Cats in Australia: Companion and Killer

CATS IN AUSTRALIA

COMPANION AND KILLER

John C. Z. Woinarski, Sarah M. Legge and Chris R. Dickman

© John Woinarski, Sarah Legge and Christopher Dickman 2019

All rights reserved. Except under the conditions described in the Australian Copyright Act 1968 and subsequent amendments, no part of this publication may be reproduced, stored in a retrieval system or transmitted in any form or by any means, electronic, mechanical, photocopying, recording, duplicating or otherwise, without the prior permission of the copyright owner. Contact CSIRO Publishing for all permission requests.

A catalogue record for this book is available from the National Library of Australia.

ISBN: 9781486308439 (pbk.)

ISBN: 9781486308446 (epdf)

ISBN: 9781486308453 (epub)

Published by:

CSIRO Publishing

Locked Bag 10

Clayton South VIC 3169

Australia

Telephone: +61 3 9545 8400

Email: [email protected]

Website: www.publish.csiro.au

Front cover: Crouching cat. Photo: Anton Darius/Unsplash

Back cover: Cat with prey. Illustration: Taylor Maggiocomo

Set in 10.5/12 Minion & Stone Sans

Edited by Joy Window (Living Language)

Cover design by Alicia Freile, Tango Media

Typeset by Desktop Concepts Pty Ltd, Melbourne

Index by Bruce Gillespie

Printed in China by Toppan Leefung Printing Limited

Acknowledgement

CSIRO acknowledges the Traditional Owners of the lands that we live and work on across Australia and pays its respect to Elders past and present. CSIRO recognises that Aboriginal and Torres Strait Islander peoples have made and will continue to make extraordinary contributions to all aspects of Australian life including culture, economy and science.

CSIRO Publishing publishes and distributes scientific, technical and health science books, magazines and journals from Australia to a worldwide audience and conducts these activities autonomously from the research activities of the Commonwealth Scientific and Industrial Research Organisation (CSIRO). The views expressed in this publication are those of the author(s) and do not necessarily represent those of, and should not be attributed to, the publisher or CSIRO. The copyright owner shall not be liable for technical or other errors or omissions contained herein. The reader/user accepts all risks and responsibility for losses, damages, costs and other consequences resulting directly or indirectly from using this information.

The paper this book is printed on is in accordance with the standards of the Forest Stewardship Council®. The FSC® promotes environmentally responsible, socially beneficial and economically viable management of the world’s forests.

Contents

Preface

Acknowledgements

Defining cats

1Domestic cat: origins

2The natural history of cats

3Cats and other predators

4The cat’s world: global distribution and impacts on biodiversity

5Cats in Australia: history, spread, distribution and population size

6Impacts of feral cats on Australian wildlife

7Pet cats in Australia and their impact on wildlife

8Economic and health impacts

9Management of cats in Australia

10 Cat ethics

11 The ill-governed cat: law and policy

12 Cats on the mind: community attitudes

13 Conclusions and outlook

References

Index

Preface

This is an account of cats in Australia. It is not a simple story, for our community has many different perspectives on cats, and there are aspects of the role of cats in Australian nature that remain poorly known or contested. As with people the world over, many Australians love cats: they admire their grace, intelligence, agility, haughty independence, companionship and occasionally given affection. Mostly in the country, some also appreciate cats for their utility, as killers of perceived pests: rats, mice and snakes. As evidence of the regard held for cats by many in our nation, Australians now spend about $4 billion annually simply to feed and care for their pet cats (Animal Medicines Australia 2016). But other Australians hate cats or at least see no place for them in Australian nature and perceive them to be a problem that needs urgently to be resolved. The issue of our regard for and management of cats polarises our community, as it does in many other countries. The way we regard and manage cats is part of a set of much broader concerns about the extent to which we value individual animals relative to wildlife more generically; whether we accept that there are pests that may have no economic detriment, and how we characterise and manage them; and how we see and value nature and naturalness. These are complex issues, further clouded by a discordant policy and legal context for the management of cats in Australia.

Our own perspective here comes from many decades of studying wildlife in tropical, temperate and arid Australia. We care deeply for Australian nature, and have striven to help secure and manage it, such that we can leave to our descendants a world that is as healthy, diverse and beautiful as that we have experienced and loved. But almost everywhere we have worked, the animal communities we have studied are now insecure, diminished or declining, buffeted by an array of threats that are incrementally subverting Australian nature. Feral cats are one of those threats. In all regions and environments where we have worked – in the Simpson Desert, in the suburbs and bushland of Sydney, in the monsoon tropics of Arnhem Land, the north Kimberley and Cape York Peninsula, on remote islands – there are cats. Other than humans, it is likely that there is now no other animal species in Australia with such an extensive distribution.

We are zoologists; we have spent most of our lives trying to understand and care for animals. We have much respect for cats, and they merit such respect. They are remarkable animals, adapting as successfully to snow-covered alpine environments as to the harshest Australian deserts; moderating their life histories to beat to the erratic rhythm of drought and flood; and being expert and clever hunters that can switch their diet opportunistically to take whatever is available.

In this book, we have sought to describe, understand and contextualise this ecology. Particularly, we have sought to weigh up the impacts of cats in Australia, especially to native wildlife; and then to explore how such impacts can be managed. This book is not a polemic. We have sought to marshal all available evidence, and to acknowledge where this evidence is threadbare or contested. We have sought to present diverse opinions, and to describe and evaluate the options for management and policy responses from ethical, economic and environmental perspectives. But, while much in this book concerns the life and ecology of cats, our fundamental focus is in the conservation of Australian wildlife – of woylies, numbats, bilbies, ground parrots and partridge pigeons – and on seeking pathways to diminish those factors that are threatening it and reducing the likelihood of recovery of its many threatened species.

