Early History of Animal Domestication in Southwest Asia
Summary and Keywords
The domestication of livestock animals has long been recognized as one of the most important and influential events in human prehistory and has been the subject of scholarly inquiry for centuries. Modern understandings of this important transition place it within the context of the origins of food production in the so-called Neolithic Revolution, where it is particularly well documented in southwest Asia. Here, a combination of archaeofaunal, isotopic, and DNA evidence suggests that sheep, goat, cattle, and pigs were first domesticated over a period of several millennia within sedentary communities practicing intensive cultivation beginning at the Pleistocene–Holocene transition. Resulting from more than a century of data collection, our understanding of the chronological and geographic features of the transition from hunting to herding indicate that the 9th millennium bce and the region of the northern Levant played crucial roles in livestock domestication. However, many questions remain concerning the nature of the earliest predomestic animal management strategies, the role of multiple regional traditions of animal management in the emergence of livestock, and the motivations behind the slow spread of integrated livestock husbandry systems, including all four domestic livestock species that become widespread throughout southwest Asia only at the end of the Neolithic period.
The origin of domestic animals is one of the crucial events of human prehistory. Livestock have long been seen as a central part of the transformative events of the Neolithic Revolution, that shift from hunting and gathering toward economies based on food production, taking place at the end of the Pleistocene. The impact of livestock, however, was not limited to providing a reliable source of protein, fat, and calories. Instead, the management and domestication of livestock touched every aspect of human culture, transforming agricultural production, views of the natural world, and political relationships as well as the demographics and genetics of both the livestock species and the humans who tended them and utilized their primary and secondary products (Evershed et al., 2008; Ingold, 1994; Tishkoff et al., 2007).
A growing body of archaeofaunal, isotopic, and paleogenomic evidence supports the conclusion that the Fertile Crescent region of southwest Asia was an early hearth for this revolutionary transition and provides an increasingly detailed picture of the geography, timing, and processes involved in animal domestication. This article describes the evidence for the early history of animal management and domestication of livestock species including sheep, goats, cattle, and pigs within the early sedentary communities of the late Epipaleolithic and Neolithic periods in this region.
History of Research
Scholars have long recognized the importance of the origins of domestic livestock in the course of human history. In Morgan’s (1877) hierarchical view of human social evolution, domestic animals were used as the defining feature of the evolutionary category of “barbarism,” responsible for bringing society out of a stage of “savagery,” while Shaler (1895) argued that animal domestication was the innovation responsible for the eventual rise of complex economies and a “civilized morality.” In these early perspectives, herding was thought to have evolved prior to the advent of farming economies, resulting in preagricultural pastoral nomads (Morgan, 1877; Roth, 1887; Shaler, 1895; Turgot, 1895). This notion of the preagricultural origins of animal domestication continued well into the 20th century, as is evident in influential works such as Toynbee’s (1934) “A Study of History.”
Detailed work on the origins of animal domestication required the collection and analysis of faunal remains from archaeological sites. Although natural historians in the 18th and early 19th centuries occasionally described skeletal remains of animals from archaeological sites (e.g., Buckland, 1822; de Perthes, 1860; Rolleston, 1876), it was not until a professional discipline of archaeology developed in the 19th century that scholars regularly collected faunal remains and deposited them in museums for analysis by zoologists and anatomists. This early generation of archaeozoology sought to identify the wild progenitors of domestic species and also made important progress in defining the morphological variation present both within and between domestic and wild taxa (e.g., Brandt, 1855; Darwin, 1868; de Mortillet, 1879; Geoffroy Saint-Hilaire, 1859; Rolleston, 1876).
Of particular importance was zoologist Ludwig Rütimeyer’s (1861) analysis of the animal remains from Neolithic lake dwellings in Switzerland. Rütimeyer’s careful work was among the first to describe skeletal differences between wild and domestic animals, and his ideas concerning the origins of early domestic cattle and pigs heavily influenced Darwin (1868) and scholars of the natural history of domestic animals for decades (Bäumler, 1921; Bökönyi, 1969; Duerst, 1908).
In the early 20th century, Pumpelly’s (1908) pioneering work in central Asia, including excavation at the site of Anau (Turkmenistan), addressed both the natural and cultural histories of this region. Pumpelly’s excavations collected faunal remains that were analyzed by zoologist J. Duerst (1908). Using skeletal features identified by Rütimeyer and others, Duerst identified the transition from hunting to herding in the stratigraphic sequence at Anau. In contrast to most previous scholarship, which argued for a preagricultural origin for animal domestication, Duerst and Pumpelly argued that livestock were locally domesticated within sedentary agricultural settlements like Anau and that climate change, particularly increased aridity, was the primary driver for this innovation. Domestication, wrote Pumpelly (1908), “may have been rendered relatively easy by the changing climatic conditions, which forced the remnants of once great herds of wild animals into close proximity to the people of the oasis” (p. 41).
This model was later incorporated into Childe’s (1934, 1936) influential concept of the “Neolithic Revolution,” that transformative series of events that led to the formation of sedentary societies practicing agriculture and livestock husbandry. Childe also viewed desiccation as the main driver of animal domestication, but he shifted the geographic focus of this process from central Asia to the semiarid Near East. Here he defined an “oasis hypothesis” in which desiccation drove animals to permanent water sources where they came in close contact with early farmers who began to control herds, culling the least docile, which they were able to support with agricultural byproducts and by grazing them in harvested fields. Early farmers quickly learned the benefits of access to dung and milk in addition to meat and skins, thus creating a productive Neolithic mixed agropastoral economy. In academic circles, this firmly set the origins of domestic animals within Neolithic agricultural settlements (e.g., Dyson, 1953), and the search for their origins began in earnest in southwest Asia.
In their popular early 20th century summary of European prehistory, Peake and Fleure (1927) reviewed biological evidence for the origins of domestic livestock, particularly knowledge of the geographic distributions of wild progenitor taxa. Their prescient conclusion that animal domestication took place “in the upper valley of the Euphrates or just where that river leaves the mountains and passes through the foothills” (p. 140) fit with Childe’s notion of southwest Asia as the hearth of both agriculture and early civilizations, a hypothesis that would be field tested by Robert Braidwood in his famous Iraq-Jarmo Prehistoric Project.
Working with a multidisciplinary team including faunal specialist Charles Reed (and also Charlotte Otten and Frederik Barth), Braidwood initiated the Iraqi-Jarmo Prehistoric Project in 1947. He collected floral and faunal remains from Paleolithic and Neolithic sites in the “natural habitat zone” in the piedmont regions of Iraq, where domestication was thought to have originated (Braidwood & Howe, 1960). During this project and the following “Iranian Prehistoric Project” in the Zagros, faunal data were systematically collected from multiple prehistoric sites, providing a rich corpus of data for identifying the transition from hunting to herding (Braidwood, 1983).
