Copyright is owned by the Author of the thesis. Permission is given for a copy to be downloaded by an individual for the purpose of research and private study only. The thesis may not be reproduced elsewhere without the permission of the Author. Responses of South Island Hector’s dolphins (Cephalorhynchus hectori hectori) to vessel activity (including tourism operations) in Akaroa Harbour, Banks Peninsula, New Zealand. A thesis presented in partial fulfilment of the requirements for the degree of Doctor of Philosophy in Marine Biology at Massey University, Auckland, New Zealand Emmanuelle Martinez 2010 © E. Martinez, 2010 DEDICATION Je dédie cette thèse A ma mère Christine Smagghe, Ma petite soeur Julie Estève, Et mon adorable filleul Martin Estève Pour avoir toujours été là pour moi et avec moi malgrè la distance. Ainsi qu’à la mémoire de ceux qui nous on quitté trop tôt: Mon père Joseph Martinez Mon grand-père Jean Smagghe Ma grand-mère Suzy Smagghe Mon arrière grand-mère Simone Brunerie Mon oncle Jean-Michel Stieber Et Josette Veray qui fut comme une tante pour moi. This thesis is also dedicated to the emblematic Hector’s dolphin, special to me in so many ways. FRONTISPIECE Photo copyright A.R.E.V.A. Project 2006 He tu te Pahu, He tu te Tai “If dolphin is well, so too are our coasts” Mäori whakatauaki (proverb) iv ABSTRACT For over 25 years, tour operators have been undertaking view and swim-with-dolphin trips in Akaroa Harbour off Banks Peninsula, east coast South Island, New Zealand. Following the international exponential growth in the commercial dolphin-based tourism industry, Akaroa Harbour is now a key eco-tourism destination in New Zealand with 32 daily permitted trips targeting Hector’s dolphin (Cephalorhynchus hectori hectori). Such a high number of trips is of particular concern given that this species is not only endemic but also endangered. Our current understanding of the effects of tourism activities on Hector’s dolphins in Akaroa Harbour is far from satisfactory. To ensure the sustainability of the economically-important and rapidly- growing dolphin-based tourism industry, there is an urgent need for sound scientific evidence on which to base management decisions. One of the challenging issues with the assessment of tourism impacts is the lack of baseline data. Prior to beginning the evaluation of the effects of disturbance on this population, data relating to the occurrence and demographics of Hector’s dolphins, as well as vessel traffic in Akaroa Harbour, were collected from land-based platforms during three consecutive austral summers (November and March), commencing in 2005. Examination of Sighting Per Unit Effort (here number of dolphin sightings per hour) and the dolphin fine-scale spatial distribution confirmed an inshore-offshore migration and, in the case of the latter, higher density patterns between the Kaik hills and the harbour entrance. However, no specific area was associated with a particular behaviour or nursery groups. The majority of groups consisted of adults only (91.2%, n = 2,000) and comprised mainly 2-5 individuals (83.2%). Group size varied with behaviour, being larger when socialising. Activity budgets within two outer bays were very comparable to Akaroa Harbour, except for socialising. In the harbour, Hector’s dolphins only spent a small proportion (14%) of their day (0600-1800 hr) in the absence of vessels. Vessel traffic in the harbour consisted mainly of recreational vessels (72.9%) although commercial vessels represented 70.4% of observed encounters and interacted twice as long with the dolphins. No Abstract v displacement was evident and as a result, Hector’s dolphins might compensate for high vessel traffic levels by adjusting their behavioural budget. To determine the effects of tourism activities on Hector’s dolphins’ behavioural budget, focal-group follows using a scan sample methodology were conducted from land-based stations and analysed using Markov chain models (n = 330 sequences). Vessel presence affected the activity budget of Hector’s dolphins by changing transition probabilities, bout durations and the time taken to return to a behavioural state once disrupted. Both diving (inferred foraging) and travelling were significantly disrupted by vessel interactions. The addition of one of more vessels during an encounter further disrupted diving. Responses of Hector’s dolphins to swim attempts were assessed from commercial tourism vessel trips (n = 420). The method of approach and swimmer placement affected the dolphins’ behaviour, with a reduction in avoidance when regulations were adhered to, i.e. using line abreast and around methods. Dolphin responses to swim encounters were also correlated with the number of successive attempts, dolphin group size and initial behaviour. Although Hector’s dolphins appear to be more tolerant of the presence of swimmers over time, some level of sensitisation to seasonally high levels of vessel interactions was also detected. The effects of swim encounters could potentially be exacerbated by the use of stones as an auditory stimulant. Specifically, swimmers who used stones had a greater probability of close and sustained approaches by dolphins than those who sang or simply floated on the surface of the water. Based on opportunistic photo-identification surveys (n = 254), 46% and 44% of the 50 identifiable individuals were infrequently and occasionally recorded interacting with commercial tourism vessels, respectively. It was also estimated that individuals using Akaroa Harbour are exposed to the highest level of cetacean-based tourism in New Zealand. This implies that dolphins that are frequent users of the harbour are likely to be more exposed to intensive tourism pressure. The high resighting rate of some individuals further suggests that frequent users are unlikely to discontinue using the harbour, even though they face increased human disturbance. Abstract vi Data presented here reveal the nature and the susceptibility of Hector’s dolphins to tourism activities, warranting the continuation of a moratorium on new permits. Furthermore, a reduction in daily trip numbers should be considered. Ongoing monitoring of this population’s response to tourism activities, combined with an integrated and adaptive approach to management, gives the best chance of ensuring the sustainability of the industry. vii ACKNOWLEDGMENTS “The road of life twists and turns and no two directions are ever the same. Yet our lessons come from the journey, not the destination” (Don William Jr). There have been an uncountable number of people involved during this journey in one way or another over the past few years, without whom this thesis would have never been completed. “Tēnā rāwā atu koe”, thank you everyone, “merci à tous”!!! Like any journey, it first starts with an idea and people who have unwavering trust in your capabilities, no matter what the potential length and difficulties ahead. My sincere gratitude goes first to Professor Mark Orams for providing the opportunity for me to do this PhD project and to continue researching the unique and endangered Hector’s dolphin. Thank you for being such a great mentor over the years (Sir Peter would have been proud) and the constant encouragement. My personal and academic growth as a researcher has been possible thanks to your willingness to let me learn from personal experience. My heartfelt thanks extend to my other main supervisor, Dr Karen Stockin, who has been such an inspiration to succeed. No words can express how grateful I am for