Ecology and Conservation of Bat Species in the Western Ghats of India
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Ecology and conservation of bat species in the Western Ghats of India Claire Felicity Rose Wordley Submitted in accordance with the requirements for the degree of Doctor of Philosophy The University of Leeds School of Biology September 2014 i The candidate confirms that the work submitted is her own, except where work which has formed part of jointly authored publications has been included. The contribution of the candidate and the other authors to this work has been explicitly indicated below. The candidate confirms that appropriate credit has been given within the thesis where reference has been made to the work of others. Chapter Two is based on work from “Wordley, C., Foui, E., Mudappa, D., Sankaran, M., Altringham, J. 2014 Acoustic identification of bats in the southern Western Ghats, India. Acta Chiropterologica 16 (1) 2014 (In press)’. For this publication I collected over 90% of the data, analysed all the data and prepared the manuscript. John Altringham and Mahesh Sankaran edited the manuscript. Eleni Foui collected the remainder of the data and prepared figures 2.1 and 2.3. Divya Mudappa assisted with data collection and manuscript editing. This copy has been supplied on the understanding that it is copyright material and that no quotation from the thesis may be published without proper acknowledgement. © 2014 The University of Leeds and Claire Felicity Rose Wordley The right of Claire Felicity Rose Wordley to be identified as Author of this work has been asserted by her in accordance with the Copyright, Designs and Patents Act 1988. ii The lesser dog faced fruit bat Cynopterus brachyotis eating a banana. iii Acknowledgements I thought this would be the easiest bit of my thesis to write, but finding the words to express my heartfelt gratitude to so many people – and wondering how many I’ve forgotten to thank – is far harder than I could have imagined. I’ll start by thanking John Altringham, who was patient, encouraging and believed I could do this far more than I ever did! John helped give me the courage to make decisions in the field, taught me that making mistakes isn’t the end of the world and (most importantly) always shared his climbing videos, malt whisky and macarons. Thanks to my co-supervisor Mahesh Sankaran who made the connections between Leeds and Valparai possible, helped me when I got snarled up in statistics and read chapters for me at lightning speed when I needed it. Thanks to everyone at the Nature Conservation Foundation in Valparai who all provided critical support. Anand Kumar sorted out so many logistical challenges, from which field assistants were free to which field sites were full of elephants. Divya Mudappa helped me sort out equipment, permission and plans when I was feeling a little lost and took some great photos of our wet, muddy fieldwork. ‘Sridhar’ Shankar Raman was a calm presence in the storm, giving valuable advice on writing and statistics, as well as cracking me up with his dry sense of humour. Divya and Sridhar also helped me to get involved in communicating about bats through articles and posters, with the help of Pavithra Sankaran and John Altringham. Ganesh Raghunathan and ‘Jegan’ Panchapakesan Jeganathan made sure I didn’t get squashed by any elephants and provided many laughs! Aditya Malgaonkar helped out in field, cooked amazing paneer parathas and was a great bird watching companion, despite saying I looked part Neanderthal. Huge thanks to Stella Kumar for feeding me, making me lovely ginger tea, fixing my trousers, teaching me Tamil, and making me giggle. I would have been lost without Sathish Kumar, who taught me to handle large bitey fruit bats and drove safely away from charging elephants and drunken motorcyclists! Anand Kumar and Dinesh T.’s knowledge of the forests, toughness and willingness to learn about bat detectors, harp traps and cameras was invaluable. Asai and co. were always remarkably cheerful about driving us about at night with piles of equipment and muddy boots! My assistants from Europe were hugely helpful - Aurelie Laurent, Ruth Angell and Kate Parker all deserve iv massive thanks, and deserving of many beers were Emma Rigby, my rock during the first field season who taught me heaps about catching bats, and Sarah Proctor who greatly improved my bat detector skills (and invented the sexy leech sock dance). Thanks to all my friends in India who made me feel welcome and showed me bits of their country; Ranjini Murali, Rahul and Chayant Gonsalves, Radhika Timbadia and Priyana Johnson. Special thanks to the people of Iyerpadi who made India feel like home by feeding me, taking me to festivals, teaching me Tamil and playing hopscotch with me until I dropped; Archana, Divya, Gayathri, Murugan, Palaniammal