Skip to main content


Recent archaeological investigations on Cyprus have unveiled unsuspected Late Glacial and Early Holocene (twelfth–tenth millennia cal BP) pieces of the island’s human history. Based on a review of the archaeological data and of the final results of the archaeozoological analyses of Sector 1 of the prepottery site at Shillourokambos, this paper examines how Cyprus improves our understanding of the process of mammal domestication in the Near East. Early introduction of controlled wild animals and then of early domestic lineages provides information about the modalities of the domestication process on the mainland. This information emphasizes the importance of technical skills, of local opportunities and adaptations, and of long-distance and increasing exchanges in the larger Near East area. Cyprus was a recipient of wild or domestic taxa from the continent through recurrent introductions, but it was fully part of the wider area of incipient farming, as seen in local innovations such as the intensive hunting/control of wild deer and boar or local domestication of wild/feral goats. The transition to farming during the tenth millennium appears to follow an unstable and opportunistic Early and Middle Pre-Pottery Neolithic B phase of low-level food production based on rapidly changing combinations of hunting, control, and breeding.


During recent decades, many new data have been accumulated by archaeology, archaeozoology, and genetics on the beginnings of mammal domestication in several regions of the world. This includes the Near East, where the study of early domestication began very early and has provided more evidence and models than many other regions (e.g., Davis 2005; Dobney and Larson 2006; Redding 2005; Zeder 2006, 2011; Zeder et al. 2006). However, even in the Near East, our knowledge remains poor, as demonstrated by the fact that many recent field projects and archaeozoological studies have led to a reconsideration of the general scenarios (cf., e.g., Bar-Yosef and Meadow 1995; Ducos 1968; Helmer 1992; Legge 1996; Zeder 2005).

Since the 1990s, Cyprus has provided startling new evidence from archaeological discoveries dating to the late eleventh and the tenth millennia cal BP (Guilaine and Le Brun 2003; Peltenburg and Wasse 2004),1 contemporaneous with the end of the Early Pre-Pottery Neolithic B (EPPNB) and the Middle Pre-Pottery Neolithic B (MPPNB) and Late Pre-Pottery Neolithic B (LPPNB) of the mainland. Because islands are clearly defined territories, because the Mediterranean islands were massively colonized beginning only in the Neolithic (Cherry 1990), because their endemic native fauna lacked any of the wild ancestors of domesticates (Vigne 1999), and because Cyprus is the only large island in the Near East, it provides unique information. As Cyprus was necessarily dependent on influences from the continent, at least at the beginning of the Neolithic, it may be thought of as an observation post recording what occurred on the nearby mainland.

In this paper, we present a synthesis from the Cypriot Pre-Pottery Neolithic (PPN) perspective on early mammal domestication in the Near East. This is based mainly on the recent results of the final analyses of the first sector of Parekklisha, Shillourokambos (Limassol district), the largest known Cypriot PPN site (10,400–9000 BP; Guilaine 2003; Guilaine and Briois 2007; Guilaine, Briois, and Vigne 2011).

Our approach to animal domestication lies within both the technological (Mauss 1947 [1967]) and the structuralist conceptual frameworks (Lévi-Strauss 1958): technoeconomic and symbolic uses of animals by human societies are part of their technical, social, and symbolic systems and thus a characteristic part of their cultural systems (Vigne 1998). However, as part of the ecosystem (another structural entity), the relationships between animals and humans also involve ecological approaches and should be considered within the structural framework of the “anthroposystem,” that is, a metasystem that groups together the cultural and ecological systems and their interactions and dynamics through time (Muxart et al. 2003; Pascal, Lorvelec, and Vigne 2006; Vigne 2011).

From this viewpoint, in addition to the different kinds of ecological affinities of animals to the human-made or human-modified ecosystems, including commensalisms, domestication appears as a process of intensification of animal-human relationships (Pascal, Lorvelec, and Vigne 2006) boosted by human intentionality.2 Consequently, we consider that control in the wild and control/protection/use of commensals are initial stages of the domestication process, although they involve animal populations that are still wild from a biological viewpoint. Such situations can be detected mostly as an increase in the frequency of the targeted species, as changes in age and sex proportions in the archaeological record (Zeder 2009, 2011), and/or as transport. We restrict the term “domestic mammals” to animal populations that show phenotypic modifications due to human breeding/herding. As some of these cannot be archaeologically detected, sometimes the domestic status cannot be determined. In the same way, we restrict the term “farming” to technoeconomic systems based mostly on animal (and plant) breeding, which appears to have started in the middle of the tenth millennium (transition between MPPNB and LPPNB) in large villages of the Near East (Vigne 2008, 2011). We distinguish farming from “low level food production” as defined by Smith (2001).

The Early Neolithization of Cyprus: A Brief Updated Review

The island of Cyprus emerged from the bottom of the Mediterranean Sea during the Miocene and has never been connected to any continent (Held 1989). As a result, the Upper Pleistocene terrestrial mammalian fauna was reduced to only four or five endemic species (Boekschoten and Sondaar 1972; Simmons 1999): mouse (Mus cypriacus; Cucchi et al. 2006), genet (Genetta plesictoides), dwarf elephant (Elephas cypriotes), dwarf hippopotamus (Phanourios minutus), and perhaps a shrew.

The upper layer of the rock shelter at Akrotiri Aetokremnos provides the earliest known evidence of visits to the island by humans, dated to 12,776–12,461 cal BP (nine wood charcoal dates; Simmons 1999; Simmons and Mandel 2007), corresponding to the Late Natufian in the northern Levant. Simmons (1988, 1999) has proposed that these human groups hunted elephants and hippos to extinction. This is, however, questioned by numerous authors (e.g., Ammerman and Noller 2005; Binford 2000; Bunimovitz and Barkai 1996; Davis 2003; Olsen 1999; Vigne 1999; Wasse 2007), who argue that the two native large mammals disappeared before that time for climatic (Bromage et al. 2002) or anthropic reasons (or both; Wasse 2007). The diet of the occupants of Aetokremnos shelter seems to have consisted mainly of fish, shellfish, and birds (Simmons 1999). Ammerman et al. (2006, 2008) recently found two other sites that may be more or less contemporaneous with Aetokremnos, but no radiometric data have corroborated their interpretations until now.

In addition to thousands of hippo bones, 18 suid bones were found at Aetokremnos, mainly in the upper layer (Simmons 1999). They have recently been directly dated to 11,746–11,396 cal BP ( BP [AA79923; degraded bone collagen]; δ13C, −25.1‰; Vigne et al. 2009). These suids were 9%–20% smaller than the Near Eastern Late Glacial and Holocene wild boars and were also significantly smaller than those of the PPN and Early Pottery Neolithic of the Near East. The most probable scenario (already partly suggested by Wasse 2007) is that wild boars were artificially introduced to Cyprus sometime at the end of the Late Glacial (Natufian on the mainland), rapidly decreased in size because of the isolation effect, and then were hunted by people who frequented Aetokremnos during the second quarter of the twelfth millennium (Late Khiamian on the mainland; fig. 1).

Figure 1. 
Figure 1. 

Evolution of large mammals on Cyprus during Neolithization, with particular focus on the data from Shillourokambos. The chronocultural frame is that proposed by Hours et al. (1994). Radiocarbon dates are given in cal BP and calibrated at 1σ (Calib Rev 5.0; Reimer et al. 2004). For each chronological phase of Shillourokambos, a box on the right-hand side contains the following information: the total number of identified specimens (NISP), the NISP of large mammals, the material dated (Ch = charcoal; Cl = bone collagen), and a reference (1, Simmons 1999 [only the nine charcoal dates]; 2, Vigne et al. 2009; 3, Vigne et al., forthcoming; 4, Manning et al. 2010; 5, Peltenburg 2003; Peltenburg et al. 2001b; 6, Guilaine 2003; Guilaine, Briois, and Vigne 2011; 7, Le Brun and Daune-Le Brun 2003 [Khirokitia, earliest level, G]). The Middle A1 and A2 phases of Shillourokambos are grouped together. Black arrows indicate the earliest evidence for the arrival of new species of fauna on Cyprus. Gray arrows indicate the probable introduction of new lineages of domestic or commensal animals to Shillourokambos and Khirokitia.

Figure 1. 
Figure 1. 

Evolution of large mammals on Cyprus during Neolithization, with particular focus on the data from Shillourokambos. The chrono-

cultural frame is that proposed by Hours et al. (1994). Radiocarbon dates are given in cal BP and calibrated at 1σ (Calib Rev 5.0; Reimer et al. 2004). For each chronological phase of Shillourokambos, a box on the right-hand side contains the following information: the total number of identified specimens (NISP), the NISP of large mammals, the material dated (Ch = charcoal; Cl = bone collagen), and a reference (1, Simmons 1999 [only the nine charcoal dates]; 2, Vigne et al. 2009; 3, Vigne et al., forthcoming; 4, Manning et al. 2010; 5, Peltenburg 2003; Peltenburg et al. 2001b; 6, Guilaine 2003; Guilaine, Briois, and Vigne 2011; 7, Le Brun and Daune-Le Brun 2003 [Khirokitia, earliest level, G]). The Middle A1 and A2 phases of Shillourokambos are grouped together. Black arrows indicate the earliest evidence for the arrival of new species of fauna on Cyprus. Gray arrows indicate the probable introduction of new lineages of domestic or commensal animals to Shillourokambos and Khirokitia.

At least three recent discoveries indicate that whether permanently or not, humans continued to frequent the island in the twelfth millennium cal BP and during the first half of the eleventh. At Ayios Tychonas Throumbovounos, François Briois and collaborators (Briois, Petit-Aupert, and Péchoux 2005; Guilaine and Briois 2007) found an eroded site with large flint series similar to Mureybet II (Khiamian of the Middle Euphrates valley, late Dryas). Unfortunately, there were no associated faunal remains (F. Briois and J.-D. Vigne “Throumbovounos (Ayios Tychônas, Chypre), rapport de sondage,” unpublished report for the Department of Antiquities, Cyprus, 2003). At Agia Varvara Asprokremnos, painted stone vessels, pendants, a shaft-straightener, and unidirectional flint débitage provide strong parallels to materials dated to the late Pre-Pottery Neolithic A [PPNA]/transitional EPPNB in the northern Levant (McCartney 2010; McCartney et al. 2007, 2008, 2010). The associated fauna is dominated by suids (Sus scrofa). A series of six radiocarbon dates ranging between BP and BP spans a likely calendar age of 10,846–10,675 cal BP (Manning et al. 2010). The last site, Ayios Tychonas-Klimonas, is also characterized by a PPN (EPPNB-type) industry (Guilaine and Briois 2007) associated with suids and dog bones (Vigne et al., forthcoming). The subsistence of the occupants of the site should have been based on hunting of only one game species, the small autochthonous wild boar, with the help of dogs. Only one wood charcoal (Prunus sp.) radiocarbon date indicates the first quarter of the eleventh millennium cal BP (11,070–10,741 cal BP, BP [AA88551]; δ13C, −27.3‰; Vigne et al., forthcoming).