Early in the second half of the 20th century, faunal specialists were regularly incorporated into archaeological field projects, and testing hypotheses concerning animal domestication became a regular part of field work in southwest Asia. This resulted in an explosion of archaeozoological work at Neolithic sites across this part of the continent, including Jericho (Zeuner, 1955), Beidha (Perkins, 1966), and El Khiam (Ducos, 1968) in the southern Levant; Ali Kosh (Hole et al., 1969), Shanidar Cave and Zawi Chemi Shanidar (Perkins, 1964), and Ganj Dareh (Hesse, 1978) in western Iran; Belt Cave in northern Iran (Coon, 1951); Suberde in central Anatolia (Perkins & Daly, 1968); and Çayönü in southeastern Turkey (Braidwood et al., 1974) (Figure 1).
Each of these projects produced detailed faunal work focused on evidence for animal domestication within early Neolithic settlements. This work spawned a body of scholarship that represents the foundation of current approaches to animal domestication in southwest Asia and resulted in innovations in the methodologies used to identify early animal management (e.g., Bökönyi, 1969; Ducos, 1968; Reed, 1960; Zeuner, 1963).
Methods for Identifying Animal Domestication
In response to the field testing of hypotheses for animal domestication in the mid-20th century, faunal specialists developed a set of methodologies for identifying domestic animals (Bökönyi, 1969; Ducos, 1969; Meadow, 1989; Uerpmann, 1973; Zeder, 2006). This included the development of conceptual frameworks for addressing the domestication process. The domestication process was conceptualized as a long-term evolutionary process in which animals adapted to living and breeding under conditions of human management (Hammer, 1984; Meadow, 1989; Price, 1984). The diachronic nature of this process resulted in two somewhat divergent uses of the term domestication: one referring to “biological domestication” as represented by the morphological changes resulting from generations of selection pressures under human management, the other referring to “cultural domestication” emphasizing the application of husbandry practices to animals even prior to the appearance of biological transformations (Bökönyi, 1969; Ducos, 1978; Dyson, 1953). These two uses of the term “domestic” have been a source of some confusion but can be clarified by the use of the term to describe animals exhibiting biological changes from their wild ancestors linked to the domestication process, while “predomestic management” can be used to describe the application of husbandry strategies on animals not yet exhibiting biological changes (previously referred to as “protoélevage,” “protodomestication,” or “cultural control” (Buitenhuis, 1997; Ducos, 1978; Hecker, 1975)).
The primary lines of evidence used to identify the domestication process, domestic animals, and animal management emerged by the mid-20th century. These include:
2. Artistic representations showing domestic animals and their use
3. Archaeological context (e.g., animal burials)
4. An increase over time in “prodomestic” taxa
5. Morphological changes associated with domestication (the “domestication syndrome”)
6. Demographic evidence
Biogeography has been an important line of evidence since the earliest generation of scholars attempted to address the origins of domestic livestock, and was a central variable in Braidwood’s choice of the “Hilly Flanks” region for the Iraq-Jarmo Prehistoric project. Using the distribution of wild progenitor taxa, early scholars hypothesized the location of the geographic origins of early domestic livestock. In a remarkably prescient statement, Brandt (1855) identified the Turkish Taurus as the potential homeland of goat domestication based on the presence of bezoar in that region and its proximity to early centers of civilization:
“Or, comme le Taurus est situe precisement dans le voisinage du siege primordial de la plus antique civilisation du monde, la Capra aegagrus a pu devenir, a l’epoque la plus reculee, l’objet de la domestication.” [Now, since the Taurus is located precisely in the vicinity of the primordial seat of the most ancient civilization of the world, Capra aegagrus could become, at the earliest time, the object of domestication.] (p. 578)
Iconography representing animal management and domestic animals provides some clues to human–animal relationships but is limited in prehistoric periods (van Buren, 1939). In addition, contextual evidence for close human–animal relationships, particularly the presence of animal skeletons in burials, may reflect human control over animals (Tchernov & Valla, 1997; Vigne et al., 2004).
Dyson (1953) suggested that a shift in reliance from wild taxa to “prodomestic” taxa, or those species known to have been eventually domesticated, is suggestive, especially in concert with other lines of evidence, of the process of domestication. This is particularly well documented at the Neolithic levels of Jericho (Tell es-Sultan), West Bank, where prodomestic goats and (nonlocal) sheep increase over time at the expense of wild gazelle and equids (Clutton-Brock, 1979).
Of these lines of evidence, the final two play central roles in most recent arguments for early animal domestication. It has long been observed (e.g., Darwin, 1868) that domestic animals exhibit a suite of morphological changes compared to their wild ancestors, known as the “domestication syndrome.” These changes include a reduction in body and brain size, changes in horn shape and limb proportions, increased variability in coat color, etc., as a result of many generations of both artificial and natural selection under regimes of human management (Arbuckle, 2005; Driscoll et al., 2009; Hammer, 1984). Rütimeyer (1861) was perhaps the first to use these changes in morphology to identify early domestic animals in the archaeofaunal record, quantifying differences in body size based on dental measurements to distinguish between the remains of large-sized wild boar and smaller domestic pigs. Decreased body size was similarly used by Winge (1900) to identify early domestic pigs in the Baltic region and was applied by Duerst (1908) to argue for the domestication status of ruminants and pigs at Anau (see also Bäumler, 1921).
In addition, Duerst (1908, pp. 370–373) used changes in the shape of horncores to identify domestic cattle, sheep, and goats. The appearance of increased variability in horncore shape, including twisting, decrease in size, or even complete loss of horns, has been used to identify domesticated animals based on both iconography and faunal remains (Bökönyi, 1977; Clutton-Brock, 1979; Hole et al., 1969; Wasse, 2002; Zeuner, 1955).
Following the work of Uerpmann (1979), postcranial articular breadth and depth measurements of prodomestic ruminants have been widely used to identify a decline in body size evident in the early Neolithic and thought to reflect human control over animal populations and the emergence of domestic livestock (Arbuckle et al., 2014; Davis, 1987; Peters et al., 2005). However, there are problems with using morphological changes, and particularly a decrease in body size, as a marker for early human management of animals. Identifying changes in body size requires robust local and regional datasets for diachronic comparison, which are often lacking. Furthermore, recent wild populations are often used as a baseline to represent the body size of ancient wild taxa without consideration of the impact of late Holocene climate- and human-induced environmental changes. This is problematic because mammalian body size responds to a host of variables including changes in diet, temperature, and predation pressure and, more generally, Bergman’s Law (Davis, 1982). In addition, many mammals are characterized by significant size differences between males and females. As a result of this dimorphism, changes in the representation of the sexes in an archaeofaunal assemblage can lead to the false impression of changes in mean or median size (Zeder, 2001).
Both Reed (1960) and Dyson (1953) recognized that evidence for morphological changes in managed animals must necessarily follow the instigation of herd management by some amount of time—perhaps several millennia. In Dyson’s view, this meant that cultural control preceded biological domestication, resulting in a significant time gap between the appearance of biological evidence for domestication and the initiation of animal husbandry. Moreover, it was recognized early on that managed animals often interbreed with wild populations, potentially dampening the effects of the “domestication syndrome” (Darwin, 1868). In order to address these issues, scholars have turned to demographic evidence for herd management and culling strategies.
