © ProQuest LLC. ProQuest LLC 789 East Eisenhower Parkway P.O. Box 1346 Ann Arbor, Ml 48106-1346 To the praise and glory of God ABSTRACT Vocalizations of the largely nocturnal Japanese Brown Hawk Owl Ninox scutulata japonica were studied, with paricular reference to the function of vocal communication. The adult owls were found to have thirteen types of vocalization and three non-vocal sounds with the possible addition of one other vocal sound. Nestlings had four vocal and one non-vocal sound. The development of the vocal repertoire was also studied from captive juveniles. Most of the vocal repertoire was recorded and sonagraphically analysed, and their behavioural contexts were investigated to give an idea of their various functions. The main function of primary calls was in interactions with nonspecific birds from a distance. The daily calling of territorial males was not time-related and appeared to be greatly influenced by the intraspecific environment. Seasonal production was concentrated during breeding activity, showing four peaks which are discussed in terms of territorial, sexual and family interactions. The unusual vocal activity of a replacing male was studied and gave valuable additional information. Primary calls were also considered to convey different types of information through variation in their physical structure. From the examination of calls in natural conditions and in aggressive responses to playback, variations of frequency and rate were related to location in the territory, the breeding cycle and the caller's motivational state. There were consistent individual variations in temporal and frequency features of the calls. Playback of neighbour and stranger's calls and that of the mate and other females' calls at various locations in the territory elicited differential responses, suggesting individual recognition by voice. A general description of the species, including their classification, distribution, migration and inter-racial differences, is also given. CONTENTS 1. Introduction 15 1.1. The aims of the study 15 1.2. The Brown Hawk Owl 18 1.3. The study areas 22 1.3.1. Zushi 22 1.3.2. Miyake-jima Island 24 2. The Vocal Repertoire 26 2.1. Introduction 26 2.2. Methods 26 2.2.1. Birds 26 2.2.2. Identification of individuals and sexes 27 2.2.3. Observation of owls 27 2.2.4. Recording sounds 28 2.2.5. Analysis of recorded sounds 28 2.2.6. Physical structure of vocalizations 29 2.2.7. Contexts of vocalizations 80 2.2.8. Interpretation of messages, meanings and functions 32 2.3. The vocal repertoire 33 2.3.1. K.hoo (the primary call) 34 2.3.2. Hoh 42 2.3.3. Guf 47 2.3.4. Oo.coo 48 2.3.5. Poo 50 2.3.6. Hyu 50 2.3.7. Krrr 52 2.3.8. Cheee 55 2.3.9. Mew and meeew 56 2.3.10. Hooh 58 2.3.11. Quay 58 2.3.12. Buf 61 2.3.13. Wing noises 61 2.3.14. Byahbyah 61 2.3.15. Pack 62 2.4. Development 63 2.5. Relationships between vocalizations *71 2.6. Comparisons "78 2.6.1. Vocal repertoire of Ninox "78 2.6.2. Vocal repertoire of Strigiformes 79 3. The Primary Call; Circadian and Seasonal Patterns of Production 83 3.1. Introduction 83 3.2. Methods 85 3.3. Nocturnal vocal activity 87 3.3.1. Time of day 87 3.3.2. Effects of the physical environment 89 3.3.3. Effects of conspecific birds 95 3.3.4. Discussion 95 3.4. Seasonal vocal activity 98 3.4.1. Seasonal peaks 98 3.4.2. Discussion 100 3.5. Vocal activity of a replacing male 103 3.5.1. Results 103 3.5.2. Discussion 106 4. The Primary Call and Aggression 108 4.1. Introduction 108 4.2. Frequency of primary calls H I 4.2.1. Methods HI 4.2.2. Results 112 4.2.3. Discussion 113 4.3. Aggressive response to playback 119 4.3.1. Methods 119 4.3.2. Results 120 4.3.3. Discussion 126 4.4. Effects of speaker's location in the territory and breeding stage 127 4.4.1. Methods 127 4.4.2. Results 130 4.4.3. Discussion 147 4.5. Motivational information 148 5. The Primary Call and Individual Recognition 150 5.1. Introduction 150 5.2. Physical basis of individual recognition by primary calls 154 5.2.1. Comparison of primary calls between two individuals 