history of cats research paper

International research transforms understanding of the history of cats in Europe

• Ancient evidence and samples from modern cats analysed to reveal a typical cat story
• Research shows, domestic and wild cats ignored each other for 2,000 years
• Then, in the 1960s, they began to mix

An international team of researchers has found new archaeological and genetic evidence which transforms our understanding of the history of cats in Europe. Domestic cats introduced from the Near East and wildcats native to Europe did not mix until the 1960s, despite being exposed to each other for 2,000 years, according to two research papers published today in Current Biology .

The team sequenced and analysed both wild and domestic cats including 48 modern individuals and 258 ancient samples excavated from 85 archaeological sites over the last 8,500 years. They then assessed the patterns of hybridisation after domestic cats were introduced to Europe over 2,000 years ago and came into contact with native European wildcats.

Wildcats and domestic cats have only hybridised very recently. It is clear that hybridisation is a result of modern threats common to many of our native species Jo Howard-McCombe, University of Bristol

The results of the studies demonstrate that, since their introduction, domestic cats and European wildcats generally avoided mating. But, about 50 years ago in Scotland, that all changed. Perhaps as a result of dwindling wildcat populations and a lack of opportunity to mate with other wildcats, rates of interbreeding between wild and domestic cats rose rapidly.

Jo Howard-McCombe from the University of Bristol and the Royal Zoological Society of Scotland explains, ‘Wildcats and domestic cats have only hybridised very recently. It is clear that hybridisation is a result of modern threats common to many of our native species. Habitat loss and persecution have pushed wildcats to the brink of extinction in Britain. It is fascinating that we can use genetic data to look back at their population history, and use what we have learnt to protect Scottish Wildcats.’

Professor Greger Larson , from the University of Oxford, says, ‘We tend to think of cats and dogs as very different. Our data suggests that, at least with respect to avoiding interbreeding with their wild counterparts, dogs and cats are much more similar to each other than they are to all other domestic animals. Understanding why this is true will be fun to explore.’

We tend to think of cats and dogs as very different. Our data suggests that...dogs and cats are much more similar to each other than they are to all other domestic animals Professor Greger Larson, University of Oxford's Schools of Archaeology

Professor Mark Beaumont , from the University of Bristol, adds, ‘The nature of the Scottish wildcat and its relation to feral domestic cats has long been a mystery. Modern molecular methods and mathematical modelling have helped to provide an understanding of what the Scottish wildcat truly is, and the threats that have led to its decline.’

Domestic animals including cattle, sheep, goats, dogs, and pigs have been closely associated with people since the emergence of farming communities more than 10,000 years ago. These tight relationships led to the human-mediated dispersal of plants and animals well beyond their native ranges.

The nature of the Scottish wildcat and its relation to feral domestic cats has long been a mystery. Modern molecular methods and mathematical modelling have helped to provide an understanding of what the Scottish wildcat truly is Professor Mark Beaumont, University of Bristol

Over the last decade, genomic sequences of modern and ancient individuals have revealed that, as domestic animals moved into new regions, they interbred with closely related wild species, which has dramatically altered their genomes. This pattern has been seen in every domestic animal, except dogs. It would be fascinating to know whether domestic cats interbred with European wildcats, but the decline of native wildcat populations across Europe, and the lack of ancient cat genomes has made it difficult to do so.

The two studies, were carried out at universities in Munich, Bristol, Oxford and the Royal Zoological Society of Scotland.

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June 1, 2009

13 min read

The Evolution of House Cats

Genetic and archaeological findings hint that wildcats became house cats earlier--and in a different place--than previously thought

By Carlos A. Driscoll , Juliet Clutton-Brock , Andrew C. Kitchener & Stephen J. O'Brien

It is by turns aloof and affectionate, serene and savage, endearing and exasperating. Despite its mercurial nature, however, the house cat is the most popular pet in the world. A third of American households have feline members, and more than 600 million cats live among humans worldwide. Yet as familiar as these creatures are, a complete understanding of their origins has proved elusive. Whereas other once wild animals were domesticated for their milk, meat, wool or servile labor, cats contribute virtually nothing in the way of sustenance or work to human endeavor. How, then, did they become commonplace fixtures in our homes?

Scholars long believed that the ancient Egyptians were the first to keep cats as pets, starting around 3,600 years ago. But genetic and archaeological discoveries made over the past five years have revised this scenario—and have generated fresh insights into both the ancestry of the house cat and how its relationship with humans evolved.

Cat’s Cradle The question of where house cats first arose has been challenging to resolve for several reasons. Although a number of investigators suspected that all varieties descend from just one cat species— Felis silvestris , the wildcat—they could not be certain. In addition, that species is not confined to a small corner of the globe. It is represented by populations living throughout the Old World—from Scotland to South Africa and from Spain to Mongolia—and until recently scientists had no way of determining unequivocally which of these wildcat populations gave rise to the tamer, so-called domestic kind. Indeed, as an alternative to the Egyptian origins hypothesis, some researchers had even proposed that cat domestication occurred in a number of different locations, with each domestication spawning a different breed. Confounding the issue was the fact that members of these wildcat groups are hard to tell apart from one another and from feral domesticated cats with so-called mackerel-tabby coats because all of them have the same pelage pattern of curved stripes and they interbreed freely with one another, further blurring population boundaries.

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In 2000 one of us (Driscoll) set out to tackle the question by assembling DNA samples from some 979 wildcats and domestic cats in southern Africa, Azerbaijan, Kazakhstan, Mongolia and the Middle East. Because wildcats typically defend a single territory for life, he expected that the genetic composition of wildcat groups would vary across geography but remain stable over time, as has occurred in many other cat species. If regional indigenous groups of these animals could be distinguished from one another on the basis of their DNA and if the DNA of domestic cats more closely resembled that of one of the wildcat populations, then he would have clear evidence for where domestication began.

In the genetic analysis, published in 2007, Driscoll, another of us (O’Brien) and their colleagues focused on two kinds of DNA that molecular biologists traditionally examine to differentiate subgroups of mammal species: DNA from mitochondria, which is inherited exclusively from the mother, and short, repetitive sequences of nuclear DNA known as microsatellites. Using established computer routines, they assessed the ancestry of each of the 979 individuals sampled based on their genetic signatures. Specifically, they measured how similar each cat’s DNA was to that of all the other cats and grouped the animals having similar DNA together. They then asked whether most of the animals in a group lived in the same region.

The results revealed five genetic clusters, or lineages, of wildcats. Four of these lineages corresponded neatly with four of the known subspecies of wildcat and dwelled in specific places: F. silvestris silvestris in Europe, F. s. bieti in China, F. s. ornata in Central Asia and F. s. cafra in southern Africa. The fifth lineage, however, included not only the fifth known subspecies of wildcat— F. s. lybica in the Middle East—but also the hundreds of domestic cats that were sampled, including purebred and mixed-breed felines from the U.S., the U.K. and Japan. In fact, genetically, F. s. lybica wildcats collected in remote deserts of Israel, the United Arab Emirates and Saudi Arabia were virtually indistinguishable from domestic cats. That the domestic cats grouped with F. s. lybica alone among wildcats meant that domestic cats arose in a single locale, the Middle East, and not in other places where wildcats are common.

Once we had figured out where house cats came from, the next step was to ascertain when they had become domesticated. Geneticists can often estimate when a particular evolutionary event occurred by studying the quantity of random genetic mutations that accumulate at a steady rate over time. But this so-called molecular clock ticks a mite too slowly to precisely date events as recent as the past 10,000 years, the likely interval for cat domestication. To get a bead on when the taming of the cat began, we turned to the archaeological record. One recent find has proved especially informative in this regard.

In 2004 Jean-Denis Vigne of the National Museum of Natural History in Paris and his colleagues reported unearthing the earliest evidence suggestive of humans keeping cats as pets. The discovery comes from the Mediterranean island of Cyprus, where 9,500 years ago an adult human of unknown gender was laid to rest in a shallow grave. An assortment of items accompanied the body—stone tools, a lump of iron oxide, a handful of seashells and, in its own tiny grave just 40 centimeters away, an eight-month-old cat, its body oriented in the same westward direction as the human’s.

Because cats are not native to most Mediterranean islands, we know that people must have brought them over by boat, probably from the adjacent Levantine coast. Together the transport of cats to the island and the burial of the human with a cat indicate that people had a special, intentional relationship with cats nearly 10,000 years ago in the Middle East. This locale is consistent with the geographic origin we arrived at through our genetic analyses. It appears, then, that cats were being tamed just as humankind was establishing the first settlements in the part of the Middle East known as the Fertile Crescent.

A Cat and Mouse Game? With the geography and an approximate age of the initial phases of cat domestication established, we could begin to revisit the old question of why cats and humans ever developed a special relationship. Cats in general are unlikely candidates for domestication. The ancestors of most domesticated animals lived in herds or packs with clear dominance hierarchies. (Humans unwittingly took advantage of this structure by supplanting the alpha individual, thus facilitating control of entire cohesive groups.) These herd animals were already accustomed to living cheek by jowl, so provided that food and shelter were plentiful, they adapted easily to confinement.

Cats, in contrast, are solitary hunters that defend their home ranges fiercely from other cats of the same sex (the pride-living lions are the exception to this rule). Moreover, whereas most domesticates feed on widely available plant foods, cats are obligate carnivores, meaning they have a limited ability to digest anything but meat—a far rarer menu item. In fact, they have lost the ability to taste sweet carbohydrates altogether. And as to utility to humans, let us just say cats do not take instruction well. Such attributes suggest that whereas other domesticates were recruited from the wild by humans who bred them for specific tasks, cats most likely chose to live among humans because of opportunities they found for themselves.

Early settlements in the Fertile Crescent between 9,000 and 10,000 years ago, during the Neolithic period, created a completely new environment for any wild animals that were sufficiently flexible and inquisitive (or scared and hungry) to exploit it. The house mouse, Mus musculus domesticus , was one such creature. Archaeologists have found remains of this rodent, which originated in the Indian subcontinent, among the first human stores of wild grain from Israel, which date to around 10,000 years ago. The house mice could not compete well with the local wild mice outside, but by moving into people’s homes and silos, they thrived.

It is almost certainly the case that these house mice attracted cats. But the trash heaps on the outskirts of town were probably just as great a draw, providing year-round pickings for those felines resourceful enough to seek them out. Both these food sources would have encouraged cats to adapt to living with people; in the lingo of evolutionary biology, natural selection favored those cats that were able to cohabitate with humans and thereby gain access to the trash and mice.

Over time, wildcats more tolerant of living in human-dominated environments began to proliferate in villages throughout the Fertile Crescent. Selection in this new niche would have been principally for tameness, but competition among cats would also have continued to influence their evolution and limit how pliant they became. Because these proto–domestic cats were undoubtedly mostly left to fend for themselves, their hunting and scavenging skills remained sharp. Even today most domesticated cats are free agents that can easily survive independently of humans, as evinced by the plethora of feral cats in cities, towns and countrysides the world over.

Considering that small cats do little obvious harm, people probably did not mind their company. They might have even encouraged the cats to stick around when they saw them dispatching mice and snakes. Cats may have held other appeal, too. Some experts speculate that wildcats just so happened to possess features that might have preadapted them to developing a relationship with people. In particular, these cats have “cute” features—large eyes, a snub face and a high, round forehead, among others—that are known to elicit nurturing from humans. In all likelihood, then, some people took kittens home simply because they found them adorable and tamed them, giving cats a first foothold at the human hearth.

Why was F. s. lybica the only subspecies of wild­cat to be domesticated? Anecdotal evidence suggests that certain other subspecies, such as the European wildcat and the Chinese mountain cat, are less tolerant of people. If so, this trait alone could have precluded their adoption into homes. The friendlier southern African and Central Asian wildcats, on the other hand, might very well have become domesticated under the right conditions. But F. s. lybica had the advantage of a head start by virtue of its proximity to the first settlements. As agriculture spread out from the Fertile Crescent, so, too, did the tame scions of F. s. lybica , filling the same niche in each region they entered—and effectively shutting the door on local wildcat populations. Had domestic cats from the Near East never arrived in Africa or Asia, perhaps the indigenous wildcats in those regions would have been drawn to homes and villages as urban civilizations developed.

Rise of the Goddess We do not know how long the transformation of the Middle Eastern wildcat into an affectionate home companion took. Animals can be domesticated quite rapidly under controlled conditions. In one famous experiment, begun in 1959, Russian scientists using highly selective breeding produced tame silver foxes from wild ones in just 40 years. But without doors or windowpanes, Neolithic farmers would have been hard-pressed to control the breeding of cats even if they wanted to. It seems reasonable to suggest that the lack of human influence on breeding and the probable intermixing of house cats and wildcats militated against rapid taming, causing the metamorphosis to occur over thousands of years.

Although the exact timeline of cat domestication remains uncertain, long-known archaeological evidence affords some insight into the process. After the Cypriot find, the next oldest hints of an association between humans and cats are a feline molar tooth from an archaeological deposit in Israel dating to roughly 9,000 years ago and another tooth from Pakistan dating to around 4,000 years ago.

Testament to full domestication comes from a much later period. A nearly 3,700-year-old ivory cat statuette from Israel suggests the cat was a common sight around homes and villages in the Fertile Crescent before its introduction to Egypt. This scenario makes sense, given that all the other domestic animals (except the donkey) and plants were introduced to the Nile Valley from the Fertile Crescent. But it is Egyptian paintings from the so-called New Kingdom period—Egypt’s golden era, which began nearly 3,600 years ago—that provide the oldest known unmistakable depictions of full domestication. These paintings typically show cats poised under chairs, sometimes collared or tethered, and often eating from bowls or feeding on scraps. The abundance of these illustrations signifies that cats had become common members of Egyptian households by this time.

It is in large part as a result of evocative images such as these that scholars traditionally perceived ancient Egypt as the locus of cat domestication. Even the oldest Egyptian representations of wildcats are 5,000 to 6,000 years younger than the 9,500-year-old Cypriot burial, however. Although ancient Egyptian culture cannot claim initial domestication of the cat among its many achievements, it surely played a pivotal role in subsequently molding the domestication dynamic and spreading cats throughout the world. Indeed, the Egyptians took the love of cats to a whole new level. By 2,900 years ago the domestic cat had become the official deity of Egypt in the form of the goddess Bastet, and house cats were sacrificed, mummified and buried in great numbers at Bastet’s sacred city, Bubastis. Measured by the ton, the sheer number of cat mummies found there indicates that Egyptians were not just harvesting feral or wild populations but, for the first time in history, were actively breeding domestic cats.

Egypt officially prohibited the export of their venerated cats for centuries. Nevertheless, by 2,500 years ago the animals had made their way to Greece, proving the inefficacy of export bans. Later, grain ships sailed directly from Alexandria to destinations throughout the Roman Empire, and cats are certain to have been onboard to keep the rats in check. Thus introduced, cats could have established colonies in port cities and then fanned out from there. By 2,000 years ago, when the Romans were expanding their empire, domestic cats were traveling with them and becoming common throughout Europe. Evidence for their spread comes from the German site of Tofting in Schleswig, which dates to between the 4th and 10th centuries, as well as increasing references to cats in art and literature from that period. (Oddly, domestic cats seem to have reached the British Isles before the Romans brought them over—a dispersal that researchers cannot yet explain.)

Meanwhile, on the opposite side of the globe, domestic cats had presumably spread to the Orient almost 2,000 years ago, along well-established trade routes between Greece and Rome and the Far East, reaching China by way of Mesopotamia and arriving in India via land and sea. Then something interesting happened. Because no native wildcats with which the newcomers could interbreed lived in the Far East, the Oriental domestic cats soon began evolving along their own trajectory. Small, isolated groups of Oriental domestics gradually acquired distinctive coat colors and other mutations through a process known as genetic drift, in which traits that are neither beneficial nor maladaptive become fixed in a population.

This drift led to the emergence of the Korat, the Siamese, the Birman and other “natural breeds,” which were described by Thai Buddhist monks in a book called the Tamara Maew (meaning “Cat-Book Poems”) that may date back to 1350. The putative antiquity of these breeds received support from the results of genetic studies announced last year, in which Marilyn Menotti-Raymond of the National Cancer Institute and Leslie Lyons of the University of California, Davis, found DNA differences between today’s European and Oriental domestic cat breeds indicative of more than 700 years of independent cat breeding in Asia and Europe.

As to when house cats reached the Americas, little is known. Christopher Columbus and other seafarers of his day reportedly carried cats with them on transatlantic voyages. And voyagers onboard the Mayflower and residents of Jamestown are said to have brought cats with them to control vermin and to bring good luck. How house cats got to Australia is even murkier, although researchers presume that they arrived with European explorers in the 1600s. Our group at the U.S. National Institutes of Health is tackling the problem using DNA.

