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http://www.elsevier.com/locate/jas

Gazelle bone fat processing in the Levantine Epipalaeolithic

Natalie D. Munroa,*, Guy Bar-Ozb

aDepartment of Anthropology, Unit 2176, 354 Mansfield Road, University of Connecticut, Storrs, CT 06269, USAbZinman Institute of Archaeology, University of Haifa, Haifa 31905, Israel

Received 7 June 2004; received in revised form 14 July 2004

Abstract

We investigate mountain gazelle (Gazella gazella) carcass processing to reconstruct resource intensification strategies during theEpipalaeolithic period of northern Israel. We adopt a multivariate taphonomic approach to identify the processes that most

influenced bone survivorship in five gazelle assemblages. All of the assemblages are characterized by significant density-mediatedbiases, yet in situ attrition played a minimal role in assemblage formation. In contrast, the survivorship of hare (Lepus capensis)skeletons is not mediated by bone density indicating that different prey taxa experienced independent taphonomic histories. Bothgazelle cortical and cancellous bone is highly fragmented and the degree of fragmentation and survivorship are strongly correlated

with fat yields. Results of multiple tests point to intensive marrow and grease extraction as the primary determinant of gazelle bonesurvivorship. Although gazelle carcasses were intensively utilized throughout the Epipalaeolithic, the intensity of use is stable acrossthe duration of the period.

� 2004 Elsevier Ltd. All rights reserved.

Keywords: Bone processing; Bone survivorship; Epipalaeolithic; Intensification; Levant; Gazelle; Grease; Marrow; Taphonomy

1. Introduction

Studies of resource intensification have compriseda major dimension of archaeological inquiry over thepast few decades [21,25,26,42,44,91]. Resource intensi-fication is frequently linked to significant transitions inhuman prehistory including the beginnings of sedentism,the origins of agriculture, and the rise of civilization.Animal resource intensification has typically beeninvestigated at the level of the community (i.e., speciesdistribution, abundance, and diversity) and taxon (i.e.,demographic profiles; average prey body size), but israrely considered at the level of the organism itself. Theintensity of carcass use provides an effective approach to

* Corresponding author. Tel.: C1 860 486 0090.

E-mail addresses: natalie.munro@uconn.edu (N.D. Munro), guy-

bar@research.haifa.ac.il (G. Bar-Oz).

0305-4403/$ - see front matter � 2004 Elsevier Ltd. All rights reserved.

doi:10.1016/j.jas.2004.08.007

reconstruct human extractive efforts especially in theface of diminishing returns [14,19,20,35,65,67,76,93].

The Epipalaeolithic cultures of the Southern Levantare frequently cited as classic examples of intensifyingforagers [5,9,47,48]. The Epipalaeolithic period (ca. 21–11.5 kya) falls along a trajectory of intensified plant usethat extends from the Palaeolithic to the Neolithic[107,108], and culminates with the adoption of agricul-ture and animal husbandry. Several community andspecies-level trends in animal exploitation also indicateintensification throughout the Palaeolithic and Epipa-laeolithic periods. One of the best documented isa change at the end of the Epipalaeolithicdsmall gameincrease in relative abundance [1,33,34,71,90,95,96,97,99], and particularly fast-moving small animals. Re-mains of mountain gazelle (Gazella gazella) also increasein abundance in relation to other ungulate taxa[31,46,55,98,99] and there are increased proportions ofjuvenile [32,71,94] and male gazelles [4,28,98,99]. This

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may point to an intensive gazelle economy during theLate Epipalaeolithic. Yet we know little about theintensity of gazelle carcass exploitation. In studies ofEpipalaeolithic faunal assemblages we [1,71] suggestthat resource intensification should also be expressed inthe intensive use of individual gazelle carcasses, in-cluding the extraction of bone marrow and grease. TheEpipalaeolithic sequence in the Mediterranean environ-ment of the southern Levant provides an opportunity toinvestigate trends in gazelle processing strategies thatmay include the beginning of grease extraction in theregion.

Here, we focus specifically on bone processing forgrease and medullary marrowdactivities that can bereconstructed from gazelle skeletal remains. Bone fat(especially grease) is often costly to procure, butprovides an essential dietary resource for humans,particularly during critical resource periods [85,86,88].Because grease and medullary marrow have differentprocessing costs (see below), identifying if and howoften these components of bone were extracted byEpipalaeolithic foragers provides an additional avenuefor investigating the intensity of individual carcass use.

1.1. Bone marrow

Bone marrow extraction is well known amonghunter-gatherer and agricultural societies worldwide[14,15,24,36,75,109]. When an animal is healthy and ingood nutritional condition, marrow is a rich, fattysubstance that is stored primarily in the large interiorcavities (medullae) of most limb bones, and themandible [29]. To access marrow, the bone casing mustbe breached, most often by striking it with a hardhammer. The concentration and types of fatty acids inbone marrow are variable. Marrow is often moreconcentrated in upper limb bones such as the femur,yet the marrow of the lower extremities contains moreoleic acid. Oleic acid has a lower melting point thanother lipids in bone marrow, and is more nutritious,storable, and reportedly even tastier than the marrow ofthe upper limbs [14, p. 24]. Because the bone cavity mustbe broken to extract the marrow, such extraction resultsin fragmentation of long bone shafts and the horizontalramus of the mandible in particular. Fragmentation canpotentially reduce bone representation by decreasing thevisibility of anatomically and taxonomically diagnosticfeatures, and by exposing more surface area to de-struction by post-depositional processes [50,62]. Bonesthat are intensively processed for marrow are thusexpected to be characterized by higher NISP:MNEratios (i.e., more fragments per element) and lowersurvivorship than non-marrow bearing bones [67].

Bone marrow yield varies by species, skeletal element,and bone portion. Not all elements contain marrow andthe marrow in those that do differs in concentration and

quality. Because humans are expected to preferentiallyopen bones with the highest marrow content, theintensity of fragmentation (i.e., NISP:MNE) should becorrelated with marrow yields. Resource intensificationcan be assessed by examining the size of marrow storesthat humans opened.

