Humans and Lions: Conflict, Conservation and Coexistence 1138558036, 9781138558038

Citation preview

“This is the natural companion to the thoughtful and elegant Ivory: while the themes broadly overlap and compliment there is also plenty of fresh research to form a panorama as wide as the African sky. These books will be debated for years to come and form touchstones for present and future generations.” — Jasper Humphrys, Director of External Relations of the Marjan Centre for War and the Non-Human Sphere, King’s College, University of London, UK “Professor Somerville has written the definitive history of the relationship between lions and humans. This meticulously and exhaustively researched book starts sixty million years ago with the evolutionary origin of Carnivores and ends with developments in lion conservation in late 2018. In between, it examines the long history of conflict between the two apex predators, documenting in agonizing detail the lion’s long spiral toward extinction at the hands of man, and the current efforts of a handful of conservationists to reverse the decline. This will be the standard reference on lion conservation for years to come.” — Laurence G. Frank, Living With Lions Project Director and research associate in the Museum of Vertebrate Zoology, University of California, Berkeley, USA “Keith Somerville eruditely rips the scales from his readers’ eyes to unveil the interacting blights that have beset lions for millennia – trade, conflict, hunting – pills all bitterly coated in the proliferation of people. Where does that leave lions? In a precarious mess.” — David Macdonald, Director of WildCRU, University of Oxford, UK “As human populations in Africa and India continue to surge into the 21st Century, placing ever increasing demands on land and resources, the future of lions, one of the world’s most iconic species, hangs in the balance. To meet the challenge of conserving these magnificent but demanding creatures in the wild, we need to grasp the complex history and nature of this issue, meticulously researched and comprehensively presented in this important book.” — Michael ‘t Sas-Rolfes, Oxford Martin Fellow, Oxford Martin Programme on the Illegal Wildlife Trade, University of Oxford, UK

Humans and Lions

This book places lion conservation and the relationship between people and lions both in historical context and in the context of the contemporary politics of conservation in Africa. The killing of Cecil the Lion in July 2015 brought such issues to the public’s attention. Were lions threatened in the wild and what was the best form of conservation? How best can lions be saved from extinction in the wild in Africa amid rural poverty, precarious livelihoods for local communities and an expanding human population? This book traces human relationships with lions through history, from hominids, to the Romans, through colonial occupation and independence, to the present day. It concludes with an examination of the current crisis of conservation and the conflict between Western animal welfare concepts and sustainable development, thrown into sharp focus by the killing of Cecil the Lion. Through this historical account, Keith Somerville provides a coherent, evidence-based assessment of current human-lion relations, providing context to the present situation. This book will be of interest to students and scholars of environmental and African history, wildlife conservation, environmental management and political ecology, as well as the general reader. Keith Somerville is a Member of the Durrell Institute of Conservation and Ecology at the University of Kent, UK, where he is also a professor and teaches at the Centre for Journalism. He is a Senior Research Fellow at the Institute of Commonwealth Studies and a Fellow of the Zoological Society of London, UK.

Routledge Environmental Humanities Series editors: Scott Slovic (University of Idaho, USA), Joni Adamson (Arizona State University, USA) and Yuki Masami (Kanazawa University, Japan)

Editorial Board Christina Alt, St Andrews University, UK Alison Bashford, University of New South Wales, Australia Peter Coates, University of Bristol, UK Thom van Dooren, University of New South Wales, Australia Georgina Endfield, Liverpool, UK Jodi Frawley, University of Western Australia, Australia Andrea Gaynor,The University of Western Australia, Australia Christina Gerhardt, University of Hawai’i at Mānoa, USA Tom Lynch, University of Nebraska, Lincoln, USA Iain McCalman, University of Sydney, Australia Jennifer Newell, Australian Museum, Sydney, Australia Simon Pooley, Imperial College London, UK Sandra Swart, Stellenbosch University, South Africa Ann Waltner, University of Minnesota, USA Jessica Weir, University of Western Sydney, Australia International Advisory Board William Beinart, University of Oxford, UK Jane Carruthers, University of South Africa, Pretoria, South Africa Dipesh Chakrabarty, University of Chicago, USA Paul Holm,Trinity College, Dublin, Republic of Ireland Shen Hou, Renmin University of China, Beijing, China Rob Nixon, Princeton University, Princeton NJ, USA Pauline Phemister, Institute of Advanced Studies in the Humanities, University of Edinburgh, UK Deborah Bird Rose, University of New South Wales, Sydney, Australia Sverker Sorlin, KTH Environmental Humanities Laboratory, Royal Institute of Technology, Stockholm, Sweden Helmuth Trischler, Deutsches Museum, Munich and Co-Director, Rachel Carson Centre, Ludwig-MaximiliansUniversität, Germany Mary Evelyn Tucker,Yale University, USA Kirsten Wehner, University of London, UK The Routledge Environmental Humanities series is an original and inspiring venture recognising that today’s world agricultural and water crises, ocean pollution and resource depletion, global warming from greenhouse gases, urban sprawl, overpopulation, food insecurity and environmental justice are all crises of culture. The reality of understanding and finding adaptive solutions to our present and future environmental challenges has shifted the epicenter of environmental studies away from an exclusively scientific and technological framework to one that depends on the human-focused disciplines and ideas of the humanities and allied social sciences. We thus welcome book proposals from all humanities and social sciences disciplines for an inclusive and interdisciplinary series. We favour manuscripts aimed at an international readership and written in a lively and accessible style. The readership comprises scholars and students from the humanities and social sciences and thoughtful readers concerned about the human dimensions of environmental change.

Humans and Lions Conflict, Conservation and Coexistence

Keith Somerville

First published 2020 by Routledge 2 Park Square, Milton Park, Abingdon, Oxon OX14 4RN and by Routledge 52 Vanderbilt Avenue, New York, NY 10017 Routledge is an imprint of the Taylor & Francis Group, an informa business © 2020 Keith Somerville The right of Keith Somerville to be identified as author of this work has been asserted by him 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 identification and explanation without intent to infringe. British Library Cataloguing-in-Publication Data A catalogue record for this book is available from the British Library Library of Congress Cataloging-in-Publication Data Names: Somerville, Keith, author. Title: Humans and lions : conflict, conservation and coexistence / Keith Somerville. Description: Milton Park, Abingdon, Oxon ; New York, NY : Routledge, 2019. | Series: Routledge environmental humanities | Includes bibliographical references and index. Identifiers: LCCN 2019007124 (print) | LCCN 2019008446 (ebook) | ISBN 9781315151182 (eBook) | ISBN 9781138558021 (hbk) | ISBN 9781138558038 (pbk) | ISBN 9781315151182 (ebk) Subjects: LCSH: Lion—Conservation—Africa. | Human-animal relationships—History. Classification: LCC QL737.C23 (ebook) | LCC QL737.C23 S5837 2019 (print) | DDC 639.97/9757—dc23 LC record available at https://lccn.loc.gov/2019007124 ISBN: 978-1-138-55802-1 (hbk) ISBN: 978-1-138-55803-8 (pbk) ISBN: 978-1-315-15118-2 (ebk) Typeset in Bembo by codeMantra

Contents

List of figure and tables viii Foreword by David W. Macdonald ix List of abbreviations xxv Introduction 1 1. Lion-human coexistence and competition from the Pleistocene to modern humans 6 2. Domestication, settlement and the effects on lions 29 3. Lions from the 14th century to colonial occupation 53 4. Hunting, conservation and the decline of the lion in colonial Africa and Asia 77 5. Contemporary coexistence and conflict in Africa 117 6. The ups and downs of Southern Africa’s lions, and the importance of the trophy hunting debate 169 Index 223

Figure and tables

Figure 5.1 African range map 118

Tables 1.1 Chart of the overlapping time spans of predator and hominin genera in Africa 16 4.1 Population estimates for the Gir region 104 5.1 Lion numbers by region and by source 119 6.1 Lion numbers by region and by source (South Africa) 170

Foreword Lion conservation: a history of could to should

The past was far from promising, the future none could tell. (My father, perhaps adapting G.K. Chesterton)

Fairness matters, something understood by both Androcles and lions over 2,000 years ago. It is also one of diverse considerations that might decide how contemporary people should treat the dwindling numbers of modern lions. Fairness, as confirmed by evolutionary psychologists, is one of a handful of evolved, adaptive, foundations of human behaviour, and indeed that of some other social mammals. Perhaps lions attend to fairness, and if they do those with long memories probably wish Androcles, Daniel and their ilk were around today. Androcles, as Pliny relates, was a Syracusan who drew a thorn from a lion’s paw, a favour reciprocated, as every game theorist might predict, when the lion later declined to kill Androcles in the Roman arena. Lions with a similar sense of justice spared Daniel (Daniel 6:6–24) but killed his false accusers. Lions, and specifically how we ought to behave towards them, turn out to be a very important 21st century topic to which I’ll return below, but meanwhile, if you need to be reminded of these edifying stories, and a myriad more, then Keith Somerville is your man. I used to think Keith was remarkable for his daunting capacity to remember and to pronounce with intimidating assurance, the names of the people and places that populate Africa’s history (the topic of an earlier book in his expanding canon). Having read every syllable of Humans and Lions: Conflict, Conservation and Coexistence, I now appreciate that this same capacity for remembering, assembling and interpreting compendious detail enables him to see far back into history (actually to the Pleistocene history of encounters between people and lions – which generally didn’t go so well as Androcles’s), to see sideways

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across the breadth of Africa’s continental history and, as a crescendo, to reflect on how those deep and broad perspectives might educate our thinking on what could, or even what should, happen next. This is an important topic of the moment because 2019 is no longer a time to study history in order to appreciate how it has, and might again, repeat itself: the stage is nowadays so radically changed for both lions and for people that another repeat of history will be terminal. The understanding informed by Keith Somerville’s remarkable book is that most of the history of modern peoples’ dealings with lions (as metaphors for biodiversity as a whole) must, at all costs, not be repeated. Further, to avoid that deathly repetition requires an understanding far beyond biology; an understanding of the transdisciplinary intersection of people and wildlife that I term Conservation Geopolitics. The 21st century is a new world, and it necessitates a new way of coexisting with nature. For this we need new science, applied within a new ethic that expands the circle of connectedness and coexistence across people, places and even more widely to encompass wildlife, not least lions.

History What, then, does Somerville tell us of the intersection of lions and history, beyond such intriguing snippets that the entire skeleton of a lion was found in the 1430 BCE tomb of Tutankhamun’s nurse? His text reveals that, more often than you might have paused to think about, quantum changes in the affairs of people have transformed, indeed often derailed, the fate of lions. Obviously, the invention of firearms was a big moment for all concerned and brought lions into the deathly power-games of, amongst other blood-spattered commodities, ivory and slaves. Colonialism catalysed a step-change in human impact. A typical Somervillian detail is that by 1888, around 100,000 guns a year were being traded through Zanzibar alone. Both local and European marksmen inflicted exponential mortality on lions (for the most part, where I write lions it is shorthand for wildlife, or at least megafauna). European occupation went into full-swing after the Berlin Conference of 1884–5 divided the continent between European powers. During the late 19th and early 20th centuries, as the cattle industry was expanding, for example into what became Namibia and Botswana, the colonial authorities employed the !Xoon (San) to hunt lions, leopards and hyenas whose predation on livestock was increasingly infuriating, as it continues to be today. Modern preoccupations with human-­ wildlife conflict over stock losses are far from new (for “there is a fox in my chicken house” comfortable European armchair conservationists should imagine “there is a lion in my cattle pen”). By the way, the fox analogy is strikingly apt (and resurfaces later in the context of sport hunting) – both fox and lion are particularly vexing when they engage in surplus killing, which both are most likely to do when triggered by sloppy husbandry; Somerville cites Wright, writing in the 1890s about Mozambique, who records one lioness that

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broke into a kraal and killed 70 goats in one night. Anyway, whether the triggers were pulled in anger or excitement, guns have been bad for lions – one amongst dozens of similar observations documented in Humans and Lions stems from 1934 when a Captain Shortridge of present-day Namibia toured Caprivi, Damaraland and the Kaokoveld and concluded the “lion range had retreated steadily in the face of European settlement”. To bring this up to date, the unwelcome corollary of the recent recovery of lion numbers in Namibia is that more are available to kill cattle – in November 2017, 171 goats and sheep were killed in a single attack in the Torra Conservancy, just a week after an attack in which 86 were killed by the same large pride – the cost to the Namibian farmers of these losses was N$450,000 (£24,822). If colonial times represent a slippery slope for lions, the period between Sudan’s independence on 1 January 1956 and Angola’s on 11 November 1975 may have brought relief to the many territories unshackled from colonialism, but it did nothing to slow the downward acceleration of lion numbers. History continued to set the environmental stage, and while those who established National Parks (NPs) imagined they would protect wildlife, war is no respecter of good intentions; as noted by Robbie Burns in conversation with a mouse, the best laid plans o’ mice and men gang aft agley. During the Rwandan civil war and genocide, the Akagera NP, home to 250 lions in 1992, was invaded by tens of thousands of hungry refugees who, understandably, killed and ate the lions’ ungulate prey; the lions that weren’t starved out were killed by the soldiers and cattle herders whose encampments followed – by 2012 none survived. In Uganda, lions were widespread at independence, but the civil war there also led to disastrous declines, worsened by incursions of armed poachers during conflicts in the neighbouring Democratic Republic of Congo (DRC) and South Sudan. The ravages of Idi Amin’s army, and then those of the Tanzanian troops that deposed him, further worsened the wildlife outcome: in Rwenzori NP, 14,000 of 46,400 large mammals were killed. In other protected areas up to 75% of prey species were killed for meat. Angola was at war from the launching of the liberation struggle in 1961 to the death of the rebel leader of the National Union for the Total Independence of Angola (UNITA), Jonas Savimbi, in February 2002. Alongside devastating humanitarian consequences, lions and wildlife were decimated. In Cuando Cubango, where UNITA was based, they and the South African army killed most things that could be eaten (buffalo and antelopes), sold (rhino horn, ivory) or, indeed, that moved (lions). Somerville reports that parts of Angola and northern Namibia became recreational hunting grounds for South African politicians and military leaders. Rural communities were displaced and relied on bushmeat to survive. The loss of ungulate prey during the Angolan war had cascading effects on lion numbers. Today, impoverished people have access to arms left over from the war and hunt for bushmeat to survive. Paul Funston (Somerville meticulously cites all his sources), a noted surveyor of lions, concluded that the war had directly and indirectly devastated Angola’s lions. The Mozambican liberation war, also against the Portuguese, exploded in 1964, raged

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for a decade, and was then replaced by Rhodesian and South African-sponsored insurgency by the Mozambique National Resistance movement against the ruling FRELIMO party. Floods of refugees fled the fighting, soon followed by economic migrants fleeing poverty. Between 1960 and 2005 an estimated 1,338,000 Mozambicans sought to enter South Africa through Kruger National Park, with 370,000 caught by the authorities. This time the boot was on the other foot: one, probably exaggerated, report estimates that 13,380 of these people were killed by lions. Even after the Mozambican war, in 1996–7 at least 11 economic migrants were killed by lions as they crossed the border into Kruger. Humans and Lions deconstructs the interwoven forces majeures that have shuddered through the fate of lions – consider disease: in 1889, Italians imported rinderpest in cows transported from India to the Eritrean port of Massawa, killing millions of wild and domestic ungulates with huge consequences for people and lions. Between 1906 and 1912 a huge campaign against rabies sought to eradicate lions – and leopards, cheetahs, hyenas and wild dogs – on European-owned farms.

Kill or be killed – trade Recently, a concern widely preoccupying conservationists has been that there is a perilous upsurge in trade in lion body parts, both for local use and for export. A modern view might think this is a new threat, but through the lens of Somerville’s history any such myopia is corrected: people have been trading the bits and pieces of lions (and indeed in the days of the Roman venationes, whole lions) for centuries. Arab traders were a force in mediaeval Africa, and a flux of Arab-­ Shirazi, Swahili and Indian merchants operating in the 7th–11th centuries along the Indian Ocean coast sold wildlife products along with spices and slaves to West Asia, India and China. In due course, and perhaps nervously, firearms were traded in return: while ivory was the greatest prize, I’d bet lions’ teeth, claws and pelts were traded too. The British, Dutch, French and Portuguese set up trading posts along the West African coast as dispersing Europeans headed south and, in 1487, the Portuguese explorer Bartolomeu Dias anchored at M ­ osselbaai on the southern Cape. In his wake, a decade later, Vasco da Gama sailed into the Indian Ocean and so Portuguese commerce developed with East Asia; ports sprung up on the Mozambican coast, trading for gold, ivory and, surely, the skins and parts of lions amidst other pelts bartered from the peoples of Mozambique, Malawi and Zimbabwe. By the end of the 16th century, Dutch, English and French had followed suit. In 1649, a Dutch expedition wintered in Table Bay and the Dutch East India Company opened the door for, three years later, Jan van Riebeeck to establish the Cape trading settlement. There is little evidence that any of these people shared 21st century qualms about either the conservation or ethics of trading the body parts of lions. Today, these qualms are prominent because of the legal trade in farmed South African lions, whose bones are exported to China and Vietnam for use in traditional medicine, hocus pocus

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health tonics and jewellery. This fear, growing in the minds of conservationists, surfaced in a 2017 report by TRAFFIC which concluded that wildlife in the Central African Republic (CAR) and Democratic Republic of Congo (DRC) was threatened by poaching, for bushmeat, ivory and skins. In this case the modern mix of poverty and conflict is ignited by invasions of heavily armed Mbororo (Fulani) herders with connections to smuggling and poaching gangs supplying lion skins, bones and claws amongst wildlife products shipped to China, Laos and Vietnam; meanwhile, bushmeat hunting by local people in southern and eastern CAR and northern DRC empties the lions’ larder. Elsewhere, over 7,800 ­captive-bred lions languish on South African farms. Their original destiny as the targets of canned hunts was substantially derailed by a 2016 US restriction on the import of captive-bred lions as hunting trophies. A number of these lions, as their financial value for their owners fall, find an alternative use in Chinese traditional medicine (where, since 2008, their bones have been exported to be used as a substitute for those of tigers). The magnitude and consequences of this trade, originally documented by the 2016 WildCRU/TRAFFIC report, Bones of Contention, raise perplexing questions at the interfaces of conservation and animal welfare and of market regulation and ethics. Following the global impact of the 2015 film Blood Lions, the trade in captive-bred lions has risen to new heights of prominence in public and political debate in South Africa, prompting its description at a 2018 colloquium of the parliamentary committee on environmental affairs as “a practice that has gained a reputation of being the most controversial subject in the conservation industry”. Somerville’s characteristically up-to-date account ends with the November 2018 recommendation from the same committee that the government reconsider the rules governing the breeding of captive lions for hunting and bone harvesting, amid fears that the trade is damaging the country’s image internationally and could affect tourism. Looking forwards, the lion trade story is as yet unfinished: a WildCRU team, tasked with understanding this complex system, continues to explore extensive and interconnected issues of empirical uncertainty and ethical concern. Contemporary trade in lion body parts thus provides a case study that illustrates the moral maze, or more accurately moral morass, that confronts modern, transdisciplinary conservationists. Last year I met a lion farmer who, understandably looking for ways of bolstering her beleaguered industry, talked enthusiastically of diversifying her product range to produce and promote lion bone cakes (a gluey cake rendered from boiled tiger bones is already popular in Vietnam Cao hổ cốt). Indeed, the increasing production of this “cake” in ­A frica from lions and leopards exemplifies the globalised nature of wildlife trade demand. It is said that you can’t have your cake and eat it; it will likely test conservationist’s mettle to confront tough decisions that weigh the needs of peoples’ cultures and traditions with the survival of species. As today’s decision-makers are forced to confront issues of criminality, economics, civil liberties and social justice, Keith Somerville concludes that those with the responsibility of deciding what ought to happen “are truly caught between a rock and a hard place”.

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Kill or be killed – man-eating Early people surely did not share the modern enthusiasm for conserving lions, or other dangerous carnivores: they were our rivals and predators. Indeed, famed carnivore biologist Hans Kruuk cleverly suggested that one reason these big ­carnivores hold our attention today is that we evolved, albeit with a very different perspective, to pay special attention to them. Because people all too often exaggerate problems with carnivores, and because that exaggeration has too often been applied to the danger to life and limb (most notably the Little Red Riding Hood view of wolves, that very rarely hurt people), modern conservationists, especially those who don’t have them in their own backyard, may too readily dismiss the risks of living with lions. You won’t make that mistake again once you’ve read Humans and Lions, peppered as it is with historical cases of lions eating people. The man-eaters of Tsavo are surely the most notorious for killing 135 railway workers – as it happens, authors of the day record that the lions’ prey had been depleted by the railway construction workers, the route followed a centuries-old slave-caravan route where lions were accustomed to eating dead or ailing slaves, and both lions were themselves debilitated. But being disabled is not a prerequisite for a lion to kill people, being hungry is reason enough. Somerville documents how lions have eaten people throughout colonial history (and, of course, since people evolved) – the first Cape settler to survive an attack was Wouter Moster in 1657; shortly afterwards, on the 12th September, Van Riebeeck noted that a Dutch soldier was found on the aptly named Lion Mountain at Cape Town: “the brain pan had already been bitten off”. In the British Uganda colony, a number of people had been “carried off by lions”; some individuals north-east of Kampala were noted for “unusual and unprovoked truculence” and along the roads walked by workmen to Lake Victoria 12 people were killed every month in 1924–5 (the government reacted by trying to poison the lions, but found hunters to be more effective). In Tanganyika in 1935, the game ranger for Lindi district reported that man-eaters were “exceedingly troublesome” and had recently killed 35–40 people. Over several years, 145 people had been killed by lions in the district. Somerville reports that at one stage five man-eaters were hunting as a group; none of the lions shot appeared to be in bad condition. Fourteen suspected man-eaters were shot in the region. At Singida in central Tanganyika, about 200 people were killed in an outbreak of man-eating in the 1920s, another outbreak erupted in the 1940s. Somerville notes that many local people claimed the lions were witchdoctors who could transmogrify into lions  – ­interestingly, exactly the belief expressed in WildCRU’s current study area in Ruaha to justify the killing of lions (they weren’t really lions – just despicable transmogrified neighbours). Between 1941 and 1946 a colonial official, Rushby, recorded that lions around Njombe (close to Tanzania’s Kipengere Mpanga Game Reserve) killed between 1,000 and 1,500 people, but a more temperate reporter, Patterson, reckoned the tally was closer to 246 (Patterson recorded a further 103 people killed in 1946–7, all generally attributed by locals to transmogrified witchdoctors).

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Humans and Lions reveals an interesting repeat of history: the colonials thought some of this man-eating arose because large numbers of wild pigs raided fields in which villagers perforce guarded their crops at night, and thus bumped into pig-hunting lions. Exactly this logic was deployed by famous lion biologist, Craig Packer, who studied the cases of 563 Tanzanians killed and 308 injured by lions between 1990 and 2004, mainly between Selous and the Mozambican border. The modernity of all these fatalities should make them vivid to the non-­A frican reader. While Tony Blair was prime minister of Britain and some readers of this Foreword were sitting in cinemas enjoying the Lord of the Rings trilogy, between 1990 and 2007, 99 citizens of Rufiji district, and 175 of Lindi were being eaten by lions. In terms of the value of life, something we awkwardly but necessarily must consider (as in the UK do the National Health Service (NHS), or the National Institute for Health and Care Excellence (NICE), for example), the ruling Chama cha Mapinduzi Party in Tanzania offered a reward of 5,000 shillings (about £4 at the time) as a bounty on each lion killed. One thing leads to another, and in the matter of proximate and ultimate causes, the pig numbers grew, and proximity of lions to people increased, as wilderness was turned to cultivation and other wild prey disappeared.

Stock-raiding and conflict I have mixed feelings that my biblical namesake, David, slayed a lion which had killed a sheep in his father’s flock (Samuel 1 17:34–36). Nonetheless, this uncomfortable link makes Somerville’s recurring historical point: human-wildlife conflict is not new. His deep slice into ultimate causes rivetingly traces the origins of plant cultivation leading to mixed farming economies, as Bantu-­speaking peoples spread from west-central Africa into Congo and the western Rift Valley and eventually on to East and Southern Africa. Bantu agro-pastoralists settled in the eastern half of southern Africa about two thousand years ago. They grew sorghum and millet, established permanent homesteads, raised livestock, and forged iron. Humans and Lions details how a predominantly pastoralist economy developed after 1000 BCE in Kenya and Tanzania’s savanna and Rift Valley regions. By 500 CE, pastoralism was expanding, and in the Zoutpansberg area of northern Transvaal communities were building homesteads of circular huts around central cattle pens to protect against large predators. Speke’s 1854 expedition documented strong enclosures designed to protect camels and other stock from attack by lions as far north as northern Somalia. Protective bomas, the conservationist’s 21st century excitement, have a long and far-flung history. These early Africans had noticed lions: there are carved lion heads and lionesses in bas-relief on stones at Axumite settlements from modern-day Eritrea and northern Ethiopia, and clues that they hunted lions. Over about three millennia, right up to the modern period, expansion of cultivated areas, livestock herds and human settlements transformed habitats to the disadvantage of wildlife and, particularly, predators like lions. The chapters on the unfolding agro-environmental

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story that made human-wildlife conflict inevitable also document that the Bantu migrants assimilated iron-working skills from Nilo-Saharan peoples who had moved south, and used them to increase the effectiveness of crop cultivation, clearing woodland and, I’d guess, of killing predators. The colonials walked right into this conflict with predators, and have been battling ever since. In December 1652, Van Riebeeck reported the first ­European case of stock predation: on the 12th of January 1653 a lion killed a sheep and was driven off with musket fire; on the 23rd January, “lions tried to storm the kraal”. The die was cast. With the compendiousness readers will by then expect of Somerville, unfolding chapters cite dozen upon dozen evidences that lions are founder members of today’s human-wildlife conflict landscape. Amongst a myriad snippets, I enjoyed that to encourage food production for the war effort in 1914–18 the British South Africa Company paid European farmers 10 shillings for every lion killed. Somerville’s interviews in Kenya suggest that between 1946 and 1952, one Laikipia game warden shot 434 lions “on control” and some individuals killed over 300 lions apiece to protect cattle ranches in Kenya in the 1970s and 1980s. In the chapter on Namibia, Andersson is quoted: “The natives highly rejoiced at the successful termination of the hunt, for this lion had proved himself to be one of the most daring and destructive ever known, having … killed upward of 50 oxen, cows and horses.” In the chapter on Sudan a British traveller, Hoskins, reported in 1835 that lions attacked livestock and camels. Vocabulary is revealing, Andersson and Hoskins both spoke of areas “infested with lions”. Another colonial, Rainsford, predicted that as game was depleted lions would become “more destructive to domestic beasts and to man”. He was right. Firearms are lethal, but poisoning is worse. The 2018 BBC1 Attenborough documentary, Dynasties, featured the poisoning of the famous Maasai Mara Marsh Pride, an all-too-common outcome resulting from conflict with people. Characteristically up to date, Somerville reports the arrest in May 2018 of three men from the Serengeti accused of poisoning or shooting 19 lions since 2015. In WildCRU’s Ruaha project in Tanzania, in February 2018, Amy Dickman records how a collared lion was poisoned after killing a cow; she tells me five other lions and 75 vultures died. In the Chobe and Linyanti areas of northern Botswana, lions frequently fall victim to poison applied to carcasses of elephants killed for their ivory, in order to prevent lions, hyenas and vultures revealing the location of the carcasses to anti-poaching patrols. In Botswana, by the way, between 2000 and 2002 reported losses of livestock to lions totalled 588 animals (leopards reportedly killed 300 more, but were less annoying insofar as they tend to take smaller stock). Poisonings are obviously linked to livestock predation, but there are also less obvious links to the illegal wildlife trade. Poisoned lion carcasses in Ruaha habitually have the paws and tails removed, but where these by-products then travel to (and how much they incentivise poisonings in the first place) is largely unknown. The extent to which such occurrences link into the wider web of contemporary international lion trade is uncertain, but provides a

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pressing challenge for the trade-investigator who must, in a recurrent interdisciplinary theme, keep their finger on the pulse of multiple conservation concerns and cultural happenings. Having worked previously on foxes, I am deeply struck by the parallels with lions (and you can check characterisations of both in the mediaeval bestiaries). Those with even a glancing acquaintance with conflicting attitudes to foxes in the UK will appreciate the resonance in the words of professional hunter, ­Sydney Downey: “[farmers] always loathed lions, classing them as vermin. Hunters have always looked upon lions in terms of impulse, zest, and challenge, as a sort of high-spot”. In colonial East Africa on unprotected farmland, lions were killed in such large numbers by pastoralists and white farmers that, by 1960, Downey presciently feared that lion numbers “could drop below the point of recovery, making these animals virtually extinct” in both Kenya and Tanzania. Fox hunters argue that their sport saves farmers stock losses. Chief Timex Moalosi of Sankuyo, Botswana, might sympathise with this proposition, insofar as he told Keith that lion predation on his community’s livestock has worsened since 2014 when a hunting ban ended the selling of hunting quotas to safari operators (which had previously earned his community $600,000 a year). Pages of evidence lead Keith Somerville to join the chorus of biologists that “human-lion conflict is a major problem across the region, deriving from rapidly increasing populations, the desire to raise more stock and encroachment on the lion estate which that entails”. What can be done? In answering this question, Keith Somerville documents the remarkable achievements of the current generation of lion conservation biologists, including such WildCRU pioneers as Andy Loveridge with his mobile bomas in Zimabwe, Amy Dickman and her porridge project in Tanzania, Guy Western and Alayne Cotterill in Kenya, and Hans Bauer in Somalia amongst a pantheon of dedicated and talented people – read the book. Amongst this cohort he dwells on the remarkable Flip Stander (who I tinglingly recall landing me in his tail-dragger in the middle of remotest Damaraland to watch desert lions). Flip founded the Desert Lion project in 1997, reputedly stimulated by the death of a large male needlessly shot while feeding on a whale carcass on Namibia’s Skeleton Coast. The home ranges of the desert prides are huge, with females having territories from 2,500 to 12,400 km 2 and males from 6,000 to 40,000 km 2. Yet again, conflict with farmers is the biggest threat to their survival. Stander relocated 75 stock-raiding lions (49 occasional cow-killers and 26 habitual ones); 22 returned to livestock areas, most were killed by herders. Between 1999 and 2017, retaliatory killing by farmers was the main cause of death for desert lions. Can conservationists make a difference? Having feared there were no lions left in Kunene in 1991, Stander’s commitment documented in Humans and Lions is surely much of the reason that by 2005 the region supported 96–165 lions. Nonetheless, between 2000 and 2010, 47 radio-collared lions died – 20 were shot or poisoned as a result of conflict with farmers. Five males, known as the Five Musketeers – film stars of The Vanishing Kings – were all killed in 2016 and 2017 by cattle farmers.

xviii Foreword

Somerville fairly draws his sources from all sides of the fence, and doesn’t pull his punches in quoting them. Founding father of the Namibian conservancy movement Garth Owen-Smith specifies the dilemmas, writing recently that there “are now too many lions in some parts of the Kunene region […] Trying to save the lions that are killing livestock […] will not serve the interests of conservation in the region […] because their present high numbers inhibit the recovery of gemsbok, zebra and kudu populations, which is essential to create a more balanced predator/prey relationship in the future … it has caused many local farmers to rethink whether conserving wildlife is a benefit or liability to their livelihoods”. Keith went on to quote farmer Andreas Ndakukamo, who lost 16 cattle to lions in April and May 2017, and complained that the Ministry had dithered in paying compensation (N$1,500 (£83.50) per head of cattle killed by a predator). He warned that if lion attacks continued, “We will kill them. We will protect our cattle, and no longer wait for the ministry to come … We will not rest until they are all dead.”

Land loss and protected areas The collected, and published, wisdom of the most informed lion biologists in the world is that, at the root of all threats, lies habitat loss and degradation. This refrain is a numbingly familiar drumbeat behind almost every major conservation issue in the world. Writing from the UK, where the losses of natural habitat were so long ago and so complete, that the meaning of that loss may be drained of its colour, Somerville’s account makes the bereavement vivid. Consider the unfamiliar bleating of the first Merino sheep to graze African grass in 1830. By 1890 there were 12 million of them, and their success was the death knell for huge herds of springbok, cauterising the annual migration of hundreds of thousands of them from the northern Cape and southern Kalahari to the Karoo. The arrival of sheep also marked the declaration of a vendetta against lions, wild dogs, leopards, caracal and staggering, if oddly unmourned, numbers of jackals. In July 1857, Richard Burton set off inland from the coast at Kaole (kitted out in Zanzibar by Omani Arab traders), chastened by warnings of night-time attack by lions and hyenas; instead of the thronging wildlife he was expecting, he went for days without seeing game, lamenting “[it] has melted away before the woodman’s axe and the hunters’ arrows”. It is interesting to think about culpability when, although hindsight renders the image starkly, it was surely apparent even as it happened that colonial development was, from the mid-17th to the mid-19th centuries, wiping out huge herds of grazers, removing the natural prey of lions, and then killing lions to remove the threat to livestock. Just like foxes in Britain, lions were viewed schizophrenically as both vermin and a sporting quarry, an ambivalence that ultimately, as the resource dwindles, leads to opposing priorities between farmer and hunter. Ironically, out of this tension arose an antidote to habitat loss: the idea of game reserves. National parks, protected areas, and the foundations of modern conservation arise

Foreword  xix

from the model provided by 19th century private shooting estates (from which African communities, much like the rural working class in Britain, were largely excluded, sometimes forcibly removed and prohibited from traditional hunting). Lions, categorised as vermin, dwindled fastest due to their role as predators of livestock, which elided also into a massive hunting toll. In the Serengeti in the late 19th and early 20th centuries, hundreds of lions might be killed by hunters on a single safari. In British territories an influential player, then as now, was the Society for the Preservation of the Wild Fauna of the Empire (now Fauna and Flora International), formed in 1903 following a meeting of colonial powers which drew up the Convention for the Preservation of Wild Animals, Birds and Fish in Africa (signed in London on 19 May 1900), calling for the conservation of endangered species and habitats. Nowadays, in Britain the managers of reserves protecting ground-nesting birds still decide to kill predators, and in 1900 in much the same spirit, Schedule 5 of the convention called for the reduction in the number of lions and other predators to preserve ungulate species which had declined since the arrival of Europeans. Consider the first warden of Kruger and Sabi NPs, James Stevenson-Hamilton (who served from 1902 until 1946), who found little game, and blamed hunting by both Steinacker’s Horse (a British unit posted there during the Boer War) and local people who had lost their stock to rinderpest. Similarly, he noted that in the absence of natural prey the lions were eating livestock and people. Rising to the call of Schedule 5, between 1903 and 1927 Stevenson-Hamilton oversaw the killing of 1,272 lions, 1,142 wild dogs, 660 leopards, 269 cheetahs and 521 hyenas. Kruger was elevated to National Park status in 1926, and the purpose of conservation began to metamorphose as it became clear that tourists would pay to see lions there. Conservation has not yet fully emerged from its chrysalis, and conversations about the purpose of conservation (and how to deliver it) continue. Much thinking in the colonial period was predicated on the assumption that wildlife could be protected only by keeping it safely separate from people, implicitly indigenous people. This view is far from exhausted, but overlooks the pre-19th century coexistence of wildlife and (admittedly much smaller num­ bers of ) people in African savannas. Today biologists – notably WildCRU’s Guy Western and Peter Tyrrell – point to the richness of biodiversity beyond protected areas and the capacity of pastoral people, such as the Maasai of Kenya’s Southern Rift, to coexist with wildlife.

Trophy hunting Encounters with dentists are notoriously unwelcome, but never in recorded history has one caused such global anguish as that between Walter Palmer, a ­M innesotan bowhunting dentist, and a Zimbabwean lion called Cecil. I direct the team that had been tracking the wanderings of Cecil for the decade before this historic encounter, so I felt first-hand the shock waves of global interest and outrage. While my colleague Andrew Loveridge attended to the bio-political

xx Foreword

ramifications in Zimbabwe, in Oxford we dealt with the media – from morning to night for eight days. At its zenith, I had 243 requests for press, radio and TV interviews backed up. Subsequently we documented that in 125 languages, there had, on the 28th July 2015 been 12,000 mentions of Cecil or the WildCRU in print media and 87,500 hits on social media. I judge this to be the greatest expression of global interest in a wildlife conservation topic in history, and in just one night it prompted an estimated four million people to visit WildCRU’s website (www.wildcru.org). I suspect that most of these people were not merely concerned with the fate of one lion, or holding an opinion (the vast majority, disgust) about one hunter; rather, I think their interest in Cecil and WildCRU’s work was an expression of a concern for wildlife more generally, and a preoccupation with how the human enterprise can and ought to live alongside nature in the 21st century. Less weightily, and mightily rectified by Keith Somerville’s scholarly book, it was plain that few people had realised that sport hunting of lions existed nowadays, far less that it had done for millennia, and I guess that almost none realised the extraordinary tallies that had been killed in historical times. Somerville details how, long before there were dentists there were lion hunters. In Greek mythology, Hercules’s first labour was to kill the Nemean Lion, which he eventually managed to strangle – thereby avoiding the problem posed by its impenetrable pelt (a property that was to later prove very useful when worn as a cloak). Three thousand years ago, Amenophis III (who ruled Egypt between 1352 and 1330 BCE) chalked up 100 lions in hunts. Alexander the Great is depicted hunting lions, a symbol of his courage and power. The Assyrian kings, who preceded Alexander, hunted lions between the 9th and 7th centuries BCE in Iraq and Syria and right up to the border of Turkey (where lions may have survived until the end of the 19th century). One king boasted: “I am Assurbanipal, the King of the World, the King of Assyria! For my regal amusement I have caught the Desert King by his tail, and on the instructions of my helpers, the Gods Ninib and Nergal, I have split his head with the twohanded sword.” Then, as now, killing lions for sport was interwoven with reprisals for their depredations; Assurbanipal (who ruled 668–627 BCE) justified his lion hunting, “They constantly kill the livestock of the fields, and they spill the blood of men and cattle. The herdsmen and the supervisors are weeping; the families are in mourning. The misdeeds of these lions have been reported to me … I have penetrated their hiding places and destroyed their lairs.” Humans and Lions describes how the Assyrians hunted lions from chariots with a charioteer, and spearmen to guard the king’s back, accompanied by “a whole host of horsemen, foot soldiers and kennel men who, at the right moment, unleashed the mastiff-like hunting dogs”. The Roman arena developed a particular approach: Pliny the Elder wrote that the first recorded lion contests in the arena were staged under ­Lucius around 80  BCE when, he says, 100 maned lions took part in fights. Under Pompey, games were held involving 600 lions, to Julius Caesar’s paltry 400. In ­Tunisia around 1249–54 CE, King Louis of France hunted lions with a

Foreword  xxi

crossbow during the crusades, but this pastime was not solely a European prerogative. Whether it’s the 16th century Mughal Emperor Babur depicted hunting, or his 17th century successor Jahangir pacifying (a political allegory) lions, hunting lions was also a preoccupation on the subcontinent. Lebna Dengel (1508–40) of the Ethiopian Solomonic Dynasty and his soldiers were enthusiastic lion hunters. The Zulu king Shaka was particularly keen too, killing driven lions with spears and axes. We lament the modern plummeting of lion numbers, some of it due to over-hunting. By the 1890s Swayne wrote that Somaliland’s hunting grounds “have lost some of their value now that so many lions have been shot”, although he adds the notable caveat that no one who had lived in the region could have sympathy for the lions because of their regular attacks on livestock and people. Kenyan settler Sir Alfred Pease (who saw 60 shot in four months in 1896–7) estimated that during 12 years at the end of the 19th century, over 1,000 lions were killed by Europeans in Somaliland. In 1900 a hunter, Charles Peel wrote: “owing to the ruthless slaughter of females by so-called sportsmen, the lion is becoming extremely scarce within 150 miles of the coast … They are totally extinct in the Golis Range, where but ten years ago they made the valleys reecho with their magnificent music. North of the Golis Range in Guban there is not a single lion track to be seen”. At the other end of the continent, the Boer settlers were avid hunters: Somerville mentions Kotje Dafel and Petrus Jacobs who reputedly killed over 100 lions each. In the tsetse-free Kenyan highlands, hunters would hunt from horseback or lions would be driven by scores of beaters towards waiting guns. When the British were given the League of Nations mandate over Tanganyika after World War I, they allowed African communities to hunt under a system of licences to preserve game while allowing stock protection. In 1929 the Serengeti was declared a Complete Game Reserve, in which hunting was banned: Somerville notes the anger of the local Nata, Ikoma, Sukuma and Musoma communities, who had hunted there for centuries prior to colonial occupation. The Kenya-based professional hunter Denys Finch Hatton, of Robert Redford Out of Africa fame, called for an end to the “orgy of slaughter” in Tanganyika, where in 1925 the Game Department concluded “that it will be necessary in the near future to limit the number of lion that may be killed on a Full Licence … otherwise this animal will soon cease to exist in some areas”. The detail, somehow mesmerising, is remarkable – read the book. A characteristic Somervillian work, it captures the interrelatedness of threats facing lions – for example in Malawi, where the “lion population was reduced under colonial rule by a combination of hunting, the spread of tea and tobacco farming and the steady growth in the human population”. So much for lion hunting in the past – what now? Before making up your mind read Chapter 6 to discover for yourself the suffocating tangle of considerations. Personally, I had to face this question in 2016 when I was asked by Rory Stewart, a British government minister, to evaluate the evidence that trophy hunting of lions has, or could, impact the distribution and abundance of lions – for better or worse. Had I at

xxii Foreword

that time read Keith’s book, I might have offered a longer historical perspective, but even reviewing contemporary evidence revealed some debilitating gaps. It has also revealed that, at least in many of the countries into which lion trophies are currently imported, large sections of society regard hunting lions for sport as an ethically inappropriate activity for the 21st century. Others take the opposite view, most notably people who actually have to live with lions in their back gardens. The proposition that rather than being a threat, hunting lions contributes significantly to their conservation through the maintenance of wild habitat, and that its cessation would worsen the species’ already deteriorating status is crucially relevant to both the discussion and its consequences. Providing facts, and identifying important gaps, is important insofar as policies applied impulsively could have perverse consequences for lion conservation. In that case, even those implacably opposed to lion hunting on ethical grounds might favour a ‘journey’ rather than a ‘jump’; that is, if society judged trophy hunting lions unacceptable, but also concluded that it benefited lion conservation, then this dilemma might be approached via a journey to find ways of replacing the benefits of hunting before jumping to end them. But the evidence is not (yet) available to draw that conclusion. Indeed, whilst evidence is necessary it is not sufficient to make decisions on conservation policy because, as in other political matters, good decisions rest upon wise judgement beyond the facts. Wise decisions are especially challenging because modern wildlife conservation strives to find mutual advantage between the well-being of wildlife and the people who live, often with difficulty and in poverty, alongside it. The ‘lion estate’ is the area of land occupied by wild lions. Any human action that diminishes the extent of that estate might be considered as inimical to lion conservation; any action that maintains or increases it might be considered a benefit. Where sport hunting sits in this balance is unresolved, and the answer seems likely to be different in different places, but Keith Somerville’s remarkable synthesis shows, at least, that its history is very long, profligate and, through the lens of 21st century conservation, often sickening.

Decline and fall So, Keith Somerville eruditely rips the scales from his readers eyes to unveil the interacting blights that have beset lions for millennia – trade, conflict, hunting – pills all bitterly coated in the proliferation of people. Where does that leave lions? In a precarious mess. Although I live in a country where no lion has prowled for 125,000 years, Herodotus reported that there were lions in Thrace in his lifetime (484–430 BCE). They survived in Greece until the 5th century BCE, and in Palestine until around the 12th century and into the 20th century in parts of Syria, Iraq and Iran (where a dead lion was reputedly found in 1943). As for North African Barbary lions, successively the Ottomans and then French colonial authorities offered bounties for killing lions, and amongst the last was shot near Taddert

Foreword  xxiii

in Western Morocco in 1942. But surely this litany of loss is from a distant, Philistine past? Considering their iconic primacy, from Landseer’s resplendent lions holding court over Trafalgar Square to MGM’s emblematic lion roaring (actually it’s a dubbed tiger’s voice) at audiences globally, surely modern people treasure wild lions as much as we revere their icons? After all, in Britain a lion knocks daily at the Prime Minister’s door, and our rugby team and Olympians sport lions. What’s left of our money is emblazoned by a coalition of three maned males of Plantagenet decent. Globally, lions are the most frequent national animal, adopted by 14 countries worldwide (of which they actually occur in only Ethiopia and Kenya). Working with WildCRU’s art historian Caroline Good, we found that in 1,000 Mughal animal artworks from 1526 to 1707, and another 1,000 from the British 1700–1900, the lion is the most depicted species. Surely, if there is one species on earth that would be safeguarded it is the lion. On the contrary, in recent history lions have disappeared from Mauritania, Gambia, Togo, Djibouti, Lesotho, probably from Sierra Leone and Eritrea, and are on the way out in Ghana, Guinea, Guinea Bissau, Ivory Coast and Mali. WildCRU’s Hans Bauer led a heart-breaking analysis that concluded that over just the last three generations of lions, their numbers overall had declined by 43%, and by a devastating 66% in West and Central Africa, with the statistics suggesting more than even odds of similar declines in the coming decades. Panthera’s Philip Henschel, a tireless searcher for needles in haystacks, reckons that only 250–587 critically endangered lions survive in the entirety of West Africa, with not a trace left in 17 of the region’s 21 large protected areas. Perhaps 24,000 lions survive across the continent, with more than a thousand in only six populations, such as Selous and Ruaha in Tanzania. In the remaining 60 or so populations, almost half number fewer than 50 individuals – hence the slogan “Secure the 6, Save the 60”.