Demographic evidence reflects the age and sex composition of the animals culled by humans. Because demographic profiles reflect the divergent goals of hunters and herders (i.e., accessing dead animals versus ensuring herd reproduction), they have been widely applied to the origins of animal domestication (Jarman & Wilkinson, 1972; Meadow, 1989; Zeder, 2012). When targeting large game, hunters often select large-bodied adult animals, whereas herding (especially of sheep, goats, and pigs) often targets surplus, juvenile males for slaughter (Payne, 1973; Stiner, 1990). Although a high frequency of juvenile animals has long been used as evidence for herd management (Bökönyi, 1969; Coon, 1951; Ducos, 1968; Perkins, 1964), diverse hunting strategies (e.g., stalking, traps, cooperative drives), prey ethology, and season of exploitation can lead to significant variation in the demographic profiles and representation of juveniles produced by hunting practices (Dyson, 1953; Wilkinson, 1976; Wright & Miller, 1976). However, the typical herder practice of targeting young males for slaughter is rarely reproduced by other exploitation methods. Because of this, Zeder has argued that sex-specific demographic profiles showing the culling of juvenile males provides a “leading edge marker” for herd management even before morphological changes are evident (Zeder & Hesse, 2000). The efficacy of this method is limited, however, in cases where multiple exploitation methods, including both hunting and husbandry, are practiced or in management regimes that are conservative in the slaughter of juveniles (Halstead, 1998; Payne, 1973).
In addition, some of the methods developed to identify the origins of domestic animals have not withstood the test of time. For example, based on collaborative work with Dexter Perkins on the early Neolithic faunas from Iran, Iraq, and Turkey, Patricia Daly and Isabella Drew (Drew et al., 1971) explored differences in the microstructure of the skeletons of wild and domestic animals. They argued that domestic status could be identified through characteristic birefringence evident in thin sections of articular ends of long bones viewed under polarized light. The pattern of birefringence was thought to reflect a reorientation of hydroxyapatite crystallites in the long bones in response to reduced activity levels associated with animal husbandry. This birefringent “blue rim” was briefly presented as a promising method for identifying domestic animals (Daly et al., 1973; Hecker, 1975; Pollard & Drew, 1975). However, critiques have pointed out a series of problems with this method, arguing that the presence or absence of a “blue rim” visible under polarized light represents differential preservation of collagen and diagenetic minerals in archaeological materials rather than domestication status (Gilbert, 1989; McConnell et al., 1971; Watson, 1975).
More recently, methodological innovations have added dimensions to understanding the origins of animal husbandry. Most important among these have been biomolecular approaches, including analysis of isotopes from dental and skeletal tissues as well as ancient DNA studies. Isotopic work has been used to identify human manipulation of animal diets associated with foddering and pasturing animals and also the early exploitation of domestic ruminants for dairy products (Evershed et al., 2008; Makarewicz & Tuross, 2012; Pearson et al., 2007).
Studies of modern and ancient livestock DNA have provided a welcome new line of evidence for the origins and movement of domestic taxa. Early generations of paleogenetic work primarily focused on recovering and identifying mitochondrial DNA haplogroups in modern and ancient populations and exploring evidence of geographic structure, population movements, and number of domestication events (e.g., Bruford et al., 2003). With more recent innovations in sequencing methods (e.g., next-generation sequencing, high-throughput sequencing (Gamba et al., 2016; Knapp & Hofreiter, 2010)), an explosion of work including nuclear and whole genome sequencing is addressing not only the spread of domestic maternal and paternal lineages but also the appearance of changes in genotype and phenotype (e.g., coat color) associated with the domestication process (Krause-Kyora et al., 2013; Librado et al., 2017; Ludwig et al., 2009; Ottoni et al., 2013).
Early Domestication of Livestock in Neolithic Southwest Asia
Four important livestock taxa were domesticated in southwest Asia in the early Neolithic: sheep, goat, cattle, and pigs. Although the origins of the management of these taxa were broadly contemporaneous, they each follow distinctive and independent trajectories. The archaeological evidence for these trajectories is therefore described separately for each taxon.
Sheep (Ovis aries)
Although it was initially difficult for early scholars to identify the wild ancestor of the domestic sheep (Ovis aries) (Hilzheimer, 1936), based on genetic evidence it is clear in the 21st century that this animal derives from the mouflon of southwest Asia (Ovis orientalis) (Chessa et al., 2009), whose habitat includes the mountains and adjacent hills and plains of the Fertile Crescent region extending from Lebanon and Syria into Turkey and eastward into the Zagros and Caucasus (Uerpmann, 1987).
In the late Pleistocene, wild sheep were intensively hunted in a broad arc from the western Taurus mountains to the northern Zagros, including the Anatolian plateau where, in combination with wild goats, they frequently represent 50%–90% of archaeofaunal assemblages. In the northern Zagros region, Perkins (1964) argued for early sheep domestication at both Shanidar Cave and the nearby open-air site of Zawi Chemi Shanidar dating to the early 10th millennium bce based on the abundance of Ovis remains, a high frequency of juvenile animals, and the locations of the sites outside of “optimal” mouflon habitat. In a recent reanalysis, Zeder (2008, 2011) confirmed that juvenile male sheep were targeted at Zawi Chemi but interprets this as a hunting strategy targeting juvenile males rather than the earliest evidence for husbandry. Wild sheep are also the dominant prey taxon in the upper Tigris basin in southeastern Turkey at the 11th and 10th millennia bce sites Hallan Çemi, Körtik Tepe, and Hasankeyf (Arbuckle & Özkaya, 2007; Miyake et al., 2012; Starkovich & Stiner, 2009; Zeder, 2012). These distinctive “round house horizon” sites exhibit patterns similar to that at Zawi Chemi in which juvenile animals (particularly larger males at Hallan Çemi and Körtik) are targeted but retain the large body size and horn morphologies typical of wild mouflon.
Although detailed interpretations of sheep exploitation practices at these early round-architecture village sites are currently lacking, it is likely that the region extending from the upper Tigris into northwestern Iran, where mouflon were heavily exploited by early populations of sedentary cultivator–hunters, saw the first movement toward intensive interaction with, and management of, wild sheep populations. However, sheep maintained a wild phenotype in this period and predomestic management, if practiced, is unclear. Also unclear is the impact of these late Epipaleoithic sheep economies on the development of sheep husbandry. The upper Tigris region, south of Diyarbakır, was abandoned in the 9th millennium bce, and the earliest evidence for the management of predomestic and early domestic sheep appears in villages of the Pre-Pottery Neolithic B (PPNB) tradition in the Turkish Euphrates valley and central Anatolia.
At the small settlement of Nevalı Çori, located within the valley of the Turkish Euphrates, sheep are relatively rare (approximately 5%, based on number of identified specimens (NISP)) in an animal economy focused on gazelle hunting, but a combination of juvenile culling, decreased body size, and isotopic evidence for foddering suggests that sheep husbandry was well under way in this farmer–hunter–herder community by the mid-9th millennium bce (Grupe & Peters, 2009; Losch et al., 2006; Peters et al., 2005). However, there is no evidence for the preferential culling of young males at Nevalı Çori, indicating that this common pastoral management practice was not utilized.