Acknowledgements

This book is based on, and inspired by, the dedication of many researchers and managers who have contributed to our understanding of the impacts of cats in Australia and who have worked to derive and implement management solutions. We pay tribute to those pioneer conservationists who expressed early concerns about the loss of Australian biodiversity and recognised the role of cats in that decline; these included Archibald Campbell, H.H. Finlayson, Colonel E.A. Le Souef, A.S. Le Souef, A.J. North and Frederic Wood Jones. Australian conservation owes much to the insight and advocacy of these pioneers: more’s the pity that they were then so little heeded.

The baton has been passed from previous generations to today’s cohort of dedicated, insightful and practical ecologists including Ian Abbott, Dave Algar, Tony Buckmaster, Andrew Burbidge, Neil Burrows, Mike Calver, Liz Denny, Tim Doherty, Glenn Edwards, Bron Fancourt, Al Glen, Aaron Greenville, Chris Johnson, Billie Lazenby, Mike Letnic, Peter McDonald, Hugh McGregor, Robyn Molsher, Keith Morris, Katherine Moseby, Brett Murphy, Thomas Newsome, Russell Palmer, Rachel Paltridge, David Paton, John Read, Euan Ritchie, Jeff Short and Adrian Wayne, and much of the information presented in this book derives from the outputs of these researchers. We are grateful also to Brett Murphy, Leigh-Ann Woolley and Hayley Geyle for their contributions to analyses of large sets of cat data. We also thank Peter Marra for helping us to better contextualise cats in Australia from a global perspective.

The Australian Government has played an important role in recognising and tackling the problem caused to Australian biodiversity by cats, particularly by developing successive Threat Abatement Plans for feral cats, and through prioritising the control of feral cats in its 2015 Threatened Species Strategy. The then Australian Environment Minister Greg Hunt, and the threatened species commissioners Gregory Andrews and Sally Box, have been instrumental in fostering a dialogue in our community about the role of cats in our country and in galvanising research and management effort to address the threat posed by feral cats; and Julie Quinn from the Department of the Environment and Energy has long overseen the development and implementation of a national Threat Abatement Plan for feral cats.

State government agencies and the non-government organisation sector have pioneered many of the advances in cat control options, from novel techniques for baiting cats to creating cat-free island arks for threatened mammals. John Walmsley of Earth Sanctuaries did much to raise public awareness of the conservation costs of cats and pioneered the use of predator exclosure fencing in Australia. Fenced exclosures have been taken up by state and local government, industry, local community groups, Indigenous groups and NGOs. Of these, the Australian Wildlife Conservancy has been instrumental in raising the public profile of fenced areas in combating the Australian biodiversity crisis caused by introduced predators.

Our compilation of this book was supported by the Australian Government’s National Environmental Science Program (Threatened Species Recovery Hub). We thank Taylor Maggiocomo for her beautiful illustrations. We are grateful also to Jaana Dielenberg and Heather Christensen for their expert guidance and help with communications and administration. We thank Jemima Amery-Gale, Kate Crossing, Milton McAllister, Hugh McGregor, Ash Millar, Keith Morris, John Read, Brooke Rankmore, Pat Taggart and Daniel White for information and checking drafts of some sections, and Trees Legge for translating some Dutch source material.

Thanks to the many people who generously shared photographs and other figures with us: Dave Algar, Nancy Brennan, Mike Calver, Gerardo Ceballos, Kate Crossing, Steve Eldridge, Matthew Godson (Sporting Shooters Association of Australia), Catherine Hall, Joanne Heathcote, Janos Hennicke, Heidy Kikillus, Caz Larcombe, Jiri Lochman, Leo McComb, Hugh McGregor, Nicholas Nunn, Reece Pedler, John Read, Katherine Tuft and Linda van Bommel.

We thank John Manger of CSIRO Publishing for his invitation to write this book, and for his patience and kind and sage advice. We also thank all those at CSIRO Publishing who supported and enhanced this book and were such a pleasure to work with, including Lauren Webb, Tracey Kudis, Briana Melideo and Joy Window.

JW acknowledges the inspiration from and support of friends, colleagues and family, particularly to Brian and Val Woinarski for immersing him in Australian nature, and to Annie Wells and Loki Woinarski for the enjoyment of life.

SL thanks her many cat-loving colleagues for discussions and adventures, but especially Hugh McGregor. She also pays tribute to Ginger and Rascal (her first cats, rescued strays); to Jackie Meowarr (secret agent of Kimberley cat research); and to Sally, Mullega and especially Brangul, her cat-tracking dogs. We learn a lot from our pets. Above all, she thanks Swanie and Koffi, for their shared nest and love of nature, and for being forgiving when she locks herself away on tasks like book-writing.

CD is indebted to his wife, Carol, and daughter, Alice, for their love and inspiration and support in varied research and writing endeavours, including this book, over many years. Carol and Alice also helped to foster his understanding of cat behaviour by introducing two feline companions to the family home and helping to sustain and entertain them – responsibly, as indoor pets – into playful old age. He has also been fortunate to work with many thoughtful and deeply knowledgeable students and colleagues who have helped to pioneer our understanding of cat biology and impacts in Australia, and whose insights have underpinned his contributions to this book. His ecological colleagues and past and present members of the Desert Ecology Research Group at the University of Sydney have been particular wellsprings of support; he is grateful to all.

Collectively, we thank in anticipation those who will follow. Although there have been some extraordinary and important advances during our lifetimes in knowledge of the ecology and impacts of cats in Australia, in public awareness of the problem, and in the effectiveness of management responses, cats are still exerting significant and detrimental impacts across most of Australia. Much remains to be done – by ecologists, by land managers, by Commonwealth, state and local governments, by pet owners, and by our community as a whole; and we hope that this book provides a robust foundation for others to contribute to more enduring pest management solutions, greater awareness of the value and rights of our country’s extraordinary wildlife, and ultimately to the recovery of Australia’s threatened species.

Defining cats

Definitions of cats are myriad, and they float in a brothy soup, with little consensus over which system of classification, or combination of systems, should take precedence. However, there are (sometimes intersecting) axes to this soup, running along the continua of the cat’s lifestyle, socialisation, degree of containment, and degree of ownership. Take ownership: cats can be owned, semi-owned (free-living, not necessarily attached to a household, but provided sometimes with food by people) or unowned. In terms of lifestyle, cats can be pets, farm cats (semi-owned cats living around farm infrastructure), strays (unowned, semi-owned or ex-owned cats living near humans), or feral cats living far from humans. For containment, they can be indoor pets or free-roaming cats (which includes pets allowed to wander outdoors as well as unowned and semi-owned cats, or feral cats living far from humans). Cats can also be socialised, semi-socialised (tolerant of humans) or unsocialised (often used interchangeably with feral). Obviously, these definitions bleed into one another, and some startlingly complex diagrams exist to explain the varied categorisations for different groups of cats (RSPCA Australia 2018a). Cats are even able to move between categories in their lifetimes: pets can transition to being strays, sometimes of their own volition, or more commonly because they got lost or their owners abandoned them. Unowned cats can be brought into shelters, socialised and adopted out as pets. Other labels often used include ‘house cat’ and ‘domestic cat’, which are the commonly used or generally accepted vernacular names for the species Felis catus (Jackson and Groves 2015; Kitchener et al. 2017), although the term ‘domestic cat’ has also confusingly been used to mean all cats that are not feral cats (RSPCA Australia 2018a).

In this book, we follow Kitchener et al. (2017) and Jackson and Groves (2015), and use ‘domestic cat’ to refer to all categories of Felis catus. While acknowledging the nuances of more complex cat classification, in this book we mostly use a relatively simple categorisation of domestic cats into one of two types: pet cats and feral cats. Pet cats are owned by people and generally cared for in a responsible and consistent manner. Feral cats are not formally owned, nor cared for in a consistent manner. Although many feral cats survive without any care from people, and indeed may never set eyes on a person in their entire lives, some feral cats do live near or among people, and may take advantage of anthropogenic resource subsidies (shelter and food). Feral cats are usually completely untamed and unsocialised, but feral cats living in towns may have had opportunities for socialisation. We sometimes use the term ‘stray’ as a shorthand for feral cats living as fringe-dwellers in towns and cities, profiting from the opportunities for shelter and food. We note that strays may be very wayward pets, ex-pets, or semi-socialised or unsocialised feral cats. Our definitions (pet cats; feral cats with one sub-category called stray cats) broadly follow the palette of management options for cats: management objectives for pet cats and feral cats are fundamentally different, and the available options for strays (feral cats living in cities, towns, farms) versus feral cats in natural landscapes (the bush) are also clearly different. Our categorisation for cats corresponds well with the definitions offered by Moodie (1995). Regardless of their categorisation, all cats are Felis catus, and all categories of cats will hunt given the opportunity, regardless of how well fed they are.

Feral cats, pet cats, and everything in between, can be hard to distinguish from looks alone. (a) A feral cat. (Photo: Hugh McGregor/Arid Recovery). (b) Pet cat. (Photo: Joanne Heathcote).

The terms for cat categories are often weaponised; for example, the term ‘community cats’ has been propagated by Outdoor Cat Advocates in the United States for any cat not contained within a household, encompassing feral cats, semi-owned cats, strays and even free-roaming pets. This label engenders a sense of shared responsibility for the wellbeing and fate of individual cats, and precipitates a different sense of the acceptable management options, compared with a term like ‘feral’. For example, social research shows that the public re-weights the relative merits of lethal versus non-lethal control, and cat welfare versus conservation value and disease risk, depending on the cat’s label (Farnworth et al. 2011; Walker et al. 2017).

Given that words are loaded, conveying different meanings and emotions to different people, we use the word pest carefully in this book, and follow the simple and inclusive definition given by the Invasive Plants and Animals Committee (2016): ‘any animal having, or with the potential to have, an adverse economic, environmental or social impact.’ We also use the words introduced, referring to species brought beyond their natural range to Australia by human agencies, deliberately or inadvertently, and invasive, referring to introduced species that can spread from their point of introduction to become pests (‘A non-native species, the establishment and spread of which threatens ecosystems, habitats or other species with economic or environmental harm’ (Invasive Plants and Animals Committee 2016)). Because they are a species distinct from their wild ancestors, domestic cats are classified as a non-native species throughout the world (Kitts-Morgan 2015).

1

Domestic cat: origins

Hear and attend and listen; for this befell and behappened and became and was, O my Best Beloved, when the Tame animals were wild. The Dog was wild, and the Horse was wild, and the Cow was wild, and the Sheep was wild, and the Pig was wild – as wild as wild could be – and they walked in the Wet Wild Woods by their wild lones. But the wildest of all the wild animals was the Cat. He walked by himself, and all places were alike to him. (Kipling 2010)

In one of Kipling’s Just So stories, the Dog is the first animal to be tamed, and the Cat is the reluctant last. Kipling’s keen supposition has been proven correct by a combination of zooarchaeological and paleaogenetic evidence. The dog Canis familiaris was indeed the first species to be domesticated (Leonard et al. 2002), and the cat one of the last (Larson et al. 2014) and, some might argue, not very well (Baldwin 1975; Clutton-Brock 1981; Driscoll et al. 2009b). Nevertheless, cats are now one of the most common pets in the world: looking across 22 countries, cats are the second-most common pet, being present in 23% of households, with dogs present in 33% of households (Animal Medicines Australia 2016). In Australia, pet cats number 3.9 million, outnumbered only by 4.8 million pet dogs. In the United States, pet cats outnumber pet dogs (86 million cats; 78 million dogs) as they do in New Zealand (1.1 million cats to 0.7 million dogs) (Animal Medicines Australia 2016).