154 5.2.1.1. Methods 154 5.2.1.2. Results 155 5.2.2. Physical features of primary calls during the breeding season 162 5.2.2.1. Methods 162 5.2.2.2. Results 164 5.2.3. Discussion 168 5.2.3.1. Consistency within individuals 168 5.2.3.2. Differences between two individuals 169 5.3. Inter-individual differences in primary calls 170 5.3.1. Methods 170 5.3.2. Results 171 5.3.3. Discussion 177 5.4. Behavioural evidence for individual recognition by primary calls 180 5.4.1. Methods 180 5.4.2. Individual recognition by male primary calls 181 5.4.3. Individual recognition by female primary calls 190 5.4.4. Discussion 198 5.4.4.1. Neighbour-stranger discrimination 198 5.4.4.2. Recognition of neighbours 200 5.4.4.3. Speed of recognition 201 5.4.4.4. Sexual discrimination and mate recognition 201 5.4.4.5. Female responses 204 Acknowledgements 205 Appendices 207 References 287 LIST OF PLATES Plate I. An adult Japanese Brown Hawk Owl 17 II. A juvenile Japanese Brown Hawk Owl 17 III. Nesting environment of the Japanese Brown Hawk Owl 25 IV. A nest of the Japanese Brown Hawk Owl 25 LIST OF TABLES Table 2-1. Categories of frequency modulation of graded vocalizations 31 2-2. Sexual differences in the primary call 'k.hoo' 39 2-3. A summary of proposed messages encoded in the adult vocal repertoire of the Japanese Brown Hawk Owl 64 2-4. A summary of proposed messages encoded in the juvenile vocal repertoire of the Japanese Brown Hawk Owl 65 2-5. Comparison of physical structure between graded vocalizations 74 2-6. Repertoire size in the Strigiformes 80 3-1. The first and last primary calls of the day 90 3-2. Effects of wind strength upon the vocal activity 93 3-3. Effects of primary calls of conspecific birds upon the seasonal vocal activity of territorial males 101 4-1. Effects of mate's presence and conspecific males's primary calls upon the frequency of primary calls 116 4-2. Number of flights during the playback experiment 122 4-3. Latency of response to the first playback at different locations of speaker in the territory and breeding stages 131 4-4. Number of flights at different locations of speaker in the territory and breeding stages 132 4-5. Effects of playback of strange male's primary calls upon the mates of the test males 145 Table 5-1. Minimum number of samples required to obtain individual means 163 5-2. Intra-individual and inter-individual variations in the physical structure of primary calls during the breeding season 166 5-3. Number of flights during the playback experiment of primary calls recorded from strange and neighbouring territorial males 164 5-4. Number of flights during the playback experiment of female primary calls recorded from different individuals 192 LIST OF FIGURES Figure 1-1. Distribution of the Brown Hawk Owl 19 1-2. Study areas 23 2-1. Sonagrams of adult calls. 1. primary calls 'k.hoo' 35 2-2. The physical structure of 'k.hoo' 36 2-3. Sonagrams of adult calls. 2. graded calls: 'hoh', 'guf', 'oo .coo', 'poo', and 'hyu'. 43 2-4. Sonagrams of adult calls. 3. 'krrr', 'cheee', 'mew' and 'meeew' 53 2-5. Sonagrams of adult calls. 4. 'hooh', 'quay', 'buf and 'pack' 59 2-6. Sonagrams of juvenile calls. 1. 'leee', 'gililili', 'chililili' and 'chewlee' 66 2-7. Sonagrams of juvenile calls. 2. 'puh', 'py.o', 'soo' and 'scheee' 68 2-8. Sonagrams of juvenile calls. 3. 'pooh' and 'beh' 70 2-9. Sonagrams of juvenile calls. 4. 'o.guf', 'k.hoo' and 'pur' 72 2-10. Distribution of graded vocalizations in temporal and frequency dimensions 75 2-11. Relationships among the graded vocalizations of males 77 Figure 3-1. Generalized nocturnal patterns of primary call production at Zushi in 1980 66 3-2. Seasonal patterns of primary call production 99 3-3. Accentuation of primary call production of the replacing male 196 4-1. Physical and behavioural continuum of primary call 'k.hoo* and its gradation H O 4-2. Effects of locations upon the frequency of primary calls H 4 4-3. Effects of mate's approach upon the frequency of primary calls H6 4-4.
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