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. The Ecology and Molecular Ecology of the New Zealand Lesser Short-tailed Bat Mystacina tuberculata A thesis presented in partial fulfIlment of the requirements for the degree of Doctor of Philosophy in Ecology at Massey University, Palmerston North, New Zealand Brian Donald Lloyd 2002 General abstract ii GENERAL ABSTRACT The distnbution, ecology, and molecular ecology of short-tailed bats My stacina tuberculata were investigated in the central North Island, New Zealand. Special emphasis was given to providing information for conservation management of the species. Seven large populations containing between 2000 and 10 000 bats were found to persist in the area. Populations require numerous large cavities in the main trunk of mature trees for colonial roosts and are therefore restricted to extensive stands of tall old-growth forest. There was no evidence of predation by introduced mammals at roosts where the bats are most wlnerable. In summer, large group size, aggressive behaviour, and frequent movement between colonial roosts reduce the probability of predation. During winter, roosting bats remain in torpor for long periods and are vulnerable to predators, but the entrances to winter roosts are usually too small for predators to enter. Although field trials demonstrated that the bats may be at risk of secondary poisoning during pest control operations, close monitoring of a bat population during a control operation revealed no mortality. The species' intraspecific phylogeny was investigated using a 2878 bp sequence alignment from mUltiple mitochondrial genes. Six sympatric phylogroups were identified with estimated divergences of 0.93-0.68 My ago. The phylogroups do not correspond to the existing subspecific taxonomy. The phylogeographic structure and demographic history of the phylogroups were investigated using control region sequences modified by removing homoplasic sites. Phylogeographic structure was generally consistent with an isolation-by-distance dispersal model. The observed pattern is not typical of microbats, which generally exlnbit low levels of genetic structure over continental ranges. Coalescent based analyses (mismatch distnbutions, skyline plots, lineage dispersal analysis, and nested clade analysis) indicated the three phylogroups found in central and southern North Island expanded before the last glacial maximum, presumably during interstadials when Nothofagus forest was most extensive. Genetic structure within a central North Island hybrid zone was consistent with range expansion from separate refugia following reforestation after catastrophic volcanic eruptions. Discrepancies between estimates of historic and c.urrent popUlation size (> 12 million cf. 50 000) indicate the species has undergone a massive popUlation decline. Acknowledgements ill ACKNOWLEDGMENTS The study was carried out under Massey University Animal Ethics Approval 96/141 and was generously funded by the New Zealand Department of Conservation. It is a pleasure to acknowledge the many people who contnouted to this work. Firstly, I want to acknowledge the importance of each of my supervisors. Dr. Murray Potter, my chief supervisor, deserves special recognition for his unfailing encouragement and good counsel throughout the study. Other supervisors each played a crucial role. Dr. Rod Hay and Dr Claire Veltman provided essential support and encouragement during the early stages and Dr. Peter Lockhart provided the advice and expertise I needed during the final stages of the study. The study would have been impossible without my field assistants: Shirley McQueen, Tom Whitford, and Rosemary Smith. I want to thank them unreservedly for every one of those too-heavy loads of equipment they carried and all the long cold hours they persevered at mist-net sites. At various times Department of Conservation field­ stafffrom throughout New Zealand also assisted with the field work, sharing the heavy loads and long cold nights. The most enthusiastic amongst them, and the most important to me, are Bryan Williams, Nigel Holland, and John Heaphy. As well as providing assistance in the field, local offices of the Department of Conservation in many areas provided support and local knOWledge. In particular, the project could not have succeeded without support from Tongariro-Taupo Conservancy staff: Harry Keys, Cam Speedy, and John Luft I treasure the time I spent flying with Elwin Greenwood, hunting for the elusive bats. His premature death saddened me and made the world a lesser place. A number of people at the Department of Conservation head office contnouted enormously to this work. My controlling officers, Don Newman and Rod Hay, ensured the project could proceed by providing funding and support. Murray Douglas and Herman Weenink kept the electronic equipment working. Chris Edkins drafted all of the best illustrations in this thesis. I am extremely grateful to Professors David Penny and David Lambert, for their generosity in providing laboratory facilities, and to my mentors in molecular ecology: Trish McLennachan, Abby Harrison, Leon Huynen, Nickie Aitken and Peter Ritchie. Special thanks goes to the staffof the Waikato University DNA sequencing facilities for Acknowledgements IV their great service. Thanks also to Colin O'Donnell for providing tissue samples from the Eglinton Valley, and Alina Arkins for providing tissue samples from the Tararuas. The most important acknowledgments of all are to the special people in my life, friends and family. My parents started me on this road many years ago with the best of childhoods, but sadly aren't here to share this completion. My brothers, Peter and Derek, and sister Doreen are all on the other side of the world, but they remain close to my heart. My daughters Sri and Judith, now grown to wonderful women, have given me love and support that has kept me going through the hardestyears. Friends like Duncan Cunningham, Rebecca Mason, Paddy O'Dea, John Brunsden, Charlotte Sunde, and many others, have given life its rich texture. Finally, I extend my love and appreciation to Lynne Huddleston who has bought so much joy to my life. Table of contents v TABLE OF CONTENTS Page GENERAL ABSTRACT .......•.....................•........................•..•••............••...........•.....•.•.. n ACKNOWLEDGMENTS.....••....•....................••..•.......•...•............•.......................••••..•.. ill TABLEOF CONTENTS ......................................•.....................••............•...........•......... V LIST OF FIGURES...............................••..................................................•••................ XI LIST OF TABLES ..................................................................................•......••••........ XIll LIST OF PLATES ....••.••.....•.........................•.•.........•....•..............•••••..................•.•.••."XV Chapter 1: General introduction ....................................................................... 1 STUDY BACKGROUND.............................................•................................•.........•.•...•• 2 INTER-POPULATION DISPERSAL ......••..••••.........•.......................................•......•......... 3 Methods of measuring dispersal................................................................... ...... 4 THESISFORMA T... •...................................•...........•.••..................•.•......................•••••. 5 REFERENCES...................................................................................................••••...... 6 Chapter 2: Advances in New Zealand mammalogy 1990-2000: Short- tailed bats ................................................................................................................. 9 SUPERFAMILY NOCTILIONOIDEA...........................•••........................••.•.....•.•.......... 10 FAMILYMYSTACINIDAE ............•.•.............•.•..................................••...................•.. 11 GENUS MYSTACINA ............................................••.•...............•.•...........••••.....•.......... 11 Description ......................................................................................................... 11 Colonisation ....................................................................................................... 14 LESSER SHORT-TAILEDBAT ...............••••.........••.•.............................•••.......••........... 15 Field sign ............................................................................................................ 16 Daytime........................................................................................................... 16 Night-time ....................................................................................................... 17 Measurements ................................................................................................... 18 Table of contents vi Variation ............................................................................................................ 18 Distribution ....................................................................................................... 18 Habitat ............................................................................................................... 21 Food ..................................................................................................................