In this chapter, we explain how the domestic cat came to be – we give a brief overview of the origins and taxonomy of the cat family (Felidae), and where small cats (including the domestic cat) sit in that taxonomy. Then we explore the fascinating history of cat domestication, which, uniquely among domesticated animals, was facilitated by nationalistically motivated and state-backed religion.

Cat family taxonomy

The cat family emerged in Asia during the Miocene, ~11.5 million years ago (Li et al. 2016). Since then members of the cat family have come to inhabit all continents except Antarctica (and Australia until very recently). Eight lineages have survived to modern times (Johnson et al. 2006; Kitchener et al. 2017). The sabre-toothed cats, Smilodon and Homotherium, were in a lineage that has not made it through, sadly. The extant eight lineages comprise the big cats – clouded leopards, snow leopards and the roaring cats (lions, tigers, leopards, jaguars); caracals; ocelots; lynx; Bornean bay cats; pumas; leopard cats; and finally, the domestic cat lineage (Felis spp.), whose ancestors split from the leopard cat lineage around 7.3 million years ago (Fig. 1.1) (Johnson et al. 2006; O’Brien et al. 2008; Li et al. 2016). The most recent revision of the classification of the cat family, carried out by the Cat Specialist Group of the International Union for Conservation of Nature (IUCN) (Kitchener et al. 2017), recognises 41 extant species of cat across these eight lineages, with two-thirds of these species listed as threatened (Vulnerable, Endangered or Critically Endangered) or Near Threatened by the IUCN (http://www.iucnredlist.org/search).

Fig. 1.1. Left: the phylogeny of the cat family, as revealed by analysis of mitochondrial (upper tree), and autosomal DNA (lower tree). Differences in the branching patterns relate to differences in inheritance patterns for maternal and biparental DNA, in a family where hybridisation has been common, and with strong male-biased dispersal. (Modified from Li et al. 2016). Right: the current distribution of species in the Felis genus, based on the taxonomy and distribution maps of Kitchener et al. 2017. Subspecies are only shown for F. lybica, the African wildcat. Felis catus is descended from F. l. lybica, from Egypt and the Fertile Crescent, encircled in red. From here, the domestic cat has spread to all continents except Antarctica.

The genus Felis contains seven species that originated around the Mediterranean basin and then spread around the Old World: European wildcat F. silvestris, sand cat F. margarita, jungle cat F. chaus, Chinese mountain cat F. bieti, black-footed cat F. nigripes, African wildcat F. lybica, and the domestic cat F. catus (Kitchener et al. 2017) (Fig. 1.1). Note that the taxonomy of the cat family has been fluid, and in some earlier classifications authors arranged the members of the Felis group differently – for example, considering lybica, cafra, ornata and sometimes bieti to be subspecies of F. silvestris (Kitchener 1991).

The domestic cat was first classified by Linnaeus in 1758 as Felis catus, before any other species in the genus. The ancestor of the domestic cat is widely recognised as the African wildcat Felis lybica (Fig. 1.2) (or Felis silvestris lybica in some classifications) (Driscoll et al. 2007; Agnarsson et al. 2010; Kitchener et al. 2017; Ottoni et al. 2017). The taxonomy of domestic animals has been very confusing, but a relatively recent ruling of the International Commission for Zoological Nomenclature (International Commission on Zoological Nomenclature 2003) means that the domestic cat is now treated as a separate taxon from its wild ancestor and regarded as a full species, Felis catus (Gentry et al. 2004; Kitchener et al. 2017).

Domestication

To understand the origins of the domestic cat, we need to first consider domestication: what is it, and how does it happen? Domestication is a complex process and lacks a consensus definition. Clearly it involves a relationship between a domesticator and a domesticate, but not all scholars define this relationship the same way. Most agree that the relationship should be multi-generational, and mutualistic. One species may control the survival and reproduction of the other for its own benefit, for example to make resources more bountiful or predictable; but the domesticated species should also benefit compared with its non-domesticated peers. Some definitions focus on the agency that domesticators have over the lives and reproduction of the domesticated species. Others view the relationship from the domesticate’s perspective – emphasising how that plant or animal has used the domesticator to give itself an advantage over its ancestral peers. (That view might resonate with many cat owners!). Domestication is accompanied by heritable genetic, morphological and behavioural change, and some definitions focus on these aspects of the process (Zeder 2015). Humans are not the only domesticators. For example, ants cultivate a variety of fungi, aphids and scale insects (Schultz et al. 2005). Although domestication can have accidental beginnings (Newsome et al. 2017b), humans have taken domestication to the next level by incorporating intentionality into the process (reviewed in Zeder 2015).

Fig. 1.2. Felis lybica, the African wild cat, which shares common ancestry with the domestic cat Felis catus. (Photo: Leon Emanuel/Wikimedia Commons, CC BY-SA 4.0).

Domestication is a process that usually begins with taming individual animals via conditioned behavioural training and ends with selection for heritable traits that genetically predispose animals towards tameness. Intensive control over breeding is what allows the artificial selection. The primary attribute that people selected for across all mammal species, including cats, was ‘tameness’ (Zeder 2012; Larson and Fuller 2014; Montague et al. 2014; Wilkins et al. 2014). Tamer animals are more approachable, they respond less fearfully to humans and novel stimuli and have a longer developmental window for socialisation (making bonding easier). As well as being tamer, domesticated species tend to share some or all of a suite of certain physical and behavioural characteristics, collectively dubbed the ‘domestication syndrome’. These include smaller brains (e.g. the domestic cat brain is 28% smaller than the wildcat brain, with most of this difference occurring in the limbic system, which controls the endocrine and autonomic nervous systems (Kruska 1988; Zeder 2012)), depigmentation of the skin or fur, foreshortening of the head, smaller teeth, curly tails, floppy ears, changes to the adreno-corticotropic hormone systems and some neurotransmitters, earlier onset of breeding and more frequent breeding cycles, and juvenile behaviour that extends into adulthood (Clutton-Brock 1981; Wilkins et al. 2014).

