Initial diversity in sheep and goat management in Neolithic south-western Asia

Initial diversity in sheep and goatmanagement in Neolithic south-western Asia

Benjamin S. Arbuckle and Levent Atici

In this paper we survey a large body of faunal data for the practice of young male culling in

Neolithic south-western Asia. Although the young male kill-off model is one of the most widely

used models for identifying animal domestication in Neolithic south-western Asia, its ubiquity has

never been addressed on a regional scale. By focusing on a combination of kill-off age and the

shape of the distributions of biometric data, we are able to address the emergence and ubiquity

of young male culling amongst Neolithic sheep and goat herders. Although the intensive culling of

young males has been presented as a ‘leading edge marker’ for the initiation of sheep and goat

herding, we find that clear evidence for young male kill-off appears in the faunal record only in the

early 8th millennium cal BC — considerably later than the origins of caprine management.

Instead, Neolithic caprine management practices appear to have been characterized by a high

degree of ‘initial diversity’, especially in the 9th and early 8th millennia, suggesting that early

management strategies may have been much more varied than previously realized. However,

after c. 7500 cal BC young male kill-off was widely practised across south-western Asia,

suggesting this efficient and effective management technology quickly replaced the diversity of

local management strategies prevalent earlier.

Keywords: Neolithic, sheep, goats, animal domestication, young male culling, management

Introduction

The origins of domesticated plants and animals

represents one of the most consequential technologi-

cal transformations in human history and has there-

fore been a major focus of archaeological research for

more than a century (Binford 1968; Childe 1936;

Pumpelly 1908). Beginning with the observation that

domesticates exhibit a distinctive set of phenotypes,

often including smaller size, compared to their wild

ancestors, early researchers focused on using skeletal

measurements to identify the origins of domestic

animals (Baumler 1921; Hammer 1984; Uerpmann

1979; Winge 1900). By the 1970s, a series of more

comprehensive and varied lines of evidence for

identifying the process of animal domestication

was developed, including the use of demographic

profiles to interpret culling practices associated with

intentional human management (Bokonyi 1969;

Ducos 1978; Hesse 1978; Meadow 1989). Recently,

increasingly sophisticated methods have been added

to the arsenal that archaeologists have at their disposal

to address the origins of animal husbandry, including

isotopic and phytolith evidence for foddering and

manipulation of herd mobility, geochemical evidence

for penning and the use of animal dung, morphometric

approaches to population dispersals, as well as the

results of paleogenetic studies (Ludwig et al. 2009;

Makarewicz and Tuross 2012; Meiggs 2010; Naderi et al.

2008; Ottoni et al. 2012; Pearson et al. 2007).

Together, the application of these methods has

resulted in tremendous progress mapping out the

broad chronological and spatial patterns for the

origins and spread of domestic animals (Peters et al.

2013; Vigne et al. 2011; Zeder 2008b). Despite this

progress, however, the early stages of the develop-

ment and spread of herding economies are still poorly

understood, and basic questions such as ‘what

husbandry methods did the earliest herders use’ and

‘how did herd management technologies spread’ are

Benjamin S. Arbuckle (corresponding author), Department ofAnthropology, University of North Carolina at Chapel Hill, CB#3115, 301Alumni Building, Chapel Hill, NC 27599-3115, USA; email: [email protected]. Levent Atici, Department of Anthropology, University ofNevada, Las Vegas, NV 89154-5003, USA; email: levent.atici@unlv.

� Council for British Research in the Levant 2013Published by ManeyDOI 10.1179/0075891413Z.00000000026 Levant 2013 VOL 45 NO 2 219

yet to be answered. This is partly the result of

limitations in the application of faunal data and

traditional lines of evidence to address the nature of

the earliest animal management strategies. For

example, researchers have shown that changes in

phenotype, including decrease in size and transfor-

mation of horn morphology in domestic bovids,

likely occurred centuries after the initiation of

intensive management regimes (i.e. herding), thus

making it difficult to use these phenotypic markers to

identify the earliest phases in the establishment of

animal husbandry practices (Zeder 2006). In addi-

tion, interpretations of demographic evidence for

herd management are often hindered by widespread

variations in the methods, localities and seasonality

of hunting practices, which may target a range of

demographic cohorts especially in wild bovids

characterized by dramatic seasonal changes in the

territoriality and demographic composition of social

groups (Schaller 1977; Simmons and Ilany 1975–77;

Watson 1978). The common, and often necessary,

analytical practice of combining faunal data into

broad multi-component assemblages, often creates

unintelligible palimpsests of evidence for animal

exploitation, thus making interpretation difficult

(Halstead 1998). Compounding these problems is

the fact that faunal researchers have been slow to

apply new methods which target evidence for specific

husbandry practices including foddering, stalling and

herd mobility, in favour of approaches seeking to

broadly characterize the ‘status’ of animal populations

as wild or domestic (although see Henton 2012;

Makarewicz and Tuross 2012; Vanpoucke et al. 2009).

Some of the most productive recent research

documenting the origins of animal management has

focused on sheep and goats, which may have been the

earliest domesticated food animals (Peters et al. 2005).

Current research suggests that these animals were

brought under human control in a process that began

sometime in the 9th millennium cal BC in a region

extending from south-eastern Turkey to north-western

Iran (Zeder 2011). One of the most effective methods

for documenting this initiation of animal husbandry has

focused on identifying a combination of sex and age

specific culling practices known as young male kill-off.

Payne (1973) has defined the most widely used

model describing young male kill-off as a typical herd

management strategy used by sheep and goat herders.

In this model, surplus immature males are the

primary target of slaughter, while females are

generally culled as adults when their reproductive

potential begins to decline. As a result of this

management system, faunal assemblages exhibit clear

biometric and demographic patterns. Because goats

and, to a lesser extent, sheep are characterized by

sexual dimorphism, and because males achieve most of

their larger body size within the first year (Davis 2000;

Zeder and Hesse 2000), young male kill-off produces

diagnostic patterns in which most immature specimens

represent large males, while most adult specimens

represent smaller females. These patterns can be

identified through the analysis of breadth and depth

measurements of postcranial skeletal elements which

bear evidence for epiphyseal fusion thus providing

evidence for both the age and sex of culled individuals.

Zeder and Hesse (2000; Zeder 2008b) have

effectively used this combined biometric-demo-

graphic approach to identify the earliest management

of morphologically wild goats in western Iran in the

early 8th millennium BC at the site of Ganj Dareh.

Expanding the use of this approach, Zeder has

succeeded in mapping out the spread of goat and

also sheep management practices across the Zagros

region. As a result of this innovative and detailed

work, it is argued that young male kill-off, identified

through this combined biometric-demographic

approach, is the best ‘leading edge marker’ for the

origins of animal management (Zeder 2006).

Curiously, and despite the effectiveness of this

approach, it has rarely been applied outside the

Zagros region (although see Arbuckle et al. 2009;

Makarewicz 2009), and there has never been a

comprehensive test for the timing and ubiquity of

evidence for young male kill-off in the archaeofaunal

record of Neolithic south-western Asia. As a result, it

is currently not clear how ubiquitous young male kill-

off was as a method for managing Neolithic herds of

sheep and goat or when and where it first emerged as

a management strategy.

In this paper, we provide the first regional scale test

for evidence for the practice of young male kill-off in

Neolithic south-western Asia. We focus on a survey

of published biometric and demographic data from

78 assemblages from across the region, particularly

those dating to the 10th through 8th millennia BC

(PPNA and PPNB periods in the Levantine chron-

ological framework), with the goal of identifying

when and where the practice of young male kill-off is

evident and characterizing its ubiquity as an early

strategy of herd management (Fig. 1; see Tables 1

and 2 for lists of assemblages).

Methods: identifying young male kill-off in thearchaeofaunal record

In order to identify evidence for the management

strategy of young male kill-off, we focus on published

Arbuckle and Atici Diversity in sheep and goat management

220 Levant 2013 VOL 45 NO 2

biometric data and, secondarily, age data for both

sheep and goats. Biometric data are ideal for identify-

ing evidence for young male kill-off in caprines for

several reasons. First, they have the advantage that

they are published for a large number of Neolithic

assemblages using a reliable and standardized format

either as raw measurements (following von den

Driesch 1976) or transformed using the Log Size

Index (LSI) method (Meadow 1999).

Secondly, in sexually dimorphic taxa such as goats

and, to a lesser extent, sheep, males and females can

be distinguished based on the breadth and depth

measurements of long bones (Davis 2000; Zeder and

Hesse 2000). For goats (Capra aegagrus/hircus),

which exhibit a relatively high degree of dimorphism,

it is often possible to attribute most measurements of

long bone epiphyses to a particular sex, and

researchers have used several statistical methods

including Mixture analysis (Monchot 1999;

Monchot and Lechelle 2002) and the Mahalanobis

distance between known male and female samples

(Zeder 2001) to achieve this result with high

confidence intervals.

Sheep (Ovis orientalis/aries) exhibit a lesser degree

of sexual dimorphism than do goats (Davis 2000;

Vigne 2011b). There is a greater degree of overlap in

the size of males and females in bone measurements

and it is therefore more difficult to attribute

individual measurements to a particular sex.