What next – ethics and coexistence Conservation scientists know the threats to lions, and thanks to Keith Somerville’s remarkable compendium they will appreciate even more the long history of those threats. The conservationist’s medical kit is bulging with diagnostic tools and ingenious analgesics and prophylactics to mitigate and to forestall the community headache of having lions in your backyard, from portable bomas to payments to encourage coexistence. Where there are evidence gaps, dedicated scientists, natural and social, strive to plug them. If the conservation doctor’s metaphorical medical kit is ever better stocked, the health service in which it functions remains nowhere near fit for purpose: global and national systems are not up to the task of integrating the conservation of lions (themselves a metaphor for biodiversity) into the 21st century human enterprise. This is largely not because we, society, do not know what to do: rather, we need to decide what we ought to do. This, essentially ethical indecision, shimmers behind almost every question Keith Somerville’s text poses. This historical account of what has always

xxiv Foreword

been does not speak to what should always be. As time runs out for wildlife, and sustainability threatens to become a receding dream, the 21st century lesson might be not to repeat history but to leave it behind, or else. Consider two of the many steps on the journey. Should lions be farmed for their bones? The answer might be informed by asking a cascade of questions, such as whether lion farming promotes, or detracts from, some aspect of biodiversity conservation, and insofar as we should promote conservation the answer could lead to the conclusion that we should or should not promote lion farming. Equally, we might ask whether lion farming generates income, maintains jobs or provides other benefits to the economy which, on the premise that we should promote benefits to the economy, could lead to the conclusion that we ought to promote lion farming. But the issue of lion bones, as Somerville’s narrative reveals, is a relatively small abscess on the wider anatomy of lion conservation. For a more challenging “what ought we do?” topic, try trophy hunting (our team, led by ethicist John Vucetich, devoted a year of concentrated thought to a paper that concluded either maybe or maybe not). Such topics led Keith Somerville, as they will lead you in the following pages, to ultimately ethical questions of deontology, consequentialism and virtue. Keith has set out for you, thoroughly and entertainingly, the grist for those mills. It is now over to you to decide what ought to happen next. Bones and hunting are side-shows to the main theatre, where the stage is set for us to decide whether humans will coexist with lions or not. Your decision will be affected by evidence, by the balance sheets of concerns for the well-being of wild nature and of developing people, and perhaps by your opinion on how inclusively to draw contours around the circle of life whose interests we are obliged to respect. And so back to fairness, Androcles and who deserves concern: are lions in or out? David W. Macdonald Oxford

Abbreviations

ADMADE

Administrative Management Design for Game Management Areas (Zambia) ANAC Mozambican National Administration of Conservation Areas APN African Parks Network AWF African Wildlife Foundation (US-based) BSAC British South Africa Company CAMPFIRE Communal Areas Management Programme for Indigenous Resources (Zimbabwe) CAR Central African Republic CBNRM Community-Based Natural Resource Management Canine distemper virus CDV Convention on International Trade in Endangered Species of CITES Wild Fauna and Flora Central Kalahari Game Reserve (Botswana) CKGR DNPWLM Department of National Parks and Wildlife Management (Zimbabwe) Democratic Republic of Congo (formerly Zaire) DRC Dallas Safari Club DSC Department of Wildlife and National Parks DWNP GLTFCA Greater Limpopo Transfrontier Conservation Area GMA Game Management Area GR Game Reserve IRDNC Integrated Rural Development and Nature Conservation (Namibia) IUCN International Union for Conservation of Nature KAZA Kavango–Zambezi Transfrontier Conservation Area KCS Kalahari Conservation Society

xxvi Abbreviations

KGR KNP KWS KTP KZN LCU LIRDP MET MMNR NP PA PCF PH PHASA PI-WMA PWMA RCP SANParks SAPBA SCI TAHOA TAWIRI TCM TWPF WAP WCTF WildCRU WMA WWF

Khutse Game Reserve (Botswana) Kruger National Park Kenya Wildlife Service Kalahari Transfrontier Park (South Africa and Botswana) – also rendered as Kgalakgadi KwaZulu-Natal (Province) Lion Conservation Unit Luangwa Integrated Rural Development Project Ministry of Environment and Tourism (Namibia) Maasai Mara National Reserve National Park Protected Area Predator Compensation Fund Professional hunter Professional Hunters’ Association of South Africa Pawaga-Idodi Wildlife Management Area (Ruaha, Tanzania) Parks and Wildlife Management Authority (Zimbabwe) Ruaha Carnivore Project South African National Parks South African Predator Breeders Association Safari Club International Tanzania Hunting Operators’ Association Tanzanian Wildlife Research Institute Traditional Chinese Medicine Tanzania Wildlife Protection Fund W-Arly-Pendjari National Park Zimbabwe Conservation Task Force Wildlife Conservation Research Unit, University of Oxford Wildlife Management Area Worldwide Fund for Nature

Introduction

Lion populations in sub-Saharan Africa have been in progressive decline over recent decades. One of the latest and most authoritative figures, from the International Union for the Conservation of Nature (IUCN), puts the possible global population between 23,000 and 39,000. Macdonald and Dickman believe the actual number is likely to be at the lower end of this wide range. The IUCN estimate represents a decline of at least 43% in estimated numbers between 1993 and 20141. They have been exterminated or expelled by human activity from at least 92% of their historic range, occurring in 24 African countries, definitely extinct in 15 and probably extinct in another seven. Outside Africa they have disappeared from Europe, West Asia, the Middle East and all but the Gir Forest and surrounding areas of Gujarat in India 2. Extinction in most of their former range is principally the result of their relationship with man. The history of their coexistence and conflict with people is vital to understanding why their numbers continue to fall and how their disappearance in the wild may be prevented in the future. This book has its origins in nearly 40 years of visiting East and Southern ­A frica as a journalist, academic, now a researcher on human-wildlife relationships and obsessive safari-goer. During this time, I’ve had the pleasure of observing hundreds of lions in many different locations, of talking to local people, game rangers, safari guides, conservation biologists, NGO campaigners and hunters about living with lions, tracking them, studying them, campaigning for their preservation and hunting them. Through this, my work as a journalist and as a magpie-like collector of data, impressions and experiences, and my academic study of human-wildlife coexistence and conflict, I built up a mass of material on lions, their habits and their millennia-old relationship with people.

2 Introduction

The spark that ignited this mass of material into the blaze that is book writing came from the media conflagration that burned fiercely for several months in 2015 after the reporting of the killing of a lion named Cecil in Zimbabwe. As a journalist and academic teaching journalism, I was both fascinated and appalled by the unbalanced, often totally inaccurate and generally misleading nature of the newspaper, online, radio, TV and social media coverage of the story. I set out to investigate what happened and why the media covered the story in the emotive, sensational and misleading way that it did. In the process I received valuable help and encouragement from Professor David Macdonald, Dawn Burnham and Amy Dickman of WildCRU at the University of Oxford, who had run the research programme that included Cecil. The result was my academic paper on Cecil and the Pride and Prejudice of the British media 3. David, who had read my book on the history of the ivory trade in Africa4, then encouraged me when I started working on a parallel book looking at the history of human-lion coexistence and conflict. Although in the historical record that I researched there were huge gaps in information on the populations and ranges of lions and their relationships with humans (particularly from the start of the Common Era up to the penetration of Africa by European hunters, traders and colonisers), I was able to find a wealth of material about early Felids (cats) and the distant hominid and hominin ancestors of humans. From this I have tried to piece together an account from the Pleistocene to the present of the evolving relationship between people and lions. This account is, at times, at best the most supportable interpretation of a mass of data and often competing explanations of hominid/hominin/human interactions with ancient and modern lions. The relationship developed from one in which humans and their ancestors were prey for lions, the scavengers of lion kills (either abandoned ones or by working in groups to drive off the lions), to becoming competing hunters of large prey. It was then transformed by the making of weapons, evolving of social organisation and the greater ability of humans to defend themselves from lions. Then, as humans domesticated livestock they needed to protect, they killed lions in defence of stock or retaliation for lion predation on domestic animals or people themselves. The expansion of human populations, development of agriculture and ever advancing technology of weaponry changed the balance of power between humans and lions, but without removing human awe, respect and fear of lions. Lions became the quarry of lords and kings, symbols of power, authority and majesty and part of human imagery of the wild. Advances from handheld stone weapons, to thrown weapons, spears, arrows, poison and then firearms gave humans the ability to kill lions in large numbers and to kill their prey in even larger numbers, making people the most deadly enemies of lions. The development of concepts of conservation and wildlife management followed which offered ways of saving lion populations, setting aside habitat for them, but often then at a cost to local communities. Linked to this is the continuing and often

Introduction  3

bitter debate over whether the hunting of lions can and should play any role in their conservation. These are the subject of the book. The chapters will take the reader through the Pleistocene period of the evolution of Felids and hominids/hominins to about 32,000 years ago (Chapter 1); through the time of the ancient empires of Assyria, Egypt, Greece, Rome, Axum and the rise of chieftaincies, kingdoms and empires in Africa, West Asia and India (Chapter 2); into the time corresponding to the renaissance period in Europe to the start of European colonial occupation of Africa in the late 19th century (Chapter 3); through the colonial period in Africa with its mixture of the decimation of wildlife and the origins of modern Western-derived conservation concepts, and the history and current status of the Asiatic lions of the Gir Forest in India (Chapter 4); and then into the contemporary period from African independence to the present (Chapters 5 and 6). While the chapters follow a broadly chronological pattern there is some overlap between them, as there is no simple division of historical periods or events that affect human-lion relationships. The final two chapters will look in detail at the modern relationship in Africa, attempts at developing conservation methods that can make coexistence possible and at the trophy hunting/­ conservation debate. A common thread throughout the chapters is the link between the rise of human civilisations, technological advance, population growth and agricultural expansion and the negative effects these have had on the range of lions across the globe. The disappearance of ancient lion species, the shrinking of the ranges and extermination of populations of modern lions will be covered in detail. The main focus will be on Africa, the original home and contemporary refuge of the lion, but the Gir lions of India will not be forgotten. The book is an historical account of human-lion coexistence and conflict and not a charting of the biological development of lions, their physical characteristics, classification of sub-species or behavioural/social traits. Nor will the book be an account of the development of the lion entertainment business through books, films, cartoons (such as The Lion King), TV documentaries (notably the drama-documentaries of Dereck and Beverly Joubert or the Big Cat Diary series on the BBC which, while not falsifying events or inventing them, did turn the stories of lion prides into quasi-human soap operas to grab prime-time audiences, with strong elements of jeopardy built into each programme to keep viewers tuned in5). The effects of such representations of lions were primarily on Western audiences and not on those communities that live alongside or are directly involved in managing or conserving lion populations. Where relevant, references to the imagery and its effects on public opinion as it affects non-governmental organisations (NGOs) and their role in conservation will be examined. A separate book is needed on the media and modern cultural imagining of lions. The source material for the book is a rich mix of: scientific/academic works on human and lion evolution and interactions over time; modern scientific studies of lions and their coexistence and conflict with people; historical accounts of

4 Introduction

human development and its effects on the environment and wildlife; the Proceedings of the Zoological Society of London; accounts by adventurers, hunters and colonial officials; the records of colonial experiences of lions and their management from the Journal of the Society for the Preservation of the Wild Fauna of the Empire and colonial game department records; diaries and blogs of lion scientists and those working in the field on conservation projects; and a mass of interview material from scientists, conservationists, game rangers, safari guides and people living alongside lions, combined with my own observations. In writing the book I have had invaluable help from many people: Professor David Macdonald, director of WildCRU at the University of Oxford, has read the manuscript and made many constructive suggestions enabling me to improve the text, and he very generously has written the foreword; he has also kindly given me access to the drafts of forthcoming studies developed at WildCRU to keep my work as up to date as possible. Dr Amy Dickman has given freely of her huge knowledge, particularly of Tanzanian lions. Dr Laurence Frank of Berkeley University and the Living with Lions/Laikipia Predator Project has read and commented in detail on all the chapters, giving me valuable advice and support. Dr Ross Barnett of Edinburgh University, Dr Bruce Patterson of the Field Museum in Chicago, Tammy Hoth-Hanssen of Africat-North in Namibia, Dr Peter Apps and Cameron Radford of the Botswana Predator project, Blondie Leathem and Matt Wijers of the Bubye Valley Conservancy in Zimbabwe, Colin Jackson of the BBC Natural History Unit, Kamogelo Pshatlele Mabowa, of the Mapungubwe National Park in South Africa, Dr Philipp Henschel of Panthera in West Africa, Professor Craig Packer (director of the Lion Research Centre at the University of Minnesota), Lee White (head of Gabon’s National Parks Department), Andrew Dunn of the Wildlife Conservation Society, Dr Vivienne Williams of the School of Animal, Plant & Environmental Sciences, University of the Witwatersrand, and Chief Timex Moalosi of Sankuyo in Botswana, have all given their time and provided valuable insights into human-lion relationships. My colleagues Dr Bob Smith and Dr David Roberts at the Durrell Institute of Conservation and Ecology at the University of Kent have been generous with their support and advice, as has Professor Adam Hart of the University of Gloucestershire. I’d also like to thank Sarah Broadhurst and Emma Milnes of the Zoological Society of London library for their help and patience in finding archive and other material for the book, and Danielle Garbouchian and Susie Weller of Panthera for giving me permission to use an up-to-date map of lion distribution in Africa. My thanks also to Liz Dawn, Leila Walker and Charlotte Endersby of Routledge for helping me turn this mass of words into a proper book. Most of all I must thank my wife Liz and son Tom for their patience, support and good humour, despite their early protestations that “I thought you weren’t writing another bloody book” when I first announced my intention to produce this tome.

Introduction  5

Notes 1 David W. Macdonald (2016), Report on Lion Conservation with Particular Respect to the Issue of Trophy Hunting, 28 November, Oxford: WildCRU, https://www.wildcru. org/wp-content/uploads/2016/12/Report_on_lion_conservation.pdf, accessed 16 March 2018, p. 1; and personal communication with David Macdonald and Amy Dickman of WildCRU, University of Oxford. 2 H. Bauer, C. Packer, P.F. Funston, P. Henschel and K. Nowell (2016), Penthera Leo, IUCN Red List of Threatened Species 2016, www.iucnredlist.org/details/15951/0, accessed 16 March 2018. 3 Keith Somerville (2017), Cecil the lion in the British media: the pride and prejudice of the press, Journal of African Media Studies, 9, 3, September, DOI: 10.1386/ jams.9.3.471_1 4 Keith Somerville (2016), Ivory. Power and Poaching in Africa, London: Hurst. 5 Personal communication with Colin Jackson (producer of numerous episodes of Big Cat Diary); see also, Jonathan Scott (2016), Big Cat Diary. An Autobiography, London: Bradt, pp. 278–286.

1 Lion-human coexistence and competition from the Pleistocene to modern humans

Ancestors of modern lions and humans developed from the earliest forms of mammals. Over tens of millions of years, mammals branched out into myriad species, including hominids (humans and apes) and the Felidae (cat) family. Between the start of the Pleistocene epoch 2.6 million years ago (million years ago henceforth rendered as ma) and 300,000 years ago, humans were refined down to one species, Homo sapiens, and lions similarly to Panthera leo (henceforth P leo), with different populations across Africa, southern Europe, the Middle East, South and West Asia. This chapter will examine the evolution of the lion alongside humans, the extinction of P leo’s ancestors and related species, and the development of patterns of human-lion coexistence and conflict. The aim is to paint a picture of how the contemporary relationship between lions and humans evolved and how early, regularly reinforced experiences may have shaped human attitudes. The narrative will cover the Pleistocene period (2.6 ma–11,700 years ago), into the Holocene epoch (from 11,700 years ago to the present).

Time periods Pliocene – from 5.33 to 2.6 ma, immediately precedes the Quaternary period and Pleistocene epoch. Quaternary period – the last 2.6 million years. Pleistocene epoch – part of the Quaternary, stretches from 2.6 ma to 11,700 years ago. Early Pleistocene – from 2.6 ma to c.781,000 years ago. Middle Pleistocene – from c.781,000 to 126,000 years ago. Late Pleistocene – from 126,000 to 11,700 years ago. Holocene – from 11,700 to present.

From the Pleistocene to modern humans  7

Glossary of other terms Acheulean – a Paleolithic period of human evolution within the Pleistocene, characterised by stone-tool making, from 1.7–1.5 million years ago to 250,000–200,000 years ago. The humans of the time “were truly primitive hunter-gatherers”1. Anthropogenic – effects related to human activity. Before Present (BP) – a timescale used to specify in broad terms when events occurred. Because the present time changes, standard practice uses 1 January 1950 as the end date. Carnivore guild – a guild is any group of species that exploit the same general type of resources; in the case of carnivores, meat-eating. Clade – a grouping that includes a common ancestor and all the descendants (living and extinct) of that ancestor. Cladogenesis – an evolutionary branching event where a parent species splits into distinct species, forming a clade. This can occur when life forms end up in new areas or when environmental changes cause extinctions, opening new ecological niches. Hominid/Hominidae – member of the group consisting of modern and extinct humans, great apes and their immediate ancestors. Hominin/Homininae – a member of the group of species (genus) that paleoanthropologists agree are human or human ancestors. This includes the Homo species (Homo sapiens, H. ergaster, H. rudolfensis, including Neanderthals­ Neanderthals, Denisovans, and Flores), all of the Australopithecines ­(Australopithecus afarensis, A. africanus, A. boisei, etc.) and other ancient forms like Paranthropus and Ardipithecus. Neolithic – relating to the Later Stone Age. This period, also sometimes called the New Stone Age, was characterised by use of stone tools produced by polishing or grinding, utilisation plants cultivated by people, domestication of animals, settlement in permanent villages, and the appearance of pottery and weaving. Oldowan – the stone tool-making culture of hominid/hominin ancestors, from 2.5–1.7/1.5 ma. First defined by Louis and Mary Leakey following tool and fossil discoveries at Olduvai Gorge, Tanzania 2. Paleolithic Period – the age in which early humans made chipped-stone tools. Sometimes called the Old Stone Age. The period started around 2.5 ma. Wisconsin Glacial period – the last cycle of cooling and glacier expansion in North America. About 100,000 years ago, the climate cooled again and a glacier, the Laurentide Ice Sheet, spread across the continent 3.

The evolution of Panthera leo – the modern lion Examination of the evolution of P leo, the modern lion, starts with early mammalian prototype carnivores, which took over the niche in the food chain occupied

8  From the Pleistocene to modern humans

by the dinosaur carnivores, which disappeared with the mass extinction of dinosaurs 65 ma. Many of the creatures were chiefly scavengers, but others hunted, and lacking the meat-shearing carnassial teeth diagnostic of modern carnivores, tore meat from their prey. A squirrel-like carnivore Cimolestes appeared around the time of the dinosaur extinction and is the ancestor from which contemporary species labelled carnivores are descended4. As Macdonald explains5, the order Carnivora evolved over millions of years from Cimolestes, developing from small animals preying on insects and small vertebrates into large ones preying on huge herbivores. Many developed carnassial teeth suited to cutting flesh from prey. These early carnivores proved more successful than the first-known meat-­ eaters between 55 and 35 ma, the Creodonts6, which had included the strikingly named Hyaenodon horribilis, a wolf-sized meat-eater, and the Megistotherium, an 800kg hyena-like predator, probably “the largest mammalian land predator ever” 7. The evolving carnivores replaced the Creodonts, and became the top land predators across the northern hemisphere by 30–20 ma. One reason for success was their adaptability. They were able to eat fruit and vegetables, as well as meat, and so survive declines in prey species in a way that the more specialised Creodonts could not8. This period coincided with climatic changes and the expansion of grassland savanna and lightly wooded areas, which started around 60–55 ma. Grasses and grazing animals appear to have evolved in tandem9. By 30–20 ma the ancestors of most of the large mammals of modern African savannas appeared, helped by the volcanic and geological activity that reconnected Africa with Asia, formed the Rift Valley and generated soils suitable for the growth of grasses. The hoofed mammals found in Africa at that time included ancient pigs, giraffids, buffalo, wildebeest, hartebeest, reedbucks, gazelles and smaller antelopes10, providing prey species enabling the evolution of carnivores, with the eventual appearance of the cats (felids). Fossil records show that one felid-related species, Dinictis, developed in America, Aeluructis, in Europe and another branch produced the Machairodontinae, the sabre- and scimitar-toothed cats11. The cats produced the Pantherinae sub-family and from it the Panthera genus (including the modern big cat species – lion, tiger, leopard and jaguar) and the Felinae, from which smaller cat species evolved. They were purely meat-eaters. The Machairodontinae produced large sabre- and scimitar-toothed species such as Homotherium, Smilodon and Megantereon.12 The sabre-toothed Smilodon was found in North and South America, but others were found in Africa, Asia and Europe alongside other cat species, and were the dominant felids around 6–5 ma, when thick forest was retreating because of declining rainfall and other climatic changes. Fossils from Kenya show evidence from 22–18 ma of early felids such as Metailurus africanus, a sabre-toothed cat. Later – in the Pliocene and early Pleistocene – the main sabre-­ tooth species, the Machairodontinae were present in Africa, with fossil finds at Langebaanweg in South Africa and the Omo Valley in Ethiopia13. Megantereon, a true sabre-tooth, was present in Africa early in the Pleistocene with fossils found at Komdraai, Schurveberg and Sterkfontein in South Africa14. Dinofelis (a genus

From the Pleistocene to modern humans  9

of sabre-tooth cats believed to be specialised ambush hunters) were widespread in Europe, Asia, Africa and North America from about 5 ma to about 1.2 ma. Dinofelis was believed by palaeontologist Brian Brain to have been a specialist primate killer that preyed on baboons and hominids, including the Australopithecus and Paranthropus human ancestors. This has been challenged but not definitively disproved by others, such as Francis Thackeray, who believe that Megantereon was most likely the chief hunter of hominids15. The evidence from fossils can be interpreted in both ways, but there is no strong reason to believe that Dinofelis could not have hunted hominids or that this would exclude Megantereon. Around 10 ma a broad shift was taking place, with the decline of dense forests and expansion of grassland, bush savanna and open woodland. This encouraged the evolution of fleeter-footed prey species, with a diversity of antelopes, gazelles and other grazers/browsers which thrived in open countryside, and fast land predators. It prompted changes in the social organisations of some predators in open countryside, developing social hunting16. For specialist meat-eaters, the evolution of prey species had a major effect on body-size, weight, strength and agility. In East and Southern Africa, “the opening of the environment heralded dramatic radiations in carnivores, ungulates and ground-dwelling primates after c. 7 Ma ago”17. As Yamaguchi et al. explain, “From the late Miocene [23.03 to 5.333ma] through the Pliocene [5.33 to 2.58 ma], African mammalian faunas experienced a great change: 76% of the land mammals were new, and of these c. 53%” were found in Africa alone. The first record of a lion-sized Panthera fossil “occurs at Laetoli in Tanzania c. 3.5 Ma ago”18. Fossil remains suggest that the earliest lion-like species originated in East Africa in the late Pliocene (5.0 to 1.8 ma). They, in a pattern uncannily like that of early humans, developed in East Africa and migrated to the Middle East and Eurasia in the middle Pleistocene, 800,000–100,000 years ago19. Lions were also believed to be present in Southern Africa at the time they are first recorded in East Africa 20. Migrating lions had reached Western Europe, including Britain, by 0.5 ma (Panthera leo fossilis). The lion in its various subspecies had, during the late Pleistocene, one of the largest geographical distributions of any terrestrial mammal 21, before combinations of human expansion, anthropogenic changes, loss of prey species and climatic/vegetation changes reduced the number and spread of species22. The early species of lions successfully colonised Europe, Asia, North and northern-South America. They included the cave lion (Panthera leo spelaea) and the modern lion (Panthera leo), the latter appearing around 0.6 ma. The cave lion “was an integral component of the late Pleistocene Holarctic ecosystem, occupying the position of apex predator”. Its distribution is known from plentiful remains preserved in cave systems in Europe and permafrost sediments of Siberia, Alaska and the Yukon 23. They were maneless in both sexes24, around 25% bigger than modern lions25. For a period, the lion species coexisted with sabre- and scimitar-toothed cats in Africa, Asia, Europe and the Americas. The lions which colonised North America via the land bridge from Eurasia (Panthera atrox) may have been essentially the

10  From the Pleistocene to modern humans

same species as modern lions (Panthera leo) though they were believed to be up to 25% larger. They lived alongside sabre-toothed Smilodon, which disappeared 9,400 years ago. These older species were dependent for prey on large, slow megafauna herbivores (including mammoths) and were better adapted to forests than open woodland or grasslands. It is very likely, based on evidence from excavations of fossils across their ranges, that during their period of coexistence with hominins, sabre-toothed cats and various Panthera species preyed on hominids and hominins. This preceded hominins developing cooperative tactics and weapons enabling them to defend themselves and then drive the predators from the caves which they both favoured as dens26. These ancient species disappeared when climatic changes led to the decline in canopy forest and the expansion of open woodland, bush savanna, grassland, steppe and desert, and when their prey disappeared as a consequence of that and human predation. Meganteron disappeared around 900,000 years ago. Homotherium (which was scimitar-toothed) survived in Europe until 500,000 years ago27. The North American and Eurasian lions later suffered extinction because of a combination of factors, including human hunting of their prey at a time when glacial fluctuations and resulting climate/vegetation change led to savanna/ steppe returning to thicker forest, in which lions and their prey struggled to survive. All the species of North American sabre-tooths and P atrox were extinct by 9,000–10,000 years ago, leaving the jaguar and puma (mountain lion or cougar) as the only large felids in the Americas. The causes of the late Pleistocene extinctions, especially in North A ­ merica, are hotly debated with potential extinction hypotheses including climate change, human overkill, or a combination of human effects during interglacial warming there. As DeSantis et al. contend, “While large carnivores like S. fatalis [Smilodon] and P. atrox are unlikely to have been directly hunted to extinction by humans, they were likely vulnerable given competition with humans for prey species”28. There is no definitive version of the extent to which megafaunal extinctions can be laid at the door of humans. The strength of causal factors varied across regions but human activities, notably hunting, were part of the mix. One factor is that the increase in numbers of Stone Age humans, the development of better weapons and methods of hunting indicate the ability to play a role in megafaunal extinctions, although that does not explain why many survived in Africa but not elsewhere29. But, as Klein contends, in Africa, where smaller numbers of megafauna disappeared, climate and vegetation change that reduced food for herbivores and so prey for carnivores was a major driver of extinctions. In ­Eurasia and the Americas changes in climate and vegetation (the spread of forests in some areas and grassland in others) were factors “in hastening extinction of large herbivores … [and with] human hunting” as something that had a “general disruptive effect on systems”. He goes on to suggest that the arrival of modern humans with advanced hunting techniques in the late Pleistocene at a time of drastic climate change may have had a particularly strong effect in Australia and the Americas30. This would help explain why towards the end of the Pleistocene,

From the Pleistocene to modern humans  11

35 genera of large mammals became extinct in North America, either in the sense ceased to exist anywhere in the world (29 genera), or that they disappeared in America but survived elsewhere (6 genera)31. Klein, along with Grayson and Meltzer, are critical of the thesis developed by Martin32 of “overkill” by human hunting during the late Pleistocene, especially in North America, being the major cause of extinctions. The overkill idea is that humans arrived in North America at a time when climate and vegetational change made prey species more vulnerable and that excessive hunting by humans had a devastating effect on prey numbers. Grayson and Meltzer argue that Martin’s position gains virtually no support from the North American late Pleistocene archaeological and paleontological records”. Among the species that disappeared, were the sabre-tooth, Smilodon, the scimitar-­tooth, Homotherium, the American cheetah, Miracinonyx, and then the American lion, Panthera atrox 33. Jerry McDonald 34 doesn’t support the overkill thesis but does advance the idea that when megafaunal extinctions took place, hunting by technologically advanced human communities was a new factor and may have hastened the collapse of vulnerable animal populations in North America. Kruuk has a similar view and suggests that “environmental change may have rendered species more vulnerable to our onslaught”, especially in a situation where humans were the new and most advanced carnivores on the block 35. Following their approach, it is not unreasonable to say that “overkill” is too strong, but that humans played a major role in extinctions in some parts of the world. The major difference between Africa and the rest of the world was the gradual evolution of efficient hunting by hominins in Africa living alongside their prey, which would have allowed prey to co-evolve anti-predator behaviour. Extinctions were slower in Eurasia than in North and South America, possibly because again, the small human population enabled time for evolution of avoidance or defence strategies by prey. When efficient human hunters and their dogs arrived in Australia and the New World, they increased rapidly in number due to consumption of prey that was vulnerable as it was unused to human hunting methods. They may also have focused on killing the young of the largest beasts, thus preventing species from recovering their numbers when hit by the effects of climate change36. In a more recent study, Sandom et al., including lion specialists Dickman and Macdonald, note the clear evidence of anthropogenic factors in species extinction between 100,000 and 1,000 years ago. But they caution that “the primary driver(s) of large mammalian predator extinction specifically remain unclear”, though “the loss of prey diversity may partly account for the extinction of large carnivores” and humans played some role in this process with “an important role in the extinction of at least 166 large continental mammal species”37. In both Africa and Europe, the disappearance of the sabre-toothed cats was matched by that of the large, early species of hyenas, leaving lions as the apex predator. In Africa, they were able to hunt the faster browsers and grazers ranging in size from buffalo to large antelopes, gazelles and small species such as

12  From the Pleistocene to modern humans

duiker, dik-dik and steenbok 38. Lions species became dominant through speed and cooperative hunting in open landscapes, social organisation of lions developing as cooperative hunting enabled them to take advantage of the opening up of the habitats. In Europe, the loss of the older forms and the shrinking of the modern lion range during and since the late Pleistocene resulted from a further series of climatic changes causing major changes in vegetation. Terrestrial mammals reacted to such changes and expanded territories or migrated, some simply disappeared being unable to adapt. Carnivora in Europe were affected by these changes as they were governed by the numbers and then by the disappearance of their major ungulate prey species and by competition from new members of the carnivore guild 39. The European cave lion became extinct when ungulate populations declined about 85,000 to 11,000 years ago. In the colder regions of Eurasia, boreal forest replaced steppe, leading to the retreat south into surviving steppe areas of prey species like the saiga antelope or north into the tundra, in the case of musk ox. Human hunting may have contributed to this process. In their study Vereshchagin and Baryshnikov suggest that while “Paleolithic people applied considerable pressure to the mammal populations of the plains,” they also scavenged carcasses and so human hunting alone is unlikely to have wiped out species40; though as David Macdonald suggests41, there was competition and hostility between members of the carnivore guild, as there is today between lions, hyenas, wild dogs, leopards and cheetahs. Examination of DNA from museum fossils and specimens of early lions suggests that from a common P leo ancestor in East Africa 124,000 years ago (given margins for error, within a span from 183,500 to 81,800 years ago), five populations emerged – the North African/Asian, West African, Central African, East-Southern African and Southern African42. The evolutionary cradle for the birth of the modern lion was in East and Southern Africa. The family tree of P leo spread out from there into the different branches, with the West African population then expanding to form the Central African population43. The North, West and Central African populations of P leo became separated during wetter periods of the Pleistocene bringing about the divergence of different subspecies or population groups in Africa from the base population. The wetter climate led to the expansion of tropical rainforest in much of West and Central Africa, isolating population groups. Around 51,500 years ago, a drier period led to the expansion of the Sahara Desert, separating North African lions from those in West and Central Africa44. Movements of lions out of Africa and into Asia and the Middle East started around 21,100 years ago. This formed the Asian, Middle Eastern and Iranian lion populations, which have been shown by DNA analysis to have descended from North African lions45. Working from Barnett et al.’s DNA analysis, the Cat Classification Task Force of the IUCN SSC Cat Specialist Group has provisionally proposed a split of current lions into two subspecies – P. l. leo of Asia and West, Central and North Africa, and P l. melanochaita from South and East Africa46. Panthera leo, which appears in the fossil evidence in Europe between

From the Pleistocene to modern humans  13

74,000 and 15,000 BCE, having migrated from Africa, retained a presence in much of Europe until at least the Bronze Age (1800–900 BCE), in some areas longer47. The most reliable accounts suggest cave lions, which had spread throughout Western Europe, disappeared approximately 14,000–14,500 years ago and were replaced in the food chain by P leo approximately 8,000 years ago. Modern lions reached the steppes of Ukraine and Hungary, but were unable to penetrate the dense forests of Central Europe48. They were found around the Black Sea littoral until at least 3000 BCE. In the Balkans, Turkey and the Middle East, the lion was still present at the beginning of the Common Era49. Sommer and Benecke note that Panthera leo went into strong decline at the end of the Iron Age (1200–600 BCE) in Europe, leading to its extinction in south-eastern Europe. The most likely factors in the extinction of European lion populations being hunting both of lions and their prey, the increase in human populations, the progressive colonisation of lion habitats by humans and the rise in stock breeding and protection of stock 50. Over millions of years during the Pleistocene and early Holocene, lion ranges expanded and contracted as climatic and vegetation changes caused fluctuations in availability of prey51. The stamina and build of lions enabled them to migrate rapidly and populate most continents before further climatic and anthropogenic changes led to a shrinking of the range to Africa, southern Europe, the Middle East, West Asia and parts of South Asia. Fossil evidence suggests that hominin interactions with the first early lion species occurred in East Africa 3.5 ma at the time of the evidence of the first lions – based on Mary Leakey’s discovery of a jawbone of a lion-like felid at Laetoli in Tanzania near to footprints of human-­ like hominins52. Hominins, like the lions, were also capable of long distance migration and also moved from East Africa into Europe, Eurasia and Asia in the same period53. As lions spread across the continents, there were changes in the composition of the carnivore guild across the Pleistocene. In East Africa this involved a decline from a peak in numbers of carnivore species in the early years of the late Pliocene period (3.6–3.0 ma), which has continued to the present 54. The decline was related to changes in habitat as forests retreated, which hastened the loss of megafaunal prey species that were the source of food for the older felids. Such human influence that there was may have been limited to h ­ ominins/early humans developing better defences against predation by carnivores and competing more effectively as scavengers and hunters. The decline in prey species and carnivore diversity was a process over hundreds of thousands of years rather than a time-restricted “event”55. This was related to a “long-term trend towards climatic cooling [that] was punctuated by stepped increases in climate variability and aridity … These changes resulted in an opening-up of the eastern African landscape, where generally forested habitats were replaced by more open grasslands”. The spread of the Homo genus of early humans may also have had an effect, as they were increasing the amount of protein derived from meat in their diet and becoming part of the carnivore guild 56.

14  From the Pleistocene to modern humans

One factor that may explain why P leo became the dominant felid in Africa and other areas while the older species failed, was its social nature, food-sharing and cooperative hunting to bring down much larger or faster prey in open landscapes. The social nature of lions distinguished it from other large cat species that survived the extinctions such as cheetahs (Acinonyx jubatus), leopards (P. pardus), jaguar (P. onca), puma/cougar (Puma concolor), tiger (P. tigris), clouded leopard (Neofelis nebulosa) and snow leopard (Uncia uncia) – all of which developed hunting methods appropriate to their size, nature of their habitat and prey; the other distinguishing feature of P leo is the mane (though it is missing in a minority of males in certain areas)57. Lion populations outside sub-Saharan Africa formed separate geographical groups – Panthera leo leo (the North African or Barbary Lion) and Panthera leo persica (the Asian Lion)58, which may be effectively the same subspecies. North African/Barbary lions are now extinct in the wild and only occur in zoos, while the Asian population is limited to 500–600 animals in the Gir Forest in Western India. There is still debate over when lions migrated into India. Most biologists who have studied the Indian lion believe it arrived around 40,000 years ago having migrated from North Africa or perhaps East Africa 59. The dissenting voice to the migration theory is that of the tiger specialist Valmik Thapur, who argues unconvincingly that lions in India were all imported from North Africa or elsewhere by Indian rulers to stock vast hunting reserves and that lions were never indigenous to the subcontinent60. The evidence assembled by Antunes et al. from DNA and other studies works against Thapur’s thesis. They posit that two major lion expansions occurred across Africa and Asia. The first was “a long-distance colonization into Central and North Africa and … West Asia… lions likely colonized West Asia from the East Africa refugia”. This migration and expansion may have been occasioned by the start of a warmer and wetter period in Africa 130,000–70,000 years ago. It is likely that multiple lion expansions occurred in the Pleistocene, as occurred with humans. “A second, more recent lion expansion probably occurred at the Pleistocene/Holocene transition, this one from Southern Africa toward East Africa.”61 The latter migration also saw lions moving north from the Kalahari Desert during Pleistocene climatic fluctuations and most likely in response to northwards movements of prey species such as the impala, kudu and wildebeest62. After the migrations from Africa and expansion into Europe and West Asia, it is very likely that there was a contiguous population at first, until human expansion, settlement and human-predator conflict fragmented the range into separate population groups63 – a process that started with early humans as they migrated from Africa into Europe and Asia and established larger settlements and cultivated the land. Lion populations were highly mobile, leading to their wide pattern of expansion out of Africa. After the disappearance of spelaea and atrox, P leo was the largest felid in its extended range, as there appears to have been little crossover in ranges with tigers, even though they were also to be found in South Asia, parts of West Asia and right up to the Caspian Sea and Turkey. The lion’s

From the Pleistocene to modern humans  15

success appears to be a function of mobility, flexibility and the development of social organisation that improved hunting ability, deterrence of competitors and scavengers (humans or hyenas)64. This may, according to Yamaguchi et al., have enabled them to colonise relatively prey-scarce environments or those where solitary hunting of large prey would not be a viable survival strategy – the group size was and is very elastic and prides will coalesce into large groups or split up into smaller ones according to availability of prey65. In sub-Saharan Africa there were mobile and quite genetically diverse populations in Southern and East Africa, while those of West and Central Africa were limited in range by the dense forests to the south and desert to the north and had less genetic diversity66. Periods of drought in West Africa between 40,000 and 18,000 years ago appear to have fragmented the smaller population there and lack of prey led to the shrinking of lion ranges. While a wetter period followed and allowed some recolonisation as vegetation recovered and prey species returned, it may help explain the historically smaller and more scattered populations there. The genetic evidence shows that the lions of West and Central Africa stretched as far down as Angola and indicate the lion population there to be more closely related to those of the West than to populations in Namibia and Botswana67.

Humans and lions – the origins of conflict, competition and coexistence Snapshot of human development chronology68 Scientists are still trying to piece together the human evolutionary puzzle based on the fossil record. 55 ma First primates evolve. 15 ma Hominids (great apes) split off from ancestors of the gibbon. 8 ma Chimp and human lineages diverge from gorillas. 7–5 ma Chimps and hominins diverge. 4.4 ma Ardipithecus appears: an early “proto-human” with grasping feet. 4 ma Australopithecines appears, with brain the size of a chimpanzee’s. 3.7–3.5 ma Australopithecus afarensis appears in East Africa.

16  From the Pleistocene to modern humans

2.3 ma Homo habilis first appears in Africa. 2 ma Homo erectus appears. Fossil evidence shows the species lived across the globe, including South Africa, Kenya, Spain, China and Java. 1.85 ma First “modern” hand emerges, according to fossil evidence. 1.6 ma Hand axes are a major technological innovation. 800,000 years ago Evidence of use of fire. 300,000 years ago Neanderthals evolve and by 200,000 years ago spread across Europe and Asia. 300,000 years ago Evidence of early Homo sapiens in Morocco. 200,000 years ago Homo sapiens found in Israel. 60,000 years ago Modern human migration from Africa that led to modern-day non-African populations. The development of modern humans (Homo sapiens) from hominid and then hominin predecessors somewhat paralleled lion evolution. There were common environmental factors in their evolution and evidence of long-term contact and familiarity. Hominid ecology, like that of lions from the Pliocene to the Holocene, Table 1.1  Chart of the overlapping time spans of predator and hominin genera in Africa Carnivora Machairodus Dinofelis Homotherium Meganteron Panthera

xxxxxxxxxxx xxxxxxxxxxx xxxxxxxxxxxx xxxxxxxxxxx xxxxxxxxxxx xxxxxxxxxxxx xxxxxxxx xxxxxxxxxxxx xxxxxxxx xxxxxxxx

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xxx xxxxxxxx xxxxxxxxx 3ma–2ma

xxxxxxxx xxxxxxxxxxx 2ma–1ma

xxxxxxxxxxx 1ma–present

Hominins Ardipithecus Australopithecus Paranthropus Homo

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6ma–5ma

xxxxx xxxx

5ma–4ma

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4ma–3ma

From the Pleistocene to modern humans  17

was affected by changes in climate, vegetation and geology. In the early period when forest was the dominant vegetation type across A ­ frica, hominid distribution and evolution relied heavily on “potable water, and animal-­based food resources and plant-based food resources, with the plant diet providing more food than animal diet”69. The early hominids were chimp-sized and needed safe sleeping sites, confining them to forests. They were preyed on by sabre- and scimitar-toothed cats in forest habitats. These were arboreal hunters more akin to leopards and jaguars than lions. “During the Pliocene Africa’s physiogeographic evolution made it possible for the continent to support a number of hominid species”; the ideal habitat being a patchwork of woodland/forest and wetland/ grassland in South-Central Africa70. Geological and climate changes, which produced drier upland areas and the spread of open woodland and grassland, encouraged the evolution of hominids adapted to living in open countryside, which occurred in East and Southern Africa. The fossil evidence leads archaeologists and anthropologists to believe that in East Africa the hominid line derived from arboreal ancestors who inhabited forests that disappeared from the areas where the hominid fossils were later found. A very early form of hominid, Ramapithecus, fossil remains of which have been found in East Africa, “must have walked upright and must have had forelimbs that could have used simple tools” 71. The earlier hominid ancestors appear to have shared a ground-dwelling, knuckle-walking existence along with apes72 . Key changes in environment took place first during the Miocene ­(15.5–12.5 ma) and then the Pliocene-Pleistocene period between 3.0 and 2.5 ma. They involved lower, seasonally restricted rainfall, tectonic changes creating highland areas and related erosion, leading to increased diversity of soil and vegetation types. Widespread volcanic activity enriched soils, providing more nutritious grasses over wide areas. The improved availability and quality of plant food supported a wide range of mammalian herbivores. The two periods of cooling and drying were followed by what Owen-Smith calls “turnover pulses” in large mammalian herbivores taking place between 10 and 5 ma and again at around 2.5 ma. The first of the pulses saw the development of the modern bovid genera and the first australopithecines, which developed from the hominid species. The second period involved the thinning out of bovid species, with those unable to adapt becoming extinct, and also the divergence from other hominids of “robust australopithecines” more adaptable to the open environments73. Relating developments back to lion evolution, this period saw the emergence and dominance of Panthera, which could cope with the hunting challenges of open areas and faster prey. Some megafaunal species disappeared as savanna (with open canopy woodland, tree parkland, low tree and shrub bush, and grassland) came to cover about 40% of Africa’s land surface. This supported a diversity of herbivores and is a strong explanatory factor in why Africa avoided the large-scale megafaunal extinctions which affected North America and Eurasia. Owen-Smith estimates that during the megafaunal extinctions at the end of the Pleistocene between 15,000

18  From the Pleistocene to modern humans

and 10,000 years ago, the Americas lost 75% of genera, Europe and Australia 45% and Africa only 13%74. One should also add, as Laurence Frank says, that African animals co-evolved with hominins as the latter developed hunting. In Eurasia and the Americas, highly skilled hunters with their dogs and fire arrived in areas populated by large herbivores unused to human hunters75. The large variety of grazers and browsers enabled large predators like lions to survive in the new habitats. The hominids that emerged from the denser forests included early predecessors of humans, like Australopithecus ramidus, whose remains have been found in Ethiopia, and Australopithecus afarensis in Ethiopia and Tanzania. The Plio-Pleistocene pulse, brought about by further climatic cooling, was the period when open savanna expanded further, and australopithecines split into two divergent lineages, both increasingly bipedal and with evidence of stone tool use76. The one lineage brought forth Paranthropus robustus and Paranthropus boisei, which disappeared around 1 ma. The other led to Homo sapiens. Owen-Smith believes that the success of the latter may have been related to the ability to adapt to a wider variety of foodstuffs, with increasing consumption of animal protein, obtained by scavenging, and perhaps the killing of slow, small or weak animals77. One adaptation to savanna living, in another parallel with lions, was that greater social organisation and expansion of groups were necessary for survival. Without comparatively safe arboreal sleeping quarters and with the presence of large predators, it follows that to survive and thrive, hominids and then hominins needed to band together. They also began to use basic tools and weapons for food acquisition, processing and defence against predators. Then, as social organisation and tool use became more advanced, the capability developed to competitively scavenge from lions and other predators, and to hunt larger prey78. Over time, they developed greater body size and strength and increased brain size – aided by a richer diet. Increased body size required the maintenance of a healthy intake of protein, favouring the more developed Homo species over the less advanced Australopithecus/Paranthropus lineages. The skull of the first known hominin, Sahelanthropus tchadensis, was discovered on the shores of Lake Chad. This specimen, dating from 6–7 ma, is thought to have stood upright and had other human characteristics while having the brain size of a chimpanzee79. At each stage of evolution, hominins became more upright in posture, more dextrous, more advanced socially and technologically, enabling them to use a greater variety and sophistication of tools and weapons. Around 2.6 ma, hominins began to fashion tools from stone, rather than just using stones and branches80. They were given the name Homo habilis and identified as part of a stone-tool making culture from 2.5–1.5 ma by Louis Leakey following tool and other fossil discoveries at Olduvai Gorge, Tanzania. The making of tools and weapons, such as spearheads, improved hunting and defensive capabilities against predators like lions. It made hunting of large herbivores and competitive scavenging possible with less risk of death or injury81. Around 1.8 ma, the species Homo ergaster evolved, with the ability to make and use more sophisticated tools; they are believed to have utilised fire. Iliffe says that Homo ergaster

From the Pleistocene to modern humans  19

developed into the more modern human species, evidence for this coming from Ethiopia’s Awash valley dated at 160,000 years ago. Their evolution is believed to have started as early as 600,000 years ago82. There is new evidence that along the Atlantic coast of North Africa and in the coastal mountain ranges, there was also a process of evolution of Homo species. Recent finds from the Jebel Irhoud in Morocco suggest that the evolution of Homo sapiens from Homo heidelbergensis or rhodiensis occurred around 315,000 years ago (fitting in with the pattern derived from fossil evidence in East ­A frica of H sapiens origin between 400,000 and 200,000 years ago)83. In addition, DNA-based research on the skeletal remains of a boy believed to have died 2,000 years ago in South Africa have helped researchers to “recalculate the time at which humans like us – Homo sapiens – first split or branched from archaic or pre-­ modern human groups to between 350 000 and 260 000 years ago. Previously, it was thought that we emerged just a little less than 200 000 years ago”84. The Moroccan fossils are from a region that supported lions, reinforcing the view that environments that enabled the evolution of humans were also viable for lions. In the case of Morocco, the lions would have been the early race of North African, which formed a separate geographical population from sub-­Saharan ­A frican lions between 203,000 and 74,000 years ago85. As Palmqvist et al. have noted, the evolving nature of the carnivore guild (and the widespread distribution and growing social grouping of lions) may have forced Homo species towards behavioural and technological improvement in the Achuelian tool-making period (1.6 ma to 200,000), which saw the demise of the sabre- and scimitar-toothed cats and dominance across Africa of P leo 86. The relationship between two species in which they coexist and/or come into conflict and influence each other’s evolution, especially in behavioural terms, is termed co-evolution. As Stiner argues, “Virtually every major period in human evolutionary history seems to provide examples of co-evolutionary processes involving animals… So much so, that we must conclude that humans are exceptionally prone to forming co-evolutionary bonds with other species.”87 This certainly seems to be the case with humans and lions. They appear to have developed a co-evolutionary relationship through “predator-prey relations, predator-­ predator competition … Zooarchaeological records testify to formative relations between Pleistocene hominins and large carnivore species as the result of a common interest in the flesh and bones of large herbivores. Species may compete … by depleting shared resources … by competing directly over them … or both”88. Stiner dates this competition and probable evolutionary influence as starting at least 2 ma in Africa and spreading as hominins and lions migrated into Eurasia and beyond89. Degrees of competition would vary over time and place but always be sharpest during any period of resource scarcity affecting one or both species. As human brain capacity increased and social behaviour became more sophisticated, humans evolved a range of strategies for defence against and competition with predators, enabling them to have greater security and compete effectively with