Actualistic studies have demonstrated that marrowextraction is a cost-effective means to obtain animal fat[14,16,20,58,63]. These studies involved the removal ofbone marrow from a variety of fresh ungulate carcassesto measure quantity, nutritional quality, seasonalvariability, and processing costs. Because the availablestudies had diverse goals and measurement strategies,the results are difficult to quantify and compare, buta few general themes emerge. First, processing timetends to increase with prey body size [58]. Second, thecomposition and/or yield of marrow vary considerablyin relation to the age, sex, and physical condition ofthe animal [86]. Third, variation in processing efficiencyis influenced more by marrow content of a givencarcass than processing costs [14, p. 25]. Finally,marrow processing has been shown to be more costeffective than grease processing due to the high cost ofextracting grease.

1.2. Bone grease

Grease provides an additional source of fat which isstored within the microstructure of cancellous (spongy)bone. One of its most valuable characteristics is its 3year storability if rendered properly [54]. This isespecially relevant in the Epipalaeolithic when foragerslikely engaged in some of the earliest storage practiceswhile becoming increasingly sedentary. Rendered fatcan be stored either in solid cakes, as a liquid in skinbags, or by mixing it with dry meat to preparepemmican [54,103]. Grease is more work to extract thanmedullary marrow, but its extraction increases theamount of nutrients that can be obtained from a givenanimal carcass.

There is good evidence for marrow extraction as earlyas the Plio-Pleistocene [16,17,22,45], but grease process-ing is a much more recent development. Exactly howrecent, however, is unknown. The extraction of bonegrease differs from marrow in that it generally requiresboiling, an elaborate and time-consuming technology.To extract grease cancellous bone must be fractured(and even pulverized in some situations) using a hammerand anvil and may subsequently be pounded andcrushed [14, p. 158, 51]. The grease is then removed byboiling the bones, often for at least 2–3 h [14,27,59,83].Following extraction, the grease is skimmed from thesurface and prepared for immediate consumption orstorage. The fat in cancellous bone may also beexploited by adding bones to stews [49]. Ethnographicevidence indicates that animals are frequently butchered

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into segments and then added to liquids and cooked[49,109]. The bones of large animals must be broken tofit them in the pot and to further facilitate greaseremoval. Prior to the invention of ceramics, boilingrequired the use of organic containers which seldompreserve in the archaeological record [14, p. 159]. Moreresilient items such as anvils and milling stones, fire-cracked rock and in some cases subterranean pits,however, may provide secondary evidence for greaseexploitation.

According to ethnographic research, the bone refusetypically produced by grease rendering consists ofnumerous small bone fragments, many of which arebroken beyond recognition [54,56,76]. Recent experi-ments suggest, however, that bones do not have to befully comminuted for efficient grease extraction to takeplace. Church and Lyman [27] discovered that greaseextraction from fragments averaging 1, 2, 3, and 5 cmwas equally efficient if the bones were boiled to the pointof diminishing returns (ca. 2–3 h). The extraction ofgrease from small segments (1 or 2 cm) was moreefficient than larger fragments (ca. 5 cm) only in the firsthour of boiling. Church and Lyman [27; pp. 1077–1084]thus conclude that ‘‘small fragments make small differ-ences in the rate or efficiency of grease extraction’’,although small fragments may be preferred if fuel is inshort supply. Cancellous bone assemblages averaging upto 5 cm in length may thus provide an archaeologicalsignature for grease production as equally plausible asfinely comminuted bone.

Unfortunately, the utility of fragmentation fordetecting grease extraction is overshadowed by thespecter of equifinalityda plethora of taphonomic pro-cesses are known to cause the fragmentation ofcancellous bone [62]. Because grease extraction isdifficult to identify in the archaeological record, yet itsappearance is crucial to our understanding of Epipa-laeolithic intensification strategies, we are especiallyinterested in refining taphonomic methods for itsidentification.

Several researchers have measured the quantity ofgrease extracted from a range of animals and elements[14,20,27,54,59]. Variation in the quantities of greaserecovered is likely attributable to the wide range ofanimals investigated, sample size, variation in sex, age,physical condition, season of death, and experimentalmethods. Despite this variability, several researchershave observed that the extraction of bone greaserequires substantially more effort than comparablequantities of bone marrow in terms of time, energeticinvestment, and firewood [14,59,83]. This difference isexacerbated when grease extraction is undertaken usingheated rocks in a hide or gut container rather thana ceramic or metal pot placed directly in the fire.Because the lipids in bone marrow and grease havesimilar nutritional values, the appearance of grease

rendering in the archaeological record represents a sig-nificant degree of intensification.

Boiling bone for grease extraction may influence itssubsequent survivorship in response to an array oftaphonomic agents [49]. For example, scavengingcarnivores tend to avoid boiled bones which are devoidof nutritional value [57,87]. Gifford-Gonzalez [39] notesthat bone boiling encourages the breakdown of bonecollagen which weakens the structure of bone andlowers its resilience to mechanical forces. Furthermore,reduction of the organic component of bone by boilingreduces the chance of microbial attack and the resultantbone mineral organization may be less chemicallyreactive in the burial environment [81]. Although someof these consequences may protect grease-processedbones from post-depositional attrition, others may in-crease bone’s susceptibility to diagenetic alteration dueto rapid increases in porosity caused by emptying voidsof fat and protein and reduced mechanical strength [81].

Clearly, a variety of attritional forces (e.g., carnivoregnawing, weathering, trampling, in situ attrition) maysimultaneously lead to the fragmentation of bonefollowing its disposal by humans. These factors maydistort or mimic patterns produced by human sub-sistence behaviors [61,62]. It is thus essential todistinguish the causes of bone fragmentation if we areto identify the most influential taphonomic agents ofassemblage formation and subsequently reconstructhuman behavior [92]. In this study we seek to separateunlikely post-depositional taphonomic processes fromthe most probable formation processes. We adopta multivariate inter-site taphonomic strategy [3,11] tocompare various taphonomic attributes of faunalassemblages originating in similar ecological and geo-graphic settings in the Mediterranean vegetation belt ofnorthern Israel. Two detailed, high-resolution databasescollected using similar research protocols are merged[1,70]. All of the sites studied date to the Epipalaeolithicand contain deep cultural deposits, high densities ofartifacts, and groundstone implements. We focus ongazelle, the most common species in the assemblages.We make comparisons to hares (Lepus capensis) andpartridges (Alectoris chukar) to highlight patterns ofbone survivorship that are exclusive to gazelles.