However, dimorphism in some measurements includ-

ing the distal breadth of the metacarpus (also scapula

SLC, tibia Bd, metatarsal Bd, radius Bp, humerus

BT, calcaneum GL) is significant within both

domestic (t-test; p,0?001; N526) (Davis 2000) and

wild populations (t-test; p50?004; N526; based on

recent specimens from Iran curated in the Field

Museum, Chicago) (Fig. 2). Although the degree of

dimorphism in sheep is limited, it is possible to

generate general estimates of the proportions of

males and females in archaeofaunal populations by

examining the shape (skewness and kurtosis) of the

distribution produced by biometric data utilizing

Figure 1 Map showing the location of sites mentioned in the text


Levant 2013 VOL 45 NO 2 221

Table 1 List of Neolithic assemblages used to address sheep (Ovis orientalis/aries) exploitation. Site names in boldfaceinterpreted as representing sheep husbandry (based on original author). Summary of skewness (positive,negative, normal, etc.) refers to the shape of the distribution of biometric data. Frequency of juveniles based inmost cases on fusion of the distal epiphysis of metapodials or tooth wear data using Payne’s (1973) categoriesA–D. * indicates percentage of juveniles calculated for combined sheep/goat. Presence or absence of evidencefor young male kill-off (** indicates cases of negative skewing interpreted as young male kill-off) and evidencefor phenotypic changes associated with the domestication process are also indicated

Site Date (cal BC) Skewness %juv Young male killoff? Morph dom.? Reference

Karain B 17,000 neg 20* No No (Atici 2009)Okuzini 1 17,000 NA 25* No No (Atici 2009)Okuzini 2 16,000 pos 26* No? No (Atici 2009)Okuzini 3–4 13,000 neg 27* No No (Atici 2009)Okuzini 5 12,000 pos 22* No? No (Atici 2009)Hallan Cemi 10,000 normal 25 No No (Starkovich and Stiner 2009)ZC Shanidar 10,000 normal 50 No No (Perkins 1973; Zeder 2008a)Palegawra 10,000 neg 25 No No (Zeder 2008a)Mureybet IB–III 9500–8400 pos 23 No No (Gourichon and Helmer 2008)Jerf el-Ahmar 9000 pos nd No? No (Vigne 2011b)Gobekli 9000 neg 40* No No (Peters et al. 2013; Vigne 2011b)Kortik Tepe 9000 neg 50 No No (Arbuckle and Ozkaya 2007)Asiab 9000 neg 33 No No (Zeder 2008a)Cayonu RG 9000–8400 neg 40 No No (Hongo et al. 2005)Cayonu CH 8400–8200 neg 50 No Yes? (Hongo et al. 2005)Cayonu CP 8200–7600 normal 32 No Yes? (Hongo et al. 2005)Nevalı Cori 8500–7600 normal c. 50* No Yes? (Peters et al. 2013)Cafer 8300–7500 neg 43* No No (Helmer 2008)Asıklı 2G 8000 neg 49* No No (Peters et al. 2013)Asıklı 2E 7900 normal 49* No No (Peters et al. 2013)Ganj Dareh 7900 neg 25 No No (Hesse 1978; Zeder 2008a)Aswad moyenne 7800 pos 48 Yes Yes? (Helmer and Gourichon 2008)Cayonu Cell 7600–7500 pos nd Yes? Yes? (Hongo et al. 2005)Abu Hureyra 2A 7500 normal 42* No? Yes? (Legge and Rowley-Conwy 2000)Aswad recente 7500 pos 48 Yes Yes? (Helmer and Gourichon 2008)Shillourokambos anc. C 7600–7500 pos 42 Yes Yes? (Vigne 2011b)Cayonu LR 7500–6900 pos 32 Yes Yes (Hongo et al. 2005)Halula moyenne 7500 pos 67 Yes Yes? (Vigne 2011b)Ali Kosh 7500–6000 neg 23 No No? (Zeder 2008a)Suberde 7500–7000 pos 40 No? No (Arbuckle 2008b)Gritille 7500–7000 pos 40* Yes Yes (Monahan 2000)Teleilat MPPNB 7500 pos 48* Yes? Yes (Ilgezdi 2008)Shillourokambos moy. A1 7500 pos 50 Yes Yes? (Vigne 2011b)Shillourokambos moy. A2 7400 pos 56 Yes Yes? (Vigne 2011b)Hayaz 7400 pos nd Yes? Yes (Peters et al. 2013)Asıklı 2B 7400 normal 49* No No (Peters et al. 2013)Catalhoyuk pre-XII 7400–7000 normal 50 No Yes (Russell and Martin 2005)Abu Hureyra 2B 7400–7100 pos 35* Yes? Yes? (Legge and Rowley-Conwy 2000)Shillourokambos moy. B 7400–7200 pos 30 Yes Yes? (Vigne 2011b)Jarmo 7300–6300 pos nd Yes? Yes (Zeder 2008a)Halula recente 7300 normal 68 No Yes (Vigne 2011b)Ras Shamra Vc 7300 bimodal 42* No Yes (Vigne et al. 2003)Shillourokambos Rec. 7200–7000 pos 19 Yes Yes? (Vigne 2011b)Ain Ghazal PPNC 7000 neg 40* No Yes (Wasse 2002)Ramad I 7000 pos nd Yes? Yes (Ducos 1993)Ghoraife 7000 normal 70 No Yes (Ducos 1993)Ras Shamra Va 7000 bimodal 38* No Yes (Vigne et al. 2003)Tepe Sarab 7000 pos nd Yes? Yes (Zeder 2008a)Gurcutepe 7000 pos 46 Yes Yes (Peters et al. 2013)Teleilat LPPNB 7000 pos 89* Yes Yes (Ilgezdi 2008)Ain Jammam 7000 pos 25 Yes Yes (Makarewicz 2009)Ba’ja 7000 neg nd No Yes (Makarewicz 2009)Ulucak VI 7000–6000 normal 55 No? Yes (Cakırlar 2012)Choga Sefid 7000–5000 pos nd Yes Yes (Zeder 2008a)Bademagacı 7000–6500 pos 31 Yes Yes (De Cupere et al. 2008)Catalhoyuk XII–VII 7000–6800 pos 50 Yes Yes (Russell and Martin 2005)Ramad II 6900–6500 neg nd Yes** Yes (Ducos 1993)Cayonu PN 6900–6300 pos 50 Yes Yes (Hongo et al. 2005)Catalhoyuk VI 6500 pos 60 Yes Yes (Russell and Martin 2005)Ulucak V 6500–6000 pos 78 Yes? Yes (Cakırlar 2012)Erbaba 6500–6000 neg 45 No? Yes (Arbuckle 2008a)Hoyucek 6400–6000 pos 41 Yes Yes (De Cupere and Duru 2003)



relatively large samples of measurements (Monchot

1999; Vigne 2011b). Although it is preferable to use

measurements from a single skeletal part (e.g.

metacarpal Bd) for this purpose, log transformed

LSI data are also effective in cases where sample sizes

are limited. Recently, both Vigne (2011b) and Zeder

(2008a) have successfully used this method to

interpret Neolithic sheep exploitation practices in

the Neolithic Near East (also see Marom and Bar-Oz

2013; Wolverton 2008).

Finally, postcranial depth and breadth measure-

ments from long bones with epiphyses record

evidence of both the size (and therefore sex) and

age of culled individuals. Since the ages of fusion for

sheep and goat epiphyses are known (e.g. Habermehl

1985; Silver 1963), the fusion status of skeletal

elements can be used to identify individuals as either

older or younger than the age range at which fusion

takes place for a particular skeletal part. Since

caprines achieve most of their skeletal growth by c.

1–2 years, even unfused specimens (especially for

later fusion skeletal parts) can effectively be used to

distinguish between males and females (Davis 2000;

Zeder 2001). Thus, by combining evidence for

epiphyseal fusion with biometric evidence for sex

ratios, we can attempt to identify assemblages that

exhibit the characteristic pattern of large-sized,

unfused specimens and, especially, the small-sized

fused specimens typically produced by the practice of

young male kill-off.

Several strategies were utilized to accommodate the

fact that this study focuses on identifying young male

kill-off through published biometric data. First,

because the samples of measurements from individual

sites identified as either sheep or goat are often

limited, we use the LSI method to combine and

standardize measurements from multiple skeletal

elements (Meadow 1999). Although allometric differ-

ences between archaeological populations and the

standard animal used to generate LSI values can

create variation which potentially blurs differences

between the sexes (Arbuckle and Makarewicz 2009;

Russell et al. 2005), previous work suggests that for

both sheep and goats LSI transformed data still

exhibit clear signals of sexual dimorphism (Vigne

2011b). Second, since specimens with unfused epi-

physes were rarely measured before Zeder’s work, we

focus on biometric data from fused specimens, which

nonetheless provide ample evidence for young male

kill-off (fused specimens will be dominated by small

females when young male kill-off is practised). In

addition, whenever possible, later fusing skeletal parts

which exhibit the highest degree of sexual dimorphism

(e.g. metacarpal Bd, metatarsal Bd, calcaneum GL,

tibia Bd) (based on Davis 2000 for sheep, and Zeder

2001 for goats) were preferentially selected for

analysis, although in some cases (due to small sample

sizes) all available measurements were utilized.

In order to identify young male kill-off, histograms

of LSI values were generated for sheep and for goats

for each of 42 archaeofaunal assemblages. The shape

of the distribution of each of these histograms was

then described in terms of its skewness and kurtosis.

Skewness describes the asymmetry of the distribution

of a variable, with positive skewing representing a

curve with a tail extending further to the right of the

mean, and negative skewing a curve with a tail

extending further to the left of the mean (Moore and

McCabe 2001). A skewness value of ‘0’ represents a

relatively symmetrical distribution. Kurtosis, on the

other hand, refers to the height of a distribution with

positive values representing a tall, peaked curve while

negative values indicate a low peak in the distribution

(Moore and McCabe 2001).