Breeding for Beauty Although humans might have played some minor role in the development of the natural breeds in the Orient, concerted efforts to produce novel breeds did not begin until relatively recently. Even the Egyptians, who we know were breeding cats extensively, do not seem to have been selecting for visible traits, probably because distinctive variants had not yet arisen: in their paintings, both wildcats and house cats are depicted as having the same mackerel-tabby coat. Experts believe that most of the modern breeds were developed in the British Isles in the 19th century, based on the writings of English natural history artist Harrison Weir. And in 1871 the first proper fancy cat breeds—breeds created by humans to achieve a particular appearance—were displayed at a cat show held at the Crystal Palace in London (a Persian won, although the Siamese was a sensation).

Today the Cat Fancier’s Association and the International Cat Association recognize nearly 60 breeds of domestic cat. Just a dozen or so genes account for the differences in coat color, fur length and texture, as well as other, subtler coat characteristics, such as shading and shimmer, among these breeds.

Thanks to the sequencing of the entire genome of an Abyssinian cat named Cinnamon in 2007, geneticists are rapidly identifying the mutations that produce such traits as tabby patterning, black, white and orange coloring, long hair and many others. Beyond differences in the pelage-related genes, however, the genetic variation between domestic cat breeds is very slight—comparable to that seen between adjacent human populations, such as the French and the Italians.

The wide range of sizes, shapes and temperaments seen in dogs—consider the Chihuahua and Great Dane—is absent in cats. Felines show much less variety because, unlike dogs—which starting in prehistoric times were bred for such tasks as guarding, hunting and herding—wildcats were under no such selective breeding pressures. To enter our homes, they had only to evolve a people-friendly disposition.

So are today’s cats truly domesticated? Well, yes—but perhaps only just. Although they satisfy the criterion of tolerating people, most domestic cats are feral and do not rely on people to feed them or to find them mates. And whereas other domesticates, like dogs, look quite distinct from their wild ancestors, the average domestic cat largely retains the wild body plan. It does exhibit a few morphological differences, however—namely, slightly shorter legs, a smaller brain and, as Charles Darwin noted, a longer intestine, which may have been an adaptation to scavenging kitchen scraps.

The house cat has not stopped evolving, though—far from it. Armed with artificial insemination and in vitro fertilization technology, cat breeders today are pushing domestic cat genetics into uncharted territory: they are hybridizing house cats with other felid species to create exotic new breeds. The Bengal and the Caracat, for example, resulted from crossing the house cat with the Asian leopard cat and the caracal, respectively. The domestic cat may thus be on the verge of an unprecedented and radical evolution into a multispecies composite  whose future can only be imagined.

The Truth about Cats and Dogs Unlike dogs, which exhibit a huge range of sizes, shapes and temperaments, house cats are relatively homogeneous, differing mostly in the characteristics of their coats. The reason for the relative lack of variability in cats is simple: humans have long bred dogs to assist with particular tasks, such as hunting or sled pulling, but cats, which lack any inclination for performing most tasks that would be useful to humans, experienced no such selective breeding pressures.

Note: This article was originally published with the title, "The Taming of the Cat".

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• Published: 09 July 2020

The earliest domestic cat on the Silk Road

• A. F. Haruda   ORCID: orcid.org/0000-0001-9660-3954 1 , 2 ,
• A. R. Ventresca Miller   ORCID: orcid.org/0000-0003-4148-4016 3 , 4 , 5 , 6 ,
• J. L. A. Paijmans   ORCID: orcid.org/0000-0002-1938-7052 7 , 8 ,
• A. Barlow   ORCID: orcid.org/0000-0002-5532-9458 9 ,
• A. Tazhekeyev 10 ,
• S. Bilalov 10 , 11 ,
• Y. Hesse   ORCID: orcid.org/0000-0002-9854-8730 7 ,
• M. Preick   ORCID: orcid.org/0000-0002-8014-1975 7 ,
• T. King 8 , 12 ,
• R. Thomas 12 ,
• H. Härke 13 , 14 &
• I. Arzhantseva 14 , 15  

Scientific Reports volume  10 , Article number:  11241 ( 2020 ) Cite this article

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• Archaeology
• Genetic variation
• Stable isotope analysis

We present the earliest evidence for domestic cat ( Felis catus L., 1758) from Kazakhstan, found as a well preserved skeleton with extensive osteological pathologies dating to 775–940 cal CE from the early medieval city of Dzhankent, Kazakhstan. This urban settlement was located on the intersection of the northern Silk Road route which linked the cities of Khorezm in the south to the trading settlements in the Volga region to the north and was known in the tenth century CE as the capital of the nomad Oghuz. The presence of this domestic cat, presented here as an osteobiography using a combination of zooarchaeological, genetic, and isotopic data, provides proxy evidence for a fundamental shift in the nature of human-animal relationships within a previously pastoral region. This illustrates the broader social, cultural, and economic changes occurring within the context of rapid urbanisation during the early medieval period along the Silk Road.

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Introduction.

Burials of domestic and wild cats remain a rarity in the archaeological record, especially in comparison to dogs, which are recovered so frequently that they have their own depositional typologies 1 , 2 , 3 , 4 , 5 . Finds of individual, articulated animals in archaeological contexts, particularly those exhibiting osteopathologies, are ideal for a social zooarchaeological approach which encourages the investigation of human animal interaction as a facet of social and cultural structures beyond productive capacity 6 . This type of integrated social-cultural approach in archaeology has largely been utilised in pre- and protohistoric contexts in which there are limited written sources in order to access and interpret the worldview of past cultures. However, there is value in utilising this framework in historical contexts as some animals, such as the domestic cat, are often not frequently and explicitly mentioned in the written record. Such animals act as signifiers of changing cultural attitudes and evidence of human-mediated species dispersal that accompanied expanding human connectivity through the end of the first millennium CE 7 , 8 . The unique and rare find of the remains of a small felid, found in the medieval urban center of Dzhankent in Kazakhstan, is ripe for this approach. Below, we combine multiple lines of scientific enquiry to: explore the ancestry of this cat and ascertain its domestic status; characterise its diet in comparison to other fauna; and reconstruct its life history. Thus, this osteobiography reflects changing human attitudes towards animals that are part of bigger social and cultural changes taking place along the medieval Silk Road.

In the semi-arid and arid steppe of Central Asia, particularly in modern day Kazakhstan, domestic cats were not widespread before the colonial period of the eighteenth and nineteenth centuries. There are no published archaeological remains of domestic felines from any prehistoric archaeological sites in the Kazakh steppe 9 , 10 , 11 , 12 despite the fact that there is increasing evidence for sedentary settlements and agricultural activity dating from the Late and Final Bronze Age within this region which would encourage the presence and utility of commensal rodent hunters such as cats 13 , 14 , 15 . Along the southern border of the Central Asian steppe, in Uzbekistan and Turkmenistan, single bones from felines of uncertain domesticated status have been found in Bronze Age urban contexts, such as at the sites of Kavat 3 and Ulug Depe, as well as in Iron Age contexts from Geoktchick Depe and Kyzyltepe (Fig.  1 ) 16 , 17 . Remains of small felines are similarly sparse throughout the Archaic and Antique periods in this region (fourth century BCE to eighth century CE) and only single bones were found at urban sites such as at Kugaitepe, Tukatepe, Tok-Kala, Toprak-Kala, Koi Krilgan Kala, Kuruk Kala and at the Gorgon Gates 16 , 17 , 18 , 19 . Osteological remains of cats dramatically increase from 1500 CE, and are found in much greater numbers at the urban site of Kunya-Urgench 20 . Indeed, there is a similar paucity of feline remains in periods prior to 1000 CE to the north in modern day Russia, after which cat remains are more commonly found at sites such as Novgorod and Tver, north of Moscow, and closer to the steppe boundary in Kazan and Toretsk 21 , 22 , 23 .

Location of Dzhankent and other archaeological sites mentioned in the text. Sites marked with squares are contemporaneous urban centres. Map generated by authors in qGIS ( www.qgis.org ) and Adobe Illustrator ( www.adobe.com ); basemap data Openstreet ( www.openstreetmap.org ), adapted under a CC BY-SA 2.0 license ( https://creativecommons.org/licenses/by-sa/2.0/ ), boundaries vector file DCW ( https://worldmap.harvard.edu/data/geonode:Digital_Chart_of_the_World?fbclid=IwAR2aFoqB1oWgay4fFXp0IifLZvsJobf0zaMYhYJo1H7TD5JHrX-RmaAA_1g ).

Felines are most frequently found in archaeological contexts in this region not as osteological remains but as artistic ornaments, and feature prominently in the iconography of animal style art, typical of Iron Age steppe cultures of the late first millennium BCE in Kazakhstan, Tuva, and Mongolia. In particular, motifs of predatory felines in twisting poses found in metalwork and in body art were likely inspired by large cats with a natural range across Central Asia and Persia, such as the snow leopard ( Panthera unica ), leopard ( Panthera pardus ), Eurasian lynx ( Lynx lynx), lion ( Panthera leo ), and caracal ( Caracal caracal ) 24 , 25 , 26 . Large feline predators were also depicted in the early first millennium CE in Persia, as seen on rhytons dating to the Parthian period (The Metropolitan Museum of Art, New York, Cat.: 1979.447) and much later in metal objects from the twelfth and thirteenth centuries CE 27 . Conversely, a number of smaller wild felids, such as the manul ( Otocolobus manul ), jungle cat ( Felis chaus ), sand cat ( Felis margarita ), Chinese mountain cat ( Felis bieti ), and the local subspecies of wild cat, ( Felis lybica ornata ), are not similarly represented in material cultures of the region and there are no published zooarchaeological finds of these taxa. Furthermore, the palaeogeograhic dispersal of these species within the medieval period remains uncertain 25 . The earliest historical mention of domestic cats originates from Persia and dates to the sixth century CE, when it was specifically noted that women kept cats as pets, dying their fur, adorning them with jewellery, and letting them sleep in their beds. In contrast, historians note that Zoroastrians from the same region held cats in contempt, associating them with the dark magic of the jinn and condemning the pungent odour of their urine 28 . These widely divergent attitudes in multicultural and cosmopolitan urban spheres of Persia, Khorasan, and Khorezm, indicates that domestic cats were widespread within human settlements and not just valued for their economic value as mousers, but also anthropomorphised and kept as pets.

Urbanisation is not a feature commonly associated with steppe societies, although there is evidence for dense conurbations in the central semi-arid steppe, dating from the Late and Final Bronze Age (c. 1500 BCE), such as Kent 29 . Steppe cultures are often characterised by a relatively dispersed settlement pattern and are largely agro-pastoralist with a component of mobility associated with seasonal resource exploitation, particularly in arid ecologies and have a distinct lack of felid remains. Instead, domestic cats are generally associated with urban agricultural contexts that proliferated across the medieval world, as they prey upon rodent and bird commensals associated with grain stores, and it is puzzling that these animals have not been found across the steppe as agricultural activity certainly was occurring, albeit at a small scale, throughout prehistorical and historical contexts 7 , 15 . Larger and more permanent settlements are characteristic of the oases and delta of the Amu Darya which belong to the Persian cultural sphere. Small defensive fortifications of the Dhzety-Asar culture first appear in the delta of the Syr Darya in the Late Iron Age/Antique periods (first to seventh centuries CE). Dzhankent was an early medieval city and one of three ‘marsh-towns’ now shown to have been founded in the seventh century CE 30 . It was located in the Syr Darya delta on a palaeochannel and close to a former inlet on the northeastern shore of the Aral Sea. The fortification of this urban site, increasing population density, and craft activities have been dated to the late ninth or tenth century CE, which is contemporary to the increasing proliferation of urban sites across Eurasia 31 . The origin of the trend of increasing density of human settlement remains uncertain, but is probably connected with intensifying long-distance trade between Khorezm and populations to the north and northwest in modern day Russia along branches of the Silk Road 31 .

Recent excavations have shown that the city of Dzhankent was a walled settlement with planned rectangular layout, with a citadel located in the northwestern corner 32 . Material culture dating from the seventh to twelfth centuries CE includes elements of intrusive nomadic Oghuz culture from the eastern steppes, the previous Dhzety-Asar culture, and neighbouring Khorezm, implying a mixture of nomadic and sedentary ‘urban’ cultures with ties to both the steppe to the north and urban cultures to the south. The Oghuz are reported as controlling the city and ruled this region until the eleventh century CE, when a significant proportion of the ruling classes were pushed south and reformed as the Seljuk Turks, conquering Persia and eventually most of the Middle East in the twelfth century 33 .

Zooarchaeological recovery and analysis

A partially articulated cat skeleton was found in a midden context at the juncture of the citadel and city walls in the fill of an abandoned house, specifically in Trench 2, Quadrant 103/99, 446–487 cm below the top of the preserved citadel wall (Fig.  2 ) 32 . The context was characterised by abundant animal and fish bone within a loose light brown clayey silt matrix. A complete skull and left and right mandible, as well as a left scapula and humerus, right humerus, ulna and radius, left pelvis, right femur, left tibia, and four vertebrae were recovered from a single individual feline (Fig.  3 , Supplementary Table 1 ). The bones were well preserved, with little evidence for weathering or post-mortem modification. These remains are categorised as an expedient burial as there is no clear grave cut and little evidence of ritual or sacred function 2 . Radiocarbon dating of the cat femur produced an uncalibrated date of 1,171 ± 15 BP (GrM-12987), with a 95% calibrated date range of 775–940 CE 34 , 35 , 36 .

Site Plan of Dzhankent with location of recovered osteological material marked by a red cross.

Skeletal elements present shaded; all recovered elements have recorded palaeopathologies. Left inset: Close up of articular fracture on left humerus. Right inset: Xray and view of 3D model of right femur.

Using epiphyseal fusion and tooth eruption, this cat was at least 58 weeks old at death; however, the presence of age-related osteological pathologies across the skeleton indicate that this cat was likely older than this lower limit estimate 37 , 37 , 38 . Comparison of greatest length data of limb bones against published contemporaneous domestic cats from Russian archaeological contexts, modern British domestic and wildcats with sexing data, as well as published standards for European domestic and wild cats indicate that this animal was a similar size to an intact (uncastrated) male domestic cat, but smaller than a European wildcat (Supplementary Table 2 ) 23 , 39 , 40 . A macroscopic examination of morphological characteristics and comparison to modern comparative zooarchaeological collections housed at the School of Archaeology and Ancient History at the University of Leicester as well as regionally specific comparative literature was inconclusive for identifying this specimen to species as extensive pathologies obscured qualitative discriminative features 41 , 42 .

Genetics: identifying ancestry

Recent genetic research has indicated that all modern domestic cat populations originate from the African wildcat, Felis lybica lybica 7 , 8 , 25 , 43 , 44 . However, disentangling the genetic structure of domestic cats and wildcats is challenging, as there is ongoing hybridisation between lineages 8 , 43 , 45 . Central Asian wildcats ( F. l. ornata ) have been reported to have a high frequency of African wildcat mitochondrial haplotypes (up to half of the sampled cats 43 ), making mitochondrial DNA unsuitable for ancestry determination. In contrast to mitochondrial DNA, nuclear data has revealed a clearer population structure, with the wildcats and domesticated cats forming distinct nuclear clades 8 , 43 . Therefore, we employed genetic analysis of nuclear DNA to explore the ancestry of the Dzhankent specimen.

DNA was extracted from the petrous bone and converted to single-stranded sequencing libraries using dedicated ancient DNA protocols 46 , 47 , 48 . Approximately 52 M reads were generated, of which 48% failed to pass the minimum length filter (< 30 bp). Of the remaining reads, 79% reads could be mapped to the domestic cat ( Felis catus ) reference genome 49 , suggesting a high endogenous content for the petrous bone of this specimen. This is in line with previous results from the petrous bones of humans 50 and cave bears 51 . The average coverage for the autosomes is approximately 0.38 – 0.43x, and 0.22 × for the X-chromosome assembly: this reduced coverage for the X-chromosome is a strong indication that the Dzhankent individual was male. The mitochondrial haplotype of the Dzankent cat falls within a clade containing domestic cat ( F. l. lybica ) haplotypes (Supplementary Fig. 1 ), but this provides weak evidence of ancestry. To further investigate this, we carried out population clustering analysis based on low coverage nuclear genome data of the Dzhankent cat and publicly available datasets representing multiple cat species. This analysis shows a clear separation of taxa 8 , 43 (Fig.  4 a; Supplementary Table 3 ), with the largest component representing more than 35% of the variation in the filtered data. The Dzhankent cat specimen clusters with the domestic cats (Fig.  4 a). A second analysis of wild European wildcat ( F. silvestris ) and domestic cats ( F. catus ) shows the same clustering (Fig.  4 b). Phylogenetic analysis of the whole nuclear genomes supports the monophyly of the domestic cats including the Dzhankent cat, which together form a sister group to the European wildcat genome (Fig.  4 c).