The existence of the domestication syndrome has been recognised, and pondered, for some time, including by Charles Darwin himself (Darwin 1868). In 1959, Dmitry Belyaev began an astonishing experiment with silver foxes (a colour phase of European red fox Vulpes vulpes), in Novosobirsk, that proved that the morphological and endocrinological features of the domestication syndrome are a by-product of selection for a single trait – tameness. Individual (and unapproachable) foxes in captivity were scored for the extent of their tameness. Only the top-scoring animals in each generation were allowed to breed. Not only did Balyaev successfully produce pet foxes after several generations, but the tame foxes also had patches of white fur, floppy ears, curly tails, foreshortened skulls, and earlier sexual maturity – none of which had been selected for, and all of which are features of the domestication syndrome (Trut 1999).

Recently, Adam Wilkins and colleagues (Wilkins et al. 2014) noted that the symptoms of the domestication syndrome all share a developmental connection: in vertebrate embryos, neural crest cells (stem cells in the dorsal part of the neural tube) migrate to various parts of the body and are involved in the development of a range of different tissues, including the adrenal glands and sympathetic nervous system (which are key components of the flight–fight response as well as other homeostatic systems), pigment cells in the skin, teeth, skull tissue (including bone and cartilage), and an area of the brain associated with learning and reward (Sánchez-Villagra et al. 2016). By selecting for animals that have a more mellow stress response that matures later (thus increasing the duration of the bonding period), people may have selected for reduced neural crest stress function, causing a cascade of other linked behavioural and morphological effects that typify the domestication syndrome. Neural crest cells are also indirectly involved in the development of the forebrain, and reductions in size and function here could relax the inhibitory control over breeding that is normally exercised by the hypothalamic–pituitary–gonadal system, in turn leading to earlier and more frequent oestrus in females. This idea is supported by very recent gene-mapping studies of domestic dogs versus foxes and wolves, which show differences in genes that relate to neural crest cell migration, differentiation and development (Pendleton et al. 2017; Wang et al. 2017).

There are alternative hypotheses to explain the domestication syndrome. For example, the endocrine system is involved in the regulation of development, so selection for tameness (which means selection for a dampened adreno-corticotropic hormone system) might cause shifts in ontogenetic processes. Parts of the domestication syndrome could be linked to the retention of juvenile characteristics (a phenomenon termed paedomorphism), such as fore-shortened skulls, floppy ears (because the neonatal cartilage fails to harden), and juvenile behaviours (Kruska 1988; Trut 1999; Zeder 2015).

Taming cats

Around 40% of all the species in the cat family have been tamed in the last 10 000 or so years, and mostly in areas that supported the earliest transitions to agriculture and settled lifestyles – that is, south-western Asia and North Africa, plus Central-South America (Faure and Kitchener 2009). With increasing settlement, people most likely tolerated or even encouraged commensal behaviour in small felids, relying on them to control rodents that threatened their grain stores, and perhaps to scavenge away discarded meat (Zeder 2012; Larson and Fuller 2014). People also used tamed felids for hunting a range of prey from birds to gazelles (e.g. cheetahs), as a short-term ‘pet’ or status symbol, as human executioners (pumas and jaguars were used for this purpose by the Incas (Guggisberg 1975)), and for entertainment. For example, in India the caracal Caracal caracal was used in betting games: people would make bets on how many birds the caracal would kill after it was placed into a group of doves, hence the expression ‘to throw a cat among the pigeons’ (Faure and Kitchener 2009). However, of all the tamed felids, only one species, the African wildcat, was domesticated (Driscoll et al. 2007; Lipinski et al. 2008; Faure and Kitchener 2009). The domestication of this wildcat was so astonishingly successful that the domestic cat spread around the human world, and its ready availability may have stifled the domestication processes of other felid species (Faure and Kitchener 2009).

Taming wildcats

It’s better to feed one cat than many mice. (Norwegian proverb (Knowles 2016))

Tracing the trajectory of domestication of the wildcat has proved challenging. Archaeologists usually recreate domestication histories by developing chronologies based on iconographies, written histories, and dated material deposits (Larson and Fuller 2014). Recreating the cat domestication process has been hampered by relatively limited zooarchaeological evidence (Serpell 1988; Lentacker and De Cupere 1993), and because the divergence in specific physical traits that characterises most domestications did not occur prominently in wildcats (Serpell 1988). Consequently, the osteology (skeletal shape and structure) of the domestic cat still closely resembles its ancestral form (Zeuner 1963; Randi et al. 2001; O’Connor 2007; Zeder 2012; Ottoni et al. 2016). Any differences in behaviour, tameness and coat colour that could distinguish domestic cats from wildcats (Kitchener et al. 2005) are not preserved in sub-fossils. The advent of modern molecular techniques has helped considerably to enrich our understanding of the cat’s domestication history, but untangling the phylogenetic and phylogeographic puzzles has been fraught because cats were moved around so comprehensively by people. Not only this, the domestic cat is capable of hybridising with all the other Felis species, so domestic cat and wildcat genes have been passed back and forth at different times and places (Clutton-Brock 1981; Daniels et al. 1998; Daniels et al. 2001; Cameron-Beaumont et al. 2002; Oliveira et al. 2008a, 2008b; Agnarsson et al. 2010; Li et al. 2016; Mattucci et al. 2016; Kitchener et al. 2017).