The practice of young male kill-off is expected to

result in diagnostic patterns in biometric data. These

patterns are represented in Figure 3 based on distal

breadth measurements for fused metacarpals for

goats from Ganj Dareh — the site with which this

model has been most clearly associated. This

characteristic distribution exhibits strong positive

skewing and positive kurtosis. The peak on the left

Site Date (cal BC) Skewness %juv Young male killoff? Morph dom.? Reference

Qdeir 6500 normal 49* No Yes (Helmer 1992)Umm El Tlel 6500 neg 60* No Yes (Helmer 1992)El Kowm 2 6500 pos 40* Yes Yes (Helmer 1992)Ain Ghazal Yarmoukian 6500 pos 46* Yes Yes (Wasse 2002)Bouqras 6500 neg nd No Yes (Wasse 2002)Khirokitia III–I 6000 normal nd No No? (Vigne 2011b)Kosk Hoyuk V 6000 pos 61* Yes Yes (Arbuckle 2006)Banahilk 6000–5000 pos nd Yes Yes (Zeder 2008a)Ilipinar X–IX 6000–5800 pos 35 Yes Yes (Buitenhuis 2008)

Table 1 Continued



Table 2 List of Neolithic assemblages used to address goat (including Capra aegagrus, C. hircus and C. ibex)exploitation. Site names in boldface interpreted as representing sheep husbandry (based on original author).Summary of skewness (positive, negative, normal, etc.) refers to the shape of the distribution of biometric data.Frequency of juveniles based in most cases on fusion of the distal epiphysis of metapodials or tooth wear datausing Payne’s (1973) categories A–D. * indicates percentage of juveniles calculated for combined sheep/goat.Presence or absence of evidence for young male kill-off (** indicates cases of negative skewing interpreted asyoung male kill-off) and evidence for phenotypic changes associated with the domestication process are alsoindicated

SiteDate(cal BC) Skewness %juv

Young malekilloff? Morph dom? Reference

Shanidar Mousterian 50,000 normal c. 10 No No (Zeder 2008a)Karain B 19,000 pos 20 No No (Atici 2009)Ksar Akil 19,000 neg 30 No No (Kersten 1987)Ucagızlı Epi 14,000 pos nd No No (Acıkkol 2006)Okuzini 5 12,000 pos 22 No No (Atici 2009)ZC Shanidar 10,000 normal 25 No No (Perkins 1964; Zeder 2008a)Es Saaide II 10,000 pos nd No No (Churcher 1994)Ain Mallaha 10,000 neg nd No No (Wasse 2002)Direkli 9000 neg 50 No No (Arbuckle and Erek 2010)Asiab 9000 neg 43 No No (Zeder 2008a)Cayonu RG 9000–8400 neg 25 No No (Hongo et al. 2005)Cayonu CH 8400–8200 normal 65 No No (Hongo et al. 2005)Cafer 8300–7500 neg 55* No No (Helmer 2008)Cayonu CP 8200–7600 pos 42 Yes Yes? (Hongo et al. 2005)Asıklı 2 8000–7500 neg 49* No No (Buitenhuis pers. comm.)Shillourokambos anc. B 8000–7600 bimodal 16 No No? (Vigne 2011a)Ujret el Mehed (ibex) 8000 pos 50 No No (Dayan et al. 1986)Wadi Tbeik (ibex) 8000 neg nd No No (Tchernov and Bar-Yosef 1982)Abu Gosh 8000–7500 neg 40 No No (Horwitz 2003)Ganj Dareh 8000 pos 66 Yes No (Hesse 1978; Zeder 2008a)Beidha 8000–7500 pos 60 Yes? No? (Hecker 1975)Aswad moy. 7800 pos 52 Yes Yes? (Helmer and Gourichon 2008)Cayonu Cell 7600–7500 pos nd Yes Yes (Hongo et al. 2005)Gritille 7500–7000 pos 40* Yes Yes (Monahan 2000)Teleilat M–LPPNB 7500–7000 pos 75* Yes Yes (Ilgezdi 2008)Abu Hureyra 2A 7500 pos 42 Yes? No? (Legge and Rowley-Conwy 2000)Ali Kosh BM 7500 pos 55 Yes Yes (Hole et al. 1969; Zeder 2008a)Shillourokambos moy. A1 7500 bimodal 41 No? No? (Vigne 2011a)Ain Ghazal MPPNB 7500 pos 29* Yes No? (Wasse 2002)Aswad recente 7500 bimodal 58 No Yes (Helmer and Gourichon 2008)Suberde 7500–7000 normal nd No No? (Arbuckle 2008b)Cayonu LR 7500–6900 pos 10 Yes? Yes (Hongo et al. 2005)Abu Hureyra 2B 7400–7100 pos 35 Yes Yes? (Legge and Rowley-Conwy 2000)Shillourokambos moy. A2 7400 pos 46 Yes No? (Vigne 2011a)Shillourokambos moy. B 7400–7200 pos 5 Yes Yes? (Vigne 2011a)Ali Kosh AK 7400–7000 pos 41 Yes Yes (Hole et al. 1969; Zeder 2008a)Catalhoyuk East 7400–6500 neg c.50* Yes** Yes (Russell and Martin 2005)Jarmo 7300–6300 pos c.16 Yes Yes (Zeder 2008a)Shillourokambos Rec. 7200–7000 pos nd Yes Yes? (Vigne 2011a)Ali Kosk MJ 7000–6000 pos 31 Yes Yes (Hole et al. 1969; Zeder 2008a)Bademagacı 7000–6500 pos 31 Yes Yes (De Cupere et al. 2008)Ulucak VI 7000–6500 pos 33 Yes Yes (Cakırlar 2012)Ramad I 7000 pos nd Yes Yes (Ducos 1993)Tepe Sarab 7000 pos nd Yes Yes (Zeder 2008a)Choga Sefid 7000–5000 pos nd Yes Yes (Zeder 2008a)Sarab 7000 pos nd Yes Yes (Zeder 2008a)Ghoraife 7000 bimodal 65 No Yes (Ducos 1995)Gurcutepe 7000 pos 39 Yes Yes (Peters et al. 2013)Ain Ghazal PPNC 7000 pos 40* Yes Yes? (Wasse 2002)Ain Jammam 7000 pos 40 Yes Yes (Makarewicz 2009)Ba’ja 7000 pos nd Yes Yes (Makarewicz 2009)Cayonu PN 6900–6300 pos 50 Yes Yes (Hongo et al. 2005)Ain Ghazal Yarmoukian 6500 normal 46* No Yes (Wasse 2002)Erbaba 6500–6000 pos 31 Yes? Yes (Arbuckle 2008a)Aswad PN 6500 pos 45 Yes Yes (Helmer and Gourichon 2008)Ras Shamra VC1 6500 normal 42* No Yes (Helmer 1989)Assouad 6500 neg nd No Yes (Helmer 1989)Qdeir 6500 pos 49* Yes Yes (Helmer 1989)Ulucak V 6500–6000 pos 80 Yes Yes (Cakırlar 2012)Hoyucek 6400–6000 neg 41 Yes** Yes (De Cupere and Duru 2003)



represents small adult females, which dominate the

assemblage, while the tail on the right represents a

small number of males allowed to survive past the age

of metacarpal fusion (c. 18–24 months). The esti-

mated distributions of females and males in the

population are further indicated in Figure 3 based on

the results of mixture analysis (PAST software),

emphasizing the highly biased sex ratio among

skeletally mature individuals reflected in this assem-

blage. This pattern, combined with age data indicat-

ing a young kill-off in which most goats (66%) were

culled by 24 months, clearly reflects a management

strategy in which young surplus males are targeted

for slaughter and females are allowed to survive into

adulthood (Zeder 2008a). This pattern is thought to

represent the archetypal management system used by

sheep and goat pastoralists (Payne 1973) and is

unlikely to be produced by hunting strategies.

Although hunting wild sheep and goats may

potentially result in a wide variety of demographic

patterns, a frequently reoccurring example of the

distribution of biometric data is presented in Figure 4.

This classic hunting model is based on measurements

of wild sheep from Ganj Dareh and exhibits strongly

negative skewing and negative kurtosis. The peak on

the right represents large adult males, which dominate

the assemblage, while the tail to the left of the peak

represents a smaller numbers of adult females. Mixture

analysis suggests that this pattern reflects a significant

sex bias with males dominating the population

(Fig. 4). In addition, the slaughter age for this

assemblage is quite high, with only 25% of sheep

slaughtered before 24 months, which is a typical,

although not universal, feature of demographic

profiles produced by hunting (Zeder 2008a).

Results

With these two models in mind, skewness and

kurtosis values were plotted for 42 Neolithic assem-

blages for sheep and goats in Figure 5. In this figure,

the NW and SW quadrants represent assemblages

exhibiting negative skewing. In accordance with the

model presented in Figure 5, many of the assem-

blages in these quadrants represent caprine exploita-

tion strategies interpreted as hunting. The majority of

these assemblages exhibit a combination of mild

negative skewing and negative kurtosis as exemplified

by the sheep data from Ganj Dareh, Gobekli Tepe,

Okuzini 3, and goats from Ksar Akil. Only three

examples fall in the NW quadrant including the

assemblages from Asiab and Zawi Chemi Shanidar

which are thought to represent sheep hunting. The

goats from Catalhoyuk, which represent morpholo-

gically domesticated animals, also fall in this quad-

rant but this is likely the result of the near absence of

large males represented in the biometric sample used

in this study (i.e. the tail to the right is missing).

Interestingly, in the SW quadrant, in addition to

assemblages representing wild caprine hunting, there

are also several Neolithic assemblages thought to

represent herding economies including the sheep and

goats from Cafer, goats from Suberde and

Khirokitia, and sheep from PPNC Ain Ghazal

(Arbuckle 2008b; Helmer 2008; Wasse 2002).

Figure 2 Measurements of the distal breadth of the metacarpal for male (black) and female (grey) recent wild sheep (Ovis

orientalis) from Iran (Field Museum, Chicago) showing clear evidence for sexual dimorphism. Curves produced

using mixture analysis accurately predicted both male and female distributions (meanmale528?7; meanfemale526?7;

mixture analysis meanmale529?3; mixture analysis meanfemale526?7)

SiteDate(cal BC) Skewness %juv

Young malekilloff? Morph dom? Reference

Banahilk 6000 pos nd Yes Yes (Zeder 2008a)Khirokitia I–III 6000 pos nd Yes? Yes? (Vigne et al. 2003)Kosk Hoyuk V 6000 pos 61* Yes Yes (Arbuckle 2006)Ilipinar IX 5800 pos 35 Yes Yes (Buitenhuis 2008)

Table 2 Continued



The NE and SE quadrants represent assemblages

with positive skewing. The NE quadrant holds the

clearest examples of young male kill-off including the

Ganj Dareh goats, which represent the earliest

evidence for goat husbandry in the Zagros. Both of

these assemblages, as well as the sheep from the 8th

and 7th millennia BC sites Gritille and Hoyucek, and

the goats from Bademagacı, are characterized by a

combination of strongly positive skewing and kurto-

sis reflecting significant sex biases in the biometric

data. Neolithic assemblages exhibiting slightly lower

levels of skewness include sheep from Aswad (phase

Recente), sheep and goats from several phases of

Shillourokambos on Cyprus, as well as Yarmoukian

Ain Ghazal, and goats from Tepe Sarab,

Bademagacı, and Erbaba. However, sheep and goat

hunting also produced similarly shaped, positively

skewed curves as evident from assemblages including

the goats from Mousterian Shanidar, Epipaleolithic

Ucagızlı Cave, Asiab, and sheep from Okuzini 2,

Hallan Cemi, and Mureybet where there is no clear

evidence for animal husbandry.