all the things you have done for me over the years. I am blessed to have first known you as a friend while doing our PhD, so I am aware that there is another side to the “slave driver” and the infamous “whip” (he he he). Here it is! Karen (aka da Boss), your friendship and commitment as a supervisor have no bound and I am eternally indebted. Thanks for bearing with me and my puns. I hope that “flip her” will get “stuck in” your mind even though it might be un-bear-able to admit it! Thanks also for Acknowledgments viii making me apart of the New Zealand Common Dolphin Project and training me in necropsy procedures, although I admit, I prefer them alive than smelly. By the end of this journey, you’ve almost turned me into a “common” girl, although I still think that the Hector’s are the cutest! It has been a privilege to be part of the Coastal-Marine Research Group (C-MRG) and see it grow first under Mark and now your leadership. I also wish to acknowledge my other co-supervisors, Associate Professors Dianne Brunton and Dr Deanna Clement for their advice and support throughout this doctorate. Particular thanks are extended to Dianne for stepping in as main supervisor during the transition period when C-MRG was without a director. Last but not least, I am grateful to Deanna for agreeing to jump on board my committee half-way through, not only providing such great support, encouragement and opportunities to do aerial surveys, but also for being such a good friend ever since our days at the University of Otago. I make no apology for the hearty serving of appreciation that follows, for I am truly indebted to many people and organisations, thanks to whom this research was possible. Big thanks go to my supervisors for providing feedback on drafts of the thesis. Even bigger thanks to Sarah Dwyer, Mike Corbett, Carol Duffield, and Krista Rankmore for patiently reading, editing, and/or commenting on final versions. In addition to my committee, I am also indebted to a number of people who contributed their time and expertise throughout the years. I am very grateful to A/P Liz Slooten, Dr Matt Pawley, Dr Will Rayment, Dr Rochelle Constantine, Dr Eric Kneist, John Harraway, Trudy Webster, and Vicky Binedell for their guidance and support. A special thank you goes to Jofe Jenkins, as always, I could not have started the fieldwork without you! A project like this could not have been established without the financial support and sponsorship of many commercial companies, organisations and non-profit groups. Significant sponsors of this project include: the Department of Conservation (Canterbury Conservancy), the Institute of Natural Sciences (Massey University at Albany), the New Zealand Federation of Graduate Women, the Helping Hand Fund, the Whale and Dolphin Conservation Society, the New Zealand Forest and Bird Protection Society and Project Aware Australasia. I also thank Sokkia Optical Services Limited, Measurement Solutions Ltd, Battery Direct, T.A. Macalister, DiGiLink, Massey University and Professor Mark Orams for sponsoring the project by providing equipment. Acknowledgments ix I am grateful to the many personnel in various departments of Massey University who have assisted with logistics and administration issues during the length of this project. Particular thanks are extended to Marleen Baling, Antonio Jalilian, Kathy Hamilton, Barbara Just, Muharram Khoussainova, Jim Lee, Simon Middleton, Sri Nagappan, Dr Gabriele Schmidt- Adam, Dr Mark Seabrook-Davidson, Rachel Summers, and Kathryn Teal. A heartfelt thanks to David Janiger (Natural History Museum of Los Angeles County) for providing invaluable references not easily accessible. You truly deserved the student award at the SMM conference in Quebec in 2009. This research was helped by the generosity and additional support of the tour operators based in Akaroa Harbour, namely, in alphabetic order, Akaroa Dolphins Ltd., the Black Cat Group Ltd., Dolphin Experience Ltd., Fox II Sailing Adventures, and Onuku Farm Hostel. I wish to acknowledge the fantastic staff and crew of these tour operators with whom I have worked over the years. They were, in alphabetic order: Alex, Andrew, Chris, Christi, Christian, Ciaran, Craig, Danielle, Daren, Desree, Ducotte, Durelle, Ed, Emily, Emma, Erena, George, Graham, Greg, Hamish, Hannah, Haley, Helen, Helena, Hugh, Ian, Jane, Jess, Julia aka Fred, Julian, Karen, Kat, Kelly, Kerstin, Klinka, Lee-Ann, Liz, Lizzie, Marko, Marky, Meike, Michelle, Nelson, Nicole, Nigel, Pascal, Patsy, Paul, Pete, Pip, Pippa, Rachel, Richard, Robby, Ron, Ross, Rosy, Rowena, Roy, Russel, Samantha, Sonia, Tobias, Tom, Tony, and Trevor. I hope I have not forgotten anyone!! I am also indebted to the proprietors of Onuku Farm Hostel, Jeff and Steve, as well as three other local land owners, Vern in particular, who provided access to the observation sites. A big thank you to Sue Nelson for letting me rent such a nice house for the team. I also Acknowledgments x appreciate the continued support of DoC Canterbury, especially Laura Allum, Alistair Hutt, John Andrews, and Derek Cox as well as several members of the Akaroa community. Of course, no project can be undertaken without the fantastic help of volunteers and friends who gave generously their time and expertise, making this project such a memorable experience! A big thank you, merci, gracias, obrigada, gracie, danke to you all. In alphabetical order, they were: Hélène Augu, Jessica Banning, Thomas Barreau, Sharon Bond, Chiara Bertulli, Daniele Cagnazzi, Carla Christie, Casey Clark, Deanna Clement, Laura Colombo, Vanessa Dantung, Nicolas de la Brosse, Soledad Esnaola Scotto, Maria Fernanda Souza, Nicki Filby, José-Henrique Garcia, Danielle Gibas, Paulina Guzman, Aidan Hubbard, Jennifer Lamb, Maryse Leguèbe, Monica Mariani, Daniela Mello, Monica Merriman, Lucy Phillips, Adam Rosenblatt, Sarah Rueff, Barbara Saberton, Ronan Scullion, Andrea Traub, Andrea van Niekerk, and Stephanie Whyte. Special thank to Monika Mariani for being such a great research assistant. “Gracie mille at te, mia amica, anche a la tua famiglia”. Congratulations to those of you who went on to pursue their own postgraduate studies. I hope that you got some valuable expertise from this experience. I would like to also acknowledge my osteopath, Dr. Bruce Miller, for fixing my thesisitis (mainly back and shoulder pain) at the most critical stage of writing up. Your hands worked like magic! Tim Cahill once said that a journey is best measured in friends rather than miles. I feel really blessed to have so many friends around the world that I can count on. “Un grand merci du fond du Coeur” (a big heartfelt thank you) is expressed to my extended family in Aotearoa, all of whom have supported me throughout the most testing times and allowed me to complete this journey. I am truly grateful to, in alphabetical order: Babette Bayet, Mike Boulic, Liz Burgess, Deanna Clement, Mike Corbett, Steve and Terry Holmes, Gabriela de Tezanos Pinto, Sarah Dwyer, Gabriel Machovsky Capuska, Monika Merriman, Laureline Meynier, the Orams, the Renwicks, Karen Stockin, Karen and Sabrina Strydom, Special thanks are also owed to Pip Journeaux, Milaan, and the Renwick family for their hospitality and friendship while in the field in Akaroa. You guys are my adopted family. Jan, “muchas gracias amiga, eres como una hermana para mi”. Mere dost, “mein aap ko dhanyavaad deti houn ki aap meri achi se saheli hokar hame hamesha achi achi salaha deti aayi hai. Mei aap aur aap ke bache ko hirday se dhanyavaad Acknowledgments