2. The Epipalaeolithic sites and their settings

We investigate gazelle grease and marrow processingacross a sequence of five Epipalaeolithic occupationsfrom the coastal plain and Mediterranean Hills of thewestern Galilee and Carmel region of northern Israel.The Mediterranean Epipalaeolithic comprises threecultural periodsdthe Kebaran (ca. 21.5–17 cal kya),the Geometric Kebaran (ca. 17–14.5 cal kya), and theNatufian (ca. 14.5–11.5 cal kya) [5].

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In the southern Levant Kebaran occupations arerestricted largely to the more hospitable coastal plainand the Mediterranean hills, particularly during the LastGlacial Maximum (LGM). The Kebarans were mobileforagers who moved seasonally between the flat coastalplain and the adjacent Mediterranean hills [9]. Geo-metric Kebaran sites share the same geographicdistribution as Kebaran sites, but extend into thesemiarid zone of the Negev, Sinai, and Jordaniandeserts. Geometric Kebaran foragers also followeda mobile hunting and gathering strategy interrupted byperiods of aggregation. The Natufian is the last of theEpipalaeolithic periods and is most often divided intotwo phases [12] (but see [100]). The Early Natufian(14.5–13 cal kya) is best known for its permanentstructures, which are generally believed to have housedthe first sedentary or semi-sedentary foragers [9,47,89].The largest of the Early Natufian sites are characterizedby rich bone tool, groundstone, and ornamentaltraditions, and the presence of cemeteries [7–9]. Al-though many aspects of these traditions continue intothe later phase, the Late Natufian (13–11.5 cal kya)corresponds to the Younger Dryas cooling and dryingevent and is characterized by increased mobility andreduced site occupation intensity and investment intopermanent features such as architecture [12,41,47,71,101].

The single Kebaran site we consider, Nahal HaderaV, overlooks the Hadera River on the coastal plain ofnorthern Israel (Fig. 1). The site is situated on the first ofthree kurkar sandstone ridges 1 km west of the modernMediterranean shoreline, although it was at least 10 kmfrom the sea at the time of occupation. Nahal Hadera Vis an open-air site with deep cultural deposits containingartifacts characteristic of the Kebaran phase. Directluminescence dating of sand from the site suggests thathuman occupation occurred between 21 and 14 cal kya[40]. The site preserved high densities of artifactsincluding lithic and groundstone tools. Excavation ofNahal Hadera V was undertaken by Jim Phillips in 1973and by Avi Gopher and Ran Barkai in 1997–1999 andyielded a well-preserved faunal sample (NISP=19,513)[2].

The Geometric Kebaran site of Hefzibah is only a fewhundred meters from Nahal Hadera V on the samesandstone ridge (Fig. 1). The faunal sample originatesfrom a 4!2 m excavation directed by Ohad Zackheimand Guy Bar-Oz between 1996 and 1998. Theseexcavations yielded a dense litter of artifactual remainsand a large faunal assemblage. Hefzibah’s stratigraphycan be divided into two components on the basis ofdifferential preservation [3]. The fauna from layers 1–6were strongly biased by taphonomic forces and are notincluded here. The assemblage from Hefzibah consistsof 8507 identifiable specimens that originated in layers7–18.

Hayonim Cave is a deeply stratified site situated inthe Mediterranean Hills of the western Galilee about 13km from the coastal plain (Fig. 1). The Natufian layercontains 9 round structures, rich artifactual assemblagesincluding lithics, groundstone, bone tools, ornamentsand portable art objects, and numerous human graves[9]. The large number of human graves in the Natufianlayer suggests that the site also served as a ritual orburial site. Hayonim Cave includes both Early and LateNatufian components that can be separated on the basisof lithic markers [5,6]. Faunas from the Early(NISP=8159) and Late (NISP=7354) Natufian layersare represented by similar numbers of identifiablespecimens.

Hayonim Cave

el-Wad

Nahal Hadera VHefzibah

Fig. 1. Map showing the location of the Epipalaeolithic sites analyzed.

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Finally, el-Wad Cave and its adjacent terrace aresituated at the foot of the western slope of MountCarmel, on the southern cliff of Nahal Me’arot (Wadi el-Mughara; Fig. 1). Dorothy Garrod excavated el-Wadover five seasons from 1929 to 1933. The Natufian layeris the best represented layer at the site. It stretches acrossthe cave and the entire terrace. Natufian deposits on theterrace yielded few architectural remains, close to 100burials [13], a rich lithic assemblage, decorative items,bone tools, ground stone implements, and a large faunalassemblage [38]. In 1980 and 1981 limited excavationswere conducted by Francois Valla and Ofer Bar-Yosef[102], while excavations in the cave were renewed byMina Weinstein-Evron in 1988 and 1989 [106]. Newexcavations were initiated on the terrace by MinaWeinstein-Evron and Daniel Kaufman starting in1995. The new excavations are located to the north-eastof Garrod’s trench over an area of 60 m2. The richmaterial culture belongs to the Late Natufian andincludes characteristic flint tools, ground-stone imple-ments, bone tools, decorative items (shells and beads)and art objects. The bone sample analyzed in this studycomes from the 1995–2000 excavations on the terrace,and includes only faunal remains that originated fromundisturbed Late Natufian contexts (NISP=2899) [4].

