COMMENTARY Cats as predators and early domesticates in ancient human landscapes COMMENTARY Fiona Marshalla,1 Cats have had diverse relationships with humans, ranging from rodent control, to household pets, to 1 cultural icons. The presence of rodents in grain stores in Neolithic farming settlements is widely thought to have led to the domestication of cats (1–3). Felid 2 5 bones are rare in archaeological sites, however, and processes of cat domestication and dispersal are not 3,4 well understood. A study by Krajcarz et al. (4) in PNAS reports on the initial spread of Near Eastern cats (Felis silvestris lybica/Felis catus), from southwest Asia into Europe ∼6,200 to 4,300 ya. Their unique data include Neolithic skeletal material from several Near Eastern cats found outside human settlements, and their analysis demonstrates that cats associated with early farmers in synanthropic or commensal relationships (5), hunting in F. silvestris silvestris agricultural landscapes. By documenting ancient ecolog- F. silvestris lybica ical relationships in humanly modified landscapes, this F. silvestris ornata F. silvestris cafra isotopic research contributes to understanding land- F. silvestris bieti scape scale human niche construction (6, 7) and to future – directions in domestication research (8 10). Fig. 1. Early interactions between humans and cats. Location of sites with The earliest cats discovered to date were buried in Near Eastern cats (F. s. lybica and F. catus) and leopard cats (P. bengalensis) early agricultural settlements, and scholars have pro- feeding in early farming food webs (1, 5), early spread of Near Eastern cats posed commensal and mutualistic domestication path- outside their wild range (1, 2), cat burials (2, 3), and household cats in Egyptian cities (4). Key: 1, Southern Polish sites; 2, Cyprus, ways for their domestication (1–3). Direct archaeological Shillourokambos; 3, Hierakonpolis; 4, Middle Kingdom Egypt, e.g., Thebes; 5, evidence for a commensal relationship between millet Quanhucan. Image credit: Lorraine Hu (Washington University in St. Louis, St. farmers and small felids feeding within the human food Louis, MO). Information on F. sylvestris distribution from ref. 1. Site locations from web was documented at Quanhucan in China (Fig. 1) refs. 3, 4, 11, and 14. (∼5,560 to 5,280 ya) (11), where leopard cats (Prionailu- rus bengalensis) (12) hunted rodents in a farming village sylvestris cf. lybica) with a person in a large farming village ecology and may have even been fed. However, this has suggested an early close relationship (3). In Egypt, the relationship did not lead to full domestication (12). elite human cemetery at the Predynastic city of Hierakonp- Evidence for commensal relationships that resulted in olis preserves evidence of subsidiary animal burials (wild domestication comes from agricultural villages and towns and domestic) including five cats (Felis sylvestris)datingto of the eastern Mediterranean and the Nile, in social ∼5,800 ya (14) (Fig. 1). Fine-grained research on birth sea- settings in which animals that were never domesticated sonality and morphology suggests phenotypic change (e.g., fox, jungle cat, and baboon) also played prominent resulting from selection on cats in villages and towns along roles. Paradigm-shifting research in Cyprus revealed that the Nile (14). It is not until 4,000 ya, when art and writing mice colonized the island with early farmers by 10,000 ya, indicate that domestic cats played a role in family life and and a range of other wild and domestic animals were state religion in Egypt, that most scholars think cats were intentionally transported, including foxes and Near East- fully domestic (15). The hieroglyphs for “female cat” used ern cats, ∼9,500 ya (8, 13) (Fig. 1). The burial of a cat (Felis as a girl’s nickname and wall paintings illustrate the role of aDepartment of Anthropology, Washington University in St. Louis, St. Louis, MO 63130 Author contributions: F.M. wrote the paper. The author declares no competing interest. Published under the PNAS license. See companion article, “Ancestors of domestic cats in Neolithic Central Europe: Isotopic evidence of a synanthropic diet,” 10.1073/pnas. 1918884117. 1Email: [email protected]. www.pnas.org/cgi/doi/10.1073/pnas.2011993117 PNAS Latest Articles | 1of3 Downloaded by guest on September 25, 2021 cats in women’s lives and households (16, 17). However, none of these food web. The authors also examine environmental change studies has explored the influence of humanly modified landscapes on through time in the trophic niche of European wildcats. Their data ancient cat diets outside of villages and towns, or the extent to which indicate that indigenous and exotic cats shared an ecological cats were provisioned or independent. This is, in part, because of a niche but that European wild cats had a broader diet and preyed focus on human intentionality in domestication and because of the lack more on migratory birds than did Near Eastern cats. of data on ancient cats. The