The site of Çayönü Tepesi, though located in the Tigris drainage, is geographically and culturally affiliated with Euphrates basin sites such as Nevalı Çori (it is located 130 km to the east) and exhibits parallel evidence for early sheep management. In the late 10th and early 9th millennia bce occupational levels at Çayönü, wild sheep, especially large rams, were occasionally hunted (Hongo et al., 2005). But in the later 9th millennium, the appearance of smaller sized sheep, combined with a bias in favor of adult females and a high frequency of juveniles, suggests that intensive but small-scale management was initiated at Çayönü.
Evidence for early sheep management extends farther north into the uplands regions of Anatolia as well, although morphologically domestic sheep are absent from this region in the early Neolithic. At the village of Cafer Höyük, located north of Nevalı Çori in the Euphrates valley, Helmer (2008) has argued that morphologically wild sheep were both hunted and herded in the 9th millennium bce. The management of sheep in the absence of changes in phenotype is also represented on the central Anatolian plateau at the site of Aşıklı Höyük. Here, sheep were the primary prey taxon, accounting for more than half of the large mammal remains in level 4 (late 9th millennium bce) and increasing in abundance through the mid-8th millennium (level 2) (Buitenhuis, 1997; Stiner et al., 2014). Although showing no signs of morphological changes or the practice of young male kill-off (similar to Cafer), sheep at Aşıklı were overwhelmingly culled as juveniles and the assemblage includes large numbers of perinatal remains, suggesting pregnant ewes were frequently present onsite (Buitenhuis, 1997; Stiner et al., 2014). Moreover, the presence of animal dung, identified as stabling deposits, indicates that herbivores (presumably sheep) were penned onsite (Stiner et al., 2014). Control over herds of morphologically wild sheep is further supported by isotopic evidence suggesting low levels of variability in the caprine (sheep/goat) diet indicative of local grazing and foddering (Pearson et al., 2007).
At the site of Shillourokambos, on the island of Cyprus, sheep were imported in small numbers by the end of the 9th millennium bce (Vigne et al., 2011). Although initially exhibiting little evidence for changes in body size or horncore shape associated with domestication, culling strategies indicate that they were intensively herded at this time (Vigne, 2011). This chronology places sheep management on Cyprus only slightly later than its emergence in the Turkish Euphrates valley and Cappadocia, but prior to its emergence on the northern Mesopotamian plains or southern Levant (Vigne et al., 2011).
In the southern Levant, robust traditions of mouflon exploitation are absent in the late Pleistocene and early Holocene (Horwitz & Ducos, 1998; but see also Yeomans et al., 2017). For most of the region, sheep appear in the archaeofaunal record in the mid-8th millennium bce, representing the introduction of domestic sheep and sheep husbandry, presumably from the north (Horwitz et al., 1999; Makarewicz, 2014; Martin & Edwards, 2013; Wasse, 2002). In support of this interpretation, it is also at this time (ca. 7500 bce) that morphologically domestic sheep appear in large numbers along the lower Turkish Euphrates at sites including Gritille, Mezra-Teleilat, and Hayaz Höyük and also Gürcütepe (Buitenhuis, 1985; Ilgezdi, 2008; Monahan, 2000; von den Driesch & Peters, 2001). Moreover, domestic sheep also appear abruptly in the occupational sequence at Tell Halula (OP8) and also at Abu Hureyra on the Syrian Euphrates, marking a rapid southern expansion of domestic sheep and ovine husbandry in the mid-8th millennium bce (Middle PPNB) (Legge & Rowley-Conwy, 2000; Sana & Tornero, 2013).
Despite an early focus on sheep exploitation, mouflon hunting declines dramatically in the Zagros in the 9th millennium bce (Zeder, 2008). Domestic sheep are abundant at Aceramic Jarmo in northern Iraq (ca. 7000 bce) (Stampfli, 1983) but are absent further south and east, where small numbers of wild mouflon, primarily adult rams, were hunted at Ganj Dareh and Ali Kosh from the 8th through the beginning of the 7th millennium bce (Hesse, 1978; Hole et al., 1969). Intensive sheep herding was adopted in the Zagros region only in the 7th millennium bce, when young male kill-off and morphological domesticates are evident at sites including Tepe Sarab and Choga Sefid (Zeder, 2008).
Goats (Capra hircus)
Domestic goats (Capra hircus) derive from the Near Eastern bezoar (Capra aegagrus) (Naderi et al., 2008), which inhabited uplands and rocky areas across southwest Asia from the Taurus mountains to the Zagros and south into portions of the southern Levant (Uerpmann, 1987). Unlike the situation with sheep, prehistoric goat exploitation has a long history in the western, eastern, and central portions of the Fertile Crescent, and early predomestic herd management emerged in each of these regions in the 9th millennium bce.
Early evidence for goat management derives from the central portions of the Fertile Crescent in southeastern Turkey. In the Epipaleolithic, hunter–gatherers exploited bezoar in the Taurus mountains at Direkli, Karain, and Öküzini caves but were a minor part of the animal economies further east in the upper Tigris drainage (Arbuckle & Özkaya, 2007; Atici, 2011; Hongo et al., 2005; Starkovich & Stiner, 2009). Goats were heavily utilized high in the Euphrates valley at Cafer Höyük and were exploited in smaller numbers at both Nevalı Çori and Çayönü in the 9th millennium. At Cafer, morphologically wild goats were the dominant prey taxon and large adult males were often targeted. Although this pattern is usually associated with hunting, a high frequency of juvenile remains suggests that goats were under human control, and Helmer (2008) has suggested both herding and hunting were practiced at Cafer.
Further downstream at Nevalı Çori, goats represent a tertiary resource following gazelle and suids. Although the earliest evidence of morphological changes in sheep is found at Nevalı Çori, comparable changes are not visible in goats (von den Driesch & Peters, 2001). However, as at Cafer, relatively high frequencies of juvenile animals suggest selective herd management (Peters et al., 2005). Moreover, Grupe (Grupe & Peters, 2009) has argued that isotopic evidence reflects human influence on goat diets associated with human control.
Additional evidence for goat management in southeastern Turkey emerges at Çayönü. Here, bezoar were a small portion of the animal economy and large adult bucks were selectively hunted in the 10th millennium bce. In the late 9th millennium (Cobble phase), goat exploitation increased, shifting toward the consistent culling of young males and the appearance of mild phenotypic changes (decrease in body size) indicating that the intensive management of small herds of goats was initiated by this time (Hongo et al., 2004).