The lithic débitage of these three sites and the radiocarbon dates of two of them clearly postdate those of Aetokremnos and predate the earliest phase (A) at Shillourokambos (Guilaine 2003; Guilaine and Briois 2007; Guilaine et al. 2000) and Kissonerga-Mylouthkia (Peltenburg 2003; Peltenburg et al. 2000, 2001a, 2001b), both of which are radiocarbon dated (fig. 1). The Early A phase of Shillourokambos, where suids are still overwhelmingly dominant (Vigne, Carrère, and Guilaine 2003), provided 12 dates (Guilaine, Briois, and Vigne 2011) ranging from 10,696 to 10,297 cal BP (mean: 10,380 cal BP). Well 116 at Mylouthkia is dated to 10,740–10,290 cal BP ( BP [OxA7460]; BP [AA33128]; BP [1133129]; Peltenburg 2003). At Shillourokambos, the blade technology of this late eleventh-millennium phase is very well documented and shows clear relationships with Early MPPNB in the Levant (Briois 2003).

From the beginning of the tenth millennium, prepottery Cypriot sites became more numerous (Guilaine and Briois 2007; Guilaine and Le Brun 2003; Peltenburg and Wasse 2004; Wasse 2007). They show a progressive shifting of all the material culture toward a local Cypriot model, which culminated during the ninth and eighth millennia BP in the Aceramic Khirokitia culture, contemporaneous with the pre-Halaf/Halaf phases of the northern Levant (fig. 1).

The Main Archaeological and Archaeozoological Characteristics of Shillourokambos

Shillourokambos is an open-air site located on the southern coastal plain of Cyprus, near Limassol (Guilaine 2003). From 1992 to 2004 it was excavated over more than 5,000 m2 under the responsibility of Jean Guilaine. Because it is located on the top of a low hill near the confluence of two small rivers, it has been strongly eroded. No building is preserved in the upper part of the site (Sector 1, ca. 3,000 m2), which is mainly composed of large sedimentary layers and numerous pits, narrow ditches, post holes, and wells. The bases of several buildings have, however, been preserved in the less eroded lower part of the site (Sector 3, ca. 2,500 m2).

Four main phases of occupation of the site have been recognized (Early A, Early B, Middle, and Late), spanning the period from ca. 10,400–10,300 cal BP to the end of the tenth millennium (fig. 1; Guilaine 2003; Guilaine et al. 2000). Thirty-seven radiocarbon dates from Sector 1 suggest that at least this part of the site had been abandoned for several decades or centuries between the Early phases A and B, at the end of the eleventh millennium (Guilaine, Briois, and Vigne 2011). However, it was occupied continuously during the tenth millennium.

Preliminary analyses of the animal remains of Sector 1 (Vigne et al. 2000) showed that dog (Canis familiaris), fox (Vulpes vulpes), pig (Sus scrofa), the Mesopotamian fallow deer (Dama mesopotamica), goat (Capra aegagrus/hircus), sheep (Ovis aries), and cattle (Bos primigenius/taurus) were already present before or at the very beginning of the tenth millennium. Cat (Felis silvestris lybica) was thought to have been introduced during the middle phases and partly controlled (“tamed”; Vigne et al. 2004). In the first preliminary study, all the ungulate species except the fallow deer were considered to have been introduced in the form of already-domesticated lineages, mainly because of their small size in comparison with their wild counterparts on the mainland. This interpretation has been questioned by Horwitz, Tchernov, and Hongo (2004), who suggested, though without any direct analytical data, that they were introduced as wild animals for stocking the island with meat sources.

Further investigation of the Early phase B at Shillourokambos suggested a more complex situation (Vigne, Carrère, and Guilaine 2003; Vigne and Guilaine 2004): suids, sheep, and cattle were small in size and might therefore be considered as resulting from early domestication; but morphological analyses and the slaughtering profiles suggest that (almost?) all the goats and approximately half of the suids were wild/feral and were obtained by hunting, while sheep and probably cattle were bred. These observations underlined the unstable status of the domestic ungulates during several centuries after the earliest domestication and suggested that the human societies on the mainland could also have released domestic (or even a mixture of wild and transported domesticated) ungulates into the wild and redomesticated them many times during the 15 centuries of the Pre-Pottery Neolithic B (PPNB), as well as later.

At that time, the question of what happened on Cyprus before the beginning of the tenth millennium (i.e., during the Early phase A) could not, however, be answered, because this earliest phase had provided less than 100 specimens. The last season of excavation at Shillourokambos in 2004 produced many more animal bones and much more archaeological information for this phase (Guilaine et al. 2008). The final study of all the animal bones of Sector 1 could be carried out (Guilaine, Briois, and Vigne 2011). Jean-Denis Vigne and Isabelle Carrère were in charge of the archaeozoological analyses and were active participants in the excavation. Consequently, they were in a good position to select the most reliable faunal assemblages.

The first important result of this final analysis is that the proportions of the different mammal species vary significantly throughout the successive occupations of the site to such an extent and in such a well-structured way that it was possible independently to find again the four phases based on the lithics only through faunal characteristics. Correspondence analysis of the faunal spectra even allows refinement of the chronology and identification of several subphases: Early A1 and A2 (the second being more or less the initial stage of the Early phase B), Early C, and Middle A1, A2, and B phases (the last more probably being the first stage of the late phase). This new chronological framework enables a more precise approach to the succession of events on this site between 10,400 and 9000 cal BP.

The final archaeozoological results presented in this paper are based on 32,500 animal remains, of which 9,225 (the number of identified specimens [NISP]) are taxonomically determined (8,864 vertebrates and 361 invertebrates; Guilaine, Briois, and Vigne 2011). On the basis of cut marks (Vigne 2006), fragmentation, and skeletal frequencies, reconstruction of the butchery and cooking processes and estimation of the distance between the kill sites and the village for each species of ungulate were possible. Nearly 11,000 bone and tooth measurements (table 1) provided statistics and enabled multivariate analyses. For deer, goat, cattle, and to a much lesser extent sheep, mixture analyses (Monchot and Léchelle 2002) provided reliable estimates of the sex ratio of adults and the decrease in sexual dimorphism due to domestication (see Helmer 2008; Helmer et al. 2005). By this technique, the evolution of the size of the different parts of the skeleton of the dimorphic ungulates could also be studied for each sex, providing reliable approaches to morphological variations through time. The tooth and epiphysal mortality profiles were based on large series of teeth and long bones (table 1) and processed according to Vigne and Helmer (2007). We did not use statistical comparisons (see Marom and Bar-Oz 2009), which are not adapted to these questions (Vigne 2000), but rather multivariate analyses (e.g., Helmer, Gourichon, and Vila 2007).

Table 1. 

Final archaeozoological analyses of the large mammals from Sector 1 at Shillourokambos

 MorphometricsCulling profiles
Animal speciesNISPNo. measurementsNo. teethMNINISP epiphyseal
Vulpes vulpes56214   
Canis familiaris1177   
Felis silvestris lybica311   
Sus scrofa1,0012,9174721131,124
Dama dama mesopotamica1,3614,0177851732,374
Capra aegagrus/hircus3941,284   
Ovis aries3781,318   
Total Caprini9913,117321108754
Bos taurus124336  26

Note. Number of measured specimens, measurements, and teeth, the minimum number of individuals (MNI) used for the culling profiles, and the number of the bones used for epiphyseal profiles. NISP = number of identified specimens.

View Table Image

These final analyses led to modification and clarification of several aspects of the preliminary interpretations. Figure 1 presents the frequencies of species according to the phases. During the Early phase A1, suids, which are the same size as those at Aetokremnos, are dominant (90%). Cats, goats, and cattle are clearly present in small numbers. The presence of dog is probable, that of fox questionable. The statistical analyses indicate that the absence of fallow deer and sheep is very unlikely to be due to the size of the sample (). It is confirmed by the absence of both species in well 116 at Mylouthkia, which is contemporaneous with Early phase A of Shillourokambos (Peltenburg et al. 2001b). Deer appeared as the dominant species at the very beginning of Early phase B. Sheep probably arrived slightly later.

Early phase B is characterized by high frequencies of cattle (8%–12% of the total NISP of the phase), which decrease to less than 1% in the middle and late phases. The Middle A phases are characterized by an increase in suids and foxes. Sheep and goats become dominant in the Middle B and late phases.

Transport and Control: The Beginnings of Mammal Domestication before 10,300 Cal BP

Faunal Turnover on the Island

Although Mus cypriacus is attested to at Mylouthkia and is present today on the island (Cucchi et al. 2006), animal bones at Asprokremnos, Klimonas, Mylouthkia, and Shillourokambos confirm that the Pleistocene endemic hippo fauna did not survive the Late Glacial–Holocene transition. According to Bunimovitz and Barkai (1996) and Ammerman and Noller (2005), this fauna would have been extinct before the human occupation of layer 2 at Aetokremnos in the first centuries of the twelfth millennium (see also Wasse 2007). The introduction of wild boar before the occupation of Aetokremnos renders it possible that people frequented the island before 12,000 BP and then caused the extinction of the hippos.

The endemic Cyprus genet (Genetta plesictoides) is present in the Aetokremnos deposits without any direct dating (Simmons 1999). It is absent from the small-mammal fauna of Mylouthkia. It is unlikely that it became extinct at the same time as hippos because its presumed main prey, M. cypriacus, did survive. The introduction of dogs (before 11,000 cal BP) or cats (mid-eleventh millennium at the latest) may have caused the extinction of the genet sometime in the twelfth–eleventh millennia.

Suids, cats, goats, cattle, and dogs were living on Cyprus before or starting from the earliest phase at Shillourokambos (fig. 1). In addition, data from Mylouthkia provide evidence for the presence of the house mouse (Mus musculus domesticus; Cucchi et al. 2002). Because they have no Cypriot Pleistocene ancestor and because such a high rate of immigration can result only from humans (Vigne 1999), these species would have been introduced between the end of the Late Glacial and ca. 10,400 cal BP.

All the large true Mediterranean islands have undergone a similar turnover of their mammal fauna. That on Cyprus is the earliest, beginning during the Late Glacial, because domestication in the Eastern Mediterranean was early (Vigne 1999).

Pre-Neolithic Wild Boar Management More than 11,400 Years Ago

The presence of small wild boar dated to 11,746–11,396 cal BP at Aetokremnos, the dominance of suids at Asprokremnos, Klimonas, and Shillourokambos Early A1, and the morphological similarities of suids at Aetokremnos and Shillourokambos are convergent evidence that small island suids were living on Cyprus and exploited by humans during the twelfth–eleventh millennia. This would explain why people frequented (or inhabited) Cyprus during this period.

