Our Wild Companions: Domestic cats in the Anthropocene

Abstract

Cats are among the most popular companion species in the world, yet there is growing evidence of the environmental impacts wrought by their large populations.The history of their domestication and association with human societies has been effectively traced with archaeological and genetic studies of contemporary and ancient DNA.We assert that the growing conflict over cat management is underpinned by an adaptive push-and-pull between the wild and the domesticated traits of cats, aligning with their dual societal roles as companions and pest controllers.Sustainable solutions require a novel ‘companion animal ecology’ and recognition that cats are not exclusively pets or pests, but are central to human societies, while having important, often adverse, environmental impacts. Cats share a long history with humans but are remarkable among domesticated species in largely retaining behavioural and reproductive independence from people. In many societies, the cat maintains liminal status as both a domestic and a wild animal. An adaptive push-and-pull between wild and domestic traits corresponds with dual roles as companions and pest controllers, and with conflicted treatment in husbandry, management, law, and public discourse. To move forward, we must proceed by understanding that cats are not exclusively pets or pests, but both a central component of human societies and an important, often adverse, influence on ecosystems. Developing a collaborative ‘companion animal ecology’, in which human–animal domestic relations link to ecological processes, will enable sustainable management of this wild companionship. Cats share a long history with humans but are remarkable among domesticated species in largely retaining behavioural and reproductive independence from people. In many societies, the cat maintains liminal status as both a domestic and a wild animal. An adaptive push-and-pull between wild and domestic traits corresponds with dual roles as companions and pest controllers, and with conflicted treatment in husbandry, management, law, and public discourse. To move forward, we must proceed by understanding that cats are not exclusively pets or pests, but both a central component of human societies and an important, often adverse, influence on ecosystems. Developing a collaborative ‘companion animal ecology’, in which human–animal domestic relations link to ecological processes, will enable sustainable management of this wild companionship. The places and roles of domestic cats Felis catus in contemporary societies and ecosystems are contentious issues. A growing body of ecological research quantifying the scale and impact of cat predation of wildlife [1.Loss S.R. et al.The impact of free-ranging domestic cats on wildlife of the United States.Nat. Commun. 2013; 4: 1396Crossref PubMed Scopus (522) Google Scholar, 2.Loss S.R. Marra P.P. Population impacts of free-ranging domestic cats on mainland vertebrates.Front. Ecol. Environ. 2017; 15: 502-509Crossref Scopus (88) Google Scholar, 3.Doherty T.S. et al.Impacts and management of feral cats Felis catus in Australia.Mammal Rev. 2017; 47: 83-97Crossref Scopus (97) Google Scholar] has led conservation advocates into clashes with cat enthusiasts, in controversies dramatically characterised as ‘Cat Wars’ [4.Marra P.P. Santella C. Cat Wars: The Devastating Consequences of a Cuddly Killer. Princeton University Press, 2016Google Scholar]. Debates about cats have indeed become combative and polarised, especially in the USA [4.Marra P.P. Santella C. Cat Wars: The Devastating Consequences of a Cuddly Killer. Princeton University Press, 2016Google Scholar], where cat and wildlife enthusiasts and their organisations level accusations at one another of making inflated or false claims about the ‘real’ impact of cats on wild animals [5.Loss S.R. Marra P.P. Merchants of doubt in the free-ranging cat conflict.Conserv. Biol. 2018; 32: 265-266Crossref PubMed Scopus (31) Google Scholar]. However, to demarcate and reinforce a neat division between ‘cat people’ and ‘wildlife people’ is to oversimplify the issue, and combative narratives may serve only to alienate the people best placed to address the challenges identified, i.e., the cat owners. Many cat owners are concerned about wildlife conservation or the welfare of prey animals, and people who identify themselves as conservationists often keep cats as pets [6.Balmford A. et al.The environmental footprints of conservationists, economists and medics compared.Biol. Conserv. 