Morphologically domestic goats subject to young male culling appear further south along the Turkish Euphrates at Teleilat, Gritille, and Hayaz Höyük, and at Gürcütepe in the mid-8th millennium bce (Buitenhuis, 1985; Ilgezdi, 2008; Monahan, 2000; von den Driesch & Peters, 1999). Goats appear well outside of their natural habitat in the earliest levels of Tell Halula, located in the Syrian Euphrates valley, dating to the early 8th millennium bce, where they surprisingly represent a central feature of the animal economy (Sana & Tornero, 2008). Here, the culling of young males combined with some evidence for decrease in body size (Helmer, 2008, Figure 17) reflects the rapid emergence of goat herding. Domestic goats appear at Abu Hureyra in the mid-8th millennium bce and at Bouqras at the Euphrates–Khabur confluence a few centuries later, further documenting the continued spread of goat husbandry southward through the Euphrates valley (Clason, 1983; Legge & Rowley-Conwy, 2000).
In the Zagros region of Iran, goats were the primary prey taxon throughout the Pleistocene and early Holocene (Perkins, 1964). At Asiab, in the Kermanshah valley, large bezoar bucks were targeted by hunters in the 10th millennium bce (Bökönyi, 1977). Hesse and Zeder (2001) have described the earliest evidence for intensive goat management at Ganj Dareh in the central Zagros dating to just after ca. 8000 bce. At Ganj Dareh, morphologically wild goats make up the vast majority of the faunal assemblage and juvenile males were subject to intensive culling, providing strong evidence for human control over herds. Although this predomestic herding of goats postdates early goat management in southeastern Turkey by several centuries, preliminary evidence from the site of Kelek Asad Morad, Luristan, hints that intensive young male culling (and therefore herd management) was practiced in the mid-9th millennium bce in the Zagros (Moradi et al., 2016). Moreover, the identification of dung in late 9th millennium bce deposits at the site of Jani is interpreted as reflecting site management and stabling of morphologically wild goats (Matthews et al., 2013). The first goats to exhibit morphological changes associated with domestication process in western Iran appear in the 8th millennium at Ali Kosh, on the Deh Luran plain, and at Tepe Sarab in the highlands. Here, a decrease in body size and changes in the shape of male horncores are associated with intensive herd management as well as isolation of managed herds from wild populations (Hole et al., 1969; Zeder, 2008).
Surprisingly early evidence for goat management also comes from the island of Cyprus. Vigne has carefully documented the introduction of morphologically wild goats to the island as early as the mid-9th millennium bce (several centuries before sheep are evident). Despite the obvious intentionality of their introduction, there is little evidence that these goats were subject to intensive husbandry until the early 8th millennium bce, when management strategies including the culling of juvenile males is apparent. This situation suggests that mainland populations somewhere in the northern coastal Levant developed close relationships with goats at an early date and is perhaps linked to traditions of goat exploitation in the late Pleistocene documented at Ksar Akil and Nachcharini in Lebanon and at Tell Qaramel in northwestern Syria, where goats were exploited in surprising numbers near Aleppo in the 10th millennium bce (Garrard et al., 2003; Grezak, 2012; Kersten, 1987).
On the northern margin of the southern Levant, Tell Aswad provides early evidence for predomestic goat management in that region contemporary with the introduction of goats to Cyprus (Helmer & Gourichon, 2017). At Aswad, goats were second only to gazelle in importance in the 9th and 8th millennia bce, and the location of the site in the Damascus basin some distance from likely bezoar habitats suggests goats may have been managed at the site since the mid-9th millennium bce. Although no changes in body size are evident through this period, biometrics and age at death suggest that herds were managed through the culling of juvenile males by the 8th millennium bce. Moreover, the appearance of torsion in male horncores in the later 8th millennium bce provides early evidence for the emergence of domestic phenotypes with parallels on Cyprus and in western Iran (Helmer & Gourichon, 2008).
Further south, although gazelle was the predominant prey species, goats were a focus of some communities in the Mediterranean hills and Jordan valley in the 9th and early 8th millennia bce. At Abu Gosh, morphologically wild goats (as well as ibex, Capra nubiana), including large numbers of adult males, were exploited but many animals were also culled as juveniles (Bar-Gal et al., 2002; Horwitz, 2003). This situation is similar to that described at contemporary sites including Cafer, Aşıklı, and Shillourokambos, suggestive of systems of predomestic goat management unique to the 9th millennium bce. Moreover, isotope evidence at Abu Gosh indicates that goats were foddered as early as 8000 bce, suggesting that intensive management likely preceded the appearance of morphological domesticates in this region (Makarewicz & Tuross, 2012). In addition, goats were the dominant species at the farming village of Ain Ghazal starting in the late 9th millennium bce, and a combination of young male culling and morphological changes including increased variation in horncore morphology and a decrease in body size suggest that goats were intensively managed by the early 8th millennium in western Jordan (Wasse, 2002).
In southern Jordan, goats represent 90% of the fauna at the late 9th and early 8th millennia site of Beidha. Young male goats were heavily culled at Beidha suggesting intensive predomestic management, although the presence of ibex (Capra nubiana), the remains of which are difficult to distinguish from goats (Capra aegagrus), complicates interpretations of early evidence for goat management (Hecker, 1975). Remains from the site of Basta and Baja clearly indicate the presence of domestic goats in southern Jordan by the late 8th millennium bce (Becker, 1991).
Pigs (Sus scrofa domesticus)
A combination of archaeological and paleogenetic work supports the interpretation that wild boar (Sus scrofa) were domesticated at an early date in southwest Asia (Larson, 2005). Control over wild boar populations is evident in the 10th millennium bce with the translocation of suids onto the island of Cyprus (Vigne et al., 2009). Here, Vigne argues, terminal Pleistocene hunters stocked the island with boar to provide a game resource. These imported pigs were the focus of the early Cypriote animal economy with evidence for pig management, in the form of morphological changes, juvenile culling, and use in funerary offerings, emerging ca. 8000 bce (Vigne et al., 2011). Although the source population for Cypriote suids is not clear, pigs were hunted at Tell Qaramel in the 10th millennium bce, perhaps indicative of a western Syrian tradition of boar exploitation that has not yet been fully explored (Grezak, 2012).
Pigs were intensively hunted in the upper Tigris basin, and the earliest archaeological evidence for pig management and domestication appears in this region. At the 10th millennium settlement of Hallan Çemi, contemporary with the movement of boar to Cyprus, Redding (Redding & Rosenberg, 1998) argued for early boar (i.e., wild phenotype) management based on an overrepresentation of males, a high frequency of juveniles, and an increase in the relative frequency of suids over time. Although subsequent work at Hallan Çemi has questioned the evidence for human management (Starkovich & Stiner, 2009), recent analysis of age of death of suids (Lemoine, 2012) shows that kill-off focused on subadult animals between 1 and 3 years of age. In addition, a huge number of neonatal suid remains (44% of the suid sample) have been described (Lemoine, 2012, Figure 15). This very specific age profile indicates that the residents of Hallan Çemi had regular and predictable access to pregnant sows/neonates and culled young adult boar—patterns that may reflect a form of predomestic management at the site.