xi deti huon kyuki aap mujhe apni pariwar ki tahara kayi mahine tak apne ghar hame apna saath rakhi. Ye mei hamesha yaad rakhu gi”. Shukriya from DLCB! “Un autre grand merci à” Magaly Chambellant, I really enjoyed all our conversations over skype when we were both in the write up stages, going through the usual ups and downs. “Merci copine”! I cannot wait for the next time we see each other, so we can do some bed jumping and drink some Monbazillac to celebrate our achievement. To all my amazing office mates over the years: Karen Stockin, Liz Burgess, Jodi Smith, Vincenzo Petrella, Gabriel Machovsky Capuska and Sarah Dwyer. You guys have been a real blessing. Thanks for the unconditional support, the good laughs, the treats to keep me going, the hugs and the shared tears. You made the final writing stages bearable and prevented me from going insane (at least more than usual) by letting me vent my frustrations and keeping my stress levels under control. Speaking about stress levels, Gabriel “mi amigo”, a special “muchas gracias” for all the soccer table games, even though the prospect of losing to Argentina (without the “hand of the frog”) was quite stressful and almost defeated the purpose of having a break! I hope that I’ll be able to give you as much support as you guys have given me. Additional thanks are extended to all my friends and colleagues at C-MRG and the Ecology Conservation Research Groups throughout the years, that I have not already mentioned, especially those from building 86: Anne-Sophie Boyer, Birgit Ziesemann, Brigette Kriegenhofer, David Gudex-Cross, Fidi Jordan, Jenny Rickett, Sarah Wells and Sarah Whitwell. Special thanks to Sarah Gardner for letting me stay at her house for the last two weeks of the write-up so I would not be homeless. I owe you one! To Danielle Cagnazzi, Jodi Smith, Dr Claire Garrigue and Aline Schaffar for a welcoming break during the winter times and giving me the opportunity to work with other beautiful cetaceans in Australia and New Caledonia. It was a pleasure to gain further experience in the field of marine mammalogy! To all my other friends whether in Aotearoa or overseas (you know who you are). “Merci” for your understanding, support and patience. I would never have been able to get to the end of the journey without you. You made the last five years of my life so special. Now that I have finally completed my thesis, I promise to keep in touch more regularly! Acknowledgments xii Last but not least, I am eternally grateful for the love and unconditional support of my family, in particular my mother Christine, my sister Julie and my godson Martin. Merci à toute ma famille pour son soutien unconditionnel, je n’aurais jamais pu y arriver sans vous! Merci tout particulièrement à maman et Juju pour m’avoir ecoutée quand j’en avais le plus besoin. Juju, merci aussi pour tous les colis, un vrai petit bonheur de les recevoir, ainsi que ton chaleureux accueil en France (et d’Olivier et sa famille)! Merci à Martin, mon filleul, un vrai rayon de soleil dans ma vie. Merci aussi d’avoir toujours cru en moi et de m’avoir laisser partir à l’autre bout du monde pour que je puisse continuer à réaliser mon rêve d’enfance. Je vous adore et je suis super fière de vous. To anyone I forgot to mention (my memory is not what it used to be), I apologise and “merci encore”. What a journey this has been. It is the interaction and time spent with you all that has truly made this a once-in-a lifetime experience for me. I will cherish all the memories for the rest of my life. xiii TABLE OF CONTENTS Title page ..........................................................................................................................i Dedication .........................................................................................................................ii Frontispiece.......................................................................................................................iii Abstract .............................................................................................................................iv Acknowledgements...........................................................................................................vii Table of contents...............................................................................................................xiii List of figures .................................................................................................................... xviii List of tables...................................................................................................................... xxii List of abbreviations ......................................................................................................... xxiv I. The South Island Hector’s dolphin, a major tourist attraction in Akaroa harbour, Banks Peninsula 1.1 Cetacean-watching ............................................................................................... 2 1.1.1 On a global scale ....................................................................................... 2 1.1.2 On a national scale, New Zealand............................................................. 3 1.1.3 On a local scale, Akaroa Harbour ............................................................. 5 1.1.3.1 The importance of tourism to the Akaroa community................ 5 1.1.3.2 The development of a dolphin-based tourism industry............... 7 1.2 The South Island Hector’s dolphin, an endemic and endangered species ........... 9 1.2.1 Status and abundance ................................................................................ 9 1.2.2 Distribution ............................................................................................... 10 1.2.3 Life history................................................................................................ 12 1.2.4 Threats to Hector’s dolphin ..................................................................... 14 1.2.4.1 Fisheries by-catch and the BPMMS ........................................... 15 1.2.4.2 Pollution...................................................................................... 17 1.2.4.3 Vessel strike ................................................................................ 18 1.2.4.4 Cetacean-watching...................................................................... 19 1.3 Management of the cetacean-tourism industry in New Zealand .......................... 23 1.4 Rationale for this thesis ........................................................................................ 25 1.5 Thesis structure and objectives ............................................................................. 26 II. Baseline fine-scale distribution, behaviour, and group dynamics of the South Island Hector’s dolphins in Akaroa Harbour, Banks Peninsula 2.1 Introduction........................................................................................................... 31 2.2 Objectives ............................................................................................................ 32 2.3 Materials and methods ....................................................................................... 32 2.3.1 Study site and observation platforms........................................................ 32 2.3.1.1 Study site..................................................................................... 32 2.3.1.2 Observation platforms................................................................. 