Associations between wildcats and people may have begun in prehistoric times, but the relationships almost certainly intensified with the development of agriculture. Humans began gathering wild cereals in the Near East over 20 000 years ago (Tanno and Willcox 2006), but agriculture – involving planted crops – developed there ~13 000 years ago, and rodents like house mice Mus musculus began to appear at human settlements around the same time (Vigne et al. 2012). Small cats were likely attracted by the surfeit of rodents raiding grain stores and dump areas. Individual cats that were less wary of humans would have been more able to take advantage of these opportunities (Baldwin 1975), leading to weak selection for tamer cats (Driscoll et al. 2009a). Some of the earliest Neolithic settlements show evidence of this human–wildcat connection. The earliest documented connection is from Cyprus. The first farmers settled the island ~11 000 years ago, bringing African wildcats, as well as house mice, goats Capra hircus, cattle Bos taurus, sheep Ovis aries, pigs Sus scrofa, fallow deer Dama dama mesopotamica, red foxes and dogs (Vigne et al. 2004, 2012). Disarticulated remains of wildcats indicate they were eaten by the early farmers, and they were presumably useful also for controlling mice. However, a deposit from Cyprus aged at 9500 years ago contains a wildcat skeleton buried with a human, and the positioning of two bodies in the joint burial site suggests a more complex cultural or emotional relationship (Vigne et al. 2004). Wildcats were also introduced to other Mediterranean islands (that lacked a native wildcat) by Neolithic farmers (Vigne 1992). Burial remains of African wildcats from Jericho dated to 8700 years ago (Zeuner 1963; Clutton-Brock 1981; Malek 1993), and old Neolithic cat figurines (from the 6th millennium BC) in Turkey (Lentacker and De Cupere 1993) reinforce the idea that, for these early farmers, cats had moved beyond being simple commensals to having some cultural significance (Vigne et al. 2004).

Cat–people relationships were not confined to the Near East, and they were not confined to African wildcats. For example, Neolithic farmers in China were associating with commensal leopard cats Prionailurus bengalensis 5500–4000 years ago (Hu et al. 2014; Vigne et al. 2016), and the remains of a jungle cat were present in a human burial site in Egypt dated at 5700 years ago (Linseele et al. 2007, 2008) (incidentally, the remains of African wildcats were found at the same location) (Van Neer et al. 2014). The bones of small cats (Felis sp.) have also been found in Harappa culture sites in the Indus valley, dated to ~4100 years ago (Lentacker and De Cupere 1993). Although people were taming small cats, and genetic analyses show that people were moving cats around (Vigne 1992; Driscoll et al. 2007, 2009b; Ottoni et al. 2017), they were not taking substantial control over the breeding of cats (the prerequisite for full domestication). This situation continued for thousands of years. The factor that finally precipitated the final stage of domestication, whereby people took control of cat breeding and applied stronger selection, was state-managed religion, and it happened in Egypt, beginning 4000 years ago but peaking from 3000 years ago (Malek 1993; Ikram 2005).

The sacred Egyptian cat

Genetic analyses unanimously indicate that the cats domesticated by Egyptians were African wildcats, from Mesopotamia in the Fertile Crescent, where agriculture first developed (Driscoll et al. 2007; Lipinski et al. 2008; Kurushima et al. 2012; Ottoni et al. 2017). All modern domestic cats can be traced, using their mitochondrial DNA, back to matrilines of African wildcats from the Fertile Crescent. The diversity of mitochondrial haplotypes in modern domestic cats is also highest in the Fertile Crescent (Driscoll et al. 2007; Lipinski et al. 2008; Kurushima et al. 2012). The genetic matrilines converge at over 130 000 years ago, even though the agriculture that facilitated domestication occurred only ~13 000 years ago. This indicates that domestication involved the repeated introgression of African wildcat genes from a wide area into the domestic cat breeding pool over a period of time, rather than being a single bottleneck event (Driscoll et al. 2007; Lipinski et al. 2008; Driscoll et al. 2009b). Cats must have been imported to Egypt from Mesopotamia regularly over the well established trading routes. It is likely that the imported cats were interbreeding with local African wildcats (Ottoni et al. 2017). There is also evidence of genetic introgression from other Felis species (Agnarsson et al. 2010). Although it is possible that people took control over cat breeding in places other than Egypt, the long-term impacts of this were swamped by the emergence of an Egyptian obsession with cats, an intensive breeding program, and then a massive diaspora of cats out of Egypt.

Ancient Egyptians associated with small cats for a long time before becoming serious about domestication. For example, the jungle cat found in a human burial site aged at 5700 years ago must have been kept for some weeks leading up to the burial, based on the presence of healed bone fractures (probably sustained during capture) in the cat (Linseele et al. 2007, 2008; Van Neer et al. 2014). The skeletons of 17 cats in a tomb at Abydos, dated at 4000–3800 years old, were accompanied by small offering bowls that had contained milk, suggesting a nurturing relationship, and concern for the cats’ wellbeing in the after-life (Malek 1993).

The growing prominence of small cats in Egyptian culture can be traced by their appearance in iconography: small cats are absent from the iconography until ~4400 years ago. The first appearance of a small wildcat (that we know of) may be on the walls of King Nyuserra’s sun temple at Abu Gurab, south-west of Cairo (~2400 BC). Unfortunately, the fragment of the wall containing the cat was taken to Germany and destroyed during World War II (Malek 1993). The oldest existing representations of cats are hieroglyphs, on a limestone wall-relief dated ~4200 years old. Two hundred years later, the limestone block was moved from its original position at Saqqara, and used in the construction of a temple wall at El-Lisht, 50 km south of Cairo (Malek 1993). Cat hieroglyphics turn up on a handful of tombs over the next 300 years or so, and indicate the name of the interred person was ‘cat’ (‘Miut’ or ‘Miit’).

Fig. 1.3. Fresco in the burial chamber of Menna, the King’s Scribe. Scene: hunting and fishing; detail: cat and mongoose. ~1422–1411 BCE, Thebes. (Photo: The Yorck Project/Wikimedia Commons).

