i i i i Experiments in Animal Behaviour Cutting-Edge Research at Trifling Cost Raghavendra Gadagkar i i i i i i i i i i i i i i i i Experiments in Animal Behaviour Cutting-Edge Research at Trifling Cost Raghavendra Gadagkar INDIANACADEMYOFSCIENCES Bengaluru i i i i i i i i All rights reserved. No parts of this publication may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, electronic, me- chanical, photocopying, recording, or otherwise, without prior permission of the publisher. © Indian Academy of Sciences 2021 Reproduced from: Resonance – Journal of Science Education Published by: Indian Academy of Sciences Production: Srimathi M Geetha Sugumaran Pushpavathi R Reformatted by: M/s. FAST BUCKS SERVICES No. 126, DR Towers, First Floor, Above Gold Palace, Dispensary Road Bengaluru - 560001 http://www.fastbucksindia.com/ Printed at: Tholasi Prints India Pvt. Ltd. Bengaluru ISBN: 978-81-950664-7-6 i i i i i i i i Foreword The Masterclass series of eBooks brings together pedagogical articles on single broad topics taken from Resonance, the Journal of Science Education, that has been published monthly by the Indian Academy of Sciences since January 1996. Primarily directed at students and teachers at the undergraduate level, the journal has brought out a wide spectrum of articles in a range of scientific disciplines. Articles in the journal are written in a style that makes them accessible to read- ers from diverse backgrounds, and in addition, they provide a useful source of instruction that is not always available in textbooks. The Eighth book in the series, ‘Experiments in Animal Behaviour - Cutting- Edge Research at Trifling Cost’, is a collection of articles by Prof. Raghavendra Gadagkar, presently DST Year of Science Chair Professor at the Centre for Eco- logical Sciences, Indian Institute of Science, Bengaluru. These articles were orig- inally published as a series in Resonance between 2018 and 2021, and provide an excellent introduction to many interesting phenomena and studies in animal behaviour while also exemplifying that cutting-edge research can often be done at relatively low cost, a consideration especially important in India. Prof. Gadagkar is a very distinguished researcher in animal behaviour, ecology and evolution, well known around the world for his research on the evolution of sociality in prim- itively eusocial wasps, and also the Founder President of the Indian Society of Evolutionary Biologists. He is also a former President of the Indian National Sci- ence Academy, New Delhi. His research is conceptually rich and experimentally elegant, yielding important insights with simple, creatively designed experiments. He is also very well known as a superb teacher and mentor. He was one of the founders of Resonance in 1996 and also served as an Associate Editor of Reso- nance for many years after its inception. iii i i i i i i i i Foreword As always, this book will be available in digital format, and will also be housed on the Academy website. Especially in view of the neglect of animal behaviour, and evolution and ecology, more broadly, as a discipline in India, it is hoped that this book will be valuable to both students and teachers as a resource with which to supplement textbook material on animal behaviour and its evolu- tion. The book should also be useful to researchers as a convenient handbook on diverse aspects of experimental design in animal behaviour. AMITABH JOSHI Editor of Publications Indian Academy of Sciences March 2021 iv i i i i i i i i About the Author Raghavendra Gadagkar is an eminent behavioural ecologist who is currently Year of Science Chair Professor (conferred by the Department of Science and Tech- nology, India) and Honorary Professor at the Centre for Ecological Sciences, Indian Institute of Science, Bengaluru. He established an outstanding research programme at the Centre for Ecological Sciences to study insect sociobiology, which, over the last three and a half decades, has provided fascinating insights into the social evolution, social organisation, and behaviour of wasps, ants, and bees, especially the primitively eusocial wasp, Ropalidia marginata. These in- sights have resulted primarily from elegantly designed experiments, careful be- havioural observations, theoretical models, and computer simulations, with ge- netic markers and biochemical techniques used when necessary. Therefore, it is fitting that Prof. Gadagkar has written this delightful book impressing upon the reader that cutting-edge research does not necessarily require expensive or sophis- ticated equipment. Prof. Gadagkar’s interest in the social behaviour of Ropalidia marginata be- gan when he was an undergraduate student at Central College, Bangalore Univer- sity. He completed B.Sc. (Hons) and M.Sc. in Zoology at Bangalore University, and joined the Microbiology and Cell Biology