3. Methods

The complete zooarchaeological and taphonomicprocedures used to collect and evaluate the data setsfor all studied assemblages are presented in Bar-Oz [1]and Munro [70]. All assemblages were meticulouslycollected by dry and wet screening through 2 mm meshand ‘‘picking’’ dry sediments for small bones. All boneswere saved and all identifiable fragments were removedfor analysis by the authors. Identifiable bones includedcranial fragments, vertebrae, long bone articular endsand shafts, and all other recognizable bone and teethfragments which were then assigned to the mostdiscriminating taxonomic level possible. If elementscould not be assigned to species, they were identifiedto body-size classes (i.e., medium mammal, small un-gulate). Bar-Oz’s medium body-size class and Munro’ssmall ungulate group are combined with gazellessince other possible contendersdroe deer (Capreoluscapreolus) and wild goat (Capra aegagrus)dcompriseless than 1% of the NISP in all assemblages. Identifiablebone specimens were assigned to element, side (i.e., left,right, or axial), and portion (e.g., proximal epiphysis,distal epiphysis, medial shaft). Shaft fragments werecoded according to the presence of specific zones (i.e.,proximal shaft, distal shaft, mid-shaft) or diagnosticfeatures (foramen, muscle attachments, and otherfeatures). In most cases identified specimens were coded

according to their fraction of completeness (i.e.,percentage of complete bone; percentage of bone shaftconnected to an epiphysis).

Frequencies of element portions were used tocalculate the minimum number of skeletal elements(MNE), the minimum number of animal units (MAU),and the minimum number of individuals (MNI)following the protocol outlined by Klein and Cruz-Uribe [51] and Lyman [62]. Different tests required theuse of different quantitative measures and bone por-tions. In some cases MNE was calculated for differentportions of the same element (i.e., proximal end orshaft), in others it was calculated for the completeelement, and in some cases juvenile bone was excludedfrom the calculation. Juvenile bone shafts were identi-fied by the high porosity of their bone structure. Todetermine MNE the frequency of each bone portion orfeature in the assemblage was tabulated, and the mostcommon portion represents the MNE. If more than onelandmark or portion was present on a bone specimen,each landmark was counted once. This includes longbone fragments comprised partly of articular ends andpartly of shafts (i.e., the bone is used to tabulate theMNE for both the end and the shaft). Recordedelements were examined for macroscopic surface mod-ifications, such as bone weathering [10], cut marks [15],percussion marks [17,89], and evidence for animalactivity such as rodent gnawing, carnivore punctures,scoring, and digestion [18,37]. Bone surfaces from theHayonim Cave gazelle assemblages were examinedfollowing the recommendation of Blumenschine et al.[18], using a microscope at 10! magnification. Finally,a sample of shaft fragments (those attached to long boneepiphyses) were selected to investigate whether boneswere broken when fresh (green) or old (dry). Themorphology of the fracture angle, fracture outline, andfracture edge were recorded following Villa and Mahieu[104] because their protocol allows distinction of freshfrom post-burial breaks. Shaft circumference was alsomeasured to demonstrate that our assemblages werefully screened and collected [23,66].

4. Results

Mountain gazelle (Gazella gazella) was the mostcommon ungulate exploited by Epipalaeolithic foragers,although it is best represented in the Natufian assemb-lages (Table 1). Earlier assemblages (Nahal Hadera Vand Hefzibah) contain notably higher proportions of thelarger Persian fallow deer (Dama mesopotamica). Small-bodied prey such as tortoises (Testudo graeca), hare(Lepus capensis), and partridge (Alectoris chukar) alsofigure prominently in the assemblages, particularly inthe Natufian.

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Table 1

Summary of taphonomic and zooarchaeological variables for Nahal Hadera V (NHV), Hefzibah (HEF), Hayonim Cave–Early Natufian (HAY EN),

Hayonim Cave–Late Natufian (HAY LN), and el-Wad Terrace (EWT)

NHV KEB HEF G-KEB HAYC EN HAYC LN EWT LN

Summary of assemblages

NISP 19513 8507 8159 7354 2899

MNI 385 140 205 118 73

Gazelle NISP (MNI) 12528 (202) 6169 (83) 2527 (18) 1773 (18) 2095 (28)

Hare NISP (MNI) 474 (20) 239 (7) 1560 (26) 417 (9) 185 (6)

Partridge NISP (MNI) – 3 (1) 818 (65) 231 (22) 42 (5)

Ungulate evenness 0.5 0.4 0.6 0.6 0.1

% Gazelle in ungulate assemblage 68 76 89 90 97

% Small game in assemblage 4.2 3.6 61 64 17.4

Preservation of gazelles

Relationship between gazelle

bone survivorship and

bone density

y ¼ 1:03x� 0:29 y ¼ 0:83x� 0:15 y ¼ 0:83xC0:09 y ¼ 1:16x� 0:19 y ¼ 0:75x� 0:09

Spearman’s r density vs.

survivorship

rs ¼ 0:50; p ¼ 0:01 rs ¼ 0:46; p ¼ 0:02 rs ¼ 0:41; p ¼ 0:04 rs ¼ 0:65; p!0:001 rs ¼ 0:40; p ¼ 0:04

Relationship between gazelle

bone survivorship and food

utility index

y ¼ �0:004xC25:01 y ¼ �0:002xC13:4 y ¼ �0:005xC54:7 y ¼ �0:0005xC48:4 y ¼ �0:003xC40:6

Spearman’s r food value vs.

survivorship

rs ¼ 0:24; p ¼ 025 rs ¼ 0:10; p ¼ 0:60 rs ¼ 0:29; p ¼ 0:18 rs ¼ 0:37, p ¼ 0:07 rs ¼ �0:17; p ¼ 0:38

Tooth: cranial bone-based MNI 92 98 65 100 88

% Complete astragalus 75 79 87 88 79

% Complete central fourth tarsal 68 60 83 88 70

% Carnivore gnawed 0.1 0 0.2 0.1 0.4

% Rodent gnawed 0.1 0 1.5 1.5 0.3

% Weathered stage 3 or higher 13 12 0.6 0.7 3

% Fresh fracture angle 64 64 60 59 52

% Shaft fragments with impact

fractures

1.4 2.5 6.3 7 2.4

4.1. Tests of attrition

There is a significant, positive relationship betweengazelle bone survivorship (following [62, p. 239] andbone structural density (based on Lam et al.’s [52]BMD1 density values for Rangifer tarandus) in the fiveassemblages (Table 1). Lam et al.’s [52] bone mineraldensity 1 (BMD1) values accurately measure a bone’sexternal shape, but do not adjust for the interior volumeof marrow cavities. These values (Lam et al.’s [53] groupC) are preferred here since they provide the mostaccurate measures of a bone’s external shape, yet theymore closely match the values [77] used in ourforthcoming analysis of hare density since they also donot account for long bone internal cavities (Lam et al.’s[53] group B). The results of the density tests indicatea pronounced density-mediated bias in gazelle skeletalpart representation in every assemblage. In contrast,there is no significant relationship between bonesurvivorship and the food utility index (FUI) at any ofthe five sites (based on the weight of useable tissue ofRangifer tarandus; Table 1 [69]), thus selective transportof high-utility body parts was not a significant factor inassemblage formation. The specific relationships be-tween bone survivorship and marrow and grease yields

are discussed below. The absence of selective transportof high-utility body parts is further supported by gazellebody part profiles which show fairly uniform represen-tation of all body parts with the exception of un-derrepresented axial elements (vertebrae and ribs). Axialelements that are present are highly fragmented.