34 2.3.2 Data collection .......................................................................................... 36 2.3.2.1 Sampling protocol....................................................................... 37 Table of contents xiv 2.3.2.2 Theodolite protocol ..................................................................... 40 2.3.3 Data analysis ............................................................................................. 41 2.3.3.1 Sighting Per Unit Effort (SPUE)................................................. 42 2.3.3.2 Distribution and density patterns ................................................ 42 2.3.3.3 Behaviour.................................................................................... 43 2.3.3.4 Group size and composition........................................................ 45 2.4 Results ............................................................................................................... 45 2.4.1 Effort .......................................................................................................... 45 2.4.2 SPUE.......................................................................................................... 46 2.4.3 Distribution and density patterns in Akaroa Harbour ................................ 47 2.4.3.1 Potential bias associated with land-based observations.............. 47 2.4.3.2 Distribution of sightings ............................................................. 48 2.4.3.3 Mean distance of sightings.......................................................... 49 2.4.3.4 Density patterns .......................................................................... 52 2.4.4 Behaviour................................................................................................... 56 2.4.4.1 Activity budget at Te Oka, Le Bons Bay, and Akaroa ............... 56 2.4.4.2 Activity budget at Akaroa Harbour............................................. 56 2.4.5 Group size and composition....................................................................... 57 2.4.5.1 Group size ................................................................................... 57 2.4.5.2 Group composition...................................................................... 58 2.5 Discussion ............................................................................................................. 61 2.6 Conclusion ............................................................................................................ 66 III. Vessel traffic levels and encounters with the South Island Hector’s dolphins in Akaroa Harbour, Banks Peninsula 3.1 Introduction........................................................................................................... 68 3.2 Objectives.............................................................................................................. 70 3.3 Materials and methods ....................................................................................... 70 3.3.1 Study area.................................................................................................. 70 3.3.2 Survey platforms, effort, and survey protocol .......................................... 71 3.3.2.1 Survey platforms........................................................................... 71 3.3.2.2 Survey effort ................................................................................. 74 3.3.2.3 Survey protocol ............................................................................. 75 3.3.3 Data analysis ............................................................................................. 78 3.3.3.1 Proportions of time dolphins observed under different condition 79 3.3.3.2 Description of vessel traffic in Akaroa Harbour........................... 79 3.3.3.3 Correlation between SPUE and vessel traffic levels..................... 80 3.3.3.4 Encounters between Hector’s dolphins and vessels...................... 80 3.3.3.5 Reaction of Hector’s dolphins to vessel speed ............................. 81 3.3.3.6 Occurrence of encounters ............................................................. 81 3.4 Results ............................................................................................................... 82 3.4.1 Proportion of time dolphins observed under different conditions ............ 83 3.4.2 Description of vessel traffic in Akaroa Harbour....................................... 85 3.4.3 Correlation between SPUE and vessel traffic levels................................. 93 3.4.4 Encounters between Hector’s dolphins and vessels.................................. 95 3.4.5 Reaction of Hector’s dolphins to vessel speed ......................................... 98 3.4.6 Occurrence of encounters ......................................................................... 100 3.4.6.1 Encounters with different vessel types ......................................... 100 Table of contents xv 3.4.6.2 Monthly encounters with commercial tour operators ................... 105 3.4.6.3 Encounters with mothers and calves............................................. 110 3.5 Discussion ............................................................................................................ 111 3.6 Conclusion ........................................................................................................... 118 IV. Short-term behavioural responses of the South Island Hector’s dolphins to interactions with vessels in Akaroa Harbour, Banks Peninsula 4.1 Introduction........................................................................................................... 120 4.2 Objectives.............................................................................................................. 122 4.3 Materials and methods.......................................................................................... 122 4.3.1 Study site and observation platforms........................................................ 122 4.3.2 Date collection .......................................................................................... 123 4.3.3 Description of interactions between vessels and Hector’s dolphins ......... 126 4.3.4 Development of Markov chains................................................................ 126 4.3.5 Data analysis ............................................................................................. 127 4.3.5.1 Assumptions................................................................................ 127 4.3.5.2 Markov chain modelling ............................................................. 128 4.3.5.3 Forecast modelling...................................................................... 130 4.4 Results ................................................................................................................... 131 4.4.1 Field effort ................................................................................................ 131 4.4.2 Assumptions of Markov chains ................................................................ 131 4.4.3 Effect of vessel presence and interactions ................................................ 132 4.4.3.1 Transition probabilities ............................................................... 132 4.4.3.2 Probabilities of staying in a given state and average time ......... required to return to a state once disrupted ................................ 135 4.4.3.3 Mean bout length ........................................................................ 