The introduction of wild boar during the Late Natufian or Early Khiamian strengthens the hypothesis of control of wild boar in the Final Late Glacial, as proposed by Redding and Rosenberg (1998), Rosenberg and Redding (1998), and Rosenberg et al. (1998; see also Redding 2005; Starkovitch and Stiner 2009; Wasse 2007) on the basis of the Hallan Çemi site in the upper Tigris basin. The evidence from Cyprus makes a strong contribution to the picture of a long span of increasingly intensive and skilled control of wild boars. This took place over a very large area, at least in eastern Anatolia (and rapidly expanded to Cyprus), but it would have extended to western Anatolia and the Aegean, as suggested by evidence of small Mesolithic wild boar on the islands of Youra and Kythnos during the eleventh–tenth millennia (Trantalidou 2008). Here and there, it developed into true pig breeding in the middle of the eleventh millennium, as observed in the Upper Euphrates valley at Nevalı Çori and Gürcütepe (Peters, von den Driesch, and Helmer 2005) and at Cafer Höyük (Helmer 2008). At the same time, control in the wild would have continued during a large part of the tenth millennium, as suggested at Çayönü (Ervynck et al. 2001) and in the Aegean.

Transport of Wild or Early Domestic Goats?

In our present state of knowledge, the earliest evidence of goats on Cyprus is about 40 bone specimens at Shillourokambos Early A. The final analyses for Shillourokambos confirmed that the morphology of the horn cores of the early phases was similar to that of the wild bezoar goat. But separate morphometric analyses of males and females has shown that contrary to our preliminary conclusions (Vigne, Carrère, and Guilaine 2003; Vigne et al. 2000), these animals were significantly smaller than the early domestic goat, as measured by Helmer (2008) at Cafer Höyük (Early MPPNB). Peters, von den Driesch, and Helmer (2005), Helmer (2008), and Hongo et al. (2009) presented convincing evidence of domestic goat in the Upper Euphrates and Tigris valleys in the middle of the eleventh millennium (late EPPNB), and Helmer and Gourichon (2008) did the same for the beginning of the MPPNB of the Damascus plain (Tell Aswad), ca. 10,300–10,200 BP. It is thus likely that the earliest goats of Cyprus were introduced as already modified domesticates.

However, on the basis of convincing theoretical inferences, Redding (2005) argued for wild-goat control before the PPNB (see also Hole 1996). Genetic investigations of the Near and Middle Eastern modern bezoar goats (Capra aegagrus) have shown that the wild lineages that mothered modern domestic goats in the Zagros were subjected to a drastic demographic increase ca. 10,000 BP, whereas the other lineages showed no similar genetic signature (Naderi et al. 2008). They also suggest that wild goats might have been transferred out of the original area of the species, though still in mountain areas. Like that of wild boar, the domestication of goats seems to have begun as control in the wild in a very large area stretching from the central Iranian plateau to southeastern Anatolia. Like the pig, morphologically domestic goat would have appeared here and there, as seen in the data from Nevalı Çori, ca. 10,500 cal BP (Peters, von den Driesch, and Helmer 2005), and later (10,000–9500 cal BP) in the Zagros (Zeder 2005, 2011; Zeder and Hesse 2000).

The final analyses at Shilloroukambos confirmed that (all?) goats were hunted not only during the Early phase B (Vigne, Carrère, and Guilaine 2003) but also during the Early phase C and the beginning of the middle phases. The bone sample for the Early phase A is too small for characterizing the mode of exploitation, but the low frequency in comparison with the middle phases (fig. 1) makes hunting more likely. It is therefore difficult to determine whether the introduction of goats to Cyprus in the course of the eleventh millennium resulted from the transport of controlled bezoar goats (as suggested by Horwitz, Tchernov, and Hongo 2004), which would have decreased in size because of insularity, or was a consequence of a rapid spread of early domestic goats. The ecology of the bezoar goat (which excludes its presence in the coastal plains, from which the visitors to Cyprus necessarily came), its introduction much later than the wild boar, and the absence of any evidence of goat control in the nearby mainland areas, however, make the latter scenario more likely. In any case, Cyprus provides strong evidence that the process of goat domestication in the Near East was well advanced in the middle of the eleventh millennium.

From Commensalism to Domestication: Mice, Cats, and Other Carnivores

Cucchi et al. (2002) demonstrated that the house mouse (M. musculus domesticus), absent from Aetokremnos (Simmons 1999), was well represented in well 116 of Mylouthkia, dated to 10,740–10,290 cal BP, together with the less commensal endemic Cypriot mouse, M. cypriacus. The earliest evidence for cat at Shillourokambos is a complete fifth right metacarpus found in the bottom of the earliest stratigraphic unit, dated to 10,322–10,288 cal BP (Guilaine, Briois, and Vigne 2011). Although there are only two cat remains in Sector 1 of Shillourokambos (fig. 1), the species is frequently present on tenth to eighth millennium Cypriot sites (review in Vigne and Guilaine 2004). The house mouse and the cat were introduced to Cyprus in the middle of the eleventh millennium at the latest, and the probably unintentional transport of the former possibly led to the introduction of its predator.

Auffray, Tchernov, and Nevo (1988) showed that house mice were already commensal in the Natufian layers at Hayonim, but nothing was known about early domestication of cats in the Near East. The presence of both house mice and cats in Cyprus during the second half of the eleventh millennium not only confirms that the house mouse spread as a pest to all the territories of the PPN complex (Cucchi and Vigne 2006), including islands, but is also evidence that cat domestication had already begun. It also supports the idea, suggested by Malek (1993) in relation to Egypt, that the most likely reason for cat domestication was use of this predator against the commensal rodents and their consumption of agricultural products. Cat domestication would have followed two steps. First, “commensalization” would have occurred, that is, cats being attracted to the villages by high concentrations of mice, whose presence was due to an increase in stocked foodstuffs. Second, the villagers would merely have provided protection or simply impunity to some individuals for the commensalism to evolve toward domestication (Pascal, Lorvelec, and Vigne 2006). This would have been all the more true if the cat had a symbolic significance for those Neolithic societies, as suggested by the numerous representations of felids in the PPN (Helmer, Gourichon, and Stordeur 2004), including in Shillourokambos Early A (Guilaine et al. 1999).

Should we expect future evidence for domestic cats in the Natufian on the mainland? This is not certain, because the proposal of Auffray, Tchernov, and Nevo (1988) has to be reevaluated with modern morphometric techniques and because, although it began during the Natufian, food storage does not seem to have developed on a large scale before the PPNA–EPPNB (Kuijt 2008; Kuijt and Finlayson 2009). In addition, the abundance of foxes in the PPNA at sites such as Jericho (Clutton-Brock 1979) and their introduction to Cyprus at the beginning of the tenth millennium (fig. 1) indicate that other species of small carnivores could also have played the same role as cats throughout the PPN.

As domestic dogs are in evidence on the mainland since the Natufian (Helmer 2008; Tchernov and Valla 1997) and on Cyprus as early as the beginning of the eleventh millennium (Vigne et al., forthcoming), it would not be surprising to find them on Cyprus during the twelfth millennium. They probably played a role in autochthonous-wild-boar hunting or control at Klimonas during the first half of the eleventh millennium. At Shillourokambos, the scarceness of their remains and the total absence of any gnawing mark on the other mammal bones suggest, however, that they lived away from the village, at least from the beginning in the Early phase B (Vigne and Guilaine 2004).

Introduction of Early Domestic Cattle during the EPPNB

Although absent at Mylouthkia, cattle were present during the Early phase A1 of Shillourokambos, represented by a complete right pelvis found at a depth of 4 m in the Early A1 fills of well 431. Cattle were introduced to Cyprus ca. 10,300 cal BP or shortly before (fig. 1), at a time when domestic cattle had begun to appear on the mainland (Helmer et al. 2005; Hongo et al. 2009).

Cattle bones are, however, too rare in the earliest phase at Shillourokambos to provide any information on their morphology and status. Their size does not seem to differ significantly from that of Early phase B cattle bones. The latter are significantly smaller than those of the aurochs of the Natufian and PPNA layers at Mureybet (Gourichon and Helmer 2008). They are very similar to those of Dja’de (L. Gourichon and D. Helmer, unpublished data), which are mainly composed of early domestic bovids (Helmer et al. 2005). Estimated through mixture analyses of log size index, the sexual dimorphism of the bovid from the early phases of Shillourokambos appears also to be significantly less than that of the aurochs of Mureybet I–IIIA and similar to that of Dja’de. In addition, the presence on the site of all the skeletal elements of the limb extremities and all the vertebrae suggests that the animals were killed very near or in the village. The epiphysal age profile and sex ratio of the early phases indicate a well-managed breeding strategy focused on the exploitation of the meat from the young adult males.

The early Cyprus cattle bred during the Early phases at Shillourokambos thus probably resulted from the introduction of already-domestic lineages such as those at Dja’de or at less distant Tell Aswad (Damascus), where Helmer and Gourichon (2008) found evidence that cattle were used for carrying heavy loads as early as the beginning of the MPPNB.

Tenth Millennium: Cyprus Reflects Continental Complexity and Intensive Exchanges

Numerous archaeozoological data from the Early phase B to the Late phase of Shillourokambos are summarized in figure 2 (after Guilaine, Briois, and Vigne 2011). This provides an exceptional case study for the evolution of the status of ungulate species and of subsistence strategy through the tenth millennium, a time of emergence and consolidation of farming on the mainland (Bar-Yosef and Meadow 1995; Vigne 2008, 2011).

Figure 2. 
Figure 2. 

Summary of the technicoeconomic and sociosymbolic characteristics of ungulates through the different chronological phases of Shillourokambos, after Guilaine, Briois, and Vigne (2011). Shading indicates the importance of hunting; darker shading indicates greater importance.

The Mesopotamian Fallow Deer

The final archaeozoological analyses provide evidence that deer were never domesticated at Shillourokambos. In all phases, age classes were slaughtered in proportion to their natural abundance without any distinction between sexes. The morphological distance between males and females remained constant, and the body size not only did not decrease but slightly and constantly increased from the tenth to the eighth millennium (Khirokitia), perhaps as a result of constant hunting pressure.

The late introduction of the Mesopotamian fallow deer to Cyprus (fig. 1) probably illustrates the persistence, several centuries after the beginning of mammal domestication, of wild-ungulate control similar to that which we have shown for wild boars during the twelfth–eleventh millennia. Even if the economic importance of the fallow deer in Cyprus appears to be unique in all the PPN and therefore specifically Cypriot, it does suggest that somewhere in the Near East, human groups were already controlling fallow deer either as a domestication experiment or as a common subsistence practice.