The SE quadrant, characterized by mild positive

skewing but a less ‘peaked’ distribution (negative

kurtosis), includes a dense cluster of assemblages.

These include those produced by hunting wild sheep

and goats such as Okuzini 5, Karain B, Direkli Cave,

Douara Cave, Kortik Tepe, Ujret el Mehed (C. ibex),

Jebel Es-Saiide II and Jerf el-Ahmar, and also those

thought to represent some of the earliest sheep

herding in the Near East including Nevalı Cori,

Aswad-Moyenne, Halula-Moyenne, and also

Suberde, Shillourokambos and Basta. Many of these

latter assemblages exhibit only very mild positive

skewness (i.e. they are symmetrical) and do not

provide clear evidence for young male kill-off.

It is possible to distinguish between assemblages,

especially those exhibiting positive skewing, produced

by hunting versus husbandry by plotting skewness

values along with the frequencies of juveniles in each

assemblage. Figure 6 compares these variables, and

shows that sheep and goat assemblages representing

early herding economies tend to exhibit positive

skewness values combined with high frequencies of

juveniles and cluster in the upper right portion of the

graph, while those representing hunting tend to fall to

the bottom and left portions of the graph. The upper

right portion of Figure 6 (shaded grey), therefore,

Figure 3 Young male kill-offmodel. Distribution of measure-ments of the distal breadth ofthe metacarpal for early domes-tic goats from the site GanjDareh, Iran (Hesse 1978).Curves estimating contributionof males and females generatedwith mixture analysis (usingPAST)

Figure 4 Hunting model.Distribution of measure-ments of the greatest lengthof the first phalanx (ante-riorzposterior) of wildsheep from the site GanjDareh, Iran (Hesse 1978).Curves estimating contribu-tion of males and femalesgenerated with mixture ana-lysis (using PAST)



provides a fairly clear signature for identifying the

practice of young male kill-off. There are, however, a

few interesting exceptions to this general pattern

including assemblages thought to represent hunting

that fall within or on the margins of this area. These

include the goats from Asiab, Direkli Cave, ibex from

Ujret El-Mehed, and sheep from Kortik Tepe. There

are also a number of assemblages which are thought

to represent herding practices but which do not show

clear evidence for young male kill-off including the

sheep from Nevalı Cori, Suberde, Cafer, Halula

Recente, PPNC Ain Ghazal and the goats from

Cafer and Hoyucek. In addition, the goats from Ganj

Dareh, which have been used as a model representing

early herd management, appear to be an outlier with

higher skewness and juvenile values than any other

Neolithic dataset.

Although raw biometric data were not available to

calculate skewness and kurtosis from many Neolithic

assemblages, the shapes of published distributions of

biometric data not presented in Figures 5 and 6 could

often be characterized in terms of positive or negative

skewness and whether they represent young male kill-

off or not (see Tables 1 and 2). For sheep, a survey of

this larger sample of assemblages showed no clear

evidence for young male kill-off based on a combina-

tion of biometric distributions and culling age prior

to c. 8000 BC, despite some evidence for the

emergence of sheep management by this time

(Peters et al. 2005; Vigne 2011a; Zeder 2008b).

The sequence at Cayonu Tepesi, located in the

Tigris drainage in south-eastern Turkey, spans the

period of sheep domestication and the faunal

assemblage has been the target of intensive scrutiny

Figure 5 Representation of skewness and kurtosis for biometric data for sheep (black squares) and goats (open

squares). Curves representing biometric distributions are shown for one typical assemblage in each quadrant.

Site abbreviations: CAF 5 Cafer Hoyuk; GD5Ganj Dareh; Catal5Catalhoyuk; SUB5Suberde; ERB5Erbaba

Hoyuk; KSAR5Ksar Akil; OK1–55Okuzini Cave; KAB5Karain Cave; Gob5Gobekli Tepe; AG-C5Ain Ghazal

PPNC; AG-Y5Ain Ghazal Yarmoukian; ZCS5Zawi Chemi Shanidar; HaLR5Halula Recente; HaLM5Halula

Moyenne; KOR5Kortik Tepe; KSK5Kosk Hoyuk; BAS5Basta; DIR5Direkli Cave; UJM5Ujret El-Mehed;

JeS5Jebel Es Saaide II; ASM5Aswad Moyenne; ASR5Aswad Recente; ShAB5Shillourokambos Ancien B;

ShAC5Shillourokambos Ancien C; ShM25Shillourokambos Moyenne B2; ShM15Shillourokambos Moyenne B1;

ShR5Shillourokambos Recente; HC5Hallan Cemi; BAD5Bademagacı; Sarab5Tepe Sarab; UCA5Ucagızlı Cave;

HOY5Hoyucek; MUR5Mureybet; GRI5Gritille; Khr5Khirokitia; NC5Nevalı Cori; Jerf5Jerf El Ahmar;

DOC5Douara Cave; SHAN-M5Shanidar Mousterian



(Hongo et al. 2005). Here, morphologically wild

sheep were hunted in small numbers in the early and

mid-9th millennium BC, producing biometric dis-

tributions exhibiting negative skewing suggesting an

emphasis on hunting large adult males. By the late

9th millennium, smaller-sized individuals, probably

representing domesticates, begin to appear at Cayonu

but the biometric distribution is symmetrical with no

indication of young male kill-off. It is not until the

Cell Room phase, dating from c. 7500 BC, that

positively skewed biometric data along with evidence

for changes in phenotype suggest that domestic sheep

were subject to young male kill-off at Cayonu.

In the upper Euphrates basin at Nevalı Cori, Peters

(Losch et al. 2006; Peters et al. 2005) has argued for

the earliest appearance of sheep husbandry, including

evidence for phenotypic changes associated with

domestication and isotopic evidence for foddering.

However, in this case the distribution of sheep

measurements is notably symmetrical with negative

kurtosis providing no clear evidence for young male

kill-off in the late 9th and early 8th millennia BC

(Peters et al. 2013). At Cafer Hoyuk, in the highlands

of eastern Anatolia, Helmer (2008) has argued for a

system of sheep husbandry combined with mouflon

hunting in the late 9th and early 8th millennia BC.

However, the sheep at Cafer seem to retain a largely

wild phenotype and biometric data exhibit negative

skewing, suggesting that large adult males were

frequently exploited providing no evidence for

the culling of young males. At Asıklı Hoyuk, in

central Anatolia, where sheep are the dominant

taxon, biometric data from levels 2G to 2B,

representing the early to late 8th millennium BC,

show either negative skewing or no skewing at all.

Despite convincing arguments for ‘proto-domestica-

tion’ management of sheep at Asıklı (Buitenhuis

1997; Matthews 1998) the biometric patterns do not

fit with expectations for young male kill-off. The

earliest available evidence for young male kill-off for

sheep comes from the northern margin of the

southern Levant at Tell Aswad, where by the early

8th millennium BC both morphological domesticates

and positively skewed biometric data suggest sheep

herds were subject to young male kill-off (Helmer and

Gourichon 2008).

Figure 6 Plot showing skewness and frequency of juveniles for Neolithic sheep (black squares) and goat (grey squares)

assemblages. Frequency of juveniles based in most cases on either fusion of metapodials or tooth wear data

using Payne’s (1973) categories A–D. Shaded grey area represents typical pattern for young male kill-off. Site

codes are the same as in Figure 5



Biometric evidence for young male kill-off becomes

much more common after 7500 BC across south-

western Asia. Out of the assemblages surveyed, 65%

(31/48) reflect fairly clear evidence for young male

kill-off compared to only 13% from earlier Neolithic

assemblages (2/15). In south-eastern Turkey, assem-

blages from Gritille and Teleilet both show evidence

for young male kill-off in the mid- to late 8th

millennium, and this management strategy is also

evident from contemporary levels at Halula and Abu

Hureyra 2B on the Middle Euphrates. At the site of

Shillourokambos, on the island of Cyprus, Vigne

(2011a; Vigne et al. 2011) has argued that domestic

sheep were introduced in small numbers from the

mainland by c. 8000 BC and positive skewing

combined with the culling of juveniles suggests that

young male kill-off was practised by 7500 BC.

Finally, in the Zagros region, central/western

Anatolia, and the southern Levant positive skewing

and young male kill-off are evident at Tepe Sarab,

Pottery Neolithic levels of Catalhoyuk, Bademagacı,

Ulucak V and Ain Jammam in the early 7th

millennium cal BC.

However, young male kill-off is not ubiquitous

even among later Neolithic sites with strong evidence

for sheep herding. For example, at Abu Hureyra on

the Middle Euphrates, domestic sheep are thought to

have appeared in level 2A (c. 7500 BC) but biometric

evidence for young male kill-off is only evident in

level 2B dating to the late 8th millennium (Legge and

Rowley-Conwy 2000). In central Anatolia, domestic

sheep are present at Catalhoyuk from the earliest

levels of the site, c. 7400 BC. However, the distribu-

tion of biometric data from these early levels is

symmetrical (Russell and Martin 2005); it is only in

the early-7th-millennium levels that the shape of the

biometric distribution suggests young male culling at

Catalhoyuk. Moreover, in western and south-western

Anatolia, neither Suberde, Erbaba, nor the earliest

levels of Ulucak, exhibit clear evidence for young

male kill-off. At Suberde skewing of fused measure-

ments is only weakly positive and measurements of

unfused specimens do not suggest that young males

were preferentially culled. At Erbaba, although sheep

with domestic phenotypes are present in this late-7th-

millennium assemblage, biometric data exhibit nega-

tive skewing and large adult males are well represented

suggesting a mixed exploitation system perhaps similar

to that described for Cafer. In the southern Levant,

where domestic sheep were imported in the late 8th

millennium (Horwitz et al. 1999), several assemblages

show, either bimodality, no skewing, or negative

skewing indicating a surprising degree of variability

in evidence for sheep management at sites including

Ras Shamra, Ghoraife, PPNC Ain Ghazal and on the

island of Cyprus at Khirokitia.