136 4.4.3.4 Behavioural budget ..................................................................... 137 4.4.3.5 Modelling the effects .................................................................. 140 4.5 Discussion ............................................................................................................. 141 4.6 Conclusion ............................................................................................................ 146 V. Effects of swim encounters on the South Island Hector’s dolphins in Akaroa Harbour, Banks Peninsula 5.1 Introduction........................................................................................................... 149 5.2 Objectives.............................................................................................................. 152 5.3 Materials and methods.......................................................................................... 153 5.3.1 Survey platforms, effort, and survey protocol .......................................... 153 5.3.1.1 Survey platforms......................................................................... 153 5.3.1.2 Survey effort ............................................................................... 154 5.3.1.3 Survey protocol ........................................................................... 155 5.3.2 Data analysis ............................................................................................. 161 5.3.2.1 Characteristics of encounters ...................................................... 162 5.3.2.2 Responses of dolphins to encounters .......................................... 162 5.4 Results .................................................................................................................. 166 5.4.1 Characteristics of encounters .................................................................... 167 5.4.1.1 Total trip time ............................................................................. 169 Table of contents xvi 5.4.1.2 Total search time to first encounter with Hector’s dolphins....... 170 5.4.1.3 Sighting cues ............................................................................... 172 5.4.1.4 Reason to end an encounter ........................................................ 172 5.4.2 Responses of Hector’s dolphins to encounters ......................................... 173 5.4.2.1 Time dolphins spent in the presence of swimmers ..................... 173 5.4.2.2 Swim encounter length according to number of swim attempts. 174 5.4.2.3 Responses to approach type ........................................................ 175 5.4.2.4 Responses to swimmer placement .............................................. 176 5.4.2.5 Orientation of Hector’s dolphins during an encounter ............... 177 5.4.2.6 Behaviour budget ........................................................................ 181 5.5 Discussion ............................................................................................................ 183 5.6 Conclusion ........................................................................................................... 194 VI. The use of auditory stimulants during swim encounters with the South Island Hector’s dolphins in Akaroa Harbour, Banks Peninsula 6.1 Introduction .......................................................................................................... 197 6.2 Objectives ............................................................................................................. 199 6.3 Materials and methods ......................................................................................... 199 6.3.1 Data collection .......................................................................................... 199 6.3.2 Sampling protocol ..................................................................................... 200 6.3.3 Data analysis ............................................................................................. 205 6.4 Results .................................................................................................................. 206 6.4.1 Field effort ................................................................................................ 206 6.4.2 Effect of swimmers’ activity on number of dolphin approaches .............. 206 6.4.3 Comparison of number of approaches towards an individual swimmer while using stones and after transferring them to another swimmer ........ 208 6.4.4 Influence of initial dolphin behaviour on interactions with swimmers .... 210 6.4.5 Occurrence of tail-slaps ............................................................................ 210 6.5 Discussion ............................................................................................................ 212 VII. Photo-identification of the South Island Hector’s dolphins associating with commercial tour vessels in Akaroa Harbour, Banks Peninsula 7.1 Introduction .......................................................................................................... 218 7.2 Objectives ............................................................................................................. 219 7.3 Materials and methods ......................................................................................... 220 7.3.1 Study area.................................................................................................. 220 7.3.2 Survey platforms, effort and survey protocol ........................................... 221 7.3.2.1 Survey platforms......................................................................... 221 7.3.2.2 Survey effort ............................................................................... 222 7.3.2.3 Survey protocol ........................................................................... 223 7.3.3 Photo-id catalogue ..................................................................................... 224 7.3.3.1 Mark quality and photograph grading......................................... 224 7.3.3.2 Matching and Hector’s dolphin catalogue .................................. 228 7.3.4 Analysis of photo-ID data .......................................................................... 229 7.3.4.1 Matching and Hector’s dolphin catalogue .................................. 230 7.3.4.2 Parameters of the sub-sampled population ................................. 230 Table of contents xvii 7.3.4.3 Site fidelity.................................................................................. 231 7.3.4.4 Estimated abundance of identifiable individuals ........................ 232 7.3.4.5 Potential exposure to vessel traffic ............................................. 237 7.4 Results .................................................................................................................. 238 7.4.1 Survey effort .............................................................................................. 238 7.4.2 Analysis of photo-ID dataset ..................................................................... 240 7.4.2.1 Comparison between the AHHD and BPHDP catalogues.......... 240 7.4.2.2 Parameters of the sub-sample population .................................. 240 7.4.2.3 Site fidelity.................................................................................. 242 7.4.2.4 Estimated abundance of identifiable individuals ........................ 250 7.4.2.5 Potential exposure to vessel traffic of identifiable individuals... 