Who were these pioneers of deer control? The Cypriot fallow deer were noticeably smaller than their northern-Levant PPNA counterparts at Mureybet (according to Gourichon and Helmer 2008). The highest density of archaeological evidence for the Mesopotamian fallow deer is located in the southern Levant (Davis 2003). The PPNB complex of cultures reached the southern Levant only at the end of the eleventh millennium BP (Khalaily, Marder, and Barzilai 2007), several centuries later than in the northern Levant. Mammal domestication was then in a more experimental phase in the south than in the north (Conolly et al. 2011). All this may suggest that the fallow deer was introduced from the southern Levant.

Sheep and Goats: Complementary Stories

The large number of bone remains (table 1) of sheep and goats and morphometric, age, and sex-ratio analyses (Guilaine, Briois, and Vigne 2011; Vigne, Carrère, and Guilaine, forthcoming) enable reconstruction of the codevelopment of sheep and goat exploitation at Shillourokambos (fig. 2). Goats were hunted throughout the early phases. Beginning in the middle phases, the villagers of Shillourokambos began to exploit them more intensively, strictly culling the young males, although no additional morphological modification appeared at that time. However, they continued to hunt wild/feral goats. Size decrease and morphological modifications of the horn cores become visible only in the late phases (i.e., 2–4 centuries later). The modification of the culling profiles through time indicates that at that time, the exploitation of goats had shifted toward their milk (Vigne, Carrère, and Guilaine, forthcoming). Even though this concerns goat lineages that may have been domesticated long before on the mainland and then became feral on the island, this is the first evidence for a domestication process on a Mediterranean island. It resulted in a “new” lineage of domestic goats ca. 9400–9000 BP, that is, more than a millennium later than in sites of the northern Levant such as Nevalı Çori, Çayönü, Tell Halula, or Tell Aswad (Helmer and Gourichon 2008; Hongo et al. 2009; Peters, von den Driesch, and Helmer 2005; Saña Segui 1999).

Sheep were introduced up to 5 centuries later than goats (fig. 1), in the form of small, horn-modified domestic animals (Vigne, Carrère, and Guilaine 2003) similar to their contemporaneous counterparts of the MPPNB at Tell Aswad. As soon as Early phase B, they were herded in a sophisticated way for meat and milk production (“milk B,” as defined by Vigne and Helmer 2007). Just after an episode of rapid size decrease and stress increase (malnutrition marks on the horn cores) that we interpreted as a breeding failure or collapse (Middle A1), new larger-sized lineages abruptly appeared, possibly introduced from the central or southern Levant. During the late phases, herding concentrated on meat production and possibly hair production. A second rapid morphological change perhaps suggests another new lineage. Davis (1994) has demonstrated an event of very rapid and significant size change between the early and late levels at Khirokitia and interprets it as the possible introduction of a new lineage. There is no evidence for feralization or sheep hunting at Shillourokambos. The exploitation of sheep on this site was strictly limited to breeding periodically strengthened by the introduction of new lineages that probably came from the mainland.

Basically, for the villagers of early Shillourokambos, goats were game animals, while sheep were the most “domestic” of all the ungulates. This probably implies quite different symbolic values for each of them. Archaeozoology tends to mix these two species only because their bones are difficult to discriminate from one another. The evidence from Shillourokambos demonstrates once again that they are really very different animals from bioecological, technoeconomic, and symbolic points of view (e.g., Balasse and Ambrose 2005). Because of these differences, the two species were able to play complementary roles in the technoeconomic system. If the rapid Middle A1 size decrease of the sheep at Shillourokambos actually resulted from a herding failure, we could consider that the domestication of goats in this village, which began at approximately that time, was developed to offset it. The complementarities between the two species are again illustrated by the end of the story, when the goats take over milk production from the sheep, which are then used for meat and hair production. This scenario provides a good illustration of both a process of domestication (goat) and the early technoeconomic complementarities of sheep and goat during the tenth millennium.

Introduction of Mammals from the Mainland

In addition to the fallow deer and several waves of domestic sheep, we also found evidence for the introduction of new lineages of domestic pig and cattle in the course of the tenth millennium (figs. 1, 2; Guilaine, Briois, and Vigne 2011). Between Early phases A and B of Shillourokambos, the size variability of suids abruptly tends toward smaller values and becomes organized into two peaks. Geometric-morphometrics analyses of the outline of the molars also suggest the presence of two distinct types of shape. During the Middle and Late phases, the two different-sized peaks tend to fuse because of interbreeding between the two lineages, but the two tooth morphotypes persist. This suggests that another lineage of pig was introduced shortly before 10,000 BP and then mixed with the local lineage that had been introduced beginning with the twelfth millennium. As the former was still smaller than the latter, we concluded that it was composed of domestic types coming from the mainland, where they are attested to several centuries earlier (Hongo et al. 2009; Peters, von den Driesch and, Helmer 2005).

The cattle of the Early C, Middle, and Late phases of Shillourokambos were much larger than those of the Early phases A and B. They are approximately the same size as the domestic cattle of the Middle and Late PPNB layers of Halula (see Saña Segui 1999) and therefore significantly larger than the cattle of Dja’de (L. Gourichon and D. Helmer, unpublished data). These data suggest the introduction of a new lineage of domestic cattle between Early phases B and C.

The tooth morphometry of the modern Cypriot house mouse M. musculus domesticus does not differ from that of the mainland mouse and has not varied since the PPN (Cucchi et al. 2002). Taking into account that genetically isolated mice diverge very rapidly from their ancestor (Pergams and Ashley 2001), this morphological stability is evidence of a constant and intensive genetic flow from the mainland to Cyprus from the Neolithic up to today. Mice would have been successfully introduced into Cyprus at least several times per year (Vigne and Cucchi 2005).

The arrows in figure 1 summarize the introductions of new species (black) and new domestic (gray) lineages to Cyprus and more specifically to Shillourokambos during the twelfth–eighth millennia. Of course, introductions are possibly underevaluated because lineages that did not morphologically differ from the autochthonous ones could not be detected. On the other hand, the appearance of new lineages at Shillourokambos did not necessarily result from introductions from the mainland, as some of them may have resulted only from transfers from other Cypriot sites. In any case, figure 1 clearly shows evidence of intensive movements of animals either from the mainland or from other Cypriot sites throughout this period, a phenomenon that has until now been largely unrecognized in the PPN of the Near East. As evidenced by several papers in this volume, mobility is observed in numerous regions of the world at the time of the Neolithization.

Shillourokambos: A Case Study for Incipient Farming in the Near East

Development of Meat Production: From Low Level to Farming

We estimated meat production on the basis of bone weights (Vigne 1992). For the species that were exploited through both hunting and breeding (suids, goats), we drew from the frequencies of age classes of the slaughtering profiles both minimal and maximal estimates of production. The final results are therefore given with a large range of uncertainty, but they show interesting tendencies (fig. 3). These estimates have two other weaknesses: (1) although attested to for sheep and goats at Shillourokambos (Vigne and Helmer 2007), dairying is not taken into account; and (2) the percentages of bone weights do not represent an actual value of meat weight (i.e., 50% does not mean that hunting provided 50% of the meat). We can take into consideration only the relative value for each phase with reference to the others. The least unreliable reference is the Late phase, where we can reasonably consider that breeding produced all the meat for all mammal species except deer, for which the rate of production was 60%, 70%, or 76%, depending on the mode of estimation (weight of meat and offal, according to Vigne [1992], NISP, or bone weight, respectively).

Figure 3. 
Figure 3. 

Evolution of meat production through the chronological phases of Shillourokambos. Bone weight percentages are not exact estimates of meat procurement. Only the relative variations between the different phases can be taken into consideration in this diagram. This is why ordinates are not graduated. EPPNB = Early Pre-Pottery Neolithic B; MPPNB = Middle Pre-Pottery Neolithic B; LPPNB = Late Pre-Pottery Neolithic B.

Figure 3 indicates that meat production increased slowly and irregularly but constantly over time, especially starting in the Middle phases. It always remained below 60%–75% and was probably less than 50% during the Early phases. However, it was probably significant as early as the earliest phases, then represented by cattle and an unclear proportion of the suids (fig. 2). Even if milk production is added, this development implies a long stage of low-level food production (see Smith 2001) with important variations that suggest frequent modifications of the technoeconomic strategies and then a slow transition to farming. The former took place during the mainland MPPNB, when numerous sites on the mainland also show a dominance of hunting/control in their meat procurement (Conolly et al. 2011; Hongo et al. 2009; Vigne and Helmer 2007). The latter closely corresponds to the end of the MPPNB and the LPPNB. This evolution fits the macroregional historical tendency rather well (see Vigne 2008). Shillourokambos can be taken as an example of the numerous local scenarios that led to farming in the Near East (Bar-Yosef and Meadow 1995) and in numerous regions, as evidenced during the Mérida conference.

Evolution of the Technical System: Instabilities and Opportunistic Strategies

Marine resources played only a small role in the animal supply of the Shillourokambos villagers. Fishing was represented only by some large groupers (Epinephelus sp.; Desse and Desse-Berset 2003), and most of the seashells collected were battered, worn, or naturally pierced—that is, already dead on the beach—for ornaments and pendants (Serrand, Vigne, and Guilaine 2005). Most of the protein and lipid supply came from large terrestrial mammals. Botanical data are scarce (Guilaine, Briois, and Vigne 2011; Willcox 2003).

During the Early phase A1, the meat supply came mainly from hunting the small autochthonous Cypriot wild boar. The kill-off profile is abnormally rich for the very old individuals, indicating that the suid population was subjected to low predation pressures. As humans were indeed the only large predator on the island, this suggests hunting of abundant and easy-to-catch populations or control, similar to that which entailed the introduction of wild boar to the island approximately two millennia earlier. Accessibility to wild/feral goats was probably lower because of their natural mountain behavior. Movement of humans between coastal and mountain game cannot be excluded. The apparently extreme specialization of the meat supply of the Early phase A appears, therefore, as an opportunistic adaptation to both the island faunal resources and the coastal-plain environments of the site. In addition, cattle were actually bred on the site, and although tenuous, the evidence suggests that people had already begun to domesticate some suids. Thus, these (perhaps mobile) human groups appear as mostly opportunistic hunters profiting from a favorable island situation and practicing hunting/control and breeding as well. Any attempt to classify them as strictly hunters or farmers would fail.

This image seems to agree with the data from the mainland (Goring-Morris and Belfer-Cohen 2011; Zeder 2011). There is no indication of animal breeding in the southern Levant for the middle of the eleventh millennium, and data collected for southeastern Anatolia and the northern Levant indicate that early domestic goats, sheep, pigs, and cattle actually first appeared at that time (Helmer et al. 2005; Peters, von den Driesch, and Helmer 2005) but that well-controlled hunting of hemiones, gazelles, wild boar, aurochs, and wild sheep and goats still provided the major part of the meat supply (Vigne and Helmer 2007). Only the absence of sheep at Shillourokambos Early A (and at Mylouthkia; Peltenburg et al. 2001b) is surprising, and it indicates that the processes of domestication of sheep and goats in the Near East resulted from two very different processes, as already shown by Zeder (2005) for the beginning of their domestication in Iran.

