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THE CHIMPANZEES OF RUBONDO ISLAND APES SET FREE Josephine Nadezda Msindai and Volker Sommer The Chimpanzees of Rubondo Island How did a random batch of chimpanzees come to populate a small island in Tanzania where apes had never lived before? Combining information gathered from feldwork, laboratory and archival research, this book tells the unique story of chimpanzee babies taken from their forest homes in West- Central Africa and sold to European zoos and circuses, to then be shipped to Lake Victoria and set free on Rubondo Island. These founder animals learnt what to eat, how to build nests, to breed and raise young – ultimately forming a chimpanzee-typical fssion-fusion society that today is thriving. The authors compare the ecology, behaviour and genetics of the Rubondo population with communities of wild chimpanzees, providing exciting insights into how our closest relatives adjust to changing environments. At the same time, a reconstruction of the historical context of the Rubondo experiment refects on its chequered colonial heritage, and the introduction is viewed against current threats to the survival of apes in their natural habitats. The book will be of interest to scholars and professionals working in primatology, animal behaviour, conservation biology and postcolonial studies. Josephine Nadezda Msindai obtained a BSc in Biological Sciences from King’s College London, UK (2005), an MSc in Primate Conservation at Oxford Brookes University, UK (2008) and a PhD in Anthropology from University College London (UCL), UK (2018). The Chimpanzees of Rubondo Island is based on her doctoral work that included almost two years of feld research in Tanzania. Volker Sommer is Professor of Evolutionary Anthropology at University College London (UCL), UK. He obtained his PhD in Anthropology at Göttingen University, Germany (1985) and has conducted extensive primatological feld studies in India (since 1981), Thailand (since 1984) and Nigeria (since 1999). The Chimpanzees of Rubondo Island Apes Set Free Josephine Nadezda Msindai and Volker Sommer First published 2023 by Routledge 4 Park Square, Milton Park, Abingdon, Oxon OX14 4RN and by Routledge 605 Third Avenue, New York, NY 10158 Routledge is an imprint of the Taylor & Francis Group, an informa business © 2023 Josephine Nadezda Msindai and Volker Sommer The right of Josephine Nadezda Msindai and Volker Sommer to be identifed as authors of this work has been asserted in accordance with sections 77 and 78 of the Copyright, Designs and Patents Act 1988. All rights reserved. No part of this book may be reprinted or reproduced or utilised in any form or by any electronic, mechanical, or other means, now known or hereafter invented, including photocopying and recording, or in any information storage or retrieval system, without permission in writing from the publishers. Trademark notice: Product or corporate names may be trademarks or registered trademarks, and are used only for identifcation and explanation without intent to infringe. British Library Cataloguing-in-Publication Data A catalogue record for this book is available from the British Library ISBN: 9780367422196 (hbk) ISBN: 9781032329178 (pbk) ISBN: 9780367822781 (ebk) DOI: 10.4324/9780367822781 Typeset in Sabon by Deanta Global Publishing Services, Chennai, India Contents List of Illustrations vi List of Tables x Acknowledgements xi Introduction xiii 1 Creating a Wilderness: The Making of an Island National Park 1 2 The Founder’s Odyssey: Captured, Caged, Released 23 3 Rubondo Island: Weather, Forests, Wildlife, Humans 52 4 Bound to Be Wild: Sociality and Ranging 86 5 Embedded: Mastering a New Environment 115 6 Apes in the Anthropocene: Lessons from a Maverick Release? 161 Bibliography 201 Appendix: Publications about Rubondo Island, Its History and Wildlife 225 Index 233 Illustrations Figures 1.1 View of Rubondo Island from a hilltop 1 1.2 S outh-western Lake Victoria region, Tanzania, East Africa 2 1.3 B ernhard Grzimek’s iconic TV series Ein Platz für Tiere commemorated on a stamp of the Deutsche Bundespost released for his 100th birthday in 2009 10 1.4 T anzanian President Nyerere and Bernhard Grzimek in 1963 14 2.1 A chimpanzee on Rubondo in her night nest – half a century after apes were released on the island 23 2.2 C himpanzee Caroline (pictured with keeper Carl Stemmler), kept at Zoo Basel, was one of the animals acquired by Grzimek to be shipped to Rubondo 31 2.3 C himpanzees arriving at Dar es Salaam port make front page news 33 2.4 R eleasing chimpanzees onto Rubondo. (a) Crates are taken to shore. (b) Chimpanzee crate opened by Sinclair Dunnett. (c) One of the frst chimpanzees set free, with Sinclair