253 7.5 Discussion ............................................................................................................ 257 7.6 Conclusion ........................................................................................................... 265 VIII. Summary and recommendations for dolphin tourism activities in Akaroa Harbour, Banks Peninsula 8.1 Introduction .......................................................................................................... 268 8.2 Summary of research findings ............................................................................. 269 8.3 Significance and contribution of research findings ............................................. 274 8.4 Application of research findings and population management implications ...... 276 8.5 Study limitations ................................................................................................... 279 8.6 Recommendations for future research ................................................................. 281 8.7 Concluding statement............................................................................................ 284 IX. References X. Appendices 1.1 Number of permitted daily trips used by commercial tour operators .................. II 1.2 Number of visitors participating in commercial tours in Akaroa Harbour ........ III 1.3 Marine Mammals Protection Act (1978) ............................................................. IV 1.4 Marine Mammals Protection Regulations (1992) ............................................... VII 3.1 Compliance of commercial tourism and recreational vessels to the MMPR ....... XI 5.1 Tolerance, sensitisation and habituation ............................................................. XIX 5.2 Akaroa tour operator data sheet .......................................................................... XXI 7.1 Definitions of markings used to catalogue individuals ........................................ XXIII 7.2 Example from the AHHD catalogue and associated database ............................. XXIV 7.3 Photo-ID catalogue of Hector’s dolphins using Akaroa Harbour ...................... XXV 7.4 Summary of photo-ID data employed for the estimation of mark rate ................ XXXIV 8.1 Management recommendations ............................................................................ XXXV 8.2 Integrated and adaptive management framework ................................................ XLIV 8.3 Akaroa Harbour Marine Reserve ......................................................................... XLVI 9 Publications .......................................................................................................... XLIX xviii LIST OF FIGURES I. The South Island Hector’s dolphin, a major tourist attraction in Akaroa harbour, Banks Peninsula 1.1 Map of the global distribution of cetacean-watching countries.............................. 3 1.2 Map of the primary cetacean-watching locations in New Zealand....................... . 6 1.3 Map of Banks Peninsula, New Zealand................................................................ . 6 1.4 Abundance of Cephalorhynchus around New Zealand ........................................ 10 1.5 Location of Banks Peninsula and the Marine Mammal Sanctuary....................... 17 1.6 Marine permit zones for commercial tour operators............................................. 22 II. Baseline fine-scale distribution, behaviour, and group dynamics of the South Island Hector’s dolphins in Akaroa Harbour, Banks Peninsula 2.1 Map of Banks Peninsula, New Zealand................................................................ 33 2.2 Map showing the location of the land-based stations in Akaroa Harbour ............ 35 2.3 View from the land-based study sites in Akaroa Harbour .................................... 36 2.4 Theodolite connected to a laptop computer .......................................................... 41 2.5 Map showing the grid system used for density patterns ....................................... 44 2.6 Proportion of sightings according to distance from land-based stations .............. 48 2.7 Changes in the distribution of Hector’s dolphin sightings.................................... 50 2.8 Mean distance of sightings recorded each month from various stations .............. 51 2.9 Monthly density patterns of Hector’s dolphin sightings in Akaroa Harbour........ 53 2.10 Density patterns of Hector’s dolphins for each behavioural activity.................... 54 2.11 Density patterns for nursery groups and mixed groups ........................................ 55 2.12 A comparison of activity budget at three different locations................................ 56 2.13 A comparison of activity budget in control and distant situation......................... 57 2.14 Proportions of Hector’s dolphin groups with at least one calf present ................. 59 2.15 Composition of groups containing mothers and calves ........................................ 60 2.16 Percentage of mutliple mother-calf pairs within nursery groups.......................... 60 III. Vessel traffic levels and encounters with the South Island Hector’s dolphins in Akaroa Harbour, Banks Peninsula 3.1 Permitted area of operation for commercial operators based in Akaroa .............. 71 3.2 Map of Akaroa Harbour showing the inner, mid and outer regions..................... 72 3.3 Commercial tour vessels used as opportunistic research platforms...................... 74 3.4 Total observation time in control, distant, and close situations ........................... 84 3.5 Total observation time per month in control, distant, and close situations .......... 84 3.6 Monthly number of vessels in the middle and outer harbour ............................... 85 3.7 Hourly number of vessels according to vessel type in middle harbour ................ 87 3.8 Hourly number of vessels according to vessel type in outer harbour................... 88 3.9 Number of vessels according to time of day ........................................................ 89 List of Figures xix 3.10 Number of vessel according to time of day for different vessel types.................. 91 3.11 Number of vessels according to day and vessel type............................................ 92 3.12 Comparison of the percentage of different types of vessels in 99/00 and 05/08 ..................................................................................................................... 93 3.13 Number of Hector’s dolphins and vessels in Akaroa Harbour between 05/08..... 94 3.14 Number of Hector’s dolphins and vessels according to time of day .................... 95 3.15 Encounter time between Hector’s dolphins and different vessel types ................ 96 3.16 Encounter time per month between dolphins and commercial vessels................. 97 3.17 Vessel speed in relation to vessel type.................................................................. 99 3.18 Reaction of Hector’s dolphin groups in relation to median vessel speed ............. 100 3.19 Distance of encounters between dolphins and vessel types from Akaroa ............ 101 3.20 Locations of swimming and dolphin-watching encounters .................................. 102 3.21 Locations of encounters with different types of recreational vessels ................... 103 3.22 Observed encounters between dolphins and different vessel categories .............. 104 3.23 Locations of commercial swim-with-dolphin encounters according to month .... 106 3.24 Locations of commercial dolphin-watching encounters according to month....... 