Early phase B differs highly from the previous one in terms of animal exploitation (fig. 2). The introduction of domestic pigs and sheep, immediately bred with sophisticated techniques, indicate that Cyprus then benefited from some of the late-MPPNB technical innovations of the mainland. As the local goats were quite abundant, the absence of goat introduction and herding reveals a selection among the potential mainland inputs that depended on local availability. Meat procurement also became more eclectic (with reference to the low taxonomic diversity) than during the Early phase A: all the species present—suids, goats, and cattle, but also newly introduced fallow deer and sheep—were exploited, and they contributed almost in the same proportion, complementing the existing subsistence system. In parallel with the Middle PPNB evolution on the mainland, animal husbandry increased in diversity and probably played an increasing buffering role in the seasonal subsistence, with a higher proportion of cattle; with the beginning of seasonal pig breeding, which very quickly mixed autochthonous and new allochthonous lineages; and in general with sophisticated sheep herding for milk and meat. The latter practice, in which lambs are kept alive but shared among the ewes for part of the time (Vigne and Helmer 2007), implies a high technical level for separate herding of young and adults, particular attention to lambs, and processing of milk and its derivatives. For the first half of the tenth millennium, this has been attested to on the mainland only for goats at Cafer Höyük, Tell Halula, and Tell Aswad (Gourichon and Helmer 2008; Helmer 2008; Vigne and Helmer 2007).

However, hunting remained the main source of meat during this Early phase B. Almost immediately after its introduction, the Mesopotamian fallow deer became the major source of meat through a very efficient strategy of exploitation, which was probably collective hunting by beating of mixed herds of males, females, and young (Vigne 2000), quick butchering of carcasses on the kill site, and selection of the most profitable joints. Wild pigs and goats were also subject to intensive hunting based on similar practices. Cooking techniques indicate this intensive exploitation (bone breakage for the consumption of marrow) with relatively standardized and refined practices (different cutting patterns for suids and ruminants, skilled cutting, roasting for some joints and boiling for others). Evidence of seasonal exploitation suggests that deer were hunted in fall and winter, suids were slaughtered in winter, and spring and early summer were devoted to lambs and milk exploitation, together with plant cultivation.

However, in that period, and again more clearly during the Early C and Middle phases, clear evidence of overexploitation of game appears mainly in the form of a demographic erosion of deer populations. This reminds us that island resources were not as diversified and abundant as those on the mainland, where MPPNB villages still practiced diversified hunting of wild equids, gazelles, and aurochs. In this context, the development of skilled breeding of cattle, pigs, and sheep in Cyprus appears not only as a counterpoint to the ongoing evolution on the mainland but also as an answer to the local necessity of dealing with both the limitation of island resources and the probably increasing population of these pioneer human communities.

This vision of the dietary history of Shillourokambos is supported by the increase in technical efforts that appear during the Early phase C. From the end of the Early phase B onward, the archaeozoological data indeed suggest an intensification of henceforth well-controlled pig breeding focused on the production of fat and meat by winter slaughtering of first- and second-year pigs. For the first time since the beginning of the tenth millennium, the NISP of pigs, which mainly result from breeding, exceeds that of deer. The presence of cattle clearly began to decline, perhaps only because cattle breeding is more difficult and less immediately profitable than pig breeding.

Until Middle phase A1, sheep herding continued at a high level. Then there was a strong and rapid decrease in the size of the sheep, together with increasing frequency of environmental stress marks on the horn cores and an extreme slenderness of the long bones. At the same time, pigs also decreased in size, and there was a significant increase in the rate of linear enamel hypoplasia, which indicates an intensification of environmental stress. New innovations and important changes in animal control seem to have emerged very rapidly to compensate for this probable breeding failure. These were based both on imports from mainland “rear bases” and on local improvements of the technical system. As mentioned above, a new, larger domestic sheep lineage arrived from the central or southern Levant as well as a new cattle lineage (which might be at the origin of the slight increase of cattle frequencies that we observed at that time). But the restoration of the system was also based on local resources, not through hunting, as the deer and then pig populations declined, but through the new domestication of the local wild goats (see above). Here again, these Cypriot populations appear to have been involved in a macroregional system of exchanges and progress, but they were also able to provide local responses to the ongoing consolidation of the new Neolithic way of life.

During the Middle phase A2, there followed a complete remodeling of the utilization of animal resources: besides deer (and secondarily goat) hunting, which persisted, seasonal pig breeding, which provided fat, is dominant; from then on, domestic goats progressively played the role of milk producers, as they usually did on the mainland (Vigne and Helmer 2007). Complementarily, the restored sheep herding tends to be devoted to meat production, then also to hair exploitation, a tendency that is also known from some pre-Halaf continental sites, such as Tell Sotto and El Kowm 2 (Helmer, Gourichon, and Vila 2007). The increase of meat production at that time is demonstrated by a 25-fold increase in the density of animal food refuse (in relation to the volume of excavated earth) in comparison with the early phases. This development appears to be a parallel to the intensification of animal breeding on the mainland at the transition between the MPPNB and LPPNB, which corresponds to the emergence of farming (Vigne 2008).

On the other hand, this emphasizes the relatively low level of animal products in the diet of the early phases at Shillourokambos. The decrease in grinding stones at the transition between the Early and Middle phases (Perrin 2011) suggests that plant cultivation, which is well documented at Mylouthkia (Peltenburg et al. 2000, 2001a), would have played a more important role in the diet of the early phases at Shillourokambos than suggested by the relatively poor set of botanical remains (Willcox 2003). The strong development of animal husbandry in the middle of the tenth millennium may be connected to a drastic change from a mostly vegetarian diet to a more carnivorous one. It is also probably connected to a population increase in the village, as seen in Sector 3 of the site, the study of which is still ongoing.

In summary, the history of meat supply at Shillourokambos comes within the scope of the PPNB Near Eastern Neolithic transition. This is seen in the exploitation of local animals by hunting and control in local domestication as well as by breeding, which led to the slow but constant intensification of food production relative to predation, and in genetic inputs from the mainland in the form of new lineages of domestic pigs, sheep, and perhaps cattle. In comparison with the earliest known Anatolian sites with domestic animals, Shillourokambos differs in its slightly later manifestation of local domestication processes (pigs and goats), which are possibly adaptations to the local conditions, because of a low and insular biological diversity. The emergence of Cypriot animal production appears to be a local version of that seen on the mainland, driven by resource shortage and human demographic growth (Bocquet-Appel 2011). This interpretation of the subsistence strategy of Shillourokambos implies a knowledge of seafaring (Broodbank 2006; Vigne and Cucchi 2005), but this was not sufficient for sustainable development of island populations; it also required a high level of control of the sources of subsistence that could be attained only through the Neolithic way of life and strong connections to continental “rear bases.”


Though still poorly documented for the thirteenth–eleventh millennia, the prepottery history of the relationships between mammals and humans on Cyprus provides substantial new information about the beginnings of domestication in the Near East. In that context, the recent Cypriot data document the following: (1) a transition to farming during the tenth millennium, after an unstable and opportunistic late-EPPBN and MPPNB phase of low-level food production based on rapidly changing combinations of hunting, control, and breeding; (2) initiation of this process during the Final Late Glacial, evidenced in Cyprus by control and long-distance transport (and thus acclimatization) of wild boars before the mid-twelfth millennium (Natufian-Khiamian) followed by the introduction of domestic dogs, commensal mice, and wild or early domestic goats, cats, and cattle during the eleventh millennium (late PPNA, EPPNB) and the subsequent addition of wild fallow deer, foxes, and domestic sheep at the beginning of the tenth millennium (MPPNB); (3) for each of the species in each of the different regions or even in each site, different scenarios and rates of intensification of exploitation existed, from control of wild/feral populations to sophisticated breeding that entailed the morphological changes that characterize domestic lineages; (4) the beginnings of domestication, that is, control in a wide area extending from the Iranian Plateau to the Mediterranean shores (though ancillary to the continent through the introduction of wild or domestic taxa, Cyprus was part of this area, with special Cypriot innovations such as intensive hunting of fallow deer and local domestication of the wild/feral goat); (5) intensive and long-distance exchanges of raw materials, including not only obsidian (Briois, Gratuze, and Guilaine 1997) but also living animals across large continental and marine areas (such interaction explains the relative homogeneity of the PPN cultural sphere); and (6) social organization and technical skills in the Near Eastern Final Late Glacial and Early Holocene societies that were more sophisticated than generally believed: in addition to rapid and high-level improvements in lithic (Astruc, Binder, and Briois 2007) and construction (Schmidt 2003; Stordeur et al. 2000) technologies, these groups were able to regularly and more and more intensively navigate to Cyprus (probably by sailing; Vigne 2009) and to control animal populations using diversified and sophisticated techniques (for meat, milk, hair, and draft products). Illustrating a part of the socioeconomic and historical complexity involved in the beginning of the Neolithic in the Near East, Cyprus also emphasizes the multifactorial causes of the transition, including, of course, climatic and environmental conditions (Bar-Yosef 2011) but also the dominant influence of demographic (Bocquet-Appel 2011), technical, and social threshold effects.

We thank the Wenner-Gren Foundation and the organizers of the Mérida conference, who invited J.-D. Vigne. We also thank all the archaeozoologists who provided us with often unpublished data: Paul Croft, Simon Davis, Sheelagh Frame, Lionel Gourichon, and Daniel Helmer. Elizabeth Willcox and Doug Price greatly improved our English-language writing. We are grateful to all the scientists and excavators who contributed to the study of this site. The French School at Athens, the Cyprus Department of Antiquities, and the French Ministry of Foreign Affairs supported our stays for fieldwork and analysis in Cyprus.


Jean-Denis Vigne is a senior researcher at the Centre National de la Recherche Scientifique (CNRS) and Director of Unité Mixte de Recherche 7209, “Archéozoologie, Archéobotanique: Sociétés, Pratiques et Environnements,” Muséum National d’Histoire Naturelle–CNRS (75005 Paris, France []). Isabelle Carrère and François Briois are researchers at Unité Mixte de Recherche 5608, “Travaux et Recherches Archéologiques sur les Cultures, les Espaces et les Sociétés” (TRACES), Ecole des Hautes Etudes en Sciences Sociales, Université Le Mirail (31000 Toulouse, France). Jean Guilaine is Professor Emeritus at the Collège de France (75005 Paris, France).

1. All the calibrated dates are given cal BP with 1σ, using Calib Rev 5.0 (Reimer et al. 2004).

2. The conceptual frame summarized in this paragraph has been strengthened, clarified, and here and there modified by discussions during the Mérida meeting “The Beginnings of Agriculture,” mainly among the archaeozoology working group: Fiona Marshall, Richard Meadow, Peter Rowley-Conwy, and Melinda Zeder.