Principal component analysis of multi-species genomic data ( A ) and domestic and wildcat genomic data ( B ; Supplementary Table 3 ), as well as a neighbour-joining phylogeny of the nuclear genomes of domestic cats, the Dzankent cat, and a wildcat ( C ). Analyses are based on 928,665 ( A ), 764,913 ( B ) and 727,798 ( C ) variable positions after filtering (see Methods ). Axis labels in A and B indicate the percentage of variance explained by each component. The neighbour-joining phylogeny is rooted with the black-footed cat ( F. nigripes ) as outgroup (not shown). Colours reflect the (sub)species as indicated in the legends.

Unfortunately, at the time of this project, no nuclear shotgun genome data was available from the Central Asian wildcat F. l. ornata , preventing a direct comparison between the modern Central Asian wildcat data and the ancient Dzankent cat. Previous studies of nuclear data (short tandem repeats and microsatellites 43 ) collected from modern cats do suggest that the Central Asian wildcat is genetically distinct from the domesticated cat, as well as from other cat lineages. Under this assumption that Central Asian wildcats and domestic cats are genetically distinct at the whole genome level, the clustering of the Dzhankent cat with domestic cats in the principal component analysis suggests that the individual has a higher affinity with the population that was domesticated in the Near East. Long-distance transportation events have been suggested previously by the presence of F. l. ornata mtDNA haplotypes in Turkey (Ottoni et al., 2017), and it is possible that the Dzankent cat may have also been moved north from the Middle East.

It should be noted that by increasing the number of analysed genetic markers from a handful of markers in previous studies to several hundreds of thousands using whole nuclear genome data, future analyses may reveal extensive admixture and incomplete lineage sorting between domestic cats and other Asian wildcats. Comprehensive geographic sampling of Asian wildcats will thus be required to assess the genetic structure of domestic cats and the Asian wildcat, and conclusively verify the assignment of the Dzhankent cat to F. l. lybica / F. catus.

Stable isotopes: feline diet, ecological context, and urban provisioning

Carbon and nitrogen stable isotopes have routinely been used to reconstruct human and animal diets in Eurasia 52 , 53 , 54 , 55 . There is also a growing body of evidence regarding the dietary intake of domesticated animals, specifically those that may have served as companions (such as dogs and cats) or eaten human refuse 56 , 57 , 58 . To clarify the dietary intake of the feline under study, we analyzed stable carbon (δ 13 C) and nitrogen (δ 15 N) isotopes of fauna (n = 34) from the site of Dzhankent and compared our results to previously published isotope values from the broader region (Fig.  5 ).

Scatterplot of stable isotope values of fauna from Dzhankent and other Asian archaeological sites.

The cat was sampled at the medial fibula, which ossifies in the first months of life, and has a δ 13 C value of − 15.1‰ and a δ 15 N of 14.4‰, while the dog from the site has values of − 14.2‰ and 9.0‰, respectively 59 . Livestock (camels, cattle, sheep, goat, horses) from Dzhankent have a wide range of δ 13 C values, from − 19.6 to − 14.7‰ (average value of − 17.4‰) and 6.9 to 14.5‰ δ 15 N values (average value of 11‰). The wide range of isotope values for fauna from this site indicates that livestock exploited resources from varied ecologies near this oasis, which was located in a relatively arid region.

The cat had similar carbon isotope values, but higher nitrogen isotope values (by 5.4‰) than the dog from the same site. When compared with zooarchaeological remains of dogs and cats from across central Asia, the cat at Dzhankent was enriched in 15  N by more than 5‰, with the exception of a cat from Geotchikdepe (16.4‰) and dogs from Chicha (range from 9.4 to 13.6‰) 52 , 56 , 60 , 61 . Large quantities of fish remains were recovered from the sites of Chicha and Geotchikdepe suggest that these largely carnivorous animals consumed aquatic taxa which enriched their nitrogen values. As fish remains were identified at Dzhankent for this period and the site was located on a palaeochannel in the delta of the Syr Darya, fish would have been readily accessible protein resource, and likely contributed to higher δ 15 N values 32 . Both the cat and dog from Dzhankent were depleted in δ 13 C relative to dogs from Quanhucun, a site which had archaeological evidence for millet, but enriched in δ 13 C relative to most livestock from Dzhankent. This suggests that both taxa were ingesting small amounts of C 4 plant resources directly, such as millet, or consuming fauna which were subsisting on C 4 plants. As the livestock from this site have only slightly lower δ 13 C values than the two carnivores, it is possible that the dog and cat ingested rodents or other commensal species which were subsisting on agricultural products.

Palaeopathologies: evidence for human care

This individual possesses extensive pathologies found across nearly all recovered skeletal elements which were recorded macroscopically using a descriptive recording protocol 62 , 63 (Supplementary Table 1 ). Osteological pathologies encompassing four nosological categories were identified: trauma, joint disease, inflammation, and dental disease. Trauma was evident on both left and right sides of the body. An oblique fracture was apparent in the distal metaphysis of the right femur (Fig.  3 ), which was in the process of healing at the time death. The fractured halves united with slight medial deviation and proximal rotation, such that the distal condyles were re-positioned 135° to the shaft. Articular contour changes to the distal femur and enthesophytes at the proximal end of the bone could represent secondary responses to altered biomechanical loading on connective tissues. The left humerus had an articular fracture on the proximal articulation, extending distally and medially from the centre of the humeral head towards the articular margin. Osteophytes and enthesophytes present on the left humerus and scapula and contralateral humerus, radius, and ulna may again be adaptations to altered loadings, although such lesions on and around the joints can also be caused by age-related joint disease. The left pelvis exhibited lesions characteristic of hip dysplasia including the flattening of the craniodorsal acetabular border and marginal osteophytes 64 . Unfortunately, the absence of the left femur meant that the adjacent lesions could not be assessed. However, the left tibia presented a change in the orientation of the proximal articular surface, ankylosis of the proximal fibula and evidence for calcification of ligament/tendon attachments, which may be a remodelling response associated with hip dysplasia in the same limb. In addition to the traumatic lesions, periosteal new bone formation was observed on the right ulna and left tibia. Ossification of the connective tissue surrounding the shafts of long bones is an inflammatory response but is relatively non-specific and can arise as a result of localised trauma, inflammation in the surrounding soft tissues, or systemic disease, and therefore cannot be directly connected to the above traumas. Some of the enthesophytes might be directly or indirectly associated with traumatic insult, however there may also be an age component, since ossification of attachment sites can occur with age. Supportive evidence for an age of the cat that is greater than one year of age is provided by the presence of marginal osteophytes and articular porosity in the mandible and distal tibia, characteristic signs of degenerative joint disease.

Significant and extensive ante-mortem tooth loss of mandibular incisors and canines with root infilling and ante-mortem loss of the second and third maxillary premolars on the left side was observed. The mandibles also presented evidence of porosity, periosteal new bone formation on the labial side and feathered alveolar edges. Calculus was present on the left maxillary fourth premolar. Enthesophytes were observed along the attachment site for the masseter muscle, while osteophytes and articular contour changes were apparent in the temporo-mandibular joint. Radiographic examination of the mandibles indicated early bone loss in the furcation of the right P4 and furcations of the left premolars with a reduction in the alveolar crest. All of these are clear signs of early stage periodontitis which could be the cause of ante-mortem tooth loss 65 . There is no radiographic evidence of feline odontoclastic resorptive lesions or tooth resorption (TR) in the mandible and maxillary dental arch 66 .

Periodontal disease arises from escalation of inflammation of the gingiva, usually following the accumulation of bacteria-rich dental plaque, and is strongly influenced by diet, with a soft diet favouring plaque and calculus accumulation and thus gingivitis; however, it also increases in frequency and severity with age 67 , 68 . Canine loss is correlated increasing age in cats and is also more prevalent in feral cat populations relative to domestic cats and may not have been solely due to periodontal disease, as TR has multiple aetiologies, and can also be caused by disturbances in vitamin D and calcium metabolism, although this is unlikely given this individuals diet 69 , 70 . The presence of intact calculus suggests that a soft sticky food was a component of the cat’s diet in the weeks prior to death; research investigating the prevalence indicates that calculus deposition peaks at four weeks, after which calculus presence decreases 71 .

The pathologies in the skeleton suggest that the cat from Dzhankent had a physically challenging life, as evidenced by three traumatic lesions: fractures to the distal right femur and articular surface of the left humerus, and dislocation of the left femur. While congenital hip dysplasia is required in the differential diagnosis of the latter, it occurs much less commonly in cats than dogs 64 . Whether the traumatic episodes occurred in the same event or independently is difficult to establish. Furthermore, the time between the trauma and death is hard to determine and healing time for fractures depends upon both biological factors, such as the age of the individual, associated traumas, and blood supply, as well as mechanical factors, such as stabilisation of the fracture site 72 . Certainly, the fracture to the distal right femur was still healing at the time of death as evidenced by the presence of lamellar and woven bone within the callus. The angle of ossification of the fracture indicates that this animal may have been restrained, as there is little evidence for torsion at the fracture site.

A fracture to all three parts of the skeleton occurring in the same event is perhaps only conceivable if the cat experienced a fall from a considerable height. Irrespective of the cause, some of the lesions observed at and around the joints in the skeleton, such as new bone formation at ligament and tendon attachment sites and the morphological changes to the proximal tibia, may reflect secondary remodelling to compensate for altered biomechanical stresses or direct stress experienced by connective tissues, such as torn ligaments. The combined effect of these lesions would have been to considerably limit the mobility of this animal prior to its death, hindering its ability to capture prey, flee to shelter, and to fight. Inflammation and tooth loss would have caused discomfort and potentially affected the cat’s ability to catch and consume food as well as defend itself 73 . The presence of both calculus on the maxillary fourth premolar as well as additional dental pathologies and isotopic data indicates that our cat likely had a soft diet that was high in protein which was consumed during bone ossification and in the weeks prior to its death.

This individual cat from Dzhankent has pushed back the date of the first archaeologically domesticated cats in this region by a millennium to 775–940 cal CE. This male adult cat likely descended from a Middle Eastern population of domestic cats, although comparative modern genetic data from regional wildcats is required to confirm this ancestry. Both stable isotope and dental palaeopathological data indicate this animal had access to a high protein diet from an early age up until the final weeks prior to death, which most likely caused gum disease, a common affliction in modern domestic cat populations. The extensive paleopathologies across the skeleton clearly demonstrate the high probability of human pet keeping behaviours which contributed to this individual’s survival. This cat lived through a number of traumas and would have had limited chances for survival without human intervention, as it would have had trouble hunting with both physical impairments and loss of canines. Furthermore, the angle of ossification of the femoral fracture indicates that this animal may have been restrained during the healing process, which was likely only possible with human care and attention.

There was a steep increase in the number of domestic felines in European and Middle Eastern contexts at the end of the first millennium CE 16 , 17 , 18 , 19 , 21 , 22 , 23 , 74 , 75 . This has been tied to the need to control commensal activity in the proliferation of new urban settlements and to increasing international trade that connected the Islamic world with northern Europe, the Mediterranean and Northern Africa 7 , 76 . Therefore, the appearance of a domestic feline at Dzhankent suggests that this urban settlement was integrated with a network of trade and commerce, despite its relatively isolated location on the steppe. Archaeological and historical evidence indicates that Dzhankent participated in global trade networks, most likely taking part in the major north/south axis of the slave trade, which linked the slave markets of the caliphate in the Amu Darya delta with two major regions where slaves were captured: the steppe to the east (Bilad al-Atrak) and eastern and northern Europe (Bilad al-Saqaliba) via the major slave route originating in Bulgar 31 , 32 , 33 , 77 .

The evidence for human care of this animal indicates a shift in trajectory in the human-animal relationships typically found on the Central Asian steppe. The city of Dzhankent was unusual, as it was ruled by the Oghuz, a people that originated from the rural hinterlands of the Central Asian steppe and had their origins as a nomadic pastoralists, who have no documented archaeological evidence for keeping domestic cats. This cat’s osteobiography demonstrates deep and pervasive cultural links to other regions, particularly to the south to the regions of Khorezm, Khorasan and Persia, where pet keeping and feline anthropomorphisation were widespread practices by the arrival of Islam in the eight century CE 28 . Throughout the early medieval Islamic world, cities proliferated as both international trade and increasing degrees of social and cultural sophistication were celebrated 77 .

While the Oghuz elite would have contributed a strong pastoral worldview to the culture of Dhzankent pet keeping and tolerance of small felids is an urban idea, one that is typically associated with permanent agricultural settlements. Therefore, evidence for human kindness and care suggests the presence of a cosmopolitan population which harboured a diversity of worldviews on animals. While it is not possible to determine whether this cat was part of a local population of cats or another import that arrived via international trade, it is clear that the people of this town had a different perspective on pet keeping than previous steppe populations. They carefully cared for and tended to this animal throughout its life, providing high quality food and medical care. Thus, this small cat is not only the earliest known domestic cat on the Silk Road, but also evidence of the complex interface between nomadic and urban cultural worldviews in a rapidly globalising world.

Methods: zooarchaeological recording

Metrics were taken according to zooarchaeological standards (Supplementary Table 2 ) 78 . Three dimensional models of the bones were recorded using a DSLR Canon 80D, an automated turntable, and Autodesk Recap® software and these models are available in an open access repository Zenodo ( https://doi.org/10.5281/zenodo.3490934 ) 79 . The bones were subjected to digital radiographic examination using an Xograph DRagon mobile x-ray unit at the University of Leicester (60 KvP; 6.40 mAs, 0.025 s).

Methods: genetic analyses

Laboratory procedures.

The petrosal part of the temporal bone was sampled from the Dzhankent cat specimen through one of the auditory channels, as this skeletal element has been found to contain high purity DNA in other mammals 50 , 51 . Extraction and high-throughput sequencing libraries were performed following strict decontamination procedures in dedicated ancient DNA facilities at the University of Potsdam. Two independent DNA extractions were performed following a protocol specifically developed for the recovery of short fragments generally found in ancient samples 46 . For each extract, one Illumina sequencing library was prepared according to a single-stranded library preparation method developed for ancient DNA 47 , 48 . UDG/Endonuclease VIII treatment was performed for all libraries to excise uracils that arise from cytosine deamination (except the uracils occurring as terminal nucleotides) that damages DNA over time 80 . Optimal library amplification cycles were estimated using qPCR, and libraries were subsequently indexed following a dual indexing protocol to detect and remove chimeric sequencing artefacts and reads 81 . Sequencing libraries generated from the extraction blanks required five cycles more than the sample libraries, indicating that there was at least 32-fold less genetic material in the blanks, supporting the absence of detectable contaminant DNA. Sequencing was performed on the Illumina NextSeq 500 platform using a 75 bp single-end strategy 82 .

Sequence data processing

Raw sequencing reads were trimmed of the adapter sequence using cutadapt v1.12 83 , using default parameters and a minimum length filter of 30 bp. Remaining sequence reads were mapped to the domestic cat genome v6.2 49 as well as the cheetah reference genome Acinonyx jubatus v1 49 , as using an intraspecific reference genome as been shown to potentially introduce biases in downstream analyses 84 , using the Burrows–Wheeler Aligner (BWA v0.7.8 85 ) ‘aln’ and ‘samse’ algorithm with default parameters. Samtools v0.1.19 86 was used to filter reads with low mapping quality (MQ < 30) and suspected PCR duplicates. Average coverage was calculated using the GenomeAnalysisTK v2.8 ( https://software.broadinstitute.org/gatk/ ). Sex determination was performed by mapping the ancient cat sequences to the domestic cat assembly, and then comparing the average coverage for autosomal regions to the coverage of the X-chromosome assembly. Nucleotide misincorporation patterns indicating typical ancient DNA damage were checked to validate sequence authenticity using mapDamage v2.0.7 87 (Supplementary Fig. 2 ).