20  From the Pleistocene to modern humans

lions90. Dinofelis and other sabre- or scimitar-toothed cats had preyed on hominids before the move into emerging savanna environments. Fossil evidence in the form of tooth marks on bones indicate carnivores preyed on them91. Status as prey for carnivores will certainly have been a major aspect of early interactions but the relationship changed with increasing hominid and then hominin intelligence, social structures and weapon acquisition. Rosell et al. describe five aspects of hominid/hominin interactions with wildlife: “1. Hominids as regular scavengers of remains of kills by other carnivores. 2. Carnivores as scavengers of remains stored by hominids in their shelters or campsites. 3. Carnivores as prey of hominids. 4. Hominids as prey of carnivores. 5. Coexistence, commensalism and domestication.”92 The move from being prey (though not one that excluded some predation on hominids, hominins and later modern humans, as evidenced by historical and contemporary evidence of lion predation on humans) to scavenging carnivore kills, defending carcasses from scavenging by carnivores and killing of carnivores, all attest to an evolving relationship and some aspects of early interactions did not disappear as the ability to compete with carnivores and even kill them improved. The need for defence against predation, growing hominin consumption of animal protein and increased brain capacity/intelligence led to “significant behavioural transformations. New subsistence strategies based on obtaining animal protein emerged, which generated changes in the ecological relationships between hominins and the other predators”93. Evidence of hunting of lions by hominins is not common but fossils from La Garma in Spain indicate that in the Middle Pleistocene (781,000 to 126,000 years ago) early lions in Europe, P leo fossilis, were hunted. The evidence suggests use of skins, tendons and teeth by the hunters, though with no evidence whether this was a vital part of a subsistence strategy 94. Cueto et al. infer some form of ritual utilisation of lion parts, noting that representation of lions in early production of images in the Upper Paleolithic “clearly places lions in a prominent hierarchic position in the early humans’ symbolic world, giving this animal an important role in human culture”95. Rituals may have included manhood rites and prestige hunting activities with inherent risks, as still conducted by the Maasai96. Hominin entry into the carnivore guild through scavenging and hunting would have a profound effect on relations with predators. These involve direct competition for resources as well as passive scavenging from predator kills by hominins. Hominin advances in technology and social organisation, the latter in the form of cooperative hunting and defensive capabilities, may have influenced the drastic reduction in numbers of some carnivores, most probably through some human role in the depletion of prey species in conjunction with climate and vegetation change, as noted above. The disappearance of sabre- and scimitar-­ toothed cats and giant hyenas on both sides of the Mediterranean in the Middle Pleistocene “are contemporaneous with the first long-term hominin settlements. At that time, hominins hunted in groups and relied on new effective weapons; these two improvements allowed them to slaughter larger gregarious preys and

From the Pleistocene to modern humans  21

to handle encounters with dangerous competitors”97. The ability to confront and kill large carnivores like lions increased with hominin evolution in the Upper Pleistocene, even though there is evidence of continuing felid predation on hominins found in bone assemblages at Sterkfontein and Cova Negra in Spain98. The “status of hominins in the Ancient Paleolithic food chain could alternate between carrion and/or prey, scavenger and/or predator, depending on the circumstances rather than on their abilities”99. Increasing hominin consumption of animal protein is indicated by evidence from 2.5 ma in Ethiopia’s Awash Valley. Bones of antelopes and extinct horses from sites of hominin habitation show tool marks from dismemberment and defleshing100. The ability to acquire carcasses (through opportunistic scavenging, competitive scavenging or hunting) and process them using tools improved with the evolution of Homo species. Clark believes that the australopithecines and early toolmakers hunted small game and scavenged larger animals. Like chimpanzees, the hunting of small antelopes, species of pig and other small game was cooperative. This form of meat acquisition became more sophisticated and organised as the species developed101. Further evidence of hominin meat consumption has been discovered in bone assemblages in Kenya from 2.2 ma, indicating that immature individuals of small or medium-sized animals were being consumed. It is not clear whether they were scavenged or hunted102. Hominins in Southern Africa developed tool-making in the 1.7–1.1 ma period and in North Africa around 1.5–1.1 ma. Scavenging and/or hunting yielded meat but also bone, horn and antler that could be used for weapon-making. Around 1.0 ma, man is believed to have started utilising fire for warmth, protection from predators at night, cooking and for the production of fire-hardened wooden weapons and hunting103, perhaps in the way that fire was used 100 years ago in southern Sudan and northern Congo by the Azande to drive, surround and kill prey in a “ring of fire”104. An increase in hominin consumption of meat put them in competition with the Felidae species remaining in Africa (plus wild dogs and hyenas). Herbivore remains in fossil finds from after 1.8 ma rose from about 15–25% of assemblages to 45%105. The increase in meat consumption at a time when the expansion of open grasslands made ambush harder suggests the development of more efficient thrown or fired weapons, which would also be effective in deterring predators. Fossil evidence also shows increasing hominin body size as Homo erectus/ergaster evolved, with improved stamina for  h ­ unting, which increased the ranges over which prey could be hunted in grasslands, and with greater strength for communal defence against predators and in competing with them for food106. There is considerable controversy over the nature of hominin and early human scavenging and hunting; whether they started as purely opportunistic scavengers, taking away and feeding from the largely eaten-out remains of animals killed by predators, or whether they were competitive scavengers, too, with the confidence and combined strength to drive lions or hyenas from kills. There is also disagreement over the extent of their hunting. What seems clear, is that the

22  From the Pleistocene to modern humans

ability developed to use large stones as hammers to smash open bones to access the marrow. As Stiner suggests, moving from using stones to crack nuts or break up tough plant foods to smashing bones and scraping meat from them107 was a small technological step but an important behavioural advance that enabled greater animal protein consumption and encouraged scavenging and hunting. De Mello describes how the stone tools of Homo habilis, living 2.5 ma, were primarily for butchering carcasses rather than hunting and killing. Homo erectus, living about 1.8 ma–300,000 years ago, has been shown to have used stone weapons for hunting. The long-term effects of this change in tool use were huge, creating greater opportunities for protein consumption with consequent growth in body size, brain capacity and the ability to compete with dangerous predators. It was the start of the process that would develop into cooperative hunting of large animals with projectile weapons. At some stage snares, pitfalls and game pits would have been added to the array of hunting technologies, as shown in bone assemblages and artefacts from early San settlements in Southern Africa108. The Homo species expanded their range as their skills advanced, spreading to Eurasia and southern Asia. Their superior size, tool-making capabilities and intelligence enabled them to marginalise less advanced hominins/hominids and may have contributed to the extinction of australopithecines just after 1.5 ma109. ­Further advances in tool-making increased the ability of the Homo species to survive and adapt. The evolutionary process saw changes in physiology, notably bone/skull structure around 180,000–100,00 years ago, culminating in the appearance between 70,000–60,000 years ago, or possibly earlier as evidenced by the Moroccan fossils, of Homo sapiens. This broad period saw advances in technology with the production of more sophisticated bladed tools and weapons110. There is evidence that in some parts of the hominin range, hafted weapons appeared, that is weapons with blades or spear/arrowheads fitted into wooden shafts. The world’s oldest known spears, some 400,000 years old and from Germany, enabled early Homo species to hunt large animals and kill predators. Hafted weapons with sharp points appear in the African archaeological record 100,000–200,000 years ago111. Evidence uncovered by Mary Leakey suggests “groups of early hominids were never very large, but comprised a sufficient number of active males to form hunting bands and to protect the females and young in case of attack”112. There is evidence of “an inexorable progression” in the relationship of humans to large predators: from being prey to being scavengers (passive or competitive) and competitive hunters across this period, with initially it being most likely “that hunted man took advantage of predator kills”. Humans could have been active scavengers by day and vulnerable to predation at night113. Humans, as Hart and Sussman neatly summarise it, could scavenge without being primarily scavengers and hunt without being fully hunters – the flexibility of diet and methods of obtaining protein were huge advantages114. Scavenging was probably the initial means of obtaining meat and there is “considerable evidence from bone assemblages at sites where hominids and early man lived that they scavenged kills – but not totally clear whether these were

From the Pleistocene to modern humans  23

examples of passive scavenging of remains or active challenging of lions and other carnivores over kills”115. Treves and Naughton-Treves studied human-lion interactions in Uganda based on official records of people killed by lions between 1923 and 1994, and concluded that while groups of unarmed or lightly armed people could drive large predators from their kills, there were also regular casualties among hunter-gatherers who obtained meat this way. They “propose that twentieth-century predation on humans is likely to differ mainly in degree, not in kind, from that faced by Pleistocene hominids … [but that] it is by no means inconceivable that hominids with little more than stone tools or long sticks might have driven carnivores from kills, particularly if they acted in a cooperative and coordinated fashion”116. From all the evidence and reflections on current primate behaviour it appears likely that hominids and early Homo species mixed passive and confrontational scavenging – bringing them into conflict with lions and other predators – u ­ sing group cooperation and early weapons to intimidate and drive off predators, methods also used to defend themselves and food resources they had acquired through scavenging or hunting. Engagement in scavenging does not exclude hunting and as Domínguez-Rodrigo and Pickering conclude, “The coincidence of all of these inferred behaviours suggests that they evolved and functioned in concert, as part of a behavioural module or network, rather than that they emerged independently at different times over the course of the evolution of the genus Homo.”117 Examination of the hunter-gatherer activities of the contemporary Hadza community in northern Tanzania, in an area with sufficient wildlife to support this lifestyle, reinforces the view that early humans could have developed a combination of strategies. Hadza engage in passive scavenging of abandoned kills, searching out recent kills and obtaining the remaining meat by driving off predators. Bunn et al.’s observations of the Hadza show them obtaining meat from lion and leopard kills. They might drive off the lions, but if they are too numerous and determined, the Hadza wait until the lions have fed enough to enable the Hadza to take the carcass without serious risk of injury118. Hawkes’s study of the Hadza confirms their ability to drive lions from kills with basic weapons such as spears or arrows and that on 11 occasions they were seen to accomplish this successfully, once killing one of the lions when it resisted119. Domínguez-­Rodrigo’s studies from the Masai Mara and Tsavo also support the view that organised groups of hominins could drive small numbers of lions from kills and appropriate the carcasses while they still had significant quantities of meat on them. He relates this convincingly to the marks left on herbivore bones in bone assemblages from the time of early humans and says it supports the hypothesis that hominids and then hominins engaged in hunting and confrontational scavenging as well as opportunistic scavenging120. One of the leading authorities on modern lions, George Schaller, also takes the view that hominin ability in competing for carcasses contributed to aggressive patterns of behaviour towards predators and they might have used weapons

24  From the Pleistocene to modern humans

to intimidate and, if necessary, attack predators, as demonstrated by the presence of predator skulls dating back as far as sabre-tooth cats in bone assemblages at Makapansgat in South Africa121. Other studies also support the ability of both social hominids and then hominins to obtain meat through “the wresting of animals killed by other predators”122. Clearly, from hominids through hominins to Homo species, a pattern of behaviour emerged combining defence against predators, strength in numbers and the confidence to compete for carcasses with lions and other large predators, and the ability and technology to hunt larger prey species and retain the carcasses. In Europe, it is very possible that increasing human hunting, scavenging and cooperative defensive capabilities may have influenced the decline of the cave lion (P spelaea). It appears to have gone into irreversible decline largely as a result of the disappearance of the European bison over much of the continent, as a result of climate/vegetation change and human hunting. Prior to their extinction, Paleolithic paintings from the Chauvet caves in the Ardeche region of southern France indicate the keen observation of cave lions by the humans who shared the area with them. Some paintings and the human remains found near them date back 40,000 years, well before the demise of the cave lion123. The paintings show that early humans lived alongside not just lions but mammoths, bison, aurochs, rhinos, giant elk and wild horses, which were prey for lions and people. ­A rchaeologists have found stone tools, reindeer-antler spearheads and evidence that humans hunted reindeer and bison, and that cave lions, wolves, leopards and hyenas may have preyed on humans, who lived in small, scattered communities124. Packer and Clotes point out in their study of the paintings the accuracy of the representations of the lions, indicating close and interested observation, achieving what they refer to as a “snapshot of lion behavior, a realistic representation of a true-life event the painter witnessed firsthand”125. The very detailed paintings, dating to around 32,000 years ago, show examples of mating behaviour and that males lacked manes. Over the millennia covered in this chapter, it is evident that the evolution of hominids into modern humans took place in parallel with the evolution of modern lions from early Panthera species and a clear relationship developed both direct through conflict between them, and indirect though human hunting of lion prey species. From being the prey of the ancient lions and P leo itself, humans became scavengers from kills, competitors trying to wrest lion kills from them and then entrants to the carnivore guild, competing for prey with lions.

Notes 1 Richard G. Klein (2000), The Earlier Stone Age of Southern Africa, South African Archaeological Bulletin 55: 107–122, p. 107. 2 Ibid. 3 https://wgnhs.uwex.edu/wisconsin-geology/ice-age/ accessed 7 June 2017. 4 David Macdonald (1992), The Velvet Claw. A Natural History of the Carnivores, London: BBC Books, pp. 22–3.

From the Pleistocene to modern humans  25

5 Ibid. 6 C.A.W. Guggisberg (1961), Simba. The Life of the Lion, London: Bailey Bros and Swinfen, p. 15. 7 Ibid., p. 23 8 Ibid., p. 26. 9 Robin S. Reid (2012), Savannas of Our Birth. People, Wildlife and Change in East Africa, Berkeley: University of California Press, p. 83. 10 Ibid., p. 83. 11 Guggisberg, 1961, p. 17. 12 See Alan Turner and Mauricio Anton (1997), The Big Cats and Their Fossil Relatives, New York: Columbia University Press. 13 R.J.G. Savage (1979), Carnivora, in Vincent J. Maglio and H.B.S. Cooke (Eds), Evolution of African Mammals, Cambridge, MA: Harvard University Press, 245–267, p. 261. 14 Ibid., p. 262. 15 See Brian Brain (1981), The Hunters or the Hunted? An Introduction to African Cave Taphonomy, Chicago: University of Chicago Press; Julia Lee-Thorp, J. Francis ­Thackeray and Nikolaas van der Merwe (2000), The hunters and the hunted revisited, Journal of Human Evolution, 39, 6, December, 565–576; and Linda Piegl and ­Bianca Bothma (2011), Dinofelis – hominid hunter or misunderstood feline?, ­Maropeng, 20 December. 16 Nobuyuki Yamaguchi et al. (2004), Evolution of the mane and group-living in the lion (Panthera leo): a review, Journal of Zoology, 263: 329–342, p. 331. 17 Ibid. 18 Ibid. 19 Ross Barnett et al. (2006), The origin, current diversity and future conservation of the modern lion (Panthera leo), Proceedings of the Royal Society, 273: 2119–2125, p. 2119. 20 Yamaguchi et al., 2004, p. 331. 21 Ross Barnett et al. (2014), Revealing the maternal demographic history of Panthera leo using ancient DNA and a spatially explicit genealogical analysis, Evolutionary Biology, 14, 70, DOI: 10.1186/1471-2148-14-70 accessed 7 June 2017, p. 1. 22 Ibid. 23 Ross Barnett et al. (2016), Mitogenomics of the extinct cave lion, Panthera spelaea (Goldfuss, 1810), Resolve its position within the Panthera cats, Open Quaternary, 2: 4. 24 Ibid. 25 Macdonald, 1992, p. 62. 26 Ibid., pp. 62–3. 27 Ibid., pp. 2–3; see also, Elaine Anderson (1984), Who’s who in the Pleistocene: a mammal bestiary, in Paul S. Martin and Richard G. Klein (Eds), Quaternary Extinctions. A Prehistoric Revolution, I, Tucson: University of Arizona Press, 40–89, pp. 57–9. 28 Larisa R.G. DeSantis et al. (2012), Implications of diet for the extinction of sabertoothed cats and American lions, PloS ONE, December, 7, 12: p. 1. 29 Richard G. Klein (1984), Mammalian extinctions and Stone Age people in Africa, in Martin and Klein, pp. 553–573, pp. 560–3. 30 Ibid., p. 568. 31 Donald K. Grayson and David J. Meltzer (2003), A requiem for North American overkill, Journal of Archaeological Science, 30: 585–593, p. 585. 32 See P.S. Martin (1958), Pleistocene ecology and biogeography of North America, in C.L. Hubbs (Ed.), Zoogeography, American Association for the Advancement of Science, Washington, DC, 375–420; P.S. Martin (1967), Prehistoric overkill, in P.S. Martin and H.E. Wright Jr. (Eds), Pleistocene Extinctions: The Search for a Cause, New Haven, CT: Yale University Press, 354–403. 33 Macdonald, 1992, p. 62; and Alan Turner and Mauricio Anton (1997), The Big Cats and Their Fossil Relatives, New York: Columbia University Press, pp. 72 and 76.

26  From the Pleistocene to modern humans

34 Jerry N. McDonald (1984), The reordered North American selection regime and Late Quaternary megafaunal extinctions, in Martin and Klein, 404–459. 35 Hans Kruuk (2002), Hunter and Hunted. Relationships between Carnivores and People, Cambridge: Cambridge University Press, p. 115. 36 The author is indebted to Laurence Frank for this view of extinctions. 37 Christopher James Sandom et al. (2017), Learning from the past to prepare for the future: felids face continued threat from declining prey richness, Ecography, April. 38 Turner and Anton, 1997, p. 72; and, Miriam Mariana Morales and Norberto ­Pedro Giannini (2014), Pleistocene extinctions and the perceived morphofunctional structure of the neotropical felid ensemble, Journal of Mammalian Evolution, 21: 395–405, p. 396. 39 Maria Rita Palombo et al. (2008), Carnivora dispersal in Western Mediterranean during the last 2.6 Ma, Quaternary International, 179: 176–189, pp. 185–9. 40 N.K. Vereshchagin and G.F. Baryshnikov (1984), Quaternary mammalian extinctions in Northern Eurasia in Martin and Klein, 483–516, pp. 497. 41 Personal communication. 42 Barnett et al., 2014, p. 4. 43 Ibid. See also, Agostinho Antunes et al. (2008), The evolutionary dynamics of the lion Panthera leo, PLoS Genetics, 1 November, 4, 11. 44 Barnett et al., 2014, p. 7. 45 Ibid., p. 5. 46 IUCN, Panthera leo (West Africa subpopulation), http://dx.doi.org/10.2305/IUCN. UK.2015-2.RLTS.T68933833A54067639.en, accessed 29 December 2017. 47 R.S. Sommer and N. Benecke (2006), Late Pleistocene and Holocene development of the felid fauna (Felidae) of Europe: a review, Journal of Zoology, 269: 7–19, p. 8. 48 M. Massetti and Paul P.A. Mazza (2013), Western European Quaternary lions: new working hypotheses, Biological Journal of the Linnean Society, 109: 66–77, p. 66. 49 Vereshchagin and Baryshnikov, 1984, 483–516, p. 497. 50 Sommer and Benecke, 2006, p. 15. 51 Yamaguchi et al., 2004, p. 332. 52 Deidre Jackson (2010), Lion, London: Reaktion Books, p. 9. 53 Bienvenido Martinez-Navarro et al. (2009), The large carnivores from ’Ubeidiya (early Pleistocene, Israel): biocrhonological and biogeographical implications, Journal of Human Evolution, 56: 514–524, p. 514. 54 L. Werdelin and M.E. Lewis (2013), Temporal change in functional richness and evenness in the eastern African Plio-Pleistocene carnivoran guild, PLoS ONE, 8, 3, p. 1. 55 Ibid., p. 8. 56 Ibid. 57 Yamaguchi et al., 2004, p. 330. 58 Annik E. Schnitzler (2011), Past and present distribution of the North African– Asian lion subgroup: a review, Mammal Review, 41, 3: 220–243, p. 224. 59 Sudipta Mitra (2005), Gir Forest and the Saga of the Asiatic Lion, New Delhi: Indus, p. 36. 60 Valmik Thapur, Romila Thapar and Yusuf Ansari (2013), Exotic Aliens. The Lion and Cheetah in India, New Delhi: Aleph, pp. 232–3. 61 Agostinho Antunes et al., 2008, pp. 6–7. 62 Ibid. 63 Yamaguchi et al., 2004, p. 333. 64 Ibid., p. 334. 65 Ibid., p. 35. 66 L.D. Bertola1 et al. (2011), Genetic diversity, evolutionary history and implications for conservation of the lion (Panthera leo) in West and Central Africa, Journal of Biogeography, 38: 1356–1367, pp. 1362–3. 67 Ibid., p. 1365. 68 Hannah Devlin (2018), Tracing the tangled tracks of humankind’s evolutionary journey, The Guardian, 12 February, https://www.theguardian.com/news/2018/

From the Pleistocene to modern humans  27

feb/12/tracing-the-tangled-tracks-of-humankinds-evolutionary-journey, accessed 13 February 2018. 69 Eileen M. O’Brien and Charles R. Peters (1999), Landforms, climate, ecographic mosaics, and the potential for hominid diversity in Pliocene Africa, in Timothy Bromage and Friedemann Schrenk (Eds), African Biogeography, Climate Change, and Human Evolution, New York: Oxford University Press, 115–137, pp. 134–5. 70 Ibid., pp. 134–5. 71 Desmond Clark (1976), African origins of man the toolmaker, in Glynn Ll. Isaac and Elizabeth R. McCown (Eds), Human Origins: Louis Leakey and the East African Evidence, Menlo Park, CA: Staples Press, 1–53, pp. 5–6. 72 Ibid., p. 6. 73 Norman Owen-Smith (1999), Ecological links between African savanna environments, climate change, and early hominid evolution, in Bromage and Schrenk, 138–149, p. 138. 74 Ibid., p. 142. 75 Author’s personal communication with Dr Laurence Frank concerning the megafaunal extinctions. 76 Owen-Smith, 1999, p. 148. 77 Ibid. 78 Ibid. 79 John Iliffe (2007), Africans. The History of a Continent, Cambridge: Cambridge University Press, 2nd edition, p. 6; and, M. Brunet et al. (2005), A new hominid from the Upper Miocene of Chad, Central Africa, Nature, 418, pp.  145–51 and C.P.E. Zollikofer et al. (2005), Virtual cranial reconstruction of Sahelanthropus tchadensis, Nature, 434, pp. 755–9. 80 Christopher Ehret (2016), The Civilizations of Africa. A History to 1800, ­Charlottesville: University of Virginia Press, pp. 16–17. 81 L.S.B. Leakey (1961), The Progress and Evolution of Man in Africa, Oxford: Oxford University Press, pp. 9 and 40. 82 Iliffe, 2007, pp. 7–8. 83 Jean-Jacques Hublin et al. (2017), New fossils from Jebel Irhoud, Morocco and the pan-African origin of Homo sapiens, Nature, 546, 289–292, 08 June, p. 289. 84 Marlize Lombard (2017), Ancient DNA increases the genetic time of modern humans, The Conversation, 13 October, https://theconversation.com/ancient-dna-­increasesthe-genetic-time-depth-of-modern-humans-84716, accessed 13 November 2017. 85 Schnitzler, 2011, p. 221. 86 Paul Palmqvist et al. (2007), A re-evaluation of the diversity of megantereon (­m ammallia, carnivora, machairodontinae) and the problem of species identification in extinct carnivores, Journal of Vertebrate Paleontology, 27, 1, pp. 160–75. 87 Mary C. Stiner (2012), Competition theory and the case for Pleistocene hominin-­ carnivore co-evolution, Journal of Taphonomy, 10, 3–4: 129–145, p. 129. 88 Ibid. 89 Ibid., pp. 130–1. 90 Ibid., p. 133. 91 C. Daujeard et al. (2016), Pleistocene hominins as a resource for carnivores: a c.500,000-year-old human femur bearing tooth marks in North Africa (Thomas Quarry I, Morocco), PloS ONE 11, 4, p. 1. 92 Jordi Rosell et al. (2012), New insights on hominid-carnivore interactions during the Pleistocene, Journal of Taphonomy, 10, 3–4, p. 127. 93 Kruuk, 2002, pp. 103–4. 94 M. Cueto et al. (2016), Under the skin of a lion: unique evidence of Upper Paleolithic exploitation and use of cave lion (Panthera spelaea) from the Lower Gallery of La Garma (Spain), PLoS ONE, 11, 10. 95 Ibid. 96 Ibid.

28  From the Pleistocene to modern humans

97 Ibid. 98 Ibid., p. 2. 99 Ibid., p. 16. 100 Lawrence Barham and Peter Mitchell (2008), The First Africans. African Archaeology from the Earliest Toolmakers to Most Recent Foragers, Cambridge: Cambridge University Press, Kindle edition, p. 95. 101 Clark, 1976, p. 24. 102 Barham and Mitchell, 2008, p. 114. 103 Francesco Berna et al. (2012), Microstratigraphic evidence of in situ fire in the Acheulean strata of Wonderwerk Cave, Northern Cape province, South Africa, Proceedings of the National Academy of Sciences, 109(20): E1215–E1220. doi:10.1073/ pnas.1117620109, accessed 28 March 2017. 104 Keith Somerville (2016), Ivory. Power and Poaching in Africa, London: Hurst, p. 82. 105 Barham and Mitchell, 2008, p. 146. 106 Ibid., p. 147. 107 Stiner, 2012, p. 134. 108 G. Mokhtar (Ed.) (1990), General History of Africa. II Ancient Civilizations of Africa, London: James Currey/UNESCO, p. 353. 109 Ehret, 2016, p. 17. 110 Ibid., pp. 20–1. 111 Reid, 2012, p. 93. 112 Mary D. Leakey (1976), A summary and discussion of the archaeological evidence from Bed I and Bed II, Olduvai Gorge, Tanzania, in Isaac and McCown, 431–459, p. 434. 113 Donna Hart and Robert W. Sussman (2009), Man the Hunted. Primates, Predators, and Human Evolution, Philadelphia, PA: Westview Press, p. xi and p. 5. 114 Ibid., p. 15. 115 A. Treves and L. Naughton-Treves (1999), Risk and opportunity for humans coexisting with large carnivores, Journal of Human Evolution, 36: 275–282, p. 275. 116 Ibid., p. 281. 117 Manuel Domínguez-Rodrigo and Travis Rayne Pickering (2017), The meat of the matter: an evolutionary perspective on human carnivory, Azania: Archaeological Research in Africa, 52, 1: 4–32, p. 23. 118 Henry T. Bunn et al. (1988), Variability in bone assemblage formation from Hadza hunting, scavenging, and carcass processing, Journal of Anthropological Archaeology, 7, 4: 412–457, p. 427. 119 Kristen Hawkes (2016), Ethnoarchaeology and Plio-Pleistocene sites: some lessons from the Hadza, Journal of Anthropological Archaeology, 44: 158–165, p. 160. 120 Manuel Domínguez-Rodrigo (1997), Meat-eating by early hominids at the FLK 22 Zinjanthropus site, Olduvai Gorge (Tanzania): an experimental approach using cutmark data, Journal of Human Evolution, 33, 6: 669–690, p. 671. 121 George B. Schaller and Gordon R. Lowther (1969), The relevance of carnivore behavior to the study of early hominids, Southwestern Journal of Anthropology, 25, 4: 307–341, p. 324; see also, R. Dart (1956), The cultural status of the South African man-apes, Smithsonian Institution Report for 1955, 317–338, p. 329. 122 Alan Turner (1988), Relative scavenging opportunities for East and South African Plio-Plestocene hominids, Journal of Archaeological Science, 15: 327–341, p. 327. 123 Craig Packer and Jean Clotes (2000), When lions ruled France, Natural History, ­November, issue 9: 52–57, p. 52. 124 Ibid., pp. 52–3. 125 Ibid.

2 Domestication, settlement and the effects on lions

Human evolution and the increasing sophistication of hunting methods substantially affected the relationship between people and lions, while domestication of animals, cultivation of crops and expansion of human settlements brought qualitative and quantitative changes in their interactions. The clearing of land for cultivation and the need to protect livestock from predators were new factors, changing the balance of conflict and power This chapter will focus mainly on Africa and how developments in human society and livelihoods shaped relationships with lions, wildlife and the environment. The decline of the lion in Europe and the Middle East will not be ignored, with analysis of the disappearance of the lion from much of its range outside Africa, as well as of the exploitation of lions for brutal forms of entertainment, as symbols of power or as a quarry hunted for pleasure and prestige.

The taming of the land and some of its beasts Around 25,000–20,000 BCE what Reid calls the “soft boundary” between people and their environment, with people gathering wild plant foods and hunting/­ scavenging for meat, was about to start evolving into a “hard boundary”, with domestication of animals and cultivation of plants for food1. As early as 19,000– 20,000 years ago, according to evidence from the Nile Valley, people started intensive exploitation of tubers and fish. Around 15,000 BCE communities along the northern Nile started harvesting wild cereals2. To the west of the Nile, peoples of the Nilo-Saharan and Niger-Kordofan language groups pursued livelihoods based on foraging, hunting and fishing. They developed tool-making, hunting with spears, bows and arrows3.

30  Domestication, settlement and lions

Over the next few millennia, as Africa’s climate became hotter and wetter, flooding in the Nile Valley encouraged settlement on surrounding, higher plains. Iliffe says by 12,000–7,500 years ago, this led communities to move away from river valleys4. Domestication of wild cattle increased availability of food; dependence on hunting or foraging declined. The raising of cattle may have spread to Egypt and the Nile Valley from the Middle East, where there is most of the early evidence of domestication5. Livestock and arable farming in the Nile ­Valley, the Horn and East Africa reduced the need to hunt but created the need to defend livestock from predators. Pastoralists killed or drove off lions, while the cultivation of land reduced grazing for lions’ wild prey. As Bertram suggests, “Herbivores which would feed on a farmer’s crops are either destroyed or kept out by fences” once more-settled cultivation developed6. Serious anthropogenic threats to lions emerged through human economic progress. Over the next 5,000 years, cattle-raising spread and the role of humans as engineers of the environment increasingly affected wildlife. Archaeological evidence from 9500 to 5000 BCE indicates people were raising animals and growing plants for food in the south-eastern Sahara, the savannas of West Africa and the Ethiopian highlands7. Archaeological finds around Lake Chad show the spread of livestock to highland areas of central Sahara with sufficient rainfall8. The subsequent drying of the climate across Africa and resulting aridity of much of the Sahara forced people and their livestock to move east, west and south. Some communities adopted a nomadic existence, utilising seasonal grazing. Ehret says by the 8th millennium BCE, Saharo-Sudanese people of the eastern Sahara were cultivating grains previously gathered from the wild and building thorn fencing to protect homesteads and cattle from predators9. Settled and nomadic communities still hunted to supplement food production. As people moved into the Sahel and along the Nile Valley, livestock-keeping communities settled in Sudan about 6,000 years ago, spreading over the next 1,000–2,000 years into Ethiopia and then Kenya, with goats and sheep raised alongside cattle10. Iliffe suggests metalworking in Africa started in the Nile Valley in the 5th millennium BCE, possibly introduced by migrants from North Africa or the Middle East11. Communities labelled as Cushitic moved into the Horn and Kenya around 5,000–4,000 years ago and pushed south into northern T ­ anzania after 3000 BCE, living alongside the resident Khoisan hunter-­g atherers12. By 1000 BCE, Nilotic peoples, including ancestors of the Kalenjin-speakers of Kenya, had moved south with cattle and other livestock into Kenya and northern Tanzania. Some communities in East and Southern Africa, the ancestors of the people who became known as Ndorobo, Hadza and San/Bushmen, retained hunter-gatherer modes of subsistence and did not adopt cultivation or livestock raising13. Some of these communities were assimilated by incoming Cushitic and then Bantu-speaking farming or pastoral communities, but many were pushed aside into areas unsuitable for crops or livestock14. Many of the pastoral communities engaged in transhumance, moving their cattle in search of grazing and water. Ehret estimates that by 7000 BCE, “the

Domestication, settlement and lions  31

typical Saharo-Sahelian extended family of the eastern Sahara region resided in a large homestead surrounded by a thick thornbush fence. The enclosed area acted as a cattle pen, protecting animals at night from predators, principally lions and hyenas”15. The growth of cultivation and pastoralism enabled increases in the size of settlements in savanna areas. Food production was more reliable than foraging, though this remained an option when drought or other factors limited food production. This substantially changed humankind’s relationship with wildlife16. Protection of crops from wild pigs, antelopes and other herbivores and defence of livestock from predators became a necessity. Defending stock, responding to attacks and pre-emptive aggression were becoming part of the picture rather than the previous need simply to defend against being preyed upon, and competition over carcasses or for wild prey. Despite gradual growth in human populations, Iliffe emphasises that until the 20th century CE, “Africa was an underpopulated continent”17. Low population density enabled nomadism, transhumance and shifting cultivation because of lack of competition for land and absence of defended territories. This limited the depletion of wildlife through hunting or the presence of large human and livestock populations. From the outset, and this is the case for much of the narrative before the arrival of Europeans in Africa, it should be noted, echoing the historian Joseph Ki-Zerbo, that a major problem facing historians of sub-Saharan Africa is that “there is scant oral, linguistic or archaeological data”18. What follows is based on what can be gleaned from a range of oral history, archaeological, anthropological and historical studies. Few directly mention human-wildlife conflict and there is inevitably a speculative element in interpreting how human societal development impinged on lion populations.

East Africa In East Africa, pastoralism preceded widespread cultivation of crops; chiefly because settlement and migration from the north and west was into grassland or woodland/savanna, where unreliable rainfall and thinner soils limited crop yields. Pastoralism enabled people to move to follow the rains. Mobility limited conflict with wildlife and did not involve the permanent exclusion of herbivores or predators from specific areas, as settled farming did. Pastoralists still came into conflict with carnivores and their prey – wild ungulates competed for food with livestock and carnivores preyed on stock. As Reid suggests, the herders who moved with their herds and flocks into eastern and then southern Africa, “likely considered wildlife more of a problem than their hunting-only neighbours”19. There is no clear archaeological or other evidence showing the extent to which the shift from hunter-gatherer to pastoral production affected the abundance or the diversity of wildlife. One barrier to the spread of pastoralism favouring the survival of wildlife were the tsetse fly belts in East and Southern Africa 20. The fly transmits sleeping sickness to humans and a disease called nagana to livestock, which can cause

32  Domestication, settlement and lions

weight loss and death. Pastoralism spread, avoiding tsetse regions, from Kenya and Uganda into Tanzania by the 2nd millennium BCE, and then south-central Africa 21. There is evidence that pastoralists were present in the Serengeti from as early as 4000 BCE. Excavations show seasonal occupation of sites on grass plains. Use of the area for seasonal grazing continued right up to the Datoga pastoralists, about 200 years ago22, before they were pushed out by the Maasai 23. Bertram believes the presence of tsetse flies in the Serengeti’s woodlands prevented largescale settlement, enabling lions and wild ungulates to thrive24. By 2000 BCE, pastoralist communities keeping sheep, goats and cattle were widespread along the Rift Valley through Kenya and into northern, western and central Tanzania 25. The area was rich in wildlife, including buffalo, a wide range of antelopes, zebra, wild pigs and predators like lions, hyenas, leopards, cheetahs and wild dogs. Predators coexisted with hunter-gatherer communities – the predecessors of the Ndorobo and Hadza competed for prey and scavenged lion kills but rarely killed lions. Archaeological excavations in south Nyanza in Kenya indicate that although pastoralism was practised there by the 2nd millennium BCE, remains of domestic and wild animals from Neolithic sites show hunting persisted and “the pattern of wild animal taxa present is suggestive of a combination of widespread hunting of the more solitary species (e.g. oribi and reedbuck) and selective hunting or scavenging of larger herd animals like zebra, hartebeest and topi”26. No evidence is provided about human-lion interactions. A reason for the paucity of evidence of the nature of the relationship, hunting and livestock defence systems at the time, as Mutundu explains, is “The mobility of both early pastoralists and later hunter-gatherers and their interactions [which] also means that the archaeological records of these groups are not always easy to find or distinguish”27. A problem associated with the study of pastoral economies in the region is the “difficulty of identifying and distinguishing archaeological sequences of hunter-gatherers with access to domestic stock or in the early phases of the adoption of herding, from those of pastoralists practising a subsistence strategy that included the utilisation of wild animals28. Robertshaw emphasised this: “Faunal bone assemblages from areas that adopted pastoralism suggest the decline but not disappearance in hunting for food”29. Boles and Lane conclude from their comparison of early pastoral settlements in Kenya and Tanzania and current practices that, “Among the Maasai, and other pastoral societies in eastern Africa, settlements are constructed first by clearing an area of vegetation, before erecting houses and fences around a central corral. Dung accumulates within this enclosure, enhancing nutrient content in local soils. This enrichment fosters distinctive sequences of vegetation succession, with rapid recolonisation of these glade sites by rich grasses that attract grazing animals, both wild and domestic,”30 suggesting early settlement didn’t have a catastrophic effect on wildlife. The climate of much of Africa became more arid from the 4th millennium BCE. Open savanna expanded at the expense of woodland, favouring pastoralism and providing suitable conditions for large ungulates and social carnivores like lions31. Crop farming did not disappear totally with this change in climate,

Domestication, settlement and lions  33

developing alongside pastoralism in areas with more reliable rainfall. This provided conditions for mixed farming economies and growth in cultivation of a variety of plant foods, which took place as Bantu-speaking peoples spread from west-central Africa into Congo and the western Rift Valley and eventually on to East and Southern Africa 32. There is evidence from the archaeological record that pastoralists, hunter-gatherers and arable farmers lived alongside a diversity of wildlife in the Kenya-Tanzania region33. Gifford et al. provide evidence of wild ungulates around Nakuru-Naivasha in the last two millennia BCE from bone assemblages showing “a diverse range of herbivores living in close proximity to the neolithic pastoralists … [with] numbers of zebra, wildebeeste, and, to a lesser degree, kongoni … impala, Grant’s and Thomson’s gazelle populations”34. The development of a predominantly pastoralist economy in Kenya and Tanzania’s savanna and Rift Valley regions occurred after 1000 BCE35. Farming, hunting and defensive capabilities improved when Stone Age technologies were replaced by iron-working over several thousand years, as detected in excavations in the Galana region north of Tsavo East National Park in Kenya 36. Iron tools enabled more extensive clearing of forests for cultivation. This was linked, in the period between 3000 and 1000 BCE, with the arrival of Bantu migrants from Central Africa in the Western Great Lakes and Rift Valley37. The Bantu migrants assimilated iron-working skills from Nilo-Saharan peoples who had moved south, and transferred to them methods of crop cultivation 38. In East Africa, this combination of migration and technological advance enabled growth in human settlements, increasing the pressure on wildlife as more land was cultivated 39. Over the next three millennia, right up to the modern period, expansion of cultivated areas, livestock herds and human settlements became the major threat to habitats, wildlife and, particularly, predators like lions. The agricultural and technological innovations in the region were complemented by a growth in inter-communal trade, and commerce along the Indian Ocean coast at the end of the BCE and beginning of the CE period. Trade with Mediterranean civilisations and the Arabian Peninsula introduced more sophisticated iron weapons and tools40, and demand for wildlife products such as skins, ivory and rhino horn. From 300–1000 CE, ancestors of the Kalenjin-speaking peoples of the Rift Valley of Kenya, often referred to as Sirikwa, moved into the region. They were primarily cattle-keepers and had societal structures based on male age-grades. As young men reached the age for transition to manhood they were initiated as muren and lived together in warrior age sets – vital both for defence of the all-­ important cattle from predators and raiders from other communities, but also for raiding others for cattle and hunting41. Around 700 CE, Maa-Ongamo people (the ancestors of the Maasai and Samburu) moved from the Lake Turkana Basin into plateau areas and plains east of the Rift Valley. Ehret says that one aspect of the development of the young warrior class among Maa and Kalenjin speakers was the emergence of rite-of-passage hunts for lions, during which initiates proved their valour42.

34  Domestication, settlement and lions

Scholars believe the Maasai were originally agro-pastoralists cultivating sorghum and millet and raising cattle and smallstock. Over time, a core group specialised in cattle-raising, developing a self-identity as pastoralists. Other Maa speakers developed into communities with a variety of economic specialities – pastoralist, agro-pastoralist, farmers and hunter-gatherers43. Steinhart warns against mixing myth and reality when portraying the Maasai. He says that they don’t routinely hunt wildlife for food and while they do kill lions that attack cattle or people – “they are pursued and killed by young warriors (moran) as part of the rites of passage into manhood, requiring the single-handed and courageous confronting of a dangerous enemy” – the extent of this is often exaggerated in Western portrayals of them, and there is no evidence to suggest that it had a substantial effect on lion numbers44. Over several centuries, Maa speakers became increasingly dominant on the Laikipia plateau and grassland areas, through the military capabilities of their age-sets and their skill in iron weapon production; making heavier socketed spears that were more effective in hunting lions than hafted spears45. The Maasai were far from the only pastoralist or mixed farming community to have to deal with the threat posed by lions, but they have been one of the most extensively studied and research into their oral history provides interesting material on lions. Kipury says Maasai customs and folk tales reveal much about the abundance of wildlife in the areas of Kenya and Tanzania to which they migrated46. She tells of the ritual lion hunts (olamayio) carried out by young men as part of their initiation as ilmurran or warriors. There was, according to their own accounts, little hunting of wild animals for food except in times of great need47. Lions and other predators would be killed to protect cattle or people or, periodically, as part of rituals, but rituals with the purpose of reinforcing the capability of the ilmurra to protect their communities48. Wildlife products were utilised widely, such as buffalo hide for shields, horn for snuff containers, lion manes and ostrich feathers for decoration and giraffe tails as fly-whisks49. Lions appear in Maasai folk stories as bullies and animals that are easily tricked. Kipury believes that for the Maasai in their oral histories and myths, “the lion represents power and authority, which tricksters strive to bring down”50. By 1000 CE, pastoralist communities occupied grassland and woodland-­ savanna areas of western Uganda and the Great Lakes, which were rich in wildlife51. Some developed systems with ownership of cattle a dominant factor in the growth of a caste of hereditary rulers (notably the Nkore of southern Uganda and Tutsi of Rwanda and Burundi)52. Between 700 CE and 1200 CE, the Bunyoro, Toro and Buganda kingdoms developed through the assimilation of Bantu migrating from the west with Sudanic pastoralists moving south53. These were more hierarchical, established permanent settlements and progressively pushed wildlife from areas of cultivation. The expanding populations of cultivator/­pastoralists cut down large areas of woodland to provide cultivable land or grassland on which to graze cattle54. This did not totally exclude wildlife from areas of human settlement but started the process of habitat loss. The establishment between

Domestication, settlement and lions  35

the 7th and 11th centuries CE of coastal trading communities along the Indian Ocean had a strong influence on human-wildlife relations. The ­Swahili trading communities, Arab and Persian merchants (trading with Arabia and West Asia) settled in ports like Lamu, Mombasa, Bagamoyo, Kilwa and Zanzibar. They traded in lion and leopard skins, hides and ivory, exporting them to markets in the East55. The value of wildlife products stimulated hunting for trade rather than just subsistence among communities like the Ndorobo, Waliangulu and Wakamba of Kenya and the Nyamwezi of central Tanzania; some became both hunters and commercial middlemen, supplying wildlife products from inland to coastal traders. West of the Serengeti and in what is now the Western Corridor of Serengeti National Park, there was considerable human settlement in the first two millennia CE. Shetler’s research shows the presence of the ancestors of the Ikoma, Nata, Ikizu, Ishenyi and Ngoreme peoples there, dispelling myths deriving from 19th century European travellers saying these areas were largely unpopulated and solely the domain of wildlife56. People coexisted with wildlife, though human settlements were never that extensive and farming/pastoralism did not involve use of fencing to keep wild ungulates or predators away from farmland. Shetler says research indicates that “Visible evidence of the long-term interaction of humans with the Serengeti environment can also be found in the very presence of the large mammal herds that tourists come to see. Humans have coevolved with the wildlife over millennia and have therefore learned to coexist”57. ­Further east, around Lake Eyasi and parts of the eastern Serengeti, lived the Hadza hunter-­ gatherers. They hunted for meat, scavenged from lion and other predator kills and occasionally killed lions in self-defence or during aggressive scavenging.