As was the case with sheep, evidence for young

male kill-off of goats is limited before c. 8000 BC

(Table 2). Biometric and survivorship evidence from

Cayonu suggest that wild males were targeted by

hunting parties in the 9th millennium BC, with a shift

towards positive skewing and morphological domes-

ticates suggesting the initiation of young male culling

in the Cobble Paved phase dating to the late 9th and

early 8th millennia BC (Hongo et al. 2005). These

patterns become clearer in the following Cell phase

dating to the mid-8th millennium. In addition, at

Cafer, Helmer (2008) has argued that goats were both

herded and hunted in the late 9th and early 8th

millennia, although morphological changes are not

evident and the biometric data exhibit strongly

negative skewing. The same pattern is evident in

central Anatolia at Asıklı Hoyuk despite arguments

for management in the early 8th millennium.

Although goats were imported to the island of

Cyprus by c. 8300 BC, Vigne (2011a) has argued

that they were fundamentally a wild resource and

were hunted. This is confirmed by the strongly

bimodal shape of biometric data from the early levels

of Shillourokambos (Ancienne B–C) and low fre-

quencies of juveniles.

The earliest and most precisely dated evidence for

the practice of culling young male goats comes from

Ganj Dareh in western Iran where it can be closely

dated to the first centuries of the 8th millennium BC

(Zeder and Hesse 2000). In the far southern Levant,

the assemblage from Beidha shows some potentially

early evidence for young male culling including a high

frequency of juveniles and positive skewing of

biometric data, although the chronology of the

assemblage is not well understood and the presence

of both goats and ibex complicate the interpretation

of biometric patterns (Hecker 1975). Roughly con-

temporaneous with Ganj Dareh, the Middle PPNB

levels of Tell Aswad provide the earliest evidence for

young male culling combined with morphological

changes in the goat population (Helmer and

Gourichon 2008). This is significant in that it

indicates that this management strategy was utilized

on opposite sides of the Fertile Crescent by the early

8th millennium.

As was the case with sheep, evidence for culling

young male goats is common in Neolithic assem-

blages across south-western Asia after 7500 BC,

where it is identified in 84% (37/44) of the assem-

blages examined in this study compared to only 35%



(5/14) of those dating to the earlier Neolithic. In the

late 8th millennium, positive skewing, high frequen-

cies of juveniles and morphological domesticates

indicate young male kill-off was practised along the

Middle Euphrates at Abu Hureyra 2A, at Gritille and

Teleilat in south-eastern Turkey, at Ali Kosh in

western Iran, at Catalhoyuk in central Anatolia,

Ulucak in western Anatolia, in southern Jordan at

Ain Jammam and Ba’ja, and it makes its appearance

on Cyprus at Shillourokambos (Moyenne A.2) by

7400 BC. However, culling young male goats is not

evident in all later Neolithic assemblages, including

Suberde in west central Anatolia; Ghoraife, Aswad

Recente and Tell Assouadin, Syria; and Yarmoukian

Ain Ghazal and Ras Shamra in the southern Levant.

Discussion

The results of this broad survey of faunal evidence for

early sheep and goat management in south-western

Asia emphasize several important features of Neolithic

animal economies. First, we are able to address the

timing of the appearance of young male kill-off as a

major herd management strategy. This is important

both for understanding the evolution of animal

husbandry, but also because Zeder’s detailed analysis

of goat exploitation in the Zagros region has shown

that young male kill-off preceded the appearance of

morphological changes and is, therefore, presented as

the best ‘leading edge marker’ for the earliest stages of

the domestication process (Zeder 2006).

The data examined here suggest that the practice of

young male kill-off was applied to goat herds at

Cayonu in the upper Tigris basin and at Ganj Dareh

in western Iran by c. 8000 BC, and to sheep herds a

few centuries later at Cayonu, Tell Aswad in the

Damascus Basin, and Shillourokambos on Cyprus.

However, these early-8th-millennium communities,

which are spread over a wide geographic area, seem

to be isolated outposts in their choice of herd

management strategies since young male culling is

not evident at contemporaneous sites in the region.

In addition, although young male culling clearly

precedes the appearance of phenotypically domestic

goats in the Zagros region, in the upper Euphrates

region the opposite pattern is evident. Here, at Nevalı

Cori, Peters (Peters et al. 2013) has argued that

phenotypic changes (size decrease) and isotopic

evidence for foddering suggest that sheep and,

perhaps, goat herding was well underway as early

as the late 9th millennium BC. Further upstream,

Helmer (2008) suggests that caprine management

may have been practised at Cafer Hoyuk from the

late 9th and early 8th millennia. However, there is no

evidence to suggest that young males were targeted

for preferential culling at either site. In fact, evidence

for young male culling is not present in the Euphrates

basin until the mid- to late 8th millennium at Teleilat,

Gritille and Gurcutepe.

Thus in the Euphrates basin, young male kill-off is

evident well after the appearance of evidence for herd

management, including phenotypic changes, and

seems not to be the leading edge marker for the

initiation of caprine management as it was for the

Zagros region. The reasons for this reversal are

unclear. It is possible that it is the result of biases in

the archaeofaunal record. For example, it is possible

that the delay in the appearance of young male

culling is the result of small sample sizes from the

earliest phases of important sites, including Aswad

(ancien), Cafer (Early phase), and Shillourokambos

(ancien A), or the common practice of ‘lumping’

faunal data from multiple archaeological phases into

one analytical unit which might obscure patterns in

the data.

However, it is also possible that this pattern

represents a different reality in the Euphrates basin,

where populations of managed caprines in the 9th

and early 8th millennia were very small, and

represented only a minor component of the animal

economy compared to the Zagros where they were

the dominant taxa (Arbuckle 2012; Legge 1996;

Vigne and Helmer 2007). In these cases where local

populations were small, access to wild caprines was

limited, and husbandry likely played as much a social

as subsistence role (Russell 2012), it may not have

made economic (or social) sense for herders to cull

large numbers of young animals. However, these

small, isolated populations of managed caprines may

have developed phenotypic changes associated with

the domestication process more rapidly than their

managed neighbours in uplands regions, which likely

maintained gene flow with wild populations.

This apparent delay in the application of young

male culling in the Euphrates basin may, therefore,

reflect differences in the historical trajectories of

caprine management systems in lowland regions

where gazelle and equids were long the focus of

animal economies, versus upland regions within the

natural habitat of the wild sheep and goats where

caprine management likely first emerged. However,

even in the upland regions of Anatolia where caprines

were heavily exploited (e.g. Asıklı, Cafer), the data

still do not suggest a pattern like that seen in the

Zagros. Here, neither evidence for young male culling

nor phenotypic changes are evident in the late 9th or

early 8th millennia BC.



The second significant finding of this study relates

to the ubiquity of young male culling as a Neolithic

pastoral management strategy. In the 9th and early

8th millennia BC, sheep and goat management

strategies seem to have been very diverse with only

a minority of communities intensively culling young

males. Young male kill-off is only evident in 30% (7/

23) of the sheep and goat assemblages dating to the

9th and early 8th millennia — it was, therefore, far

from ubiquitous as a management strategy in the

early Neolithic.

However, a major change took place in the mid-7th

millennium BC, after which young male kill-off

became the dominant pastoral management strategy

across south-western Asia. Among sheep and goat

assemblages dating between 7500–6000 BC, clear

evidence for young male kill-off was recognized in

almost 75% (62/84) of cases. That young male kill-off

shifted from an experimental management strategy

used in only a handful of communities to become the

dominant mode of herd management in the mid-8th

millennium BC, is no accident. This period represents

the major inflection point in the evolution of

Neolithic animal economies across south-western

Asia. The frequencies of sheep and goats — which

now exhibit domestic phenotypes — increase drama-

tically at this time, becoming the mainstay of animal

economies across the region, with domesticates

appearing outside of their natural habitat including

goats on the middle Euphrates and sheep in the

southern Levant (Arbuckle 2012; Legge 1996; Vigne

and Helmer 2007; Zeder 2011). This new emphasis on

herding domestic caprines seems to represent the

widespread adoption of intensive and large-scale

mixed-caprine pastoralism throughout much of the

Near East. It was in these new economies of large-

scale sheep and goat pastoralism, combined with

mature agriculture systems (Asouti and Fuller 2012),

that young male kill-off emerged from the cacophony

of diverse local practices as the preferred strategy for

herd management in the late 8th and 7th millennia.

However, even though the majority of later

Neolithic assemblages exhibit clear evidence for

young male kill-off there is still a surprising number

(c. 25%) that do not. This degree of variation in

evidence for herd management, although greatly

reduced compared to the early Neolithic, is interest-

ing. If culling young males is the most efficient and

productive method of herd management, what, then,

is the source of this continued variation?

There are several possibilities. First, with the

development of large-scale pastoral systems seasonal

mobility may have increased in the later Neolithic

(Meiggs 2010). In Neolithic France, Helmer et al.

(2005) have shown that seasonal mobility can result

in truncated demographic profiles, with faunal data

from village settlements reflecting only a portion of a

larger, spatially diverse, exploitation system, thus

blurring evidence for young male culling. This may

explain the lack of evidence for young male culling in

the later Neolithic at sites including Ain Ghazal,

Umm el-Tlel and Tell Assouad.

Second, variation in evidence for management may

be the result of the application of multiple exploita-

tion strategies or types of management systems with

no recent parallels. This is perhaps especially the case

within the natural habitat zone of caprines in upland

regions of north-western Iran and central and eastern

Anatolia, where management may have been com-

bined with hunting to produce biometric and demo-

graphic palimpsests which are difficult to interpret.

For example, at 7th-millennium Erbaba Hoyuk,

negative skewing in sheep biometrics combined with

the presence of phenotypically wild sheep suggests

that local mouflon were the target of hunting parties.

However, the presence of small-sized domesticates

and the fact that the mean LSI value for unfused

specimens is larger than that for fused specimens

suggests that management, including young male kill-

off, may have been practised as well (Arbuckle 2008a).

In addition, continued inter-breeding between mana-

ged and wild populations likely resulted in the long

delay in the appearance of morphological changes

evident at upland sites including Cafer, Ganj Dareh,

Asıklı and Suberde.