Cats make their first appearance in a clearly domestic setting in a tomb painting at Beni Hassan ~3970 years ago, where a cat, accompanied by a man, is hunting a large rat, possibly alluding to the cat’s role in rodent control (Malek 1993). A slightly later (~3400 years ago) portrayal in a tomb painting at Menna shows a striped tabby, possibly a north African wildcat, beating a mongoose to a bird’s nest (Fig. 1.3) (Malek 1993). From ~4000 years ago, cat shapes also began to be used as vessels for cosmetic oil, and as decorations on jewellery (Clutton-Brock 1981; Malek 1993). From 3500 years ago, small cats became increasingly common in art, portrayed as sitting under chairs (usually under women, rather than men), eating, playing, and hunting birds in the rushes. Their inclusion in these domestic settings indicates their status as pets, possibly even domesticated, rather than tamed commensals (Serpell 1988; Linseele et al. 2007). Cats also featured in Egyptian medicine: the fat of tomcats could be used to deter rodents, cat placentas were an ingredient in a lotion that stopped hair from going grey, and female cat fur folded into human milk and resin was applied to burns (Malek 1993).

Small cats became increasingly integral to religious life, featuring more prominently than large felids like lions which had been more common up to that point (Gautier 1999). At first, cats were a symbol of protection. Magic knives, inscribed with images of animals, including small cats, served to protect the dead person from dangers, and began appearing in tombs from 4000 years ago. The cats were sometimes shown holding a knife, a symbol for cutting off the head of enemies (Malek 1993). Cats got a promotion in the religious hierarchy around 3600 years ago, when they became involved in helping Ra, the sun-god, survive each night (Malek 1993), by killing the serpent (Apep) who tried every night to swallow the sun (Fig. 1.4) (Serpell 1988). The next upgrade in the cat’s religious status was facilitated by politics. The Libyan Pharaoh Sheshonq I took power ~945 BC after a prolonged period of political instability and invasions by Nubians and Assyrians. He made Bubastis his capital, thus raising the local deity Bastet, to national status (Serpell 1988; Malek 1993; Gahlin 2014). By this time, Bast, ‘female devourer’, the protector goddess who had a lion’s head (Gahlin 2014), had morphed into Bastet, a gentler goddess of family, fertility and birth, who had a small cat head (Fig. 1.5) (Wilkinson 2003; Zivie and Lichtenberg 2005). The two extremes of leonine character were opposite manifestations of a linked deity, and Bastet was connected to other gods and goddesses including Sekhmet and Hathor, as well as Ra (Malek 1993; Wilkinson 2003). Small cats were perceived to be nurturing mothers, and women sometimes wore amulets of Bastet with kittens, indicating how many children they wished for. Small cats owned by royalty wore golden jewels and ate off their owners’ plates (Malek 1993).

Fig. 1.4. The Great Cat as Ra, killing Apep the serpent at Thebes. Papyrus of Hunefer, 1300 BCE. (Photo: British Museum/Wikimedia Commons).

Fig. 1.5. (a) The goddess Bast with a lion’s head. (Photo: Walters Art Museum, Boston/Wikimedia Commons). (b) Bastet in her later form as a cat-headed woman. (Gunawan Kartapranata/Wikimedia Commons, CC BY-SA 3.0).

Cats took their last step towards sacredness from around 660 BC, when they became integral to the animal cults. Animal cults started in the predynastic period, before 5000 years ago, gained popularity from 3500 years ago, but reached their peak from ~2780 years ago (Malek 1993; Ikram 2005; Zivie and Lichtenberg 2005). The surge in ‘devotion’ may have been driven by reactive nationalism and a need to appease gods following invasions by Libyans, Nubians, Assyrians and Persians (Nicholson 2005). In addition, there was deliberate state policy to control religion, because it was lucrative – cults were taxed, priests bought their positions, and the populace bought offerings from the temples (Malek 1993). By the time that Herodotus visited Egypt 2500 years ago, the cat cult was at fever-pitch. Family cats who died were mourned: ‘in whatever house a cat dies a natural death, all the family shave their eyebrows …’ and then ceremonially buried: ‘all cats that die are carried to sacred houses, where, being first embalmed, they are buried in the city of Bubastis’ (Herodotus 1920, II: 66–67). Killing a cat was a crime punishable by the priests. Yet ironically, priests killed cats – by the hundreds of thousands or even millions, in order to prepare mummies for votive offerings in areas sacred to Bastet across Egypt (Malek 1993; Zivie and Lichtenberg 2005; Gahlin 2014).

Fig. 1.6. The sarcophagus of the cat of the Crown Prince Djhutmose. (Photo: Larazoni/Flickr, CC BY 2.0).

Egyptians were creating elaborate mummies of animals, as well as people, by 4000 years ago. At first, cat mummies were of beloved pets. For example, 3360 years ago Prince Djhutmose’s pet cat (Tamyt) was interred with its image (sporting a necklace) carved on the outside of its limestone sarcophagus (Fig. 1.6) (Ikram 2005). But by the first millennium BC, when cats had become the manifestation of deities (especially Bastet), cat mummification became an industry. By Herodotus’ time, cats, mostly domestic cats but also other small cat taxa, were the most frequently mummified animal in Egypt (Armitage and Clutton-Brock 1981; Kurushima et al. 2012), offered up in reverence to Bastet, especially during annual festivals (Fig. 1.7) (Malek 1993; Gahlin 2014). Pilgrims would buy the cat mummies at temples and then have them buried in catacombs. Bastet’s temple at Bubastis contained many thousands of mummified cats, as did other cat cemeteries, for example at Saqqara (Zivie and Lichtenberg 2005). An example of the scale of cat burials is that the contents of a single burial ground, unearthed in 1888 at Beni Hassan, were purchased by a businessman and then shipped to England for conversion to fertiliser. A single shipment of 19 t contained ~180 000 cat mummies (Zeuner 1963; Armitage and Clutton-Brock 1981; Malek 1993; Zivie and Lichtenberg 2005).