Clearer evidence for suid management and domestication has been described at Çayönü, where suids are the most abundant mammalian taxon in the 10th and 9th millennia bce occupations, suggesting an unusually intensive human–suid relationship (Hongo & Meadow, 1998). At Çayönü, gradual decreases in body size and tooth size and in the age of kill-off as well as isotopic evidence for changes in diet from the 9th and 8th millennia suggest that management and population isolation led to the in situ development of a domestic population (Ervynck et al., 2001; Pearson et al., 2013).
In the Turkish Euphrates basin at Cafer Höyük and Nevalı Çori, suids were initially a small component of the 9th millennium bce economies, but by the mid-8th millennium bce they increase to approximately 34% and 20% respectively of the mammalian faunas at those sites (Helmer, 2008; von den Driesch & Peters, 2001). At Cafer Höyük, juveniles were heavily culled but suids do not exhibit changes in body size, likely indicating a system of predomestic management was practiced at the site (Helmer, 2008). Combined with the appearance of some small-sized individuals with isotope values interpreted as indicating foddering (Losch et al., 2006), suid remains from Nevalı Çori have been interpreted as reflecting intensive management and population isolation. Further reductions in suid body size indicating the emergence of a domestic phenotype are widely apparent in the later 8th millennium bce in southeastern Turkey and in the Syrian Euphrates valley (Peters et al., 2005).
In central Anatolia, boar were frequently hunted at Pinarbaşı A and Boncuklu Höyük on the Konya Plain in the 10th and late 9th millennia bce (Baird, 2012; Carruthers, 2003). However, this tradition of boar hunting declines in the 8th millennium when suid remains become rare at Aşıklı and Çatalhöyük. In fact, there is no evidence for suid management or domestic pigs in central Anatolia for the entirety of the Neolithic (Arbuckle, 2013). However, domestic pigs are evident at sites such as Bademağacı and Ulucak, representing the expansion of Neolithic lifeways into southwestern Anatolia in the early 7th millennium bce (Çakırlar, 2012; De Cupere et al., 2008; see also Atici et al., 2017).
Finally, early pig management has been suggested at the northern margin of the southern Levant at Tell Aswad. A decrease in the size of suids is evident in the early 8th millennium bce, which Helmer (Helmer & Gourichon, 2008) has interpreted as evidence for the management of domestic pigs. In the later 8th millennium, these smaller sized individuals were slaughtered at young ages (6–12 months primarily), suggesting that they represent an intensively managed population.
Further south in the southern Levant, pig management was initiated in the 7th and 6th millennia, the latter part of the Neolithic, as seen at Wadi Shu’eib in Jordan, Atlit Yam, Hagoshrim, and also at Ras Shamra in Lebanon (Haber & Dayan, 2004; Helmer, 1989; Makarewicz, 2016). This slow adoption of pig management is also evident in the Zagros region. Here, although the management of domestic pigs is evident at Jarmo in northern Iraq in the 7th and possibly late 8th millennium bce, domestic pigs were not raised in the intermontane valleys of western Iran until the 6th millennium bce and perhaps even later in the southern Zagros region (Flannery, 1983; Price & Arbuckle, 2015).
Cattle (Bos taurus)
Domestic taurine cattle (Bos taurus) derive from the wild aurochs (Bos primigenius), a large ruminant that inhabited a wide range of forest, grassland, and riverine environments across Eurasia (Bradley et al., 1998; Uerpmann, 1987). Although aurochs were occasionally hunted throughout southwest Asia in the late Pleistocene and early Holocene, they were intensively utilized in the upper Euphrates valley and also on the central Anatolian plateau. It is thought that early management was initiated in the former region, with evidence for cattle management and morphological changes occurring significantly later in surrounding regions.
Aurochs were heavily hunted along the Syrian Euphrates at the site of Mureybet, where morphologically wild cattle make up an unusually high frequency (approximately 30%) of the mammalian fauna in the 9th millennium (layer IV) (Gourichon & Helmer, 2008). Cattle are also abundant at contemporary Jerf el-Ahmar and Djade in the Syrian Euphrates valley, at Tell Qaramel near Aleppo, and at Göbekli Tepe in southeastern Turkey (Grezak, 2012; Helmer et al., 2005). At both Göbekli and Jerf el-Ahmar, hunters preferentially focused their hunts on young adult and female animals rather than large adult males (Peters et al., 2013; Peters et al., 2005).
The earliest evidence for morphological changes associated with intensive human management comes from the site of Djade, where Helmer has argued a decrease in the size of bulls reflects a reduction in sexual dimorphism linked to the process of domestication (Helmer et al., 2005). However, additional evidence for herd management is limited at this time (early 9th millennium bce). Cattle were translocated to the island of Cyprus in the mid-9th millennium, where Vigne argues they represent morphological domesticates subject to the preferential culling of young males (Vigne et al., 2011).
More dramatic evidence for morphological changes associated with intensive management is available from the earliest layers at Halula dating to the early 8th millennium bce, where cattle husbandry was a central feature of the economy, and also at mid-8th millennium deposits at both Teleilat and Gritille on the Turkish Euphrates (Ilgezdi, 2008; Monahan, 2000; Sana & Tornero, 2013). At Cafer, Helmer has argued that a slight decrease in dimorphism (similar to the situation at Djade) and the presence of pathologies reflect cattle herding, although aurochs hunting is thought to have continued as well (Helmer, 2008).
In the Tigris basin, aurochs were hunted in the 10th millennium at Çayönü, Körtik Tepe, and Nemrik 9 in northern Iraq but were very rare at Hallan Çemi and Hasankeyf (Arbuckle & Özkaya, 2007; Hongo et al., 2004; Lasota-Moskalewska, 1994). At Çayönü, changes in stable isotope values from bovine teeth, along with the appearance of gracile individuals in the late 9th millennium bce, have been argued to reflect manipulation of cattle diets, population isolation, and intensive management in the late 9th millennium bce (Hongo et al., 2009).
In central Anatolia, aurochs were heavily exploited in the 9th and 8th millennia bce but there is little evidence that they were under intensive human management. Large-sized aurochs are abundant at Boncuklu in the 9th and early 8th millennia, and in the earliest reported levels of Aşıklı (level 4e) they represent the most important food animal based on meat contribution (Baird, 2012; Stiner et al., 2014). Morphologically wild cattle are also the most abundant taxon at the nearby special purpose site of Musular, where aurochs feasting may have been a central activity (ca. 7500 bce) (Russell et al., 2005). In the early phases at Çatalhöyük (7300–6500 bce), cattle continued to be a major economic and symbolic focus, with the remains of adult bulls frequently incorporated into architecture as commemorative deposits (Russell et al., 2005, 2013b). However, by the mid-7th millennium this focus on large bulls is no longer evident, and a predominance of the remains of females and an increase in the culling of juvenile–subadult cattle suggest major changes in cattle exploitation, perhaps reflecting a shift toward predomestic management. Domestic cattle exhibiting small body size, altered horncore morphologies, and evidence for young male culling appear suddenly at Çatalhöyük, and also at Erbaba Höyük in the adjacent Beyşehir basin, in the second half of the 7th millennium, likely reflecting the importation of domesticates from neighboring regions (Arbuckle, 2013; Arbuckle & Makarewicz, 2009; Russell et al., 2013a; Twiss & Russell, 2009). In addition, residues of dairy fats have been identified on ceramic sherds dating to the second half of the 7th millennium bce, suggesting that domestic cattle were used for secondary as well as primary products at this time (Evershed et al., 2008; Pitter, 2013).
