This electronic thesis or dissertation has been downloaded from Explore Bristol Research, http://research-information.bristol.ac.uk Author: Venugopal, Parvathy Title: An integrated approach to the taxonomy of hipposiderid bats in South Asia General rights Access to the thesis is subject to the Creative Commons Attribution - NonCommercial-No Derivatives 4.0 International Public License. A copy of this may be found at https://creativecommons.org/licenses/by-nc-nd/4.0/legalcode This license sets out your rights and the restrictions that apply to your access to the thesis so it is important you read this before proceeding. Take down policy Some pages of this thesis may have been removed for copyright restrictions prior to having it been deposited in Explore Bristol Research. 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An integrated approach to the taxonomy of hipposiderid bats in South Asia Parvathy Venugopal A dissertation submitted to the University of Bristol in accordance with the requirements for award of the degree of Doctor of Philosophy (PhD) in the Faculty of Life Sciences School of Biological Sciences January 2020 39,380 words Abstract Cryptic diversity has been well documented in several bat families and particularly in the Old-World families such as the Hipposideridae and Rhinolophidae which exhibit high levels of acoustic divergence. The genus Hipposideros is the most speciose in the family Hipposideridae and is well known for its taxonomic complexity due to the presence of several morphologically cryptic lineages. This study aims to unravel the taxonomic uncertainty in two hipposiderids, Hipposideros pomona Andersen, 1918 and Hipposideros lankadiva Kelaart, 1850 from South Asia. H. pomona from southern India has recently been identified as a distinct species. Meanwhile, all specimens from northeast India and Southeast Asia have been assigned to H. gentilis Andersen, 1918. Currently, three subspecies are recognised in H. lankadiva: H. l. lankadiva (Sri Lanka), H. l. indus (peninsular India) and H. l. gyi (northeast India and Myanmar). To date, no study has reassessed the taxonomic status of these taxa using an integrated taxonomic approach throughout their geographic extent. Therefore, an integrated taxonomic approach was applied using multiple lines of evidence, namely: morphometrics, bioacoustics and molecular phylogenetics. In addition, a presence- only modelling approach (MaxEnt) was used to better understand the geographic distribution of the targeted taxa. Results showed that H. pomona is distinct from H. gentilis sensu lato based on morphometrics, bacular and molecular data and its distribution is confined to the south of peninsular India. Hence, the recent species status of H. pomona is valid. Although there is a significant variation in the size and echolocation call frequency of H. lankadiva from Sri Lanka and northeast India- Myanmar compared with bats from mainland India, the taxon exhibited moderate to low genetic divergence in both mitochondrial and nuclear datasets. Therefore, the current subspecies status is appropriate in H. lankadiva. Species distribution models predicted that H. pomona is restricted to southern areas in peninsular India, though suitable conditions exist for its presence in Sri Lanka. The range of H. gentilis s.l. potentially overlaps with that of H. pomona in some areas of peninsular India. Models predicted that the three subspecies of H. lankadiva do not overlap in range. In conclusion, the present results reiterate the importance of using integrated approaches in bat taxonomy. i Dedication പ്രിയപ്പെട്ട വല്ല്യ, വവപ്പെയമ്മ, സൂര്യ, നീെിക്കാട്ട .... ഞാൻ ഈ രി. എ്. ഡി. തീസിസ് നിങ്ങൾക്കായി സമർെിക്കുന്നു. ഞാൻ ഈ യാപ്ത തുടങ്ങിയവൊ എൻ്്പ്പെ കൂപ്പട ഉണ്ടായിര്ുന്നവര്ാണ് നിങ്ങൾ. എന്നാൽ ഈ അഞ്ച് വർഷങ്ങൾക്ക് ഇടക്ക്, ഒര്ു യാപ്ത വരാലുԂ വ ാദിക്കാപ്പത, ഒര്ുരിടി സ്വനഹവയാർമകൾ തന്ന് നിങ്ങൾ ഓവര്ാര്ുത്തര്ായി എപ്പന്ന വിട്ട് വരായി. എങ്കിലുԂ എനിക്കെിയാԂ അകപ്പല എവിപ്പടവയാ ഇര്ുന്ന് നിങ്ങളുԂ എനിക്കായി സവതാഷിക്കുന്നുണ്ടാവുԂ. നന്ദി വല്ല്യ! സവതԂ മകപ്പള വരാപ്പല നിെപ്പയ സ്്‌വനഹി് നിെമുള്ള കുട്ടിക്കാല ഓർമ്മകൾ തന്നതിന്. നന്ദി വവപ്പെയമ്മ! കകരിടി് കളര്ിയിൽ പ്പകാണ്ട് വിട്ടതിന്, രിപ്പന്ന പ്പരവേ എന്ന് വിളി് സ്വനഹിു കലഹിതിന്. നന്ദി സൂര്യ! കാടുകയെി ഇവൊളുԂ രഠിവാണ്ടിര്ിക്കുന്ന നിവന്നാപ്പടനിക്ക് പ്പരര്ുത്ത അസൂയ ആണ് രാെു എന്ന് നിെപ്പയ സ്വനഹവത്താപ്പട രെഞ്ഞു എൻ്്പ്പെ ആത്മാഭിമാനപ്പത്ത ഉണർത്തിയതിന്. നന്ദി നീെിക്കാട്ട! നന്നായി രഠി് സവതԂ കാലിൽ നിൽക്കാൻ എപ്പന്നന്നുԂ ഓർമിെിതിന്, മുെത് വയസ് കഴിഞ്ഞിട്ടുԂ നാട്ടുകാര്ുԂ വീട്ടുകാര്ുԂ കൂട്ടുകാര്ുԂ കലയാണപ്പത്ത രറ്റി വ ാദിവൊ 'കുവഞ്ഞ ഈ രഠിത്തԂ കഴിഞ്ഞാ നല്ല് വ ാലി കിട്ടിവല്ല്' എന്ന് മുവന്നാട്ടുള്ള എൻ്്പ്പെ ീവിതപ്പത്ത രറ്റി ഒര്ുരടി മുവന്ന ആശങ്കപ്പെട്ടതിന്. (I would like to dedicate my PhD wholeheartedly to few wonderful persons of my life who had been there with me from the very outset of my journey, supporting me and applauding me throughout my thick and thin but too early bid goodbye to this world. Today when I am taking the final step in this journey my heart sinks mourning on your absence, but I wish to believe that you all are proudly embracing this achievement. I can't express enough of my gratitude towards you uncle, my godfather, for loving me as a daughter of yours and adding more colours to my childhood. I fall short of words when expressing my love and affection towards you Grandma, the strongest lady I have witnessed in my life. Those silly fights and warm hugs of yours are still missed!! I can't find enough words for you Grandpa. You were the one who has always motivated me to look beyond the stereotypes of the society and engage in the fruits of success. Last but not the least, dear Surya you are always remembered. My long journey towards this stage of PhD has always been intertwined with your generous and inspiring words which has always boosted and uplifted my morale). ii Acknowledgements My five-year journey, from a tiny village in southern part of India, to a doctorate degree from one of the most prestigious universities in the world is almost complete. A million thanks to my supervisor Prof. Gareth Jones for lending a hand when I stumbled, for picking me up when I struggled and for his words of wisdom when I thought I could not move anymore. I am proud to be a member of the Gareth Jones's Bat Lab. I am grateful to Prof. Paul Racey, whose kindness and timely intervention, brought me in touch with Prof. Gareth. I am deeply indebted to Dr. Paul Bates, who was involved in the project from the beginning. His constant presence, advice and encouragement were invaluable to me. Words fail me to express my sincere gratitude to him for his support and concern. Much love to Paul and Beatrix for looking after me whenever I came to the Bowerwood House. I am grateful to my main sponsor, the 'Commonwealth Scholarship Commission (CSC)' for the studentship and research grant that made this herculean undertaking possible. Special thanks to Dr. P. O. Nameer who introduced me to the world of bats. Thanks to Dr. Animon Illias, Dr. Praveen Karanth, Dr. Janhvi Joshi, and Dr. Felix Francis for helping me piece together a successful project proposal and CSC application. The project would have been incomplete without my collaborators Dr. Adora Thabah, Dr. Manuel Ruedi, Dr. Paul Bates, Tharaka Kusuminda and Dr. Wipula Yapa. The best parts of my work have the footprints of their wholehearted effort. I feel proud and honoured in working with them across borders and I look forward to the same in future. I cannot imagine finishing my project without the excellent training I received in lab work, data processing and analysis at the expert hands of Dr. Angelica Menchaca, Edmund Moody, Dr. Matt Zeale, Dr. Orly Razgour, Dr. Rachael, C. B., Dr. Rupert Collins, Dr. Stephen Cross, Dr. Tom Davis and Dr. Tom Williams. Their time, patience and willingness to share knowledge were extremely precious to me. I would like to express my sincere thanks and appreciation to Dr. Kailash Chandra, Dr. Gourav Sharma, Dr. Venkatraman C., Dr. Kamalakannan, M. (ZSI, Kolkata), Dr. Uttam Saikia (Northeastern Regional Centre, ZSI), Rahul Khot (BNHS, Mumbai), Roberto Portela Miguez (NHM, London) and Dr. Görföl Tamás (HNHM, Budapest) for permitting me to work on the bat collections in the respective museums. I thank Dr. Adora Thabah, Kiran Thomas, Nilantha Vishvanath, Tharaka Kusuminda Shyam Bhayya, Venkidesh, D. and Vishnu Satheesan for their immense support and dedication during the tiring field-work days in India and Sri Lanka. I also thank iii everyone who took care of me by providing a roof over my head and food on my plate, during my research travels throughout India. Harpreet Kaur, Jasmin Purushothaman, Kavitha Subin, Kumari, Mini Anil, Smrithu Mohan and Tariq Ahmed. My entire journey would have fallen apart without the unflinching support of my parents, Rema Devi and Venugopal, R. They gave their all in every choice and decision made by me, often sacrificing themselves.