Overall, these results suggest that the representationof gazelle skeletal elements was most strongly influencedby density-mediated attritional processes that occurredprior to or following deposition in the sites. Density-mediated biases may arise from a variety of pre- andpost-depositional processes ranging from butchery tocarnivore ravaging and decomposition [50,60,62]. Notall taphonomic processes are expected to affect all taxaequally. For example carcass processing is likely to varyfrom species to species, whereas postdepositional pro-cesses like chemical dissolution may not discriminatebetween taxa. To assess whether the density-mediatedbias is species-specific or characteristic of the assemb-lages in general, we compared bone survivorship withbone density in a small game species (hare) common inEpipalaeolithic assemblages. Hare is the only Epipa-laeolithic non-ungulate species for which comparabledensity values are available (based on Lepus californicus)[77], but only the Early and Late Natufian hare samples

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from Hayonim Cave are large enough for analysis. Weuse Pavao and Stahl’s [77] shape adjusted volumedensity measures (VDSA) (Lam et al.’s [52] Group B]which are corrected for surface shape, but not shaftvolume. The surfaces of the hare bones were adjustedusing a geometric model which is not as accurate asvalues measuring using computed tomography (CT)which was used to capture the reindeer BMD1 valuesused above. Although they are not directly comparableto the measures used in our gazelle tests they are theclosest available [52, Fig. 2]. Unlike the density tests forgazelle, the relationship between hare bone density andsurvivorship is not significant in the Early (p ¼ 0:16;rs ¼ 0:31) and Late Natufian (p ¼ 0:46; rs ¼ 0:17) fromHayonim Cave (Fig. 2). This suggests that the twoanimals experienced independent taphonomic historiesin the same depositional setting. The destruction ofgazelle bone is likely attributable to pre-depositionalfactors that are more likely than processes of in situattrition to impact taxonomic groups or body sizesdifferentially.

We explore the preceding using two measures of insitu bone attrition. First, we use Stiner’s [89] tooth- tobone-based MNI to examine the relative representationof elements with different structural densities. Thetooth- to bone-based ratio measures preservation ona scale from zero to onedthe closer the ratio is to one,the better the quality of preservation. The Epipalaeo-lithic assemblages all display high tooth to cranial boneratios (ranging from 0.65 to 1.00; Table 1), indicatingminimal loss of identifiable bones by decompositionand/or advanced fragmentation. Similar results wereobtained using Marean’s [64] completeness index whichinvestigates the quality of preservation of high-density

elements including gazelle astragali and central fourthtarsals (navicular-cuboid). The percentage of completeastragali and central fourth tarsals ranges from 60–70%at the open-air sites of Hefzibah and Nahal Hadera V to92% in the Early Natufian assemblage from HayonimCave (Table 1). These results attest to good preservationof all studied assemblages, and demonstrate that post-depositional sources of in situ attrition played a minorrole in the formation of the Epipalaeolithic assemblages.Weathering beyond Behrensmeyer’s [10] stage 2 (ofstages 0–5) is most widespread in the open air sites ofNahal Hadera V and Hefzibah (!13%; Table 1), but inno case compromises specimen identifiability.

Fluvial action can cause density-mediated biases bywinnowing low-density bones such as vertebrae and ribsout of archaeological assemblages [105]. We find noevidence for fluvial action in our assemblages. Forexample, although low-density transportable bones aremissing, the high-density astragalus is also easily trans-ported by water, but is one of the best representedelements in our assemblages. None of the bone edgesfrom Nahal Hadera V, el-Wad, and Hayonim Cave andonly 2.6% of bone edges from Hefzibah display round-ing and/or smoothing of their break surfaces [84],suggesting that rolling caused by water transport wasinsignificant. Furthermore, evidence for striationscaused by abrasion [51] is rare to non-existent in allassemblages. These results are in accordance with theabsence of sedimentological evidence for fluvial trans-port in the coastal archaeological sites [82].

Non-human biotic agents had similarly low impactson the assemblages. Rodent gnawing reaches a maxi-mum frequency of 1.5% in all assemblages and evidencefor carnivore activity (i.e., scoring, gnawing, digestion)

y=0.83x + 0.09; rs=0.41; p=0.04 y=1.16x - 0.19; rs=0.65; p<0.001HAYC EN

100806040200

0 0.2 0.4 0.6 0.8 0 0.2 0.4 0.6 0.8

Bone density

0.0 0.2 0.4 0.6

Bone density0.0 0.2 0.4 0.6

Bone density

Bone density

Surv

ivor

ship

100

80

60

40

20

0

Surv

ivor

ship

100

80

60

40

20

0

Surv

ivor

ship

100806040200

Surv

ivor

ship

HAYC LN

y=0.54x + 0.36; rs=0.31; p=0.16 y=0.07x + 0.40; rs=0.17; p=0.46HAYC EN HAYC LN

Fig. 2. Relationship between bone density and bone survivorship (%MNI) in gazelle (top) and hare (bottom) from Early and Late Natufian Hayonim

Cave.

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occurs on no more than 0.4% of gazelle bones in anyassemblage (Table 1). The near absence of carnivoredamage may be surprising given the causal role it hasbeen assigned to explain patterns of density-mediatedattrition in many Palaeolithic assemblages [78 andreferences therein]. Carnivore damage is uncommon inLevantine assemblages [80] (and see discussion in [79]),particularly those dating to the Epipaleolithic periods.This likely results from one or a combination of factorsincluding the composition of the carnivore community,intensified site use, and cooking practices that extractdesirable fats from bone prior to disposal. Regardless, itis very interesting and significant that given the virtualabsence of carnivore activity, and the minor role of insitu attrition, all five of the Epipalaeolithic assemblagesshow significant density-mediated biases.