References Cited

  • Ammerman, Albert J., Pavlos Flourentzos, R. Smadar Gabrieli, Thomas Higham, Carole McCartney, and Tim Turnbull. 2008. Third report on early sites on Cyprus. Report of the Department of Antiquities, Cyprus 2008:1–32.

  • Ammerman, Albert J., Pavlos Flourentzos, Carole McCartney, Jay S. Noller, and Daniel Sorabji. 2006. Two new early sites on Cyprus. Report of the Department of Antiquities, Cyprus 2007:1–22.

  • Ammerman, Albert J., and Jay S. Noller. 2005. New light on Aetokremnos. World Archaeology 37(4):533–543.

  • Astruc, Laurence, Didier Binder, and François Briois, eds. 2007. Systèmes techniques et communautés du Néolithique précéramique au Proche-Orient. Antibes, France: APDCA.

  • Auffray, Jean-Christophe, Eitan Tchernov, and Eviatar Nevo. 1988. Origine du commensalisme de la souris domestique (Mus musculus domesticus) vis-à-vis de l’homme. Comptes Rendus de l’Académie des Sciences de Paris sér. III: Sciences de la Vie 307(9):517–522.

  • Balasse, Muriel, and Stanley H. Ambrose. 2005. Distinguishing sheep and goats using dental morphology and stable carbon isotopes in C4 grassland environments. Journal of Archaeological Science 32(5):691–702.

  • Bar-Yosef, Ofer. 2011. Climatic fluctuations and early farming in West and East Asia. Current Anthropology 52(suppl. 4):S175–S193.

  • Bar-Yosef, Ofer, and Richard H. Meadow. 1995. The origin of agriculture in the Near East. In Last hunters, first farmers. T. Douglas Price and Anne Birgitte Gebauer, eds. Pp. 39–94. Santa Fe, NM: School of American Research Press.

  • Binford, Lewis. 2000. Review of Faunal extinctions in an island society: pygmy hippopotamus hunters of Cyprus, by Alan H. Simmons. American Antiquity 65(4):771.

  • Bocquet-Appel, Jean-Pierre. 2011. The agricultural demographic transition during and after the agriculture inventions. Current Anthropology 52(suppl. 4):S497–S510.

  • Boekschoten, G. J., and Paul Y. Sondaar. 1972. On the fossil mammalia of Cyprus. Proceedings of the Kononklijke Nederlandse Akademie van Wetenschappen B 75(4):306–308.

  • Briois, François 2003. Nature et évolution des industries lithiques de Shillourokambos. In Le Néolithique de Chypre: actes du colloque international organisé par le Département des Antiquités de Chypre et l’École Française d’Athènes, Nicosie 17–19 mai 2001. Jean Guilaine and Alain Le Brun, eds. Pp. 121–133. Bulletin de Correspondance Hellénique, supplément 43. Athens: École Française d’Athènes.

  • Briois, François, Bernard Gratuze, and Jean Guilaine. 1997. Obsidiennes du site néolithique précéramique de Shillourokambos (Chypre). Paléorient 23(1):95–112.

  • Briois, François, Catherine Petit-Aupert, and Pierre-Yves Péchoux. 2005. Histoire des campagnes d’Amathonte. I. L’occupation du sol au Néolithique. Études Chypriotes 16. Athens: École Française d’Athènes.

  • Bromage, Timothy G., Wendy Dirks, Hediye Erdjument-Bromage, Mathias Huck, Ottmar Kulmer, Rasime Öner, Oliver Sandrock, and Friedemann Schrenk. 2002. A life history and climate change solution to the evolution and extinction of insular dwarves: a Cypriot experience. In World islands in prehistory: international insular investigations. William Waldren and Josep Ensenyat, eds. Pp. 420–427. British Archaeological Reports, International Series 1095. Oxford: Archaeopress.

  • Broodbank, Cyprian. 2006. The origins and early development of Mediterranean maritime activity. Journal of Mediterranean Archaeology 19(2):199–230.

  • Bunimovitz, Shlomo, and Ran Barkai. 1996. Ancient bones and modern myths: ninth millennium BC hippopotamus hunters at Akrotiri Aetokremnos, Cyprus. Journal of Mediterranean Archaeology 9(1):85–96.

  • Cherry, John F. 1990. The first colonization of the Mediterranean islands: a review of recent research. Journal of Mediterranean Archaeology 3(2):145–221.

  • Clutton-Brock, Juliet. 1979. The mammalian remains from the Jericho Tell. Proceedings of the Prehistoric Society 45:135–157.

  • Conolly, James, Sue Colledge, Keith Dobney, Jean-Denis Vigne, Joris Peters, Barbara Stopp, Katie Manning, and Stephen Shennan. 2011. Meta-analysis of zooarchaeological data from SW Asia and SE Europe provides insight into the origins and spread of animal husbandry. Journal of Archaeological Science 38(3):538–545.

  • Cucchi, Thomas, Annie Orth, Jean-Christophe Auffray, Sabrina Renaud, Laurent Fabre, Josette Catalan, Eleftherios Hadjisterkotis, François Bonhomme, and Jean-Denis Vigne. 2006. A new endemic species of the subgenus Mus (Rodentia, Mammalia) on the Island of Cyprus. Zootaxa 1241:1–36.

  • Cucchi, Thomas, and Jean-Denis Vigne. 2006. Origin and diffusion of the house mouse in the Mediterranean. Human Evolution 21(2):95–106.

  • Cucchi, Thomas, Jean-Denis Vigne, Jean-Christophe Auffray, Paul Croft, and Edgar Peltenburg. 2002. Introduction involontaire de la souris domestique (Mus musculus domesticus) à Chypre dès le Néolithique précéramique ancien (fin IXe et VIIIe millénaires av. J.-C.). Comptes Rendus de l’Académie des Sciences, Palévol 1(4):235–241.

  • Davis, Simon J. M. 1994. Even more bones from Khirokitia: the 1988–1991 excavations. In Fouilles récentes à Khirokitia (Chypre) 1988–1991. Alain Le Brun, ed. Pp. 305–334. Éditions Recherches sur les Civilisations. Paris: ADPF.

  • ———. 2003. The zooarchaeology of Khirokitia (Neolithic Cyprus), including a view from the mainland. In Le Néolithique de Chypre: actes du colloque international organisé par le Département des Antiquités de Chypre et l’École Française d’Athènes, Nicosie 17–19 mai 2001. Jean Guilaine and Alain Le Brun, eds. Pp. 253–278. Bulletin de Correspondance Hellénique, supplément 43. Athens: École Française d’Athènes.

  • ———. 2005. Why domesticate food animals? some zoo-archaeological evidence from the Levant. Journal of Archaeological Science 32(9):1408–1416.

  • Desse, Jean, and Nathalie Desse-Berset. 2003. Les premiers pêcheurs de Chypre. In Le Néolithique de Chypre: actes du colloque international organisé par le Département des Antiquités de Chypre et l’École Française d’Athènes, Nicosie 17–19 mai 2001. Jean Guilaine and Alain Le Brun, eds. Pp. 279–291. Bulletin de Correspondance Hellénique, supplément 43. Athens: École Française d’Athènes.

  • Dobney, Keith, and Greger Larson. 2006. Genetics and animal domestication: new windows on an elusive process. Journal of Zoology 269(2):261–271.

  • Ducos, Pierre. 1968. L’origine des animaux domestique en Palestine. Bordeaux: Institut de Préhistoire de l’Université.

  • Ervynck, Anton, Keith Dobney, Hitomi Hongo, and Richard Meadow. 2001. Born free? new evidence for the status of pigs from Çayönü Tepesi, eastern Anatolia. Paléorient 27(2):47–73.

  • Goring-Morris, A. Nigel, and Anna Belfer-Cohen. 2011. Neolithization processes in the Levant: the outer envelope. Current Anthropology 52(suppl. 4):S195–S208.

  • Gourichon, Lionel, and Daniel Helmer. 2008. Etude archéozoologique de Mureybet. In Le site néolithique de Tell Mureybet (Syrie du Nord): en hommage à Jacques Cauvin. Juan José Ibañez, ed. Pp. 115–227. British Archaeological Reports International Series 1843. Oxford: Archaeopress.

  • Guilaine, Jean. 2003. Parekklisha-Shillourokambos. Périodisation et aménagements domestiques. In Le Néolithique de Chypre: actes du colloque international organisé par le Département des Antiquités de Chypre et l’École Française d’Athènes, Nicosie 17–19 mai 2001. Jean Guilaine and Alain Le Brun, eds. Pp. 3–14. Bulletin de Correspondance Hellénique, supplément 43. Athens: École Française d’Athènes.

  • Guilaine, Jean, and François Briois. 2007. Shillourokambos and the Neolithization of Cyprus: some reflections. Eurasian Prehistory 4(1–2):159–175.

  • Guilaine, Jean, François Briois, and Jean-Denis Vigne, eds. 2011. Shillourokambos: un établissement néolithique pré-céramique à Chypre: les fouilles du secteur 1. Paris: Errance–École Française d’Athènes.

  • Guilaine, Jean, François Briois, Jean-Denis Vigne, and Isabelle Carrère. 2000. Découverte d’un Néolithique précéramique ancien chypriote (fin 9e début 8e millénaires cal. BC), apparenté au PPNB ancien/moyen du Levant nord. Comptes Rendus de l’Académie des Sciences, sér. IIA: Sciences de la Terre et des Planètes 330(1):75–82.

  • Guilaine, Jean, François Briois, Jean-Denis Vigne, Thomas Perrin, Yann Béliez, Isabelle Carrère, Patrice Gérard, and Maxime Rémicourt. 2008. Shillourokambos (Paerekklisha): l’établissement néolithique précéramique: études et rapports 2.1. Bulletin de Correspondance Hellénique 128–129(2):1006–1021.

  • Guilaine, Jean, Philippe Devèze, Jacques Coularou, and François Briois. 1999. Tête sculptée en pierre dans le Néolithique précéramique de Shillourokambos (Parekklisha. Chypre). Report of the Department of Antiquities, Cyprus 1999:1–12.

  • Guilaine, Jean, and Alain Le Brun, eds. 2003. Le Néolithique de Chypre: actes du colloque international organisé par le Département des Antiquités de Chypre et l’École Française d’Athènes, Nicosie 17–19 mai 2001. Bulletin de Correspondance Hellénique, supplément 43. Athens: École Française d’Athènes.

  • Held, Steven O. 1989. Colonization cycles on Cyprus. 1. The biogeographic and paleontological foundations of early prehistoric settlement. Report of the Department of Antiquities, Cyprus 1989:7–28.

  • Helmer, Daniel. 1992. La domestication des animaux par l’homme préhistorique. Paris: Masson.