2017; 214: 260-269Crossref Scopus (24) Google Scholar]. Furthermore, the roles of cats in both human societies and wider ecosystems are complex and not reducible to claims that cats are either an innocent blessing or an invasive scourge. Here, we argue that the fraught contemporary relationships between cats and human societies reflect a longstanding, important symbiosis that is both maintained and challenged by the resistance of the ‘domestic’ cat to complete domestication, and by the duality of its roles as autonomous predator and ostensibly dependent companion. Modern domestic cats are descended from the Near Eastern wildcat Felis silvestris lybica [7.Driscoll C.A. et al.The Near Eastern origin of cat domestication.Science. 2007; 317: 519-523Crossref PubMed Scopus (367) Google Scholar]. It has been proposed that cats ‘self-domesticated’ through a fortuitous combination of ecological and sociocultural circumstance, whereby individual cats that tolerated humans were able to take advantage of the hunting and scavenging opportunities provided by early settlements [8.Driscoll C.A. et al.From wild animals to domestic pets, an evolutionary view of domestication.Proc. Natl. Acad. Sci. U. S. A. 2009; 106: 9971-9978Crossref PubMed Scopus (325) Google Scholar]. A degree of variation among people in their acceptance of cat presence is likely in this ‘self-domestication’, especially in light of more commonplace antagonistic relations between humans and sympatric carnivores. Cats seen killing increasingly problematic rodents around food stores would have been appreciated as low-maintenance pest controllers [9.Hu Y. et al.Earliest evidence for commensal processes of cat domestication.Proc. Natl. Acad. Sci. U. S. A. 2014; 111: 116-120Crossref PubMed Scopus (98) Google Scholar] and may have conferred significant advantages upon the individual people expressing tolerant behaviour towards them. The value of domestic cats as predators-in-residence certainly played a significant part in their global spread, as they were employed for rodent control on trade ships and in the outbuildings of emerging civilisations [10.Ottoni C. et al.The palaeogenetics of cat dispersal in the ancient world.Nat. Ecol. Evol. 2017; 1: 0139Crossref Scopus (81) Google Scholar]. The special and multiple roles of the cat in ancient Egypt are well known, with cats frequently depicted as commensals, pest controllers, religious icons, and sacrifices. It is also in Egypt that the role of the cat as a domestic companion emerged, with house cats clearly portrayed in artefacts from the Middle Kingdom [11.Faure E. Kitchener A.C. An archaeological and historical review of the relationship between felids and people.Anthrozoös. 2009; 22: 221-238Crossref Scopus (49) Google Scholar]. Genetic lineages with Egyptian origins have recently been identified as having contributed significantly to the international domestic cat gene pool, indicating that Egyptian cats enjoyed considerable popularity, potentially linked to desirable characteristics, such as increased tameness and sociality [10.Ottoni C. et al.The palaeogenetics of cat dispersal in the ancient world.Nat. Ecol. Evol. 2017; 1: 0139Crossref Scopus (81) Google Scholar]. While, in some regions, cats are still treated more as commensals than pets [12.Gray P.B. Young S.M. Human-pet dynamics in cross-cultural perspective.Anthrozoös. 2011; 24: 17-30Crossref Scopus (47) Google Scholar], companionship has become the cat’s primary role in many societies. As household pets, cats provide company, affection, and enjoyment for millions of people, and, in some cases, cat ownership may also confer health benefits (although evidence is mixed) [13.Amiot C. et al.People and companion animals: it takes two to tango.Bioscience. 2016; 66: 552-560Crossref Scopus (45) Google Scholar]. The expansion of the pet-food industry and increased availability of cheap protein has enabled growing numbers of people to provide for the demanding diet of an obligate carnivore, and the mid-20th century invention of cat litter has allowed urban residents to keep pets largely or wholly indoors. Nevertheless, today’s cats are unusual among domesticated species in that they (i) exhibit comparatively few morphological changes compared with their wild ancestors; and (ii), apparently unlike dogs [10.Ottoni C. et al.The palaeogenetics of cat dispersal in the ancient world.Nat. Ecol. Evol. 2017; 1: 0139Crossref Scopus (81) Google Scholar], have not been purposively selected to fulfil particular roles in human societies. Excepting pedigree breeds, most of which have been developed within the last 150 years [14.Montague M.J. et al.Comparative analysis of the domestic cat genome reveals genetic signatures underlying feline biology and domestication.Proc. Natl. Acad. Sci. U. S. A. 2014; 111: 17230-17235Crossref PubMed Scopus (192) Google Scholar], most cats continue to self-select through unregulated breeding. They remain capable of sustaining themselves without human assistance, as demonstrated by the global success of feral cats, both living in colonies near reliable food