In other cases variation may simply reflect the

application of management systems with no modern

analogues. This variation may reflect a period of

early experimentation in caprine management

regimes, a common stage in the development of

technological systems (Skibo and Schiffer 2008). The

evidence for variation in herd management strategies,

especially in the early Neolithic, may represent a

period of ‘initial diversity’ in management technolo-

gies, in which individual communities were in the

process of developing husbandry systems that fit into

their unique local environmental, social, historical

and economic contexts. It was only after c. 7500 BC

that this diverse array of strategies, reflecting funda-

mentally local answers to the problems associated with

caprine husbandry, began to be replaced as herders

settled on young male culling as the most effective

answer to their needs — a shift perhaps fuelled by the

development of large, regional populations of produc-

tive domesticates and increased mobility and inter-

community interaction. Even so, pockets of variation



continued into the late Neolithic with some commu-

nities still exhibiting evidence for local adaptations to

their subsistence needs (Asouti and Fuller 2012).

Finally, variations in herd management may have

been affected by the biological limitations of herding

phenotypically wild sheep and goats, especially in the

9th and 8th millennia prior to the widespread

availability of caprines with domestic phenotypes.

Domestic livestock exhibit a series of developmental,

morphological, and behavioural changes, known as

the ‘domestication syndrome’ (Hammer 1984), which

include rapid development, earlier sexual maturity,

and, in males, larger testes and greater sperm

production (Kenagy and Trombulak 1986; Lincoln

et al. 1990; Preston et al. 2003). In recent pastoral

societies practicing young male culling, adult males

often represent only 1–10% of the adult population,

requiring males to inseminate a large number of

females (Redding 1981). In the earliest herding

economies this may have posed a problem for

phenotypically wild male caprines in the early stages

of the domestication process, thus potentially creat-

ing a biological limitation to the effectiveness of

young male kill-off. In addition, Ryder (1960) has

suggested that wild sheep, and presumably also goats,

exhibit significantly slower weight gain compared to

domesticates, which may have also made slaughtering

young animals less attractive to early herders.

Although the impact of these and other issues

associated with herding phenotypically wild animals

are not well understood, they must have affected

Neolithic management regimes. In this context, the

rapid increase in phenotypically domesticated sheep

and goats across south-western Asia in the mid- to

late 8th millennium, may represent a response to the

increased productivity of these new and ‘improved’

domesticates, which spurred the widespread adoption

of intensive caprine pastoralism as well as systems of

young male culling.

Conclusion

In this paper we have surveyed a large body of faunal

data focusing on evidence for the practice of young

male culling in Neolithic south-western Asia. By

focusing on a combination of kill-off age and the

shape of the distributions of biometric data, espe-

cially those from skeletal parts that exhibit the

greatest levels of sexual dimorphism, we are able to

address the emergence and ubiquity of young male

culling amongst Neolithic sheep and goat herders.

Although the intensive culling of young males has

been presented as the best ‘leading edge marker’ for

the initiation of sheep and goat herding, this survey

of available data suggests that young male kill-off

appears in the faunal record only in the early 8th

millennium BC — considerably later than the origins

of caprine management. Instead, Neolithic caprine

management practices appear to have been charac-

terized by a high degree of ‘initial diversity’,

especially in the 9th and early 8th millennia,

suggesting that early management strategies may

have been much more varied than previously realized.

However, after c. 7500 BC young male kill-off was

widely practised across south-western Asia, suggest-

ing this efficient and effective management technol-

ogy quickly replaced the diversity of local

management strategies prevalent earlier — although

some local management systems continued into the

7th millennium BC. This mid- 8th-millennium transi-

tion represents an important shift in the evolution of

animal economies in the Near East, characterized by

the widespread appearance of phenotypically domes-

tic caprines and the spread of intensive and large-

scale sheep and goat pastoralism.

Finally, by highlighting diversity in early caprine

management strategies, this survey also suggests that

local-scale variability was also present in the Late

Pleistocene and earliest Holocene animal economies

that preceded herding economies. This suggests that

10th- and early-9th-millennia assemblages within the

natural habitat zone of wild caprines, including

Hallan Cemi and Kortik Tepe in south-eastern

Turkey, Asiab and Zawi Chemi Shanidar in the

Zagros, and sites such as Jeftelik and Qarassa in Syria

and Nachcharini in Lebanon (Garrard et al. 2003;

Ibanez et al. 2010; Rodrıguez Rodrıguez et al. 2010),

may warrant closer scrutiny in order to identify the

origins of local histories of caprine exploitation that

likely led to the management regimes of the later 9th

millennium (Zeder 2012).

Moreover, new data indicating the complexity and

time depth of human manipulation of wild animals

(e.g. Gifford-Gonzalez and Hanotte 2011; Vigne et al.

2003), suggests that to further clarify the picture of

early animal management, future work needs to focus

on a combination of high-resolution methods and

locally oriented models that seek to identify specific

management practices and define their histories of

use, perhaps extending back to the Pleistocene-

Holocene boundary in particular regions. Although

this includes addressing evidence for young male

culling, it is also necessary to explore evidence for

other management practices, including foddering,

control over mobility, penning and manipulation of

weaning age (Balasse and Tresset 2002; Brochier

1993; Makarewicz and Tuross 2012; Meiggs 2010;



Upex et al. 2012). Only with a better understanding

of these specific management practices will we be able

to develop a detailed understanding of the local

histories from which the earliest systems of animal

husbandry emerged.

Acknowledgements

The authors would like to thank Bill Finlayson and

Cheryl Makarewicz for the invitation to present an

earlier draft of this paper at a session of the Society

for American Archaeology annual meeting in 2012

and to participate in this volume. Financial support

has been provided to BA by Baylor University.

BibliographyAcıkkol, A. (2006) Ucagızlı Magarası Faunasının Zooarkeolojik

Acıdan Incelenmesi: Capra, Capreolus, Dama ve Cervus ların

Morfometrik Acıdan Analizi. Unpublished PhD thesis, Ankara

University.

Arbuckle, B. S. (2006) The evolution of sheep and goat pastoralism and

social complexity in Central Anatolia. Unpublished PhD thesis,

Department of Anthropology, Harvard University.

— (2008a) Caprine exploitation at Erbaba Hoyuk: a Pottery Neolithic

village in Central Anatolia. Pp. 345–65 in L. Gourichon and E.

Vila (eds), Archaeozoology of Southwestern Asia and Adjacent

Areas VIII. Paris: Travaux de la Maison de l’Orient.

— (2008b) Revisiting Neolithic caprine exploitation at Suberde,

Turkey. Journal of Field Archaeology 32, 219–36.

— (2012) Animals in the ancient world. Pp. 201–19 in D. T. Potts (ed.),

A Companion to the Archaeology of the Ancient Near East. Oxford:

Wiley-Blackwell.

— and Erek, C. M. (2010) Late Epipaleolithic hunters of the central

Taurus: faunal remains from Direkli cave, Kahramanmaras,

Turkey. International Journal of Osteoarchaeology 22, 694–707.

— and Makarewicz, C. A. (2009) The early management of cattle (Bos

taurus) in Neolithic central Anatolia. Antiquity 83, 669–86.

— and Ozkaya, V. (2007) Animal exploitation at Kortik Tepe: an early

Aceramic Neolithic site in southeastern Turkey. Paleorient 32,

198–211.

—, Oztan, A. and Gulcur, S. (2009) The evolution of sheep and goat

husbandry in central Anatolia. Anthropozoologica 44, 129–57.

Asouti, E. and Fuller, D. Q. (2012) From foraging to farming in the

southern Levant: the development of Epipaleolithic and Pre-

pottery Neolithic plant management strategies. Vegetation History

and Archaeobotany 21, 149–62.

Atici, L. (2009) Implications of age structures for Epipaleolithic

hunting strategies in the western Taurus mountains, Southwest

Turkey. Anthropozoologica 44, 13–40.

Balasse, M. and Tresset, A. (2002) Early weaning of Neolithic domestic

cattle (Bercy, France) revealed by Intra-tooth Variation in

Nitrogen Isotope Ratios. Journal of Archaeological Science 29,

853–59.

Baumler, H. (1921) Die morphologischen Veranderungen des

Schweineschadels unter dem EInfluss der Domestikation. Archiv

fur Naturgeschichte 87, 140–78.

Binford, L. R. (1968) Post-pleistocene adaptations. Pp. 313–41 in S.

Binford and L. R. Binford (eds), New Perspectives in Archaeology.

Chicago: Aldine.

Bokonyi, S. (1969) Archaeological problems and methods of recogniz-

ing animal domestication. Pp. 219–29 in P. J. Ucko and G. W.

Dimbleby (eds), The Domestication and Exploitation of Plants and

Animals. Chicago: Aldine.

Brochier, J. E. (1993) Cayonu Tepese. Domestication, rythmes et

environnment au PPNB. Paleorient 19, 39–49.

Buitenhuis, H. (1997) Asıklı Hoyuk: a ‘protodomestication’ site.

Anthropozoologica 25–26, 655–62.

— (2008) Ilipinar: The faunal remains from the late Neolithic and early

Chalcolithic periods. Pp. 299–322 in E. Vila, L. Gourichon, A.

Choyke and H. Buitenhuis (eds), Archaeozoology of the Near East

VIII. Lyon: Maison de l’Orient et de la Mediterranee.

Cakırlar, C. (2012) The evolution of animal husbandry in Neolithic central-

west Anatolia: the zooarchaeological record from Ulucak Hoyuk

(c. 7040–5660 cal BC, Izmir, Turkey). Anatolian Studies 62, 1–33.

Childe, V. G. (1936) Man Makes Himself. London: Watts and

Company.

Churcher, C. S. (1994) The vertebrate fauna from the Natufian level at

Jebel es-Saaıde (Saaıde II), Lebanon. Paleorient 20, 35–58.

Davis, S. J. (2000) The effect of castration and age on the development

of the Shetland sheep skeleton and a metric comparison between

bones of males, females and castrates. Journal of Archaeological

Science 27, 373–90.

Dayan, T., Tchernov, E., Bar-Yosef, O. and Yom-Tov, Y. (1986)

Animal exploitation in Ujrat El-Mehed, a Neolithic site in

southern Sinai. Paleorient 12, 105–16.