In order to compare the nuclear data from the Dzhankent cat to publicly available genome data, 37 shotgun sequencing datasets were selected from NCBI to reflect a range of breeds and species (Supplementary Table 3 ). Data was downloaded from the European Nucleotide Archive FTP server, and subsampled to 50–100 million reads using seqtk v1.2-r101 ( https://github.com/lh3/seqtk ) (Supplementary Table 3 ). Adapter sequences were trimmed using Skewer v0.2.2 88 , and overlapping reads were merged using flash v1.2.10 89 using default parameters for both. Mapping of the published data was performed using the same pipelines as described above, except for using BWA ‘sampe’ instead of ‘samse’. For mitochondrial DNA analysis, reads were mapped to the domestic cat mitochondrial genome (GenBank accession number FCU20753 90 ); following the same procedure as the nuclear genome, except for duplication removal which was performed taking both mapping coordinates into account (‘MarkDupsByStartEnd.jar’ ( https://github.com/dariober/Java-cafe/tree/master/MarkDupsByStartEnd ). For the phylogenetic analysis of mitochondrial DNA, 233 short sequences (ND5 & ND6 genes)—mainly from domestic cat and wildcat—were downloaded from GenBank and aligned with the Dzhankent cat, and the published individuals included in the genome analysis (Supplementary Table 3 & 4 ). Columns with gaps or missing data were removed, resulting in a 2,363 bp alignment of 264 taxa. A median-joining network (Supplementary Fig. 1 ) was generated using PopArt v1.7.0 91 , based on 214 segregating sites and 89 parsimony-informative positions.

Population clustering and phylogenetic analyses

Population structure was tested based on the genotype likelihood method (major/minor allele) in ANGSD v0.914 92 using random base sampling. Data was filtered for low base quality (-MinQ 30) and mapping quality (-minMapQ 30). Only reads mapping to cheetah scaffolds over 1 Mb were used in this analysis. As -MaxDepth parameter, we applied the 95th percentile of the global coverage as recovered using ANGSD doCounts. The analysis was restricted to transition sites (-rmTrans 1), and to sites where all individuals were covered by at least one read (-minInd). Singletons were excluded by applying a minimum allele frequency filter of 2/N of individuals. Two analyses were performed: one including only domestic cat and wildcat data, and one multi-species analysis with three additional cat species ( F. margarita , F. nigripes and F. chaus ). Principal component analysis of the resulting data was performed in R v3.4.4 and v3.5.1 93 . The nuclear phylogeny was estimated by calculating a pairwise distance matix using indentical data filters for all domestic and wildcat data, including a representative of F. nigripes to serve as outgroup. A rooted neighbour-joining tree was then calculated using the nj function in the R package ape 94 .

Methods: isotopes

Carbon and nitrogen isotope samples were prepared in the Archaeological Stable Isotope Laboratory (ASIL) at Christian Albrechts University at Kiel. Samples of bone weighing ~ 0.5 g were demineralized in 0.5 M EDTA (pH 8.0) with a change of acid every other day until collagen pseudomorphs were translucent and flexible. The resulting collagen pseudomorpha were rinsed in distilled water seven times, with an overnight soak on to remove residual EDTA solution, and then rinsed a further eight times 95 . The samples were then rinsed in 0.1 NaOH to remove humic acids and rinsed again five times in distilled water. The collagen samples were freeze-dried and weighed for analysis. Stable carbon (δ 13 C) and nitrogen (δ 15 N) isotope analysis was undertaken at the Boston University Stable Isotope Laboratory using a EuroVector Euro EA elemental analyser coupled with a GVI IsoPrime in continuous flow mode. Analytical error was 0.1‰ and 0.2‰ for δ 13 C and δ 15 N, respectively. Isotopic values are reported in permil (‰) relative to the Vienna Pee Dee Belemnite (VPDB) standard for δ 13 C and atmospheric nitrogen (AIR) for δ 15 N. Collagen quality was assessed using %C, %N, and C:N ratios 96 , 97 .

Data availability

Raw sequence data has been deposited in the European Nucleotide Archive (Study Accession Number: PRJEB38002). The consensus mitochondrial genome sequence is available from GenBank (Accession number: MT499915). Three dimensional models of postcranial bones and metrical data are available at Zenodo ( https://doi.org/10.5281/zenodo.3490934 ).

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Acknowledgements

We wish to thank P.D. Wordsworth for assistance with cartography, M. Stache with photogrammetry, and K. Schneider for assistance within the comparative mammal collections at the Central Natural Science Collections, Martin Luther University Halle-Wittenberg. We would also like to thank E. Lightfoot, E. Çoraman, and E. Evin for comments on an earlier draft of this paper. Funding for this research was provided by Wenner Grenn Grant No. ICRG – 10, Deutsche Forschungsgemeinschaft (DFG): GS 208 (AVM) and the University of Leicester and the Max Plank Society. We acknowledge the financial support from the DFG under the programme Open Access Publishing for the publication of this work.

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A.H. and A.V.M. conceived and designed the study; I.A. and H.H. directed sample collection by A.T. and S.B., and provided the archaeological and historical context; A.H. and R.T. conducted palaeopathological analysis; A.V.M. conducted stable isotope analysis; T.K., J.L.A.P., A.B., M.P. and Y.H. conducted genetic analyses; A.H., R.T., A.V.M., H.H., I.A., T.K., J.L.A.P. and A.B. wrote the manuscript.

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Haruda, A., Ventresca Miller, A.R., Paijmans, J.L.A. et al. The earliest domestic cat on the Silk Road. Sci Rep 10 , 11241 (2020). https://doi.org/10.1038/s41598-020-67798-6

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Science News Explores

Dna tells tale of how cats conquered the world.

Analyzing 9,000 years of cat remains points to two waves of feline migration

Before cats achieved world dominion, they spread in two waves, a DNA study suggests. Early Middle Eastern farmers probably brought kitties and agriculture to Europe over land. Egyptian cats may have been transported by boat to far-reaching parts of Europe, Africa and Asia.

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By Tina Hesman Saey

July 10, 2017 at 6:05 am

When it comes to revealing when and how wild felines became couch kitties, the cat is starting to come out of the bag. Cats were likely first tamed in the Middle East. Later, they spread — first by land, then by sea — to the rest of the world, researchers now report.

Early farmers brought cats with them to Europe from the Middle East by 6,400 years ago. That’s the conclusion from looking at DNA from 352 ancient cats. A second wave of migration, perhaps by ship, appears to have occurred some 5,000 years later. That’s when Egyptian cats quickly colonized Europe and the Middle East.

Researchers describe how they came to these dates in a new study. It was published June 19 in Nature Ecology & Evolution .

Domestication (Doh-MES-ti-kay-shun) is the long and slow process by which people have adapted wild animals or plants to be tame and useful. Wolves became dogs, for instance. Wild ox became cattle. And wildcats became house cats.

Exactly where and when this happened to cats, though, has been a matter of great debate. Researchers had only the DNA from modern cats to work with. These data showed that house cats had been tamed from African wildcats. What was not clear was when domesticated cats began to spread around the world. Now, new ways of studying ancient DNA are pointing to some answers.

Eva-Maria Geigl and Thierry Grange are behind this deepest dive yet into the genetic history of cats. They are molecular biologists. Both work at the Institute Jacques Monod in Paris, France. Mitochondria (My-tow-KON-dree-uh) are tiny energy-producing structures inside cells. They contain a bit of DNA. Only mothers, not fathers, pass mitochondria (and its DNA) to their offspring. Scientists use slightly different varieties of mitochondrial DNA, called mitotypes , to track the female side of families.

Geigl, Grange and their colleagues collected mitochondrial DNA from 352 ancient cats and 28 modern wildcats. These felines spanned 9,000 years. They came from regions stretching across Europe, Africa and Southwest Asia.

Story continues below image.

About 10,000 to 9,500 years ago, African wildcats ( Felis silvestris lybica ) may have tamed themselves. They would have hunted rodents and scavenged scraps from the homes of early farmers in the Middle East. People probably encouraged the cats to hang around as a way for these farmers to control mice, rats, snakes and other vermin. The arrangement would have been “mutually profitable for both sides,” explains Grange.

No one really knows how friendly people and cats were with each other at the beginning of cat domestication. Some people may have been very close to their pet cats. Indeed, one person on the Mediterranean island of Cyprus, 9,500 years ago, was buried with a cat. Says Geigl, this suggests that some people, back then, already had close ties to cats.

Before early farmers started migrating from the Middle East to Europe, European wildcats ( Felis silvestris silvestris ) carried one mitotype. It’s called clade I. A 6,400-year-old Bulgarian cat and a 5,200-year-old Romanian cat had a different type of mitochondrial DNA. They both had mitotype IV-A*. That mitotype was previously seen only in domesticated cats from what is now Turkey.

Cats are territorial and usually don’t roam far. This suggests people must have transported cats to Europe.

Mummies (and more) tell another story

Domesticated cats in Africa — including three cat mummies from Egypt — had yet another mitotype. It’s known as IV-C. Until about 2,800 years ago, that type was found mostly in Egypt. But then it began showing up in Europe and the Middle East. And between 1,600 and 700 years ago, it spread far and fast. By then, seven of nine of the ancient European cats the researchers tested now carried this Egyptian type of DNA. Among them was a 1,300- to 1,400-year-old cat from a Viking port far to the north, on the Baltic Sea.

Thirty-two of 70 cats from Southwest Asia also had that mitotype. That rapid spread may indicate that sailors traveled with cats, some of which could have jumped ship to find a new home.

The speedy spread of the Egyptian cats’ DNA could mean that something made these animals especially attractive to people, Geigl and Grange say. House cats aren’t much different from wildcats. The big difference is that domestic cats tolerate people. And the Egyptian cats may have been particularly friendly. They may have more resembled the type of purring pet found in homes today, the researchers speculate. Earlier house cats might have been more comfortable with people than wildcats were, but still have qualified as scaredy cats.

There’s not enough evidence to say that, counters Carlos Driscoll of the National Institutes of Health in Bethesda, Md. Working at its Laboratory of Comparative Behavioral Genomics, he studies the genetic bases of some behavioral traits. And Driscoll now suggests another reason why Egyptian cats got popular so fast: They may have lived along shipping and trade routes. That would have made hopping a boat to some new port easy, especially if they offered to work as mousers on the ship.

Earlier cats may have been just as popular, Driscoll says, but moving them would have been harder. Those early cats, he says, would have been “dependent on somebody putting a bunch of kittens in a basket and walking across a desert with them.”

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A review of over three decades of research on cat-human and human-cat interactions and relationships

Affiliation.

• 1 Animal Behavior, Dept. of Evolutionary Biology and Environmental Studies, University of Zurich, Winterthurerstrasse 190, CH-8057 Zurich, Switzerland. Electronic address: [email protected].
• PMID: 28119016
• DOI: 10.1016/j.beproc.2017.01.008

This review article covers research conducted over the last three decades on cat-human and human-cat interactions and relationships, especially from an ethological point of view. It includes findings on cat-cat and cat-human communication, cat personalities and cat-owner personalities, the effects of cats on humans, and problems caused by cats.

Keywords: Cat-human relationships; Communication; Ethology; Personality; Socialization.

Copyright © 2017 Elsevier B.V. All rights reserved.

Publication types

• Behavior, Animal / physiology*
• Cats / psychology*
• Communication*
• Human-Animal Bond*
• Personality*

Developments in Research on Cat Cognition and Personality

• First Online: 28 August 2021

Cite this chapter

• Saho Takagi 3 , 4  

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Cats, along with dogs, are one of the most popular companion animals for humans. Across the world, increasing numbers of cats are being kept as pets. Despite their familiarity, cats’ cognition has long been shrouded in mystery, mainly because cats were considered largely unsuitable for psychological studies in laboratory settings. The “Cats Team” in Kazuo Fujita’s lab has developed several innovative and useful methods for studying cat cognition. In this chapter, I review findings from some of the team’s studies of cat cognition, including physical inference, use human social cues, incidental memory, cross-modal integration, jealousy, and third-party social evaluation. I also briefly describe some ongoing work on the relation between genes and personality, and suggest directions in which behavioral and cognitive studies of cats might go.

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Takagi, S. (2021). Developments in Research on Cat Cognition and Personality. In: Anderson, J.R., Kuroshima, H. (eds) Comparative Cognition. Springer, Singapore. https://doi.org/10.1007/978-981-16-2028-7_17

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Cats: The strange and fascinating history of our feline friends

Cats have associated with humans for up to 10,000 years.

When were cats domesticated?

• Are there different breeds?

How smart are cats?

Do cats feel emotion.

• Do cats see color?

How do cats help people?

Additional resources.

Domestic cats ( Felis catus ) are small carnivorous members of the family Felidae — the only member of that family that has deigned to join humans in domesticated bliss. 

Cats have lived among humans for thousands of years. They probably started hanging around human grain stores, attracted by mice and other vermin, and eventually spread around the world as sailors brought them aboard ships.

Today, cats still help humans control vermin and also provide companionship. In the past few centuries, humans have bred some cats to display certain traits, like hairlessness, establishing dozens of cat breeds. With their charming mix of aloofness and goofiness, cats amuse and fascinate humans in equal turns.

Cats were domesticated around 10,000 years ago, research shows. A 2017 genetic study found that today's domestic cats descend from Felis silvestris lybica , a wild cat subspecies from the Near East. Genes from cats found in archaeological sites in the Near East, Europe and Africa reveal that about 10,000 years ago in modern-day Turkey, cats started to associate with humans and split from their wild relatives. 

Despite having relatively small natural ranges, Felis silvestris lybica started showing up in eastern Europe by 4400 B.C., according to telltale genes from cats found in archaeological sites. This spread strongly suggests that cats were hitching rides aboard ships with traders, who probably appreciated that cats kept rats in check. Cats certainly traveled long distances: A 2016 study found DNA from Egyptian cats at a Viking site in northern Germany dating to between A.D. 700 and 1000.

The oldest known burial of a domesticated cat comes from Cyprus, where a human and a cat were buried together 9,500 years ago, researchers reported in 2004 . Cat bones also have been found buried in 5,300-year-old refuse pits in China , suggesting that the felines were a part of human life in the Far East, too.

Even if cats joined people in the Near East, it was in ancient Egypt where they took on a starring role. Scientists aren't yet sure whether the Egyptians domesticated cats separately from the Near East lineage, or whether the cats spread from Turkey to Egypt. Either way, Egyptians treasured cats' mixture of protectiveness and independence and saw the traits of their gods in cats, which were sometimes mummified lovingly next to their deceased owners but were also sacrificed in large numbers as part of religious rituals. Bastet, a feline-headed goddess, was worshipped as a protector and as a deity of pregnancy and childbirth.

Related: Why were the ancient Egyptians obsessed with cats?

Are there different breeds of cats?

Cats come in many breeds, though not with the same degree of diversity as dogs. The Cat Fanciers' Association , a nonprofit dedicated to cats, recognizes 45 pedigreed breeds as well as the "companion cat," otherwise known as "most regular cats."

These breeds include relatively well-known varieties, like the slender Siamese cat and the fluffy Persian, as well as the leopard-spotted Bengal, the short-tailed American bobtail and the hairless Sphynx. While dogs have been bred over centuries to carry out different tasks, leading to a variety of sizes and shapes, domestic cats have only two jobs: controlling pests and being pets. Thus, most pedigreed cats are bred for traits such as color and fur length rather than, say, a waterproof coat for swimming or a strong herding instinct for guarding sheep. Most cat breeds date to less than 100 years ago, Leslie Lyons, a professor at the University of Missouri College of Veterinary Medicine, told Live Science in 2017 .

Cats are intelligent, and the widely held notion that dogs are smarter than cats may be unfounded, given that each species has cognitive abilities best suited to their lifestyle, Live Science previously reported. Felines display object permanence , or the ability to realize that something still exists when it's out of sight — an ability humans develop around the age of 8 months. Object permanence is an important skill to have when you're a hunter operating at night, listening for the telltale patter of tiny mice feet. Cats also recognize their owners' voices , according to a 2013 study in the journal Animal Cognition .

But cats don't always make their smarts easy to measure. "They're a nightmare to work with in the lab," said Julia Meyers-Manor, a psychologist at Ripon College in Wisconsin who studies animal cognition.

Compared with other animals, such as rats and dogs, few studies have been done on cat intelligence, Meyers-Manor told Live Science. Cats hate strangers and unfamiliar places, like labs, she said; they typically bury their faces in their owners' arms and refuse to cooperate with the task at hand.

Unlike dogs, which are pack animals, cats evolved from a solitary wild lifestyle, thus making them less attuned to social cues   In a 2021 study , researchers had cats watch as their owners struggled to open a container. In some cases, an actor helped the owner open the container. In others, that actor rudely turned away. In still others, an actor sat by neutrally, neither helping nor refusing to help.

The actor then offered the watching cat a treat. Previous studies had found that dogs avoided taking treats from an actor who had refused to help their owners, but cats didn't care; they took treats from anyone. According to the researchers, this may not have been cold-hearted behavior by the kitties; instead, cats may not have understood the difference between someone who helped and someone who didn't. They simply haven't been bred for hyper-cooperativity with humans, the way dogs have.

In other words, cats are plenty smart, at least at tasks that a solitary hunter needs for survival. They just don't necessarily care if you know it.