sources and reverting to predatory, solitary lifestyles (Box 1). Yet, modern cats are also well adapted to living in close association with humans, and most rely on human provisioning or anthropogenic environments (Figure 1). They are predisposed to form attachments with people during early developmental stages, tolerate the presence of humans, other cats, and other domestic animals far better than wilder felids, and exhibit distinctive behavioural traits (including vocalisations and body language) that facilitate effective interspecies communication [15.Turner D.C. Bateson P. The Domestic Cat: The Biology of its Behaviour. Cambridge University Press, 2013Crossref Scopus (19) Google Scholar]. Contemporary societies, especially in Europe, are often broadly tolerant of roaming pet cats, and suggestions that cats should be kept indoors can be strongly resisted (although, again, cat husbandry practices are culturally variable [15.Turner D.C. Bateson P. The Domestic Cat: The Biology of its Behaviour. Cambridge University Press, 2013Crossref Scopus (19) Google Scholar,16.Hall C.M. et al.Community attitudes and practices of urban residents regarding predation by pet cats on wildlife: an International comparison.PLoS ONE. 2016; 11e0151962Crossref PubMed Scopus (73) Google Scholar]). The territoriality of cats, and their attachment to place over people, may be the distinguishing feature that makes this unusual arrangement possible; they are independent, yet relatively self-restricting in their roaming behaviour.Box 1Feral CatsCats exist on a spectrum of human responsibility and control over their movement, feeding and reproduction (see Figure 1 in the main text). Cats living independently of human provisioning and behavioural management are usually called ‘feral’ (although this can also refer to any unowned cat). In Australia, distinguishing between companion, stray, and feral cats has been important for designating feral cats as pests (in some jurisdictions) and implementing management accordingly [36.Deak B.P. et al.The significance of social perceptions in implementing successful feral cat management strategies: a global review.Animals. 2019; 9: 617Crossref Scopus (13) Google Scholar]. Latest estimates that has a feral cat of S. et a many feral cats are in Conserv. 2017; Scopus Google Scholar]. the ecological impacts of cats on are increasingly well T.S. et al.Impacts and management of feral cats Felis catus in Australia.Mammal Rev. 2017; 47: 83-97Crossref Scopus (97) Google et in and Killer. Google et et in and Killer. Google Scholar] that feral cats have been a in at of and is a relatively level of public for feral cat management, control C.M. et al.Community attitudes and practices of urban residents regarding predation by pet cats on wildlife: an International comparison.PLoS ONE. 2016; 11e0151962Crossref PubMed Scopus (73) Google et in and Killer. Google Cats in the Environ. 2017; Scopus (13) Google Scholar] in However, in the there is significant public feral cat management [4.Marra P.P. Santella C. Cat Wars: The Devastating Consequences of a Cuddly Killer. Princeton University Press, 2016Google Scholar], in urban where may be Europe, and stray, and feral cat are often in their and, while for may not be by et challenges and conservation for urban cat management in Conserv. Biol. 2016; Scopus Google on and some of suggestions effectively cat not to feral Nevertheless, feral cat research and management should still be in a more ‘companion animal and often it is to feral from and cats can between over the of their urban are a and domestic cats can et al.The origins and expansion of feral cats in PubMed Scopus Google who value cats as may be as concerned for the and welfare of feral cats as for their pets C. et conservation management of feral domestic Ecol. Evol. 2019; PubMed Scopus Google Scholar]. in Australia, evidence some resistance to management of cats and about cat owners to be of both than Cats in the Environ. 2017; Scopus (13) Google Scholar]. The presence of free-ranging cats also for feral cats, to of ownership status in or the of or cats in and Cats exist on a spectrum of human responsibility and control over their movement, feeding and reproduction (see Figure 1 in the main text). Cats living independently of human provisioning and behavioural management are usually called ‘feral’ (although this can also refer to any unowned cat). In Australia, distinguishing between companion, stray, and feral cats has been important for designating feral cats as pests (in some jurisdictions) and implementing management accordingly [36.Deak B.P. et al.The significance of social perceptions in implementing successful feral cat management strategies: a global review.Animals. 2019; 9: 617Crossref Scopus (13) Google Scholar]. Latest estimates that has a feral cat of S. et a many feral cats are in Conserv. 