From So Simple A Beginning... The Expansion Of Evolutionary Thought #1 #2 From So Simple A Beginning... The Expansion Of Evolutionary Thought Compiled and Edited by T N C Vidya #3 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, mechanical, photocopying, recording, or otherwise, without prior permission of the publisher. c Indian Academy of Sciences 2019 Reproduced from Resonance–journal of science education Published by Indian Academy of Sciences Production Team: Geetha Sugumaran, Pushpavathi R and Srimathi M Reformatted by : Sriranga Digital Software Technologies Private Limited, Srirangapatna. Printed at: Lotus Printers Pvt. Ltd., Bengaluru #4 Foreword The Masterclass series of eBooks brings together pedagogical articles on single broad top- ics 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 readers from diverse backgrounds, and in addition, they provide a useful source of instruction that is not always available in textbooks. The sixth book in the series, ‘From So Simple A Beginning... The Expansion Of Evolu- tionary Thought’, is a collection of Resonance articles about scientists who made major con- tributions to the development of evolutionary biology, starting with Charles Darwin himself, collated and edited by Prof. T. N. C. Vidya of the Evolutionary and Organismal Biology Unit of the Jawaharlal Nehru Centre for Advanced Scientific Research, Bengaluru. Prof. Vidya is a widely recognized expert on Asian elephant social organization, behaviour, and ecology and, in addition, teaches and carries out research in diverse areas of ecology, evolution and behaviour. Her research is an elegant and effortless blend of painstaking field ecology and conceptually rich theory in several areas within ecology and evolutionary biology. She has also been a con- tributor of pedagogical articles to Resonance, and has been an Associate Editor of Resonance since 2017. 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 evolutionary biology 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 evolution. The book should also be useful to experts as a convenient handbook on diverse aspects of the historical development of evolutionary thinking. Amitabh Joshi Editor of Publications Indian Academy of Sciences March 2019 #5 #6 About the Editor T N C Vidya, PhD, is currently Associate Professor and Chair of the Evolutionary and Inte- grative Biology Unit (earlier the Evolutionary and Organismal Biology Unit) at the Jawaharlal Nehru Centre for Advanced Scientific Research, Bengaluru. With her interest already kindled in wildlife biology, Vidya enrolled in the Integrated PhD programme of the Indian Institute of Science, Bengaluru in 1997. She investigated the role of ecology in shaping the intestinal parasite loads of elephants for her master’s thesis. For her PhD, Vidya pioneered the study of the phylogeography of the Asian elephant, drawing attention to the role of Pleistocene climatic oscillations in shaping patterns of gene flow among populations. At the Jawaharlal Nehru Centre for Advanced Scientific Research, Vidya has established a flourishing research programme focusing on behavior, ecology, evolution and conservation bi- ology, while extending her taxonomic reach beyond elephants to include other large mammals such as gaur, sambar, chital, dhole, and Nilgiri tahr, and birds such as mynas. Vidya’s research seamlessly combines methods of traditional ethology with those of molecular population ge- netics, and also some theory. One of her studies, for example, involves monitoring over 800 individually identified elephants in Nagarahole and Bandipur Tiger Reserves in Southern In- dia, to understand the effect of ecology on social organization and behavior. Another example involves using elephant dung samples to extract DNA and estimate population sizes and sex ratios - parameters with obvious management value. Vidya has held prestigious research fellowships at the Indian Institute of Science, Columbia University, USA, and Stellenbosch University, South Africa, as also the Ramanujan Fellowship of the Department of Science and Technology, Government of India. She is a recipient of the Indian National Science Academy medal for young scientists, the Prof. C N R Rao Oration Award Lecture, and has represented the country in the Indo-American Frontiers of Science Forum and in the Young Leaders Programme of the Science and Technology in Society Forum, Japan. She has also served as invited member of the IUCN SSC (World