107 3.25 Maps showing path and tracks of commercial vessels in December 2006........... 108 3.26 Encounters between commercial tours and Hector’s dolphins ............................. 109 3.27 Location of encounters between commercial vessels and mother-calf groups..... 110 3.28 Encounters with mother-calves according to months and location ...................... 111 IV. Short-term behavioural responses of South Island Hector’s dolphins to interactions with vessels in Akaroa Harbour, Banks Peninsula 4.1 Map showing the location of the land-based stations in Akaroa Harbour ............ 124 4.2 Transition probabilites in control, distant, and close chains ................................ 133 4.3a Effect of vessel presence on transitions in behavioural state of dolphins (control vs. distant) ............................................................................................... 134 4.3b Effect of vessel presence on transitions in behavioural state of dolphins (control vs. close).................................................................................................. 135 4.4 Mean bout length for control, distant, and close chains ....................................... 136 4.5 Effect of vessel presence on the behavioural budget of Hector’s dolphins .......... 138 4.6 Effect of the presence of commercial and non-commercial vessels on the behavioural budget of Hector’s dolphins .............................................................. 139 4.7 Effect of the presence of one or more vessels on the behavioural budget of Hector’s dolphins .................................................................................................. 139 4.8 Differences in percentage time spent in a given state between cumulative budget and control behavioural budget................................................................. 140 V. Effects of swim encounters on the South Island Hector’s dolphins in Akaroa Harbour, Banks Peninsula 5.1 Permitted area of operation for commercial tour operators based in Akaroa ....... 153 5.2 Strategies used by commercial vessels to approach a dolphin group and place swimmers .............................................................................................. 158 5.3 Distribution of dolphin group size during swim-with-dolphin trip ...................... 167 5.4 Distribution of total number of swim attempts per trips....................................... 168 List of Figures xx 5.5 Trip duration of commercial trips according to months ....................................... 169 5.6 Trip duration of commercial trips according to departure time............................ 170 5.7 Search time taken to encounter the first dolphin group ........................................ 171 5.8 Occurrence of different sighting clues used to locate Hector’s dolphins ............. 172 5.9 Reasons for terminating a dolphin-watching or swimming encounter ................. 173 5.10 Encounter duration of successive swim attempts with a same group ................... 174 5.11 Proportion of behavioural change in dolphin groups in relation to vessel approach when considering initial behaviour of dolphins .................................... 176 5.12 Hector’s dolphin responses to swimmers as a function of placement .................. 176 5.13 Encounter duration as a function of swimmer placement..................................... 177 5.14 Probability of a dolphin group heading towards swimmers/vessels as a function of time into the encounter ....................................................................... 179 5.15 Probability of a dolphin group heading away swimmers/vessels as a function of time into the encounter ..................................................................................... 180 5.16a Behavioural budget of Hector’s dolphins in the presence of viewing and swimming vessels ................................................................................................. 182 5.16b Behavioural budget of Hector’s dolphins in the presence of small and large groups of swimmers .............................................................................................. 182 5.17 Behavioural budget of Hector’s dolphins in relation to discrete and staggered departure times ..................................................................................... 183 VI. The use of auditory stimulants during swim encounters with the South Island Hector’s dolphins in Akaroa Harbour, Banks Peninsula 6.1 Map showing Banks Peninsula and Akaroa Harbour, New Zealand.................... 200 6.2 Photos showing the different swimmer activities and types of dolphin approaches............................................................................................................. 204 6.3 Mean approach rate according to swimmers’ activity .......................................... 207 6.4 Mean sustained approach rate according to swimmers’ activity .......................... 207 6.5 Mean frequency rate of close approaches according to swimmers’ activity ........ 208 6.6 Mean approach rate with and without stones for the same swimmer ................... 209 6.7 Mean frequency of sustained approaches according to a swimmer’s activity...... 209 6.8 Mean time of first interaction; interaction rate; interaction time; and time dolphins present; according to the initial behaviour of the dolphins .................... 211 6.9 Mean frequency rate of tail-slaps according to swimmers’ activity ..................... 210 VII. Photo-identification of the South Island Hector’s dolphins associating with commercial tour vessels in Akaroa Harbour, Banks Peninsula 7.1 Permitted area of operation for commercial tour operators .................................. 221 7.2 Map showing the positions of each surveys where photo-ID was conducted ...... 239 7.3 Distribution of the minimum age of identifiable individuals............................... 241 7.4 Distribution of the number of months an individual was captured....................... 243 7.5 Distribution of the number of days an individual was captured ........................... 243 7.6 Frequency of the number of days between re-sightings ....................................... 244 7.7 Site fidelity of Hector’s dolphins in Akaroa Harbour........................................... 245 7.8 Identified dolphins found to be infrequent, occasional, and frequent users ......... 246 7.9 Map showing the positions of sightings of the five most frequent users.............. 248 List of Figures xxi 7.10 Map showing the positions of sightings over consecutive days ........................... 249 7.11 Number of individuals captured and discovery curve .......................................... 250 7.12 Photograph of the dorsal fin of individual SIDH023............................................ 251 7.13 Comparison between the monthly percent change in estimated abundance......... 254 xxii LIST OF TABLES I. The South Island Hector’s dolphin, a major tourist attraction in Akaroa harbour, Banks Peninsula 1.1 The cetacean-watching industry in New Zealand between 1994 and 2008............ 4 1.2 The cetacean-watching industry in New Zealand in 2008...................................... 5 1.3 Permitted trips allocated to target Hector’s dolphins in Akaroa Harbour .............. 8 1.4 Examples of short-term responses of cetaceans...................................................... 