Dunnett on top an empty crate 35 2.5 T eam members of the author’s research team (Jul13) 45 2.6 P hylogenetic relationships among chimpanzees and bonobos, including haplotypes of the Rubondo chimpanzees 49 3.1 A erial view of Rubondo Island National Park’s coastal area and forest cover 52 3.2 R ubondo Island National Park and important locales of its main island 54 3.3 A nnual rainfall 1900–2006 at meteorological stations on Lake Victoria. Top line Entebbe (Uganda), bottom line Mwanza (Tanzania) 55 Illustrations vii 3.4 W eather on Rubondo, 2004–14. (a) Yearly values. (b) Averaged across 11 years 58 3.5 T otal annual rainfall over the 18-yr period 1996–2013 59 3.6 I nterior forest on Rubondo. (a) Section with trees of smaller diameter. (b) Trees with tangling lianas. (c) Open growth with buttressed trees 62 3.7 D iameter at breast height (DBH) of transect trees 64 3.8 F ruits and fowers of transect plants on Rubondo 69 3.9 F lowering schedule of transect plants (Jan–Dec13) and annual rainfall 71 3.10 F ruit-bearing transect plants (Jan–Dec13) 73 3.11 T ransect plant fruiting schedule (Jan–Dec13), expressed as a comparison between (i) fruit index and (ii) percentages of fruiting transect plants 73 3.12 P roportions of primate and non-primate frugivores as assessed from feedings remains under transect trees 74 3.13 L arge animals on Rubondo Island: (a) Nile crocodile; (b) hippopotamus; (c) guereza; (d) vervet monkey; (e) otter; (f) elephant; (g) sitatunga 75 3.14 Illegal and legal fshing on Rubondo. (a) Poachers with their catch apprehended by TANAPA rangers (1994). (b) Tourists with their catch (2012) 77 4.1 Rubondo chimpanzee female 86 4.2 Phylogenetic relationships amongst the Hominini 88 4.3 The genus Pan: Geographical distribution of chimpanzee subspecies and bonobos 90 4.4 Two main study regions in the island’s north, with trail grids for chimpanzee surveys and straight-line botanical transects (Chapter 3) 94 4.5 Chimpanzee party sizes (Apr12–Mar14). (a) Day party frequencies. (b) Night party frequencies. (c) Annual distribution of day parties. (d) Annual distribution of night parties 97 4.6 Locations of evidence for chimpanzee presence (Apr12– Mar14). (a) Tree night nests (n = 1,224). (b) Ground night nests (n = 22). Daytime sightings (n = 146) 108 4.7 Home-ranges calculated via a minimum convex polygon (MCP), adjusted to exclude Lake Victoria 109 4.8 Main areas of chimpanzee activity as discerned during different study periods: 1966–84, 2002–08, 2012–14 111 viii I llustrations 5.1 N est, roughly one month old, built by a Rubondo chimpanzee 115 5.2 N est types on Rubondo. (a) One of the rare ground nests (2013). (b) Three arboreal nests (indicated by arrows) built in the same tree 118 5.3 H eight distribution classes of arboreal nests during three study periods: 1994, 2002–07, 2012–14 120 5.4 N ight nest height as a function of tree height (n = 690) 122 5.5 N est tree DBH (diameter at breast height) as a function of tree height (n = 803) 122 5.6 S hares of preferred species of nesting trees compared against their occurrence on random transects (Chapter 3) 126 5.7 A ltitude at which nests were built (n = 801) as a function of monthly rainfall 127 5.8 M ean (+/– SD) density of tree and liana fruit patches on Jan–Dec03 transects in relation to rainfall 131 5.9 G rouping of obtained DNA sequences of the ciliad Troglodytella abrassarti in chimpanzee subspecies and bonobos 142 5.10 Q uantifcation of beta diversity in chimpanzee microbiomes via PCoA plots. (a) Weighted Unifrac distances. (b) Unweighted Unifrac distances 147 5.11 B oxplot of weighted Unifrac distances for chimpanzees, based on pairwise-calculated distance values 148 5.12 P otential cultural traits of Rubondo chimpanzees. (a)A-frame grooming with overhead hand-clasp. (b) Consumption of a sitatunga antelope; note pattern of hide. (c) Buttress-beating with rock; note mark on trunk of Ficus tree, and rock lying at base. (d) Discarded grass stems used for insect-fshing 156 6.1 R ubondo chimpanzee 161 6.2 D evelopment of great ape numbers kept in sanctuaries across Africa 165 6.3 C aretaker feeding orphaned chimpanzees at Tacugama Sanctuary in Sierra Leone 167 Box Figures B1.1 Gustl Anzenberger watching a burrow of a carpenter bee 19 B1.2 Markus Borner 21 B2.1 Monica Borner with one of her children 41 B3.1 Guido Müller – on yet another rainy day on Rubondo 60 Illustrations ix B3.2 J ames Leonard with Josephine N. Msindai 65 B4.1 M ichael A. Huffman on Rubondo 93 B4.2 L iza Moscovice with her team of trackers, including Joseph Mgwesa 98 B5.1 K lara Petrželková during feldwork 150 B5.2 Mwanahamisi Issa Mapua (r., holding a child), with feld assistant Giulia Graziani (m.) and parl warden Hobokela Mwamjengwa (l.) 154