remaining question of whether the cats in southern Poland were wild camp followers (synanthropic and commensal) or a feral Cats Living Outside Settlements population with domestic counterparts in settlements might—as Krajcarz et al. (4) used an isotopic approach to examine diets and Krajcarz et al. (4) suggest—be addressed in the future by discov- ecological and social relationships among people and the earliest eries of cat remains in farming settlements. It is not at all clear, Near Eastern cats during the spread of farming into Europe in the however, what biometric or genetic difference might be expected Neolithic (Fig. 1). Their findings bear on the question of whether as early as 6,000 ya between wild living (feral) domestic cats (Felis the cats studied in southern Poland were house cats, feral cats, catus) and synanthropic wild F. s. lybica hunting in an agricultural or whether they were wild Near Eastern cats that associated landscape far from its wild Asian range. Phenotypic change may be synanthropically with farmers and their fields and moved into the minimal, given the lack of evidence for strong directional selection wild range of European wild cats as an invasive species. Stable in cat domestication. carbon (δ13C) and nitrogen (δ15N) isotope signatures were col- From a binary perspective, whether the cats are considered lected for a broad range of Late Neolithic (ca. 4200 to 2300 BCE) wild or domestic matters. However, given our current un- taxa including cats, humans, dogs, birds, and rodents as well as derstanding of the significance of relationships to domestication comparative information from contemporary European wildcats (18), the prevalence of gene flow (19), and the lag in subsequent and from domestic cats in Roman contexts. These data docu- morphological or genomic shifts generally (e.g., goats and don- ment food webs in which cats fed and allow sophisticated dis- keys) (10, 18–20), the question of classification is less interesting crimination among potential cat prey taxa with varied isotopic than understanding of the influence of selection in human envi- signatures. The context of the cat specimens and the size of the ronments on cat domestication and spread. sample of rodents and birds studied are unprecedented in cat domestication research. Since cats are rarely eaten and their Cats in Ancient Human Landscapes bones seldom found in trash dumps even in villages, the chance Krajcarz et al.’s (4) findings change the conversation regarding cat of finding them in human landscapes distant from human set- domestication by showing that Near Eastern cats, loosely associ- tlements is extremely low. The six Late Neolithic Near Eastern ated with early farmers, were able to expand their range into cat skeletons they studied were recovered from nonanthropogenic Europe by moving with settlements, livestock, agriculture, and contexts in four cave sites distant from human villages. These forest clearance. Instead of focusing on temporal shifts from provide a unique window into human relationships with cats and Neolithic villages to later indoor or household cats like those living selection processes in agricultural landscapes outside households in Middle Kingdom Egypt, these isotopic data on cats in wider and villages. agricultural settings allow systematic consideration of the signifi- Krajcarz et al.’s (4) research reveals that the Late Neolithic Near cance of outdoor cats to humans through time. Eastern cat diets differed considerably from those of people and Extending this approach to cats in earlier human landscapes will dogs living in this ecosystem. They also differed from Later Roman be a priority for future research. Recent studies show that the cats. With isotopic signatures similar to those of people and dogs, ecological effects of long-term hunter-gatherer occupation attrac- the authors suggest that Roman cats were probably fed by their ted house mice to Natufian hunter-gatherer dwellings in the Levant owners. by 15,000 ya (8, 9), revealing that rodent food sources for cats in human settlements preceded grain storage. These mouse pop- Krajcarz et al.’s research emphasizes the ulations varied with climate and human mobility (9), drawing at- potential of integrated isotopic, morphological, tention to fluctuating opportunities for cats in human settlements through time and the possibility of nondirectional selection. and genetic approaches for examining the So far, cats have not been found in forager settlements or process of cat domestication. landscapes, but the appearance of Near Eastern cats with early farming communities in Cyprus by 10,000 ya supports a long Interestingly, the diets of the Late Neolithic Near Eastern cats time frame for interactions with domestic, feral, or wild cats in studied also differed from those of Pre-Neolithic–Early Neo- farming settlements in the eastern Mediterranean (3, 8). Krajcarz lithic European wild cats that lived far from village communi- et al.’s (4) findings also raise the possibility that the appearance ties.