Cat remains from Poland dated to 4,200 to 2,300 y BCE are currently the earliest evidence for the migration of the Near Eastern cat (NE cat), the ancestor of domestic cats, into Central Europe. This early immigration preceded the known establishment of housecat populations in the region by around 3,000 y. One hypothesis assumed that NE cats followed the migration of early farmers as synanthropes. In this study, we analyze the stable isotopes in six samples of Late Neolithic NE cat bones and further 34 of the associated fauna, including the European wildcat. We approximate the diet and trophic ecology of Late Neolithic felids in a broad context of contemporary wild and domestic animals and humans. In addition, we compared the ecology of Late Neolithic NE cats with the earliest domestic cats known from the territory of Poland, dating to the Roman Period. Our results reveal that human agricultural activity during the Late Neolithic had already impacted the isotopic signature of rodents in the ecosystem. These synanthropic pests constituted a significant proportion of the NE cat's diet. Our interpretation is that Late Neolithic NE cats were opportunistic synanthropes, most probably free-living individuals (i.e., not directly relying on a human food supply). We explore niche partitioning between studied NE cats and the contemporary native European wildcats. We find only minor differences between the isotopic ecology of both these taxa. We conclude that, after the appearance of the NE cat, both felid taxa shared the ecological niches. Synanthropization of the wildcat developed from occasional commensalism, initiated by availability of synanthropic rodents in agricultural landscapes. Through times, it led to nearly full dependency on anthropogenic resources and behavioral The cat’s way to domestication is a complex and still unresolved topic with many questions concerning the chronology of its dispersal with agricultural societies and the nature of its evolving relationship with humans. The Near Eastern wildcat Felis silvestris lybica is the only subspecies of wildcat that has been domesticated ( 15 ). It is native to Northern Africa and the Near East. This subspecies is the ancestor of all modern domestic cats, Felis silvestris catus . Both wild and domesticated forms are very close genetically and cannot be discriminated with mitochondrial DNA (mtDNA) analysis ( 15 ). We expect a similar effect in all synanthropic animals foraging on plants from manured fields, including domesticated ungulates fed with grains or straw ( 47 ) or grazing in manured pastures, in dogs eating similar foods as their owners ( 48 ), and in rodent pests foraging on the crop grains. Manuring of fields by Late Neolithic farmers has been identified at several sites ( 49 , 50 ) situated about 20 to 65 km from sites with remains of Late Neolithic NE cats. Humans and domestic dogs seemed to be very close isotopically to each other and showed the highest δ 15 N among all analyzed samples. Wild birds and herbivorous/omnivorous mammals (domestic mammals, rodents, and leporids) showed high variability in δ 15 N and δ 13 C signals ( Fig. 3 and SI Appendix , Table S2 ). Among the suite of species we analyzed, we assumed that birds, rodents, and leporids were potential prey of felids (both NE cats and European wildcats), which is in agreement with known dietary habits of modern felids ( 57 – 59 ). The first isotopic report on commensal behavior of ancient wild felids examined Neolithic cats from China ( 60 ). Based on δ 13 C and δ 15 N values of bone collagen, Hu et al. ( 60 ) identified a substantial consumption of millet-based food by humans, rodents, and cats, which suggested a possible commensal or even mutualistic behavior of Neolithic cats. However, lack of data for contemporary rodents and other possible prey limited the interpretative potential of those results ( 61 ). Furthermore, morphometric verification of Chinese cat remains revealed that the study involved a leopard cat ( Prionailurus bengalensis ) rather than a wildcat or domestic cat ( 62 ). The high proportion of prey from cluster B is especially relevant because this group likely represents pests of farmed crops. These NE cats certainly lived in a human-modified environment and were involved in the synanthropic food web. Drawing a broader picture of relationships between the NE cats and Neolithic people is limited by taphonomic factors and depositional contexts of the sampled remains. Firstly, the number of yet discovered remains is low. With only several specimens in hand, we have to consider that our results may represent individual-related behaviors rather than population-scale trends. In the alternative scenario assuming that NE cats were still-wild synanthropes following the farmlands, our specimens may represent the part of the population which was more oriented toward exploiting forest resources. However, each scenario implies that NE cats coexisted and shared the ecological niches with native European wildcats as early as the Neolithic Period. Conclusions One of the key issues for understanding the process of domestication is to determine the type of ecological relationships that existed between humans and a given species. Among the wild animals which have been domesticated, the cat ancestors were unique due to their solitary, territorial behaviors. This provides serious implications for the history of wildcat gene pool contamination by NE/domestic cats and for the conservation of this species. However, the full understanding of this past hybridization requires further nuclear DNA studies of fossil specimens. How close the relationship was between Late Neolithic NE cats and humans that once inhabited present-day Poland, and whether those cats were already domesticated, is still an open question. Searching for cat remains among archaeozoological material from Neolithic settlement sites may provide an insight into the human/cat relationship.