De Cupere, B. and Duru, R. (2003) Faunal remains from Neolithic

Hoyucek (SW-Turkey) and the presence of early domestic cattle in

Anatolia. Paleorient 29, 107–20.

De Cupere, B., Duru, R. and Umurtak, G. (2008) Animal husbandry at

the Early Neolithic to Early Bronze Age site of Bademagacı

(Antalya province, SW Turkey): evidence from the faunal remains.

Pp. 367–406 in E. Vila, L. Gourichon, A. Choyke and H.

Buitenhuis (eds), Archaeozoology of the Near East VIII. Lyon:

Maison de l’Orient et de la Mediterranee.

Ducos, P. (1978) ‘Domestication’ defined and methodological

approaches to its recognition in faunal assemblages. Pp. 53–56 in

R. H. Meadow and M. Zeder (eds), Approaches to faunal analysis in

the Middle East. Cambridge: Peabody Museum Bulletin 2.

— (1993) Proto-elevage et elevage au Levant Sud au VIIe millenaire BC

les donnees de la Damascene. Paleorient 19, 153–73.

— (1995) Note preliminaire dur les faunes d’Aswad et Ghoraife. Pp.

339–49 in H. de Contenson (ed.), Aswad et Ghoraife. Sites

Neolithiques en Damascene (Syrie), aux IXeme et VIIIeme

millenaires avant l’ere Chretienne. Beyrouth: Bibliotheque

Archeologique et Historique.

Garrard, A., Pirie, A., Schroeder, B. and Wasse, A. (2003) Survey of

Nachcharini Cave and prehistoric settlement in the northern Anti-

Lebanon highlands. Bulletin d’archeologie et d’architecture liba-

naises 7, 15–48.

Gifford-Gonzalez, D. and Hanotte, O. (2011) Domesticating animals in

Africa: implications of genetic and archaeological findings.

Journal of World Prehistory 24, 1–23.

Gourichon, L. and Helmer, D. (2008) Etude archeozoologique de

Mureybet. Pp. 115–228 in J. J. Ibanez (ed.), Le site neolithique de

Tell Mureybet (Syrie du Nord). Oxford: BAR International Series

1843.

Habermehl, K.-H. (1985) Altersbestiummung bei Wildund Pelztieren.

Hamburg: Verlag Paul Parey.

Halstead, P. (1998) Mortality models and milking: Problems of

uniformitarianism, optimality and equifinality reconsidered.

Anthropozoologica 27, 3–20.

Hammer, K. (1984) Das Domestikationssyndrom. Die Kulturpflanze

32, 11–34.

Hecker, H. (1975) The faunal analysis of the primary food animals

from the Pre-Pottery Neolithic Beidha (Jordan). Unpublished

PhD thesis, Columbia University.

Helmer, D. (1989) Le developpement de la domestication au Proche-

Orient de 8500 a 7500 BP: les nouvelles donnees d’El Kowm et de

Ras Shamra. Paleorient 15, 111–21.

— (1992) La domestication des animaux par les homes prehistoriques.

Paris: Masson.

— (2008) Revision de la faune de Cafer Hoyuk (Malatya, Turquie):

apports des methodes de l’analyse des melanges et de l’analyse de

Kernel a la mise en evidence de la domestication. Pp. 169–96 in E.

Vila, L. Gourichon, A. Choyke and H. Buitenhuis (eds),

Archaeozoology of the Near East VIII. Lyon: Maison de l’Orient

et de la Mediterranee.

— and Gourichon, L. (2008) Premieres donnees sur les modalites de

subsistance a Tell Aswad (Syrie, PPNB Moyen et Recent,

Neolithique Ceramique Ancien) - Fouilles 2001–2005. Pp. 119–

51 in E. Vila, L. Gourichon, A. Choyke and H. Buitenhuis (eds),

Archaeozoology of the Near East VIII. Lyon: Maison de l’Orient et

de la Mediterranee.

—, Gourichon, L., Sidi Maamar, H. and Vigne, J.-D. (2005) L’elevage

des caprines neolithiques dans le Sud-Est de la France: saisonna-

lite des abattages, relations entre grottes bergeries et sites de plein

air. Anthropozoologica 40, 167–90.



Henton, E. (2012) The combined use of oxygen isotopes and microwear

in sheep teeth to elucidate seasonal management of domestic

herds: the case study of Catalhoyuk, central Anatolia. Journal of

Archaeological Science 39, 3264–76.

Hesse, B. (1978) Evidence for husbandry from the early Neolithic site of

Ganj Dareh in western Iran. Unpublished PhD thesis, Columbia

University.

Hole, F., Flannery, K. V. and Neely, J. A. (1969) Prehistory and human

ecology on the Deh Luran Plain. Museum of Anthropology,

University of Michigan, Memoir No.1. Ann Arbor: Museum of

Anthropology, University of Michigan.

Hongo, H., Meadow, R. H., Oksuz, B. and Ilgezdi, G. (2005) Sheep

and goat remains from Cayonu Tepesi, Southeastern Anatolia. Pp.

112–23 in H. Buitenhuis, A. Choyke, L. Martin, L. Bartosiewicz

and M. Mashkour (eds), Archaeozoology of the Near East VI.

Proceedings of the Sixth International Symposium on the

Archaeozoology of Southwestern Asia and Adjacent Areas.

Groningen: ARC Publication 123.

Horwitz, L. K. (2003) The Neolithic fauna. Pp. 87–101 in H. Khalaily

and O. Marder (eds), The Neolithic Site of Abu Gosh. The 1995

Excavations. Jerusalem Israeli Antiquities Authority Reports vol.

19.

—, Tchernov, E., Ducos, P., Becker, C., Driesch, A. v. d., Martin, L.

and Garrard, A. (1999) Animal domestication in the southern

Levant. Paleorient 25, 63–80.

Ibanez, J. J., Balbo, A., Braemer, F., Gourichon, L., Iriarte, E.,

Santana, J. and Zapata, L. (2010) The early PPNB levels of Tell

Qarassa North (Sweida, southern Syria). Antiquity 84.

Ilgezdi, G. (2008) The domestication process in Southeastern Turkey:

the evidence of Mezraa-Teleilat. Unpublished PhD thesis,

Geowissenschaftlichen Fakultat der Eberhard-Karls-Universitat

Tubingen.

Kenagy, G. J. and Trombulak, S. C. (1986) Size and function of

mammalian testes in relation to body size. Journal of Mammalogy

67, 1–22.

Kersten, A. M. P. (1987) Age and sex composition of Epipaleolithic

fallow deer and wild goat from Ksar ‘Akil. Palaeohistoria 29, 119–31.

Legge, A. J. (1996) The beginning of caprine domestication in

southwest Asia. Pp. 238–62 in D. Harris (ed.), Origins and

Spread of Agriculture and Pastoralism in Eurasia. London: UCL

Press.

— and Rowley-Conwy, P. A. (2000) The exploitation of animals. Pp.

423–71 in A. M. T. Moore, G. C. Hillman and A. J. Legge (eds),

Village on the Euphrates. Oxford: Oxford University Press.

Lincoln, G. A., Lincoln, C. E. and McNeilly, A. S. (1990) Seasonal

cycles in the blood plasma concentration of FSH, inhibin and

testosterone, and testicular size in rams of wild, feral and

domestication breeds. Journal of Reproduction and Fertility 88,

623–33.

Losch, S., Grupe, G. and Peters, J. (2006) Stable isotopes and dietary

adaptations in humans and animals at Pre-Pottery Neolithic

Nevalı Cori, Southeast Turkey. American Journal of Physical

Anthropology 131, 181–93.

Ludwig, A., Pruvost, M., Reissmann, M., Benecke, N., Brockmann,

G. A., Castanos, P., Cieslak, M., Lippold, S., Llorente, L.,

Malaspinas, A.-S., Slatkin, M. and Hofreiter, M. (2009) Coat

color variation at the beginning of horse domestication. Science

324, 485.

Makarewicz, C. (2009) Complex caprine harvesting practices and

diversified hunting strategies: Integrated animal exploitation

systems at Late Pre-Pottery Neolithic B ‘Ain Jammam.


— and Tuross, N. (2012) Finding fodder and tracking transhumance:

Isotopic detection of goat domestication processes in the Near

East. Current Anthropology 53, 495–505.

Marom, N. and Bar-Oz, G. (2013) The prey pathway: a regional history

of cattle (Bos taurus) and pig (Sus scrofa) domestication in the

northern Jordan Valley, Israel. PLOS One 8, e55958.

Matthews, W. (1998) Appendix 2. Micromorphological analysis of

occupation sequences at the Aceramic Neolithic settlement of

Asikli Hoyuk: an assessment, and comparison to depositional

sequences at Catalhoyuk. Catalhoyuk 1998 Archive Report.

Meadow, R. H. (1989) Osteological evidence for the process of animal

domestication. Pp. 80–96 in J. Clutton-Brock (ed.), The Walking

Larder: Patterns of Domestication, Pastoralism, and Predation.

London: Unwin Hyman.

— (1999) The use of size index scaling techniques for research on

archaeozoological collections from the Middle East. Pp. 285–300

in C. Becker, H. Manhart, J. Peters and J. Schibler (eds), Historia

animalium ex ossibus: Beitrage zur palaoanatomie, archaologie,

agyptologie, ethnologie, and geschichte der tiermedizin. Rahden:

Verlag Marie Leidorf.

Meiggs, D. (2010) Investigation of Neolithic ovicaprine herding

practices by multiple isotope analysis: a case study at PPNB

Gritlle, Southeastern Turkey. Unpublished PhD thesis,

Department of Anthropology, University of Wisconsin.

Monahan, B. H. (2000) The organization of domestication at Gritille, a

Pre-Pottery Neolithic B site in southeastern Turkey. Unpublished

PhD thesis, Northwestern University.

Monchot, H. (1999) Mixture analysis and mammalian sex ratio among

middle Pleistocene mouflon of Arago Cave, France. Quaternary

Research 52, 259–68.

— and Lechelle, J. (2002) Statistical nonparametric methods for the

study of fossil populations. Paleobiology 28, 55–69.