20 II. Baseline fine-scale distribution, behaviour, and group dynamics of the South Island Hector’s dolphins in Akaroa Harbour, Banks Peninsula 2.1 Definitions of behavioural state categories........................................................... 39 2.2 Definitions of the different categories used to classify sightings ......................... 39 2.3 Summary of effort and data collected in the field................................................. 46 2.4 Summary of SPUE according to several variables ............................................... 47 2.5 Summary of mean group size and group classes at different locations ................ 58 2.6 Variance in Hector’s dolphin group sizes at Akaroa Harbour.............................. 58 III. Vessel traffic levels and encounters with the South Island Hector’s dolphins in Akaroa Harbour, Banks Peninsula 3.1 Summary of permitted commercial vessels that operated in Akaroa Harbour ..... 73 3.2 Definitions of the different categories used to classify sightings ......................... 76 3.3 Definitions of vessel types using Akaroa Harbour ............................................... 76 3.4 Definitions of sampling protocol terms ................................................................ 77 3.5 Summary of opportunistic vessel observations onboard vessels .......................... 83 3.6 Percentage of time dolphin groups were observed under control situation.......... 83 3.7 Percentage of vessel traffic and encounters observed, mean interaction time with Hector’s dolphins according to vessel type in Akaroa Harbour ................... 95 3.8 Descriptive statistics of vessel speed according to vessel type............................. 98 3.9 Results of Pearson’s χ2 tests calculated for encounters with each vessel type ..... 104 IV. Short-term behavioural responses of South Island Hector’s dolphins to interactions with vessels in Akaroa Harbour, Banks Peninsula 4.1 Definitions of the behavioural state categories ..................................................... 125 4.2 Descriptive statistic of control, distant and close sequences ................................ 131 4.3 Chain order selection using Bayes Information Criterion .................................... 132 4.4 Probability of being in a particular state, average number of time units taken to return to a behavioural state, and average time required to return to a state .... 136 4.5 Comparison of average bout length (control vs. distant)...................................... 137 4.6 Comparison of average bout length (control vs. close) ........................................ 137 List of tables xxiii V. Effects of swim encounters on the South Island the Hector’s dolphins in Akaroa Harbour, Banks Peninsula 5.1 Definitions of encounter ratings of swim-with-dolphin trips ............................... 157 5.2 Definitions of the behavioural state categories ..................................................... 160 5.3 Definitions of sampling protocol terms ................................................................ 161 5.4 Summary of opportunistic vessel observations from commercial tours ............... 167 5.5 Statistics of swim-with-dolphin trips .................................................................... 168 5.6 Hector’s dolphin headings as towards, neutral, and away ................................... 178 5.7 Orientation of Hector’s dolphins towards swimmers and/or vessels.................... 178 5.8 Analysis of deviance for assessing goodness-of-fit of different models .............. 178 5.9 Orientation of Hector’s dolphins away from swimmers and/or vessels ............... 179 5.10 Analysis of deviance for assessing goodness-of-fit of different models .............. 180 5.11 Chain order selection using Bayes Information Criterion .................................... 181 VI. The use of auditory stimulants during swim encounters with the South Island Hector’s dolphins in Akaroa Harbour, New Zealand? 6.1 Definitions of sampling protocol terms ................................................................ 201 6.2 Definitions of behavioural state categories........................................................... 202 6.3 Definition of the different types of swimmers’ activity........................................ 203 VII. Photo-identification of the South Island Hector’s dolphins associating with commercial tour vessels in Akaroa Harbour, Banks Peninsula 7.1 Definition of categories used in the photo-ID catalogue ...................................... 226 7.2 Description of attribute criteria used to judge the quality of photographs ........... 227 7.3 Scale of photo-quality and attributes used for the catalogue and database........... 227 7.4 Summary of effort conducted in the season 2006/2007........................................ 238 7.5 Summary of effort conducted in the season 2007/2008........................................ 238 7.6 Comparison of mark rates between different species in New Zealand................. 242 7.7 Sightings of the eight frequent users per month during each field season............ 247 7.8 AIC estimates from different CJS models ............................................................ 253 7.9 Monthly estimates of abundance of marked Hector’s dolphins............................ 253 7.10 Estimated exposure levels of different delphinids in New Zealand ..................... 256 7.11 Estimation of the total number of trips Hector’s dolphins are exposed to between November and March in Akaroa Harbour.............................................. 256 xxiv LIST OF ABBREVIATIONS AAGR Average Annual Growth Rate AEI The Acoustic Ecology Institute AHHD Akaroa Harbour Hector’s Dolphin catalogue AIC Akaike’s Information Criterion A.R.E.V.A. Akaroa Research on the Effect of Vessel Activity project BPHDPC Banks Peninsula Hector’s Dolphin Photo-ID Catalogue BPMMS Banks Peninsula Marine Mammal Sanctuary BWR Bow riding BSS Beaufort Sea State ca. circa, approximately C.I. Confidence Interval CJS Cormack-Jolly-Seber (model) cm Centimetre COC Code of Conduct COM Commercial C.V. Coefficient of Variation d Days Dec December d.f. Degrees of freedom DIV Diving DOC Department of Conservation DW Dolphin-Watching e.g. exempli gratia, for example EIA Environmental Investigation Agency et. al. et alii, and others etc. et cetera, and other things Feb February Fig. Figure GIS Geographic Information System GLM Generalised Linear Model GOF Goodness Of Fit GPS Global Positioning System hr Hour ID IDentification i.e. id est, in other words IFAW International Fund for the Animal Welfare IUCN International Union for Conservation of Nature IWC International Whaling Commission Jan January kHz KiloHertz kg Kilograms km Kilometre km2 Square kilometre km/hr Kilometre/hour kts Knots LAC Limit of Acceptable Change List of abbreviations xxv LR Logistic Regression Ltd. Limited m Metre min Minute Mar March MFish Ministry of Fisheries MIL Milling MMPA Marine Mammals Protection Act MMPR Marine Mammals Protection Regulations MPA Marine Protected Area MRA Marine Reserves Act MSR Maritime Safety Regulations NGOs Non-Governmental Organisation NIWA National Institute of Water and Atmospheric research n Sample size nm Nautical mile Nov November NRC National Research Council s Second S South SIHD South Island Hector’s Dolphin S.E. Standard Error of the mean SOC Socialising SPUE Sighting Per Unit Effort SST Sea Surface Temperature SW Swim-With-dolphins TMP Threat Management Plan TRAV Travelling UNCED United Nations Conference on Environment and Development UK United Kingdom USA United States of America UTM Universal Transverse Mercator vs. versus, against