The morphology of the fracture angle, fractureoutline, and fracture edge (following [104]) wererecorded for all shaft fragments that were connected toa portion of a gazelle long bone epiphysis. Shaftcircumference data was recorded following Bunn [22]from an additional sample from Nahal Hadera V,Hefzibah, and el-Wad Terrace, and included fragmentsfrom both the midshaft and those attached to thearticular ends. Comparable data were not available fromthe Hayonim Cave assemblages. Our fracture data werecompared to data derived from two French Neolithicsites studied by Villa and Mahieu [104]. The sitesprovide good comparisons since one assemblage wasfractured when fresh and the other was broken whendry, and their taphonomic histories were independentlyassessed using multiple lines of evidence. Our fiveEpipaleolithic assemblages closely match the breakagepatterns of the butchered human remains from Font-bregoua, but differ substantially from those from theSarrians burials which were crushed in situ after thebones had dried. The shaft fragments in our assemblagesare dominated by high frequencies of oblique, V-shaped,and jagged breaks (Table 2)dfractures that Villa and

Mahieu’s [104] study and actualistic experiments haveshown to occur when bone is broken while fresh [36,76].Only a small percentage of long bone shafts exhibitevidence of dry breaks (i.e., transverse, right-angledbreaks with uneven break surfaces). Most of thefragmentation of the Epipalaeolithic assemblages oc-curred when the bones were still freshdeither just priorto deposition or shortly thereafter [76,104]. Thefrequency of green long bone fractures points to pre-depositional human activities as the primary source ofgazelle bone fragmentation rather than post-deposition-al attrition. In addition, the shaft circumference datafrom the three open-air sites indicate that the majorityof shaft fragments are represented by less than half ofthe complete shaft circumference (68–71%), and a shaft’sfull circumference is rarely complete (0.5–16.5%). Thesevalues are in accordance with assemblages that werefully screened and collected [66].

The results seem to eliminate post-depositional forcesas major processes in the formation of the Epipalaeo-lithic assemblage and point to humans as the primaryagent of bone breakage. To further test this assertionand to explore the nature and importance of humanbone-breaking activities, we performed a series of teststo determine whether patterns of breakage and body-part representation are consistent with marrow process-ing, grease extraction, both, or neither. All of these testsassume that marrow and grease processing will prefer-entially fragment and destroy bone portions with highfat contents regardless of bone density.

4.2. Tests for marrow processing

Marrow indices are not available for gazelle so weused the closest available analogue, Binford’s [14] valuesfor domestic sheep, to explore variation in bonefragmentation and survivorship across the gazelleskeleton. First, we examined the relationship betweenthe marrow index and the intensity of bone shaft

Table 2

Frequencies of fresh and dry fracture angles, fracture outlines, fracture edges, and shaft circumferences for gazelle long bone shafts from the

Epipalaeolithic assemblages

Fracture angle (%) Fracture outline Fracture edge Shaft circumference

Oblique

(fresh)

Right

(dry)

V shaped

(fresh)

Transverse

(dry)

Jagged

(fresh)

Smoothed

(dry)

Less

than !1/2

More

than O1/2

Complete

Nahal Hadera V Kebaran 83 (74) 30 (26) 82 (68) 38 (32) 102 (82) 23 (18) 89 (68) 24 (21) 13 (11)

Hefzibah Geometric Kebaran 121 (80) 30 (20) 117 (78) 33 (22) 145 (86) 23 (14) 174 (91.1) 16 (8) 1 (!1)

Hayonim Cave Early Natufian 55 (87) 8 (13) 55 (86) 9 (14) 78 (86) 13 (14) n.d. n.d. n.d.

Hayonim Cave Late Natufian 54 (84) 10 (16) 39 (80) 10 (20) 72 (80) 18 (20) n.d. n.d. n.d.

el-Wad Terrace Late Natufian 78 (58) 57 (42) 77 (59) 53 (41) 85 (65) 45 (35) 113 (72) 18 (11) 26 (17)

Fontbregoua 114 (71) 47 (29) 134 (59) 92 (41) 163 (62) 98 (38) 115 (76) 23 (15) 13 (9)

Sarrians 22 (11) 176 (89) 106 (35) 193 (65) 111 (31) 247 (69) 16 (7) 10 (4) 200 (89)

Comparative data are provided from two French Neolithic sites with known taphonomic historiesda sample of butchered human remains from

Fontbregoua and a sample of human bones from crushed after drying from Sarrians [104]. Relative percentages are given in parentheses. Comparable

data were not available for Hayonim Cave (n.d.=no data).

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fragmentation. A fragmentation index was derived formarrow-bearing bones (long bone shafts, mandible,phalanx 1 and 2, and calcaneum) by calculating theNISP:MNE ratio for each element. We found significantand positive correlations between the fragmentation ofthese skeletal elements and their respective bone marrowindex values in all five assemblages (Fig. 3). The longbone shafts encasing the largest marrow stores (distaltibia shaft, proximal radius shaft, and proximal meta-carpal and metatarsal shafts) are the most highlyfragmented, while elements containing the smallestmarrow stores (the first and second phalanges) havethe lowest fragmentation indices.