Cats might not be the most socially savvy, but there is evidence that they form bonds with their humans. A 2002 study in the Journal of the American Veterinary Medical Association found that cats can develop separation anxiety, often displayed by peeing or pooping somewhere they shouldn't. Cats also seemed to defer to their owners in a 2015 experiment in which they were presented with an unfamiliar object (in this case, a fan). Of these cats, 80% looked between the fan and their owners while their owners talked about the fan in either a reassuring or alarmed tone of voice. Cats who heard the alarmed tone were more likely than cats whose owners were calm to look toward the room's exit, suggesting they understood the negative emotion in their humans’' voice and were responding to it. They also interacted with their owners more, suggesting they were seeking reassurance.

Cats also seem to recognize the emotions of both other cats and humans. In a 2020 study in the journal Animals , researchers showed pet cats pictures of angry or happy human faces alongside recordings of humans laughing or growling angrily. They also showed the cats pictures of angry and contented cats alongside recordings of cats hissing and cats purring. In some cases, the pictures matched the emotional sounds; in others, they were mismatched. The researchers then measured how long the cats looked at each image while the matched or mismatched sound played.

The results showed that cats looked longer at the pictures when the emotional sounds matched. They also showed more stress when exposed to the stimuli representing both human and cat anger. This finding suggests that cats can process basic emotions, even when displayed by another species.

"These findings demonstrate that cats have developed social skills that allow them to understand human emotional signals, which is a key factor for the maintenance of interspecies relationships and for strengthening the human-cat bond," the researchers concluded.

Do cats see color? (And other feline senses)

Cats have excellent vision, according to the Merck Veterinary Manual . Their eyes are loaded with sensing cells called cones, which provide acuity in bright light. They're also equipped with a high number of rods, cells in the retina that are good at capturing dim light. This allows cats to see six times better than humans can in the dark, according to Merck. A reflective layer in the eye called the tapetum lucidum also helps collect extra light at night. This layer is what makes cats' eyes shine green if a flashlight or headlight catches their eye in the dark.

Research from 2014 suggests that cats can see ultraviolet light , blue light outside the range that humans can see. Cats likely don't see color as well as humans, though. They have two types of cones that can detect short- and long-wavelength light, according to a 2009 study , while humans have three, giving people greater sensitivity at distinguishing between colors. In 2013, an artist created comparisons of scenes from the perspectives of cats and humans , showing that cats don't see long distances as well as people do and that their world looks more desaturated. However, cats have a broader field of view than humans do: 200 degrees compared with 180 degrees for people.

Cats aren't particularly strong smellers, but their sense of hearing is very sensitive. According to a 1985 study in the journal Hearing Research , cats can hear frequencies between 55 hertz and 78 kilohertz, a broad range that is topped only by porpoises and cattle. (For comparison, the deepest sound a human can hear is 20 hertz, and the most piercing high sound is around 20 kilohertz.)

Cats still work to control rats and mice around farms, nurseries and warehouses. But most of the time, a cat’s job is to provide companionship for people.

Science suggests they do their job well. A 2016 study in the journal Pet Behaviour Science found that spending time with a cat reduced heart rate and blood pressure in both owners and volunteers who didn't own cats. A study published online in the journal Anthrozoös in 2015 found that cats reduce their owners' negative moods.

Interacting with cats activates the prefrontal cortex, the front part of the brain that is associated with complex tasks. A 2020 study in the journal PLOS One found that playing with, petting, training and feeding cats all activated a region of the prefrontal cortex linked to empathy and nonverbal communication.

The autonomy and independence of the cats seemed to be key to what makes the cat-human relationship work: Getting a normally uncooperative cat to cooperate with play or training provided a big mood jolt, the researchers found. It turns out that for cats, playing hard to get pays off.

Learn more about caring for cats and avoiding their diseases at the Centers for Disease Control and Prevention . Check out the The Humane Society of the United States for useful information on how to care for cats and solve common problems. And this Library of Congress page has a fascinating description of how cats became domesticated.

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Stephanie Pappas is a contributing writer for Live Science, covering topics ranging from geoscience to archaeology to the human brain and behavior. She was previously a senior writer for Live Science but is now a freelancer based in Denver, Colorado, and regularly contributes to Scientific American and The Monitor, the monthly magazine of the American Psychological Association. Stephanie received a bachelor's degree in psychology from the University of South Carolina and a graduate certificate in science communication from the University of California, Santa Cruz. 

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A Short Natural History of the Cat its Relationship with Humans

The creation of cat breeds has been a significantly different process than breed development in other companion animal and agricultural species. These nuances are important for an understanding of the appropriate genetic tools, resources, and techniques that will be the most beneficial and efficient for cat genetic research and health programs.

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A Brief History of Cats: From the Wildcat to the House Cat

• by Ljubica Cvetkovska
• May 5, 2020

There are 600 million cats globally , so chances are good that one sits on your lap as you read this fascinating article.

As you look at your sleeping cat, have you ever wondered how much we, mere humans, know about the history of cats and where they came from?

How long have they been a part of our lives?

Is it the history of domesticated cats, or did they simply decide to include us in their lives ?

What caused the transformation from the majestic and feral wildcat to your furry friend and cuddly companion?

These are questions that have puzzled scientists for years. They have led to much research and studies into the origin of cats and their bond with humans.

The Natural History of Cats: In the Beginning

For a long time, it was a widely held belief that cats originated in Egypt some 4,000 years ago.

However, there’s now DNA evidence showing that cats and humans have been living together for 12,000 years.

Talk about a long cat domestication history . 

An analysis of the genetic composition of 79 house cats and their ancestors from all over the world concluded that all domestic cats actually have a common descendant, the African wildcat called Felis silvestris lybica , or “cat of the woods.”

As the name suggests, the ancestors of your precious furball probably made the transition from the forest to people’s homes, from there to their hearts.  

There’s even more proof to dispute the Egyptian theory.

In 2004, a cat was discovered deliberately buried with a human in a 9,500-year-old neolithic grave in Cyprus.

As there are no native wildcats on this Mediterranean island, it’s safe to assume that cats must have been brought to the island. Thus, cats’ evolutionary history and  domestication started much earlier than was initially thought. 

Today, it’s believed that wildcats started living with humans in the Middle East’s Fertile Crescent. This is an area of fertile land along the Tigris and Euphrates rivers.

The people who lived there settled into a life of agriculture—transitioning from being hunters and nomads to becoming farmers.

Farms and crops attracted rodents and vermin, which in turn attracted cats. Thus a beautiful relationship was born between felines and humans.

The History of Cats in Ancient Egypt : Gods and Cats

Everyone knows that the Egyptians worshipped felines as gods, but did you know that cats were so revered there that killing one was punishable by death? 

Like in the Middle East, Egyptian cat history shows they were “employed” by the ancient Egyptians as mouse-catchers.

They were in charge of protecting crops and stopping diseases from spreading among the people. 

They did a great job and raised the quality of life of Egyptians to a whole new level. Thus marking a turning point in the history of cats so much so that felines soon started to be seen as more than just useful animals, but as sacred creatures that represented life and prosperity.

And it wasn’t just ordinary people who held cats in high esteem.

Pharaohs were commonly buried with their cats. They believed that felines brought good luck and a safe journey to the afterlife. 

Moreover, archaeologists are still discovering hieroglyphics, pictures, and carvings of cats shown wearing expensive jewelry or taking up prominent positions, thus demonstrating their importance to this ancient civilization.

So even cat history proves it: felines were born to be adored.  

According to legend, cats were so beloved in Egypt that the Persian army carried cats with them when they fought the Egyptians, knowing that their archers would not fire and risk hurting the precious felines. 

The Egyptian goddess most associated with cats is Bastet, the goddess of love. She had the head of a cat and could also turn into one.

The festival dedicated to this goddess was one of the biggest and most celebrated in all of Egypt. According to the history of cats in Egypt , people showed their devotion to the goddess by leaving sculptures of cats in front of her temple. They even did so by mummifying felines. 

Based on archeological findings, there were many necropolises dedicated to cats. In fact, one discovered in Beni-Hassan was found to contain around 300,000 cat mummies. 

Cats Traveling the World: From Egypt to China and Beyond

Enjoying their popularity in Egypt wasn’t enough for the majesty of the cat.

The domestic cat evolution spread beyond Egypt, and felines soon started being sold to Greece, Persia, and even China. 

However, cat history facts note that China already had its own kind of domesticated cat 5,300 years ago, known as the leopard cat. This was a distant cousin of the African wildcat and thus one of the ancestors of the kitties we have and love today. 

Chinese felines, like their relatives, weren’t kept just for their looks.

As with the history of cat domestication on other continents, they earned their keep by protecting villages, food, and religious manuscripts from pests.

However, modern cats in China are descended from the wildcat. This means that at some point in the cat history timeline, F. silvestris lybica replaced the leopard cat in China.

This led to the spread of domestic cats in China, with other breeds such as the Siamese and the Burmese .

From China, cats moved to Japan and India as stowaways on ships. That is where the popular Burmese and Siamese cat history began . 

But Egyptian merchants didn’t just trade with the Far East. They’re also credited with bringing felines to Rome, where they were also used to ward off rats and disease. 

It was the Romans who introduced cats to Britain around 100 AD.

It seems that the role of cats throughout history has remained the same, because like other nations before them, the Britons also saw the value of felines as crop protectors and mouse catchers.

Like the Egyptians, they worshipped them as sacred animals, particularly in Wales, where felines were protected by law and beloved by kings. 

When the Romans left Britain, they also left a couple of cats behind who were picked up by the Vikings and taken to Norway.

From there, the cat traveled all over Europe. 

A Brief History of Cats in the Middle Ages: A Bad Time for Felines

Sadly, cats started to be associated with superstition and witchcraft during the Middle Ages. They were even seen as the “devil’s pet,” and black cat history was especially difficult.

No one is really sure why, but many believe that people distrusted the way cats hunt. 

Also, a cat’s wild nature couldn’t have been too helpful either.  This attitude particularly intensified during the Black Plague.

As people all over Europe were dying from this terrible disease, the history of the cat took a downturn.

Felines were pronounced as the culprits, and millions were killed in bonfires, hung, or tortured. 

Ironically, the mass killing of cats actually helped spread the disease because it allowed rats to thrive and thus worsened the epidemic. Scholars now suggest that countless people could have survived the plague if cats had been spared. 

This was an unfortunate time for all cats, but black kitties  had it the worst .

These felines have always had an image problem, but the history of black cats was darkest during the Middle Ages. 

During this difficult history of house cats , people believed that black cats were witches’ familiars who helped them do dark magic.

Some even thought that witches could turn into cats, so it wasn’t uncommon to burn felines along with those suspected of witchcraft. 

It wasn’t until the 1600s that cats stopped being seen as doing the devil’s work and were once again in people’s good graces all over Europe.

By then, felines had already made the brave journey to the New World. 

History of Cats in America: Settling in the New World

Early colonists started seeing the value of cats once again in the late 15th and throughout the 16th Century—despite still being persecuted in Europe . They gave them jobs on ships as rodent catchers, once again putting the cat evolution timeline back on track. 

It’s believed that Columbus brought cats with him to America, particularly the British Shorthair, which is rumored to be the ancestor of today’s very popular American Shorthair.

Cats seemed to have taken a liking to the New World, where they thrived and flourished. 

Felines even moved to the White House—a pinnacle in the history of domestic cats .

Abraham Lincoln was the first president to take two felines to Washington. His two companions, Tabby and Dixie, were the first two White House cats. 

He is also said to have loved kitties so much he even took in strays.

This started a tradition of other presidents bringing their cats to the White House with them.

Teddy Roosevelt’s cat, Slippers, known to fall asleep in the hallway, was also a famous White House cat. He enjoyed forcing guests at banquets to detour around her so as not to wake her.

Another famous White House cat was George W. Bush’s cat, India, the family pet cat. 

The History of Cats in the 20th Century

Regardless of their popularity, cats still weren’t treated as pets until the early 19th Century.

In fact, like dogs, they were employed by cowboys and even the US army.

The evolution of the cat from wandering mouse killers to furry companions happened gradually as more and more people started bringing them indoors. 

But the invention of certain technological and scientific developments, like fridges and kitty litter, was a turning point in the history of cats as pets. They turned felines into an integral and beloved member of households and families everywhere.

When did cats first appear on Earth?

No one is really sure how cats first appeared. Some theories indicate that both cats and dogs share a common ancestor, Miacis , a weasel-like animal that lived about 40 or 50 million years ago. 

The cat evolution tree reveals that cats belong to the Felidae family, whose evolution began 25 million years ago.

It produced 37 different species of cats, including lions, tigers, cheetahs, and the African wildcat, the ancestor of all domestic cats. 

Where did the first cat come from?

The first contact between humans and cats occurred almost 12,000 years ago when wildcats started hanging around farms in the Fertile Crescent, searching for food and shelter.

Farmers saw their skills and value in getting rid of rodents and protecting crops. Thus, an unbreakable bond was born between felines and people.

How are wildcats and domestic cats similar?

Despite thousands of years of cat evolution, wildcats and domestic cats are very similar .

In fact, the only thing setting them apart is the tabby coat pattern, which is characteristic of domestic felines only. 

Of course, domestic cat evolution has also changed them in other ways.

They aren’t as solitary as their cousins in the wild. They put up with humans and other animals (more or less), but some of that feral animal is still in them and comes out when they play or hunt. 

If you think that cat evolution has wholly removed your pet from its ancestors, just try disturbing it while it’s napping.

You’ll see that there’s no need for DNA evidence to determine his connection to the African wildcat.

Are cats given jobs?

Throughout history, cats have managed to hold down some jobs. Admittedly, not as many as dogs, but cats’ careers are still pretty diverse.

Cats have worked as station masters, astronauts, and spies. They’ve been elected mayor and even tried to deliver mail in Belgium (unsuccessfully).

But still, the primary role of cats throughout history is catching mice.

In fact, the British government employs around 10,000 cats to keep Downing Street a rat-free area.

There’s even the title of Chief Mouser, given to the cat that resides with the Prime Minister.

There’s been a Chief Mouser in England since the 1500s.

However, perhaps the most famous of them all is Larry. He was fired from his position for sleeping on the job, spending time with a neighborhood cat, and making just one confirmed kill. 

When did cats become house pets?

Although cats have been domesticated—i.e., they can live with humans—for as long as 12,000 years, the cats of early civilizations could hardly be called pets.

Felines started to be kept for company, in addition to their usefulness, around the start of the 19th Century.

By the end of World War I, they were found in many households worldwide. 

What are some of the myths and legends connected to cats?

Many people love cats, but perhaps none as much as the Japanese.

The history of cats in Japan is closely connected to the myth of the Maneki-Neko, or beckoning cat, which is commonly found at the entrance of shops, restaurants, and other businesses.

According to Japanese legend, one day a landlord saw a cat beckoning to him with its paw.

When he came close to the cat, lightning struck the place where he had just been standing.

The landlord believed that the cat was responsible for his good luck, and since then, the Maneki-Neko is believed to bring good fortune. 

Not all legends throughout the entire history of cats reflect favorably on them.

For instance, people in England believed for centuries that cats stole the breath of babies.

At one time it was even thought that felines could make beer go sour or that stepping on a cat’s tail means you won’t get married that year. 

Bottom Line 

Cats have been through a lot: from being revered as gods and good luck charms to becoming associated with disease and witchcraft.

However, if there’s one takeaway from the rich and captivating history of cats , it’s that felines have always had a special place in the lives of humans and will likely continue to be a big part of their homes and hearts in the future. 

Ljubica Cvetkovska

Ljubica is a writer and researcher who enjoys spending most of her time between the pages of her favorite books or immersed in her writing. With a background in English literature, she prides herself on delivering content that is well-researched and backed up by relevant data. When she’s not working, she’s known to binge-watch a TV show or two or hit the gym, which doesn’t happen that often.

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• 1 Department of Population Health and Reproduction, School of Veterinary Medicine, University of California, Davis, Davis, CA, United States
• 2 School of Animal and Comparative Biomedical Sciences, University of Arizona, Tucson, AZ, United States
• 3 Institute for applied Ethology and Animal Psychology, Horgen, Switzerland
• 4 Department of Veterinary Medicine and Surgery, College of Veterinary Medicine, University of Missouri, Columbia, MO, United States
• 5 Department of Anatomy, Physiology, and Cell Biology, School of Veterinary Medicine, University of California, Davis, Davis, CA, United States

Editorial on the Research Topic Exploring cats: their behaviors and human-cat interactions

Introduction

Domestic cats are immensely popular companion animals in households around the world ( 1 ). Over 45 million US households contain at least one companion cat ( 2 ); in the European Union, the population of pet cats is estimated to be 113 million (outnumbering the estimated 92 million dogs) ( 1 ). Despite this global popularity, research into the behavior and welfare of cats living in private homes is still limited; and arguably, even less is known about the mechanisms of human-cat interactions within the home. Outside the home, cats allowed uncontrolled outdoor access alongside free-roaming cat colonies outside of human ownership (but not always without human care), still generate considerable controversy between animal advocates and conservationists concerned about cats' impact on wildlife. This Research Topic presents 12 new papers that shed light on these issues and more. The goal of this Research Topic is to improve our understanding of companion cats, with particular focus on their interactions with humans, and human attitudes toward these animals. The twelve manuscripts in this Research Topic on cat behaviors and the development of the human–cat bond cover a wide variety of themes.