The ancient Egyptians mummified an abundance of cats during the Late Period (664 - 332 BC). The overlapping morphology and sizes of developing wildcats and domestic cats confounds the identity of mummified cat species. Genetic analyses should support mummy identification and was conducted on two long bones and a mandible of three cats that were mummified by the ancient Egyptians. The mummy DNA was extracted in a dedicated ancient DNA laboratory at the University of California - Davis, then directly sequencing between 246 and 402 bp of the mtDNA control region from each bone. When compared to a dataset of wildcats (Felis silvestris silvestris, F. s. tristrami, and F. chaus) as well as a previously published worldwide dataset of modern domestic cat samples, including Egypt, the DNA evidence suggests the three mummies represent common contemporary domestic cat mitotypes prevalent in modern Egypt and the Middle East. Divergence estimates date the origin of the mummies' mitotypes to between two and 7.5 thousand years prior to their mummification, likely prior to or during Egyptian Predyanstic and Early Dynastic Periods. These data are the first genetic evidence supporting that the ancient Egyptians used domesticated cats, F. s. catus, for votive mummies, and likely implies cats were domesticated prior to extensive mummification of cats.

Cat domestication likely initiated as a symbiotic relationship between wildcats (Felis silvestris subspecies) and the peoples of developing agrarian societies in the Fertile Crescent. As humans transitioned from hunter-gatherers to farmers ~12,000 years ago, bold wildcats likely capitalized on increased prey density (i.e., rodents). Humans benefited from the cats' predation on these vermin. To refine the site(s) of cat domestication, over 1000 random-bred cats of primarily Eurasian descent were genotyped for single-nucleotide variants and short tandem repeats. The overall cat population structure suggested a single worldwide population with significant isolation by the distance of peripheral subpopulations. The cat population heterozygosity decreased as genetic distance from the proposed cat progenitor's (F.s. lybica) natural habitat increased. Domestic cat origins are focused in the eastern Mediterranean Basin, spreading to nearby islands, and southernly via the Levantine coast into the Nile Valley. Cat population diversity supports the migration patterns of humans and other symbiotic species. Archeological discoveries of human remains and artifacts in the Near East and the middle Yangtze and Yellow Rivers in China indicate the earliest emergence of complex civilizations (Baldwin 1975 ; Bar-Yosef 1998 ; Hu et al. 2014 ). The Indus Valley of modern-day Pakistan is also argued as a historical center for agricultural development (Bellwood et al. 2005 ). Recent studies of Chinese random-bred cats and the local wildcat species/subspecies ( F.s. bieti ) suggest the noted introgression of this wildcat with random-bred cats in China does not explain the distinctive genetics of Far Eastern and Western European random-bred cats; further, the agricultural center near the middle Yangtze and Yellow Rivers is likely not a second domestication site for cats (Yu et al. 2021 ). While these previous studies all support the domesticated F.s. catus arose from F.s. While they have a low survival rate in the wild, their high reproductive capacity increases population size (Nutter et al. 2004 ). As apex predators, this reversion capability has often been exploited to eradicate invasive animals from island populations, whereas later, the cats themselves became invasive alien species (Rendall et al. 2021 ; Plein et al. 2022 ). Here, the random-bred cats of the study represent semi-domesticated animals that lie somewhere between “habituation” and “commercial breeds and pets” on the commensal domestication trajectory (Zeder 2012 ; Larson and Burger 2013 ). For cats, human assistance is not necessarily required for mating, shelter, safety, or the procurement of food (Driscoll et al. 2009 ). The cat’s semi-domesticated behavioral state is consistent with weaker human-influenced artificial selection pressures on the species. Although cats may have been domesticated at approximately the same time as many agricultural species, ~8000–10,000 years ago, cats have scavenged refuge piles and curbed vermin populations during their symbiotic relationship with humans (Clutton-Brock 1988 ). Therefore, for the past several thousand years, cats have not been transformed drastically in form or function, unlike dogs and economically important species. Only for the past ~200 years, cat breeds, not random-bred cats, have been selected for mainly monogenic aesthetic traits undergoing novelty selection on a small number of loci and likely a small portion of the genome. Minor structural differences and no functional behavioral differences were present in cats when the first cat show took place in 1871 (The Cat-Show 1871 ). The semi-domesticated nature of random-bred cats makes them an excellent resource to understand cat population origins, domestication, and dispersal. Studies using ancient DNA of domesticated cats may reveal a more complicated process (MacHugh et al. 2017 ), but the pattern revealed in random-bred cats is striking and agrees with archeological evidence. Unlike many domestication studies that must use modern breeds for comparisons, these random-bred cats have likely had less selection, weaker founder effects, and lower genetic loss by drift since cats are under fewer constraints by humans. The cat diaspora is relatively more recent than for humans or canines. Although cats and agricultural species serve