A second way to investigate the relationship betweenfragmentation and marrow content is to compare therelative completeness of adult and juvenile gazelle longbone shafts. As an animal ages the primary function ofthe marrow cavity shifts from red blood cell productionto fat storage. Red marrow which is low in fat content isgradually replaced by fatty yellow marrow as the animalmatures [16,29]. Juvenile bone is also spongier and lessdense than adult bone and becomes more mineralized asthe animal approaches adulthood. It has been estab-lished that juvenile bone is more subject to breakage andattrition by post-depositional agents than adult bone

[72]. If post-depositional processes are the major causeof bone breakage, juvenile bone should therefore, bemost severely affected. In contrast, if marrow extractionis the major cause, we expect fewer complete adult longbone shafts. This test compares the proportion ofcomplete juvenile with complete adult gazelle long boneshafts. It is performed only for the Natufian layers atHayonim Cave which contain substantial quantities ofjuvenile gazelles (O33%) [71]. In general, the faunalassemblages from the Natufian layers at Hayonim Caveare highly fragmented (gazelle average fragment lengthis 2.7 cm for Early Natufian and 2.8 cm for LateNatufian), and thus complete long bones are rare (EarlyNatufian NISP=40, Late Natufian NISP=28). Longbone shafts are considered complete if at least 90% ofthe shaft is present, this enables bones that are onlyslightly eroded or broken to be included in the completegroup to ensure that they are distinguished from brokenbones that were breached for bone marrow. Themajority of complete gazelle long bone shafts in theassemblage belong to juveniles (72.5% in the Early and60.0% in the Late Natufian). Fragmented gazelle longbone shafts show the opposite patterndonly 21.4% and27.2% belong to juveniles. In sum, most broken gazellelong bone fragments belong to adults whereas most

y=0.025x + 1.60; rs=0.63; p<0.05 y=0.031x + 1.80; rs=0.84; p<0.001NHV

0

2

4

6

0 20 40 60 80 100 0 20 40 60 80 100

Marrow Index Marrow Index

0 20 40 60 80 100 0 20 40 60 80 100

Marrow Index

0 20 40 60 80 100

Marrow Index

Marrow Index

HEF

0123456

y=0.017x + 1.67; rs=0.54; p=0.05 y=0.029x + 1.02; rs=0.80; p<0.001 HAYC EN

0

2

4

6

HAYC LN

0

2

4

6

y=0.043x + 1.05; rs=0.80; p<0.001 EWT

0123456

NIS

P/M

NE

NIS

P/M

NE

NIS

P/M

NE

NIS

P/M

NE

NIS

P/M

NE

Fig. 3. Relationship between the marrow index [14] and the fragmentation (NISP/MNE) of gazelle long bone shafts in the five Epipalaeolithic

assemblages.

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complete shafts belong to juveniles. In this case bonebreakage corresponds to marrow content but not tobone density, and implies that adult bones wereintentionally selected and broken for their marrow whilejuvenile bones were neglected. Preservation of completejuvenile long bone shafts strengthens the argument thatthe breakage of adult shafts did not result from density-mediated post-depositional processes.

Gazelle phalanges 1–3 and tarsals (astragalus andcalcaneum) contain little if any marrow, and thusprovide a simple gauge of the thoroughness ofEpipalaeolithic processing. Table 3 presents the extentof fragmentation (percent of complete elements) [62, p.333–334] of gazelle compact foot bones that havevariable marrow contents. Extent of fragmentation(COMP) was calculated as the number of wholeelements represented relative to each bone’s MNE. Toensure that bones that were eroded or had recent nicksand breaks were included, a bone is considered completewhen at least 90% of its original mass is intact. In allassemblages extent of fragmentation correlates roughlywith marrow content. The first phalanx has the highestmarrow content and is complete least often (on average40.2% for all assemblages), while the second phalanxand the calcaneum with moderate marrow content haveintermediate completeness values (62.3% and 49.8%),and the third phalanx and astragalus which contain nomarrow are complete most often (74.6% and 81.6%).The low proportion of complete first phalanges suggeststhat although small, this source of bone marrow wasregularly tapped (see [30] for an alternate explanation offoot bone preservation). Less effort was invested incracking the second phalanx and calcaneum whichcontain less marrow. Data on impact fractures was notcollected from the gazelle phalanges. Compellingevidence that the fracture of first and second phalangeswas driven by marrow extraction is instead provided bycomparing the frequency of complete (O90% intact)juvenile and adult toes in the Hayonim Cave assemb-lages. In both the Early and Late Natufian assemblages

the combined sample of juvenile first and secondphalanges are complete more than 80% of the time(juvenile MNE for Early Natufian=73, Late Natu-fian=57). In contrast, adult first and second phalangeswere complete less than 65% of the time (adult MNE forEarly Natufian=139, Late Natufian=127). Juvenilemarrow-bearing toes are much less dense yet arecomplete substantially more often than their adultcounterparts suggesting that the denser adult boneswere broken to extract bone marrow.

Another test for marrow exploitation compares theproportion of complete gazelle long bone shafts to thosefrom smaller animals that yield either less (hares) or nobone marrow (partridges). Degree of shaft completenessrefers to the proportion of marrow-bearing long boneshafts that are more than 90% complete. These tests areperformed only for the Hayonim Cave assemblageswhich contain adequate samples of small game. There isa significant difference in the completeness of long boneshafts between the three groups in both the Early andLate Natufian phases (Kruskal–Wallis H ¼ 17:91,p!0:001). Gazelle long bone shafts are complete ca.18% of the time, whereas at least 30% of hare and 70%of partridge long bone shafts are complete (Table 4).This result confirms once again that bones with moremarrow are more fragmented than those with less,despite greater fragility of the non-marrow bearingbones.

Finally, impact fractures typical of human percussionappear on 1.4–7.0% of gazelle long bone shafts (Table1). The frequency of impact fractures per skeletalelement correlates with marrow content in gazelle longbones from the Hayonim Cave assemblages (rs ¼ 0:636,p!0:05, n ¼ 10) [71] indicating once again, that humanspreferentially butchered bones with the highest marrowcontent.