Southern Africa Constructing an evidence-based account of the relationship between the humans in Southern Africa and lions and other wildlife is not easy and again involves piecing together fragments to try to provide viable interpretations. The historian, David Beach, warns that sources on the development of communities and their relationship to their environment are thin and one has to be careful not to adopt “an over-enthusiastic use of traditions to reconstruct the past”58. Traditions and myths develop that are used in contemporary nation-building that are not verifiable and may be exaggerations, distortions or even inventions59. Many communities in the region, like the Shona of Zimbabwe, had no strongly established system of traditional historians or custodians of the past60. Looking at the settlement of southern Africa since 12,000–10,000 BCE, Khoikhoi peoples moved south from near Lake Malawi through eastern Namibia and the Okavango. Others migrated from the northern end of Lake Malawi via the Zambezi and Limpopo. Later contact with other communities, including early Bantu migrants, may have influenced Khoikhoi adoption of pastoralism61. The San and Khoikhoi had a culture of cooperative foraging, hunting and

36  Domestication, settlement and lions

scavenging from lion and other carnivore kills62. Late Stone Age hunter-gatherer cultures disappeared when contact with Bantu communities migrating southwards around 300–200 BCE replaced stone-tool making with metalworking, producing better weapons and tools63. The San, though, remained dependent on hunting and foraging for wild foods64. They lived in constant contact with wildlife, whether as prey, competitors for carcasses or threats to people. Despite regular contact with them, San oral traditions and tales have few references to lions, whereas the mantis, the mongoose, the hartebeest and baboon are frequently important characters in stories65. Over time, the Khoikhoi of southern Africa became owners of large herds of cattle and smallstock, with hunting a supplementary activity66. Bantu agro-pastoralists settled in the eastern half of southern Africa about 2,000 years ago. They grew sorghum and millet, established permanent homesteads, raised cattle, sheep and goats and forged iron67. The western half of South Africa was not suitable for settlement and the San and Khoikhoi communities were left largely undisturbed there until Bantu groups moved south through Angola, and the arrival of European settlers with the Dutch establishment of a provisioning station at Cape Town in 1652. The forging of iron tools enabled more efficient hunting and facilitated the clearance of woodland, expansion of agriculture and establishment of larger settlements in south-east Africa by the end of the 2nd century CE68. By 500 CE, pastoralism was expanding and in the Zoutpansberg area of northern Transvaal, communities were building homesteads of circular huts around central cattle pens to protect against large predators69. To the south-west, in Botswana, sufficient rainfall enabled the development of a pastoral culture known as the Toutswe tradition70, based on similar settlements around cattle pens – defence of the all-important cattle vital to food security. To the west of the Kalahari, nomadic pastoralism was dominant in the 1st millennium CE, in an area too dry for extensive cultivation71. By 700 CE, Bantu-speaking communities had established settled communities in south-west Zimbabwe/north-eastern Botswana/ northern South Africa in what became known as the Leopard’s Kopje culture. They combined pastoralism and cultivation with iron-working72. By 1075, the Leopard’s Kopje communities had grown into a kingdom based on Mapungubwe on the border of Botswana, Zimbabwe and South Africa; the area is now a South African National Park with a healthy wildlife population, including lions73. The Leopard’s Kopje people hunted as a supplement to livestock and arable farming. There is no evidence of the exclusion or widespread destruction of either herbivores or predators. In kingdoms like this and the Manekweni community of Bantu migrants in the coastal lowlands of southern Mozambique, a site occupied from the 12th to the 16th centuries, meat from hunting made up a relatively small part of food intake74. Much the same was the case for the larger and more intensively researched settlement at Great Zimbabwe75. In eastern Botswana, Zimbabwe and Mozambique, the Mapungubwe and Great Zimbabwe states came and went with no substantial ecological effect, according to the evidence available. The end of Great Zimbabwe led to the growth

Domestication, settlement and lions  37

in power of the Monomutapa kingdom further north. Both had mixed cultivation, livestock, supplementary hunting and trade in commodities like gold and ivory, but also skins, horn and hides. One area that was under Monomutapa control is what is now the Mufurudzi Safari Area and is believed by Pikirayi to have been a prime hunting area for the Mutapa76. At the time of Monomutapa rule, elephant and lion were there in large numbers. The size of the communities was such that while hunting might have been used to supplement food production, population levels were “beyond the point where any community could survive entirely upon hunting and gathering if its crops failed” 77. In the period leading up to the arrival of European traders, settlers and hunters, the southern African environment was savanna in eastern and southern ­A ngola, eastern Zambia and northern Namibia, often mixed with dry woodlands as in northern Botswana and Zimbabwe, and semi-desert in parts of ­Botswana, ­Namibia and South Africa78 – supporting a great diversity of ungulates preyed on by resident populations of lions and other predators. In South Africa, the southern Kalahari and the Karoo were arid but had sufficient rainfall to support blaubok, blesbok, bontebok, gemsbok, hartebeest, quagga, springbok and zebra, which fed healthy populations of lions. Human settlement was sparse, with minimal effect on wildlife numbers. The 16th and 17th centuries saw the establishment of Bantu settlements in northern and eastern South Africa that would become Xhosa, Nguni, Sotho and Tswana kingdoms.

Sudan, the Horn of Africa and Central Africa In southern Sudan, there was continuity of human habitation over the last ­m illennium BCE and into the first two millennium CE. Communities mixed cultivation and pastoralism or, in grasslands less suitable for crops, relied heavily on pastoralism79. In mixed systems, cattle grazed on seasonal floodplains for much of the year, then moved to fields used for cultivation after harvesting, to feed on stubble and manure the fields. The main crops were sorghum, cowpeas and groundnuts80. Among the pastoral peoples, the ancestors of the Dinka, large cattle camps would be formed on the floodplains, where they could be protected from lions and other large predators by herdsmen during the day and in bomas at night. Bone assemblages from settlement sites in the region show cattle, smallstock and a wide variety of wild faunal remains, including large and small ungulates. Cattle-rearing cultures continued to expand in the grassland and woodland-savanna area of South Sudan through the 15th–17th centuries among the Jyang (Dinka), Nuer and Ocholo (Shilluk) of southern Sudan, supplemented with hunting and fishing81. In all these communities there would have been a need to protect livestock and people from predators, but there is no oral, archaeological or other evidence detailing this or whether they developed into attempts to wipe out lion or other predator populations through pre-emptive extermination. Lions were present in large numbers when Arab and European traders, hunters and raiders travelled to the area.

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To the east, the 4th century BCE saw the rise of the Axumite Empire in northern Ethiopia and Eritrea. It became the dominant regional power, with strong regional trading links. Situated on a plateau, which has become increasingly arid over the centuries, it was a centre of human expansion with a strong agricultural and commercial economy. Hunting was the source of one of the main commodities traded, ivory. Axumite merchants traded as far north as A ­ lexandria in Egypt and via Mediterranean routes with Rome. They traded across the Red Sea to Yemen and the Arabian Peninsula and south to Somalia82. That lions were present in the area and respected by the Axumites is evidenced by the carved lion heads and lionesses in bas-relief on stones at Axumite settlements. Tefera believes that the archaeological finds at Axumite excavation sites “illustrate that lions were of symbolic importance in the life of the Axumite peoples”. There are also suggestions from finds that lions were hunted by the Axumites83. In western Ethiopia excavation of sites at Ajilak (in the Gambella region) indicate extensive hunting of wild fauna in the 1st millennium CE84. The region’s long-grass plains provided ideal habitat for a variety of ungulates. In 1973, sufficient wildlife remained there for the government to establish the Gambella N ­ ational Park. Lions are present in the park and surrounding region today, though in small numbers, as a result of habitat loss and hunting. Other wildlife includes buffalo, giraffe, waterbuck, roan antelope, zebra, warthog, hartebeest and elephant85. In the first two millennia CE wildlife was more abundant with antelope, giraffe and buffalo providing abundant prey for lions – there are no records from that period estimating the size of the lion population. Historically, lions were found throughout savanna, open woodland and other suitable habitats in Ethiopia, Eritrea, Somalia and Djibouti86. Lions were exterminated in the arid areas of Djibouti and Eritrea in the second half of the 19th century87. That there were substantial numbers with wide ranges in Ethiopia at least until 1900 is evident from the accounts available and the key roles that lions play in Ethiopian culture and imperial symbolism. Lion hunting was a widespread and highly regarded occupation, demonstrating bravery and enabling the collection of skins and other trophies. Lion skins were used in ceremonial headdresses and military apparel, and were an important form of tribute to the emperors. Early hunting methods included driving lions towards pits where they would be speared. In the 19th century, hunters in the Tigre region wore an earring in the right ear to indicate they had killed a lion88. Early hunting was probably not vastly damaging to overall lion numbers, that changed with the spread of firearms from the late 16th and early 17th centuries, when muskets were imported for the imperial army. The Zagwe dynasty ruled much of Ethiopia from 900 to 1270 CE, when the Solomonic dynasty (claiming descent from Solomon and the Queen of Sheba and their supposed son M ­ enelik89) succeeded. Lebna Dengel (1508–1540) raised an army said to have numbered 16,000 cavalry and 200,000 infantry to fight invaders90. The existence of well-organised, trained military forces with access to firearms denotes an enhanced capability to hunt lions. In Ethiopia, dating from the Axumite Empire, lions were symbols of b­ ravery – the term anbasa was used to refer to someone courageous who could be compared

Domestication, settlement and lions  39

to a lion. References to brave noblemen of this sort can be found in the land charters of emperors Zara Yagob (1399–1468) and Lebna Dengel. In the 16th century, coronation ceremonies involved the emperor striking at a lion – p­ resumably a captive one – and lions were kept by the royal family at Axum. Lebna Dengel was said to be preceded in procession by four lions when touring his domain91. In the 18th century, captive lions were set on bulls for the entertainment of nobles and other dignitaries92. Christian abbots and monks also had captive or tame lions associated with them – notably Gabra Manfus Qeddus, the Ethiopian saint and founder of the Zugualla monastery in Oromia. Often referred to as a 14th century monk, myths surrounding him say he lived for 562 years, and that he dwelt in the Egyptian desert for years with 60 lions and 60 leopards93. The Solomonic dynasty was Christian and adopted the biblical title, Lion from the Tribe of Judah. A royal seal of Menelik from the 19th century shows the crowned lion holding a cross94. The importance of the legitimacy and authority of the Lion of Judah title was developed in the reign of Menelik II, the modernising emperor at the end of the 19th century 95. Live lions were seen as embodiments of the power of the Ethiopian monarchs. Across the Sahel region from the Red Sea, through Sudan and into what is now Chad, the Central Africa Republic and Cameroon, there is evidence of the presence of lions dating back several thousand years in rock paintings in areas of the southern Sahara and the Sahel. They show a variety of antelopes, ostriches, buffalo, giraffe, elephants and rhino96. There are no published sources this author could trace on the size or ranges of lions across the region during this period or references to the nature of the relationship between settled or nomadic communities and lions. The probable range of lions in this region was limited by the Sahara to the north and dense forest south of Lake Chad and in the Congo Basin. Where there were pockets of grassland with herds of herbivores, the presence of lions is likely, as on the Bateke Plateau of Gabon on the border of the Republic of Congo. Lions were present there in reasonable numbers to the 1960s until, as Lee White head of Gabon’s national parks told me, the French poisoned most of them as part of a project to start livestock rearing on a large scale97. Lions were present in northern Cameroon and the savanna areas north of the Congo ­Basin in present-day Democratic Republic of Congo (DRC), where they survive but in rapidly dwindling numbers. An early Portuguese traveller in southern Congo and Angola reported in the late 16th century that lions were to be found in the Ambriz area of Angola on the coast north of Luanda. Filippo Pigafetta said that they were present near human settlements, adding the strange statement that “they do not molest white men, but only black ones, and even kill and eat black men whilst asleep, sparing white ones”98.

West Africa Grasslands, dry woodland and semi-arid savanna in West Africa provided the habitat and prey species necessary for lion survival. They were found across an

40  Domestication, settlement and lions

area covering northern Cameroon, northern Nigeria, the Lake Chad Basin, ­Niger, northern Benin, Burkina Faso, northern Ghana and Ivory Coast, Mali, small areas of Guinea and Guinea-Bissau, eastern Sierra Leone to Senegal and, before the spread of the desert, Mauritania. Small, scattered populations still survive in some of these countries99. They were absent from the dense forests inland from the Atlantic coast. In the period from around 4000 BCE to the Common Era, the grasslands of West Africa had a diversity of ungulates, which supported carnivores and human hunters100. The drying out of the region and increased desertification occurred around 2000 BCE101 and the effects of human expansion and land degradation combined with deforestation progressively reduced wildlife ranges and numbers up to the present102. Early Stone Age populations had limited ability to clear thick forest, limiting expansion of cultivated areas or open woodland suitable for livestock but also for lions and their prey. Better tools and techniques emerged around 500 BCE, as demonstrated by the Nok culture of Nigeria, according to Davidson103. This enabled an expansion of agriculture as well as improved weapons for hunting and protection against dangerous animals. Settled agriculture and more extensive production of yams, millet and rice developed in the river valleys and in the areas cleared of forest in the better watered areas south of the savanna. In areas with less reliable rainfall and poorer soils, shifting agriculture was practised alongside pastoralism104. Transhumant pastoralism was ubiquitous across the Sahel and savanna zone of West Africa. The population groups that coalesced to become the pastoralist Fulbe (called Peuhl in Senegal and Fulani in Nigeria) moved west and south as the Sahara dried out and grazing land was lost. They traversed the whole region of West Africa between the desert and the dense forest with their livestock105. As Arab, Berber and Tuareg peoples expanded their territories southwards and gained in technological and military capability, the Fulbe/ Peuhl/Fulani and other pastoral communities were pushed south106. At the beginning of the 2nd millennium CE, populations increased, settled communities grew and became kingdoms or large polities, like the states of Ghana, Kanem, Songhay and Takrur in the Sahel107. These states, exacting tribute from weaker surrounding communities, mixed cultivation in fertile valleys and floodplains with pastoralism and trade. In the better watered, more southerly regions extensive cultivation suggests the likely exclusion or diminution of large herbivore populations, with the consequent effect on predators. The depletion of wildlife and especially of predators would have been a long drawn out process in savanna and Sahel areas. The states of the region had well-developed iron-­ working industries, used horses or camels for herding and warfare, and were capable of mounting a far greater threat to wildlife and ability to kill predators. The Mali kingdom, established in the 13th century CE by the Mande leader, Sundiata Keita, stretched from the Atlantic Coast to present-day Mali. To emphasise his power, Sundiata had praise names such as Simbon Salata (Master Hunter) and Mari Diata (Lord Lion)108. Extensive crop cultivation and pastoralism in the Mali Empire pushed out wildlife, which survived only in regions

Domestication, settlement and lions  41

unsuitable for cultivation and marginal for pastoralism109. Hunting was not recorded as being of importance, though elephants were killed to feed demand for ivory. The period from the 12th up to the 19th centuries was characterised in West and Sahelian Africa by the rise and fall of empires based on the power of horse-borne armies and the economic production of slave-reliant systems110, systems in which wildlife did not figure in available records as a source of food or worthy of preservation. The ill-fated explorer Mungo Park in his accounts of travels along the Gambia River and in Mandingo districts makes mention of reports by local people of hyena, leopard, elephant and antelope species hunted for meat, but not of lions being present during his travels in 1795, suggesting absence or rarity111. The sole reference from that area is to reports from “many years ago” of a hunt for a lion in eastern Gambia that was said to have been killing cattle. Attempts to kill it failed and ended up with the lion killing and “devouring” two hunters112. Park’s only recorded encounter with a lion was on the Niger River in Mali between Bamako and Mopti. He saw the lion when riding along the river bank, but the lion made no attempt to attack. Park mentions that in this area of Mali, the Fulani keep their cattle on fertile islands in the river to protect them from predators113.

North Africa Lions were present across the coastal plains and mountains of North Africa throughout the period under review in this chapter, though they were gradually thinned out by the combined effects of climate change, human settlement, agricultural expansion and hunting to kill them or to capture lions for use in the Roman arenas. The states and dynasties that developed here were based on cultivators and pastoralists, who had been domesticating animals since around 7000 BCE along the Nile114. The productivity of agriculture enabled the expansion of populations, growth of settlements and formation of larger, hierarchical societies with sophisticated economic, trade and government systems. It was from these developing economic systems and civilisations that farming and livestock cultures spread across North Africa, along the Nile and into areas inland such as the Hoggar Mountains of Algeria and Lake Chad Basin115. The presence of diverse wildlife is evidenced by the images of lions, elephants, buffalo, giraffe, zebra and other large herbivores painted or carved on rock by Neolithic peoples in areas north of the Sahara in the 5000–3000 BCE period116. At this time North Africa had a wetter climate and lusher vegetation supporting a rich fauna of herbivores including elephant, giraffe, buffalo and zebra, which supported populations of predators117. Rock engravings including lions were found in the mountainous areas at Djanet, and Akatous in Algeria, Messak Settafet in Libya, Atlas in Morocco and Tibesti in Chad. Lions were also painted on rocks at Djebel Silsila and North Kom Ombo, east of the Nile. These paintings date from 4400 BCE to 3125 BCE118. Excavations at Beni Salama in Lower Egypt from the Merimde culture (4800–4300 BCE) revealed the burned

42  Domestication, settlement and lions

bones of lions, though with no indication of how and why they were killed and came to be burned – whether for consumption or rituals119. Remains of lions were found in a royal tomb at Um el-Qaab north-west of Thebes (Upper Egypt about 675 km south of Cairo, and the capital of Egypt under the 18th dynasty from 1550 BCE to 1290 BCE) and the entire skeleton of a lion was found in the tomb of Tutankhamun’s nurse (1430 BCE)120. The Great Sphinx has a human head and a lion’s body. It is located at Giza and appears to guard the pyramid of the Pharaoh Khafra. The date of its construction is uncertain, though Khafra ruled for 26 years around 2570 BCE. At the site, a 1400 BCE inscription on a stele of the 18th dynasty names three manifestations of the sun deity of that period, Khepera–Rê–Atum. The inclusion of these figures in tomb complexes was a tradition and many pharaohs ordered that carvings of their heads be put on the guardian statues of their tombs to show their close relationship with the powerful solar deity, Sekhmet, a lioness, sometimes depicted as a woman with a lioness’s head. In the New Kingdom (16th–11th century BCE) lions were a major part of symbolism. Bast (the cat goddess) was depicted as a lioness. The Egyptians believed that a sacred lioness was responsible for the annual flooding of the Nile, which created the fertile floodplains on which the kingdom’s abundance of food and great wealth was based121. Texts dating from the reign of Amenhoteb II around 1400 BCE refer to him hunting lions in Egypt122. Amenophis III (1352–1330 BCE) recorded that he had killed 100 lions in hunts. Schnitzler’s research suggests that lions were killed on royal hunts in Nubia (southern Egypt and northern Sudan). Lions appear to have been “relatively common” in Egypt from 1500 to 1300 BCE, being found along the coast of the western desert, in the eastern desert and Sinai123. Along the Mediterranean coast, the rise of the Carthaginian Empire on the foundations of the Phoenician settlement there, dating from 814 BCE, introduced a powerful state with large towns and more developed agriculture, not to mention a strong army124. Prior to the Punic Wars with Rome, starting at the end of the 3rd century BCE and ending with the third phase of war in 146 BCE, Rome traded with Carthage. Live animals (including lions) and wildlife products such as skins and ivory were part of the commerce. Carthage sent trading caravans into the interior of Africa exchanging its manufactured and agricultural goods for salt, gold, timber, ivory, live animals, skins and hides125. By the 5th century BCE, the desert traffic – in animals such as monkeys, lions, leopards and elephants, ivory, precious stones and slaves – had become so important that the Carthaginians began to conduct major expeditions across the Sahara to control the commerce in the live animals they sought126. This trade network was disrupted by the defeat of Carthage in the Third Punic War, when Rome was victorious and made the region a province of Rome. Romans would then have come into direct contact with wild lions in the parts of their growing empire from Mauritania through Numidia, Cyrenaica and into Egypt. Rome now controlled the Mediterranean end of trans-Saharan trade and continued the substantial commerce in dangerous animals bound for the arenas of the empire127.

Domestication, settlement and lions  43

The Romans gained control of Egypt in 30 BCE, taking advantage of the revolts against the Ptolemaic rulers (who had established a dynasty when one of Alexander the Great’s officers declared himself pharaoh in 305 BCE). Roman officials like Manlius described Egypt as a barbarian country inhabited by “big elephants and furious lions”. Texts from Herodotus (c.484–425 BCE) and Saint Augustine of Hippo (354–420 CE) suggest that over hundreds of years, bridging the BCE and CE eras, lions were common in Libya and, to a lesser extent, in Egypt128. The populations of lions and other large animals, including elephants, provided the Romans with animals that gladiators fought or hunters massacred in the circuses for the sensation-hungry Romans. Schnitzler says the capture and export of lions and other wild animals by the Romans continued for several centuries129. The wildlife population of North Africa at this time was abundant, with evidence of wildebeest, hartebeest, barbary sheep, lions and leopards130. The Roman writer Pliny the Elder wrote that the first recorded lion contests in the arena were staged under Lucius around 80 BCE when, he says, 100 maned lions took part in fights. He adds that under Pompey, games were held involving 600 lions and under Julius Caesar there was one with 400 lions131. Pliny developed a respect for lions, which he described as courageous and noble, and was one of those who helped establish the myth that lions would not attack the weak, who, he said, were seen by lions as “prey unworthy of their dignity”132. This merciful aspect, to be oft repeated in coming centuries, was illustrated by Pliny with the story of Androcles, a Syracusan who took a thorn from the paw of a lion, which later came face to face with him in the Roman arena, where Androcles had been thrown to the lions. The lion, according to the fable, recognised him and refused to attack – leading to the release of both Androcles and the lion133. The Roman games frequently involved staged fights between gladiators and lions or massive animal hunts, known as venationes, involving a variety of dangerous animals (including lions, elephants, rhinos, leopards, bears and wolves), which were released into the arena to be hunted by archers and spearmen. There were also the occasions when criminals and, later, Christians, were herded into the arena or tied to stakes to be killed by lions, bears and other predators134. The last animal hunts in the arenas in Rome took place in 523 CE but they continued for several more decades in the Eastern Roman Empire135. Beard suggests that the massive animal hunts and the slaughter of lions from across the empire were emblematic of the power of Rome over its subject peoples136. The widespread capture of lions stopped with the disappearance of the R ­ oman Empire, but hunting of them to protect livestock or for sport continued. Schaller notes that they were still present in Tunisia around 1249–54 CE, when King Louis of France hunted lions with a crossbow during the crusades137. The North African or Barbary lion survived across the region for several centuries. They were abundant in the lightly wooded plains along Morocco’s Atlantic coast until the 16th century138 and were present in Libya until the 18th century. In some areas they survived until the end of the 19th century and possibly a few decades beyond, notably in Morocco’s Atlas and Rif mountains, though records are sparse139.

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Europe, West Asia and South Asia After the extinction of the cave lion (P spelaea) in southern-western Europe, about 14,000–14,500 years ago and their replacement by modern lions (P leo) approximately 8,000 years ago, lion populations in Europe gradually declined and disappeared because of human expansion and the disappearance of their prey, the latter resulting from human hunting, agricultural expansion and changes in climate. Open grassland and light woodland was replaced by thicker forests as the climate became colder and wetter. Records of modern lions from the Iberian peninsula show they were present there from 8000–7000 BCE140. They were also present from that time across southern Europe and as far north as Hungary and Ukraine. They were found around the Black Sea until about 3000 BCE. In the Balkans, Asia Minor and the Middle East, the lion survived until the early years of the Common Era141. P leo went into decline during the Iron Age (c.1100 BCE–400 CE) in Europe, leading to its disappearance from south-eastern Europe. The most likely factors in the extinction of European lion populations are the expansion of human settlement and agriculture, stock protection and hunting142. Lions survived in Greece until the 5th century BCE, and Herodotus reported that there were lions in Thrace in his lifetime (484–430 BCE). He wrote that when the Persian king Xerxes marched his army through Macedonia and Thrace, “night after night lions used to come down from their usual hunting grounds and attack the camels in the baggage train – they didn’t attack anything but the camels … there were huge numbers of lions in these parts”, which normally preyed on wild cattle, wild boar and deer143. In the next century, Aristotle wrote that lions were still to be found in Thrace and other parts of Greece but were becoming scarce. Human hunting and loss of habitat led to the extermination of the European lion in most areas by the first two centuries of the CE period, with total disappearance by the end of the 2nd century144. No skeletal remains of Greek lions have been found, which makes it difficult to date or explain their extinction. Lions were an important part of the stories, legends and symbolism of ancient Greece, representing power, nobility and wealth145. To slay a lion was seen as an act of the greatest bravery and strength. In Greek myths, Hercules had as his first labour (assigned by the gods for killing his wife and children) the killing of the Nemean Lion, which by legend had fur made of gold and claws sharper than swords. Hercules strangled it and thereafter wore its skin146. Alexander the Great, was often depicted wearing a lion’s head and a lion skin. There are artistic representations of Alexander hunting lions, a symbol of his courage and power147. The theme of the lion attack on prey “occurs prominently in architectural sculpture” at Delphi and other parts of mainland Greece. Usually, one or two lions are depicted attacking a bull, calf, goat, boar or deer; “the common denominator, however, is the vanquishing lion itself, who is clearly the aggressor and, in nearly every instance, the obvious victor in the struggle”148. Evidence suggests that lions survived in Macedonia and surrounding areas at this time and

Domestication, settlement and lions  45

that lion hunting was a popular sport among the rulers149. It is thought that the last Greek lions died out between 80 and 100 CE150. In 6th century BCE, Lydia and Babylon lions were symbols of authority, with official weights for trade made in the form of lions151. The Assyrian kings, who preceded Alexander, were avid hunters of lions between the 9th and 7th centuries BCE in Iraq and Syria and right up to the border of Turkey (where lions may have survived until the end of the 19th century). One king, who ruled from 668 to 627 BCE, declared: “I am Assurbanipal, the King of the World, the King of Assyria! For my regal amusement I have caught the Desert King by his tail, and on the instructions of my helpers, the Gods Ninib and Nergal, I have split his head with the two-handed sword”152. The lion-hunting exploits of the Assyrian rulers were recorded on friezes and stelae. Assurbanipal’s chroniclers recorded that lions were plentiful in forest areas and in the reed thickets along major rivers. The king justified his killing of lions saying that they had become too bold in the killing of cattle, smallstock and men: “They constantly kill the livestock of the fields, and they spill the blood of men and cattle. The herdsmen and the supervisors are weeping; the families are in mourning. The misdeeds of these lions have been reported to me … I have penetrated their hiding places and destroyed their lairs”153. The ­Assyrians hunted from chariots with a charioteer, and spearmen to guard the king’s back, accompanied by “a whole host of horsemen, foot soldiers and kennel men who, at the right moment, unleashed the mastiff-like hunting dogs … a lion, after having been found by the trackers, was driven into the open and surrounded”. The king would fire his arrows and the spearmen would finish the lion off 154. The rulers of the Persian Empire hunted lions, using beaters to drive lions towards the hunters. Lions were common throughout the plains and woodlands of Persia before the Common Era, but gradually declined over the next 1,800 years, surviving in some places until the mid-19th century. The last lion was said to have been shot in 1923, but a dead lion was reported to have been found in Iran as late as 1943155. Lions survived in Palestine until around the 12th century and into the 20th century in parts of Syria, Iraq and Iran156. Lions are believed to have been common in Azerbaijan and possibly elsewhere in the foothills and coastal plains of the Caucasus until the 10th century CE157. Further east, according to Divyabhanusinh, they survived in Afghanistan and Baluchistan until the end of the 19th century158. Lions are mentioned more than 150 times in the Bible, and in early Jewish literature, demonstrating a familiarity with them and use of lions as symbols of power, nobility but also anger159. Among the best known biblical references are Samson’s killing of a lion with his bare hands when it roared at him and seemed about to attack. There are references to David slaying a lion and a bear which had killed a sheep in his father’s flock (Samuel 1 17:4) and to lions killing people and domestic animals (Kings 1 13:24–8). They are generally presented as aggressive, threatening and wrathful – the anger of kings is compared with that of lions (Proverbs 19:12). They are also symbols of justice. When the wrongly accused Daniel is thrown to the lions, they do not touch him but they kill his false accusers and their families (Daniel 6:6–24).

46  Domestication, settlement and lions

India, now home to the last surviving wild lion population outside Africa, had a widespread population in northern, western and some central areas, where there were open habitats or lightly forested areas supporting sufficient prey. What is now identified as P leo persica originated as the populations of lions of north, central and western Africa and is believed to have entered the Indian subcontinent via West Asia, probably through Persia and Baluchistan or down the Indus Valley sometime between the latter part of the 3rd millennium and the end of the 2nd millennium BCE, moving across northern and western India between 1500 and 600 BCE160. This included colonisation of the Kathiawar Peninsula of Gujarat (here the Gir Forest, the last refuge of the Asiatic lion, is found), which is believed to have occurred 2,680 years ago. Soon after, the peninsula was temporarily separated from mainland India by rising sea levels, leading to a genetic bottleneck for the lions there lasting several generations161. The arrival of lions in these areas coincided with the rise of the Indus Valley civilisations in India around 3300–1300 BCE, equivalent to the Bronze Age cultures of Egypt and Mesopotamia. More advanced agricultural and societal forms spread with the peoples of this culture and reached Gujarat from the north162. The human migrants came from Aryan communities from the north and west. This was true of many of the dynasties that developed in northern and western India, such as the Yadavas, the Mauryan, the Kshatrap dynasty, the Gupta ­Empire and then the Pratiharas (which succeeded one another and ruled parts of the region from the 2nd millennium BCE through to the 9th century CE)163. They had experience of coexistence and conflict with lions, which were present in their original homelands and in territories through which they had migrated. They had brought religious beliefs with them which included imagery of lions – the lion-mounted Hindu Goddess Durga, for example, is said by Mitra to be derived from the goddess Ishtar worshipped by the Sumerians, Assyrians and Babylonians164. Ishtar was often depicted as riding a lion. Lions are frequently mentioned in the religious Vedic texts, such as the Rig Veda, in which the “great roar of the lions in the hills” and their strength are lauded. There is controversy over the exact age of the text, with some Hindu yogis and scholars putting it before 4000 BCE and Western scholars nearer to 1500 BCE165. Its mention of lions, along with scientific evidence, supports the idea that the modern race of Indian lions was present at least from the 2nd or 3rd millennia BCE. Divyabhanusinh notes that descriptions of lions according to the colour of their manes are to be found at the time of the Mahabharata, which was written or compiled between the 4th century BCE and the 4th century CE166. The wide range of lions across northern and western India, the large numbers known to have been present (testified to by the huge slaughter of lions in the late 18th and first 60 years of the 19th century by Indian rulers, visiting British royalty and administrators/officials of the British Raj) points to the migration of lions from west and north-west Asia into the areas of India with suitable prey and habitat. There are, though, long gaps in historical, literary or other accounts of lions between their arrival and the establishment of the Maurya dynasty in Gujarat,

Domestication, settlement and lions  47

when Chandragupta Maurya conquered Saurashtra in around 323 BCE167. The Maurya rulers invaded and controlled land that had been ruled or tributary to Alexander the Great’s Empire in western Asia. Lions in India now have an historical association with the Maurya rulers, who ruled Saurashtra, including the Gir Forest, and were avid lion hunters, organising huge royal hunts and establishing hunting grounds reserved for royalty. They allowed farmers and hunters to kill wildlife that invaded farmland or killed stock168. Hunting had up until this period been mainly a form of supplying meat, now it mixed sport hunting, hunting for meat, crop and livestock protection. This may have had the effect of limiting safe habitats available to wildlife and reducing the range and numbers of lions, but there is no suggestion of the extermination of wildlife around inhabited areas. The empires that succeeded the Mauryan and ruled over much of this area appear to have followed similar attitudes of respect, veneration and laudatory depiction of lions while also hunting them and representing royal hunts as part of the majesty and power of rulers. Saurashtra was ruled over by the successors to the Mauryans (the Guptas and Pratiharas – who ruled up until the 9th century CE). They were succeeded by the Chavada dynasty and the Solankis until the powerful Mughal dynasty, with its origins in the Muslim polities of Central Asia, became the dominant power in this part of India from 1526 until its disintegration under the weight of pressure from the Marathas and British colonial expansion in 1758169. The wide range of lions in northern, western and parts of east-central India, their importance in religious and artistic symbolism indicate a lasting acquaintance of many Indian peoples, empires and religions with the subcontinent’s lions and their long-­ standing symbolism as majestic, powerful and courageous creatures. At the time of the Mughals, lions were widely distributed across the present-­ day states of Rajasthan, Gujarat, Punjab, Haryana, Uttar Pradesh, Madhya Pradesh and Bihar, and many reports of their presence can be found in the literature170. The later Mughal rulers respected but also avidly hunted lions171. The Mughal emperor Jahangir was said to have hunted lions from horseback and killed them with spears or arrows172. Once firearms became available, the emperors are recorded as having hunted from horseback with guns. Under the Mughal emperor Aurangzeb, lions were killed in huge military-style hunts. The lions were driven from cover by huge numbers of beaters towards the royal ­hunters – Aurangzeb was said to be a particularly keen and regular lion hunter173. When Mughal power declined and British occupation of much of India began in the 18th and early 19th centuries, the new rulers took up lion hunting in similar style and with similar enthusiasm, leading to the extermination of lions in much of the subcontinent174.

Notes 1 Robin S. Reid (2012), Savannas of Our Birth. People, Wildlife and Change in East ­Africa, Berkeley: University of California Press, 2012, p. 95. 2 Christopher Ehret (2016), The Civilizations of Africa. A History to 1800, Charlottesville: University of Virginia Press, p. 37. 3 Ehret, 2016, pp. 41–2.

48  Domestication, settlement and lions

4 John Iliffe (2007), Africans. The History of a Continent, Cambridge: Cambridge University Press, 2nd edition, pp. 12–13. 5 For a more detailed examination of the origins of domesticated cattle, see Jared E. Decker et al. (2014), Worldwide patterns of ancestry, divergence, and admixture in domesticated cattle, PLOS Genetics, 27 March. 6 Brian Bertram (1978), Pride of Lions, London: J.M. Dent, p. 240. 7 Ehret, 2016, pp. 25–33. 8 Ibid., p. 13. 9 Christopher Ehret (1998), An African Classical Age. Eastern and Southern Africa in World History 1000 BC to AD 400, Oxford: James Currey, p. 6. 10 Reid, 2012, p. 96. 11 Iliffe, 2007, p. 17. 12 Reid, 2012, p. 96. 13 Juliet Clutton-Brock (1994), The legacy of Iron Age dogs and livestock in Southern Africa, Azania: Archaeological Research in Africa, 29/30, 1: 161–167, p. 164. 14 Frank W. Marlowe (2010), The Hadza Hunter-Gatherers of Tanzania, Berkeley: University of California Press, p. 18. 15 Ehret, 2016, pp. 73 and 75. 16 Ibid., p. 27 and R. Woodroffe and L.G. Frank (2005), Lethal control of African lions (Panthera leo): local and regional population impacts, Animal Conservation, 8: 91–98. 17 Iliffe, 2007, pp. 1–2. 18 Joseph Ki-Zerbo and Djibril Tamsir Niane (1997), Africa from the Twelfth to the Sixteenth Century. General History of Africa IV, London: James Currey/UNESCO, p. 199. 19 Reid, 2012, p. 101. 20 Juliet Clutton-Brook (1993), The spread of domestic animals in Africa, in Thurstan Shaw et al., The Archaeology of Africa. Food, Metals and Towns, London: Routledge, pp. 61–70, p. 66–7. 21 Ibid. 22 J.R.F. Bower (1973), Seronera: excavations at a stone bowl site in the Serengeti ­National Park, Azania, 8: 71–101. 23 T.M. Cato (1994), Cheetahs of the Serengeti Plains. Group Living in an Asocial Species, Chicago: University of Chicago Press, p. 16. 24 Bertram, 1978, p. 14. 25 Paul Lane (2004), The ‘moving frontier’ and the transition to food production in Kenya, Azania: Archaeological Research in Africa, 39, 1: 243–264, p. 248. 26 Ibid., p. 252. 27 Kennedy K. Mutundu (2010), An ethnoarchaeological framework for the identification and distinction of Late Holocene archaeological sites in East Africa, Azania: Archaeological Research in Africa, 45, 1: 6–23, p. 7. 28 Ibid. 29 Peter Robertshaw (1993), The beginnings of food production in southwestern Kenya, in Thurstan Shaw et al., 358–371, p. 365. 30 Oliver J.C. Boles and Paul J. Lane (2016), The green, green grass of home: an archaeo-­ ecological approach to pastoralist settlement in central Kenya, Azania: Archaeological Research in Africa, 51, 4: 507–530, p. 509. 31 See Ehret, 2016, p. 105, on the drying of the climate and its effects. 32 Ibid. 33 Philip Curtin et al. (1995), African History from Earliest Times to Independence, London: Longman, 2nd edition, p. 104. 34 Diane P. Gifford et al. (1980), Evidence for predation and pastoralism at Prolonged Drift: a Pastoral Neolithic site in Kenya, Azania: Archaeological Research in Africa, 15, 1: 57–108, pp. 64–5. 35 Curtin et al., 1995, p. 104. 36 David Wright (2003), Archaeological investigations of three Pastoral Neolithic sites in Tsavo National Park, Kenya, Azania: Archaeological Research in Africa, 38, 1: 183– 188, p. 187.

Domestication, settlement and lions  49

37 Ibid., pp. 107–9. 38 Ehret, 1998, p. 31. 39 Ibid., p. 109. 40 Ehret, 2016, p. 161. 41 Ehret, 2016, p. 269. 42 Ibid., p. 270. 43 Dorothy L. Hodgson (2001), Once Intrepid Warriors: Gender, Ethnicity, and the Cultural Politics of Maasai Development, Bloomington: Indiana University Press, pp. 23–4. 44 Edward I. Steinhart (2006), Black Poachers White Hunters. A Social History of Hunting in Colonial Africa, Oxford: James Currey, p. 21. 45 Anonymous (1998), Hyrax Hill and the later archaeology of the Central Rift Valley of Kenya, Azania: Archaeological Research in Africa, 33, 1: 73–112, DOI: 10.1080/00672709809511465, accessed 15 June 2017, pp. 81–2. 46 Naomi Kipury (1983), Oral Literature of the Maasai, Nairobi: East African Educational Publishers, p. 2. 47 Ibid. 48 Ibid. 49 Ibid., p. 4. 50 Ibid., p. 24. 51 Curtin et al., 1995, p. 134. 52 Ibid. 53 Ki-Zerbo and Niane, 1997, pp. 200–201. 54 Ehret, 2016, p. 272. 55 I. Hrbek (1992), Africa from the Seventh to the Eleventh Century, London: James Currey/UNESCO, p. 293. 56 Jan Bender Shetler (2007), Imagining Serengeti. A History of Landscape Memory in Tanzania from Earliest Times to the Present, Athens: Ohio University Press, p. 3. 57 Ibid., p. 33; and, A.R.E. Sinclair and P. Arcese (1995), Serengeti II: Dynamics, Management and Conservation of an Ecosystem, Chicago: University of Chicago Press, p. 3. 58 D.N. Beach (1980), The Shona and Zimbabwe 900–1850, Gweru, Zimbabwe: Mambo Press, p. 53. 59 See Benedict Anderson (1983), Imagined Communities, London: Verso; and, Eric Hobsbawm and Terence Ranger (Eds) (1992), The Invention of Tradition, Cambridge: Cambridge University Press. 60 Beach, 1980, p. 53. 61 Alan Barnard (1992), Hunters and Herders of Southern Africa. A Comparative Ethnography of the Khoisan Peoples, Cambridge: Cambridge University Press, pp. 30–2. 62 Ibid., p. 54. 63 Ehret, 2016, p. 165. 64 Ibid., pp. 170–2. 65 W.H.I. Bleek and L.C. Lloyd (1911), Bushman Folklore, republished 2007, London: Forgotten Books, Kindle edition, loc 349. 66 Thembi Russell and Faye Lander (2015), ‘What is consumed is wasted’: from foraging to herding in the southern African Later Stone Age, Azania: Archaeological Research in Africa, 50, 3: 267–317, p. 268. 67 Ibid., p. 271. 68 Ki-Zerbo and Niane, 1997, p. 209. 69 Iliffe, 2007, pp. 100–101. 70 Ibid., p. 101. 71 John Kinahan (1994), A new archaeological perspective on Nomadic Pastoralist expansion in south-western Africa, Azania: Archaeological Research in Africa, 29/30, 1: 211–226, p. 211. 72 Hrbek, 1992, p. 318. 73 Personal communication with Kamogelo Pshatlele Mabowa, environmental monitor at Mapungubwe National Park.

50  Domestication, settlement and lions

74 Graeme Barker (1978), Economic models for the Manekweni Zimbabwe, ­Mozambique, Azania: Archaeological Research in Africa, 13, 1: 71–100, p. 71. 75 Ibid., p. 86. 76 Innocent Pikirayi (1991), The Archaeological Identity of the Mutapa State. Towards an Historical Archaeology of Northern Zimbabwe, Uppsala: Societas Archaeologica Upsaliensis, p. 30. 77 Beach, 1980, pp. 26–8. 78 Curtin et al., 1995, p. 214. 79 Peter Robertshaw and Ari Siiriäinen (1985), Excavations in Lakes Province, Southern Sudan, Azania: Archaeological Research in Africa, 20, 1: 89–161, p. 152. 80 Ibid., p. 91. 81 Ehret, 2016, p. 375. 82 M. Tefera (2004), Recent evidence of animal exploitation in the Axumite Epoch, 1st–5th centuries AD, Tropical Animal Health and Production, 36: 105–116. 83 Ibid., pp. 108 and 113. 84 Alfredo González-Ruibal et al. (2014), Late hunters of western Ethiopia: the sites of Ajilak (Gambela), c. AD 1000–1200, Azania: Archaeological Research in Africa, 49, 1: 64–101, p. 95. 85 Selamta (2018), Gambella National Park, www.selamta.net/gambela_national_ park.htm, accessed 31 January 2018. 86 Siegbert Uhlig (Ed.) (2007), Encyclopaedia Aethiopica, Volume 3, Wiesbaden: Harrassowitz Verlag, p. 571. 87 Ibid. 88 Ibid. 89 Richard Pankhurst (Ed.) (1967), The Ethiopian Royal Chronicles, Addis Ababa: Oxford University Press, p. xi. 90 Ibid., p. 49. 91 Ibid. 92 Uhlig, 2007, p. 572. 93 Haile Getachew (no date), Ethiopian Saints, The Coptic Encyclopaedia, Claremont Colleges Digital Library, http://ccdl.libraries.claremont.edu/cdm/ref/collection/ cce/id/840, accessed 1 February 2018; see also, G.W.B. Huntingford (1989), The Historical Geography of Ethiopia, London: British Academy, p. 85. 94 Ibid. 95 Ibid. 96 C.A. Guggisberg (1961), Simba. The Life of the Lion, London: Bailey Bros and ­Swinfen, p. 29. 97 The author is indebted to Lee White, head of Gabon’s national parks department for this information in an e-mail of 21 January 2018. 98 Duarte Lopes and Filippo Pigafetta (translated by Maragerite Hutchinson) (1881 reprinted), A Report of the Kingdom of Congo: And of the Surrounding Countries, London: John Murray (reprinted by Leopold Classic Library), p. 49. 99 See H. Bauer et al. (2001), Status and Needs for Conservation of Lions in West Africa and Central Africa. An Information Exchange Workshop, Limbe, Cameroon, 2–4 June, ­Apple Valley Minnesota: Conservation Breeding Specialist group (IUCN/SSC), and, P. Henschel et al. (2014), The lion in West Africa is critically endangered, PLoS ONE, 9, 1. 100 Oliver Davies (1967), West Africa Before the Europeans. Archaeology and Prehistory, London: Methuen, pp. 148–9. 101 Basil Davidson (1998), West Africa Before the Colonial Era. A History to 1850, London: Longman, p. 10. 102 Keith Somerville (2017), Africa’s Long Road Since Independence. The Many Histories of a Continent, London: Penguin, pp. 167–70. 103 Davidson, 1998, pp. 12–13. 104 Davies, 1967, p. 149. 105 Curtin et al., 1995, pp. 84–5.

Domestication, settlement and lions  51

106 Hrbek, 1992, p. 67. 107 Ibid., p. 69. 108 Ki-Zerbo and Niane, 1997, p. 56. 109 Ibid., p. 65. 110 Iliffe, 2007, pp. 72–3. 111 Mungo Park (2002), Travels in the Interior of Africa, Ware, Herts: Wordsworth Classics of World Literature, 1795, p. 10. 112 Ibid., p. 26. 113 Ibid., pp. 191–2. 114 Lawrence Barham and Peter Mitchell (2008), The First Africans. African Archaeology from the Earliest Toolmakers to Most Recent Foragers, Cambridge: Cambridge University Press, Kindle edition, p. 344. 115 Ehret, 2016, pp. 79 and 86. 116 Annik E. Schnitzler (2011), Past and present distribution of the North African– Asian lion subgroup: a review, Mammal Review, 41, 3: 220–243, p. 224. 117 Ibid. 118 Ibid., p. 225. 119 Ibid. 120 Ibid., p. 225. 121 See for a fuller account of Egyptian symbolism involving lions and other animals, Garai, Jana (1974), The Book of Symbols, New York: Simon & Schuster. 122 Andrew J. Loveridge, Craig Packer and Adam Dutton (2008), Science and the recreational hunting of lions, in Barney Dickson et al. (Eds), Recreational Hunting, Conservation and Rural Livelihoods, Oxford: Wiley-Blackwell, 108–123, p. 109. 123 Schnitzler, 2011, p. 198. 124 Robin Derricourt (2011), Inventing Africa. History, Archaeology and Ideas, London: Pluto, p. 11. 125 Peter I. Bogucki (2008), Encyclopedia of Society and Culture in the Ancient World, New York: Facts On File, p. 389. 126 Adu Boahen (1962), The caravan trade in the nineteenth century, Journal of African History, 2: 349–359, p. 349. 127 Ibid. 128 Herodotus (1998), The Histories (trans. Robin Waterfield), Oxford: Oxford University Press, pp. 107 and 299. 129 Schnitzler, 2011, p. 198. 130 Susan Raven (1969), Rome in Africa, London: Evans Brothers, p. 3. 131 Pliny the Elder (2004), Natural History. A Selection, London: Penguin, revised edition, p. 115. 132 Ibid. 133 Ibid., p. 116. 134 Ingvild Sælid Gilhus (2006), Animals, Gods and Humans: Changing Attitudes to Animals in Greek, Roman and Early Christian Ideas, New York: Routledge, p. 12. 135 Ibid., p. 36. 136 Mary Beard (2015), SPQR. A History of Ancient Rome, London: Profile, p. 504. 137 George B. Schaller (1972), The Serengeti Lion. A Study of Predator-Prey Relations, Chicago: University of Chicago Press, p. 5. 138 Ibid., pp. 231–7. 139 S.A. Black, A. Fellous, N. Yamaguchi and D.L. Roberts (2013), Examining the extinction of the Barbary Lion and its implications for felid conservation, PLoS ONE 8, 4: 2–3. 140 Ibid., p. 224. 141 N.K. Vereshchagin and G.F. Baryshnikov (1984), Quaternary mammalian extinctions in northern Eurasia, in Paul S. Martin and Richard G. Klein (Eds), Quaternary Extinctions. A Prehistoric Revolution, I, Tucson: University of Arizona Press, 483–516, p. 497. 142 R.S. Sommer and N. Benecke (2006), Late Pleistocene and Holocene development of the felid fauna (Felidae) of Europe: a review, Journal of Zoology, 269: 7–19, p. 815.