Department (MCBL) at the In- dian Institute of Science (IISc), Bengaluru, in 1974 to carry out Ph.D. research on a Mycobacteriophage. Although he loved both molecular biology and animal behaviour, once he completed his Ph.D. in 1979, he chose to switch to animal behaviour (read the book to understand why) and has carried out world-class re- search using the locally present paper wasp, Ropalidia marginata. He demon- strated the importance of demographic factors in social evolution with his theory of Assured Fitness Returns, and the inadequacy of genetic relatedness alone in v i i i i About the Author explaining the social evolution of eusocial insects. He and his students have also found self-regulating age polyethism of workers, pheromonal inhibition of worker reproduction by queens, and seamless, conflict-free queen successions through queueing, suggesting that this wasp is more socially advanced than the typical primitively eusocial wasp. Prof. Gadagkar’s outstanding research has been recognized by numerous awards in India and abroad - the Shanti Swarup Bhatnagar Prize, B.M. Birla Sci- ence Prize, the Third World Academy of Sciences Award in Biology, and Cross of the Order of Merit of the Federal Republic of Germany – to name a few. He is an elected Fellow of the Indian National Science Academy, of which he was also President, and other Indian science academies, and International member of the National Academy of Sciences, USA, the American Academy of arts and Sciences and the German National Science Academy Leopoldina, among others. He is a Non-resident Permanent Fellow of the Wissenschaftskolleg zu Berlin. He is also Founding President and Fellow of the Indian Society of Evolutionary Biologists and has held numerous other academic and advisory positions. Prof. Gadagkar is a teacher par excellence, cultivating logical and critical thinking and a love for science amongst students/audience in his classes and the hundreds of outreach talks he has given. Further, his book Survival Strategies simply and succinctly explains concepts in behavioural ecology to a wide audi- ence. He was also involved in setting up the pedagogic journal Resonance to cater to undergraduate students. He has mentored a very large number of research students, many of whom have gone on to establish independent research groups. Prof. Gadagkar also pioneered the establishment of the Centre for Contemporary Studies at IISc to impart an appreciation for the interdisciplinarity of knowledge. The present volume is a compilation of a series of articles by Prof. Gadagkar that appeared in Resonance. Although focused on experiments in animal be- haviour, the message of the book applies to other subjects as well; all are en- couraged to read it. T N C Vidya, JNCASR, Bengaluru vi i i i i Contents Preface ix 1. How Wasps Find Their Nests 1 2. Do Bees Have Colour Vision? 15 3. How Do Ants Find the Shortest Path? 29 4. How Do Bees Estimate the Distance Flown? 43 5. How Do Ants Estimate the Distance Walked? 55 6. Why are Male Wasps Lazy? 69 7. How Do Wasps Decide Who Would Be the Queen? – Part 1 87 8. How Do Wasps Decide Who Would Be the Queen? – Part 2 107 9. Why Do Wasps Fight? – Part 1 131 10. Why Do Wasps Fight? – Part 2 145 11. Fighting Fish—Does Experience Matter? 165 12. Male Frogs Sing for Sex but Why Don’t They Do Their Best? 191 vii i i i i i i i i Contents 13. Harmless Snakes Mimic Venomous Snakes 213 14. Cuckoos Lay Their Eggs in Others’ Nests 241 15. Why Do Parents and Offspring Quarrel? 277 16. Cutting-Edge Research at Trifling Cost 311 viii i i i i i i i i Preface In The Ten Most Beautiful Experiments (2008) *, the science writer George John- son laments that “Science in the twenty-first century has become industrialized. The experiments so often celebrated in the newspapers—sequencing the genome, proving the existence of the top quark, discovering a new planet by analyzing the wobble of a distant star—cost millions of dollars. They generate terabytes of data to be analyzed by supercomputers: calculating factories spewing so much heat that they are equipped with cooling stacks that consume the energy of small towns. The experiments are carried out by research teams that have grown to the size of corporations”. Reviewing Johnson’s book in the New York Times† Peter Dizikes, then an in- dependent writer, said “Johnson’s lively book nicely evokes the lost world of the tabletop experiment”, and asked “But are all remaining advances really beyond the reach of individual hands and minds, as he supposes? Might we still attribute major ideas to ingenious individuals, even if the ideas are tested by teams?”. Fi- nally, Dizikes concluded sardonically that “Certainly, Johnson is entitled to his nostalgia. Still, if lone scientists rarely push knowledge forward today, they rarely impede it, either.” I think that all of this pessimism is misplaced.