The mechanics of human-cat interactions

Turner(a) presents a mini-review of the available literature on a number of topics relevant to our understanding of human-cat interactions, such as the importance of kitten socialization, how cats communicate with their humans, and the mechanics of social interactions between cats and humans (such as the influence of who initiates contact, and of symmetry in compliance, or lack of compliance, with the partner's “wishes”). Noting the importance of ensuring the animals' wellbeing during human-cat interactions (and the scarcity of research into this issue with companion cats), Haywood et al. present Human-Cat Interaction guidelines designed to improve the comfort and welfare of companion cats during such interactions. They developed and tested the efficacy of these new guidelines with 100 shelter cats, interacting with 120 novel members of the public, and report their results here. Nagasawa et al. examine physiological (urinary oxytocin and cortisol) responses to interaction with humans, by comparing these variables in cats during positive interactions with a familiar caretaker (including physical contact, play, etc.), vs. when such interactions were removed. In another study using physiological (fecal cortisol) measures of stress, along with weight and behavior, Carlisle et al. investigate stress levels of cats adopted by families of children with autism spectrum disorder (ASD). Cats in the Carlisle et al. study were specifically selected for sociability and calmness using the validated Feline Temperament Profile, and the adopters provided with education on cat behavior; the authors discuss the importance of these factors to successful adoption into these homes.

Understanding interactions between cats

Gajdoš Kmecová et al. review and seek to extend the existing research on play in cats, much of which has focused to date on object play [e.g., ( 3 )], by looking at social play between cats. They suggest using a psychobiological approach to the study of play, which considers the motivational and emotional states of the cats; and present an ethogram (synthesized from the literature) and common terminology for use in future studies of cat social play. Khoddami et al. also seek to extend the existing literature on interactions between cats in multi-cat homes, by focusing specifically on two-cat households. They note that previous studies frequently lack focus on any particular group size, limiting our understanding of social dynamics in specific group sizes, despite the fact that most multi-cat households in the US and Canada consist of two cats ( 4 , 5 ).

Free-roaming cats and wildlife

Four papers in this issue focus on free-roaming cats, with two exploring the often-contentious issue of domestic cats' impact on wildlife. Tan et al. summarize the arguments for and against allowing cats outdoor access, and identify several owner- and cat-related factors associated with allowing companion cats uncontrolled access to the outdoors. Kim et al. investigate attitudes of different demographic groups toward feral cats in Seoul, South Korea, following the establishment of government-supported cat feeding stations around that city. They report distinct and sometimes complex differences between the groups in their attitudes toward cats and their preferred management approach for feral cat populations [e.g., trap-neuter-release (TNR) vs. culling]; they also discuss the possible impact of the feeding stations on these results. Turner(b) takes a critical look at the literature on cats' impacts on wildlife, in light of recent media reports of the “alarming predation of house cats on prey populations.” Turner(b) cautions that researchers should avoid bias and misinterpretation of field data, by considering what is known about predatory behavior in domestic cats and reporting estimates of total prey species population sizes. In their paper examining human-cat interactions involving free-living cats, Wandesforde-Smith et al. note the “moral pluralism” involved in the emphasis (even requirement) for humane care and protection of owned companion cats, alongside the systematic culling of large numbers of cats supported by public policy.

Use of technology in research on cats

The final two papers discuss research into new use of technology in the study of domestic cats. Xu et al. apply and advocate for machine learning techniques (in contrast to the more traditional biomechanical experiments with living cats or cat cadavers) for improving our understanding of the feline “athletic ability.” Given recent work using heart rate variability (HRV) as an indicator of emotion in non-human animals [e.g., ( 6 )], Grigg et al. compare HRV data collected using an affordable, commercially available cardiac monitoring system (Polar H10) against data from a traditional ambulatory electrocardiogram, to assess whether the Polar monitors could be used for this purpose in unrestrained cats.

This Research Topic tackles a broad range of topics relevant to domestic cats. Many of the papers add particular insight into our understanding of human-cat, and cat-cat, interactions. Others report on issues important to cat welfare, such as controversies surrounding outdoor cats and wildlife. Our understanding of domestic cat behavior and human-cat interactions continues to improve, as these papers demonstrate.

Author contributions

EG: Writing – original draft, Writing – review & editing. DT: Writing – review & editing. LL: Writing – review & editing. BH: Writing – review & editing. LH: Writing – review & editing.

T he author(s) declare that no financial support was received for the research, authorship, and/or publication of this article.

Conflict of interest

The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.

The author(s) declared that they were an editorial board member of Frontiers, at the time of submission. This had no impact on the peer review process and the final decision.

Publisher's note

All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors and the reviewers. Any product that may be evaluated in this article, or claim that may be made by its manufacturer, is not guaranteed or endorsed by the publisher.

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5. Larkin M,. Pet Population Still on the Rise, With Fewer Pets Per Household. American Veterinary Medical Association. (2021). Available online at: https://www.avma.org/javma-news/2021-12-01/pet-population-still-rise-fewer-pets-household (accessed February 13, 2023).

6. Von Borell E, Langbein J, Despres G, Hansen S, Leterrier C, Marchant-Forde J, et al. Heart rate variability as a measure of autonomic regulation of cardiac activity for assessing stress and welfare in farm animals – a review. Physiol Behav . (2007) 92:293–316. doi: 10.1016/j.physbeh.2007.01.007

Keywords: cats, welfare, human-animal interactions, free-roaming cats, human-animal bond, companion animals

Citation: Grigg EK, Turner DC, Lyons LA, Hart BL and Hart LA (2023) Editorial: Exploring cats: their behaviors and human-cat interactions. Front. Vet. Sci. 10:1329398. doi: 10.3389/fvets.2023.1329398

Received: 28 October 2023; Accepted: 21 November 2023; Published: 05 December 2023.

Edited and reviewed by: Marta Hernandez-Jover , Charles Sturt University, Australia

Copyright © 2023 Grigg, Turner, Lyons, Hart and Hart. This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY) . The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.

*Correspondence: Emma K. Grigg, ekgrigg@ucdavis.edu

This article is part of the Research Topic

Exploring Cats: Their Behaviors and Human-Cat Interactions

Cat’s Cradle by Kurt Vonnegut Research Paper

Kurt Vonnegut is one of the most well-known writers of the twentieth century who speaks about social and political problems using satire, acute irony, and black humor. Vonnegut is a science fiction writer who tells about Cold War fears and the threat of the Bomb, the lurking dangers of overpopulation and food shortage on the one hand, and on the other government’s efforts to assuage the population.

The threats to the individual, of being dehumanized in an anonymous technological world, of loss of identity, purpose, or power of choice, are implied repeatedly in even lightly humorous stories (Allen 65). Science fiction plots provide the perfect mode in allowing Vonnegut to treat these topics without becoming bogged down in the quagmires of logic that often inhibit their more serious discussion. His novel Cat’s Cradle (1963) vividly portrays his unique style of writing and vision of the world typical for Vonnegut. Critics underline that like many of his works, Cat’s Cradle is an autobiographical novel reflecting his life events and ideals.

Kurt Vonnegut was born in Indianapolis, Indiana, on November 11, 1922. His pedants, Kurt Vonnegut, Sr, and Edith Lieber Vonnegut were third-generation Americans. Vonnegut was a son of an architect, and his family never suffered the severe hardships that affected many families during the Depression. Kurt Vonnegut’s earliest publications appeared in his high school and college newspapers. In themselves, they do not appear exceptional in the quality of either their prose style or their ideas (Allen 12). That is natural enough since they are intended primarily to entertain student readers by being catchy and topical.

He wrote for The Daily Echo and the Cornell Daily Sun . The suicide of his mother was a terrible event in his life that had a great impact on the themes and symbols used in many novels. During WWII, Vonnegut was imprisoned in Dresden and spent about a year gathering bodies for mass burial. These events had a profound impact on his themes and were reflected in the novel Slaughterhouse-Five. The novel Cat’s Cradle was accepted as his Master’s thesis.

After the war, he worked as a police reporter and further a teacher at the University of Iowa. The second wife of Vonnegut was a famous photographer, Jill Krementz. He had seven children, and four of them were adopted. He spent most of his life in Barnstable, Massachusetts (Allen 43). Vonnegut died in 2007 in Manhattan. Two of his children become writers. and a daughter became an artist.

Cat’s Cradle is centered on the themes of science, religion, and technology. The protagonist of the novel, John, describes that he was going to write a novel about Hiroshima, and during his research, he got acquainted with the family of Felix Hoenikker, a fictional developer of the atomic bomb. He develops a substance called ice-nine, but now it becomes a possession of his children, the events take place n a fictional Caribbean island ruled by a dictator.

Their religion, Bokononism, is based on irrational actions aimed to bright joy to the community. In order to save his life and receive ice-nine, the dictator kidnaps the son of Felix Hoenikker. Instead of recovery, the dictator turns into solid ice. Because of an airplane crash, Monzano’s frozen body sends into the sea, which immediately freezes along with most of the water on earth. Extinction of life on the Earth is inevitable. At the end of the story, Bokonon suggests that he would write “a history of human stupidity” (Vonnegut 287).

The main character of the novel is John or “Jonah”. Cat’s Cradle is presented as if told by an almost anonymous narrator, who begins by trying to write the history of total destruction, with which Vonnegut himself was still wrestling in vain. John-Jonah moves among the heirs who share the invention–old Hoenikker’s children, along with their lovers and friends–learning slowly, painfully how to become yet one more Vonnegut victim: the patsy and reluctant messiah of yet one more true, i.e., false, religion (Schatt 54).

At the book’s close, he lies frozen for all eternity, his thumb to his nose and history of the world clasped to his side. He has learned this sacred gesture of contempt for the God or not-God behind the universe from Bokonon, a Black Prophet who is Vonnegut’s most impressive rebel-guru; and who, just before his own suicide, composed the final sentence of his Scriptures, as if for John-Jonah’s special benefit:

Vonnegut portrays that Dr. Hoenikker’s children suffer from the failure of their father and his scientific discoveries. Three of them have some physical disabilities that are only the sign of psychic problems. they are depicted as weak characters unable to resists the temptation to use ice-nine.

The Prospero who regulates the actions of everyone else is dead before the fiction begins; a certain Dr. Felix Hoenikker referred to throughout as “the father of the Atomic Bomb”. “Hoenikker, father of a bomb, father of three children, father of ice-nine. He was a little person” (Vonnegut 114). Franklin “Frank” Hoenikker is depicted as a “fox-faced, immature young man,” and a “pinch-faced child” (80, 194). He is cruel and unsympathetic, light-minded. and egoistic character. Frank obtains power on San Lorenzo by giving its dictator ice-nine, a decision that leads to the end of life on the Earth.

Bokonon, a priest on the island, dreams to turn the island they found into a utopia. He created a religion of Bokononism, Vonnegut invents new languages and creates terms like karass and Boko-maru , which seem to survive, in the heads of his “readers, his plots and even his jokes” (Allen 43).

Bokonon understands that all religions are founded on lies, but he tries to find some good in order to attract followers. Bokonon is a debunker, a demystifier, a mocker, an alternative voice through which Vonnegut can find the freedom to be as iconoclastic as he pleases. Bokonon is perhaps both more cheerful and more cynical. His ultimate philanthropic gesture, while judged “insane” or at least highly eccentric by others in positions of power or responsibility when looked at in light of either its limited but positive results or its ethical implications, appears eminently sane and even highly commendable (Bloom 33).

The novel takes place largely on an island paradise in the Caribbean, which stirs in us once more memories of that Master of Illusion, Prospero. Vonnegut describes the island as perfectly rectangular as a township in Kansas. Any restless soul, any soul seeking to find what lay beyond…” (Vonnegut 74). It is possible to say that this perfect and ideal setting is used in opposition to anarchy and false morals existing in society.

The Pioneers’ Day parade is another setting that helps to unveil false values and traditions preached by society. There is stock humor in this story, such as the descriptions of the bodies amphibians choose and how they behave in them, the Pioneers’ Day parade, and the trial, with its parody of a McCarthy hearing. Science fiction is not always as benign in its comic contributions. Variable gravity is one of the science-fiction devices that set up much of the plot of the novel, much as Ice-Nine does in Cat’s Cradle . Both of these transformations of nature have their comic potential as well as their catastrophic consequences (Bloom 36).

The main symbol of the book is ice-nine can be interpreted as an atomic bomb that can destroy everything in a second. For John-Jonah, however, it was to be a book about Hiroshima rather than Dresden, and in the end, he does not even manage that–his imagination (and Vonnegut’s) pre-empted not by the Atomic Bomb, which did not quite end the world, but by Hoenikker’s next, posthumous invention, which did: not by the final fire, but the final ice–a kind of super-nice, called Ice-Nine , which melts at 114 degrees Fahrenheit, and with which Hoenikker was playing like a child at the moment of his death (Giannone 23). Vonnegut depicts the substance as “containing a seed of ice-nine, a new way for the atoms of water to stack and lock, to freeze” (47).

Bokononism can be seen as a symbol of false truth and values followed by society. Indeed, the not-quite nihilism of the book’s close is a product of the tension between the religion of Bokononism, which advocates formulating and believing sacred lies, and the vision granted to the dwarfed son of the Father of the Bomb of the emptiness behind all lies, however sacred (Reed 42). Vonnegut depicts the religion: “Bokonon’s theory … is what he called “Dynamic Tension,” his sense of a priceless equilibrium between good and evil….” (102).

At any rate, is revealed as having experienced two great joys before his tale is told: one slow and long-continued, as he learns who are the other members of his karass , the handful of others in the world with whom, willy-nilly, he must work out the pattern of his destiny: one intense and momentary, as he plays footsie with the blonde Negress, Mona, whom he, and everyone else, loves: their naked soles touching in the union called by Bokononists “ Boko-maru .” “One of the central ironies of Cat’s Cradle is that through “the bittersweet lies” of the true-false religion Bokononism, the novel thematically quizzes itself” (Schulz 19).

Cat’s Cradle symbolizes the ability of a human race to change and shape the world around, and transformations caused by our actions. Through all historical periods, people try to shape and structure the world using religion and science. Once again, the central science fiction ingredient in the plot of this story is a throwaway device (Schulz 7).

The few absentminded steps necessary to set the would-be amphibian walking out of a body have about the same plausibility as Barnhouse’s aligning his brain cells. at the end of the novel, Bokonon comments: “If I were a younger man, I would write a history of human stupidity; and I would climb to the top of Mount McCabe… and I would make a statue of myself, lying on my back, and thumbing my nose at You Know Who” (Vonnegut 299). Vonnegut presents the unresolvable conflict of mutually exclusive theories; namely, the possibility of actual joy.

Writing books can be interpreted as a symbol of future hopes and the creation of history by humans. For instance, Bokonon, priest-philosopher writes his Books of Bokonon , whose tenets and calypsos are quoted frequently in the novel. He declares that all of his truths are shameless if harmless lies, once again raising the issue of the ethics of the writer. Bokonon invents versions of history and explanations of various of life’s mysteries with alacrity. In so doing he shares disregard for distinctions between fact and fiction (Goldsmith 87).

In many reviews, Cat’s Cradle is often described as a “black humor” novel in which Vonnegut created a new language and a new religion to convey how language and religion help to invent beliefs that provide meaning and purpose in the face of life’s paradox (Reed 45).

When the narrator begins the story with the quietly loaded statement, “Call me Jonah,” Vonnegut launches a literary irony of several dimensions. Critics (Giannone 43) admit that One aspect is of course that of the Old Testament prophet who was punished for his failure to carry God’s message of mercy to the Assyrians by being cast off a ship into a storm, swallowed by a whale, and then coughed up on dry land. Following Allen (69) Vonnegut touches upon issues of free will, population control with hyperbole and humor. He can thus express a philosophical point of view or make a moral judgment in a manner that may avoid the resistance argumentation might invite (Klinkowitz 45).