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143 Herodotus, 1998, p. 446. 144 Marco Masseti and Paul A. Mazza (2013), Western European Quaternary lions: new working hypotheses, Biological Journal of the Linnean Society, 109: 66–77, p. 67. 145 http://platos-academy.com/lions-in-ancient-greece/ accessed 24 March 2017. 146 Robert Graves (1955), Greek Myths, London: Penguin revised edition 1992, pp. 465–469. 147 http://platos-academy.com/lions-in-ancient-greece/ 148 Glenn Markoe (1989), The “lion attack” in Archaic Greek art: heroic triumph, Classical Antiquity, 8, 1: 86–115, p. 86. 149 J.K. Anderson (1985), Hunting in the Ancient World, Berkeley and Los Angeles: ­University of California Press, pp. 80–1. 150 Ibid., and Maureen Alden (2005), Lions in paradise: lion similes in the Iliad and the lion cubs of IL. 18.318-22, The Classical Quarterly, New Series, 55, 2: 335–342, p. 335. 151 Ibid., pp. 103 and 110. 152 Guggisberg, 1961, p. 158. 153 Ibid., and Anderson, 1985, p. 6. 154 Guggisberg, 1961, p. 158. 155 Divyabhanusinh (2008), The Story of Asia’s Lions, Mumbai: Marg Publishers, p. 45; see also Guggisberg, 1961, p. 42. 156 C.D. Lynch (1988), Lion: a historical account, Culna, Issue 35, October, pp. 20–21. 157 Asiatic Lion, http://web.archive.org/web/20090206173029/http://www.asiatic-­ lion.org/distrib.html, accessed 24 March 2017; see V.G. Heptner and A.A. Sludskii (1972), Mammals of the Soviet Union, Vol. 2, Brill: Brill Publishers. 158 Divyabhanusinh, 2008, p. 49. 159 Gideon R. Kotze (2015), Lion imagery in 1 Maccabees 3:4: Septuagint conference articles (LXXSA 2014), Journal for Semitics, 24, 1, January: 326–351, pp. 326–7. 160 Sandeep Kumar and R.L. Meena (2014), Foresters Diary. Asiatic Lion Landscape, ­Gujarat Forest Department, p. 3. 161 Kausik Banerjee and Yadvendradev V. Jhala (2012), Demographic parameters of endangered Asiatic lions (Panthera leo persica) in Gir Forests, India, Journal of Mammalogy, 93,. 6, December: 1420–1430, p. 1420. 162 Sudipta Mitra (2005), Gir Forest and the Saga of the Asiatic Lion, New Delhi: Indus Publishing, p. 43. 163 Ibid. 164 Ibid. 165 Ibid. 166 Ibid., p. 70. 167 Mitra, 2005, p. 43. 168 Ibid. 169 See John F. Richards (2007), The Mughal Empire, Cambridge: Foundation/Cambridge University Press; and Mitra, 2005, p. 14. 170 Ibid. 171 Ibid., p. 44. 172 Guggisberg, 1961, p. 171. 173 Ibid. 174 Ibid.

3 Lions from the 14th century to colonial occupation

By the 15th century CE, lions had disappeared from Europe but were still found in West Asia, northern and western India and North Africa. Sub-Saharan A ­ frica was clearly the key region for lion survival, with populations spread across savanna, open woodland and semi-desert regions; only absent from the central Sahara and rainforest in West Africa and the Congo Basin. Africa will be the focus of this chapter, dealing with the consequences of the arrival of Europeans, firearms and expansion of human populations and settlements. One aspect examined of human-lion coexistence and conflict will be the depiction of lions, aspects of which are still relevant today, in what Kusimba calls the European myth of “a wild Africa teeming with riches and savagery” which became imprinted in European minds1.

Africa in the centuries before colonial occupation Although there were growing centres of population, the rise and fall of empires and kingdoms from the 14th century to European penetration in the 19th century, sub-Saharan Africa remained underpopulated. Most people lived in rural communities dependent on pastoralism, crop cultivation and supplememtary hunting, or mixtures thereof, with the persistence in some areas of hunter-­ gatherer lifestyles2. Advances in livestock husbandry and settled forms of agriculture had an effect on wildlife, chiefly through habitat loss. This was magnified by the arrival of European traders and hunters, who ventured into the interior for profit and adventure, and the consequent spread of firearms from the 5th century onwards. European traders encouraged increased hunting by indigenous people for ivory, horn and skins to supply European demand. On the Indian Ocean coast, the continuing commerce of Arab-Shirazi, Swahili and a growing

54  From 14th century to colonial occupation

number of Indian merchants fed markets for wildlife products in West Asia, India and China. Ivory, horn, animal skins, spices and slaves were exported, manufactured goods and firearms imported. Arab traders sold muskets and gunpowder in North Africa, parts of West Africa, along the Red Sea and the kingdoms of the Horn of Africa and in parts of East Africa 3. The Portuguese, and later the Dutch and British, were wary of supplying firearms, but began to trade them in return for ivory and other produce. The British, Dutch, French and Portuguese set up trading posts along the West African coast, and the Dutch created their refuelling settlement at Cape Town.

Southern Africa – settlement, expansion and the European-led destruction of wildlife In the 15th century, the region’s population was a mixture of San hunter-­ gatherers, Khoikhoi hunter-pastoralists and increasing numbers of Bantu migrants from the north, who brought crop cultivation with them4. The region was sparsely populated in the west, but migration increased in north-eastern and eastern areas, which had better rainfall. Wildlife was abundant, with a diversity of ungulates and predators, including large populations of lions. Human settlement and agriculture had at this point only a minor effect on numbers and ranges of animals. There was conflict between pastoralists and predators, but the low density of human population limited conflict with lions and the abundance of natural prey limited lion predation on livestock. In 1487, the Portuguese explorer Bartolomeu Dias rounded the Cape, anchoring at Mosselbaai on the south coast. Ten years later, Vasco da Gama sailed into the Indian Ocean, opening the way for Portuguese trading expeditions. Commerce developed with East Asia and ports were established on the Mozambican coast, trading for gold, ivory and skins with the peoples of Mozambique, Malawi and Zimbabwe5. By the end of the 16th century, Dutch, English and French ships had rounded the Cape to trade in Asia. In 1649, a Dutch expedition wintered in Table Bay and the Dutch East India Company decided to occupy the area to reprovision ships sailing to Asia. In 1652, Jan van Riebeeck arrived as commander of the expedition to establish a settlement to supply fruit, vegetables and meat to the trading fleets. The landscape and vegetation encountered by the first Europeans at the Cape was a mixture of fynbos (from the Dutch fijnbosch, meaning fine-leafed bush6), wooded mountains and grassland in the coastal plains. Good sources of water supported a diversity of herbivores and predators, including lions, leopards and hyenas. Company workers and settlers grew fruit, vegetables and kept cattle and sheep. The enclosure of land by settlers led to conflict with Khoikhoi pastoralists, who opposed fencing of what had been their grazing land, and with San who had hunted there for centuries. The settlers also hunted the game herds for food and skins, and began to shoot predators to protect livestock. The possession of firearms and horses gave the Dutch huge combat advantage over the Khoikhoi

From 14th century to colonial occupation  55

and San7 but also huge advantages in hunting wildlife, of which they made full and destructive use8. The first administrator of the settlement for the Dutch East India Company, Van Riebeeck, kept a detailed journal, which provides information about the presence of lions. He also recorded the widespread killing of antelopes for food and for skins to export to Japan and other markets. In December 1652, Van ­R iebeeck reported the first case of stock predation, writing that seven or eight “wild beasts” killed a sheep. On 12 January 1653, he is more specific saying that a lion killed a sheep and was driven off with musket fire; another lion was seen near the carcass9. On 23 January, “lions tried to storm the kraal” at the fort, in which the Dutch kept their livestock at night10. Wildlife was a resource for food and commerce, and game around the settlement was soon depleted. Whether this led to lions raiding the livestock or whether lions were attracted by enclosed, docile prey, is unclear. But lion attacks continued and in 1657 there was the first recorded attack on a settler, Wouter Mostert, who was nearly killed when a lion entered his farm close to the fort11. On 12 September, Van Riebeeck records that after a search on Lion Mountain at Cape Town (named for its shape), the body of a missing Dutch soldier was found: “the brain pan had already been bitten off, so that it is presumed that he must have been killed by a lion, from which animal our people sometimes suffer loss and attack”12. In July 1661, a Dutch hunter, Carel Broers, brought to the fort the skin of a lioness he had shot near the cattle kraal of the Caapman Khoikhoi community. Van Riebeeck says they would be given “the reward” to which they were entitled, suggesting the Dutch were giving a bounty for killing lions. On 31 October, he writes that two lions broke into the kraal behind the fort, killing an ox and calf and wounding several others13. While most settlers stayed near Cape Town under the control of the Dutch East India Company, some sought to evade its jurisdiction and went on hunting expeditions as far east as the Great Fish River, where they encountered Xhosa communities. In 1702, one party went there and reported large populations of wildlife with lions, elephants, rhino and buffalo14. By 1707, there were over 700 company employees at the Cape, a settler community of 2,000, a large population of servants, local pastoralists who traded with the Dutch and slaves captured during conflicts with indigenous groups or imported from Benin and Angola. As early as 1710, the Dutch East India Company established outposts along the Atlantic Coast past Saldanha Bay and the Elephant River, inland as far northeast as Graaf-Reinet, and in the east to the Fish River beyond Algoa Bay (Port Elizabeth)15. These were areas rich in wildlife. The free burghers, as the white settlers were known, were harsh and racist in their treatment of indigenous people but rebellious towards the Dutch East India Company – starting the tradition of the trekboers, the families who moved to new land to avoid regulation (first by the Dutch and then the British). They lived by stock farming, cultivation and extensive hunting for meat, hides, ivory, rhino horn, lion skins, claws, etc. to sell16. As they moved east and north, the trekboers

56  From 14th century to colonial occupation

relied on killing game for meat and hides, to preserve the livestock which was their basic resource for establishing farms and pulling wagons. From the mid17th to the mid-19th centuries, they wiped out the huge herds of grazers in the Cape and Karoo, removing the natural prey of lions, which were then killed to remove the threat to livestock. But for 50 years after Dutch settlement, lions were common across the Cape. In 1707, one Dutch settler, Pete Kolb, said lions were still seen right up to the edge of the town, while 18th century travellers in the Cape like Sparrman, Paterson, Thunberg and le Vaillant all reported regular sightings within easy reach of Cape Town. Lions were especially numerous in the Karoo and east along the Olifant River17. The governor of the colony in 1776–7, Hendrik Swellengrebel, warned that when crossing the mountains east of the town, travellers needed to take spare oxen, as animals frequently fell prey to lions at night18; this crops up repeatedly in the narratives, with lions recorded as attacking camps at night to prey on oxen, donkeys and horses. Mostert writes that the Dutch hunter and cattle dealer, Joachim Prinsloo, who traded for livestock with Xhosa communities near the Fish River, had to arm the Khoikhoi herders on his farm with muskets, “as the lions were very bad there”19. The British occupied Cape Town in 1795 during the Napoleonic Wars, retaining it under the 1814 peace settlement. It was an ideal revictualling station for ships going to British possessions in India. The British inherited a fractious community of Dutch settlers, company employees, their slaves and Khoikhoi communities. By this period, as hunters and traders penetrated eastwards and northwards, lions disappeared around Cape Town and adjacent areas. In farming districts they were seen as vermin, with trekboers and settled farmers across the colony complaining of the depredations of lions on their trek oxen 20. In 1823, a former British naval officer, Francis Farewell, led a trading expedition to the east coast. It anchored at the bay that became Durban. Farewell did not have initial success in opening trade, but saw the potential for commerce with the thriving Zulu kingdom. Commodities like ivory, skins and other wildlife products were among the goods they sought. In 1824, Farewell and other traders visited the Zulu king, Shaka, taking him presents and obtaining permission to establish a trading post on land at the bay, which was named Port Natal by the British. British traders and hunters proceeded to conduct commerce and hunted in areas where Shaka granted permission. Meanwhile, from the Cape they pushed eastwards, coming into contact and conflict with the Xhosa kingdoms21. The arrival of travellers and traders led to the spread of firearms across the region. British hunters and traders provided some of the most interesting accounts of encounters with lions in what became South Africa. Few had scientific backgrounds and their observations must be treated with care, though they do give an idea of where lions occurred and interactions with them. Sir John Barrow, private secretary to the first British governor of the Cape, gave extensive accounts of his travels across the Cape22. Journeying to Paarl, the Karoo and then GraaffReinet (north-east of the Karoo), he describes seeing zebra, quagga, gemsbok, kudu, springbok and ostriches. In the Swartruggens area of the Karoo he first

From 14th century to colonial occupation  57

encounters lions, noting “The prints of the feet of this destructive animal were every where fresh on the road, and every night we heard them roaring around us.”23 At a farm near Swartkop salt pan, a Boer farmer tells him he is moving to a new location because of attacks by lions on his oxen and horses. The night before Barrow arrived, two of the farmers’ horses were eaten by lions24. In the eastern areas around the Sneeuwbergen escarpment, near present-day Cradock, the farmers and local people kept their livestock in strong enclosures at night because of lions. Travelling north from Cape Town along the Atlantic Coast into Namaqualand, Barrow says Boer farmers lived in fear of lion attacks. He is told that just before his arrival, a Khoikhoi herder had tried to drive cattle to a watering point when a lion appeared and, bursting through the cattle, pursued the herder until he climbed a tree, where he had to stay for 24 hours until the lion moved off. When he came down from the tree and ran back to the farm where he was employed, the lion followed him to within 100 yards of the farm. Local people told him that, “It seems to be a fact well established, that the lion prefers the flesh of a Hottentot [Khoikhoi] to that of any other creature. He has frequently been singled out from a party of Dutch. The latter being disguised in clothing, and the former going generally naked, may perhaps account for it. The horse, next to the Hottentot, seems to be his favourite food”25. More than a decade after Barrow, William Burchell (after whom the southern African subspecies of zebra is named) went to South Africa to gather natural history specimens, including skins. He shot 289 mammals during his trip. On 7 September 1811, the indigenous hunters accompanying him kill a quagga and wound a lion near Spionberg, on the eastern edge of the colony. Up until that point he had expressed disappointment at how little wildlife he had seen crossing the Cape and the Roggeveld Karoo, noting instead the large flocks of sheep and goats26. On 13 November 1811, returning from the Ky-Gariep River (a tributary of the Orange), he describes a stormy night during which lions circle the camp, saying, “Such nights I already knew, by dear-bought experience favour the prowling lion, and seem to give him a spirit of daringness which he seldom evinces at other times”27 Other European hunters who travelled through this and neighbouring regions, including Gordon-Cumming and Selous, said lions were inclined to raid livestock in camps on stormy nights, taking advantage of the confusion caused by the storm. Burchell is one of those hunters who seem keen to kill lions but have little respect for them, saying they are an “indolent and skulking animal”28. Twenty-five years later, the hunter William Cornwallis Harris undertook a long expedition from Cape Town, through the Eastern Cape into what became Transvaal and the territory of the Ndebele king, Mzilikazi. Although wildlife numbers had been reduced by the trekboers as they deserted the Cape, Harris reported seeing large herds of springbok and black wildebeest between Grahamstown and Graaff-Reinet and in the Sneeuwbergen, where lions were frequently seen hunting antelope29. In the region of the Maritsane River, to the west of

58  From 14th century to colonial occupation

where the Kruger National Park’s south-western border stands, Harris expresses concern about the lions prowling round his camp at night and says three lions broke through a thorn fence, killing two sheep30. When he visits Mzilikazi’s kraal in northern Transvaal, he says the king’s residence is “strewed with the skulls, paws and tails of lions, some of them quite fresh”31. Harris says the Ndebele are “fierce” in the protection of their herds from lions and if a lion attacks cattle, then the herder either kills the lion or is killed by the lion when he tries to hunt it down32. Journeying from Ndebele country down through Tswana areas of what is now South Africa’s North-West province, he repeatedly refers to lions trying to raid his camp at night to kill the oxen. During much of the 19th century, European hunters were active in the Eastern Cape, North-West and Northern Cape hunting for meat, hides, skins and ivory, and trading with local communities, who were encouraged to hunt. Kuruman in South Africa’s Northern Cape was a centre for trading in wildlife products. The Tswana of this region and in Botswana hunted to supplement food from livestock and cultivation, as attested to by finds at archaeological sites, showing varied bone assemblages including local fauna 33. Hunting large antelope and carnivores would have involved cooperative effort, given the Tswana didn’t use horses or firearms to hunt until the mid-19th century. Some of the assemblages include remains of zebra, elephant, buffalo, kudu, impala and giraffe, “with regularity and relative abundance”34. Lion skins were used by the Tswana for making cloaks, tails for ornamentation, claws and teeth for necklaces, bones for pipes, and body parts for medicine or charms (muti)35. Hunting by them increased in the mid-19th century to supply European demand – it was part of the “transformation of the non-human environment into a commodity [and] was the entry of the cash economy and the depletion of game from most areas of the subcontinent”36. During travels in the Kalahari from 1849 to 1863, James Chapman often mentioned seeing large numbers of jackal, lion, leopard, cheetah and other prepared skins, and ivory; some of these had been hunted by the San to trade to the Tswana, who sold them to Europeans37. Morton and Hitchcock record that in the mid- to late 19th century there was “specialised hunting in the Kalahari for high value animals (i.e., animals with high value body parts). They included elephant, rhinoceros, lion, leopard, cheetah, blackbacked jackal, ostrich for feathers, gemsbok for horns”38. One of those who used Kuruman as a way station was Roualeyn Gordon-­ Cumming. He undertook hunting and trading expeditions for ivory, skins and ostrich feathers, starting in 1843 in the Eastern Cape. As he moved into the interior he met Boer farmers and found that they not only killed lions to protect stock but to obtain their skin, which they used to make the straps to harness horses to wagons, believing it to be stronger and more enduring than other hides39. Passing through farming areas, he expressed surprise at the large numbers of wildebeest and springbok skulls in farmyards, showing the extent of killing for meat and hides40. After crossing the Great Fish River into sparsely populated areas, he saw huge migratory herds of springbok. Nearer Colesberg

From 14th century to colonial occupation  59

and the Orange River, the herds diminished but springbok were still plentiful along with black wildebeest and quagga – he makes no mention of seeing lions41. Moving into the main part of the Karoo, he reports seeing increasing numbers of gemsbok, being told that the region’s lions preyed mainly on them. Later in his journey, near the Riet and Vaal Rivers to the north, he comes across a party of Boer hunters who had recently killed a lion and lioness which had attacked their stock. Gordon-Cumming said the Boer way of killing lions was to fill them full of bullets before daring to approach; the carcasses of the two lions he sees had their heads “shot to pieces”42. The image he conveys is of hunting to exterminate lions and not just remove stock raiders. Gordon-Cumming held lions in high regard. He wrote: “There is something so noble and imposing in the presence of the lion, when seen walking with dignified self-possession, free and undaunted, in his native soil, that no description can convey an adequate idea of his striking appearance.”43 He, too, warned that they were the greatest danger to travellers on rainy or stormy nights, when they took advantage of the noise to seize stock44. Gordon-Cumming b­ elieved that man-eating by lions mainly involved old male lions experiencing difficulty hunting wild prey. Among the Tswana of the region, he said man-eating was encouraged by the custom of not burying the dead, leaving bodies to be scavenged by lions, accustoming them to eating people45. Gordon-Cumming, despite his admiration for the lion and awareness of their scarcity, was keen to hunt them and first recorded killing a lioness on 23 March 1844, having hunted it on horseback. During this trip to Bamangwato (Tswana) territory he killed several lions and skinnned them all, to sell the skins when he returned to Grahamstown. I won’t detail all the lion hunts, but they become a major part of the narrative in both books referenced, with Gordon-Cumming sitting over waterholes and killing the lions as they came to drink or pursuing them on horserback before dismounting to shoot them. Harris, Gordon-Cumming, Selous and other hunters wrote exhaustive accounts of their hunting, stressing the rigours, danger and implied heroism of it all. The books and articles in newspapers and journals became part of the creation of an image of Africa for British people. Along with the Livingstone legends and other missionary stories, this helped form “An emotion culture – performed publicly, exercised privately, highly symbolic and creating an imperial community of feeling” that encouraged the development of colonialism and the concept of Africa as a dark and dangerous continent where European men could prove their heroism as hunters, missionaries and, ultimately, occupiers46. Returning hunters, Selous in particular, held public lectures boasting of their adventures and displaying lion skins and skulls. Their exploits, and the less romanticised but even more widespread hunting by Boer trekkers, had a huge effect on southern Africa’s wildlife, with the extinction of some species and near extinction of others during the 19th century. Accounts by hunters like Selous – who became what the wildfowler-turned-­ naturalist, Peter Scott, described as “penitent butchers”, bewailing the depletion

60  From 14th century to colonial occupation

of game and becoming enthusiasts for conservation47 – shifted blame from Europeans to indigenous African communities. The attribution of responsibility for the depletion of game in southern Africa became a highly politicised issue. English-speakers blamed Boers and overall “there was a tendency in the colonial period to blame Africans for the destruction of nature … Africans were characterized as being cruel to animals”48. Africans were criticised for using pits, snares and poisoned arrows rather than the more “sporting” guns. But as Beinart and Hughes concluded, “it is clear that firearms were far more significant in the overall destruction”49. European hunters and later settlers saw wildlife as something for commercial exploitation, sport or to be exterminated in farming areas. African communities hunted, as they had always done, but not with the destructive effect seen in the 19th century when the combination of European hunting, global demand for wildlife products and the spread of firearms led to a massacre of wildlife. In southern Africa, large areas were cleared of all but the smallest animals. The introduction of sheep to South Africa by settlers led to the exclusion of wildlife and systematic killing of predators. Merino sheep were first brought to South Africa in 1830 and by 1890 there were 12 million. The establishment of sheep farms was a key determinant in the decline in the huge springbok herds and the ending of the annual migration of hundreds of thousands from the northern Cape and southern Kalahari to the Karoo. They were shot in huge numbers, as were the lions, wild dogs and other predators preying on them 50. Human-wildlife conflict was in an extreme phase and led to the depletion of lions and their prey over most of the Cape Colony and then what became the Orange Free State and Transvaal. Hunters had to travel further afield into Botswana, Zimbabwe and Zambia to shoot game for sport or gain. Trophies were sought, both for personal pride or to sell to wealthy collectors in E­u rope. As Mackenzie explains, “The possession and public display of hunting trophies, particularly of spectacular or exotic animals, became a fashion during the 19th century and a symbol of wealth and status … Great cat skins became the thing to have on the floor of your mansion or great house. People basked in the reflected glory of hunting prowess”51. There was also a scientific side to it and a desire to increase knowledge. Hunters, like Gordon-Cumming, Oswell and Selous, were on commission from museums to collect specimens, and the ­Zoological Society of London in its Proceedings published learned papers on the specimens collected. Some African communities engaged in large, communal, ritualistic hunts. Beaters drove animals into a fenced area to channel them towards hunters. These were used by the Ndebele, Xhosa and Zulu. The Zulu king Shaka was particularly keen on them and would kill the driven animals with spears and even axes. He liked to kill the most dangerous animals – elephants, rhino, buffalo, lions and leopards52. One hunt, thought to have taken place in August 1819, involved the driving of hundreds of animals towards Shaka and his hunters. There is no way of verfying the accounts, but they say that three prides of lions were driven towards the king. He speared one male and then a group of lions were trapped in

From 14th century to colonial occupation  61

an enclosed area into which Shaka and other warriors entered and, according to Ritter, engaged in single combat with lions, armed just with spears53. Until the middle of the 19th century the fauna of southern Africa was “richly diverse, highly prolific, and widely dispersed”, according to Mackenzie. Throughout history, people had exploited it as a food source and defended their stock and crops against animal raiders. Game meat was a supplementary source of protein for pastoral or farming communities and vital for hunter-gatherers, but was not exploited on a scale that decimated game herds. It was the expansion of white settlement and hunting that caused the fauna of southern Africa to be “sent into rapid retreat”54. Lions and other wildlife were plentiful in Zululand and Natal throughout much of the 19th century but by the end were severely depleted by human expansion, the development of large-scale farming, the killing of predators to protect livestock, and sport and commercial hunting. Guggisberg believes the last lion in the Cape colony was killed by a trap-gun in 1850, north of Port Alfred55. Prior to settlement in Natal, wildlife had been abundant and indigenous agriculture had only a minor effect, being based on livestock and small-scale cultivation around homesteads. The terrain of steep hills, deep gorges and broken ground limited expansion56. Even Shaka’s huge hunts had hardly dented wildlife numbers. Settlement by Europeans took off in the 1840s, encouraged by the British government, and there was a substantial increase in cultivation and pastoralism. Hunting was widespread for meat, ivory, skins, sport and livestock protection. As Ellis writes, “The carnivores which were feared the most were the lion and the leopard, for they were considered to be dangerous to persons and their livestock. The settlers therefore attempted to eliminate these animals from the environs of the town. Lion seem to have moved away from the settlement [Port Natal] fairly early on.” The last record of a lion there was recorded in the Natal Commercial Advertiser on 8 July 185457. The lion was gone from British-­ settled Natal by 186558. In Tswana and Sotho areas, hunting was common with snares and pitfall traps commonly used to catch antelopes and buffalo for meat and hides59. Lions tended to be hunted only when they became a threat to people or livestock. Despite the European colonial narrative of wasteful and excessive African hunting, accounts by early European travellers show it was the arrival of European settlers and hunters that led to the accelerated disappearance of game. There is evidence from those hunters that Mzilikazi and Lobengula of the Ndebele tried to limit European hunting in their territory, especially by Boers, who gained a reputation for excessive killing to produce biltong and hides60. The Ndebele were pushed out of northern Transvaal by the arrival of large numbers of Boer settlers in 183761. Forced to retreat north of the Limpopo, Mzilikazi established a new kingdom in southern Zimbabwe, where he again attempted to limit European hunting 62, though hunters like Selous were able to gain permission to hunt by applying to the king. The Tswana chief, Khama, also limited European hunting in Bamangwato lands, because of the danger of depletion of game, which his own subjects hunted for meat and hides63.

62  From 14th century to colonial occupation

Settlement by Boer farmers north and west of the Zulu kingdom, north of the Orange River and then between the Vaal and the Limpopo, led to the spread of livestock and arable farming, the rapid increase in population numbers of white settlers, and a consequent decline in wild herbivores and predators dependent on them64. In the Orange Free State and Transvaal, where Harris and Gordon-­ Cumming shot many lions in the 1830s and 1840s, there were few if any left by the end of the century65. A study of published references to wildlife in the 19th century in the highveld region of the Free State and Transvaal records that eight large herbivore species (quagga, Burchell’s zebra, eland, black wildebeest, blue wildbeest, red hartebeest, blesbok, springbok) occurred in large numbers in the highveld, supporting a substantial predator population66. Accounts from the 1850s speak of lions following migrating herds of antelope across the Drakensburg from the Free State into northern Natal and Zululand. But the large herds began to disappear by the 1870s and were gone within two decades67. Boer settlers were avid hunters, some killing huge numbers of lions and game. Kotje Dafel and Petrus Jacobs are credited with killing over 100 lions each. In the 1870s, Frederick Selous engaged in his hunting expeditions in southern Africa, with the aim of making money, partly by supplying specimens for museums and collectors in Britain. His biographer says that Selous’s and other contemporary accounts reveal that by 1870 the movement of settlers into the Orange Free State and Transvaal had “cleared the Highveld of big game all the way to the Limpopo”68. By 1884, the Zoological Society of London recorded that one of its Fellows reported from South Africa that while leopard and cheetah could be found in the vicinity of Grahamstown in the Eastern Cape, the lion had been exterminated years before69. In December 1871, Selous travelled along the Orange River between Kimberley and Upington, recording that there was no game in the region, not even springbok70. It is only when Selous talks of his hunts in Bamangwato territory in Botswana that he records seeing and hunting lions. Moving on to Zimbabwe, he meets the famous Boer lion hunter, Petrus Jacobs, along the Sebakwe River. As Selous recounts, “Eight days before, this old Nimrod, who has probably shot more lions than any man that ever lived, had been terribly mauled by one of these animals” 71. It was three years into his hunting trips that Selous shot his first lioness, between Victoria Falls and Wankie in Zimbabwe in December 1874. Indicating how rare they had become, even in Zimbabwe, Selous says it was the first lion he had seen since July 187272. His second lion kill was in 1876 on the Shashe River, where Zimbabwe meets Botswana73. Hunting near Serule in Central Botswana in December 1876, he recorded, “All over the interior of South Africa, wherever game still exists in sufficient quantities to furnish them with food, lions are to be met with, and are equally plentiful on the high open downs of the Mashuna country … in the dense thorn-thickets to the west of the Gwai River, or in the marshy country in the neighbourhood of Linyanti” 74. Selous had less than complimentary things to say about lions, despite his supposed heroics in killing them, stating that, “It has always appeared to me that the

From 14th century to colonial occupation  63

word ‘majestic’ is singularly inapplicable to the lion in its wild state, as when seen by daylight he always has a stealthy, furtive look that entirely does away with the idea of majesty.” But when standing at bay, Selous says the low-held head, the twitching tail and growling mean “no animal can look more unpleasant than a lion”. Selous writes that when lions are hungry they fear nothing and will do anything for food even though fired at. He recounts an episode when he was camping in 1880 on the Umfule River and a lioness repeatedly tried to get at cattle in a strongly built kraal. Despite keeping fires burning, the lioness persisted and attacked the people in the camp when Selous was away, seizing one of his servants. When driven off, it returned and immediately attacked another man. The lion was eventually shot when she came back and tried to attack again to obtain a meal. The coat and teeth of the lion were in good condition, but it was very thin, indicating that it was hunger that drove her on75. In his Hunter’s Wanderings, Selous describes a dozen different lion hunts or encounters. In November 1883 in Mashonaland, his camp is attacked at night by a lion, which one of his servants kills. The following day he hunts a large male lion, with a very good mane. He fatally wounds it but before it dies, one of his European companions shoots it again. Selous is furious because “he had knocked a hole through the lion’s skin about the size of a shilling”. Selous says he abused him “very freely,” as he was collecting skins to sell and wanted them as little damaged as possible76. Later in his account, Selous gives details of an old lion that killed several people from a village in Mashonaland in 188677. In this area of Zimbabwe, Selous says game was scarce and lions became very dangerous. They were said to have carried off so many women while they were working in their cornfields, that the Mashona families living in the district fled. Stevenson-­ Hamilton, the famous warden of Kruger National Park, said that the rinderpest epidemic of the 1890s in eastern and southern Africa had a strong effect on the propensity of lions to attack stock and people. Lions benefited from the deaths of huge numbers of cattle and game from disease, feeding off the carcasses with little need to hunt. By the time the epidemic was over in late 1896, lions had increased in numbers because of the abundance of food. But after the epidemic there was less game for them to hunt and they relied increasingly on stock-­raiding and killed more people, because they could get no other prey78. By the end of his hunting career Selous had killed 31 lions out of about 1,000 head of game79.

Malawi, Mozambique, Namibia and Angola European hunters and traders moved into Malawi towards the end of the 19th century, leading to the establishment of the Nyasaland and District Protectorate in 1891. The protectorate developed from the African Lakes Company’s trading activities, mainly to access the substantial amounts of ivory from the region80. The Portuguese had by this time expanded their coastal and riverine trading settlements into a large colony. Hunters like Vaughan Kirby used Malawi as a starting point for expeditions down the lakeshore and the Shire Valley, and along

64  From 14th century to colonial occupation

the Zambezi into Mozambique. During one hunting trip in 1894, he laments the few sightings of lion or their spoor in southern Malawi or the Zambezi region of Mozambique. Two years later, he sees only the spoor of one male and one lioness near the confluence of the Shire and Zambezi81. Moving into the gamerich Gorongosa region, he hunts lions and, in attempting to kill two lionesses he tracks, one of his “kraal-boys”, as he calls his locally employed servants, is killed by a lioness wounded by Vaughan Kirby. He finds more lions along the coast than inland82. Robin Wright, who wrote of his experiences for the British Country Life magazine, started big-game hunting when he was working on a plantation near Zomba in Malawi. This forested and hilly area had a population of lions which, as settler agriculture took land to grow cash crops and settlers shot game for meat and sport, began to regularly attack domestic stock. Wright says that on one occasion a lioness broke into the goat kraal at night and killed 70 animals – mass killing being quite common when predators break into a kraal from which the stock cannot escape. The lioness was shot after killing the goats83. Wright notes that Malawi was badly affected by the rinderpest epidemic, with cattle and large amounts of game dying in the mid- to late 1890s. Around this time, local communities started complaining to the British about attacks on stock and people by lions84 – a result of the depletion of prey by rinderpest. In Namibia, lions were found across much of the country, including the arid pre-Namib, Kaokoveld and Damaraland. Game was plentiful, with large herds of gemsbok, springbok. zebra, giraffe, wildbeest, hartebeest, kudu and other antelope. The density of the human population – a mixture of San, Nama, Herero, Damara, Ovambo and other communities, plus Boer farmers who had trekked north (known as the Thirstland Trekkers) – was low and the main forms of livelihood were pastoralism, hunting and some cultivation in Ovamboland and the Caprivi. The Nama, Damara and other peoples of Khoikhoi descent mixed livestock husbandry with hunting and foraging. The Nama had been pushed north by Dutch settlement in the Cape in the 18th century and had moved into Namibia from the Fish River. By the time of their settlement in Namibia many of them had horses and guns, which increased their ability to conduct raids for cattle against neighbouring communities and protect their stock against predators 85. The Nama and Herero rarely hunted for meat, except in times of particular need, but killed predators to protect their stock86. Lions were recorded as being encountered by travellers, hunters and missionaries in Namibia throughout the 19th century. Charles Andersson, accompanying the British geographer Francis Galton on a Royal Geographical Society expedition to Namibia in 1850, records seeing his first lion at Scheppmansdorf, south of Walvis Bay. The lion entered the village in which he was staying and killed a dog. The villagers killed it. Andersson said, “The natives highly rejoiced at the successful termination of the hunt, for this lion had proved himself to be one of the most daring and destructive ever known, having … killed upward of 50 oxen, cows and horses … every successive attack made upon him only served to increase his ferocity”87. Journeying north, he reported that an area called

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Daviep, near the Swakop River, was said to be a favourite place for lions, rearing their young there. Andersson’s party found the tracks of numerous lions. During the night, one of their mules and one of their horses were killed and eaten by lions. Later in the day they spotted six lions sitting on rocks from which they had intended to try to stalk the killers of their animals88. Further along the Swakop River, they are told by locals that lions are present in large numbers and are a constant danger to people in the area, even though game is plentiful. Travelling along the river, he meets a Danish hunter, Hans Larsen, who recounts how he was riding one of his oxen when a lion attacked and tried to pull him off it. He said he had been hunting there for a while and admitted denuding the area of its game by shooting extensive numbers of rhino, lions, giraffes, wildebeest and gemsbok89. Near Richterfeldt, Andersson is told by Afrikaner settlers that lions are very bold and approach houses searching for prey. While camped there, Andersson hears a lion killing a zebra close to his camp. The lion later enters the camp, despite them keeping a fire burning, and kills one of their dogs90. Despite the confidence of the lions, he is told that Damara and other local people drive lions from fresh kills to take the meat 91. Stock-raiding by lions, hyenas and leopards is such a problem that many farmers set up gun traps to kill predators. The regular killing of substantial amounts of game in some areas and the availability of cattle, smallstock, horses, donkeys and trek oxen may have combined to encourage stock killing by lions and the overcoming of fear of people through hunger. Back in the Richterfeldt area, Andersson says local farmers told him that lions had become particularly numerous and that one local hunter and his sons had killed more than 20 in the last year 92. The San of Namibia and the western Kalahari lived alongside lions and hunted much of the same prey, but did not appear to have an adversarial relationship. ­Suzman notes that among the Ju/’hoansi of eastern Namibia, with whom he lived while studying their culture, lions were considered to be tolerable neighbours and they shared space, waterholes and prey species with them. Ju/’hoansi hunters would only appropriate lion kills when lions had fed and there was unlikely to be serious conflict. They told Suzman that when they dismembered prey they had killed there was always part of the carcass left behind if it was a large animal, and this the lions would take93. None of the San communities hunted lions for food and as they did not keep domestic stock, until persuaded to do so by Europeans in the 20th century, they did not come into conflict over stock-raiding. The relationship between people and wildlife in Namibia began to change in the 1880s when Germany laid claim to the region, with the first German outpost established at Luderitz in 1882. This enabled German Chancellor Bismarck to lay claim to what became known as German South-West Africa at the Berlin Conference of 1884, at which the major European powers divided up Africa between them. The influx of German administrators, traders, settlers and soldiers completely altered the nature of the region, with commercial farming, the search for valuable minerals and hunting for profit and to protect livestock taking its toll on wildlife in many areas.

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To the north in Angola, Portuguese colonisation had started in the late 15th century. Over the next three centuries, their presence grew, mainly due to the slave trade, which provided labour for Portuguese plantations in Brazil. Plantation agriculture developed in the north and centre of the country, where rainfall and soils were better suited for growing coffee, cotton and other cash crops. In the south and south-east, population density was low, Portuguese settlement was minimal and the initial effects on wildlife consequently limited. The areas adjacent to Namibia along the Kunene River were rich in wildlife, especially the coastal areas and inland from the coast north of the Kunene, in what became Iona National Park. To the east, the arid savanna and dry woodland supported a similar range of antelope as the nearby areas of Namibia and northern Botswana, and large numbers of predators. But there is little recorded evidence of the numbers and distribution. One of the first written accounts of Angola’s wildlife came from Joachim John Monteiro, a Portuguese colonial official and naturalist. He worked in Angola from 1858 to 1876 and was a keen collector of natural history specimens. Travelling widely across the territory he noted that in Cuanza Norte, east of Luanda, cattle were predominant with the result that wildlife of all sorts was scarce. He doesn’t mention any sightings of lions or their tracks94. Further south, beyond the Kwanza River mouth, he wrote that the road to the town of Calumbo “used to be formerly much infested with lions, but with the greatly increased traffic they are seldom now seen or heard”. Giving no dates, he said lions used to be found very close to Luanda and in recent memory had taken cattle from walled enclosures near the capital95. Further south in Cuanza Sul province, near Sumbe, he was told the area was infested with lions, though he only saw spoor and no lions. He was warned by other travellers not to stop long at a waterhole along the road, as it was known by local people to be visited regularly by lions at dusk. A Cabindan traveller they met along the way was reported to them as having been killed later by lions at the waterhole96. At the settlement of Dobe Grande, further south, Monteiro was told that lions regularly tried to break into enclosures containing sheep and cattle. The lions were said to move into the area after the first rains had brought forth a good crop of grass and herds of antelope arrived to take advantage of the grazing. In this area lions were said by colonists to have killed and eaten slaves sent out into the fields to harvest crops97. Boer settlers who trekked into southern and central Angola in the mid- to late 19th century hunted game for biltong and hides, and killed lions to protect their livestock. In central and eastern Angola, game numbers were reduced heavily by Boer settlers, many of whom worked as transport drivers when the Portuguese were building roads and later the Benguela Railway. In 1881 there were still large herds of game and plenty of predators in southern and south-east Angola. The hunters Axel Eriksson and William Chapman travelled across these areas in that year on a hunting expedition and reported that they were teeming with game including lions, leopards, black rhino, buffalo, elephant and a diversity of antelopes98.

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East Africa – European penetration and the growth in human-lion conflict In much of East Africa, especially savanna areas of central-southern Kenya and northern Tanzania, human populations were in a state of flux in the 18th and 19th centuries, with movements of pastoralist communities across the region. Maasai dominance was established in many areas around Laikipia and south towards Narok and the Mara. The spread of the Maasai southwards brought new factors into play in human-lion coexistence there, as the Maasai who settled in the Serengeti-Mara ecosystem in the mid-19th century were fierce protectors of their livestock and killed stock-raiding lions. Out of necessity, lion hunting became an important duty for men in communities, like the Maasai, living chiefly as pastoralists, leading to the tradition of ritual lion hunts as part of manhood initation rituals, according to Schaller 99. Maasai social organisation and an individual’s responsibilities within the community are based on gender and on age-sets. Young men from around 15 to 30 make up the warrior set and among them bravery is seen as a virtue bringing respect within their age group and the wider community – a major measure of bravery being participation in lion hunts or protecting cattle from human raiders. As Hazzah et al. set out, the relationship between Maasai and lions is multifaceted, and “includes positive and negative attitudes” and must be evaluated carefully in order to understand lion killing100. Historically, killing a human enemy or a lion were comparable feats in gaining prestige. As European occupation of land and later imposition of colonial laws changed Maasai relations with other communities, killing human enemies became rarer, so the killing of lions became more important as part of a young warrior’s attainment of prestige101. In an almost counterintuitive way, through the ritualised killing, the Maasai “have historically valued lions (except when they have attacked livestock) because they provide warriors with a cultural role that reasserts their power and strength as they protect their communities”102. The killing of lions was a duty not a sport and it went beyond mere responses to instances of predation by lions on cattle. But it did not happen so regularly as to seriously threaten lion numbers. Wildlife was still abundant in the grassland and woodland savanna regions populated by these pastoralists. To the west of the Maasai, the Nandi and other Kalenjin-speaking groups settled on the highlands of the Rift Valley. They combined livestock husbandry with crop production, supplemented with hunting when necessary. The Nandi developed male age-sets and through manhood rituals established them as warrior groups tasked with protecting cattle against raiders (chiefly Maasai), but also from attack by predators. Having organised units of men made them more effective in deterring or hunting down dangerous predators like lions. To the south-west, on what is now the western border of the Serengeti National Park, communities such as the Ishenyi and the Ikoma lived alongside the considerable concentrations of game animals and the lions preying on them. They, too, developed age-sets of young men and rituals of manhood involving hunting lions. In a

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ritual called aghaso young men hunted lions or leopards and the young man who killed the animal would take trophies from it and eat a portion of the heart103. The continued development of trade on the Indian Ocean littoral, controlled by Arab, Afro-Shirazi or Swahili traders, had an effect on the hunting practices of communities across the region, as demand continued to grow for ivory, but there was also a market for the skins, teeth and claws of lions and leopards104. The trade brought firearms into the region, a commodity much in demand in exchange for slaves, ivory and wildlife products105. The expansion of commerce attracted European hunters and traders. From the mid-19th century onwards there was a growth in British expeditions seeking to establish trade routes and map out the river and lake systems. In July 1857, Richard Burton set off inland from the coast at Kaole having kitted out the caravan in Zanzibar with the help of the Omani Arab traders. Just inland from the coast he reports seeing areas of grassland and herds of zebra as well as numerous kudu and says his local guides warned of the danger of attacks by lions and hyenas at night106. In the populated districts of the regions he traversed, “game has melted away before the woodman’s axe and the hunters’ arrows,” and he went for days without seeing game. Burton says lion, and leopard skins were prized by Zanzibari traders107. Burton was accompanied in travels in the Great Lakes region by an Indian Army officer, John Hanning Speke. Speke planned to use his journeys with Burton to chart the territory and collect specimens to establish his own museum. In December 1856, passing through the Usumbara Mountains inland from Tanga he said lion tracks were often seen, along with those of a variety of antelope108. He periodically expressed disappointment at the scarcity of game, noting that the animals spotted could all be seen in South Africa in far greater numbers109. Further north, around Mount Kilimanjaro, lions were said by the colonial official Sir Harry Johnston to be “very abundant and very bold”, but that leopards were feared far more by the local villagers110. One of the major and most ecologically damaging consequences of European penetration and the role of trading communities, was the introduction of large numbers of guns into East Africa. The use of firearms had a huge and lasting effect on wildlife. Traded for ivory, slaves and skins sought by the merchants, they were often antiquated muzzle-loaders but had a major effect on the ability of communities to hunt for food, skins and ivory, to protect themselves and raid for slaves. By 1888, around 100,000 guns a year were being traded to people through Zanzibar alone111. European hunters also took an increasingly heavy toll as both commercial and sporting hunters saw East Africa as a limitless source of profit and quarry. Sport hunting, especially later in British colonies, “became a mark of class distinction”. Ironically, it was through sport hunting by wealthy and titled British settlers and visitors that eventually the idea of game reserves “based on the model of the nineteenth-century private shooting estates” came to form the basis for conservation. African communities, like the rural working class in Britain, were seen as impediments to “sport” and their hunting was frequently banned and criminalised112.

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Sudan, the Horn of Africa and Central Africa In the early 19th century, Sudan was controlled by Ottoman-ruled Egypt. The Ottomans had extended their hegemony in north-east Africa into Sudan in the 1820s113. Egyptian control was strongest near the Nile and fortified trading posts called zaribas were set up along its banks as far south as what is now South ­Sudan. The sought-after commodities for Arab traders were slaves, ivory, cattle and gold – they traded and raided for them using horse-borne Baggara and other Arab retainers, equipped with firearms. They didn’t occupy large areas but dominated the Dinka, Nuer and Shilluk communities, and encouraged them to hunt to supply wildlife products. Firearms, which had been scarce in the region, were introduced by the Arab trading and raiding expeditions. The arid regions of of northern and central Sudan had desert-adapted wildlife, including gazelles and lions, but wildlife was progressively exterminated in the heavily cultivated and populated areas along the Nile and pushed into areas unsuitable for cultivation. In 1821–2, a French engineer and administrator for the Ottomans, Linant de Bellefonds, reported that lions could still be found in the area south of Wadi Halfa, on the border with Egypt, and north of Dongala on the Nile. They inhabited woodland areas114 and preyed on antelopes but also entered settlements at night to kill cattle or camels. A British traveller called Hoskins reported in 1835 that despite the attacks on livestock and camels, the local people rarely killed the lions, which could still be found as far as 120km north of Khartoum. He added that they “infested” roads to Senaar and Atbara, above the fifth cataract on the Nile115. In 1861, the Khartoum-based ivory trader, John Petherick, said lions were common around the area of Abba island in the Nile, south of Khartoum116. In eastern Chad, Darfur and Kordofan they were still numerous in the 18th and 19th centuries. In the arid savannas and thornbush around Lake Chad, the German explorer Gustav Nachtigal was warned by his local guides on an expedition in 1869 of the presence of lions, which preyed on domestic animals and attacked people. They were to be found in good numbers in the Wadai region, and the forested valleys of Butiha and Batha north of Fort Lamy (Ndjamena)117. In more arid areas to the east and into Darfur and Kordofan in Sudan, the shortage of natural prey (resulting from desertification and human hunting) meant lions were a threat to nomadic pastoralists and their herds. Caravans that camped for the night near waterholes or rivers were particularly vulnerable118. Some of the nomadic groups – the Tuareg in Niger and Mali and the Baggara/Rizeigat of Darfur – hunted lions on horseback with spears. They later adopted firearms and reduced lion numbers in areas with little natural prey and so greater predation on stock119. Abyssinia, whose culture was closely bound up with the symbolism of the lion, had a declining population of lions by the 19th century. Lions and other wildlife were pushed into areas with low human settlement that were unsuitable for farming. A correspondent of the Zoological Society of London (ZSL) reported in 1868 that in western Abyssinia, near the border with Sudan, lions were mainly

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to be found in river valleys and thick bush in mountainous areas120. The hunter and traveller Lord Lovat obtained specimens of lions from a hunting expedition described by the ZSL as taking place “north of Addis Ababa”, which were sent to the Society121. The eminent historian of Ethiopia, Richard Pankhurst, wrote that lions were widespread up to the 16th and 17th centuries122. They then declined due to expansion of human settlements and agriculture and, particularly, to the increasing availability of firearms, which led to them being confined to the south-west by the end of the 19th century123. By the early part of the 21st century, they had been exterminated in 82% of their Ethiopian range and survived in only a few national parks and remote border areas. Somalia and what is now Somaliland (former British Somaliland) had a large and widespread lion population in the 19th and early 20th centuries and became a popular hunting ground for British and other European hunters. Extensive sport shooting and the regular killing of lions by Somali pastoralists, who by the mid-19th century had access to firearms, took a heavy toll by the turn of the century. Prior to the incursions by European hunters, lions had only been killed by local communities to protect cattle, camels and donkeys. In some Somali clans that hunted lions for these purposes, young men would wear the mane on their heads signifying that they had killed a lion124. Although Speke, recounting his expedition to Somalia in 1854, talks of the scarcity of game, he notes that northern Somali nomads built strong bomas to protect their camels and other stock from attack by lions125. The main accounts of lions in Somalia emerged in the 1890s, as more and more British settled in Kenya and used the territory to the north as a lion-­hunting ground. Lord Delamere, the prominent Kenyan settler, made many hunting trips to Somalia and Somaliland. His biographer, Huxley, says his letters and diary entries show that in December 1891 he and his companions killed five lions in a couple of days around Hargeisa, and that by 24 January 1892, they had killed ten126. Two years later on a safari there, Delamere was badly mauled by a lion and only saved by his gun-bearer, who was also injured by the lion127. ­Harald Swayne, a British army officer, was another who hunted extensively in the region between 1885 and 1893. He recorded his own lion-shooting expeditions but also noted that the Midgan people pursued lions with dogs, killing them with arrows128. They skinned the lions and used or sold the skins. In his journeys across northern Somalia, Swayne records lions being present in most districts, from near the port of Berbera to east of Hargeisa near the border with Abyssinia. As travellers in East and Southern Africa found, hunting camps attracted lions at night129. On the border of Somaliland and southern Abyssinia, Swayne said the local Malingur people told him that there was a problem with man-eating lions in their district and that a man had been carried away just before he arrived130. On an expedition in 1893 near Berbera, Swayne comes across a man dying from wounds received after being attacked by a lion. Shortly after, at a camel zariba, he is told that another man had been badly injured by a lion. He had been sleeping with 20 other herders inside the thorn fence used to protect the camels. A lion jumped the fence and seized the man rather than attacking the camels131.