very different purposes to humans, the geographic patterns of admixture in cats are a near-perfect reflection of admixture and migration in cattle populations, such as along the Silk Road and in the Americas (Decker et al. 2014 ). Along with archeological and genetic data, even the cat’s prey, house mice, have also represented bio-proxies for human migration patterns (Rajabi-Maham et al. 2008 ; Jones et al. 2013 ; Cucchi et al. 2020 ; Li et al. 2020 ). catus is suggested as the coastal regions of the Mediterranean Basin of the Fertile Crescent where cats have high observed heterozygosity and a short genetic distance to the progenitor subspecies. As highly agrarian societies developed, domesticated cats then spread down into the Nile Valley where cultural integration of felines into society slightly decreased heterozygosity and increased the genetic distance from the initial founders. The slightly lower diversity could also be an influence of ancient cultural selections. ornata from Iraq, Iran, the Indus Valley region, and Northwestern India could further explain the genetic variation seen in cat populations. Genetic and archeological studies from pre-farming cats would be an important addition in further clarifying the cat domestication process. The patterns of genetic diversity and differentiation observed in worldwide random-bred cats parallel those of other species, especially humans once they became farmers, suggesting human history is written in the DNA of domesticated species. Supplementary information Supplementary Tables Supplementary Figures Supplementary File 1 Supplementary File 2

The domestication of plants and animals permitted the development of cities and social hierarchies, as well as fostering cultural changes that ultimately led humanity into the modern world. Despite the importance of this set of related evolutionary phenomena, scholars have not reached a consensus on what the earliest steps in the domestication process looked like, how long the seminal portions of the process took to unfold, or whether humans played a conscious role in parts or all of it. Likewise, many scholars find it difficult to disentangle the cultural processes of cultivation from the biological processes of domestication. Over the past decade, the prevailing views among scholars have begun to shift towards unconscious and protracted models of early domestication; however, the nomenclature used to discuss these changes has been stagnant. Discussions of early domestication remain bound up in prevailing definitions and preconceived ideas of what the process looked like. In this paper, we seek to break down definitions of domestication and to construct a definition that serves equal utility regardless of the views that researchers hold about the process.This article is part of the theme issue 'Unravelling domestication: multi-disciplinary perspectives on human and non-human relationships in the past, present and future'.

s. lybica) includes domesticated cats, and that wild cats from this group were almost indistinguishable from domesticated cats. Phylogenetic analysis The domestication of the cat can be traced back to Near-Eastern and Egyptian populations of the African wildcat, Felis lybica, from which emerged the domestic cat (F. catus). Both belong to the Felis lineage of the family Felidae, which is thought to have first diversified around 12 to 13 million years ago. Several investigations have shown that all domestic varieties of cats come from a single l The domestication of the cat can be traced back to Near-Eastern and Egyptian populations of the African wildcat, Felis lybica, from which emerged the domestic cat (F. catus). Both belong to the Felis lineage of the family Felidae, which is thought to have first diversified around 12 to 13 million years ago. Several investigations have shown that all domestic varieties of cats come from a single lineage of F. lybica, although early interactions between wildcats and Neolithic human settlements occurred across a broad geographic range. Originating in the Near East, the first domestic cats spread from the ancient Mediterranean world via maritime trade as well as the Silk road, and later worldwide through European colonization. A separate but ultimately unsuccessful domestication of Leopard cats in ancient East Asia demonstrates that prolonged proximity to humans does not inevitably lead to domestication. The domestication process was protracted and geographically complex, rather than being a single discrete event. The earliest interactions between wildcats and humans… A 2014 study compared cat genomes with tiger and dog genomes. Genomic regions under selection in domestic cats included those involved in neuronal processes (fear and reward behavior) and in homologous recombination (increased recombination frequency). In addition, the KIT mutations responsible for the white-spotted phenotype were identified. Domestic cats have unusually high single nucleotide variant diversity compared with most other domestic animals, approximately 9.6 million in an average specimen, compared with the 4 to 5 million SNVs typically found in a single human. This peculiarity reflects the unusual nature of cat domestication, as rather than undergoing strong selective breeding and population bottlenecks typical of domestic species, cats effectively "self-domesticated," without losing ancestral traits, such as their hunting behavior. Intensive selective breeding in cats only began approximately 200 years ago and has focused almost entirely on aesthetic genetic traits. The blotched tabby cat trait (Aminopeptidase Q mutation) arose in the Middle Ages. Wild-type cats have a mackerel pattern.