In sum, Cat’s Cradle does not offer readers a Happy Ending: the book begins and ends with a vision of the total destruction of mankind, to which only an eternal gesture of contempt is an adequate response. Vonnegut questions the role and importance of science in everyday life and its threats to humanity. Vonnegut seeks to make a fairly wide-ranging commentary in Cat’s Cradle , he faces the challenge of creating broad or multiple plot situations and categories of characters to permit his plausibly portraying many aspects of life. He minimizes the danger of the reader’s becoming confused and distracted. Cat’s Cradle vividly portrays the unique style of Vonnegut and his ability to join humor into the science fiction elements.

Works Cited

• Allen, William Rodney. Understanding Kurt Vonnegut . Columbia: University of South Carolina Press, 1991.
• Bloom, Harold, ed. Modern Critical Views: Kurt Vonnegut . Philadelphia, PA: Chelsea House, 2000.
• Giannone, Richard. Vonnegut: A Preface to His Novels . Port Washington, NY: Kennikat, 1977.
• Goldsmith, David H. Kurt Vonnegut: Fantasist of Fire and Ice . Bowling Green, KY: Bowling Green University Popular Press, 1972.
• Klinkowitz, J. Kurt Vonnegut, Jr. and the Crime of His Times. Critique , 12 (1971). 38-53.
• Reed, Peter J., and Marc Leeds, eds. The Vonnegut Chronicles: Interviews and Essays . Westport, CT: Greenwood Press, 1996.
• Schatt, S. The World of Kurt Vonnegut, Jr. Critique , 12 (1971), 54-70.
• Schulz, M. F. The Unconfirmed Thesis: Kurt Vonnegut, Black Humor, and Contemporary Art. Critique 12 (1971), 5-26.
• Vonnegut, K. Cat’s Cradle . New York: Holt, Rinehart and Winston, 1963, 1998.
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IvyPanda. (2021, September 7). Cat’s Cradle by Kurt Vonnegut. https://ivypanda.com/essays/cats-cradle-by-kurt-vonnegut/

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IvyPanda . 2021. "Cat’s Cradle by Kurt Vonnegut." September 7, 2021. https://ivypanda.com/essays/cats-cradle-by-kurt-vonnegut/.

1. IvyPanda . "Cat’s Cradle by Kurt Vonnegut." September 7, 2021. https://ivypanda.com/essays/cats-cradle-by-kurt-vonnegut/.

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When humans picture corals, we tend to think of words like “colorful,” “intricate,” and “bustling.” But the shallow tropical reefs that come to mind in the vivid style of “Finding Nemo” are only a small percentage of coral diversity on planet Earth.

Coral reefs have been around for hundreds of millions of years. Today, there are more than six thousand known species of coral that can be found almost anywhere in the ocean. Deep-sea corals can thrive as many as 10,000 feet below the surface, a realm of the ocean only accessible to researchers through remotely operated vehicles.

According to research zoologist Andrea Quattrini , the curator of corals at the National Museum of Natural History , corals are a great system to understand evolution in the deep sea because they occur across such a broad range of depth. Quattrini’s newest paper, published this week in the journal Proceedings of the Royal Society B: Biological Sciences , pushes back the evolution of bioluminescence in corals to at least 540 million years ago. This is much earlier than previously predicted and provides clues into how these ecosystem engineers colonized the abyss.

Quattrini also studies the origin of coral skeletal types and the current state of coral biodiversity worldwide. Understanding corals past and present provides the necessary knowledge to support coral conservation in the future.

I had no idea there were so many coral species living in the deep ocean. What is it about deepwater corals that captivates your research interests?

There’s something about the remoteness and the unknown of the deep sea that absolutely intrigues me. I fell in love with the ocean at a very young age and by the time I was twelve, I decided I wanted to be a marine biologist.

In 2003, I went on my first dive in a submersible, called the Johnson Sea Link. We went to a deepwater coral reef off South Carolina.  When I was 700 meters down under the surface of the ocean, exploring this deepwater coral reef that had been around far longer than I had, I felt humbled that I had the opportunity as a human to visit this place and see all the incredible life. And then we saw all this fantastic bioluminescence on the way back up. It made me think: “this world is an amazing place.”

After that, I told myself that I would study these ecosystems for as long as I could.

Why are corals such good subjects for studying evolution?

When people picture corals, they think of shallow, warm, sunny environments with these colorful coral reefs. But that’s actually not the majority of corals. There’s so much morphological diversity amongst corals. Some are arboreal, tree-like shapes. Some are sea fans. Others are soft corals, and some are solitary cup corals. The diversity of corals is much more than just reef-building corals.

They’re a great system to understand evolution in the deep sea because of this diversity and because they occur across this broad range of depths. They occur from shallow waters to the deep abyss, from the poles to the tropics. They face various environmental variables that change across depth. Because of this, they can help us answer questions about evolution in the deep sea.

Your recent finding about the origin of bioluminescence is making a big splash. Can you tell us how that research started?

About a decade ago, I was working on a coral sample while on a research cruise. The coral released a light, and that was amazing to see. Since then, I’ve had an interest in bioluminescence. In 2014, I started testing various coral species for bioluminescence, and since then I’ve been compiling a list. So now, every time I go out to sea, I test various corals for this trait.

I worked with a team of collaborators to use those observations of living corals to get a sense of the evolution of bioluminescence, through a process called ancestral state reconstruction. We don’t have any fossil records of bioluminescence, so you take the information from today in extant species, and you move backwards in time. The more living species that share a trait, the more likely it is that their ancestors are going to share that trait as well.

By working backwards, we found that the ability to bioluminesce has been in the genomes of these corals for hundreds of millions of years.

What do you hope people will learn from this paper?

The key point of our study is that bioluminescence arose 540 million years ago and has been retained all this time. That tells us that it must have been important for these organisms’ fitness.

What is also interesting is that it arose when animals were exploding and diversifying across the planet during the Cambrian period. It is likely that bioluminescence either enabled diversification of corals in the deep sea or was retained in those families that are most diverse in the deep sea.

Now that the paper is out, I’m excited that people will know that corals are pretty cool animals that can communicate with light. We're hoping that our paper will help also get more people to look for bioluminescence and further explore why it has been so important for so long.

Why is it important to have a detailed picture of coral diversity?

When I started my career, we didn’t really know a lot about deepwater corals, and we didn’t know a lot about coral genetics. But biodiversity is critical for ecosystem functioning. Biodiverse systems equal healthy ecosystems.

This past summer, there were huge heat waves that occurred off Florida, where corals went locally extinct. If a coral species goes locally extinct in one place, you want to be able to understand that species distribution to help protect it somewhere else. Species’ identities are fundamental to this understanding.

In addition, NMNH is leading efforts to help characterize that biodiversity across the northern Gulf of Mexico at sites where corals were injured from the oil spill. We're part of this project to help characterize that biodiversity so that scientists can restore corals and areas that were injured by the spill.

"Biodiversity is critical for ecosystem functioning. Biodiverse systems equal healthy ecosystems." 

— Andrea Quattrini, curator of corals NMNH

What do you like about working as a curator at the museum?

I like the fact that I can incorporate the coral collection into my research, and I can help grow the collection as well. But I also think being at a museum is so unique in terms of education and outreach for the public. And we have people from all over the world that come to our collections, so I get to interact with collaborators from across the world.

In our lab, we’re working toward a collective goal of better understanding coral systematics, but also just working to become better human beings and better scientists. 

This interview has been edited for length and clarity.  

Meet a SI-entist : The Smithsonian is so much more than its world-renowned exhibits and artifacts. It is a hub of scientific exploration for hundreds of researchers from around the world. Once a month, we’ll introduce you to a Smithsonian Institution scientist (or SI-entist) and the fascinating work they do behind the scenes at the National Museum of Natural History   

Related Stories  Smithsonian Expedition Yields a New Species of Deep-Sea Coral    Summer Summary: A Mysterious Fossil Tooth, Metallic Planet and Marine Hitchhikers   Meet the Smithsonian Scientist Who Has Spent Decades Exploring Ocean Depths    Scientists Cryopreserve and Revive Coral Fragments in a World First for Conservation

Naomi Greenberg | READ MORE

Naomi Greenberg is a Science Writing Intern with the Smithsonian’s National Museum of Natural History. She translates natural history research for general consumption in her writing for Smithsonian Voices as well as for the Smithsonian Ocean Portal. She is a senior at Georgetown University, where she founded and led the science section of the campus newspaper, The Hoya, in addition to studying biology and journalism. You can find more of her work  here .

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Pet Ownership and Quality of Life: A Systematic Review of the Literature

Kristel j. scoresby.

1 College of Social Work, University of Tennessee, Knoxville, TN 37996, USA; ude.ktu.slov@bserocsk

Elizabeth B. Strand

2 Veterinary Social Work, Colleges of Veterinary Medicine and Social Work, University of Tennessee, Knoxville, TN 37996, USA; ude.ktu@dnartse

Zenithson Ng

3 College of Veterinary Medicine, University of Tennessee, Knoxville, TN 37996, USA; ude.ktu@gnz (Z.N.); ude.ktu.slov@zlitsc (C.R.S.); ude.ktu.slov@lebortsk (K.S.)

Kathleen C. Brown

4 Department of Public Health, University of Tennessee, Knoxville, TN 37996, USA; ude.ktu@nworbck (K.C.B.); ude.ktu@osorrabc (C.S.B.)

Charles Robert Stilz

Kristen strobel, cristina s. barroso, marcy souza, associated data.

Data was not generated in this study.

Pet ownership is the most common form of human–animal interaction, and anecdotally, pet ownership can lead to improved physical and mental health for owners. However, scant research is available validating these claims. This study aimed to review the recent peer reviewed literature to better describe the body of knowledge surrounding the relationship between pet ownership and mental health. A literature search was conducted in May 2020 using two databases to identify articles that met inclusion/exclusion criteria. After title review, abstract review, and then full article review, 54 articles were included in the final analysis. Of the 54 studies, 18 were conducted in the general population, 15 were conducted in an older adult population, eight were conducted in children and adolescents, nine focused on people with chronic disease, and four examined a specific unique population. Forty-one of the studies were cross-sectional, 11 were prospective longitudinal cohorts, and two were other study designs. For each of the articles, the impact of pet ownership on the mental health of owners was divided into four categories: positive impact ( n = 17), mixed impact ( n = 19), no impact ( n = 13), and negative impact ( n = 5). Among the reviewed articles, there was much variation in population studied and study design, and these differences make direct comparison challenging. However, when focusing on the impact of pet ownership on mental health, the results were variable and not wholly supportive of the benefit of pets on mental health. Future research should use more consistent methods across broader populations and the development of a pet-ownership survey module for use in broad, population surveys would afford a better description of the true relationship of pet ownership and mental health.

1. Introduction

Throughout history, animals have played a significant role in society including in agriculture and pet ownership. A recent survey conducted in the United States estimated that approximately 67% of homes had at least one pet, equaling about 63 million homes with at least one dog and 42 million homes with at least one cat [ 1 ]. Pets can constitute a connection to nature, function in recreational and work activities, and provide companionship in our homes [ 2 , 3 , 4 ]. The importance of animals in our lives is founded on the human–animal bond concept, which is the “mutually beneficial and dynamic relationship that exists between people and other animals that is influenced by behaviors that are essential to the health and well-being of both” [ 5 ]. This concept has championed animals as companions and family members, leading to their essential part of everyday life for many. The human–animal bond has additionally driven the common belief that pets are good for human health, both physical and mental [ 6 , 7 , 8 ].

While there are some qualitative [ 9 , 10 ] studies that claim that pet ownership benefits people, particularly in regard to improved mental health, there are few studies with substantial evidence from large, diverse population samples to support this theory. The studies that have been published are often not substantiated with regard to study populations or methods, making broad conclusions difficult. Furthermore, some studies that have investigated the correlation between pet ownership and mental health have revealed no effect, or even worse, negative effects of pet ownership [ 11 , 12 , 13 , 14 , 15 ]. The inconsistencies in the literature and limitations of these studies warrant a thorough exploration of the effect of pet ownership on mental health outcomes among large, diverse population samples.

Two previous systematic reviews of the literature did examine the relationship between pet ownership and mental health/well-being [ 16 , 17 ]. Islam and Towel [ 16 ] did not find a clear relationship between pet ownership and well-being in the 11 studies included in their review. Similarly, Brooks et al. [ 17 ] examined the role of pets in owners with diagnosed mental health problems and found mixed results across the 17 studies included in the review. The purpose of this study was to perform a systematic review of the peer-reviewed published literature containing original research that examined the relationship between pet ownership and mental health for people in any population. Previous reviews included a smaller sample of research articles, often limited to a specific population of pet owners. By describing the relationship between pet ownership and mental health across all examined populations, this study will better inform whether pets could be recommended to help with mental health and whether promotion of the human–animal bond is generally beneficial.

2. Materials and Methods

The systematic review process involved a literature search, screening, extraction, and an assessment of the remaining articles by four researchers and three graduate students. For the purpose of this study, pet ownership was limited to dogs and cats. Our research team sought to answer, “How does ownership of a dog or cat influence the mental health or quality of life of pet owners?”

In May of 2020, the following databases were searched for peer-reviewed articles on pet ownership and mental health: PubMed and Web of Science. Utilizing Boolean search terms, the literature search was conducted using the terms: anxiety OR depressi* OR bipolar OR (mental* AND (health OR disease* OR disorder* OR condition* OR ill*) for the problem, (dog OR dogs OR cat OR cats OR canine* OR feline*) AND ((pet OR pets)) AND (owner* OR companion* OR interact* OR bond* OR “human animal bond” OR “animal human bond” OR “animal assisted”) for the intervention and health* AND (impact* OR outcome* OR status OR effect* OR affect* OR consequen* OR result*) for the outcome.

Although there was not an approved PRISMA protocol, the research team used Covidence (Melbourne, Australia), a software program that tracks the systematic review screening process. Identified articles were imported into Covidence, duplicates were removed, and the remaining articles were screened by the research team. Through random assignment, each article was independently reviewed by one faculty member and one graduate student. Each reviewer indicated in Covidence if the article should be included or excluded according to established criteria ( Table 1 ). When there was a conflict between reviewers, a third reviewer (non-student) resolved the conflict. The full review process is shown in Figure 1 . At the final review stage, two researchers independently extracted specific information ( Table 2 ) from each article. The type of impact on mental health was determined based on the results reported in each article.

Following a literature search, articles were reviewed for adherence to inclusion and exclusion criteria. A total of 54 articles were identified to meet all criteria.

Inclusion and exclusion criteria used for evaluation of research articles that examined the relationship between pet ownership and mental health.

At the extraction stage, the following information was used for evaluation of research articles that examined the relationship between pet ownership and mental health.

In addition to extracting the information outlined in Table 2 , an index ( Appendix A ) was created to assess article quality. The index was based on two previous systematic reviews of mental health in veterinary science [ 17 , 18 ]. Each dichotomous index question assigned a 0 if the article did not meet criteria and a 1 if the article did meet criteria. The higher the score an article received (0–9 points), the higher the quality of the article.

Interventionary studies involving animals or humans, and other studies that require ethical approval, must list the authority that provided approval and the corresponding ethical approval code.

The article review process and number of articles in each step are shown in Figure 1 . A total of 54 articles met the inclusion and exclusion criteria ( Table 1 ) and were systematically extracted ( Table 2 ). These articles were then divided into four categories based on the type of overall impact pets had on the mental health of owners: (1) positive impact (n = 17); (2) mixed impact ( n = 19); (3) no impact ( n = 13); and (4) negative impact ( n = 5). Factors that influenced mental health include (a) age (middle-aged female caregivers had more psychological stress than young female and male caregivers), (b) obedience and aggressiveness of the pet, (c) marital status (single women who owned a dog were less lonely and socially isolated than women without pets), and (d) attachment to the pet (high level of bonding has lower anxiety and depression scores than lower level of bonding) [ 19 , 20 , 21 , 22 , 23 , 24 ]. A few representative studies with mixed results include one examining the general population, which found that unmarried men who live with a pet had the most depressive symptoms and unmarried women who live with a pet had the fewest [ 19 ]. Another study examining the impact of companion animals on cancer patients found that mental health was associated with the status of cancer treatment, with those receiving intense treatment having poorer mental health [ 20 ]. In addition to overall impact, the study population, study type, population size, year of publication and article quality are reported ( Appendix B ).