Conservation Union’s Species Survival Commission) Asian Elephant Specialist Group since 2007. Vidya is a passionate teacher and committed to spreading interest in animal behavior and evolution among students and the general public, having mentored a large number of under- graduate, masters and PhD students and delivered many lectures for general audiences. She is also actively associated editorially with several Indian and international journals. In particu- lar, as Associate Editor of Resonance – journal of science education, published by the Indian Academy of Sciences, Vidya is continuing her endeavours to inculcate enthusiasm for organis- mal biology in Indian academia. Raghavendra Gadagkar Centre for Ecological Sciences, Indian Institute of Science, Bengaluru March 2019 #7 #8 Contents 1. Revolutions in Evolutionary Thought: Darwin and After 1 Renee Borges 2. The Origin of Species after 150 Years – One Hundred and Fifty Years without Darwin are Enough 15 Vidyanand Nanjundiah 3. Not Saint Darwin 35 John S Wilkins 4. Co-discoverer 47 Vidyanand Nanjundiah 5. Ernst Heinrich Philipp August Haeckel: 16 February 1834 to 09 August 1919 53 Amitabh Joshi 6. Vignettes of Haeckel’s Contributions to Biology 63 Amitabh Joshi 7. Walter Frank Raphael Weldon (1860–1906) 83 Amitabh Joshi 8. Weldon’s Search for a Direct Proof of Natural Selection and the Tortuous Path to the Neo-Darwinian Synthesis 91 Amitabh Joshi 9. From ‘Particulate Factors’ to ‘Designer Genes’: A Hundred Years of Genetics 107 Amitabh Joshi 10. Sir R A Fisher and the Evolution of Genetics 111 Amitabh Joshi 11. J B S Haldane 115 Vidyanand Nanjundiah 12. Haldane’s Contributions to Biological Research in India 121 Partha P Majumder 13. Sewall Wright: A Life in Evolution 125 Amitabh Joshi ix #9 14. The Shifting Balance Theory of Evolution 135 Amitabh Joshi 15. Dobzhansky and Evolutionary Cytogenetics: Pioneering Evolutionary Studies with Chromosomes 143 Bimalendu B Nath 16. Theodosius Dobzhansky: A Great Inspirer 149 Amitabh Joshi 17. Conrad Waddington and the Marriage of Genetics, Development, and Evolution 153 John Tyler Bonner 18. C H Waddington, Canalisation and Genetic Assimilation 155 Vidyanand Nanjundiah 19. J Maynard Smith: From Engineering to Evolution 165 Amitabh Joshi 20. The Logic of Animal Conflict 169 Raghavendra Gadagkar 21. John Maynard Smith (1920 – 2004): “One of the Last Grand Evolutionary Theorists of the 20th Century” 171 Vidyanand Nanjundiah 22. Bill Hamilton – The Greatest Darwinian Since Darwin 179 Raghavendra Gadagkar 23. The Origin and Resolution of Conflicts in Animal Societies: The Case of the Bees and the Birds 183 Raghavendra Gadagkar 24. What Do Ethologists Wish to Know? 197 Raghavendra Gadagkar 25. Nikolaas Tinbergen: The Careful Scientist 201 Sindhu Radhakrishna 26. Supernormal Stimuli and Responses 207 T N C Vidya x #10 27. Amotz Zahavi: Tales of a Swamp and Peacock Tails 215 Manjari Jain 28. Evolutionary Biology Today: 1. The Domain of Evolutionary Biology 223 Amitabh Joshi 29. Evolutionary Biology Today: 2. What do Evolutionary Biologists do? 231 Amitabh Joshi Author Profile 243 xi #11 #12 Preface I am delighted to present to you this compilation of articles, published previously in Reso- nance, as a Biology Masterclass. Arranged roughly in chronological order, these articles give us insights into the lives and work of several great thinkers who made outstanding contributions to the idea of evolution and the growth of evolutionary biology. As this book includes only pre- viously published articles, it is not a comprehensive list of all great evolutionary biologists, but does include many of them. We begin the book with a broad account, by Renee Borges, of the history of evolutionary thought prior to Darwin, Darwin’s thoughts on evolution through natural selection and on in- heritance, and the subsequent Neo-Darwinian synthesis. She then describes the thinking about the levels of selection, kin selection and reciprocal altruism, and the origin of sexual reproduc- tion. This chapter is followed by Vidyanand Nanjundiah’s description of the course of events that led to the publication of the Origin of Species and an exposition of its contents. Nanjundiah explains the crux of evolution through natural selection and clarifies common misconceptions. As Nanjundiah points out, it is impressive that Darwin formulated his theory without a knowl- edge of either the origin of variations or the mechanism of heredity. However, as he also points