2017; Scopus Google Scholar]. the ecological impacts of cats on are increasingly well T.S. et al.Impacts and management of feral cats Felis catus in Australia.Mammal Rev. 2017; 47: 83-97Crossref Scopus (97) Google et in and Killer. Google et et in and Killer. Google Scholar] that feral cats have been a in at of and is a relatively level of public for feral cat management, control C.M. et al.Community attitudes and practices of urban residents regarding predation by pet cats on wildlife: an International comparison.PLoS ONE. 2016; 11e0151962Crossref PubMed Scopus (73) Google et in and Killer. Google Cats in the Environ. 2017; Scopus (13) Google Scholar] in However, in the there is significant public feral cat management [4.Marra P.P. Santella C. Cat Wars: The Devastating Consequences of a Cuddly Killer. Princeton University Press, 2016Google Scholar], in urban where may be Europe, and stray, and feral cat are often in their and, while for may not be by et challenges and conservation for urban cat management in Conserv. Biol. 2016; Scopus Google Scholar]. on and some of suggestions effectively cat not to feral Nevertheless, feral cat research and management should still be in a more ‘companion animal and often it is to feral from and cats can between over the of their urban are a and domestic cats can et al.The origins and expansion of feral cats in PubMed Scopus Google Scholar]. who value cats as may be as concerned for the and welfare of feral cats as for their pets C. et conservation management of feral domestic Ecol. Evol. 2019; PubMed Scopus Google Scholar]. in Australia, evidence some resistance to management of cats and about cat owners to be of both than Cats in the Environ. 2017; Scopus (13) Google Scholar]. The presence of free-ranging cats also for feral cats, to of ownership status in or the of or cats in and Cats and humans have long from close the dynamics of the relationship have over It is that many people cats with a to a but feature of the maintained the role of cats as pest However, between people and cats are not a of most human an to cats [15.Turner D.C. Bateson P. The Domestic Cat: The Biology of its Behaviour. Cambridge University Press, 2013Crossref Scopus (19) Google Scholar], and, without cats not effectively with people Cat Scholar]. The relationship also for both living in close to cats human of with 2013; PubMed Scopus Google et with dogs and cats in 2016; PubMed Scopus Google Scholar], while humans their to cats Nevertheless, the relationship has and over of years and is best as a than an or The domestication of Scopus Google Scholar] (Figure cats are humans are some such as from which they have been often living at They are a of the of global pet and have and significance that and for by their and that cats have global environmental their for for animal the and of which have environmental impacts in of and and (although pet food may from of human et of pet Rev. 2013; 4: PubMed Scopus (49) Google Scholar]. cats can as and of with 2013; PubMed Scopus Google Scholar], from but to S. 2017; PubMed Scopus Google Scholar]. the significance of domestic cat impacts on wildlife on and and a are large numbers of free-ranging and feral cats will large numbers of and S.R. Marra P.P. Population impacts of free-ranging domestic cats on mainland vertebrates.Front. Ecol. Environ. 2017; 15: 502-509Crossref Scopus (88) Google T.S. et al.Impacts and management of feral cats Felis catus in Australia.Mammal Rev. 2017; 47: 83-97Crossref Scopus (97) Google et of wildlife by domestic cats Felis catus in Rev. Scopus Google Scholar]. Cats on to other can have on adapted such as et global review of the impacts of invasive cats on Biol. 2011; Scopus Google Scholar]. Domestic cats often in than other wild et and domestic cat in urban Scopus Google Scholar]. Cats can also have on the success of other through of and behaviour et impacts of invasive cats on not only a of Conserv. 2014; Scopus Google Scholar]. and to humans cats and their behaviour are important with for both and for ecosystems Cats a as both wild are largely and domestic have close with and on this duality is to understanding the social to which cats are of relationship and relatively early during the association of lybica with humans [8.Driscoll C.A. et al.From wild animals to domestic pets, an evolutionary view of domestication.Proc. Natl. Acad. Sci. U. S. A. 2009; 106: 9971-9978Crossref PubMed Scopus (325) Google C. et al.The palaeogenetics of cat dispersal in the ancient world.Nat. Ecol. Evol. 2017; 1: 0139Crossref Scopus (81) Google Scholar], yet for the traits roles they one To be effective pest controllers, cats require and hunting while to be humans cats that more behaviour and between the cat as and as are in human behaviour and attitudes towards Cat management