In the southern Levant, scholars have suggested that a decrease in the size of bulls, a kill-off focused on juvenile animals, and the presence of skeletal pathologies indicative of mechanical stress indicate intensive cattle husbandry was practiced at the site of Aswad in the 8th millennium bce (Helmer & Gourichon, 2008). Further south, there is no clear evidence for intensive management of cattle until the 7th millennium, when morphologically domestic cattle appear at sites on the coastal plain and Jordan Valley (Horwitz & Ducos, 2005; Horwitz et al., 1999; Marom & Bar-Oz, 2009). A similarly late appearance of domestic cattle is evident in the Zagros region, where a dramatic decrease in body size has been interpreted as representing the beginnings of cattle husbandry in the 6th millennium bce (Arbuckle et al., 2016).
History of Animal Domestication
The rich archaeofaunal record in prehistoric southwest Asia provides an increasingly detailed view of the complex and long-term history of animal domestication. A preponderance of evidence supports the conclusion that the first steps toward livestock domestication took place within the context of settled communities at the Pleistocene–Holocene transition. Intensive relationships likely developed between humans and ungulate prey taxa at sedentary settlements practicing predomestic plant cultivation in the late Epipaleolithic, at sites such as Hallam Çemi, Körtik Tepe in southeastern Turkey, Mureybet in northern Syria, Chogha Golan and Sheik-e Abad in western Iran, and Wadi Faynan 16 in Jordan (Finlayson et al., 2011; Gourichon & Helmer, 2008; Matthews et al., 2013; Starkovich et al., 2016). These manipulative relationships, however, have been difficult to identify using traditional zooarchaeological methods since they resulted in no morphological changes in animal populations and likely were combined with traditional hunting practices. However, the discovery that wild boar and wild goats were transported to the island of Cyprus in the 10th and early 9th millennia bce both confirms control over wild animal populations at this time and indicates that human manipulation of those populations was taking place on the mainland as well.
Despite the likelihood of increasing and widespread human manipulation of wild animal populations in the 10th millennium bce, clear evidence of herd management is lacking until the 9th millennium when the cultivation of domestic crops becomes widespread, providing strong (if circumstantial) support for the relationship between plant cultivation, sedentism, and the origins of animal management (Zeder, 2012). In this period, the systems of predomestic animal management that did emerge were highly diverse and localized, and developed from previous long-term relationships with local taxa within their natural habitats.
In this period of “initial diversity” in animal management (Arbuckle & Atici, 2013), animals retained a wild phenotype, and likely wild behaviors, as predicted by Dyson (1953). This must have resulted in significant challenges to early herders since animals were not yet adapted to living and breeding around humans, and pastoral practices were likely developed in an extended process of “learning by doing” (Peters et al., 2015). Although the process of developing experimental management regimes was widespread across the Fertile Crescent in the 9th millennium bce, there is no indication that it originated in one “nuclear zone” spreading outward. Instead, it appears that unique initial management systems emerged contemporaneously and in situ across a wide region. Each of these systems was unique in the taxa exploited and the management strategies applied, and they generally lack analogs in later pastoral economies.
In the Turkish Euphrates valley, small-scale intensive management of sheep, perhaps including foddering but not young male culling, was practiced as a complement to a hunting economy focused on gazelle, while in the Syrian Euphrates, it was aurochs that came under human control with a focus on culling adult females. At Çayönü, on the upper Tigris, boar was the central focus of early management strategies. In the uplands of eastern Turkey, a combination of juvenile goats and pigs were heavily exploited but culling focused on large adult males as well. In central Anatolia, sheep were penned and herded locally on a large scale. In western Iran, unique early management systems focused exclusively on goats and the intensive culling of young males—a practice perhaps developed in order to reduce the aggressive behavior of adult rams in large herds. In the southern Levant, small-scale management of local goat populations likely emerged within hunting economies, while in the far south, ibex (Capra nubiana) may have been subject to experimental management (Ducos, 1997).
That these systems were highly localized and experimental is supported by the fact that at settlements such as Nevalı Çori and Çayönü, animal management was initially a small part of animal economies that continued to rely on hunting. But even in communities that applied management practices on a large scale, such as Aşıklı in central Anatolia and Ganj Dareh in western Iran, large numbers of fetal and perinatal caprine remains suggest that captive animals were kept under biologically stressful conditions resulting in spontaneous abortions and/or high infant mortality. The general lack of morphological changes in predomestic managed animal populations is another sign of the difficulties posed by early herding experiments. The lack of evidence for changes in phenotype suggests that managed herds were not isolated from their wild counterparts and instead reflects regular recruitment from free-living wild populations. Although some of this introgression may have been unintentional (especially for pigs (Ottoni et al., 2013)), it is also possible that early herd management practices were simply not very productive when applied to phenotypically wild animals and that herds were unstable and vulnerable to collapse and failure.
A Pastoral Revolution
By the early 8th millennium bce, changes in the faunal record indicate a major transformation in animal economies across southwest Asia. Beginning in the early 8th millennium bce, but really taking hold by mid-millennium, there is evidence for a massive shift in regional animal economies toward large-scale sheep and goat management with a concomitant decline in evidence for hunting large game (Arbuckle, 2012). This shift is evident in increases in the relative frequency of sheep and goats across the Fertile Crescent but is especially dramatic in the lowland regions dominated by earlier traditions of gazelle hunting (Figure 2).
This period also witnesses the first widespread appearance of morphological changes, including a decrease in body size and changes in horn morphology, signaling the appearance of mature populations of biologically domestic sheep and goats (and also cattle and pigs). In addition, the practice of culling young male animals—a common pastoral management practice that was uncommon in the earliest management regimes—became widespread, as does evidence for animal dung in archaeological sites (Arbuckle & Atici, 2013; Brochier, 1985, 1993; Miller & Smart, 1984). These changes reflect increased investment in a more intensive and homogenous caprine pastoralism, in contrast to the often small-scale and diverse strategies of the 9th millennium, likely reflecting increased access to productive and stable herds of domestic livestock, well adapted to living and breeding under human management, along with the application of mature and effective husbandry techniques. Widespread evidence for the use of dung suggests that these early pastoral systems were tightly integrated into agricultural systems, with herds likely grazing on and fertilizing harvested fields.