and all the rest. You are like family to me and will always have a special place in my heart. Funding was provided by NERC (Natural Environment Research Council), the University of Leeds and UKIERI (UK India Education and Research Initiative). Thanks to landowners for permissions: Tamil Nadu Forest Department, the Tamil Nadu Electricity Board, Peria Karamalai Tea Company, Bombay Burmah Trading Corporation, Tata Coffee Ltd, Parry Agro Industries Ltd, Altaghat Estate and Thalanar Estate. Thanks to Eleni Foui for being brilliant on my first trip to India and sharing data, and to Anita Glover and Anna Berthinussen from whom I have learned a lot catching bats in the Yorkshire Dales, Poland and Latvia. Thanks to Felicity Edwards who introduced me to functional diversity and Peter Steward who helped with some of the graphs, Richard Gunton, Chris Hassall, John Graham and Amanda Bretman who all gave me advice on statistics when I needed it. Chloe Bellamy, Chiara Polce and Ellie Brown gave me some good ArcGIS/MaxEnt advice, and Chris Scott shared his automated extraction software. My friends in Leeds have supported me when the going got tough, mostly with merciless ‘character building’ teasing and beer – they are all awesome mates who I hope to stay in touch with for years to come. Becky Surtees and Corinne Shaw have kept me sane(ish) by being brilliant climbing partners. Jamie Kelly has had to put up with more stressing about my thesis than anyone else, and deserves huge thanks for cooking dinner when I was climbing, walking me even in bad weather and for being patient with all my woes. Finally my family have put up with me having no interest in anything other than bats and my PhD for years, missing many special occasions and showing up sporadically wanting food, money or lifts to airports – and yet they’ve encouraged me to follow my passion. Thank you. v Abstract The Western Ghats of India are a globally important biodiversity hotspot, but around 90 % of the land has been converted to agriculture. Little is known about how bats respond to the conversion of native rainforest to different plantation types in the Western Ghats. This thesis examines the response of bats to coffee and tea plantations, and to riparian habitats, in the southern Western Ghats. Most bat assemblages in the tropics have been studied by catching bats, but many studies have shown that catching alone can give biased and incomplete results. In order to use acoustic data as well as catching data in this landscape I made a library of the echolocation calls of fifteen echolocating species in the landscape. Comparisons of the data from each method showed that combining catching and acoustic data gave the most complete picture of the assemblage, but that acoustic data alone detected more species than catching data alone. Acoustic and catching data were used to build habitat suitability models for ten species. Scales of 100 m – 500 m were the most important for predicting bat presence. Several species showed a positive response to habitats containing native trees and habitat richness, and a negative response to tea plantations and distance to water. Coffee plantations did not differ significantly from forest fragments in terms of bat species or abundance, but did differ in species composition. Bat assemblages in coffee plantations were functionally very similar to those in forest fragments. Tea plantations had the lowest bat species richness of all habitats and differed in species composition from all other habitats. Bat assemblages in tea plantations had lower functional richness and specialisation than other habitats, and the bats remaining were open- adapted species. Rivers with riparian corridors did not have significantly greater bat species richness than rivers without corridors, but differed functionally in several ways. Rivers without riparian corridors had reduced functional specialisation and functional divergence. Rivers with riparian corridors supported more forest adapted species than rivers without riparian corridors. vi Table of contents Acknowledgements .......................................................................................................... iv Abstract ............................................................................................................................ vi Table of contents ............................................................................................................ vii Table of figures ................................................................................................................ xv Table of tables ................................................................................................................. xx Chapter One: General Introduction .................................................................................