Moore, D. S. and McCabe, G. P. (2001) Introduction to the Practice of

Statistics. New York: W.H. Freeman and Company.

Naderi, S., Rezaei, H.-R., Pompanon, F., Blum, M. G. B., Negrini, R.,

Naghash, H.-R., Balkiz, O., Mashkour, M., Gaggiotti, O. E.,

Ajmone-Marsan, P., Kence, A., Vigne, J.-D. and Taberlet, P.

(2008) The goat domestication process inferred from large-scale

mitochondrial DNA analysis of wild and domestic individuals.

PNAS 105, 17659–64.

Ottoni, C., Flink, L. G., Evin, A., Georg, C., De Cupere, B., Van Neer,

W., Bartosiewicz, L. S., Linderholm, A., Barnett, R., Peters, J.,

Decorte, R., Waelkens, M., Vanderheyden, N., Ricaut, F. O.-X.,

Rus Hoelzel, A., Mashkour, M., Karimlu, A. F. M., Seno, S. S.,

Daujat, J., Brock, F., Pinhasi, R., Hongo, H., Perez-Enciso, M.,

Rasmussen, M., Frantz, L., Megens, H.-J., Crooijmans, R.,

Groenen, M., Arbuckle, B., Benecke, N., Vidarsdottir, U. S.,

Burger, J., Cucchi, T., Dobney, K. and Larson, G. (2012) Pig

domestication and human-mediated dispersal in western Eurasia

revealed through ancient DNA and geometric morphometrics.

Molecular Biology and Evolution 30(4), 824–832.

Payne, S. (1973) Kill-off patterns in sheep and goats: the Mandibles

from Asvan-kale. Anatolian Studies 23, 281–303.

Pearson, J. A., Buitenhuis, H., Hedges, R. E. M., Martin, L., Russell,

N. and Twiss, K. (2007) New light on early caprine herding

strategies from isotope analysis: a case study from Neolithic

Anatolia. Journal of Archaeological Science 34, 2170–79.

Perkins, D. (1964) Prehistoric fauna from Shanidar, Iraq. Science 144,

1565–66.

— (1973) The beginnings of animal domestication in the Near East.

American Journal of Archaeology 77, 279–82.

Peters, J., Buitenhuis, H., Grupe, G., Schmidt, K. and Pollath,

N. (2013) The long and winding road. Ungulate exploitation

and domestication in early Neolithic Anatolia (10,000–7,000

cal BC). In S. Colledge, J. Connolly, K. Dobney, K. Manning

and Shennan, S. (eds), Origins and Spread of Domestic Animals

in Southwest Asia and Europe. Walnut Creek: CA Left Coast

Press, 83–114.

Peters, J., von den Driesch, A. and Helmer, D. (2005) The upper

Ephrates-Tigris basin: cradle of agro-pastoralism? Pp. 96–124 in J.-

D. Vigne, J. Peters and D. Helmer (eds), The First Steps of Animal

Domestication: New Archaeological Approaches. Proceedings of the

9th ICAZ Conference, Durham 2002. Oxford: Oxbow.

Preston, B. T., Stevenson, I. R., Pemberton, J. M., Coltman, D. W. and

Wilson, K. (2003) Overt and covert competition in a promiscuous

mammal: the importance of weaponry and testes size to male

reproductive success. Proceedings of the Royal Society B 270, 633–40.

Pumpelly, R. (1908) Explorations in Turkestan; Expedition of 1904.

Prehistoric Civilizations of Anau; Origins, Growth and Influence of

Environment. Washington DC: Carnegie Institution of

Washington Publication No. 73.

Redding, R. (1981) Decision making in subsistence herding of sheep and

goats in the Middle East. Unpublished PhD thesis, Departments of

Anthropology and Biology, University of Michigan.

Rodrıguez Rodrıguez, A., Haıdar-Boustani, M. G. U., Ibanez, J. J., Al-

Maqdissi, M., Terradas, X. and Zapata, L. (2010) Jeftelik: a new

Early Natufian site in the Levant (Homs Gap, Syria). Antiquity 84.

Russell, N. (2012) Social Zooarchaeology: Humans and Animals in

Prehistory. Cambridge: Cambridge University Press.



— and Martin, L. (2005) The Catalhoyuk mammal remains. Pp. 33–98

in I. Hodder (ed.), Inhabiting Catalhoyuk: Reports from the 1995–

1999 seasons. Cambridge: McDonald Institute for Archaeological

Research.

—, Martin, L. and Buitenhuis, H. (2005) Cattle domestication at

Catalhoyuk revisited. Current Anthropology 46 Supplement, S101–8.

Ryder, M. L. (1960) A study of the coat of the mouflon Ovis musimon

with special reference to seasonal change. Proc. Zool. Lond.135,

387–408.

Schaller, G. B. (1977) Mountain Monarchs: Wild Sheep and Goats of the

Himalyah. Chicago: University of Chicago Press.

Silver, I. A. (1963) The aging of domestic animals. Pp. 250–68 in D.

Brothwell, E. Higgs and G. Clark (eds), Science in Archaeology.

London: Thames and Hudson.

Simmons, A. H. and Ilany, G. (1975–77) What mean these bones?

Paleorient 3, 269–74.

Skibo, J. M. and Schiffer, M. B. (2008) People and Things: A Behavioral

Approach to Material Culture. New York: Springer.

Starkovich, B. M. and Stiner, M. C. (2009) Hallan Cemi Tepesi: high-

ranked game exploitation alongside intensive seed processing at

the Epipaleolithic-Neolithic transition in southeastern Turkey.


Tchernov, E. and Bar-Yosef, O. (1982) Animal exploitation in the Pre-

Pottery Neolithic B Period at Wadi Tbeik, Southern Sinai.

Paleorient 8, 17–37.

Uerpmann, H.-P. (1979) Probleme der Neolithisierung des

Mittelmeeraums. Wiesbaden: Verlag.

Upex, B., Balasse, M., Tresset, A., Arbuckle, B. and Dobney, K. (2012)

Protocol for recording enamel hypoplasia in modern and

archaeological caprine populations. International Journal of

Osteoarchaeology, DOI: 10.1002/oa.2227.

Vanpoucke, S., Mainland, I., De Cupere, B. and Waelkens, M. (2009)

Dental microwear study of pigs from the classical site of

Sagalassos (SW Turkey) as an aid for the reconstruction of

husbandry practices in ancient times. Environmental Archaeology

14, 137–54.

Vigne, J.-D. (2011a) Histoire de la gestion des chevres et des moutons.

Pp. 1039–57 in J. Guilaine, F. Briois and V. Jean-Denis (eds),

Shillourokambos: Un etablissement neolithique pre-ceramique a

Chypre. Les fouilles du secteur 1. Paris: Editions Errance.

— (2011b) Le mouton (Ovis aries). Pp. 1021–38 in J. Guilaine, F. Briois

and V. Jean-Denis (eds), Shillourokambos: Un etablissement

neolithique pre-ceramique a Chypre. Les fouilles du secteur 1.

Paris: Editions Errance.

—, Carrere, I., Briois, F. and Guilaine, J. (2011) The early process of

mammal domestication in the Near East. Current Anthropology

52, S255–71.

—, Carrere, I. and Guilaine, J. (2003) Unstable status of early domestic

ungulates in the Near East: the example of Shillourokambos

(Cyprus, IX–VIIIth millennia cal BC). Pp. 239–51 in J. Guilaine

and A. Le Brun (eds), Le Neolithique de Chypre. Actes du colloque

international organise par le Department des Antiquites de Chypre

et l’Ecole Francaise d’Athenes, Nicosie 17–19, Mai 2001. Athens:

Ecole francaise d’Athenes.

— and Helmer, D. (2007) Was milk a ‘secondary product’ in the Old

World Neolithisation process? Its role in the domestication of

cattle, sheep, and goats. Anthropozoologica 42, 9–40.

von den Driesch, A. (1976) A guide to the measurements of animals

bones from archaeological sites. Peabody Museum Bulletin 1.

Cambridge, MA: Peabody Museum of Archaeology and

Ethnology, 1–137.

Wasse, A. (2002) Final results of an analysis of the sheep and goat

bones from Ain Ghazal, Jordan. Levant 34, 59–82.

Watson, J. P. N. (1978) The interpretation of epiphyseal fusion data.

Pp. 97–101 in D. R. Brothwell, K. D. Thomas and J. Clutton-

Brock (eds), Research Problems in Zooarchaeology. London:

Institute of Archaeology Occasional Publication No. 3.

Winge, H. (1900) Bløddyr-skaller. Knogler af dyr. Plante Levninger. Pp.

158–63.

Wolverton, S. (2008) Harvest pressure and environmental carrying

capacity: an ordinal-scale model of effects on ungulate prey.

American Antiquity 73, 179–99.

Zeder, M. (2001) A metrical analysis of a collection of modern goats

(Capra hircus aegagrus and C. h. hircus) from Iran and Iraq:

Implications for the study of caprine domestication. Journal of

Archaeological Science 28, 61–79.

— (2006) Archaeological approaches to documenting animal domestica-

tion. Pp. 171–80 in M. Zeder, D. Bradley, E. Emshwiller and B. D.

Smith (eds), Documenting Domestication: New Genetic and

Archaeological Paradigms. California: University of California Press.

— (2008a) Animal domestication in the Zagros: an update and

directions for future research. Pp. 243–78 in E. Vila,

L. Gourichon, A. Choyke and H. Buitenhuis (eds),

Archaeozoology of the Near East VIII. Lyon: Maison de l’Orient

et de la Mediterranee.

— (2008b) Domestication and early agriculture in the Mediterranean

Basin: Origins, diffusion, and impact. Proceedings of the National

Academy of Sciences 105, 11597–604.

— (2011) The origins of agriculture in the Near East. Current

Anthropology 52, 221–35.

— (2012) The Broad Sprectrum Revolution at 40: resource diversity,

intensification, and an alternative to optimal foraging explana-

tions. Journal of Anthropological Archaeology 31, 241–64.

— and Hesse, B. (2000) The initial domestication of goats (Capra hircus)

in the Zagros mountains 10,000 years ago. Science 287, 2254–57.



Initial diversity in sheep and goat management in Neolithic south-western Asia

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