Of the 54 articles, 19 (35%) were studies conducted in the general population, 15 (28%) were studies in older adult individuals, eight (15%) were in children and adolescents, six (11%) focused on people with some type of chronic physical illness/disease, three (6%) were studies in people with severe mental illness, and three (6%) studies examined unique populations. Of the 15 studies that had only older adult participants, none of them reported a positive impact. Seven of the articles reported mixed impact based on type of pet, gender, companionship, or another demographic. Six of the studies had no impact and two had a negative impact. Of the eight studies that involved children and adolescents, six of them indicated a clear positive impact, one indicated mixed impact, and one indicated no impact. Of the three studies that involved those with severe mental illness, two indicated clear positive impact and one indicated mixed impact.

Research studies either compared mental health outcomes in pet owners versus non-pet owners ( n = 41) or with regard to owner attachment to the pet ( n = 13). Similar to the overall distribution, the outcomes within these two different types of studies were distributed across all four categories ( Table 3 and Table 4 ). In 38% (five of 13) of the studies, attachment to a cat or dog was associated with a positive impact on mental health in 38% of the studies. Four of the 13 studies (31%) indicated mixed results, meaning that human–animal attachment sometimes was associated with better mental health and sometimes it was not. One example of higher attachment leading to worse mental health was for those amid cancer treatment [ 20 ]. There was no clear trend towards attachment and better mental health.

Outcomes of 41 studies that examined mental health outcomes in pet owners compared to non-pet owners.

Outcomes of nine studies that examined mental health outcomes in relationship to the pet owner’s attachment bond with their pet.

The study types included 41 (76%) cross-sectional studies, 11 (20%) prospective cohort longitudinal studies, and two (4%) other study designs. Of the cross-sectional studies, 27 (66%) found that companion animals had no or negative impact on mental health and 14 (34%) found mixed or positive impact on mental health. Of the 11 articles that reported on a longitudinal study design, five (45%) demonstrated no or negative impact and six (55%) demonstrated mixed or positive impact. Among the 54 studies, sample size ranged from 30 to 68,362.

To measure mental health constructs, 75 different validated scales were used ( Table 5 ). Eight scales were used to measure human attachment to pets. The most common scales used across studies were the CES-D (13 studies) to measure depression and the ULS (10 studies) to measure loneliness. Two scales were used by four studies each (DASS and any variation of GHQ). Three scales were used by three studies each (GDS, CABS, and any variation of PHQ). The remaining scales were used only once or twice across the studies assessed.

The scales used across studies to measure mental health.

Regarding the study quality scores ( Appendix A ), no articles received a quality score of 9, six (11%) received a score of 8, 11 (20%) received a score of 7, 20 (37%) received a score of 6, and 17 (31%) received a score of 5 or below. Of the articles with a quality scale score of 5 or lower, 18% (3) articles had no or negative impact and 82% ( n = 14) had mixed or positive impact on owner mental health. Articles with a quality scale score of 6 or higher, 43% ( n = 16) showed no or negative impact and 57% ( n = 21) showed mixed or positive impact.

4. Discussion

Understanding the nature of the relationship between mental health and pet ownership is important for both human and animal welfare and to better determine the impact of human–animal interactions. Over the years, the perspective that “pets are good for you” has become an assumption [ 25 ] and when negative implications are recognized it often relates to zoonotic diseases rather than human–animal interactions [ 26 ]. This belief in the positive aspects of the human–animal bond is strengthened by marketing tools used by the pet industry [ 27 ]. While there certainly is evidence that supports the benefits of the human–animal bond to people’s mental health [ 28 , 29 ], there is also clear and consistent evidence that the relationship is complex and sometimes negative [ 30 , 31 ]. The question of whether pets should be prescribed by health professionals is an especially important one. Recent qualitative research supports that attending to a pet can help a person manage mental health crises [ 32 ], however, doing so can also cause a person to rely on the pet instead of other evidenced based methods of seeking mental health support. The recommendation of obtaining a pet in the presence of mental illness ought to be coupled with other evidenced based strategies for mental health recovery such as increasing social support and engaging in third wave behaviorally based interventions such as Acceptance and Commitment Therapy or Dialectical Behavior Therapy.

The broad perspectives that pets are good for mental health may cause people to place false expectations on the role a dog or cat must play in their lives [ 33 ]. The anthropomorphism of pets (people placing human cognitive motivations on pets’ behavior and treating pets as people) can in fact have a negative impact on the animal’s welfare [ 34 ]. The untreated stress of people who turn to their pets instead of their human social supports and health professionals may in fact be causing pets to be more stressed [ 35 ]. Although initial data suggest relinquishment rates were not higher after COVID-19 lockdowns were lifted [ 36 ], some still have concerns that the recent increase in pet adoptions from shelters may result in pet relinquishment once the pandemic is more managed and people return to their daily work environments [ 37 ] (J. Schumacher personal communication, 5 May 2021). Developing clear guidelines about the benefits and liabilities of pet ownership and mental health is important to mitigate the public halo effect that suggests that simply acquiring a pet will improve your mental health.

Previous systematic reviews of the literature have found mixed results regarding the relationship between mental health and pet ownership [ 16 , 17 ]. Our search and review methodology was similar to Islam and Towel [ 16 ], which yielded 11 studies compared to the 54 studies compiled in this review. Although the Brooks et al. [ 17 ] review yielded 17 studies, they limited their search to studies only including people diagnosed with mental health conditions. While the current study did examine a larger body of research that covered broader populations and more recent publications than previous reviews, the findings were similar in that results varied across outcomes including positive, negative, mixed, and negligible. Unlike previous studies, this review also differentiated studies that compared pet owners to non-pet owners and studies that examined the level of attachment with a pet as a predictor of the mental health of the owner. Islam and Towel [ 16 ] argued that the definition of pet ownership needs to be defined across all studies, including aspects of length of ownership, time spent with the animal, and perceived quality of the interaction. Within these two categories of study types, the outcomes still varied and showed no consistent evidence that pet ownership is a positive contributor to mental health. The lack of consensus from these studies was not surprising. While popular literature and media consistently highlight the positive, it rarely highlights the negative aspects of pet ownership. In fact, studies with negative or non-significant findings are often subject to the “file drawer” effect, in which authors ultimately decide not to publish their studies [ 15 ]. In this review, we did find and include studies that reported negative or mixed findings.

The authors made the decision a priori to divide the results into categories based on the type of impact each study had on mental health. Among the 17 studies that were determined to have positive results, most of the studies were with children and adolescents ( n = 6) and the general adult population ( n = 6). There were some challenges to identifying these studies as clearly positive. Because a variety of different variables and a variety of different methodologies were used based on the specific purpose of each study, they could not be directly or easily compared to one another. Many of the positive impact studies investigated additional variables that could be better predictors of positive mental health than dog/cat ownership. For example, several studies indicated that children or adolescents with a dog had less depression and/or less anxiety than peers without a dog. However, family dynamics such as single parent or two parent households, time parents spend at work, presence of siblings, and family dysfunction [ 2 , 8 ] may be more significant contributors to child mental health than dog ownership.

The 19 mixed impact studies were easier to categorize because of conflicting outcomes, particularly for studies with an older adult or general adult population. In each of these studies, the direction of the outcome was influenced by demographic variables (such as gender) or the type of pet (cat or dog). For example, one general population study determined that women with pets had lower levels of depression whereas men with pets had higher levels of depression [ 19 ]. Another example is that pet-owning individuals with severe mental illness had less psychiatric hospitalizations than non-pet owning peers, however, they also had higher levels of substance use [ 38 ]. Another reason why a study would be categorized as mixed impact is if mental health was assessed using multiple instruments and yielded conflicting results. For instance, one study indicated that when compared to people without pets, those with pets had no difference in anxiety or stress scores yet had higher depression scores [ 22 ].

For the 13 studies that had no impact, most were with the older adult ( n = 6) and general adult ( n = 4) population. These studies concluded that when comparing pet ownership to non-pet ownership or when comparing attachment levels, the pet had no correlation with positive or negative mental health. Many of these studies controlled for demographic variables such as age, gender, and socioeconomic status in their statistical models. One challenge to categorizing the studies was that study participants subjectively believed their pets were helpful to their mental health despite what validated measures showed. The inclusion of these biased observations in an attempt to still put a positive spin on the study may reflect the conflict a researcher has in publishing negative results. An additional challenge is that studies that included non-mental health measures (such as physical health) showed that those with pets did better than those without. Expert reviews of pet ownership on cardiovascular health have demonstrated a significant challenge to reach a definitive conclusion of the impact of pet ownership on health based on the current evidence [ 39 ].

Five studies demonstrated a clear negative impact between pet ownership and mental health. The sample populations were general ( n = 2), older adults ( n = 2), and single adults living alone ( n = 1). In these studies, pet ownership was associated with higher levels of depression, loneliness, and other psychological symptoms across all demographic variables and type of pet (dog or cat). Again, the challenge to classifying these studies as negative impact suggests that pet ownership causes increased levels of mental health illnesses, when in reality, the studies are about correlation, not causation. There may be other factors that cause the samples in these studies to have worse mental health. As indicated by Mullersdorf et al. [ 40 ], the presence of a psychological condition could predispose individuals to become pet owners, making it difficult to truly know if pet ownership causes a negative impact on mental health. These studies, regardless of type of outcome, only indicate association of pet ownership and mental health.

Another challenge in comparing the 54 studies was the difference in methodology and quality of each study. Due to this, our methods did not evaluate the individual and overall power and effect sizes of study results. Quantitative methodologies are warranted in this field, particularly prospective, randomized, double-blind, placebo-controlled intervention trials that are longitudinal in design to provide evidence of the impact of animal ownership over time while eliminating as many extraneous and confounding variables as possible [ 41 ]. Ideally, this truly experimental model of pet ownership would include random assignment of companion animals in a closed system to eliminate as many sources of error variance as possible [ 42 ]. However, due to the nature of pet ownership being integrated as a part of daily life on a voluntary basis, this experimental model would be difficult to achieve. Perhaps the most compelling of all studies that comes closest to this design was a prospective interventional study in which 71 previous non-pet owners were given a cat or dog; results demonstrated mild benefits in mental health and behavior after 10 months of pet ownership compared to the 26 non-pet owners [ 43 ]. While noteworthy, there was lack of randomization, so the pet ownership group consisted of a relatively small number of subjects who were searching for a pet to adopt rather than receiving it on random chance. Regardless, this study still reports an improvement in mental health in this specific population. Future studies should strive to achieve this prospective, controlled, experimental methodology to more compellingly connect pet ownership with mental health.

A quality index attempted to rate the rigor of each study, but the index was subjective and based on questions that could be asked without statistical analysis (e.g., does this study include a comparison population?). The higher the score on the quality index, the more likely the study was scientifically rigorous. The lower the score, the more likely the study was to demonstrate a positive or mixed impact on the pet owner’s mental health. While both previous literature reviews critiqued the rigor of the studies reviewed and remarked upon the consistent methodological flaws, Islam and Towel did not assign objective scores to the 11 studies reviewed. Brooks et al. [ 17 ] did assign quality scores to each of the 17 studies reviewed but did not evaluate the impact of the quality of the study on its results. The quality scores in the current review varied across all four outcome categories and did not give any indication of quality impacting the overall outcome. Still, it is important that researchers strive for higher quality research that carries more weight in the question of whether pet ownership truly impacts mental health. Additionally, we recommend that studies be replicated in an attempt to corroborate previous findings, which contribute to the overall understanding of the phenomenon.

Lastly, this study also examined how mental health was evaluated across the studies. For the 54 studies included in this review, 75 different scales ( Table 5 ) were used with many research studies implementing more than one scale ( Appendix B ). While most of the scales used have been previously validated, the inconsistent use of scales makes comparison of results across studies challenging. While it is common to utilize an instrument that is a validated self-report of depression, it is likely that researchers often utilize other scales because they are investigating other aspects of mental health such as loneliness, stress, and anxiety. Many scales also rely on self-reporting of mental health indicators, which can be affected by inherent bias, especially when completing a survey regarding mental health and pet ownership. To allow for better comparison of future studies, researchers should attempt to use consistent measures of mental health across studies, such as the CES-D [ 44 ], which was the most commonly used scale in 13 of the 54 examined studies.

In addition to consistent use of mental health scales across studies, the development of a module for use in wide-scale population surveys with a focus on pet-ownership would benefit future research examining the relationship between pet ownership and health. The Behavioral Risk Factor Surveillance System (BRFSS) [ 45 ] is an annual questionnaire administered by the US Centers for Disease Control and Prevention. There are 14 core sections that are administered to all participants and 31 optional modules [ 45 ]. None of these modules focuses on pet ownership and the addition of such a module would allow for a more in-depth evaluation of the relationship between pet ownership and health, both mental and physical, across large populations. While pets can play a significant role in the owner’s health, it can be difficult to differentiate the effects of pet ownership from the many other factors that contribute to one’s mental and physical health. The addition of a pet ownership module to the BRFSS would allow researchers to examine the role of pet ownership in tandem with other factors that contribute to health. On a smaller scale (approximately 3000 participants), the General Social Survey (GSS) is a representative survey that monitors trends in opinions, behaviors, and demographics among Americans [ 46 ]. Though not a main focus, the GSS does include pet ownership and mental health variables. Including pet ownership allows researchers who study the relationship of ownership with humans to have a large, representative dataset to analyze correlations. For example, a recent study used the GSS 2018 to examine demographics of pet ownership [ 46 ]. In their conclusion, the authors of this study indicated that the strengths of using the GSS to study pet ownership characteristics are high quality data, multiple covariates, sound methodology, and easy access [ 47 ]. Including pet ownership questions in multi-wave, representative studies would further the work of human animal relationship research.

This systematic review was limited due to only searching two databases and only evaluating research published in English. The majority of studies focused on pet-owners in Western cultures. The human–animal bond may differ across cultures and future studies should include pet-owners in non-Western cultures. However, a large number of articles were identified, and the total number of articles included in final extraction was greater than similar previous systematic reviews. More consistent methods across research that evaluates the relationship between pet ownership and mental health might allow for more extensive comparison of studies.

5. Conclusions

Previous research examining the impact of pet ownership on mental health has shown mixed results and the results of this study were the same. While there were more absolute numbers of studies to demonstrate a positive impact ( n = 17) compared to negative impact ( n = 5) on mental health, the overall results indicate a much more complicated picture. While 17 of the 54 studies had a clear association of pet ownership and positive mental health, the remaining 37 articles show a mixed association, no association, or a negative association. Comparing these studies is quite challenging due to the number of measures used to assess mental health, the differences in study quality, and the variety of variables that were controlled for. While research studies can be improved by addressing limitations as described, a more comprehensive evaluation of behavior and its association with health outcomes is warranted. We also cannot ignore that mental health is multifactorial. Pet ownership and the resulting human–animal interaction is a single factor; other factors that also contribute to mental health should be examined in large populations of pet-owners and non-pet-owners. The addition of a pet-ownership specific module to the BRFSS, as previously described, would allow for prospective research that can be replicated, and eventually retrospective research, that will also allow for inclusion of other factors that contribute to health.

Following a literature review and data extraction of research articles that examined the relationship between pet ownership and mental health, the following articles were found to meet inclusion and exclusion criteria as outlined in Table 1 .

Author Contributions

Conceptualization, K.J.S., E.B.S. and M.S. methodology, E.B.S. and M.S.; validation, all; formal analysis, all; data; writing—original draft preparation, K.J.S., E.B.S., M.S., Z.N.; writing—review and editing, K.J.S., E.B.S., Z.N., K.S., K.C.B., C.R.S., C.S.B. and M.S.; supervision, E.B.S., Z.N., K.C.B., C.S.B. and M.S.; project administration, E.B.S. and M.S.; funding acquisition, E.B.S. and M.S. All authors have read and agreed to the published version of the manuscript.

This research was funded by Maddie’s Fund.

Institutional Review Board Statement

Not applicable.

Informed Consent Statement

Data availability statement, conflicts of interest.

The authors declare no conflict of interest.

Publisher’s Note: MDPI stays neutral with regard to jurisdictional claims in published maps and institutional affiliations.

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Title: the collective use and evaluation of generative ai tools in digital humanities research: survey-based results.

Abstract: The advent of generative artificial intelligence (GenAI) technologies has revolutionized research, with significant implications for Digital Humanities (DH), a field inherently intertwined with technological progress. This article investigates how digital humanities scholars adopt, practice, as well as critically evaluate, GenAI technologies such as ChatGPT in the research process. Drawing on 76 responses collected from an international survey study, we explored digital humanities scholars' rationale for GenAI adoption in research, identified specific use cases and practices of using GenAI to support various DH research tasks, and analyzed scholars' collective perceptions of GenAI's benefits, risks, and impact on DH research. The survey results suggest that DH research communities hold divisive sentiments towards the value of GenAI in DH scholarship, whereas the actual usage diversifies among individuals and across research tasks. Our survey-based analysis has the potential to serve as a basis for further empirical research on the impact of GenAI on the evolution of DH scholarship.

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