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The widespread hunting of lions reduced their numbers and on later trips in the 1890s, Swayne said the hunting grounds of Somaliland “have lost some of their value now that so many lions have been shot and the elephants driven away”132. He added, though, that no one who had lived in the region could have sympathy for the lions because of their regular attacks on livestock and people. He evinced no recognition of the fact that the widespread hunting of wildlife will have been one factor in forcing lions to change their diet, nor does he show any regret over killing so many lions, which he said only survived in any numbers in the Haud and Ogaden regions on the border of Somaliland and Abyssinia133. This estimation of their surviving range is supported by the naturalist Drake-Brockman who wrote in 1910 that they used to be found all over Somaliland, but were now “seldom seen far distant from the Haud, Nogal valley and Ogaden … Lions are rare in British Somaliland”134. Cloudsely-Thompson in his book describing the depletion of game across East Africa quoted the politician and Kenyan settler Sir Alfred Pease as estimating that, over 12 years at the end of the 19th and beginning of the 20th centuries, over 1,000 lions were killed by European hunters in Somaliland. Pease himself saw 60 shot on hunting trips between December 1896 and March 1897135. Because of the heavily forested terrain of much of French Central and Equatorial Africa, there are few accounts of lions there. A few lived in more open areas on the border of present-day Gabon and the Republic of the Congo, while grassland areas of Cameroon, the Central African Republic (CAR) and northern Democratic Republic of the Congo (DRC) (around Garamba and the border with Sudan) had healthy populations of lions. Paul Belloni Du Chaillu explored much of French Equatorial Africa in the 1850s and 1860s, and said that lions only occurred to the north of the Congo Basin and then to the south of the rainforest region136.

West Africa The rise and fall of empires in the grassland, riverine areas and semi-desert regions of Senegal, Mali, Niger and northern Nigerian region of West Africa through the first 750 years of the second millennium CE were accompanied by growth in the population of the region, the rise of large settlements and development of cultivation and pastoralism in river valleys and areas with sufficient water for livestock. Wildlife suitable as prey for lions was limited to the areas between the dense forest and the Sahara. Wild ungulates competed for grazing and water with domestic stock and lions threatened the stock. Both were progressively depleted as human populations grew. The late 18th and early 19th centuries saw the rise of new polities, notably the Kanem and Borno empires of northern Nigeria, Niger and western Chad. These states had strong armies of horse-borne warriors armed both with traditional spears and swords but also growing numbers of firearms. One account of the rise of the Borno empire records that some areas were very thinly populated and

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were impenetrable or arid bushland, “the domain of the elephant and lion, with no human inhabitants except a few scattered herdsmen or cattle-breeders”. The Tuareg of the northern Sahel and Sahara, who were famous for their mobility and raiding of more sedentary communities, hunted lions to protect their livestock, doing so from horseback, with spears, until firearms became available. In northwest Nigeria, the Sokoto Sultanate established itself in 1804 and lasted as the major power in the region until British overlordship was established at the beginning of the 20th century137. A centralised Islamic polity, it again had strong, horse-borne armies and large herds of livestock. Protection of those herds by well-armed herders kept lion and other predator numbers low. The fragmented nature of the shrinking lion population of the region had its origins in both the habitat destruction and human settlement patterns that were well established by the advent of colonial rule.

Notes 1 Chapurukha M. Kusimba (2004), Foreword, in Bruce D. Patterson, The Lions of Tsavo. Exploring the Legacy of Africa’s Notorious Man-Eaters, New York, McGraw-Hill, p. xi. 2 Keith Somerville (2016), Ivory. Power and Poaching in Africa, London: Hurst, pp. 12–24. 3 Ibid., p. 345; see also Philip Curtin et al. (1995), African History. From Earliest Times to Independence, London: Longman, pp. 206 and 360; and, John Iliffe (2007), Africans. The History of a Continent, Cambridge: Cambridge University Press, p. 74 for West Africa, p. 187 for East Africa, p. 171 for Ethiopia, p. 127 for southern Africa. 4 Leonard Thompson (2000), A History of South Africa, New Haven, CT: Yale University Press, pp. 10–11. 5 Ibid., pp. 32–3. 6 www.fynboshub.co.za/fynbos-conservation/what-is-fynbos/ accessed 23 March 2018. 7 Iliffe, 2007, p. 127. 8 Noël Mostert (1992), Frontiers. The Epic of South Africa’s Creation and the Tragedy of the Xhosa People, London: Jonathan Cape, p. 123. 9 Ibid., pp. 55 and 59. 10 Ibid., p. 64. 11 Mostert, 1992, p. 141. 12 C.A.W. Guggisberg (1961), Simba. The Life of the Lion, London: Bailey Bros and Swinfen, p. 76. 13 Ibid., pp. 282 and 297. 14 Mostert, 1992, p. 153. 15 Thompson, 2000, p. 35. 16 Ibid., p. 45. 17 Guggisberg, 1961, pp. 38–9. 18 Thompson, 2000, pp. 47–8. 19 Mostert, 1992, p. 226. 20 Guggisberg, 1961, pp. 38–9. 21 Christopher Ehret (2016), The Civilisations of Africa. A History to 1800, ­Charlottesville: University of Virginia Press, p. 423. 22 Sir John Barrow (1802), An Account of Travels Into the Interior of Southern Africa in the Years 1799 and 1798, London: G.F. Hopkins, Kindle edition, loc 154–60. 23 Ibid., loc 1461–5. 24 Ibid., loc 1546.

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25 Ibid., loc 4614. 26 William John Burchell (1822, reprinted 2016), Travels in the Interior of Southern Africa, Vol. I, London: Longman, Rees, Orme, and Brown, pp. 243 and 289. 27 Ibid., p. 450. 28 William John Burchell (2015, originally 1824), Travels in the Interior of South Africa, Vol. II, London: Longman, Hurst, Rees, Orme, Brown and Green, reprinted by Cambridge University Press, p. 191. 29 Sir William Cornwallis Harris (1852), The Wild Sports of Southern Africa, Kindle edition (originally published by H.G. Bohn), loc 311. 30 Ibid., loc 987. 31 Ibid., p. 1739. 32 Ibid., loc 1852. 33 Fred Morton and Robert Hitchcock (2014), Tswana hunting: continuities and changes in the Transvaal and Kalahari after 1600, South African Historical Journal, 66, 3: 418–439, pp. 419–20. 34 Ibid., p. 420. 35 Morton and Hitchcock, 2014, p. 423. 36 Nancy Jacobs (2002), The colonial ecological revolution in South Africa: the case of Kuruman, in Stephen Dovers, Ruth Edgecombe and Bill Guest (Eds), South Africa’s Environmental History. Cases & Comparisons, Athens: Ohio University Press, 19–33, p. 21. 37 J. Chapman (1971 from 1868 original), Travels in the Interior of South Africa 1849– 1863. Hunting and Trading Journeys from Natal to Walvis Bay, and Visits to Lake Ngami and Victoria Falls, Vol. 1. Cape Town: A.A. Balkema, p. 71. 38 Morton and Hitchcock, 2014, p. 433. 39 Roualeyn Gordon-Cumming (2011), Five Years’ Adventures in the Far Interior of South Africa. With Notices of the Native Tribes and Savage Animals, Milton Keynes: Lightning Source, p. 15. 40 Ibid., p. 16. 41 Ibid., pp. 18–19. 42 Roualeyn Gordon-Cumming (2015), The Lion Hunter, in the Days when All of South Africa Was Virgin Hunting Field, Larnaca Press. Kindle edition, loc 50. 43 Gordon-Cumming, 2015, loc 228. 44 Ibid., loc 273. 45 Gordon-Cumming, 2011, p. 43. 46 Joanna Lewis (2018), Empire of Sentiment. The Death of Livingstone and Myth of ­Victorian Imperialism, Cambridge: Cambridge University Press, pp. 4–5. 47 Richard Fitter and Sir Peter Scott (1978), The Penitent Butchers. 75 years of Wildlife Conservation, London Collins, pp. 7–9. 48 William Beinart and Lotte Hughes (2004), Environment and Empire, Oxford: Oxford University Press, p. 70. 49 Ibid. 50 Ibid., pp. 71–2. 51 John M. Mackenzie (1988), The Empire of Nature. Hunting Conservation and British Imperialism, Manchester: University of Manchester Press, p. 29. 52 Ibid., p. 61. 53 E.A. Ritter (1978), Shaka Zulu, London: Penguin, pp. 232–3. 54 Mackenzie, 1988, p. 86. 55 Guggisberg, 1961, pp. 38–9. 56 John Lambert (2002), ‘The titihoya does not cry here any more’: the crisis in the homestead economy in colonial Natal, in Dovers, Edgecombe and Guest, 48–60, pp. 49–50. 57 Beverley Ellis (2002), White settler impact on the environment of Durban, 1845– 1870, in Dovers, Edgecombe and Guest, 4–47, p. 43.

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58 J.L. Cloudsely-Thompson (1967), Animal Twilight: Man and Game in Eastern Africa, London: G.T. Foulis, p. 55. 59 Mackenzie, 1988, p. 63. 60 Ibid., pp. 64–7. 61 R. Kent Rasmussen (1978), Migrant Kingdom, London: Collings; and, Thompson, 2000, p. 90. 62 Curtin et al., 1995, p. 245. 63 Somerville, 2016, pp. 50–1. 64 Elna Kotze (2002), Wakkerstroom: grasslands, fire and war – past perspectives and present issues, in Dovers, Edgecombe and Guest, 174–190, pp. 182–3. 65 Guggisberg, 1961, p. 39. 66 Andre F. Boshoff and Graham I.H. Kerley (2015), Lost herds of the Highveld: ­evidence from the written historical record, African Journal of Wildlife Research, 45, 3: 287–300, p. 187. 67 Ibid., pp. 288–9 and 296. 68 Norman Etherington (2016), A Biography of Frederick Courteney Selous. Big Game Hunter, Marlborough, Wilts.: Robert Hale, p. 31. 69 Dr A Gunther FZS (1884), Note on a supposed Melanotic variety of leopard from South Africa, Proceedings of the Zoological Society of London (hereafter referred to as Proceedings), 3 March, pp. 243–247. 70 Frederick Courtney Selous (2001, Originally published in 1881), A Hunter’s Wanderings in Africa, Alexander, NC: Alexander Books, p. 14. 71 Ibid., p. 42. 72 Ibid., pp. 228–9. 73 Ibid., p. 249. 74 Ibid., pp. 259–60. 75 Selous, 2001, pp. 276–80. 76 Ibid., pp. 128–33. 77 Ibid., pp. 182–3. 78 James Stevenson-Hamilton (1937), South African Eden, London: Penguin Modern Classics, Kindle edition, loc 4014–17. 79 F.C. Selous (1894), The Lion in South Africa, reprinted from Clive Phillipps-­Wolley, Big Game Shooting, Volume 1, London: Longmans, Kindle version, loc 109; and Etherington, 2016, p. 141. 80 Somerville, 2016, pp. 36–7. 81 F. Vaughan Kirby (1899), Sport in East Central Africa. Being an Account of Hunting Trips in Portuguese and Other Districts of East and Central Africa, London: Rowland Ward, p. 118. 82 Ibid., p. 311. 83 Keith L. Chiazzari (Ed.) (2012), Bwana Mkubwa: Big Game Hunting and Trading in Central Africa 1894 to 1904 (Compiled from Robert Wright’s contributions to Country Life magazine, Copyright ©Keith Chiazzari 2012), Kindle edition, loc 309. 84 Ibid., loc 750. 85 Garth Owen-Smith (2010), An Arid Eden. A Personal Account of Conservation in the Kaokoveld, Jeppestown, SA: Jonathan Ball, p. 77. 86 Ibid., p. 58. 87 Charles John Andersson (1856, reprinted 2012), Lake Ngami, Memphis, TN: General Books, p. 9. 88 Ibid., p. 13. 89 Ibid., p. 19. 90 Ibid., p. 24. 91 Ibid., p. 32. 92 Ibid., p. 102. 93 James Suzman (2017), Affluence without Abundance. The disappearing world of the Bushmen, New York: Bloomsbury, pp. 195–6.

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94 Joachim John Monteiro (1875), Angola and the River Congo, Volume II, London: ­Macmillan, p. 79. 95 Ibid., p. 116. 96 Ibid., p. 161. 97 Ibid., p. 209. 98 Brian J. Huntley (2017), Wildlife at War in Angola. The Rise and Fall of an African Eden, Pretoria: Pretoria Book House, p. 136. 99 George B. Schaller (1972), The Serengeti Lion. A Study of Predator-Prey Relations, ­Chicago: University of Chicago Press, p. 7. 100 L. Hazzah et al. (2017), From attitudes to actions: predictorsof lion killing by Maasai warriors, PLoS ONE, 12, 1. 101 Ibid. 102 Ibid. 103 Jan Bender Shetler (2003), Interpreting rupture in oral memory: the regional context for changes in Western Serengeti age organization (1850–1895), Journal of ­African History, 44, 3: 385–412, p. 404. 104 Curtin et al., 1995, pp. 90–3. 105 Ibid., p. 393. 106 Sir Richard Burton (2001, originally published 1860), The Lake Regions of Central Africa. From Zanzibar to Lake Tanganyika. Volume 1, Santa Barbara, CA: Narrative Press, pp. 61 and 64. 107 Sir Richard Burton (2001, originally published 1860), The Lake Regions of Central Africa. From Zanzibar to Lake Tanganyika. Volume II, Santa Barbara, CA: Narrative Press, p. 12. 108 John Hanning Speke (1864), What Led to the Discovery of the Source of the Nile, originally published by William Blackwood, London: Public Domain Book, Kindle version, loc 1705. 109 Ibid., loc 3222–7. 110 H. Johnston (1885), General observations on the fauna of Kilimanjaro, Proceedings, 3 March, p. 216. 111 Robin S. Reid (2012), Savannas of Our Birth. People, Wildlife and Change in East ­Africa, Berkeley: University of California Press, p. 105. 112 Jan Bender Shetler (2007), Imagining Serengeti. A History of Landscape Memory in ­Tanzania from Earliest Times to the Present, Athens: Ohio University Press, p. 17. 113 Curtin et al., 1995, pp. 302–5. 114 Captain H.C. Brocklehurst (1931), Game Animals of the Sudan, London: Gurney and Jackson, cited by Cloudsely-Thompson, 1967, p. 59. 115 Ibid. 116 Ibid., p. 60. 117 Annik E. Schnitzler (2011), Past and present distribution of the North African– Asian lion subgroup: a review, Mammal Review, 41, 3: 220–243, p. 231. 118 Ibid. 119 Guggisberg, 1961, pp. 30–1; and Schnitzler, 2011, pp. 231–7. 120 L. Sclater (1868), Notes on Baker’s Antelope (Hippotrugus bakeri), Proceedings, 26 March, p. 215. 121 W.E. de Winton (1898), On the Mammals obtained in Southern Abyssinia by Lord Lovat during an Expedition from Berhera to the Blue Nile, Proceedings, January 1900, p. 83. 122 See Richard Pankhurst (1964), Wildlife and forests, Ethiopia Observer, 7: 240–255. 123 Siegbert Uhlig (Ed.) (2007), Encyclopaedia Aethiopica, Volume 3, Wiesbaden: ­Harrassowitz Verlag, p. 572. 124 John Hanning Speke (1864), What Led to the Discovery of the Source of the Nile, originally published by William Blackwood, London: Public Domain Book, Kindle version, loc 159. 125 Ibid., loc 566.

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126 Elspeth Huxley (1935), White Man’s Country. Lord Delamere and the Making of Kenya, Volume One: 1870–1914, London: Chatto and Windus, pp. 13–14. 127 Ibid., p. 20. 128 Harald George Carlos Swayne (1895), Seventeen Trips Through Somaliland: A Record of Exploration & Big Game Shooting, 1885 to 1893: Being the Narrative of Several Journeys in the Hinterland of the Somali Coast Protectorate, London: Rowland Ward. Kindle edition, loc 104–9. 129 Ibid., loc 828. 130 Ibid., loc 3745. 131 Ibid., loc 4359–65. 132 Ibid., loc 4860–2. 133 Ibid., loc 4884. 134 Ralph Evelyn Drake-Brockman (1910), The Mammals of Somaliland, London: Hurst and Blackett, p. 11. 135 Cloudsely-Thompson, 1967, p. 70. 136 Paul Belloni Du Chaillu (1868), Explorations and Adventures in Equatorial Africa, with accounts of the manners and customs of the people, and of the chase of the gorilla, the crocodile, leopard, elephant, hippopotamus, originally published London: Harper, Kindle edition, loc 5102. 137 Stephanie Zehnle (2016), War and wilderness – the Sokoto Jihad and its animal discourse, Critical African Studies, 8, 2: 216–237, p. 217.

4 Hunting, conservation and the decline of the lion in colonial Africa and Asia

Colonial occupation magnified the consequences for wildlife of European penetration of Africa. The desire of colonising countries to make their African possessions pay, through the development of settlement and cash crop agriculture, reduced habitat available to wildlife. Ungulates were seen as competitors with livestock or sources of disease, and were exterminated and excluded from areas of cultivation. They were also used as a source of meat and hides to consume or sell, tiding settlers over until their crops could be marketed. Lions were treated both as vermin and a quarry for sports hunters. The combination of wildlife habitat loss and depletion, and the very direct persecution of lions had a significant and lasting effect. This was mirrored in India, where lion ranges across northern and western areas were reduced to a single, small population.

Africa European occupation went into full swing after the Berlin Conference of 1884–5 divided the continent between European powers1. Under colonial rule, African communities in many territories were banned from or restricted in their hunting. The occupiers worked to profit from their new territories through exploitation of land and natural resources. Elspeth Huxley, the daughter of settlers in Kenya, wrote that from the start there was an intention to end the tribal system of communal land ownership by consolidating fragmented holdings and registering their titles in the names of individual owners; and to complete the momentous change-over from a primitive system of ‘shifting cultivation’ and nomadic cattle-herding, to the intensive production of permanent crops, and the controlled care of livestock 2.

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This had immense repercussions for indigenous peoples and for the environment, wildlife and the relationship of people to the wildlife, hardening the boundaries between them. Settlement brought a direct onslaught against wildlife as settlers, colonial officials, soldiers and traders used the opportunities in colonies to hunt on a massive scale for food, to clear land, for commercial gain or sport. Settlers in Kenya hunted for meat, ivory and skins, and recreation. Those like Lord Delamere, to quote his biographer Huxley, killed prodigious numbers of lions, elephants, rhino and other game in Kenya, Somaliland and Abyssinia 3. Sport hunting reduced lion numbers in many areas, but the designation of lions as vermin had the greatest effect. This ensured a high toll of lions wherever they were a threat to livestock. Hunting resulted in a massive rate of killing, with evidence that in areas like the Serengeti in the late 19th and early 20th centuries hundreds of lions might be killed by hunters on a single safari4. This was magnified during the establishment of reserves or national parks by the killing of predators to preserve large populations of ungulates, a practice that continued up to the 1950s5. Hunting was viewed as ideal training for those running the imperial project. Mackenzie sums it up: “The importance of the Hunt can be identified at every level of the theory and practice of the imperial ethos”6, and hunting in Africa and India was an important aspect of “popular imperialism” in Britain and helped romaticise it. Hunting that regal symbol of African nature, the lion, was a powerful symbol of dominance not just over nature but over territories and peoples colonised7. As the anthropologist Chapurukha Kasimba wrote, “European fascination with Africa can be traced back to the Victorian period when the myth of a wild Africa teeming with riches and savagery was firmly imprinted”8 and mastery over this “savagery”, perfectly represented by the lion, was a sign of imperial might. The narratives and practices of hunting and wildlife management, as Laurence Frank told me9, came straight from European farming and hunting traditions. European hunting was presented as noble sportsmanship, African hunting as cruel and wasteful, using methods labelled abhorrent – snares, traps and poisoned arrows10. Europeans, amid their widespread hunting, depicted themslves as guardians of wildlife11. Even when cattle diseases depleted herds and caused starvation across East and Southern Africa, Europeans viewed African subsistence hunting for bushmeat as destructive rather than necessary for survival12. Europeans wiped out thousands of zebra, buffalo and other grazers to clear grazing for cattle, seeing it as progress. Keen hunter and arch-colonialist Lord Lugard later admitted, “We were butchers not sportsmen”13. The butchery, whether by sportsmen, settlers or those Africans encouraged to hunt for meat, hides and ivory by Europeans, was enabled on a huge scale by the availability of modern firearms. In 1889, Europeans imported something as damaging as firearms – rinderpest, the cattle disease which wiped out millions of cattle and wild ungulates. The disease came from Italian imports of cattle from India to the Eritrean port of Massawa14. The effects of the disease had a roller-coaster effect on lions. At first, lions multiplied rapidly, feasting on wildlife

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and livestock killed by the epidemic and those of people who died of starvation. When the epidemic was over, prey availability was reduced significantly, leading to livestock killing and even man-eating. Pastoralist communities across East and Southern Africa reverted to hunting to survive, reducing the lions’ prey and increasing human-lion conflict15. Soon after the end of the epidemic, wildlife numbers recovered but European hunting increased with the expansion of settlement in East and Southern Africa. This led to a realisation on the part of some hunters or administrators that hunting, along with rinderpest, had “produced such a marked diminution of game that conservation measures seemed necessary”16. This wasn’t an altruistic measure to preserve wildlife, it was fear that the decline deprived settlers of income and sport. The form “preservation took was shaped by the social and economic realities of empire … Access to animals was to be progressively restricted to the elite; animals were to be categorised … some were to be specially protected for their rarity, others shot indiscriminately as vermin”17. Lions were catergorised as vermin and efforts to conserve other species justified killing them. Much of the impetus for the creation of reserves, and later national parks, in British territories came from the formation in Britain of the Society for the Preservation of the Wild Fauna of the Empire in 1903 by a group of hunters and colonial administrators. This followed the meeting in London of colonial powers which drew up the the Convention for the Preservation of Wild Animals, Birds and Fish in Africa, which was signed in London on 19 May 190018. It called for concerted efforts to conserve endangered species and habitats19; schedule five of the convention called for the reduction in number of lions and other predators to preserve ungulate species in decline since the arrival of Europeans. At the heart of its ethos was that only Europeans could be trusted to manage wildlife. What was totally ignored, for reasons of self-interest and imperial myth-­ making, was that the inhabitants of the territories had coexisted – albeit with exploitation for food, hides and killing for livestock protection or ritual ­purposes – with wildlife without depletion on a catastrophic scale20. This history and culture of coexistence and moderate utilisation was replaced by a European one of selfish consumption and the linked policy of conservation to perpetuate utilisation or appreciation by Europeans. Central to it was the end or limiting of African control over wildlife and the exclusion of Africans from hunting or even living in certain areas. As Reid argues, “Since the late 1800s, this ancient coexistence of people and wildlife in African savannas has often been ignored, often replaced by a modern practice of conservation which assumes that wildlife are best conserved in landscapes with no people.” Colonial governments carved out large areas to be reserves with limited or no hunting; later making them national parks from which people and livestock were excluded 21. Some colonisers had flashes of insight into the effects of European penetration of Africa on wildlife. The hunter and settler, Robert Foran, admitted that The relentless advance of civilization into the unspoiled wastes of the earth has driven the wild fauna further and further from their age-old habitat …

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rare species of animal life have been wiped out completely; others have been brought to the brink of extinction. Modern deadly weapons, coupled with improved means of transport and easier communications, have all played their parts in bringing about this ruthless extermination of the fauna 22. Another hunter and colonial official, Charles Hobley, an advocate of conservation and of restricting African hunting, admitted that the slaughter of thousands of lions in sanctuaries established by colonial administrations in Africa created massive imbalances between predators and herbivores23. Loveridge emphasises that protected areas were far from protective towards lions and that since the early 20th century they’ve been exterminated in over 80% of their range and in a major protected area, Kruger NP, 3,031 lions were shot, trapped or poisoned between 1902 and 196924. The effects of colonisation of Africa and its effects on lion will be examined region by region, followed by the recent history of the surviving Asiatic lions in the Gir Forest in India.

East Africa – settlement, big-game hunting and conservation Hunting was ubiquitous before colonial occupation. As Steinhart says, “It would not be far wrong to answer the question of who hunted in precolonial Kenya with one word: everyone”25. The arrival of British administrators and settlers changed this. Hunting was criminalised, unless you were white. Many communities ignored colonial laws and contined to hunt, hidden from the gaze of colonial officials. But professional white hunters, settlers and visiting hunters killed huge numbers of lions, elephants and game. It is no exaggeration to say that. “virtually all the Europeans who came to Kenya in their capacities as explorers, missionaries, administrators, soldiers or settlers also hunted regularly and sometimes prodigiously”26. Hunting was a source of income and a subsidy (through ivory, skin and hide exports, and meat for the table) for the establishment of colonial rule. Exploitation was largely unrestrained by regulation and led to prodigious killing of lions and game. It escalated in the early 20th century with the extermination of game to make way for cash crops or livestock 27. Through biographical, narrative and fictional accounts of settlement and hunting in East Africa, Kenya was presented as a colonial paradise with teeming wildlife and fortunes to be made for hard-working settlers, who could then spend their spare time hunting game. Books such as Karen Blixen’s Out of Africa, Elspeth Huxley’s The Flame Trees of Thika, Beryl Markham’s West with the Night and accounts of big-game hunting, created a popular mythology about colonial Kenya and a romanticised view of hunting. Just before and after the First World War, the commercial exploitation of big-game by hunting operators became lucrative. Safaris became popular among wealthy Europeans and Americans. Professional hunters who had shot elephants

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and other game for profit, now guided rich clients in pursuit of trophy animals, especially. This gave rise to the image of the great white hunter, Bror Blixen-­ Finecke and Denys Finch Hatton being prime examples (familiar from their representation in the film of Blixen’s Out of Africa). Lions were shot in unlimited numbers. Until 1926, they were labelled vermin with no need for licences or limits to off-take. The thrill of a lion hunt, for those inclinded to enjoy it, was heightened by the image of the lion as powerful, majestic and highly dangerous. The latter quality being emphasised by the attacks by man-eating lions on labourers building the railway from Mombasa to the Uganda Protectorate. The story of the Man-Eaters of Tsavo is one that may shed light not just on why those lions predated on humans, but also on why lions kill people. The main source of information on the man-eaters is Lt-Col. J.H. Patterson, who was charged with overseeing work on the line near what is now Tsavo NP. He managed the workforce that was being killed by the lions and he eventually shot the the man-eaters. His account, supplemented by contemporaneous reports by officials in Kenya and studies since, indicates that two maneless male lions were reponsible for repeated attacks on labourers over nine months in 1898. At one stage the workforce was so petrified that work stopped for three months. When attacks started in 1898, the lions weren’t always successful in taking their victims. But experience soon enabled them to take a regular toll of workers, raiding camps at night. As more men were killed, Patterson tightened security. Attempts by him and others to shoot the lions were unsuccessful and, as Patterson observed, many of the Indian labourers began to believe that they weren’t ordinary lions but “the angry spirits of two departed native chiefs” who wanted to stop the building of the railway28. Patterson would sit up at night watching where attacks had taken place only to hear the lions raiding camps nearby to take more victims29. After one attempt to shoot the lions at night, there was a gap with no attacks, but then they resumed. A lion broke into a camp and workers threw firebrands and rocks at it, but it still killed a man and carried him 30 yards before stopping to eat part of the body. The next night, a camp two miles away was raided by the lions and a man eaten within sight of the camp30. When the colonial District Officer, Whitehead, came with armed askaris to help Patterson kill the lions, he was attacked and one of his men killed and eaten31. On 9 December 1898, Patterson succeeded in wounding one of the animals32. Soon after, he sat up over the carcass of a donkey killed by the lions and killed one of them, described as a huge male33. The attacks continued until Patterson was able to kill the second lion on 29 December; 135 workers had been killed by the lions during the attacks. One theory about why the lions started attacking the workers’ camps, put forward by a historian of the railway, Ronald Hardy, was that large numbers of domestic stock were butchered to feed the workers, while European overseers and engineers hunted game to provide meat. The fresh carcasses in or around the camps, attracted predators. The regular hunting concentrated around camps would also have thinned out prey in the area 34. In addition, the railway followed the route of historic slave caravans from the Indian Ocean coast to the

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interior. The bodies of slaves who died or were killed along the route had been left in the bush. There was also a high rate of death from injury or disease among the Indian workers. Over time, this may have accustomed resident lions to feed on human bodies35. Bruce Patterson in his study of the man-eaters noted that there were numerous accounts of man-eating in the area preceding the building of the railway, particularly around villages with permanent water supplies, which were crucial for people, livestock and wildlife during the 1887–8 drought 36. That lions in the area would have fed on corpses over a long period of time, or even killed those weakened by famine, could explain the seeming specialisation of the lions in the area in hunting people. Lion specialist Brian Bertram suggests that while man-eating is relatively rare among lions, “It seems once a lion has for some reason killed and fed on humans, and so knows that they are harmless, easy prey and good edible meat, he is liable to continue to do so.”37 This could explain why healthy lions in certain areas may take to regular predation on humans, rather than man-eaters being only old, sick or injured lions killing out of dire need 38. But, in most cases, as Patterson details, old age and chronic infirmity were causes of man-eating, including injuries sustained hunting dangerous prey, porcupine quills embedded deeply in the flesh or wounds caused by snares39. He points out that the two Tsavo man-eaters and a man-eating lion that killed many people in Mfuwe (Zambia) in the 1970s had broken teeth and jaw damage, which made it harder for them to kill wild prey40. At the turn of the century, the high commissioner for the East African colony, Sir Charles Eliot, charted a new phase in economic development which had a major impact on wildlife. Eliot was committed to settlement as an economic necessity to prevent the colony being a drain on the British Treasury. He believed that only large-scale European food and cash crop cultivation would enable the colony to pay its way41. Occupation and fencing of land by settlers resulted, with the extensive slaughter of herbivores and predators, and the creation of “tribal” reserves to which pastoralist and other local communities were consigned. Those encouraged by the British government to settle in Kenya could apply for land. The settler and hunter, Lord Delamere, obtained large areas of land to raise imported cattle and sheep42. To protect them, he became a determined exterminator of lions, and killed zebra and other ungulates he saw as competition for his livestock43. When he established a sheep farm near Lake Elmenteita, he brought in another hunter, Paul Rainey, to help him get rid of the lions – they killed 12 in three months and fenced off the farm to prevent wildlife getting in44. At the start of the 20th century, despite inroads made by European hunting, lions were found across Kenya. Daphne Sheldrick, who lived in Tsavo and then near the Nairobi wildlife sanctuary bearing her name, wrote that when her grandparents settled in Maasailand in 1907, game was pentiful and lions were to be found in prides of 20 or more, but “slowly those great herds began to dwindle, eroded by the impact of civilization, and with each year that passed, the numbers grew fewer, until people suddenly wondered in astonishment where all the animals had gone!” Game was treated as “a source of free meat and free sport

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for anyone who could hold a gun”45. In 1908, 150 lions were shot on licence in Laikipia alone; the scale of killing across the colony must have been huge46. As Frank et al. wrote, “an example of the zeal with which lions were shot, safaris to the Serengeti area in the early part of the last century sometimes shot over 100 lions, clients of just one safari company killed 700–800 lions in 1911”47. Frank, Director of Living with Lions and prime mover of the Laikipia Predator Project, told me that the scale of slaughter was not exclusive to the early days of settlement, and between 1946 and 1952, one Laikipia game warden shot 434 lions “on control” and some individuals killed over 300 lions apiece to protect cattle ranches in Kenya in the 1970s and 1980s48. Much of this shooting, according to Frank, was part of an attempt “to open a huge region of NE Kenya to modern cattle ranching in a virgin area with abundant predators… they were not simply maintaining low numbers on long-established ranches, but attempting to reduce a dense population to allow free-range grazing of cattle across something like a million acres”49. In 1902, a government ordinance created two game reserves – the Northern (from the Ugandan border to Marsabit, south of the Guaso Nyiro River and to Lake Baringo) and the Southern (from the Uganda Railway in the north to the border with German-ruled Tanganyika, and from the Tsavo River in the east to Uganda in the west)50 – in which hunting was supposedly regulated. High Commissioner Eliot admitted that settlers ignored hunting restrictions and felt the laws “may be violated without loss of moral character when they can be violated with impunity”51. The reserves were unfenced and wildlife moved in and out, often venturing into populated areas in search of grazing and water during dry periods. The colonial official, Charles Hobley, wrote to the Zoological Society of London that during a drought in 1910, zebra and hartebeest were seen in large numbers in Nairobi and on the surrounding plains as they left the Southern Reserve in search of grazing. They were followed by lions. Hobley said lions killed prey right in the centre of Nairobi. He believed that as long as the reserves were maintained and plains game survived there, lions were in no danger of extinction in Kenya52. Hobley noted that during the First World War, when sports hunting declined as settlers joined army units formed to fight the Germans in Tanganyika, lions increased rapidly in numbers and, in some areas, became livestock killers as there was extensive hunting of game during the war to supply meat for the troops, depleting natural prey53. After the war, the Game Department, under the leadership of Blayney Percival, started shooting lions in the reserve to reduce numbers. In the early decades of the 20th century, increased settlement led to the removal of the original inhabitants from areas earmarked for European farming. The Maasai, weakened as a community by the effects of rinderpest and smallpox54, were pushed into signing an agreement to move from the Maasai reserve around Laikipia into the Narok area. The areas vacated were given to European farmers for livestock raising55. They were intolerant of lions and had guns with which to eradicate them. The fencing of land, killing of herbivores and treatment

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of lions as vermin, progressively reduced their numbers. The relocation of the Maasai and their herds restricted their freedom of movement and concentrated people and cattle into smaller areas. Concentration of pastoralists around Narok and the Southern Reserve created conditions for enhanced conflict with the large herds of plains game and resident lions through competition for grazing and predation of stock. The resident and nomadic lions in these areas threatened Maasai cattle and tempted young warriors keen to prove their manhood at a time when British occupation had more or less ended cattle raiding as a way young men could demonstrate courage. As Hazzah et al. pointed out, killing a lion has a great traditional significance in Maasai society, notably among the warrior group of young men from 15 to 30 years old, though this did not mean large numbers of lions were killed or that they sought to exterminate lions56. Guggisberg emphasised that the relatively low off-take from Maasai, Nandi and other indigenous hunters engaging in coming-of-age rituals or killing livestock predators, paled into insignificance compared with the effects of commercial farming and hunting for profit and sport by settlers and visiting hunters57. He gave examples to illustrate his point: two cousins, Harold and Clifford Hill, farmed just outside Nairobi. They were credited with killing 300 lions in the early years of settlement. In the tsetse-free Kenyan highlands, hunters would hunt from horseback or lions would be driven by scores of beaters towards waiting guns and, at night, hunters sat in a machan to shoot lions over baits. Some farmers simply poisoned lions, including the settlement of the White Fathers missionaries on the shores of Lake Tanganyika, who are recorded as having killed 37 lions using poisoned antelope carcasses – for no obvious reason, as the lions were not said to be either cattle thieves or man-eaters58. Blayney Percival, the first head of the British East Africa Game Department, personally killed 50 male lions59. Extensive hunting reduced lion numbers across Kenya. From being ubiquitous in suitable habitats (most of Kenya excepting thick or high montane forest), they became less common and wary of man. The hunter Abel Chapman recorded in 1908 that, while he believed lions were still numerous in many areas of East Africa, during a three-month-long expedition in 1904–5, he only saw lions once. On a safari in December 1905, he noted that after several weeks’ searching around the Athi Plains (once teeming with lions) he had failed to find a single one60. Another settler and hunter, W.S. Rainsford, said attacks by lions were common in the Ol Donyo Sabuk area north-east of Nairobi. He said that 20 white men had been killed or badly mauled by lions there in recent years along with numerous local people (who it was presumably not worth counting)61. Most of the Europeans received their injuries following up wounded lions62. Rainsford believed lions to be common along the railway, around Voi and in the Rift Valley from the Mau Escarpment to Nakuru and Naivasha63. Rainsford also believed that as game was depleted lions would become “more destructive to domestic beasts and to man”, adding that “the lion that jumps a boma to stampede a herd is on the road to be a manslayer”64 In this comment he

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unwittingly identified one of the key reasons for increasing human-lion conflict in colonial and post-colonial Africa – the disappearance of the lions’ natural prey as a result of the actions of man. The accounts of hunters and settlers of this period leave little doubt about the extent of killing of game for economic reasons and for sport65. Richard Tjader estimated the number of lions killed in a 15-month period before 1910 on the Sorik and Loita Plains, south-west of Nairobi, at 160, with one man killing 21 in two weeks66. Glorifying these hunters in his writings, Tjader admitted that by 1910, “the white man with his modern firearms has almost exterminated the big cat”67. The former American president Theodore Roosevelt and his son killed 512 mammals on their safari in 1909, including 17 lions68. Prolific hunter and Lieutenant-Governor of the East African Protectorate, Sir Frederick Jackson, denounced the Roosevelt expedition for exceeding “all reasonable limits of hunting”69. But Jackson took no measures to reduce hunting, especially of “vermin” like lions. Another American hunter, Paul Rainey, shot 74 lions, on one occasion killing nine in 35 minutes and 27 in 35 days on a hunting safari70. The official record of game shot in Kenya in 1911–12 lists 76 male lions and 43 lionesses, adding that lions, not being included on game licences, were shot in greater numbers than shown in official figures. The report goes on to say that recorded killings totalled 206 for the 1910–12 period but evidence suggests a further 648 lions were killed by shooting, trapping or poisoning by settlers that don’t appear in official statistics71. The settler and hunter Sir Alfred Pease noted that in this period an estimated 60,000 game animals were shot in Kenya by hunters, not counting those shot by settlers on their own farms72. The toll from hunting and from the rinderpest epidemic in neighbouring Tanganyika was somewhat reversed after the First World War, when the British, who took over the territory, expanded game reserves and wildlife repopulated areas in northern Tanzania denuded of cattle by disease outbreaks73. The recovery of wildlife in northern Tanganyika led to it becoming a popular safari hunting destination in the 1920s. Two professional hunters, Simpson and White, led one group in 1925 which killed 50 lions. Other hunting parties followed, some accounting for as many as 100 lions in a single trip. There was no regulation by the Tanganyika Game Department, which allowed hunters to engage in what were considered unethical practices, such as shooting from vehicles and excessive killing74. In 1928, a motorised safari in the Serengeti was credited with killing 60–65 lions75. The notoriety of these motorised safaris led the Kenya-based professional hunter Denys Finch Hatton to call for an end to the excessive killing of lions in Serengeti76. He wrote an article for the London Times extolling the virtues of photographic safaris and deploring the “orgy of slaughter” in Tanganyika. He proposed the creation of closed districts and demanded that the administration in Tanganyika limit motorised safaris and stop indiscriminate shooting of lions, referring to butchery by “tourist hunters” 77. He gained the support of the Prince of Wales, who he had guided on a safari, and that of Lord Onslow of the Fauna Preservation Society78. Under pressure from such well-known figures,

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the government sent the scientist Julian Huxley to examine the situation in northern Tanganyika. The result was the declaration of a 900-square mile closed reserve in the Serengeti area from Banagi Hill westwards between the Grumeti and Mbalagati Rivers and protection for lions in surrounding areas79. The 1920s–1930s was the era of the big-game hunters, with men like Arthur Hoey, R.J. Cunninghame, Bill Judd and Leslie Tarleton succeeding the generation of Blixen and Finch Hatton in leading safaris, on which lions were at the top of the menu for visiting hunters. They would track lions for their clients or ride them down on horseback80. One, who wrote copiously about his “adventures”, was Sir Alfred Pease, who admitted to killing 14 lions in one day near his ostrich ranch at Kitanga in Kenya. Pease says in his defence that at the time lions were still plentiful and prides of up to 20 could be seen on the Kapiti plains south-west of Nairobi81. Kenya’s first Game Department warden, Blayney Percival, wrote in 1924 that lions were still to be found in most areas, but that they were beginning to inhabit thick bush rather than open plains as in the past82. On the overall lion population, Percival concluded: Reviewing the whole country within my knowledge, I think there are far more lions in the Southern Game Reserve than in any other district … they swarm there … protected from the sportsman’s rifle and with an unlimited supply of game, they live in peace on the fat of the land. Their only foes are the Masai. When a lion takes to cattle-killing these people rise in wrath and turn out to hunt him. Also a few are killed by the young Moran to blood their spears, but the numbers thus slain cannot make any appreciable difference. He said that in settled districts “the case is different; where stock-farming and ostrich farming are extensively carried on the lions have been steadily killed down.”83 In the 1920s, game ordinances still encouraged the killing of lions without a licence and allowed those with travellers’ licences to kill unlimited numbers of lions outside the reserves84. The writer Karen Blixen, who hunted regularly, said the availability of vehicles meant that residents of Nairobi, “shop-people” as she called them, would drive out into the country at weekends and shoot anything they could find, including lions85. When hunters like Finch Hatton and the renowned elephant hunter W.D.M. Bell began to express concern over excessive hunting of lions it was not purely altruistic but, as Bell stated, because if protection was given to them in Kenya and Tanganyika this “should ensure good sport for many years to come”86. By 1925, concern was expressed by the Game Department “that it will be necessary in the near future to limit the number of lion that may be killed on a Full Licence … otherwise this animal will soon cease to exist in some areas where he does no harm” 87. The following year, the chief game warden, Captain A.T.A. Ritchie, said in his annual game report that lions had been taken off the “unlimited list” and the number

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allowed to be shot on a licence reduced to four 88. In his report for 1930, Ritchie lamented that there had been over-hunting by motorised safaris in the Mara River area with the result that lions were “comparatively rare here”89, when they had been numerous. Other practices continued that reduced the chances of lions surviving in many areas. In the Narok region, north of the Mara, a professional hunter called Jack Bonham shot 8,000 zebra and 5,000 wildebeest to clear land for livestock90. The extermination of prey species led both to the disappearance of lions and increases in the taking of livestock by surviving lions deprived of their natural prey. George Adamson, of Born Free fame, worked for the Game Department at the beginning of the Second World War and was charged with killing thousands of zebra, oryx and other antelope in northern Kenya to preserve grazing for cattle. At first this increased lion numbers, as the carcasses were left where they fell91. But in time, the decimation of prey species increased lion predation on livestock. Adamson, who later developed an obsession with lions, regularly shot stock-killing lions and man-eaters. He also used poison extensively to try to kill stock raiders, particularly hyenas92. Kenyan Game Department records record that while a game warden, Adamson killed 50 lions93. At the end of the Second World War, conservation policies led to the creation of fully protected national parks, in which hunting was prohibited. Nairobi NP, which contained several prides of lions, and Tsavo NP with a very large lion population, were the first. During the campaign by conservationists to establish the parks, Mervyn Cowie (who became head of the National Parks Board and was director of Nairobi NP) tried to ensure that the Nairobi park was established by feeding lions there, so that when Kenya’s new governor visited the proposed area of the park he was able to see lions. A drawback was that some of the lions became accustomed to people, leading them to kill a game guard and pursue tourists visiting the park94. There were also periodic problems of lions moving out of the park into heavily populated areas, which usually resulted in them being shot or speared. Lions were the biggest draw for the increasing numbers of tourists who visited the park after it opened in 194895. Outside protected areas, there was a steady toll of lions resulting from conflict with people. The former game warden, Ian Parker, wrote that by the 1950s large areas of Maasailand in Kenya and Tanzania were devoid of predators because of widespread poisoning by the Maasai with a pesticide used to keep their cattle free of ticks96. The taste of the pesticide was liked by lions, who would lick it from old dip tanks and die, although there was no actual intent to kill in those cases. According to the professional hunter, Sydney Downey, farmers “always loathed lions, classing them as vermin. Hunters have always looked upon lions in terms of impulse, zest, and challenge, as a sort of high-spot”97. In unprotected farming or rangeland areas, lions were killed in large numbers by pastoralists and white farmers. By 1960, Downey feared that lion numbers “could drop below the point of recovery, making these animals virtually extinct” in both Kenya and Tanzania98.