  • ———. 2008. Révision de la faune de Cafer Höyük (Malatya, Turquie): apports des méthodes de l’analyse des mélanges et de l’analyse de Kernel à la mise en évidence de la domestication. In Archaeozoology of the Near East VIII: proceedings of the 8th International Symposium on the Archaeozoology of Southwestern Asia and Adjacent Areas. Emannuelle Vila, Lionel Gourichon, Alice M. Choyke, and Hijlke Buitenhuis, eds. Pp. 169–195. Travaux de la Maison de l’Orient 49. Lyon: Maison de l’Orient et de la Méditerranée.

  • Helmer, Daniel, and Lionel Gourichon. 2008. Premières données sur les modalités de subsistance dans les niveaux récents (PPNB moyen à Néolithique à Poterie) de Tell Aswad en Damascène (Syrie): fouilles 2001–2005. In Archaeozoology of the Near East VIII: proceedings of the 8th International Symposium on the Archaeozoology of Southwestern Asia and Adjacent Areas. Emannuelle Vila, Lionel Gourichon, Alice M. Choyke, and Hijlke Buitenhuis, eds. Pp. 120–151. Travaux de la Maison de l’Orient 49. Lyon: Maison de l’Orient et de la Méditerranée.

  • Helmer, Daniel, Lionel Gourichon, Hervé Monchot, Joris Peters, and Maria Saña Segui. 2005. Identifying early domestic cattle from Pre-Pottery Neolithic sites on the Middle Euphrates using sexual dimorphism. In The first steps of animal domestication: new archaeobiological approaches. Jean-Denis Vigne, Joris Peters, and Daniel Helmer, eds. Pp. 86–95. London: Oxbow.

  • Helmer, Daniel, Lionel Gourichon, and Danielle Stordeur. 2004. À l’aube de la domestication animale: imaginaire et symbolisme animal dans les premières sociétés néolithiques du nord du Proche-Orient. Anthropozoologica 39(1):143–163.

  • Helmer, Daniel, Lionel Gourichon, and Emmanuelle Vila. 2007. The development of the exploitation of products from Capra and Ovis (meat, milk and fleece) from the PPNB to the Early Bronze in the northern Near East (8700 to 2000 BC cal.). Anthropozoologica 42(2):41–69.

  • Hole, Frank. 1996. The context of caprine domestication in the Zagros region. In The origins and spread of agriculture and pastoralism in Eurasia. David R. Harris, ed. Pp. 263–281. Washington, DC: Smithsonian Institution.

  • Hongo, Hitomi, Jessica Pearson, Banu Öksüz, and Gülçin İlgezdi. 2009. The process of ungulate domestication at Çayönü, southeastern Turkey: a multidisciplinary approach focusing on Bos sp. and Cervus elaphus. Anthropozoologica 44(1):63–73.

  • Horwitz, Liora K., Eitan Tchernov, and Hitomi Hongo. 2004. The domestic status of the Early Neolithic fauna of Cyprus: a view from the mainland. In Neolithic revolution: new perspectives on Southwest Asia in light of recent discoveries on Cyprus. Edgar Peltenburg and Alexander Wasse, eds. Pp. 35–48. Levant Supplementary Series 1. Oxford: Oxbow.

  • Hours, Francis, Olivier Aurenche, Jacques Cauvin, Marie-Claire Cauvin, Lorraine Copeland, and Paul Sanlaville. 1994. Atlas des sites du Proche-Orient (ASPO): 1400–5700 BP. Travaux de la Maison de l’Orient 24. Lyon: Maison de l’Orient Méditerranéen.

  • Khalaily, Hamudi, Ofer Marder, and Omry Barzilai. 2007. An early Pre-Pottery Neolithic B blade cache from Motza, west of Jerusalem, Israel. In Systèmes techniques et communautés du Néolithique précéramique au Proche-Orient. Laurence Astruc, Didier Binder, and François Briois, eds. Pp. 269–276. Antibes, France: APDCA.

  • Kuijt, Ian. 2008. Demography and storage systems during the southern Levantine Neolithic demographic transition. In The Neolithic demographic transition and its consequences. Jean-Pierre Bocquet-Appel and Ofer Bar-Yosef, eds. Pp. 287–313. New York: Springer.

  • Kuijt, Ian, and Bill Finlayson. 2009. Evidence for food storage and predomestication granaries 11,000 years ago in the Jordan Valley. Proceedings of the National Academy of Sciences of the USA 106(27):10966–10970.

  • Le Brun, Alain, and Odile Daune-Le Brun. 2003. Deux aspects du Néolithique pré-céramique récent de Chypre: Khirokitia et Cap Andreas-Kastros. In Le Néolithique de Chypre: actes du colloque international organisé par le Département des Antiquités de Chypre et l’École Française d’Athènes, Nicosie 17–19 mai 2001. Jean Guilaine and Alain Le Brun, eds. Pp. 45–59. Bulletin de Correspondance Hellénique, supplément 43. Athens: École Française d’Athènes.

  • Legge, Anthony J. 1996. The beginning of caprine domestication in Southwest Asia. In The origins and spread of agriculture and pastoralism in Eurasia. David R. Harris, ed. Pp. 238–262. Washington, DC: Smithsonian Institution.

  • Lévi-Strauss, Claude. 1958. Anthropologie structurale. Agora 7. Paris: Plon.

  • Malek, Jaromir. 1993. The cat in ancient Egypt. London: British Museum.

  • Manning, Sturt W., Carole McCartney, Bernd Kromer, and Sarah T. Stewart. 2010. The earlier Neolithic in Cyprus: recognition and dating of a Pre-Pottery Neolithic A occupation. Antiquity 84(325):693–706.

  • Marom, Nimrod, and Guy Bar-Oz. 2009. Culling profiles: the indeterminacy of archaeozoological data to survivorship curve modelling of sheep and goat herd maintenance strategies. Journal of Archaeological Science 36(5):1184–1187.

  • Mauss, Marcel. 1947 (1967). Manuel d’ethnographie. Petite Bibliothèque Payot. Lausanne, Switzerland: Payot.

  • McCartney, Carole. 2010. Outside the corridor? the Neolithisation of Cyprus. In The development of pre-state communities in the ancient Near East: studies in honour of Edgar Petenburg. Diane Bolger and Louise Maguire, eds. Pp. 185–196. British Association for Near Eastern Archaeology Publication Series 2. Oxford: Oxbow.

  • McCartney, Carole, Sturt W. Manning, Sandra Rosendahl, and Sarah T. Stewart. 2008. Elaborating Early Neolithic Cyprus (EENC): preliminary report on the 2007 field season: excavations and regional field survey at Agia Varvara-Asprokremnos. Report of the Department of Antiquities, Cyprus 2008:67–86.

  • McCartney, Carole, Sturt W. Manning, David Sewell, and Sarah T. Stewart. 2007. The EENC 2006 field season: excavations at Agia Varvara-Asprokremnos and survey of the local Early Holocene landscape. Report of the Department of Antiquities, Cyprus 2007:27–44.

  • ———. 2010. Reconsidering Early Holocene Cyprus within the eastern Mediterranean landscape. In Landscapes in transition. Bill Finlayson and Graeme Warren, eds. Pp. 133–146. Levant Supplementary Series 8. Oxford: Oxbow.

  • Monchot, Hervé, and Jacques Léchelle. 2002. Statistical nonparametrics methods for the study of fossil populations. Paleobiology 28(1):55–69.

  • Muxart, Tatiana, Franck-Dominique Vivien, Bruno Villalba, and Joëlle Burnouf, eds. 2003. Des milieux et des hommes: fragments d’histoires croisées. Paris: Elsevier.

  • Naderi, Saeid, Hamid-Reza Rezaei, François Pompanon, Michael G. B. Blum, Riccardo Negrini, Hamid-Reza Naghash, Özge Balkız, et al. 2008. The goat domestication process inferred from large-scale mitochondrial DNA analysis of wild and domestic individuals. Proceedings of the National Academy of Sciences of the USA 105(46):17659–17664.

  • Olsen, Sandra L. 1999. Investigation of the Phanourios bones for evidence of cultural modification. In Faunal extinction in an island society: pygmy hippopotamus hunters of Cyprus. Alan H. Simmons, ed. Pp. 230–238. New York: Kluwer Academic/Plenum.

  • Pascal, Michel, Olivier Lorvelec, and Jean-Denis Vigne. 2006. Invasions biologiques et extinctions: 11,000 ans d’histoire des vertébrés en France. Paris: Belin.

  • Peltenburg, Edgar. 2003. The colonisation and settlement of Cyprus: investigations at Kissonerga-Mylouthkia (1976–1996). Studies in Mediterranean Archaeology. Sävedalen, Sweden: Aströms.

  • Peltenburg, Edgar, Sue Colledge, Paul Croft, Adam Jackson, Carole McCartney, and Mary Anne Murray. 2000. Agro-pastoralist colonization of Cyprus in the 10th millennium BP: initial assessments. Antiquity 74(286):844–853.

  • ———. 2001a. Neolithic dispersals from the Levantine corridor: a Mediterranean perspective. Levant 33:35–64.

  • ———. 2001b. Well-established colonists: Mylouthkia 1 and the cypro-pottery Neolithic B. In The earliest prehistory of Cyprus: from colonization to exploitation. Stuart Swiny, ed. Pp. 61–93. Boston: American Schools of Oriental Research.

  • Peltenburg, Edgar, and Alexander Wasse, eds. 2004. Neolithic Revolution: new perspectives on Southwest Asia in light of recent discoveries on Cyprus. Levant Supplementary Series 1. Oxford: Oxbow.

  • Pergams, Oliver R. W., and Mary V. Ashley. 2001. Microevolution in island rodents. Genetica 112–113:245–256.

  • Perrin, Thomas. 2011. Les données du macro-outillage. In Shillourokambos: un établissement néolithique pré-céramique à Chypre: les fouilles du secteur 1. Jean Guilaine, François Briois, and Jean-Denis Vigne, eds. Paris: Errance–École Française d’Athènes.

  • Peters Joris, Angela von den Driesch, and Daniel Helmer. 2005. The upper Euphrates-Tigris basin: cradle of agro-pastoralism? In The first steps of animal domestication: new archaeobiological approaches. Jean-Denis Vigne, Joris Peters, and Daniel Helmer, eds. Pp. 96–124. Oxford: Oxbow.

  • Redding, Richard W. 2005. Breaking the mold: a consideration of variation in the evolution of animal domestication. In The first steps of animal domestication: new archaeobiological approaches. Jean-Denis Vigne, Joris Peters, and Daniel Helmer, eds. Pp. 41–48. Oxford: Oxbow.

  • Redding, Richard W., and Michael Rosenberg. 1998. Ancestral pigs: a New (Guinea) model for pig domestication in the Middle East. In Ancestors for the pigs: pigs in prehistory. Sarah M. Nelson, ed. Pp. 65–76. MASCA Research Papers in Science and Archaeology 15. Philadelphia: University of Pennsylvania Museum.