To satisfy the enormous demand for cat mummies, priests constructed catteries, breeding up cats to be turned into votive offerings (Malek 1993; Gahlin 2014). The priests killed the cats by massive blows to the head that fractured the skull, or by strangling, often breaking their necks (Armitage and Clutton-Brock 1981; Zivie and Lichtenberg 2005). The demand for votive cat mummies was so high that some mummies were faked, with the outer linen and plaster coverings containing sand or other bits of animals, rather than a cat. Priests found selling fake cat mummies were tried and punished, and the practice led to the rule ‘one god in one vessel’ to prevent the sale of vessels without any cat remains inside (Malek 1993; Ikram 2005). Although domesticated African wildcats were the primary source for cat mummies, other small, undomesticated cats were also mummified at a lower rate, including the jungle cat, and the serval Leptailurus serval, which must have been imported to Egypt from the south (Baldwin 1975). The religiously and politically motivated, highly managed nature of cat breeding was the key circumstance that made artificial selection and thus domestication possible.

Fig. 1.7. Mummified cats. (Source: E.A.W Budge, H.R. Hall (1904) A Guide to the Third and Fourth Egyptian Rooms, p. 88. British Museum, London, UK. Photo: Internet Archive Book Images/Flickr).

Out of Egypt

Domesticated Egyptian cats were tightly controlled: there were restrictions on exporting cats, and there were even systems for finding cats that had been smuggled out of Egypt, and bringing them back (Serpell 1988; Donalson 1999). Nonetheless, cats began spreading (or being spread) out of Egypt (Ottoni et al. 2017). Phoenician traders (otherwise known as ‘cat thieves’ by the Egyptians) took cats north over the Mediterranean (Beadle 1977), including to Greece. The exodus from Egypt was initially slow, partly because of the rules against moving cats, and possibly also because one of the cat’s primary tasks, rodent control, was already being carried out in parts of Europe and Africa by other tamed species, especially mustelids Mustela spp. and genets Genetta genetta (Lentacker and De Cupere 1993; Donalson 1999; Lewis and Llewellyn-Jones 2018). Notably, of these alternative pest controllers, only polecats Mustela putorius were also domesticated.

Cats begin to appear regularly in Greek iconography from 2500 years ago, and especially after the Ptolemaic dynasty took control of Egypt from 330 BC (Baldwin 1975; Lewis and Llewellyn-Jones 2018). The cat goddess was exported to Greece along with the cat; Bastet evolved into the Greek goddess Ailuros, and a version of Artemis (moon goddess). Archaeological evidence indicates that domestic cats were transported widely across Europe around this time. For example, skeletons of a family of cats (implying domestic animals rather than wildcats) have been found at an Iron Age settlement in Dorset that was occupied 2500–2000 years ago (Harcourt 1979), and in human graves as far north as the Orkney Islands, Scotland, in deposits dated at 2400 years old (Smith et al. 1994). The Orkney cats were identified as domestic animals rather than European wildcats based on biometric measurements (Smith et al. 1994; O’Connor 2007).

Cats were present in the countryside around Rome by the time of the Republic around 2500 years ago, but it is possible they were mostly feral, or tolerated commensals, rather than loved companions. In the 4th century BC, the agricultural chronicler Palladius noted that ‘it is of service against moles, to have cats in the middle of the [artichoke] plantation’ (Owen 1807; Lewis and Llewellyn-Jones 2018). It took a few hundred years before cats were common in the city of Rome (Donalson 1999; Faure and Kitchener 2009). Ironically, Rome now has an enormous stray cat population (Natoli 1985). Nevertheless, the Romans and especially its armies, were a conduit for cats travelling as stowaways, mascots and rodent killers, especially as Roman expansion ramped up from the 3rd century BC (Clutton-Brock 1981; Malek 1993; Donalson 1999). The cat diaspora accelerated further when the Roman Empire annexed Egypt in 30 BC, after which the religious significance of cats to Egyptians, and the protectionism that accompanied this status, was weakened (Malek 1993; Ikram 2005).

Through a combination of spreading themselves, and being spread by people, domestic cats became common across continental Europe by 500 AD (Zeuner 1963; Faure and Kitchener 2009). For example, by 900 AD in Wales, cats were so normalised that a hamlet was defined as a place with nine buildings, one herdsman, one plough, one kiln, one churn, one bull, one cock and one cat (Sunquist and Sunquist 2002). Cats continued to be moved and mixed around. The Vikings were responsible for a wave of cat-spreading (8th–11th centuries), bringing them home to northern Europe from raids to the south around 1200 years ago. This has been inferred both from genetic evidence (Ottoni et al. 2016) and also the contemporary distribution of pelage patterns (Todd 1977). During the medieval period, returning Crusaders may have moved cats from the Near East and North Africa to areas of Europe, as records of cats in Europe increase from that time (Sunquist and Sunquist 2002).

Domestic cats were present in India from 2200 years ago, possibly earlier, and probably spread east from there quickly, thanks to the opening of the Silk Road from Persia to China (Baldwin 1975). The first record of domestic cats in China is dated to almost 2100 years ago (Baldwin 1975); cats were common there by 1300 years ago, being used as pets and rodent controllers, especially to kill rats that preyed on silk-moth cocoons (Vigne et al. 2016). They were used for a similar purpose in Japan by 1400 years ago, as well as to protect manuscripts from rodents (Faure and Kitchener 2009). Cat traffic between East and West must have slowed after these initial invasions, as east Asian cats are genetically distinct, suggesting a period of isolation following their introduction (Lipinski et al. 2008).

European colonists and traders brought domestic cats to the Americas 500 years ago (Lipinski et al. 2008), and to many Pacific islands from 500–200 years ago (Baldwin 1990; Duffy and Capece 2012). Cats arrived in Australia with the First Fleet in 1788. Smaller expansions radiated