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When the British were given the League of Nations mandate over Tanganyika after the First World War, they allowed African communities to hunt under a system of licences to preserve game while allowing stock protection. There was also a growing trophy hunting industry but not the settler presence that was the main factor in the ban on African hunting in Kenya. Hunting of wildlife was governed by Game Department ordinances99. Following the limited protection given to areas of the Serengeti and restrictions on lion hunting, in 1929 the Serengeti was declared a Complete Game Reserve, in which hunting was banned, much to the anger of the local Nata, Ikoma, Sukuma and Musoma communities, who had hunted there for centuries prior to colonial occupation100. An area of 2,286 sq km was protected in 1929, given full Protected Area status in 1940, covering southern Serengeti and Ngorongoro. In 1959, it was expanded to include the area between Banagi and the Kenyan border, while the Ngorongoro Conservation Area was removed from the park101. Within a few years of the ban on hunting in areas of the Serengeti in the late 1920s, lion numbers began to recover. In 1935, protection was extended through more stringent licensing of the hunting of lions in central Tanganyika, around Iringa, in the Northern, Lake and Tanga provinces – limits of six for a full visitor’s licence or resident’s licence or two for temporary licences102. Charles Hobley was quoted as saying in 1936 that in “a certain reserve in Tanganyika there had been a vast increase of wildebeest”, partly brought about, he believed, by the fact that during the last decade three or four thousand lions had been shot in that region. To restore the balance the killing of lions had been stopped in the area103. In southern Tanganyika, the problem was less the disappearance of lions than the persistent problem of man-eaters. In 1935, the game ranger for Lindi district reported that man-eaters were “exceedingly troublesome” and had recently killed 35–40 people. Over several years, 145 people had been killed by lions in the district. The ranger said that at one stage five man-eaters were hunting as a group; none of the lions shot appeared to be sick, thin or in bad condition104. One cause was thought to be the large number of wild pigs in the area, on which the lions preyed. This brought them into contact with villagers protecting their fields at night from the pigs. A total of 14 suspected man-eaters were shot in the region105. At Singida in central Tanganyika, there were outbreaks of man-eating in the 1920s and the 1940s. In the first period, around 200 people are thought to have been killed. Many local people claimed the lions were witchdoctors who could change themselves into lions, rather than real lions, a superstition that occurs regularly during outbreaks of man-eating in Tanzania. These beliefs were often inspired by unscrupulous local healers who would use lion attacks to extort money from local people, threatening to turn themselves into lions106. According to the elephant hunter turned game warden, George Rushby, this happened in the Iyayi area of south-west Tanganyika in 1932, when a local healer extorted money during an outbreak of man-eating, claiming he could stop the lions with magic but would only do so if paid. Some villages paid him cash or cattle to stop the lions attacking them107. Rushby was sent to the area and killed several of the

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lions and the rest were eventually exterminated by the Game Department. Between 1941 and 1946, Rushby said the man-eating lions around Njombe (close to the Kipengere Mpanga Game Reserve) were far more rapacious than those of Tsavo108. They killed possibly between 1,000 and 1,500 people in areas with very thick bush but limited wild game; Patterson is sceptical of Rushby’s accounts but says the number was very high even though only 246 were confirmed as victims109. In 1946–7, there was another outbreak of attacks by lions in the same region, with 103 people killed. Patterson told the author that it is certainly possible, given the recurrence of man-eating in the wider region of south-west Tanganyika, that lions there had adopted it as a part of their routine prey acquisition110. During the 1940s, reports of man-eating by lions in that area covered three sub-chiefdoms encompassing 1,500 square miles of very dense bush – in all the instances local communities became convinced that the lions were either witchdoctors or men dressed in lion skins. The predator specialist Hans Kruuk said that there were many folk tales in Tanzania of men turning into lions and being able to transform back into men, with these tales coming to the fore during outbreaks of man-eating111. In 1948, the Tanganyika administration passed an ordinance creating national parks. On 1 June 1951, Serengeti became a national park, with a total ban on sport and commercial hunting112 , though with a persistent problem of bushmeat hunting by local communities such as the Sukuma and Ikama, who had always used this to supplement farming113. The creation of the park and protection accorded wildlife in the adjacent Ngorongoro Conservation Area and Crater, reduced the killing of lions. But the first Serengeti warden, Myles Turner, was concerned that the wider region’s lion population was being reduced by the continuation of legal sport and commercial hunting on the Sabora Plains, where he said there was unlimited hunting of lions, with excessive killing by professional hunters and their clients in 1959 and 1960, wiping out most of the population there114. In the British Uganda colony, lions were plentiful in areas that became the Kidepo Valley, Murchison Falls and Queen Elizabeth NPs, with populations spread along the Nile and in suitable savanna or woodland habitats. Hunting was regulated but lions could be shot if they endangered livestock or people. Reports to the Zoological Society of London suggested that lions were even common in areas with little game, where they were a danger to people and their domestic stock, and noting that a number of people had been “carried off by lions” in some areas115. In Bukanga, north-east of Kampala, lions had a bad reputation among local people – some of whom had fallen victim to the animals – and were noted for “unusual and unprovoked truculence”116 Serious manifestations of man-eating were also reported in 1925 in Ankole, west of Lake Victoria. A report in the Fauna Preservation Society’s Journal mentioned that this had been a problem for generations, suggesting it was more than just predation by sick or old animals. But in the mid-1920s the problem had increased, with 12 people

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being killed by lions every month in 1924–5, most victims taken along the region’s roads as they travelled towards Lake Victoria to find work. Many of the travellers made themselves vulnerable by sleeping by the road at night with no shelter or protection117. The government reacted by trying to poison the lions, with little success, and the slower but more effective method of bringing in hunters to shoot the lions was the only way of stopping the predation118. Around the same time there were outbreaks of predation on people between Mubende and Entebbe in western Uganda. The attacks, thought by government officials to be by the same lions, spread to Sanga (80 miles away) and Masaka, with 33 people being killed in 1924119. Treves and Naughton-Treves examined the records of human-lion conflict in Uganda, finding regular references to attacks across the colonial period. They noted that agricultural settlements where attacks took place were scattered in what they called “a matrix of bush and forest, specially in Western Uganda” and were close to game reserves and, later, national parks120. Record-keeping was not systematic and there was little reference to the age or sex of victims. Between 1923 and 1994, records showed 275 lion attacks, 74.9% of the attacks resulted in deaths. Interestingly, nine of the reports of casualties from lion attacks said that the victims had been scavenging from lion kills, while others (no numbers given) occurred when the victims were out hunting121. During the same period 376 lions were recorded as being shot as problem animals by game wardens. As Uganda approached independence, it still had a largely well-distributed population of lions, particularly in national parks and reserves. Guggisberg believed there were significant populations in Murchison Falls and Queen Elizabeth NPs, which had been established in 1948, while outside the parks there were lions in Karamoja, northern Teso, Acholi, West Nile, Ankole and along the shores of Lake Albert. He said their range had been reduced steadily by human settlement, agricultural expansion and hunting since the late 19th century122 .

Southern Africa – conservation, lion control and recovery in depleted populations As the 19th century neared its end, the depletion of wildlife since European settlement spawned a feeling among some government officials and hunters that unregulated hunting and the killing of game herds to make way for farming would lead to the disappearance of wildlife across southern Africa. As early as 1884, President Paul Kruger of the South African Republic (Transvaal) proposed the establishment of a reserve with restrictions on hunting along the Sabi and Shingwedzi rivers. Kruger had been a keen elephant and lion hunter123. There had been precolonial ecological practices (including Khama’s and Lobengula’s attempts to limit European hunting in their territories), best described as customary modes of behaviour, but the development of “a formal conservation ethic” was a product of European concern about the extermination of species124.

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The moves to establish protected areas laid the foundations of a conservation system that included forced removal of African communities from their lands and the prohibition of traditional hunting125. Outside protected areas, white farming and the creation of reserves for communities like the Xhosa, Nguni and other peoples led to deforestation and the disappearance of game and the predators which preyed on it, particularly lions. Natal and Zululand are good examples of where settler policies led to serious declines in wildlife126. This was hastened by hunting for subsistence when rinderpest killed cattle and other stock. Lions had been wiped in Orange Free State and Transvaal by the end of the 19th century. Guggisberg says there was considerable surprise when a lion was reported to have been killed at Springs (50 km east of Johannesburg) in 1898 and another in Heidelberg (50 km south-east of Johannesburg) in 1899127. Practically all large wildlife species had been exterminated in the Orange Free State by the 1890s as farmers killed herbivores to reserve grazing for livestock and killed predators on sight128. The great herds of springbok, zebra, eland, black wildebeest and hartebeest were gone; bontebok and blesbok only survived on a few farms, whose owners saved them from extinction. The quagga and blaubok were long gone129. The lions which preyed on them and which were recorded as following the migrations had gone, too130. Within the territory that is present-day South Africa, lions survived in small numbers around Sabi, Komatipoort, Malelane and Kaapmuiden in Transvaal province. Some survived in northern Zululand, bordering Mozambique131, and possibly in the northern Cape on the border with Bechuanaland (Botswana) in largely uninhabited areas of the Kalahari. A few survived in the Mkuzi Game Reserve in northern Zululand. The reserve’s warden believed there were at most 12 in the reserve in 1933132. The area with the most surviving lions was around the Sabi River, in what became the Sabi Game Reserve and later Kruger NP. The first warden, James Stevenson-Hamilton (who served from 1902 until 1946), found little game in much of the reserve at first, which he blamed on hunting by members of Steinacker’s Horse (a British unit posted there during the Boer War), and the local people, who had lost much of their stock to rinderpest133. Lions were present and were known to attack both livestock and people, because of the shortage of natural prey. Several Africans serving with the British forces during the war and a British soldier had been killed at night by lions in the years before the warden took up his post134. One aspect of the establishment of Sabi and similar reserves, is that they were originally intended to encourage the recovery of game so that sport hunting could be resumed135. When he first surveyed the reserve in 1902, the new warden found a few zebra, waterbuck and impala and not much else, though he did find the spoor of two lions near an abandoned railway construction site136. He doesn’t give an estimate in his early accounts of the numbers of lions or other predators, but signals early on his intention to cull predators to encourage the recovery of game herds and allay fears of local white farmers that the reserve would be a breeding ground for carnivores that would take their stock137. It was only much later, when the

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reserve became a national park in 1926, when the Sabi and Shingwedzi reserves were joined, that it was realised that tourists visting the park wanted to see lions, which then became an asset not a nuisance138; even then lion culling continued in the park for three more decades. Stevenson-Hamilton did not want the complete extermination of predators in the park, unlike the farmers along the reserve boundary, but saw lions as a threat to the expansion of game numbers and wanted to keep numbers below their natural level, in line with the colonial approach that predators were vermin139. Under Transvaal’s 1908 game laws, lion were classed as vermin, despite being all but wiped out140. The warden spent much of his early years in Sabi trapping, baiting, shooting or hunting lions with dogs141. The annual game reports in the 1910s and 1920s all report progress in eradicating lions, and the use of gintraps as part of the programme to reduce predator numbers – with such success that there were only about 30 lions left between the Crocodile and Olifants rivers (no figure is given for the original number). Between 1903 and 1913, 190 lions were killed by rangers in Sabi142. The warden wrote that throughout the 1910s and into the 1920s there was agitation by local farmers and politicians against the reserve and the idea of making it into a national park, believing it to be a breeding ground for stock-raiding lions, hyenas, wild dogs and leopards. Stevenson-­Hamilton admitted that during the winter months when zebra, kudu and other prey species crossed the Crocodile River out of the reserve and fed on farmers’ crops, lions followed them and killed stock. This angered the farmers, who in letters to the Farmers’ Weekly newspaper in South Africa accused Stevenson-Hamilton of “breeding lions for the last twenty-five years”, ignoring his annual cull143. The warden believed this not only represented the vehement opposition of farmers to predators being present in the reserve but was a move on behalf of hunters who wanted to be able to shoot lions there144. To avert the threat to the reserve he tried to ensure there were no lions in the reserve sections bordering the farmland across the Crocodile River. The Sabi and Shingwedzi reserves became the Kruger NP in 1926, and the minister responsible for the bill, Grobler, urged the warden to stop the extensive killing of lions, a request with which he complied, though the shooting of problem lions continued, with 57 shot in 1931, because they were a menace to visitors or killed livestock outside the park. In his report for 1927, Stevenson-Hamilton estimated that there were about 400 lions in the Sabi area and 150 in the Shingwedzi section. He recorded that 159 lions had been killed by rangers in southern areas of the reserve in 1924 and 153 in 1925145. Altogether, between 1903 and 1927, Stevenson oversaw the killing of 1,272 lions, 1,142 wild dogs, 660 leopards, 269 cheetahs and 521 hyenas, according to a Senior Research Officer for Kruger NP, G.L. Smuts146. Smuts adds that control killing, removal of problems animals and other reasons for culling meant that between 1902 and 1969, 3,031 lions were killed in the park and another 1,000–2,000 were killed outside the park boundaries147. Even though lions were not being killed specifically to reduce numbers after 1926, it is estimated that between 1954 and 1958, 450 lions were killed as problem animals148.

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Despite the continued killing of lions, lion numbers grew steadily when full-scale culling ended. Stevenson-Hamilton was reconciled to this, as it became clear that lions were the major draw for tourists visiting the park. The development of hand-held cameras that tourists could use from their vehicles helped increase the popularity of lions among visitors – this was magnified by the growing interest in affluent societies (including white South Africans) in wildlife films, both documentaries and films for entertainment based on hunting or safaris by people like the British cameraman Cherry Kearton (who filmed Maasai lion hunts and the Roosevelt safari in Kenya) and Martin and Osa Johnson, the American film-makers who recorded their expeditions to Africa149. By 1945, Stevenson-Hamilton believed the population of lions in the park, estimated from the number of lion kills observed, was in the region of 800150. In south-western South Africa, lion numbers recovered following the creation in 1931 of the Kalahari Gemsbok NP, on the border with Botswana. It was established to protect migrating herds of gemsbok, but the park contained lions that moved back and forth into Botswana. The official parks board report for 1939 did not put a figure on the park’s lion population but mentioned prides in the plural and talked of one being made up of six adults. It added that lions had taken donkeys and cattle on the boundaries of the park and on occasions the rangers had to drive them back into the park with gunshots151. The lion population of Namibia (South-West Africa) declined during German colonial rule and then South African occupation. The range shrank and numbers fell, as both livestock protection and hunting took their toll. At the beginning of the 20th century, lions could be found close to Windhoek and Walvis Bay but, as Guggisberg wrote, “the settlers have waged an unceasing war on the big cats, completely exterminating them in many areas”, and they were restricted to the Etosha region, Damaraland, Kunene and the Caprivi Strip, where there were fewer settlers or it was too arid for farming or large-scale pastoralism152. During the late 19th and early 20th centuries, the colonial authorities employed the!Xóô San to hunt lions, leopards and hyenas preying on livestock where the cattle industry was being expanded in eastern Namibia and Western Botswana153. In 1934, Captain Shortridge surveyed the Namibian mammal population and reported that lions were still widely distributed, but with few in the south-west, a scattered population in the south-east but increasingly common the further north you went, with high concentrations in the north-east and Caprivi154. They were present but not numerous in Damaraland but were widely distributed in Kaokoveld and Outjo districts. He observed that the “lion range had retreated steadily in the face of European settlement”155. Lion numbers and distribution were affected in Bechuanaland (Botswana), and the Kalahari regions shared with South Africa and Namibia, by the arrival of Europeans and the increasing availability of horses, guns and iron gintraps to Tswana and other communities, who supplemented pastoralism with hunting. The combination of “transformations in technology and concomitant hunting

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strategies led to serious game depletion” and, by the early 20th century, to declines in trading of wildlife products156. This in turn led to a decline in hunting and wildlife utilisation in the early 20th century, with migrant labour taking its place along with a revival of the livestock industry after the runderpest epidemic of 1896–7157. Where hunting continued it often involved a return to older methods, avoiding the use of guns to prevent discovery by colonial wildlife officers and police implementing hunting laws introduced by the British when they established a protectorate in Bechuanaland158. This allowed a gradual recovery in wildlife numbers, particularly in the central Kalahari and Okavango Delta, where the habitat was unsuitable for livestock production. The livestock industry was widespread in the territory and was the mainstay for rural Tswana communities. The major development in wildlife policy came in the final years of British rule, when concerned white residents formed the Bechuanaland Fauna Preservation Society, calling for regulation of commercial and sport hunting, which was beginning to take a heavy toll of plains game, elephants and lions in northern Bechuanaland and the Delta159. There was a fear among local conservationists that expansion of the hunting industry would deplete wildlife, as safari operators moved to southern Africa following the independence of Kenya and Tanzania, fearing that their days were numbered in the newly independent East African states160. The conservationists worked with Tswana communities to agree a way forward to conserve wildlife from over-hunting and assist local communities to benefit from wildlife either through income from hunting or from tourism. The Batawana regent, Elizabeth Moremi, persuaded her community to vote in favour of the formation of the Moremi Wildlife Reserve in 1963 and to limit and regulate hunting in areas surrounding the reserve161. The creation of the reserve and later of other reserves in the central Kalahari, Chobe and the southern Kalahari paved the way for the protection of wildlife, which led to the resurgence in populations of lions, gemsbok, zebra, wildebeest and elephants. In Angola, lions were widely distributed in colonial times along the eastern border areas with Congo and Zambia and in the country to south of Benguela on the Atlantic coast, along the southern and south-eastern border with Namibia and south-western Zambia162. Until the 1950s and early 1960s, the Portuguese did little to conserve wildlife. Colonial officials, Portuguese settlers and soldiers all hunted extensively with no meaningful regulation in place. The British hunter, Colonel F.C.R. Statham, said that the greatest threat to game in Angola in the 1920s was “that great enemy of all wild things, the Boer hunter and his wagon”163. Boer settlers and transport drivers were among the keenest hunters in the colony. The Portuguese established what is now the Bicuari NP as a hunting reserve in 1938; it became a game reserve in 1964 with greater restriction of hunting. In 1957, reserves to protect mammal species had been established by the Portuguese colonial authorities at Iona in Namibe province (bordering northwest Namibia along the Kunenhe River and Atlantic coast), Casmeia in Moxico province in the south-east and Kissama in Bengo province164.

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In Malawi, the widespread lion population was reduced under colonial rule by a combination of hunting, the spread of tea and tobacco farming and the steady growth in the human population. A game census for the British protectorate in 1904 listed 543 lions, 751 leopards, 385 hyena, 530 wild dogs and over 100,000 antelopes and other herbivores165. Hunting by local people was not completely banned in the protectorate, perhaps because of the low level of European settlement, but a gun tax had been introduced in 1891 to restrict firearm ownership and there were attempts to restrict the previously unregulated killing of wildlife166. Game returns for the colony list regular shooting of lions on licence, with five shot in 1907 and 17 shot in 1912167. By the 1920s, most large game species had retreated to tsetse fly-infested areas, such as around Kasungu and in the Shire Valley. Many species of herbivore and lions were to be found on the Nyika Plateau in the north. In some districts, wildlife was still a problem. In the Misuku hills of northern Nyasaland, lion and leopard predation on cattle was a major problem – hundreds of cattle were killed every year168. Man-eaters were a regular problem in southern Malawi and in neighbouring areas of Mozambique, notably Benga. According to the trader and hunter John Taylor, who tracked down some of the man-eaters there, they occurred in dry thorn-bush country, which had little in the way of natural prey. The lions he shot for attacking people were not all old, sick or injured aniumals, but were driven by hunger169. Taylor hunted a group of five man-eating lions near the Lupata Gorge on the Zambezi, where the borders of Mozambique, Zambia and Zimbabwe meet. As with the Tsavo man-eaters, they would kill in one district and move several miles before killing again. There was sufficient game in surrounding areas to support the lions but they seemed to have found hunting people easier170. Similarly, man-eating developed among lions in an area of western Mozambique near Mangoche in Nyasaland. Taylor said that there was little game in the area apart from warthog and bushpigs, which regularly raided farmers’ fields, thereby bringing the lions into well-populated regions, where they regularly killed people for food171. To the west in Northern Rhodesia (Zambia), there were periodic problems recorded with lions killing local people or their livestock172. As in most other British colonies, lions were listed in game schedules as vermin and could be killed without a licence or any regulation of numbers. Protection only began in certain areas declared reserves in the 1940s, with Kafue, Mweru Marsh, the Luangwa Valley, Sumba and Lukusuzi all declared protected reserves by 1942173. A report by the wildlife authorities in 1950 noted that the Kafue reserve had been redesignated as a national park and listed lions among the mammal species present174. In Southern Rhodesia (Zimbabwe), there was a long history of hunting among the Ndebele but also visiting European hunters. Colonisation by the ­British South Africa Company (BSAC) of Cecil Rhodes and then self-government as a settler-dominated British territory saw an influx of English and A ­ frikaner settlers and the establishment of huge tobacco, wheat and livestock farms, which

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involved the extirpation of wildlife, including lions, in most farming areas outside what became reserves and national parks such as Hwange, Victoria Falls, Mana Pools, Matusadona and Gonarezhou. Wildlife policy under company rule and then self-government allowed for the mass slaughter of game for a variety of reasons from livestock protection, land clearance, rabies prevention and tsetse fly eradication. Between 1906 and 1912 thousands of predators, other wildlife and over 100,000 dogs were killed to prevent the spread of rabies175. There was a concerted campaign to eradicate vermin, which included lions, leopards, cheetah, hyenas and wild dogs, on European-owned farms. Over one million game animals were killed on and around European farms to create buffer zones to prevent tsetse fly176. The BSAC paid compensation to European farmers who spent time eradicating lions, with 10 shillings paid for every lion177. This programme was enthusiastically implemented, especially during a drive during the First World War to increase beef production for the British war effort. After the war and the granting of self-government to Southern Rhodesia there was a gradual shift towards conservation, with the government in Salisbury realising that if some species were to survive there would have to be habitat preservation and regulation of hunting. Tourism began to be seen as a source of income. The first reserve was established at Matopos in 1926, followed in 1928 by Wankie (Hwange NP) and then Victoria Falls in 1933178. There was growing state regulation of wildlife and hunting for conservation but also with an eye on income from hunting permits. Hunting by African communities was increasingly restricted. Conservation did not end the tsetse eradication programmes outside reserves or the fencing of European farmland, both of which hastened the disappearance of game and predators. The warden of the Wankie reserve reported in 1934 that lions were not numerous there, but there had been a small increase in eastern districts since it was gazetted. He recorded that several lions had been shot or poisoned by farmers at Ngamo and Sinamatella Ranch along the reserve boundary, despite the lack of evidence of any attacks on stock outside the reserve179. Reports published by the Fauna Society’s journal at the end of the Second World War indicated continuing antipathy of European farmers in Southern Rhodesia to predators in the vicinty of their farms180. Between 1942 and 1965, the Southern Rhodesian administration expanded Hwange and then made it, along with Victoria Falls, Matopos, Gonareazhou, Mana Pools and Matusadona, into a national park181. There was a drive to increase tourism domestically, from South Africa and Europe. The building of hotels/lodges and improvement of roads helped increase income from safari tourism, giving an economic value to conservation. Income also increased through the state monopoly over the issuing of permits to hunt outside reserves182. Hunting outside reserves included large-scale culling of game to clear land for new settlers being attracted from Britain after the war. This led one correspondent, who disguised his or her identity, to the Fauna Society’s journal to lament the disappearance of wildlife outside the reserves and the lack of value placed on

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wildlife by European settlers – the article estimated that between 1948 and 1951, 102,025 game animals were shot183. By the 1960s, most game, and certainly most lions, was restricted to reserves and national parks. Blondie Leathem, veteran game warden and general manager of Bubye Valley Conservancy, told me that lions had been completely eradicated in cattle ranching areas, other than a few migratory lions dispersing from protected areas184.

Sudan, the Horn of Africa and Central Africa British hegemony in Sudan as part of the Anglo-Egyptian joint rule, instituted following the defeat of the Mahdist state by Lord Kitchener’s army in 1898, led to the treatment of the territory as part of the British empire. British traders and hunters had been active in Sudan in the 19th century, prior to the overthrow of the Egyptian overlords and their British-led forces at Khartoum by the Mahdi in 1885. The restoration of colonial rule led to an influx of officials, soldiers and hunters, who engaged in commercial and sports hunting, especially in the southern regions with the greatest concentrations of wildlife. The British authorities established the Sudan Game Department to regulate hunting and grant licences. The report for 1904 recorded that 23 lions had been killed on licence. Lord Cromer, the British Consul-General in Egypt and effective ruler of Sudan, said he believed these figures greatly underestimated the number killed185; no doubt many were killed by local communities to protect livestock. Cromer said the game law, as it stands, “is legally applicable to all natives of the Sudan alike. But it is quite clear, from what Mr. Butler [head of the Sudan Game Department] says, that it cannot be fully enforced. ‘I have not attempted,’ Mr. Butler says, ‘to collect any statistics relating to game killed by natives owing the impossibility of obtaining anything approaching to accurate information. A great deal of destruction undoubtedly went on during the year’”186. Lions were numerous in the Sennar region (near the border with Ethiopia, where Dinder NP is situated), Kassala, Kordofan and Nile provinces. Dozens were shot annually on licence in these areas. Butler told Sir Alfred Pease that the northern limit of the lion in Sudan was between 16 and 17 degrees north (north of Khartoum) and south of there lions occured at Atbara to Gallabat, at Filik, Khor Baraka, Setit and the Bahr el Salaam; on the Blue Nile they were plentiful south of Sennar in south-east Sudan towards the Ethiopian border; on the White Nile they ranged north to Jebelein (north of the current border with South Sudan). Butler said they occurred in good numbers around Sobat, Pibor and Akobo, were absent in the Sudd areas of Bahr el Jebel but present on the drier country away from the swamps, while west of Upper Nile they ranged “practically all over the Bahr el Ghazal province”187. Game records from 1901 to 1908 record the killing of 201 lions188. Pease estimates the real number at well in excess of 300 in the first decade of the century, not counting lions killed by local communities defending their cattle189.

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Brocklehurst, who published an account of the fauna of Sudan, said that, “The lion is only hunted as a defensive measure. So long as he confines his attention to game, he is accorded due respect and left severely alone. But once he begins to apply his strength and cunning to the killing of native cattle, he becomes an expensive pest to be destroyed as rapidly as possible”190. Villagers armed with spears surrounded the area where the lions were thought to be. When the lion charged it was speared, though often hunters were injured or even killed. Scars from lion hunts were treated as coveted trophies191. The Game Department viewed lions and other predators as vermin and game wardens and hunters were encouraged to kill them to preserve game numbers192. Butler believed that the diet of Sudan’s lions regularly took livestock and domesticated camels, breaking into zaribas at night. Man-eating, be believed, occurred periodically in many areas193. Despite persecution encouraged by the Game Department and hunting by local communities, in the 1930s lions were still common. The British colonial official, traveller and writer, Wilfred Thesiger, wrote of his service in Sudan from 1935 that lions were plentiful and and a major problem for pastoralists in ­Darfur194 – suggesting a substantial presence in areas of Sudan that supported wild ungulates and had large livestock populations. Thesiger said the Bani Hussain community of northern Darfur hunted lions on horseback with broad-bladed spears – Thesiger often hunted with them and records shooting 30 lions there195. They were part of the wider Baggara Arab-speaking population of nomadic pastoralists and traders, including the Rizeigat, who became skilled hunters from horseback and were later notorious as the Janjaweed militia in the Darfur civil war, and were involved in poaching and ivory smuggling across large areas of Central Africa and the Sudanic Sahel for centuries196. To the east of Sudan in French Equatorial Africa (Chad), lions were widespread bordering Kordofan and Darfur and around the Lake Chad Basin. Brocklehurst’s survey of Sudan’s mammals noted that lions were found in the border areas with Darfur and in the arid semi-desert regions bordering the Sahara197. Lions were hunted by nomadic pastoralists such as the Zaghawa of eastern Chad and to the far north-west the Kanuri and Tuareg. Tuareg hunters killed lions from horseback with spears, padding the flanks of their horses to protect them198. The elephant hunter W.D.M. Bell reported seeing lions in large numbers and “some of immense size” when hunting game along the Chari River in Chad in the late 1920s199. In the east and south-east of Chad, lions were numerous in the reserve created by the French at Aoukl and around Zakouma (Chad’s most important wildlife area). Lions were also regularly sighted in the savanna regions of what became the Central African Republic, the Republic of Congo and Rwanda’s Akagera area. The Cameroonian population was cut off from the small populations in Congo and Gabon by dense forest200. British and other European hunters hunted lions extensively in Somaliland in the late 19th and early 20th centuries. Guggisberg talks of thousands having been shot by the end of the first decades of the 20th century201. As early as 1900 hunters complained of their scarcity. One, Charles Peel, wrote: “owing to

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the ruthless slaughter of females by so-called sportsmen, the lion is becoming extremely scarce within 150 miles of the coast … They are totally extinct in the Golis Range, where but ten years ago they made the valleys re-echo with their magnificent music. North of the Golis Range in Guban there is not a single lion track to be seen.”202 A Captain Mosse recorded that by 1912 much of the wildlife, including lions, had disappeared in the interior. But he said that on his journey from the port of Berberas to Hargeisa he saw lions, adding that they were neither uncommon nor abundant203. Nevertheless, at one stage on the journey he shoots five lions in the space of two days and then three more over the next few weeks204. In 1913, the British Commissioner for Somaliland estimated that there could be as many as 3,000 lions in the territory, but that they were threatened by local people hunting the animals they preyed on205; the killing of thousands by European hunters over 20 years was clearly not considered a threat. In Ethiopia, lions survived in areas with suitable habitat and limited human population, including on the border with Sudan adjacent to the Dinder region, in the south-west in the Omo district and Nech Sar, the Bale Mountains, the Genale River district of south-east Ethiopia, the Awash Valley and the Ogaden. The main threats in Ethiopia were not hunting but human population expansion and habitat loss. During his journeys in the early 1930s through the Danakil Depression and Awash Valley, Thesiger said local people told him that lions were still to be found in the wooded mountains along the Awash Valley. He believed there had only ever been a few in the Danakil and the access of local people to firearms had led to their extermination 206. In 1952, it was reported that lions were “very prevalent” in the hills of Nech Sar region around Lake Chamo. They were said to be smaller than lions found on the plains with heavier manes and darker colouration; no estimates of number were given 207.

West Africa – progressive decline and fragmentation of populations Because of the small regional lion population, the lack of European settlers and of a developed hunting or safari sector, information on lion numbers and ­coexistence/conflict with local people in West Africa is as sparse and as fragmented as the lion range itself. Lions were found in small numbers in suitable savanna, woodland or semi-desert habitats across many of the territories, but rarely in large numbers and always subject to pressures of human expansion, agricultural development and habitat loss. They were present from the colonial territories that would become Guinea and Senegal on the Atlantic coast through Mali, Ghana, Benin, Burkina Faso, Togo and Nigeria into Niger and Cameroon. Nowhere numerous and in increasingly isolated populations, lions were very vulnerable by the early 20th century. In Nigeria, there was suitable habitat in the north, around the Lake Chad Basin and along parts of the eastern border with Cameroon, but low volumes of prey species and growing human populations kept numbers down. Some areas, such as Yola between the Benue and Gongola

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rivers, and Yankari, west of the border with Cameroon, had “a few families of lion” and plenty of prey species such as roan, buffalo, hartebeest, waterbuck, kob and reedbuck, according to a survey of conservation in Nigeria by Richard Oakley in 1931208. The Lake Chad Basin and border areas with Cameroon had isolated lion populations. Cameroon had a larger population in the grasslands of western and northern districts. One of the largest concentrations was in the region now home to the Waza NP, which was estimated in the 1950s to be just 50 animals, while smaller groups occurred between Yaounde and Bemenda 209. A few lions were also reported to be present in northern Ghana 210. There were reports of lions killing livestock in northern Ghana on the border with Burkina Faso (which had a larger lion population in savanna areas) in 1950211. None of the reports give estimates of lion numbers. The area of West Africa with the largest recorded lion population was where the borders of Benin, Burkina Faso, Niger and Togo met, now home to the W, Pendjari and Arly National Parks and a number of smaller reserves and hunting concessions. In the 19th and early 20th centuries lions were recorded to be present in large (but unquantified numbers) in the Air Mountains of Niger, though a British expedition led by Angus Buchanan in 1922 to collect specimens for the Rothschild natural history museum at Tring found none there, blaming their disappearance on desertification rather than hunting.212 In the early 19th century, there were reports of lions surviving in semi-desert areas from Niger, through Mali and into Mauritania, which had a reputation for preying on camels and other livestock of passing caravans and nomadic pastoralists. Reports of lions in Mauritania continued up to the 1940s, in border areas with Mali and Senegal 213. Lions were also reported as far north in the Sahara region as the Adfrar district of south-central Algeria 214.

North African and Asian lions disappear from most of their range Relatively small and often isolated populations of Barbary lions survived in North Africa in coastal regions and the mountainous hinterland into the 19th century, despite the encouragement of lion hunting by the Ottoman Empire. The decline of the Ottomans led to colonisation by the French (from Tunisia to Morocco), with Spanish occupation of northern Morocco, Italian colonisation of Libya and the hegemony of the British in Egypt. The Ottomans and then French colonial authorities offered bounties to hunters to kill lions. The increasing number of explorers and hunters in the region in the 19th century opened the way for sport hunting, particularly by French colonial officials and soldiers. This started the progressive decline of Barbary lions and their eventual extinction in the wild. In her survey of the fate of the North African lion, Schnitzler says that into the mid- and late 1890s, lions were still found in less populated and more mountainous areas of Morocco, with recorded shootings of lions in such regions across the 1830 and 1840s. She believes the last lion in the Rif Mountains

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was shot in 1895, but lions survived in the Middle Atlas and the forested area of Budaa as late as 1930, and in the High Atlas until 1935. A lion was shot near Taddert in Western Morocco in 1942, which was the last recorded wild lion in Morocco215. In Algeria, lions survived in the wild during most of the 19th century, with one estimate of the population size in 1840 being as high as 1,000, with the largest number in the forests of eastern Algeria and mountainous areas around Aures216. Pressure of human population expansion, the spread of the desert and hunting combined to reduce numbers to a few hundred by 1860. This did not stop hunting and 176 lions were killed in Algeria between 1873 and 1884217, with another 26 between 1884 and 1893. The last two lions known to have been killed in Algeria were shot in 1891 and 1893, the latter near Batna 218. Reports of lions in remote areas continued into the 1900s219. The lions of Tunisia had been largely killed off by the time of French occupation in the mid-19th century, many as a result of conflict with livestock farmers. Small numbers survived into the 1880s. The British colonial official, Sir Harry Johnston, reported that three lions had been shot near where he was camping in Tunisia in late 1880220. The last known lion was shot in 1891 in the region of Tamerza 221. Rumours of their survival persisted into the 1900s in the Khmir Mountains and near Feriana, but there were no verified sightings222. In Libya, the coastal plain population disappeared in the 18th century, while some animals survived in the far south on the border with Chad and near the Sudanese border223. Black et al. believe that even though firm evidence is lacking, small population groups may have survived in remote, probably mountainous areas of the Maghreb into the 1950s224. The eventual extinction was most likely a result of the isolated nature of surviving populations, over-hunting, loss of prey and human expansion. As Black et al. argue, “As a pride-forming species, P. leo populations are prone to collapse, whereas other felids may survive at lower local population densities by not living in social groups.” They go on to warn that this is a lesson for lion conservation in West and Central Africa, where small populations “persist, even if rarely seen, in fragmented remnants, yet clearly exist at the edge of a precipitous drop into extinction”225. In the Middle East and West Asia, lions survived in remote parts of Syria, in riverine and marsh areas of Iraq and southern Iran. By the end of the 19th century, they had disappeared from Palestine and survived in very small numbers in eastern Turkey and Syria 226. In the early 1900s, Sir Alfred Pease travelled through eastern Turkey and concluded that the lion had been exterminated there but was present in Syria west of Aleppo227. The British archaeologist Sir Austen Layard recorded seeing lions regularly in Iraq in the 1840s and having to light fires to keep them from his camp near Mosul on the river Tigris; he also reported their presence around Nineveh, south of Baghdad 228. The marshes along the Tigris and Euphrates in southern Iraq and the plains of Khuzestan in Iran were strongholds of the West Asian lions, where abundant wild boar and gazelles were the main prey species. A British surveyor named St John, working in Iran

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in the 1870s, said that in the Shiraz region of Iran, four to five lions were shot every year and that cubs were often captured 229 to be sold to wealthy Iranians and Arabs from the Gulf for private zoos. By the middle of the 20th century, the lions of the marshes of southern Iraq had been killed by hunters230. There is no clear date for their disappearance, but it is likely to have been around the end of the First World War. The lions of central Iran had been exterminated or had migrated south by the middle of the 19th century and were found south of Shiraz and in Baluchistan (bordering Pakistan). In Khuzestan, British colonial officials saw them as late as 1908 near Ahwaz, and one was killed in the Khark Valley in 1918231. Later observations date from 1942, north of Dezful in Khuzestan, and 1957 in the Dez Valley232. To the east in Pakistani Baluchistan, bordering southern Afghanistan, a British admiral reported in 1935 that he had seena lion near the Bolan pass on the railway line to Quetta. It had killed a goat and was lying next to the line, this despite the last lion in Pakistan having been recorded as shot in 1810233. They were also present along the Amu Darya River in northern Afghanistan, where lion predation on local people was reported as late as 1838234.

The survival and resurgence of India’s Gir Forest lions In the 18th century, north and central India had a widespread lion population, found across Bihar, Uttar Pradesh, Madhya Pradesh, Haryana, Rajasthan and Gujarat. They were exterminated or had disappeared from most areas by the end of the 19th century, “strong human and wild animal interface, human-wildlife conflict” being the major causes235. Rapid growth in population and agricultural expansion forced lions from much of their range, but extermination as a stock killer and extensive sports hunting by British colonial officials and Indian rulers took a heavy toll. In 1833, Walter Smee, a British soldier serving in western India, recorded extensive predation by lions on livestock 236. He said that he had shot 11 lions in Gujerat in one month 237. Extensive hunting and habitat loss meant that lions had disappeared from most of India by 1880 – exterminated in the Delhi region in 1834, Bihar 1840, Bhagalpur 1842, Eastern Vindyas and Bundelkhand 1865, central India and Rajasthan in 1870. The last lion outside Gujarat was killed in 1884. Lions survived in the Saurashtra district of Kathiawar peninsula, chiefly the Gir Forest, in Gujarat. Their survival is due largely to the Nawabs of Junagadh, who ruled from 1748. The Junagadh nobles hunted in Gir and invited British officials and Indian royalty to hunt there, killing many lions238.The decline was halted in the late 19th century by the Nawab of the time, in whose traditional royal hunting ground most remaining lions resided 239. In the Gir Forest and adjacent areas the main human communities were Maldhari pastoralists and Siddhis, primarily arable farmers. Lions were a potential threat to the Maldharis’ cattle. The reduction of habitat for prey animals and lions was the main consequence for the cats of Siddhi cultivation. A census in 1880 found only 12 lions in the area. In 1900, the Viceroy, Lord Curzon, was

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invited to shoot a lion there, but declined because of their scarcity. A year later he wrote that “the cause of the diminution of wild fauna in India are the steady increase of population, the winding area of cultivation and the improvement in the means of communication”240. With Curzon’s backing, the Nawab declared Gir a royal game reserve, but did not totally ban shooting. By the start of the 20th century, the Gir lions had increased to 31. A drought in the early 1900s stopped the recovery and led to increased human-lion conflict, through the movement of some lions out of the core forest area. Livestock losses and the mauling of 31 people (three of whom died) in 1902 in Junagarh resulted in local people killing three lions241. The Nawab paid compensation and attempts were made to drive the lions back into the Gir area. When lions moved into Baroda district and killed a woman and a child, a British officer shot one and local villagers two others. Soon after, two cattle herders were killed by lions there, leading to a conflict between the Nawab and neighbouring rulers. The loss of lions during conflict with people and through hunting outside the protected area angered the Nawab of Junagadh and he complained to the British administration about the failure of neighbouring rulers to preserve the lions. By 1913, the number was down to 20, mostly in Gir but some in Girnar, Mitiyala, Barda and the Alech hills, joined to Gir by forest corridors242. Conservation efforts bore fruit and by 1920 it was estimated that there could be 50 spread across the whole range on the Kathiawar peninsula 243. Efforts to protect habitat, lions and their prey continued, and numbers increased. But hunting was still permitted and 23 lions were shot between 1920 and 1946, mainly by British hunters244. The lion population had grown to 287 by 1936. Pressure from the Fauna Preservation Society, whose chairman Lord Onslow was the brother-in-law of the British Viceroy, led to strict regulation of the killing of lions, with sport hunting banned in the Nawab’s territory. But neighbouring rulers still refused to cooperate fully, seeing lions as a danger to livestock and people245. Human-lion conflict remained a problem for the remainder of British rule in India, with the lions taking stock owned by the Maldharis and other farming communities246. The East African wildlife specialist, Captain Keith Caldwell, visited Gir in 1938. He downgraded the estimate of the population to 200 and expressed concern that the shortage of natural prey in the region was behind stock killing. Caldwell said 4–5 lions were being killed annually with permission, but at least another 12 were killed outside the Nawab’s jurisdiction 247. A census in 1948–9 by M.A. Winter-Blyth concluded that the wider Gir region contained 219–227 lions, with the possibility of an absolute maximum of 250 in Kathiawar as a whole248. Indian independence brought new problems. The Nawab was a Muslim and during the violent period of partition, he fled to Pakistan. The removal of his protection led to an outbreak of poaching of sambur deer. On one occasion, 70 were shot in one night by poachers. It is thought some lions were shot, too249. The situation began to improve after 1952, when the Indian Board of Wildlife was formed and gradually, fighting the resistance of traditional rulers, began to

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exert control over wildlife refuges250. The Board was fairly ineffective at first because of this resistance, and it took until 1972 for the Indian Wildlife Act to be passed by parliament establishing legal protection and sanctuaries. India’s first prime minister, Jawarhalal Nehru, ordered the Saurashtra authorities right from independence in 1947 to prevent the hunting and illegal killing of lions. This had some success, but Mitra believes that lions were being poached in the Gir region for several years following independence251. A Conservator of the Gir Forest was put in place in 1956 and Nehru’s declaration of the lion as the national animal and a national emblem had a symbolic effect. Initial attempts to preserve lions focused on moving some of the Maldhari cattle camps or nesses out of the forest. Further surveys by Winter-Blyth show the population at 227 in 1950, rising to 293 in 1955. From then the numbers began to rise steadily, though with a dramatic fall in the mid- to late 1960s, possibly as a result of more efficient survey methods giving a more realistic picture, but perhaps also reflecting the effects of over-grazing of domestic stock in the Gir Forest, which reduced natural prey numbers and increased conflict. One of the major problems for the conservation of the lions and the avoidance of human-lion conflict, was the shortage of natural prey and the lions’ reliance on predation of cattle. A study in 1959 showed that the lions’ prey was 25% wild ungulates and 75% livestock 252. Wild prey was scarce due to human encroachment, the persistence of poaching, the results of past hunting and continued legal hunting in areas to the west of Gir Forest253. A major problem during the late 1950s and early 1960s was that police units sent into the forest to stop poaching were themselves involved in poaching. A further problem was the growing use of pesticides like DDT and Folidol by local farmers, believed to have killed between 80 and 100 lions in a ten-year period 254. A long-term issue is that being the last Table 4.1  Population estimates for the Gir region

Year

Male

Female

1936 1955 1963 1968 1974 1979 1985 1990 1995 2001 2005 2010

143 144   82   60   40   52   66   99   94 101   89   97

  91 100 134   64   52   68   75   95 100 114 124 162

Sub-adult Male Sub-adult Female

13 13

18 20 23

Cubs

53 49 69 51 25 50 14 58 48 63 21 71 18 55 72 unidentified 74 23 + 29 unidentified 77

Total 287 293 285 175 180 205 189 257 304 308 359 411

Source: Figures take from R.I. Meena, Sandeep Kumar and Shamshad Alam (2014), Action Plan for the Conservation of the Asiatic Lion (Pantheraleo persica Meyer), Junagadh: Gujarat Forest Department.

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remaining population of wild Asiatic lions and confined to a very small area, genetic diversity is limited with no chance of recruitment, leading to possible infertility and low resistance to disease255. The problem of reliance on livestock as prey increased in the 1960s as more domesticated buffalo and cattle grazed in the sanctuary, reducing vegetation and cover for deer and antelope256. The lion population crashed as a result, falling to 160–170 animals by 1969. In 1970, in the small lion range centred on the forest (500 square miles) there were 112 villages and nesses, 5,000 people and 21,000 domestic animals, with an additional 30,000 brought in during the monsoon when forage outside was scarce. There were a further 450,000 livestock in areas bordering the sanctuary. The presence of so many stock and people worsened the possibility of human-lion conflict 257. The Maldharis lost hundreds of cattle and buffalo annually to the lions, while members of the Harijan (Dalit) community regularly aggressively scavenged lion kills to take the meat and hides; with lions losing some 64% of their prey in this way258. One partially successful measure taken to restrict cattle encroachment was the building of a metre-high wall around the sanctuary area of the Gir Forest, which was declared a national park in 1975. This, combined with the removal of 500 of the 845 families living in the reserve, reduced pressure on the lions and their prey. Vegetation recovered and the wild ungulate population increased. The development of tourism brought in income, which encouraged the state government to conserve the lions259. In the early 1980s, there were the first suggestions that some of the lions should be relocated to reserves elsewhere in India to prevent the possibility that a natural disaster or disease could wipe out the population. By 1981, herds of chital (spotted deer) numbering 30–40 were regularly being seen and larger sambar deer and nilgai antelopes were present in groups scattered throughout the forest 260. It should be noted that in contrast with the retaliatory killing of lions by communities in African lion range states where livestock predation occurred, killing of stock-raiding lions was relatively low and the conflict tended be more related to encroachment and loss of wild prey. The Maldharis, the pastoral community making up most of the human population of the region, are vegetarian Hindus, who raise livestock purely for production of dairy produce. They have been remarkably tolerant of predation. This appears to be because those allowed to live or graze cattle in the protected area are financially better off than those outside the forest, as they don’t have to buy fodder for their stock. As it is more profitable to live alongside the lions than move out, they tolerate high levels of stock loss – compensation paid for losses is an added incentive to tolerate predation 261. There were also very few recorded attacks on Maldharis by lions and evidence that lions took weak, sick and old stock, with a low economic value. The payment of compensation for poor quality animals meant owners received income for animals that might have died of disease or natural causes. This, combined with strict legal enforcement by the Gir Park management, has limited human killing of lions

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As numbers recovered in the late 1980s and 1990s, because of reduced human and livestock pressure and the increase in natural prey, more lions began to disperse outside the protected area of the Gir Forest towards Kamaleshwar Dam and surrounding forest and farmland areas. This led to increased predation on domestic stock as wild prey is scarcer outside the sanctuary. There have been cases of lions breaking into enclosures at night on farms in the dispersal areas262 . But in the forest, the increase in wild prey meant that by the 1990s, Chellam and Johnsingh had found that chital and sambar were making up the majority of prey, instead of livestock 263. Even then, livestock losses were still high with 1,650 taken in Gir Forest and immediate surrounding areas in 1995, 1,171 in 1996 and 1,474 in 1997; buffalo making up 24–27%, cattle 49–54% and smallstock the rest 264 – down from the previously recorded levels of 75% dependence on livestock. Attacks on humans occurred but were not at a high level despite the close proximity, with only 19 human fatalities recorded between 1988 and 1991. Saberwal et al. estimated that there had been 193 attacks on people between 1977 and 1991, many of them a result of chance encounters rather than attacks by lions to prey on people, though during the 1987–8 drought lions were observed to feed on the corpses of people who had died 265. The majority of lion attacks took place outside the protected area in farmland where cultivation and stock-keeping have driven out natural prey, making human-­lion conflict more likely, and involved sub-adults dispersing away from their birth prides266. Dispersal outside the national park and the value attached to conservation of the lions has led the Gujarat government to gazette parts of Girnar, Pania and Mitiyala (forest land outside the NP) as lion sanctuaries, while all the areas of the region that have resident lions have been declared part of the Asiatic Lion Landscape and have some measure of protection 267. This was necessitated as lions in Gir East and West, Sasan, Porbandar, Bhavnagar and Junagadh under Junagadh Wildlife; Junagadh Social Forestry, Amreli Social Forestry, Bhavnagar Social Forestry and Rajkot Social Forestry are not within the remit of the Gir Forest NP and fall under Rajkot Social Forestry, which is not a wildlife division 268. A new danger to the lion population comes from human action to provide water for stock. Six or seven lions have been reported as drowning annually in water troughs or deep wells that provide water for livestock. To prevent further loss, the state government encouraged farmers to build parapets around the wells, and some success was achieved in cutting losses269. By 2010, the lion population in Gir and the dispersal areas had reached 411, with a significant number of young adults among the population with breeding potential for the future270. The predation on livestock was also falling with cattle down to 18.2% of prey in Gir Forest, only increasing dramatically in dispersal areas like Bhavnagar (35.5%) and the Junagadh Social Forest (47.1%), where natural prey was more scarce271. In these areas, there was an oberserved increase in human fatalities in the 2013–16 period, with 14 killed, six in 2016272. There has also been an increase in lions being killed accidentally, six being run over on railways in 2014.