  • Reimer, Paula J., Mike G. L. Baillie, Édouard Bard, Alex Bayliss, J. Warren Beck, Chanda J. H. Bertrand, Paul G. Blackwell, et al. 2004. Intcal04 terrestrial radiocarbon age calibration, 0–26 cal kyr BP. Radiocarbon 46(3):1029–1058.

  • Rosenberg, Michael, R. Mark Nesbitt, Richard W. Redding, and Brian L. Peasnall. 1998. Hallan Çemi, pig husbandry, and post-Pleistocene adaptations along the Taurus-Zagros arc (Turkey). Paléorient 24(1):25–41.

  • Rosenberg, Michael, and Richard Redding. 1998. Early pig husbandry in Southwestern Asia and its implications for modeling the origins of food production. In Ancestors for the pigs: pigs in prehistory. Sarah M. Nelson, ed. Pp. 55–64. MASCA Research Papers in Science and Archaeology 15. Philadelphia: University of Pennsylvania Museum.

  • Saña Segui, Maria. 1999. Arqueología de la domesticacíon animal: la gestión de los recursos animales en Tell Halula (Valle del Éufrates, Siria) de 8.800 al 7.000 BP. Treballs d’Arqueologia del Pròxim Orient 1. Barcelona: Universitat Autònoma de Barcelona.

  • Schmidt, Klaus. 2003. “Kraniche am See”: Bilder und Zeichen vom frühneolithischen Göbekli Tepe (Südosttürkei). In Der Turmbau zu Babel: Ursprung und Vielfalt von Sprache und Schrift. Wilfried Seipel, ed. Pp. 23–29. Vienna: Kunsthistorischen Museums.

  • Serrand, Nathalie, Jean-Denis Vigne, and Jean Guilaine. 2005. Early Preceramic Neolithic marine shells from Shillourokambos, Cyprus (late 9th–8th mill. cal BC): a mainly ornamental set with similarities to mainlands PPNB. In Archaeomalacology: molluscs in former environments of human behaviour. Daniella Bar-Yosef Mayer, ed. Pp. 122–129. Oxford: Oxbow.

  • Simmons, Alan H. 1988. Extinct pygmy hippopotamus and early man in Cyprus. Nature 333(6173):554–557.

  • ———, ed. 1999. Faunal extinction in an island society: pygmy hippopotamus hunters of Cyprus. New York: Kluwer Academic/Plenum.

  • Simmons, Alan H., and Rolfe Mandel. 2007. Not such a new light: a response to Ammerman and Noller. World Archaeology 39(4):475–482.

  • Smith, Bruce D. 2001. Low level food production. Journal of Archaeological Research 9(1):1–43.

  • Starkovich, Britt M., and Mary C. Stiner. 2009. Hallan Çemi Tepesi: high-ranked game exploitation alongside intensive seed processing at the Epipaleolithic-Neolithic transition in southeastern Turkey. Anthropozoologica 44(1):41–61.

  • Stordeur, Danielle, Michel Brenet, Gérard Der Aprahamian, and Jean-Claude Roux. 2000. Les bâtiments communautaires de Jerf el Ahmar et Mureybet, Horizon PPNA, Syrie. Paléorient 26(1):29–44.

  • Tchernov, Eitan, and François R. Valla. 1997. Two new dogs and other Natufian dogs from the southern Levant. Journal of Archaeological Science 24(1):65–95.

  • Trantalidou, Katerina. 2008. Glimpses of Aegean Island communities during the Mesolithic and Neolithic Periods: the zooarchaeological point of view. In A colloquium on the prehistory of the Cyclades. Neil Brodie, Jennifer Doole, Giorgos Gavalas, and Colin Renfrew, eds. Pp. 19–27. Cambridge: McDonald Institute.

  • Vigne, Jean-Denis. 1992. The meat and offal weight (MOW) method and the relative proportion of ovicaprines in some ancient meat diets of the north-western Mediterranean. Rivista di Studi Liguri A 57(2):21–47.

  • ———. 1998. Faciès culturels et sous-système technique de l’acquisition des ressources animales: application au Néolithique ancien méditerranéen. In Production et identité culturelle: actualité de la recherche. André D’Anna and Didier Binder, eds. Pp. 27–45. Antibes, France: APDCA.

  • ———. 1999. The large “true” Mediterranean as a model for the Holocene human impact on the European vertebrate fauna? recent data and new reflections. In The Holocene history of the European vertebrate fauna: modern aspects of research. Norbert Benecke, ed. Pp. 295–322. Archäologie in Eurasien 6. Berlin: Deutsches Archäologisches Institut.

  • ———. 2000. Outils pour restituer les stratégies de chasse au cerf en Europe au Mésolithique et au Néolithique: analyses graphiques, statistiques et multivariées de courbes d’âges d’abattage. Ibex Journal of Mountain Ecology 5/Anthropozoologica 31:57–67.

  • ———. 2006. Découpe du cerf (Cervus elaphus) au Mésolithique moyen, à Noyen-sur-Seine (Seine-et-Marne): analyses tracéologique et expérimentale. Revue de Paléobiologie vol. spécial 10:69–82.

  • ———. 2008. Zooarchaeological aspects of the Neolithic diet transition in the Near East and Europe, and their putative relationships with the Neolithic demographic transition. In The Neolithic demographic transition and its consequences. Jean-Pierre Bocquet Appel and Ofer Bar-Yosef, eds. Pp. 179–205. New York: Springer.

  • ———. 2009. Introductions et réintroduction de mammifères à Chypre aux IXe et VIIIe millénaires av. J.-C. (Néolithique précéramique): indices indirects de l’usage de la voile au Néolithique? In De Méditerranée et d’ailleurs…: mélanges offerts à Jean Guilaine. Pp. 807–820. Toulouse, France: Archives d’Écologie Préhistorique.

  • ———. 2011. The origins of animal domestication and husbandry: a major change in the history of humanity and the biosphere. Comptes Rendus de l’Académie des Sciences, Biologie. 334(3):171–181.

  • Vigne, Jean-Denis, François Briois, Antoine Zazzo, Isabelle Carrère, Julie Daujat, and Jean Guilaine. Forthcoming. Preliminary data on a new Early Pre-Pottery Neolithic site on Cyprus (Ayios Tychonas, Klimonas, ca. 9000 cal. BC). Paléorient.

  • Vigne, Jean-Denis, Isabelle Carrère, and Jean Guilaine. 2003. Unstable status of early domestic ungulates in the Near East: the example of Shillourokambos (Cyprus. IX–VIIIth millennia cal. B.C.). In Le Néolithique de Chypre: actes du colloque international organisé par le Département des Antiquités de Chypre et l’École Française d’Athènes, Nicosie 17–19 mai 2001. Jean Guilaine and Alain Le Brun, eds. Pp. 239–251. Bulletin de Correspondance Hellénique, supplément 43. Athens: École Française d’Athènes.

  • ———. Forthcoming. Evolution of the use of sheep and goat during the Pre-Pottery Neolithic at Shillourokambos (8400–7000 cal. BC). In Archaeozoology of the Near East IX: Proceedings of the 9th International Symposium on the Archaeozoology of Southwestern Asia and Adjacent Areas. Marjan Mashkour and Mark Beech, eds. Oxford: Oxbow.

  • Vigne, Jean-Denis, Isabelle Carrère, Jean-François Saliège, Alain Person, Hervé Bocherens, Jean Guilaine, and François Briois. 2000. Predomestic cattle, sheep, goat and pig during the late 9th and the 8th millennium cal. BC on Cyprus: preliminary results of Shillourokambos (Perkklisha, Limassol). In Archaeozoology of the Near East IV: proceedings of the 4th International Symposium on the Archaeozoology of Southwestern Asia and Adjacent Areas. Marjan Mashkour, Alice M. Choyke, Hijlke Buitenhuis, and François Poplin, eds. Pp. 52–75. Publicaties 32. Groningen: Centre for Archaeological Research and Consultancy.

  • Vigne, Jean-Denis, and Thomas Cucchi. 2005. Premières navigations au Proche-Orient: les informations indirectes de Chypre. Paléorient 31(1):186–194.

  • Vigne, Jean-Denis, and Jean Guilaine. 2004. Les premiers animaux de compagnie 8500 ans avant notre ère? … ou comment j’ai mangé mon chat, mon chien et mon renard. Anthropozoologica 39(1):249–273.

  • Vigne, Jean-Denis, Jean Guilaine, Karyne Debue, Laurent Haye, and Patrice Gérard. 2004. Early taming of the cat in Cyprus. Science 304(5668):259.

  • Vigne, Jean-Denis, and Daniel Helmer. 2007. Was milk a “secondary product” in the Old World Neolithisation process? its role in the domestication of cattle, sheep and goats. Anthropozoologica 42(2):9–40.

  • Vigne, Jean-Denis, Antoine Zazzo, Jean-François Saliège, François Poplin, Jean Guilaine, and Alan Simmons. 2009. Pre-Neolithic wild boar management and introduction to Cyprus more than 11,400 years ago. Proceedings of the National Academy of Sciences of the USA 106(38):16131–16138.

  • Wasse, Alexander. 2007. Climate, economy and change: Cyprus and the Levant during the Late Pleistocene to mid-Holocene. In On the margins of Southwest Asia: Cyprus during the 6th to 4th millennia BC. Joanne Clarke, ed. Pp. 44–63. Oxford: Oxbow.

  • Willcox, George. 2003. The origins of Cypriot farming. In Le Néolithique de Chypre: actes du colloque international organisé par le Département des Antiquités de Chypre et l’École Française d’Athènes, Nicosie 17–19 mai 2001. Jean Guilaine and Alain Le Brun, eds. Pp. 231–238. Bulletin de Correspondance Hellénique, supplément 43. Athens: École Française d’Athènes.

  • Zeder, Melinda A. 2005. A view from the Zagros: new perspectives on livestock domestication in the Fertile Crescent. In The first steps of animal domestication: new archaeobiological approaches. Jean-Denis Vigne, Joris Peters, and Daniel Helmer, eds. Pp. 125–146. Oxford: Oxbow.

  • ———. 2006. Central questions in the domestication of plants and animals. Evolutionary Anthropology 15(3):105–117.

  • ———. 2009. The Neolithic macro-(r)evolution: macroevolutionary theory and the study of culture change. Journal of Archaeological Research 17(1):1–63.

  • ———. 2011. The origins of agriculture in the Near East. Current Anthropology 52(suppl. 4):S221–S235.

  • Zeder, Melinda A., Eve Emshwiller, Bruce D. Smith, and Daniel G. Bradley. 2006. Documenting domestication: the intersection of genetics and archaeology. Trends in Genetics 22(3):139–155.

  • Zeder, Melinda A., and Brian Hesse. 2000. The initial domestication of goats (Capra hircus) in the Zagros mountains 10.000 years ago. Science 287(5461):2254–2257.