is often by both animal welfare and wildlife with on the of the cat and its ownership in for unowned cats are to control as feral pet cats are but must be and (in some et in and Killer. Google Scholar]. are also in husbandry practices for cats kept as animals on where they often in outbuildings and are to and and as cats can be a to their and hunting In societies where pet cats are more popular in their hunting are increasingly more as a than a about behaviour are as that cats should be and that their are the responsibility of their owners. is also among cat about the welfare and of cats indoors to a that the and of wild independence and of domestic and et behaviour in domestic an of and responsibility among cat 2019; 1: Scopus Google Scholar]. However, the most contemporary the impact of cats on and is by two cat on the one of urban and other free-ranging cat and, on the wildlife conservation advocates in and, in Australia, advocates link with numbers of free-ranging and feral cats and to management Cat advocates argue that domestic cats are the for environmental changes to other anthropogenic The dual status of the species as both wild predator and domestic companion of this division among people. 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It is not only in public about management that research on cats between and animal that on largely and behavioural of health and and research in and conservation biology that hunting behaviour and environmental impacts. a for of research to be through and there are of for with both welfare and ecological feral cat or is also for research (see that and for the between wild and which cats that the of cats is certainly to their global and there is for an ‘companion animal ecology’ where anthropogenic are as to ecological processes, as in and urban in this take novel to the of roaming cats with other domestic animals, and human their and of and anthropogenic and their roles as and of The to research cat impacts and their that cat and behaviour are and by human cat owners as research will be to understanding of and to the and of sustainable solutions to the environmental challenges by the global of people and their research has provided important into of cat behaviour et behaviour in domestic an of and responsibility among cat 2019; 1: Scopus Google E. et towards impacts on Scopus Google Scholar, et to cat owners in Australia, to identify the and to cat PubMed Scopus Google Scholar, et of public on cat predation and the of cat management in 2017; Scopus Google Scholar] and the of for communication and behaviour et cat for a to wildlife Sci. 2019; Scopus Google Scholar, et the impact of on the and behaviour of cat a control 2017; PubMed Scopus Google Scholar, et communication for domestic cat 2017; Scopus (19) Google Scholar]. However, behaviour between cat owners and wildlife conservationists, and will be to understanding and and to on human to and for domestic environmental cat and must towards changes to practices and that will the relationship between people and cats a sustainable a ‘companion animal the domestic of companion animals with their wild and of other and unowned companion animals of cats and with humans, and other domestic animals, in a of ecological and social in their domestic husbandry the environmental impacts of companion animal between multiple of and social research be around or animal husbandry or cat and (i) roaming and hunting and the of companion species in animal and with significant for companion animal health and environmental companion animal and research and cat owners of other companion become research of owners in and research of and are and and for companion and practices will be to and can management best be between to is the role of and in sustainable management of companion pet and societies their individual and to other animals, both wild and a ‘companion animal the domestic of companion animals with their wild and of other and unowned companion animals of cats and with humans, and other domestic animals, in a of ecological and social in their domestic husbandry the environmental impacts of companion animal between multiple of and social research be around or animal husbandry or cat and (i) roaming and hunting and the of companion species in animal and with significant for companion animal health and environmental companion animal and research and cat owners of other companion become research of owners in and research of and are and and for companion and practices will be to and can management best be between to is the role of and in sustainable management of companion pet and societies their individual and to other animals, both wild and and and the provided review and The Cat and of is by are to of the for their on an of this

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