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The population increase and the growing conflict with humans outside protected or sanctuary areas led to a more concerted attempt to find relocation sites in other states. These included the Sita Mata Wildlife Sanctuary, Darrah-Jawahar Sagar Wildlife Sanctuary and Kumbhalgarh Wildlife Sanctuary in Rajasthan and Kuno Palpur Wildlife Sanctuary in Madhya Pradesh. Ravi Chellam and a team of biologists studied these and other sites to try to relocate lions and prevent either overcrowding or a high level of conflict in the unprotected areas surrounding Gir. But most were rejected as having too high a human population or insufficient prey. Kuno Palpur was the most promising site and the Madhya Pradesh government prepared a scheme to move people out of the reserve to make way for the lions, spending £3.4m to upgrade the sanctuary to make it safe for the lions273. But so far nothing further has happened as the Gujarat state government is opposed to “its” lions leaving the state and losing its status as the only area of India with wild lions. Every time Madhya Pradesh makes improvements to Kuno Palpur, the Gujarat authorities bring up new reasons why they can’t be moved, despite a decision by the Indian Supreme Court in April 2013 that some lions must be relocated 274. The court ordered the Ministry of the Environment to take urgent steps for the relocation of some of the lions from Gir to Kuno. It told the authorities to carry out the order within six months. The International Union for Conservation of Nature (IUCN) also urged the Indian government to move some of the lions to ensure population survival and genetic diversity275. The need for relocation was demonstrated in September 2018, when 23 lions from a 26-member pride died from canine distemper virus, thought to have been introduced by domestic dogs276. The opposition Congress Party leader Ahmed Patel said that the deaths were a result of government failure to have a workable plan for the future of the lions; he said 180 lions had died since 2016277. The Gujarat state government still dragged its heels on relocation to Kuno and instead the state’s environment minister said they would fast-track relocation to Barda Dungar sanctuary at Porbandar in Gujarat state, to prevent lions leaving the state278. In 2016, the Zoological Society of London, which is assisting the conservation of Asiatic lions, put the wild population at about 500, up from the 2010 estimate of 411279, with another estimate at the time suggesting a population of 523280. The latter report noted that the IUCN was urging the Indian government to find at least one relocation site with some urgency to avoid overcrowding in Gir. The danger of overcrowding was emphasised when the Gujarat Forest department issued an estimate in July 2017 that the population had reached 650 lions281.

Notes 1 Keith Somerville (2017), Africa’s Long Road Since Independence. The Many Histories of a Continent, London: Penguin, pp. 6–7. 2 Elspeth Huxley (1935), White Man’s Country. Lord Delamere and the Making of Kenya, Volume One: 1870–1914, London: Chatto and Windus, reprinted new edition 1980, p. ix.

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3 Ibid., p. 22. 4 Andrew J. Loveridge, Craig Packer and Adam Dutton (2008), Science and the recreational hunting of lions, in Barney Dickson, Jon Hutton and William M. Adams (Eds), Recreational Hunting, Conservation and Rural Livelihoods, Oxford: Wiley-­ Blackwell/ZSK, 108–123, p. 109. 5 Ibid. 6 John M. Mackenzie (1987), Chivalry, social Darwinism and ritualised killing: the hunting ethos in Central Africa up to 1914, in David Anderson and Richard Grove (Eds), Conservation in Africa: People, Policies and Practice, Cambridge: Cambridge University Press, 41–61, p. 41. 7 Ibid. 8 Chapurukha M. Kusimba (2002), Foreword, in Bruce D. Patterson (2004), The Lions of Tsavo. Exploring the Legacy of Africa’s Notorious Man-Eaters, New York: ­McGraw-Hill, p. xi. 9 Personal communication with Laurence Frank. 10 Jan Bender Shetler (2007), Imagining Serengeti. A History of Landscape Memory in Tanzania from Earliest Times to the Present, Athens: Ohio University Press, p. 180. 11 Ibid. 12 Ibid., p. 188. 13 F.D. Lugard (1893), The Rise of Our East African Empire, Vol. I, London: Blackwood, p. 352. 14 Ibid. 15 Ibid. 16 John M. Mackenzie (1988), The Empire of Nature. Hunting, Conservation and British Imperialism, Manchester: University of Manchester Press, p. 201. 17 Ibid. 18 Ibid., pp. 208–9. 19 IUCN Environmental Law programme, An Introduction to the African Convention on the Conservation of Nature and Natural Resources, www.sprep.org/attachments/Legal/ IUCNApia.pdf, accessed 29 May 2018. 20 Keith Somerville (2016), Ivory. Power and Poaching in Africa, London: Hurst, p. 60. 21 Robin S. Reid (2012), Savannas of Our Birth. People, Wildlife and Change in East ­Africa, Berkeley: University of California Press, p. 105. 22 Major W. Robert Foran (1933), Kill or be Killed. The Rambling Reminiscences of an Amateur Hunter, London: Hutchinson, reprinted 2017, p. 17. 23 W. Hobley (1937), The conservation of wildlife retrospect and prospect, Part I, ­Journal of the Society for the Preservation of the Fauna of the Empire, vol. 32, p. 42. Henceforth will be rendered as Journal. 24 Andrew Loveridge (2018), Lion Hearted: The Life and Death of Cecil & the Future of Africa’s Iconic Cats, Kindle Edition, loc 1688. 25 Edward I. Steinhart (1989), Hunters, poachers and gamekeepers: towards a social history of hunting in colonial Kenya, Journal of African History, 30, 2: 247–264, p. 248. 26 Ibid., p. 251. 27 Ibid., p. 252. 28 Lt-Colonel J.H. Patterson (1907), The Man-Eaters of Tsavo, reprinted by Natraj ­Publishers, Dehra Dun, 1999, p. 20. 29 Ibid., p. 25. 30 Ibid., pp. 65–6. 31 Ibid., pp. 77–8. 32 Ibid., p. 84. 33 Ibid., p. 90. 34 Ronald Hardy (1965), The Iron Snake, London: Collins, pp. 120–1. 35 Ibid., p. 313.

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36 Bruce D. Patterson (2004), The Lions of Tsavo. Exploring the Legacy of Africa’s Notorious Man-Eaters, New York: McGraw-Hill, p. 28. 37 Brian Bertram (1978), Pride of Lions, London: J.M. Dent, p. 238. 38 Patterson, 2004, p. 67. 39 Ibid., pp. 68–9. 40 Ibid., p. 68. 41 Elspeth Huxley (1935), White Man’s Country. Lord Delamere and the Making of Kenya, Volume One: 1870–1914, London: Chatto and Windus, reprinted new edition 1980, p. 79; see also, Joanna Lewis (2000), Empire State-Building. War and Welfare in Kenya 1925–42, Oxford: James Currey, p. 29. 42 Huxley, 1935, p. 85. 43 Ibid., pp. 149–50. 44 Ibid., p. 306. 45 Daphne Sheldrick (1973), The Tsavo Story, London: Collins and Harvill Press, p. 14. 46 Laurence G. Frank et al. (2005), People and predators in Laikipia District, Kenya, in Rosie Woodroffe, Simon Thirgood and Alan Rabiniwitz (Eds), People and Wildlife. Conflict or Coexistence, Cambridge: Cambridge University Press, 286–304, p. 303. 47 Laurence Frank et  al. (2008), Lions, conflict and conservation, in Barbara Croes, Ralph Buij, Hans de Iongh and Hans Bauer (Eds), Management and Conservation of Large Carnivores in West and Central Africa. Proceedings of an international seminar on the conservation of small and hidden species CML/CEDC, 15 and 16 November 2006, Maroua, Cameroon, eofoundation.org/wp-content/uploads/2015/03/2008-Management-and-­ conservation-of-large-carnivores-....pdf, accessed 17 July 2018, 81–98, p. 82. 48 Personal communication with Laurence Frank. 49 Ibid. 50 Somerville, 2016, p. 67. 51 Thomas P. Ofcansky (2002), Paradise Lost: A History of Game Preservation in East ­Africa, Morganstown: West Virginia University Press, p. 15. 52 C.T. Hobley (1922), The fauna of East Africa and its future, Proceedings of the Zoological Society of London, 7 February, pp. 1 and 5–6. (Henceforth will be referred to as Proceedings.) 53 Ibid. 54 Dorothy L. Hodgson (2001), Once Intrepid Warriors: Gender, Ethnicity, and the Cultural Politics of Maasai Development, Bloomington: Indiana University Press, pp. 36–8. 55 David Collett (1987), Pastoralists and wildlife: image and reality in Kenya Maasailand, in Anderson and Grove, 129–148, pp. 138–9. 56 L. Hazzah, A. Bath, S. Dolrenry, A. Dickman and L. Frank (2017), From attitudes to actions: predictors of lion killing by Maasai warriors, PLoS ONE, 12, 1: 3. 57 C.A.W. Guggisberg (1961), Simba. The Life of the Lion, London: Bailey Bros and Swinfen, p. 185. 58 Ibid. 59 Ibid., p. 186. 60 Ibid., loc 2899. 61 W.S. Rainsford (1908), The Land of the Lion, Nairobi, Waxkeep Publishing, Kindle edition, loc 571. 62 Personal communication with Laurence Frank. 63 Ibid., loc 989. 64 Ibid., loc 1435. 65 See, for example, the books by Chapman and others, like Edward John House (1909), On Safari: East African Big Game Hunting on the Veldt, Mount Kenia, and Guaso Maru, reprinted from: A Hunter’s Camp-fires, London: Harper & Brothers, Kindle edition; John T. McCutcheon (1910), In Africa: Hunting Adventures in the Big Game Country, reprinted by Read Books, Kindle edition; and, Richard Tjader (1910), The Big Game of Africa, London: D. Appleton, Kindle edition.

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66 Tjader, 1910, loc 329. 67 Ibid., loc 534. 68 Theodore Roosevelt (1910), African Game Trails, New York: Charles Scribner’s Sons. Kindle edition. 69 Cited by Mackenzie, 1988, p. 162. 70 Deirdre Jackson (2010), Lion, London: Reaktion Books, pp. 168–9. 71 Return of game killed in the East Africa Protectorate during 1911–12, Journal, 1913, vol. 6, pp. 21–23. 72 Sir Alfred Pease (1913), The Southern Game Reserve, British East Africa, Journal, vol. 6, p. 24. 73 Helge Kjekshus (1996), Ecology Control and Economic Development in East African ­History, London: James Currey, pp. 176–7. 74 Anthony R.E. Sinclair (2012), Serengeti Story. Life and Science in the World’s Greatest Wildlife Region, Oxford: Oxford University Press, Kindle edition, pp.  120–3; and, George Schaller (1972), The Serengeti Lion. A Study of Predator-Prey Relations, ­Chicago: University of Chicago Press, p. 6. 75 Guggisberg, 1961, p. 190. 76 Sara Wheeler (2007), Too Close to the Sun. The Life and Times of Denys Finch Hatton, London: Vintage, p. 188. 77 Sinclair, 2012, p. 120. 78 Wheeler, 2007, pp. 203–4. 79 Ibid., p. 123; and, Guggisberg, 1961, p. 191. 80 Brian Herne (1999), White Hunters. The Golden Age of African Safaris, New York: Henry Holt/Owl Books, pp. 9–10. 81 Ibid., p. 71; and, also, Sir Alfred E. Pease (1913), The Book of the Lion, London: John Murray, reprinted by Forgotten Books, London, p. 41. 82 A. Blayney Percival (1924), A Game Ranger’s Note Book, Read Books, Kindle edition, loc 303–308. 83 Ibid., loc 303–308. 84 Colony and Protectorate of Kenya, an ordinance No 58 of 1921, Journal, 1922, vol. 2, p. 36. 85 Karen Blixen (1937), Out of Africa, London: Century Publishing, reprinted 1985, p. 13. 86 W.D.M. Bell (1960), Bell of Africa. The Greatest Elephant Hunter of Them All, Sudbury, Suffolk: Neville Spearman, p. 225. 87 Extracts from Kenya Colony Game Warden’s Report, 1925, Journal, 1926, vol. 6, p. 57. 88 Journal, 1928, vol. 8, pp. 67–8. 89 Journal, 1931, vol. 15, pp. 79–80. 90 Ofcansky, 2002, p. 69. 91 Ian Parker (2004), What I Tell You Three Times is True, Kinloss, Moray: Librario, p. 58. 92 George Adamson (1968), Bwana Game. The Life Story of George Adamson, London: Fontana, pp. 75 and 100. Ian Parker, who worked with Adamson in the Game ­Department, told the author Adamson was an avid user of strychnine, which indiscrimintaely killed lions, leopards, hyenas and other predators when put in carcasses of stock killed by predators. 93 Parker, 2004, p. 5. 94 Edward L. Steinhart (2006), Black Poachers White Hunters. A Social Hisatory of Hunting in Colonial Kenya, Oxford: James Currey, p. 184. 95 Captain Keith Caldwell (1951), Report on a visit to East Africa, Journal, vol. 4, p. 178. 96 Parker, 2004, p. 45. 97 Anthony Cullen and Sydney Downey (1960), Saving the Game, London: Jarrolds Publishers, p. 15.

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98 Ibid. 99 Shetler, 2007, p. 180. 100 Ibid., p. 181. 101 A.R.E. Sinclair (1995), Serengeti past and present, in A.R.E. Sinclair and Peter Arcese, Serengeti II. Dynamics, Management, and Conservation of an Ecosystem, Chicago: University of Chicago Press, 3–30, p. 4. 102 Tanganyika Territory, Game Department Report, 1935, Protection of lion, leopard and cheetah, Journal, 1936, vol. 27, p. 26. 103 Journal, 1936, vol. 2, p. 15. 104 Tanganyika Territory Game Preservation Department Annual Report, 1935, Journal, 1937, vol. 30, p. 81. 105 Ibid. 106 Guggisberg, 1961, p. 256. 107 George Rushby (1965), No More the Tusker, London: W.H. Allen, pp. 182–4. 108 Ibid., p. 185. 109 Patterson, 2004, p. 81. 110 Correspondence with Bruce Patterson, 6 June 2018. 111 Hans Kruuk (2002), Hunter and Hunted. Relationships between Carnivores and People, Cambridge: Cambridge University Press, p. 185. 112 Alan Root, History of the Serengeti, www.serengeti.org/footprints_mh_hi.html, accessed 6 June 2018. 113 Myles Turner (1987), My Serengeti Years, London: Elm Tree Books, p. 27. 114 Ibid., p. 80. 115 Proceedings, 20 January 1903, http://onlinelibrary.wiley.com/doi/10.1111/j.14697998.1903.tb08253.x/epdf, accessed 11 January 2018. 116 Colonel Delme Radcliffe (1905), Notes on the game in the Nile province and in South-Western Uganda, Journal, 2. 117 W.H. Nelson (1925), A Uganda lion problem, Journal, vol. 5, p. 36. 118 Ibid., p. 38. 119 Uganda Protectorate, Extracts from the Annual Report of the Game Warden for 1925, Journal, 1927, vol. 7, pp. 41–4. 120 Adrian Treves and L. Naughton-Treves (1998), Risk and opportunity for humans coexisting with large carnivores, Journal of Human Evolution, 36: 275–282, p. 276. 121 Ibid., pp. 277–8. 122 Guggisberg, 1961, pp. 35–6. 123 Somerville, 2016, p. 63. 124 Farieda Khan (2000), Environmentalism in South Africa: a sociopolitical perspective, Macalester International, 9, article 11. Available at: http://digitalcommons.­ macalester.edu/macintl/vol9/iss1/11, accessed 13 February 2018, p. 157. 125 Ibid., p. 158. 126 John Lambert (2002), ‘The titihoya does not cry here any more’: the crisis in the homestead economy in colonial Natal, in Stephen Dovers, Ruth Edgecombe and Bill Guest (Eds), South Africa’s Environmental History. Cases and Comparisons, Athens: Ohio University Press, 48–60, pp. 56–7. 127 Guggisberg, 1961, p. 39. 128 P. Steyn (2000), The greening of our past? An assessment of South African environmental history, Historiography Series, www.mail-archive.com/ecofem@csf. colorado.edu/msg07917.html, accessed 12 February 2018, p. 7. 129 Andre F. Boshoff and Graham I.H. Kerley (2015), Lost herds of the Highveld: ­evidence from the written historical record, African Journal of Wildlife Research, 45, 3: 287–300, p. 287. 130 Ibid., p. 295. 131 Guggisberg, 1961, p. 39. 132 Province of Natal, Report of Game Conservation (Capt. Potter) for 1933, Journal, 1934, vol. 23, p. 66.

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133 James Stevenson-Hamilton (2012), South African Eden. From Sabi Game Reserve to Kruger National Park, London: Penguin (originally printed in 1937), Kindle edition, loc 124–8. 134 Ibid., loc 288–313. 135 Jane Carruthers (2001), Wildlife and Warfare. The Life of James Stevenson-Hamilton, Pietermaritzburg: University of Natal Press, p. 84. 136 Stevenson-Hamilton, 2012, loc 446–55. 137 Ibid., loc 899–916. 138 Ibid. 139 Carruthers, 2001, pp. 91–3. 140 Handbook of the Game Laws of the Transvaal, 1908, Journal, 1909, vol. 5, p. 71. 141 Ibid., p. 108. 142 Cited by Pease, The Book of the Lion, 1913, p. 115. 143 Stevenson-Hamilton, 2012, loc 3098–120. 144 Ibid. 145 Transvaal Game Reserve (now Kruger National Park) – Report of Game Warden, Journal, 1928, vol. 8, p. 56–7. 146 G.L. Smuts (1982), Lion, Johannesburg: Macmillan, p. 174. 147 Ibid., p. 186. 148 Laurence G. Frank and Rosie Woodroffe (2001), Behaviour of carnivores in exploited and controlled populations, in John L. Gittleman, Stephan M. Funk, ­David W. Macdonald and Robert Wayne (Eds), Carnivore Conservation, Cambridge: ­Cambridge University Press, 419–442, p. 424. 149 David Bunn (2003), An unnatural state. tourism, water and wildlife photography in early Kruger National Park, in William Beinart and Joann McGregor (Eds), Social History and African Environments, 199–218, pp. 200–1. 150 Kruger National Park, Journal, 1946, vol. 53, pp. 26–7. 151 Report of the National Parks Board of Trustees of South Africa, 1939, Journal, 1940, vol. 40, pp. 17–18. 152 Guggisberg, 1961, p. 39. 153 Robert K. Hitchcock (2016), Discontinuities in ethnographic time: a view from Africa, Journal of Anthropological Archaeology, p. 8. 154 Captain G.C. Shortridge (1934), The Mammals of South West Africa, Volume 1, ­L ondon: William Heinemann, p. 76. 155 Ibid., p. 78. 156 Fred Morton and Robert Hitchcock (2014), Tswana hunting: continuities and changes in the Transvaal and Kalahari after 1600, South African Historical Journal, 66, 3: 418–439, DOI: 10.1080/02582473.2013.855809, accessed 20 April 2018, p. 436. 157 Jeff Ramsay (2014), Firearms in nineteenth-century Botswana: the case of Livingstone’s 8-bore bullet, South African Historical Journal, 66, 3: 440–469, p. 444. 158 Morton and Hitchcock, 2014, p. 436. 159 Robert and June Kay (1962), Preservation in N’Gamiland, Oryx, 5, October, pp. 285–6. 160 J.J. Mallinson (1962), Dangers involved in the exploitation of game in N’Gamiland, Oryx, 5, October, p. 288. 161 Somerville, 2016, p.  151; and, June Kay (1963), Moremi Wildlife Reserve, ­Okavango, Oryx, August, p. 2. 162 Guggisberg, 1961, p. 36. 163 Brian J. Huntley (2017), Wildlife at War in Angola. The Rise and Fall of an African Eden, Pretoria: Pretoria Book House, p. 139. 164 Soki Kuedikuenda and Miguel N.G. Xavier (2009), Framework Report on Angola’s Biodiversity, Luanda: Republic of Angola Ministry of the Environment, p. 19. 165 Game Census of the British Central Africa Protectorate, Journal, 1904, vol. 1, appendix 13.

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166 John McCracken (1987), Colonialism, capitalism and the ecological crisis in Malawi: a reassessment, in David Anderson and Richard Grove (Eds), Conservation in Africa: People, Policies and Practice, Cambridge: Cambridge University Press, 63–78, p. 69. 167 Summary of game killed in various parts of the Nyasaland Protectorate under licences issued in the Protectorate of Nyassaland during the year ending March 31, 1912, Journal, 1913, 6. 168 Ibid., p. 71. 169 John Taylor (1959), Maneaters and Marauders, London: Frederick Muller, p. 16. 170 Ibid., p. 46. 171 Ibid., p. 131. 172 Correspondence with the British South Africa Company, Journal, 1905, vol. 2. 173 Somerville, 2016, p. 63. 174 A National Park in Northern Rhodesia, Oryx (formerly Journal), 1950, vol. 1, p. 15. 175 Roben Mutwira (1989), Southern Rhodesian wildlife policy (1890–1953): a question of condoning game slaughter? Journal of Southern African Studies, 15, 2, Special Issue on Politics of Conservation in Southern Africa, January, pp. 250–262, p. 250. 176 Ibid. 177 Ibid., p. 254. 178 Ibid., p. 258. 179 South Rhodesia. Wankie Game Reserve. Warden’s Report, 1934, Journal, 1935, vol. 26, p. 38. 180 H.E. Horny (1945), A note from Southern Rhodesia, Journal, vol. 52, pp. 23–5. 181 Somerville, 2016, p. 87. 182 Mutwira, 1989, p. 260. 183 I.D.M., The Wildlife Situation in Southern Rhodesia, Oryx, 1952, 7, p. 352. 184 Personal communication. 185 Extract from Lord Cromer’s Report for Egypt and the Sudan for the Year, Journal, 1904, vol. 1, appendix 10. 186 Ibid. 187 Pease, 1913, pp. 121–3. 188 Soudan, table of game killed under licences 1901 to 1908, Journal, 1909, vol.  5 p. 134. 189 Pease, 1913, p. 132. 190 Captain H.C. Brocklehurst (1931), Game Animals of the Sudan, London: Gurney and Jackson, cited by J.L. Cloudsely-Thompson (1967), Animal Twilight: Man and Game in Eastern Africa, London: G.T. Foulis, p. 24. 191 Ibid. 192 Ibid., p. 123. 193 Pease, 1913, pp. 124 and 129. 194 Wilfred Thesiger (1979), Desert, Marsh and Mountain. The World of a Nomad, London: Collins, pp. 26–7. 195 Ibid., p. 28. 196 Somerville, 2016, pp. 234–5. 197 Cited by Guggisberg, 1961, p. 31. 198 Ibid., p. 160. 199 Bell, 1960, p. 213. 200 Ibid. 201 Ibid., p. 35. 202 Cited by Guggisberg, 1961, p. 35. 203 Captain A.H.E Mosse (1913), My Somali Book. A Record of Two Shooting Trips, London: Sampson Low, Marston and Company, reprinted by Forgotten Books, pp. 10–11. 204 Ibid., pp. 92, 106 and 142–4.

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205 Pease, 1913, p. 104. 206 Wilfred Thesiger (1998), The Danakil Diary. Journeys through Abyssinia, 1930–34, London: Flamingo, pp. 47–8 and 88–9. 2 07 Captain D.S. Sorrell (1952), Wildlife in Southern Ethiopia, Oryx, 1952, 6, pp. 288–9. 208 Richard Oakley (1931), Game preservation in Nigeria, Journal, vol. 14, pp. 34–5. 209 Ibid., p. 33. 210 Gold Coast. Preservation of Wildlife, Report by Colonel A.H.W. Haywood, May 1932, Journal, 1933, vol. 18, p. 33. 211 The Gold Coast, report on the Game Department for the Year 1950–51, Oryx, 1952, 6, p. 291. 212 Guggisberg, 1961, pp. 29–30. 213 Ibid. 214 Ibid., p. 31. 215 Annik E. Schnitzler (2011), Past and present distribution of the North African– Asian lion subgroup: a review, Mammal Review, 41, 3: 220–243, pp. 231–7. 216 Ibid. 217 Ibid. 218 Guggisberg, 1961, p. 24. 219 S.A. Black et al. (2013), Examining the extinction of the Barbary Lion and its implications for felid conservation, PLoS ONE, 8, 4: 2–3. 220 Sir Harry Johnston KCB and FZS (1898), On the larger mammals of Tunisia, ­P roceedings, May, p. 351. 221 Schnitzler, 2011, pp. 231–7. 222 Black et al., 2013, pp. 2–3. 223 Schnitzler, 2011, pp. 231–7. 224 Black et al., 2013, p. 10. 225 Ibid. 226 Colin Matheson (1947), Man and lion, Journal, vol. 55, p. 33. 227 Pease, 1913, p. 110. 228 Cited by Schnitzler, 2011, pp. 231–7. 229 Ibid. 230 Thesiger, 1979, p. 170. 231 Ibid. 232 Ibid. 233 Guggisberg, 1961, p. 43. 234 Schnitzler, 2011, pp. 231–7. 235 R.I. Meena, Sandeep Kumar and Shamshad Alam (2014), Action Plan for the Conservation of the Asiatic Lion (Panthera leo persica Meyer, 1826), Junagadh: Gujarat Forest Department, p. xi. 236 Captain Walter Smee, Bombay Army, FZS (1833), Some account of the maneless lion of Guzerat, Proceedings, 10 December, p. 171. 237 Ibid. 238 Sudipta Mitra (2005), Gir Forest and the Saga of the Asiatic Lion, New Delhi: Indus Publishing, p. 19. 239 Ibid., p. 16. 240 Ibid. 241 Ibid., p. 22. 242 Meena et al., 2014, pp. 12–13. 243 Ibid., p. 10. 244 Mitra, 2005, p. 24. 245 India, Preservation of the Gir Lion, Journal, 1937, vol. 32, p. 27. 246 Mitra, 2005, p. 27. 247 The Gir Lions, Journal, 1938, vol. 34, pp. 62–3.

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248 The Indian Lion (from a report on the lions of the Gir Forest by M.A. Winter-Blyth), Journal, 1951, vol. 3, pp. 100–1. 249 Mitra, 2005, pp. 27–8. 250 Ibid. 251 Ibid. 252 Meena et al., 2014, p. 15. 253 Mitra, 2005, pp. 27–8. 254 Ibid., pp. 201. 255 Kausik Banerjee and Yadvendradev V. Jhala (2012), Demographic parameters of endangered Asiatic lions (Panthera leo persica) in Gir Forests, India, Journal of Mammalogy, 93, 6: 1420–1430, p. 1420. 256 Lion v Buffalo in the Gir (1969), Oryx, 10, 2, September, p. 111. 257 Wildlife Problems in India, Report from the IUCN Assembly (1970), Oryx, 10, 4, May, pp. 230–1. 258 Ibid. 259 Oryx, 16, 1, May 1981, p. 4. 260 Oryx, 16, 3, February 1982, pp. 206–7. 261 K. Banerjee, Y.V. Jhala, K.S. Chauhan and C.V. Dave (2013), Living with lions: the economics of coexistence in the Gir Forests, India, PLoS ONE, 8, 1: 4. 262 Mitra, 2005, p. 95. 263 Ravi Chellam and A. Johnsingh (1993), Management of Asiatic Lions in the Gir Forest India, Symposia of the Zoological Society of London, London: Zoological Society of London, p. 414. 264 Mitra, 2005, p. 96. 265 Vasant K. Saberwal, James P. Gibbs, Ravi Chellam and A.J.T. Johnsingh (1994), Lion-human conflict in the Gir Forest, India, Conservation Biology, 8, 2: 501–507, pp. 503–4. 266 Ibid. 267 Meena et al., 2014, p. xi. 268 Ibid. 269 Ibid., p. xii. 270 Ibid., p. 11. 271 Ibid., p. 17. 272 M. Venkataraman et al. (2014), Managing success: Asiatic lion conservation, interface, problems and people’s perceptions in the Gir Protected Area, Biological Conservation, 174, 6: 120–126, pp. 124–5. 273 A.J.T. Johnsingh et  al. (2007), Preparations for the reintroduction of Asiatic lion Panthera leo persica into Kuno Wildlife Sanctuary, Madhya Pradesh, India, Oryx, 41, 1, January, 94–5. 274 Himanshu Kaushik (2015), Times of India, 22 April 2015, https://­t imesofindia. ind iat imes.com /cit y/ah med abad/Kuno-ex pansion-must-to-house-l ions/­ articleshow/47007365.cms, accessed 13 September 2018. 275 Mayank Aggarwal (2018), How politicking and state apathy has put the survival of the Asiatic lion at risk in India, https://scroll.in/article/876349/how-­politickingand-state-apathy-has-put-the-sur vival-of-the-asiatic-lion-at-risk-in-india, ­accessed 2 October 2018. 276 Times of India, 4 October 2018, https://timesofindia.indiatimes.com/india/23-­ lions-dead-in-20-days-whats-killing-gujarats-lions/articleshow/66067049.cms, ­accessed 7 October 2018. 277 Times of India, 7 October 2018, https://timesofindia.indiatimes.com/­india/startscheme-on-lines-of-project-tiger-to-save-lions-says-ahmed-patel/­a rticleshow/ 66104201.cms, accessed 7 October 2018. 278 Times of India, 6 October 2018, https://timesofindia.indiatimes.com/india/gujarat-toshift-lions-from-gir-to-barda/articleshow/66093394.cms, accessed 7 October 2018.

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279 Julian Godfrey (2016), Protecting the Asiatic Lion, Asia conservation programme ZSL, 21 March, https://www.zsl.org/blogs/asia-conservation-program/­protectingthe-asiatic-lion, accessed 28 August 2018. 280 Mayank Aggarwal (2018), Is the King of the Jungle being hedged in by man-made boundaries? Mongabay, 2, https://india.mongabay.com/2018/04/20/is-the-kingof-the-jungle-being-hedged-in-by-man-made-boundaries/, accessed 28 August 2018. 281 Times of India, 4 August 2017, https://timesofindia.indiatimes.com/city/­a hmedabad/ lion-population-roars-to-650-in-gujarat-forests/articleshow/59907625.cms, ­accessed 29 August 2018.

5 Contemporary coexistence and conflict in Africa

Between Sudan’s independence on 1 January 1956 and Angola’s on 11 November 1975, most territories gained independence from their colonisers1. With independence came “an inherited system of wildlife management and a developing system of conservation through national parks and reserves … systems established to benefit colonial powers, along with the measures instituted to conserve habitats and species”2. The legacy, retained unaltered by most states, included criminalisation or tight regulation of hunting by indigenous peoples. Areas gazetted by colonial regimes as national parks or reserves, from which local communities were expelled or limited in their access, remained exclusionary and were often extended, as governments saw the potential for tourist income, with lions as a major attraction. No verifiable figures exist for lion populations at the time. An educated guess would be that they numbered in the hundreds of thousands3. It is evident that in the 60 years since then, as Riggio et al. identify, “numbers have declined precipitously … Given that many now live in small, isolated populations, this trend will continue”4. A wide range of estimates, many educated guesses, exists: from about 400,000 in 1950 to between 30,000 and 100,000 in the mid-1990s5. Over that period the range over which lions are found shrank dramatically. Funston et al. suggest that they now occupy only 8% of their original historical range, and have declined by 45% since 19956. There are now thought to be between 23,000 and 32,000 lions remaining in the wild in Africa in 59 different populations, over half of which have under 100 lions and only six of them have at least 1,000 lions, according to an extensive study being carried out by Amy Dickman, David Macdonald and others at WildCru, University of Oxford7. This chapter will focus on the recent history of human-lion coexistence in sub-Saharan Africa as a whole, then look in greater detail at West, Central, ­Sudan, the Horn of Africa and East Africa.

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Africa – the big picture Lions have lost most of their habitat in West and Central Africa, and are becoming restricted even in their more expansive ranges in East and Southern Africa. In 2002, Chardonnet projected the total area in which lions resided at 2,950,367 km 2 – East Africa 39%, Southern Africa 35%, Central Africa 22% and

Figure 5.1 

A frican range map

Source: IUCN Red List, Panther, Natural Earth © Panthera 2016

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West Africa 4%8. Riggio et al. said the likely lion range in 2012 was down to 25% of the original savannah area they inhabited9. Lions have disappeared from Mauritania, Gambia, Togo, Djibouti, Lesotho and almost certainly Sierra Leone and Eritrea, with concern that there are no longer viable populations in Ghana, Guinea, Guinea Bissau, Ivory Coast or Mali10. Estimating the remaining population is not easy. Access to lion range areas may be limited by conflict, inaccessibility of terrain or unwillingness of governments to permit research11. Chardonnet’s 2002 extrapolation from a number of surveys gave a range of 28,854–47,312, with a mean of 39,37312. In 2004, Bauer and van der Merwe gave a range of 16,500–30,000, with a probable population around 23,00013. They emphasised the difficulty of reaching an accurate figure that took account of all populations, noting that unsurveyed areas across Africa may contain “substantial numbers of lions”14. The major exercise in population estimation by Riggio et al. stressed the importance of “lion strongholds”, areas containing at least 500 lions which are in protected areas (PAs) or designated hunting areas and have stable or expanding populations, for the survival of the lion in the wild15. Their 2012 estimate is given as two figures, one aggregating data on populations including estimates given by the hunting organisations, the Safari Club International (SCI) and International Foundation for the Conservation of Wildlife (IGF), and a separate figure omitting the numbers from the hunting organisations. They produced this comparison of their figures with those of Chardonnet, Bauer and van der Merwe and an IUCN estimate from 2006: The likely remaining wild population in Africa lies somewhere between 23,000 and 32,000 but probably at the bottom end of that scale16. Amy Dickman told me in August 2018 that around or just under 24,000 would be realistic and that in the vast majority of the 59 populations lions are present in numbers well below the carrying capacities of the ranges they occupy averaging 37% of capacity across all ranges.17 Table 5.1  Lion numbers by region and by source

Region

Chardonnet (2002)

Bauer and Van Der IUCN Merwe (2004) (2006)

Present review Present review No SCI/IGF

West Central East South Total

  1,213   2,765 20,485 13,482 37,945

   701    860 11,167   9,415 22,143

   480   2,419 19,972 12,036 34,907

  1,640   2,410 17,290 11,820 33,160

   525   2,267 18,308 11,160 32,260

Note: Riggio et  al.’s 2012 estimate is given as two figures, one aggregating data on populations including estimates given by the hunting organisations, the Safari Club International (SCI) and International Foundation for the Conservation of Wildlife (IGF), and a separate figure omitting the numbers from the hunting organisations. Source: J. Riggio et al. (2012), The size of savannah Africa: a lion’s (Panthera leo) view, Biodiversity Conservation, 22, December, pp. 26–7.

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Across the remaining lion range there is a patchwork of national parks, game reserves or conservation areas (where some human presence and hunting may be allowed), conservancies, hunting blocks and unprotected areas, where wildlife exists alongside humans and their livestock. The major, entirely human-­generated threats are loss of habitat and prey through human encroachment, human-lion conflict and bushmeat poaching, and the lesser problems of direct poaching and the effects of poorly regulated trophy hunting. The factors affecting lions are paralleled by and linked with declines in wildlife populations across Africa, with 69 large mammal species in 78 protected areas across West, East and Southern Africa falling from a 1970 index figure of 1 to 0.41 in 2005; with West Africa’s index going from 1 to 0.15, East Africa from 1 to 0.48 but Southern A ­ frica increasing to 1.24. Lions have declined by 43% across Africa, with a 60% decline in West Africa; while in Botswana, Namibia, South Africa and Zimbabwe, home to between 24% and 34% of Africa’s lions, they have increased by 12%18. The once continuous lion range and network of linked populations, as Dolrenry et al. describe, “exhibits a metapopulation structure: distinct populations within a wider landscape with limited migration between them”19. Their study indicated increasing separation of populations, with limited dispersal and recruitment between them, with the danger of declining genetic diversity, increased vulnerability to disease and lower fertility. They emphasise how important dispersal is to population health and replenishment, particularly through the regular flow of male dispersers in areas where adult male survival may be low through human action 20. Lions do well, in the short term, within fenced reserves, but while fences may protect lions from people and vice versa and may increase tolerance by people of expanding lion populations within fenced areas, they can lead to population fragmentation and genetic isolation 21. The loss of habitat to growing human populations, with the continual expansion of agriculture eating away at lion habitat, is the underlying factor that fuels conflict between people and lions. It is a process that will continue “as Africa’s human population grows exponentially—from 1.2 billion currently to 2.47 billion in 2050”. As more land is needed to produce the food, the cultivated land area is expected to increase by 21% and livestock by 73%22. The need to feed the expanding human population will shrink lion ranges, increase competition and conflict with livestock farmers, and increase demand for bushmeat for subsistence and commercial sale in towns. Hunting for bushmeat drives declines in wildlife and is ubiquitous throughout the lion’s range23. This leads to the depletion of lions as ungulates disappear and increasing conflict as lions prey on domestic stock, resulting in retaliatory killings. Lions may thrive in fenced and well-managed areas and have a greater chance of protection from anthropogenic factors than elsewhere. Populations in substantial fenced areas buck the trend of decline across Africa and show increases24. Fenced reserves or conservancies in Southern Africa have had the greatest success in increasing lion numbers, with South Africa enjoying the highest rate of growth, including in populations re-established on land formerly used

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for farming 25. The downside, as will be examined in the regional sections below, is that the more successful the fenced area is in increasing lion populations, the more heavily lions have to be managed to avoid overpopulation and promote genetic diversity. This may involve culling, trophy hunting or attempts to relocate them elsewhere – this is being carried out at the Bubye Valley Conservancy in Zimbabwe, for example, where major population growth is controlled by trophy hunting and culling 26. Unfenced protected areas are more vulnerable and prone to human-lion conflict, but are crucial to the survival of substantial and genetically diverse lion populations27. State-owned PAs in Africa account for 14.7% of land area, though many effectively exist on paper only and have minimal effect in reality because of inadequate funding or insufficient, poorly motivated staff. Some countries have set aside huge areas for national parks or protected reserves – Botswana 40%, Zambia 38% and Tanzania 32% (though this latter figure may fall as areas are de-gazetted by President Magufuli)28 – and many of these have buffer zones around them with private safari tourism or hunting concessions, increasing the areas of contiguous wildlife habitat. These areas, such as Selous and Ruaha in Tanzania, Serengeti-Mara (Tanzania-Kenya), the Kavango-Zambezi area (in Southern Africa) and the Greater Limpopo Transfrontier Park (linking South Africa, Mozambique and Zimbabwe), contain the largest lion populations. One important aspect of the debates over lion and other wildlife conservation is the role of sustainable or consumptive approaches. Lindsey et al. note that lion populations fare less well in areas where there is no economic utilisation of wildlife that brings benefits to local people from the presence of wildlife29. This lack of economic benefit generates resentment of wildlife, retaliatory killings of lions and greater poaching for bushmeat 30. The majority of people who coexist with wildlife populations adjacent to PAs are poor pastoralists and small-scale farmers pursuing lifestyles which may have changed little for hundreds of years. They have coexisted with wildlife for centuries with regular but relatively limited utilisation of wildlife resources through hunting, without substantially reducing numbers, even of predators. But massive human population growth, the need for food and the increasing fencing-off of farmland have increased conflict with wildlife. Conflict with humans is the most important cause of adult lion mortality31. The reasons for the conflict are complex and will be dealt with region by region.

West and Central Africa – decline and fragmentation The lion populations in these areas are smaller, more fragmented and vulnerable than those in East and Southern Africa. The key reasons are high human population density, habitat loss and hunting (bushmeat, retaliatory killings, poaching for body parts and poorly regulated sport hunting)32. A 2005 workshop on the status of West African lions concluded that it had been reduced to 17 lion conservation units over 5.8% of its former range33. Riggio et al. in 2012 listed

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just eight populations in West Africa, with a probable total of 480: 50 lions in Senegal (shared with Guinea, Guinea Bissau and Mali), 6 in Guinea, 350 in the W-Arly-Pendjari National Park (WAP) straddling Benin, Burkina Faso and Niger, another 40 across three other areas of Benin, and 34 in Nigeria. In a later study, Henschel et al. identified 21 large PAs, amounting to about 37% of the surviving regional lion range, but concluded that lions were only definitely present in four of them, while in two in Guinea, which used to have lions, they had not been seen for ten years34 – though the shooting of a male lion in Guinea in August 2018 (with a picture of the hunter and the dead lion put on Facebook) indicated that some had survived 35. The estimate of the population in 2014 was pessimistic, only three PAs having populations of around 50 individuals and only the WAP park with a large population, put at 356. The total number of lions remaining in the region was a probable 406, within a range of 250–587. The region’s lions are critically endangered. The plight of West African and Central African lions – the latter with an estimated 2,419 remaining in eight populations across Cameroon, the Central African Republic (CAR), Chad, the Democratic Republic of Congo (DRC), and at least one lion on the border of Congo and Gabon – is an extreme example of the threats facing lions across Africa 36. The severity of human pressure has caused massive decline in all wildlife in the two regions over the last 50 years, with habitat destruction and bushmeat hunting being the main causes37. To make the chances of survival of populations worse, conservation is underfunded. The PAs containing lions or with the potential to support large populations have inadequate budgets and management 38. Tourism income and potential is low, compared with East and Southern Africa, and contributes little to conservation. In the areas surrounding WAP (in Benin and Burkina Faso) and in Cameroon, one of the options for funding lion conservation and anti-poaching measures to preserve prey species is trophy hunting, though with the need for strict regulation given low lion numbers. Poorly regulated trophy hunting would be a threat rather than a possible conservation tool39. National parks and reserves need improved funding and management and must work to ensure the recovery of prey species, which declined by 85% across West African savanna PAs between 1970 and 2005. The depletion is continuing through extensive bushmeat hunting in and around PAs, as well as encroachment by nomadic herders40. The WAP complex of Benin, Burkina Faso and Niger contains the only stronghold of the West African lion that numbers in three figures. In 2002 Chardonnet put the population there at 450 and suggested there could be other small groups surviving outside WAP. Benin was thought to have fewer than ten in the Mount Kouffé-Wari Maro Forestry Complex and two in the Keto-Dogo Forest Reserve. Niger had its part of the WAP population, with possibly 100 in the region around the W park. Lions have completely disappeared from Niger’s Air Massif. Riggio et al. in their 2012 survey reduced the WAP number to 350, with other small groups believed to number 18 in Alibori Superior, 19 in Trois Rivieres and 3 at Mount Kouffé-Wari Maro, with none listed for other parts of

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Burkina Faso or Niger. The decline in the three countries results from human encroachment on PAs, loss of habitat outside PAs, bushmeat hunting and prey loss. In the 1970s, the populations in Niger outside the WAP, especially those in the north on the border with Algeria and in the Air district, were decimated by military personal, mining prospectors and local government officials hunting for meat and so wiping out prey and starving lions, and shooting lions for sport41. At the time, the W National Park component of WAP was in decline with little funding from the government, inadequate numbers of rangers and an inability to stop bushmeat poaching or killing of lions by pastoralists illegally grazing livestock in the park42. WAP covers 33,000 km 2 and is adjacent to a ring of designated hunting areas, forming a larger ecosystem. The lion range is divided between the park (58.6%), hunting areas (40.6%) and game reserves (0.8%). The area contains 88% of the West African lion population43. Hunting areas are leased by the governments to private operators for 20 years. Legal hunting only takes place in Benin and Burkina Faso. Prior to 1996, hunting was poorly regulated, lacking well-defined leases and obligations on the hunting operators. Over-hunting was prevalent and little was done to conserve prey species44. The system was changed in 1996 to allow operators longer leases, encouraging them to invest in the concessions, develop access roads, waterholes, manage quotas efficiently (set by governments, but administered by operators) and anti-poaching units. Game numbers recovered, which benefited the adjoining WAP45. The extended leases and scope for investment encouraged a more long-term approach by hunting companies, rather than short-term returns through excessive hunting46. Quotas have been set at lower levels to prevent depletion of adult male lions. Of more concern is the level of human-lion conflict inside and outside the PA, with lions attacking livestock and being killed as a result. This takes place when herders take their cattle into the park, and when lions move outside and prey on stock in villages or pastoralists’ camps. According to Sogbohossou, this problem has not been addressed seriously in Benin47. The average annual loss from carnivore predation for a Fulani herder has been estimated at $365, while smallstock owners in villages lose $204 48. The seasonal pattern of attacks replicates that observed across the lion range, with more stock taken during the rainy season when prey disperses. In villages on the boundaries of the park, predation is a major problem for local farmers. Lions killed 335 cattle between 2000 and 2006, out of a total of 3,271 animals killed by predators49. Villagers retaliate and kill predators, often using poison. The most recent recorded case being the poisoning of three lionesses inside WAP in May 2018 by farmers who said the lions had eaten one of their cattle50. A partial aerial survey of the regional W National Park in the dry season of 2012 revealed that in comparison with one in 2003, numbers of livestock illegally in the park had increased fourfold to >50,000 cattle, compared with a