Unlike the local development of myriad, predomestic animal management strategies in the early stages of animal domestication, the 8th millennium bce “pastoral revolution” seems to have begun in the Turkish Euphrates region, where it is characterized by a massive increase in the remains of morphologically domestic sheep and goats (Figure 2). Domestic sheep and goats appear to spread from this region into adjacent areas where one or the other was not heavily exploited in the past. For domestic goats, we can follow their movement southward out of their natural habitat in the Turkish uplands into the lowlands of the Syrian Euphrates valley, appearing first at the site of Tell Halula (ca. 7800 bce), slightly later at Abu Hureyra, and then at Bouqras and into central Syrian oasis settlements such as Umm El Tlel around 7000 bce (Clason, 1983; Moore et al., 2000; Sana Segui, 2000). Moreover, Makarewicz and Horwitz have argued that despite evidence for traditions of indigenous predomestic goat management in the southern Levant, morphologically domestic goats appear suddenly in the region in the mid-8th millennium bce, suggesting that productive populations of domestic goats may have been introduced from the north (Horwitz et al., 1999; Makarewicz, 2014; Makarewicz et al., 2016; Makarewicz & Tuross, 2012). Morphologically domestic goats also appear in western Iran at this time as evident at Tepe Sarab and Ali Kosh (Zeder, 2008).
Domestic sheep, the wild ancestors of which are absent from the southern Levant, spread into that region for the first time in the mid-8th millennium bce, likely also reflecting their diffusion from the north (Makarewicz, 2014; Martin & Edwards, 2013). On the island of Cyprus, where sheep bearing a wild phenotype appear ca. 8000 bce, Vigne (2011) has noted changes in sheep husbandry in the mid-8th millennium bce including a massive increase in the relative frequency of sheep remains as well as dramatic changes in body size, which he interprets as evidence for the importation of an entirely new stock of domestic sheep from the mainland. At Jarmo in northern Iraq, we have evidence for the spread of domestic sheep into the eastern wing of the Fertile Crescent by the late 8th millennium where previously predomestic goat management was dominant (Stampfli, 1983). Interestingly, the spread of sheep husbandry into the Zagros region—with its long history of intensive goat exploitation—is delayed by a half millennium; there, sheep management only appears in the Pottery Neolithic (7th millennium bce), at which point it quickly became a central part of local pastoral economies (Zeder, 2008).
Even though sheep and goat pastoralism became the dominant components of southwest Asian animal economies in the 8th millennium bce, local and regional variation is still evident. The chronological and spatial patterns in the spread of domestic sheep and goats clearly indicate that these livestock did not travel as a package but instead followed independent trajectories across Neolithic southwest Asia (Arbuckle et al., 2014; Atici et al., 2017). This is also the case for domestic cattle and pigs. Although regularly referred to as part of a widespread “Neolithic package,” the integrated husbandry of domestic sheep, goats, cattle, and pigs is only evident in a highly restricted region in the early Neolithic (PPNB) (Peters et al., 2005). This integrated “barnyard” system emerges in the late 9th and early 8th millennia bce in the westernmost upper Tigris basin (i.e., Çayönü) and the Turkish Euphrates but is absent in adjacent regions, where animal husbandry focused only on sheep and/or goats. It is this unique, diverse, and integrated agropastoral system centered in the northern Levant that spread westward with the expansion of Neolithic communities in the 7th millennium bce. However, at this time when Neolithic agropastoralists were driving herds of domestic sheep, goat, cattle, and pigs into western Turkey and southeastern Europe, cattle and pig management were only just beginning to infiltrate into the southern Levant (Horwitz et al., 1999). On the central Anatolian plateau, domestic cattle are similarly introduced only in the mid-7th millennium and pig husbandry was never practiced throughout the entirety of the Neolithic (Arbuckle, 2013; Russell et al., 2013a). Similarly, in western Iran, cattle husbandry was not widely adopted until the 6th millennium bce and pig husbandry remained a rarity in the Zagros into the Chalcolithic (Arbuckle et al., 2016; Price & Arbuckle, 2015). Thus, the faunal data reflect a situation in which the emergence of the livestock component of the “Neolithic package” developed over a period of more than two millennia in southwest Asia and was not fully in place until the end of the Neolithic in some areas—despite the position of this region at the geographic epicenter of the process of early animal domestication.
After more than two centuries of scholarly attention, we can piece together the main features of the history of animal domestication in southwest Asia. Current evidence indicates that the histories of domestic livestock, including sheep, goats, cattle, and pigs, have their origins in the increasingly intensive and manipulative human–animal relationships of the 10th millennium bce within communities of sedentary cultivator–hunters across the Fertile Crescent region. These histories are interlinked yet independent and spanned a period of several millennia straddling the late Pleistocene and early Holocene transition. From the earliest initiation of predomestic animal management to the emergence of the first generations of domestic sheep, goats, cattle, and pigs, to the development of the husbandry methods necessary to successfully management them, to the spread of pastoral systems throughout the region, the early history of animal domestication is spread over a period of almost three millennia revealing a remarkably complex tangle of local and regional processes.
Although we know a great deal about the identity of the wild ancestors of domestic sheep, goats, cattle, and pigs and the general chronological and geographic patterns of their domestication, going forward many questions still remain unresolved. Of particular interest is further exploring the methods of control applied to predomestic animal populations in the 10th millennium bce. Here we must decouple ideas of early animal management from the domestication process, which evidence from the island of Cyprus indicates were independent processes (i.e., not all predomestic management systems resulted in animal domestication). Identifying the precise nature of local and diverse early management regimes is necessary in order to understand which practices led to animal domestication and why. This may also lead to the discovery of entirely new traditions of animal exploitation within previously undescribed late Epipaleolithic cultural traditions, which must have existed in areas such as western Syria and Lebanon but which have received little scholarly attention.
A second set of questions remain concerning the motivations for the origins of animal management and why some forms of management spread quickly while others did not (especially cattle and pig management). Early notions of a climate trigger as the motivation for domestication do not adequately explain the diversity of new evidence for animal management, and both the ecological and social contexts of animal exploitation need to be explored more deeply in order to make sense of these changes.
A final set of questions revolve around identifying the complex movements of Neolithic livestock populations across southwest Asia. Evidence from Cyprus clearly shows that Epipaleolithic and early Neolithic peoples moved animals around the landscape, but scholars have had less success identifying these movements on the mainland. Current projects utilizing ancient DNA, isotopes, and methods such as geometric morphometrics have the potential to identify both the movement of specific livestock populations across geographies and also the contributions (or not) of local wild animals on livestock populations, which will have a revolutionary impact on future understanding of the processes involved in the histories of early domestic livestock in southwest Asia. Building on the rich foundation provided by earlier scholars such as Rütimeyer, Duerst, Reed, Perkins, Bökönyi, and Ducos, the next generation of faunal studies will continue to develop our understanding of this crucial and complex transformation, the variables behind it, and its enormous impact on subsequent generations of both humans and animals.
The author wishes to thank the following individuals and institutions for assistance and support during the writing of this article: Cheryl Makarewicz, Levent Atici, Hitomi Hongo, Richard Meadow, Sarah Kansa, Max Price, Hijlke Buitenhuis, Joris Peters, Lionel Gourichon, C. Merih Erek, and Aliye Öztan; and Baylor University, Harvard University, Ankara University, and the University of North Carolina at Chapel Hill. Financial support for some of the research described in this article was provided by grants from the National Geographic Society, the National Science Foundation (BCS-1311551; BCS-0530699), and the American Research Institute in Turkey.
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