Earliest economic exploitation of chicken outside East Asia: Evidence from the Hellenistic Southern Levant Lee Perry-Gal 1 , Adi Erlich, Ayelet Gilboa, and Guy Bar-Oz Zinman Institute of Archaeology, University of Haifa, Mount Carmel, Haifa 3498838, Israel Edited by Melinda A. Zeder, National Museum of Natural History, Santa Fe, NM, and approved June 5, 2015 (received for review March 4, 2015) Chicken (Gallus gallus domesticus) is today one of the most wide- spread domesticated species and is a main source of protein in the human diet. However, for thousands of years exploitation of chickens was confined to symbolic and social domains such as cockfighting. The question of when and where chickens were first used for economic purposes remains unresolved. The results of our faunal analysis demonstrate that the Hellenistic (fourth–second centuries B.C.E.) site of Maresha, Israel, is the earliest site known today where economic exploitation of chickens was widely prac- ticed. We base our claim on the exceptionally high frequency of chicken bones at that site, the majority of which belong to adult individuals, and on the observed 2:1 ratio of female to male bones. These results are supported further by an extensive survey of faunal remains from 234 sites in the Southern Levant, spanning more than three millennia, which shows a sharp increase in the frequency of chicken during the Hellenistic period. We further ar- gue that the earliest secure evidence for economic exploitation of chickens in Europe dates to the first century B.C.E. and therefore is predated by the finds in the Southern Levant by at least a century. We suggest that the gradual acclimatization of chickens in the Southern Levant and its gradual integration into the local econ- omy, the latter fully accomplished in the Hellenistic period, was a crucial step in the adoption of this species in European husbandry some 100 y later. chicken | Gallus gallus | zooarchaeology | Hellenistic | Levant I n the modern world, the chicken (Gallus gallus domesticus) is one of the most widespread livestock species and is a major source of animal protein in the human diet. The ancestor of the domestic chicken is the red jungle fowl (Gallus gallus), origina- ting in Southeast Asia, with possible genetic contributions from closely related species through hybridization (1–5). Intensive hybridization between the modern chicken and its wild ancestor caused a loss of the wild progenitor genes (6, 7). Consequently, recent studies usually have focused either on the genetics of the chicken progenitor (8–12) or on zooarchaeological evidence for the domestication of chickens (13–15). The dispersal trajectory of chickens to West Asia, to the Mediterranean, and to Europe following its initial domestication in Southeast Asia remains largely unknown. Moreover, there are only very partial data, and thus there is great uncertainty regarding the place and time of the earliest economic exploitation of chickens: When and where did chickens move from being an exotic species, used only sporadically for symbolic and ritual purposes, to an important livestock species in the Mediterranean and European economies (16, 17)? Our study of chicken remains from the Southern Levant (Israel, the Palestinian Authority, and Jordan) and particularly from the Hellenistic site of Maresha in Southern Israel sheds new light on these issues. We define three main phases in the cultural history of chicken use, based on archaeological, historical, and iconographic evi- dence (Fig. 1). The early phase (Fig. 1, phase A) may have already begun around the sixth millennium B.C.E. when the chicken was initially domesticated during several independent domestication events in Southeast Asia and China (1, 2, 4, 11, 12). On the Indian subcontinent, which also constitutes a part of the natural dispersal range of the jungle fowl, chicken remains were recorded at a few second millennium B.C.E. sites, and it is commonly assumed that domestication occurred there independently (1, 14, 15, 18, 19). The second phase took place in the third–second millennia B.C.E. and includes the dispersal of the chicken out of its natural distribution range to West Asia (Fig. 1, phase B). The earliest chicken remains in the Near East were retrieved in Iran, Anatolia, and Syria and dated to the third millennium B.C.E. or slightly earlier (20). In Egypt, the oldest known chicken remains are possibly even earlier (16). At this early phase, chicken re- mains in archaeological sites are very sparse and often are not associated with domestic contexts. Historical and iconographic records demonstrate an acquaintance with the chicken from the mid-second millennium B.C.E. in Egypt, Mesopotamia, and the Levant (21). All these sources relate to chickens (almost ex- clusively cocks) as an exotic bird, used inter alia for cockfighting and displayed as exotica in royal zoos. The third phase includes its introduction to Europe (Fig. 1, phase C1) and the intensi- fication of its use mainly on this continent (Fig. 1, phase C2). Archaeologically, chicken remains are first observed in Europe only in late ninth and eighth century B.C.E. contexts. The in- troduction of chickens to this region usually is attributed to the Phoenicians who brought chickens from their homeland to their colonies in the West (17, 22). This hypothesis is based on the fact that the earliest chicken remains in Europe were retrieved from Phoenician sites, mostly (although not only) in Iberia (23–25). The oldest reliable dated remains of chickens from central Europe (in the Czech Republic) are from the eighth century B.C.E. (26). The continued presence of chickens has been confirmed in Iberia (27, 28), as well as in southern France and Greece (24, 29), during the second half of the first millennium B.C.E. (Fig. 1, phase C1). However, a survey of the zooarchaeological literature of Europe demonstrates that before the first century B.C.E. the proportion Significance This study offers new evidence on the cultural history of the chicken, a species that until recently received limited attention compared with other domesticated animals. We provide evi- dence for the earliest known economic exploitation of the chicken outside its original distribution. This intensified use is first documented in the Southern Levant during the Hellenistic period (fourth–second centuries B.C.E.), at least 100 y before chickens spread widely across Europe. We explore the mecha- nisms for the spread of chickens as an important species in livestock economies from Asian to Mediterranean and European economies in antiquity to become one of the most widespread and dominant domesticates in the world today. Author contributions: L.P.-G., A.E., and G.B.-O. designed research; L.P.-G., A.E., A.G., and G.B.-O. performed research; A.E., A.G., and G.B.-O. contributed new reagents/analytic tools; L.P.-G. and G.B.-O. analyzed data; and L.P.-G. and G.B.-O. wrote the paper. The authors declare no conflict of interest. This article is a PNAS Direct Submission. 1 To whom correspondence should be addressed. Email: leeper1980@gmail.com. This article contains supporting information online at www.pnas.org/lookup/suppl/doi:10. 1073/pnas.1504236112/-/DCSupplemental. www.pnas.org/cgi/doi/10.1073/pnas.1504236112 PNAS Early Edition | 1 of 6 ANTHROPOLOGY