All of the tests indicate significant correlationsbetween bone fragmentation and marrow content. Whatmakes this result compelling is that fragmentation doesnot correlate with bone density in any of these cases, as

Table 3

Percent completeness (COMPZMNE bones at least 90% complete/total MNE) of gazelle compact foot bones in the five Epipalaeolithic assemblages

Phalanx 1 Phalanx 2 Calcaneum Phalanx 3 Astragalus

(33.77) (25.11) (23.11) (0) (0)

Nahal Hadera V COMP/MNE 165/840 451/882 70/173 518/663 402/535

% Completeness 29.7 51.1 40.5 78.1 75.2

Hefzibah COMP/MNE 40/170 171/311 26/69 162/226 79/201

% Completeness 23.5 55.0 37.7 71.7 79.2

Hayonim Cave Early Natufian COMP/MNE 70/120 86/103 24/36 69/81 27/31

% Completeness 58.3 83.5 66.7 85.2 87.1

Hayonim Cave Late Natufian COMP/MNE 55/89 71/92 15/24 49/65 21/24

% Completeness 61.7 77.2 62.5 75.4 87.5

el-Wad Terrace COMP/MNE 23/83 41/92 5/12 30/48 50/64

% Completeness 27.7 44.6 41.7 62.5 78.9

Average % completeness of all assemblages 40.2 62.3 49.8 74.6 81.6

Marrow values are in parentheses and are based on Binford’s [14] values for domestic sheep.

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Table 4

Percentage of complete (O90% complete) gazelle, hare, and partridge long bone shafts from the Early and Late Natufian layers at Hayonim Cave

Complete long bone shafts % Gazelle % Hare % Partridge

HAYC EN HAYC LN HAYC EN HAYC LN HAYC EN HAYC LN

Humerus shaft 10 (20) 0 (27) 11 (38) 0 (9) 100 (18) 100 (11)

Radius shaft 0 (8) 7 (14) 12 (34) 53 (17) 50 (2) 50 (2)

Metacarpal shaft 38 (8) 38 (8) 66 (92) 48 (25) 80 (5) 33 (3)

Femur shaft 38 (8) 20 (5) 7 (15) 25 (12) 100 (13) 0 (1)

Tibia shaft 0 (20) 14 (7) 46 (28) 56 (9) 100 (16) 71 (7)

Metatarsal shaft 13 (15) 50 (10) 46 (77) 30 (23) 100 (11) 100 (3)

Numbers in parentheses represent the MNE of each shaft. Juvenile and fetal gazelle bones were removed from the analysis since these bones contain

little or no fat-rich yellow marrow. For partridge the metacarpal and metatarsal categories refer to the carpometacarpus and the tarsometatarsus.

would be expected if attritional processes were re-sponsible. Even in cases where non-marrow bearingbones are significantly more fragile than marrow-bearing bones, the fragile bones are less fragmented.Furthermore, juvenile gazelle bones which lack yellowmarrow are consistently more complete than their adultcounterparts, despite their substantially lower bonemineral density and greater fragility. This confirms thatdespite the overriding pattern of density-mediatedattrition, culturally meaningful sub-patterns can stillbe detected in the data. Results of the four tests stronglysupport the interpretation that Epipalaeolithic foragersroutinely processed gazelle long bone cavities formarrow and invested much effort to process elementswith the highest marrow yields.

4.3. Tests for grease processing

Grease processing is more difficult to detect in thearchaeological record since grease concentrates in low-density cancellous bone, and thus its extraction canmimic the signatures of other agents of density-mediatedattrition. Additional tests are performed to see if greaseprocessing can be distinguished from other taphonomicforces in the Epipalaeolithic assemblages. Althoughother taphonomic processes may produce similarsignatures, bones processed for grease are expected tobe highly fragmented and to have low survivorship incomparison to those that are not. The following analysiscompares the %survivorship of grease-rich, cancellousgazelle bones against their fragmentation ratios (NISP/MNE). Long bone shafts were excluded from theanalysis to eliminate the potentially confounding effectsof medullary marrow processing. Fig. 4 shows a strongnegative relationship between %survivorship and frag-mentation in all five Epipalaeolithic assemblages. Thissuggests that cancellous bones may be underrepresentedin gazelle assemblages because they were alreadyfragmented when they were deposited in the archaeo-logical record, possibly because they were processed forgrease or stewed.

4.4. Intensification

Evidence indicates that Levantine Epipalaeolithicforagers intensively utilized gazelle carcasses, yet welack the comparative data to pinpoint when this began.Furthermore, the preceding results reveal no temporalchanges in gazelle carcass use across the Epipalaeolithicperiod itself. In all cases, the test results from the fiveassemblages are similar and provide uniform answersdall results of a test are statistically significant or all arenot. Pinpointing the beginning of intensified carcassprevalent throughout the Epipalaeolithic examples is animportant avenue for further research.

5. Discussion

Four conclusions can be drawn from our multivariateanalyses. First, attrition in the Epipalaeolithic gazelleassemblages is clearly density-mediated, although this isnot the case for the smaller-bodied hare, at least atHayonim Cave. Second, density-mediated bone loss isunlikely to have been the consequence of post-de-positional attrition or abiotic post-depositional factors.Third, patterns of bone survivorship in the gazelleassemblages were most likely created by pre-deposition-al processing by humans. In particular, breakage andattrition of long bone shafts is attributable to marrowprocessing, whereas grease extraction or cooking likelyaccounts for patterns of cancellous bone survivorship.We conclude that human processing of bones formarrow and possibly grease is the primary determinantof bone survivorship in the five Epipalaeolithic assemb-lages. The case for medullary marrow extraction isstrong and in the case of long bone shafts is supportedby significant correlations between fragmentation andmedullary marrow content, but not with bone densityand fragility. Fourth, although we lack data to comparethe intensity of fat extraction with the Palaeolithicrecord, there is no evidence for intensified fat extractionduring the Epipalaeolithic.

The case for grease processing is less conclusive.What makes the argument for grease extraction

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NHV

0

2

4

6

8

0 20 40 60 80 100

Survivorship

0 20 40 60 80 100

Survivorship

0 20 40 60 80 100

Survivorship

0 20 40 60 80 100

Survivorship

0 20 40 60 80 100

Survivorship

HEF

0

2

4

6

8

HAYC EN

0

2

4

6

8

HAYC LN

0

2

4

6

8

el-Wad Terrace

0

2

4

6

8

NIS

P/M

NE

NIS

P/M

NE

NIS

P/M

NE

NIS

P/M

NE

NIS

P/M

NE

Fig. 4. Relationship between gazelle bone survivorship (%MNI) and fragmentation (NISP/MNE) in the five Epipalaeolithic assemblages.

compelling, at least by the Natufian, is the taxon-specificnature of the density-mediated bias. It applies to gazelle,but not to hare, and thus does not characterize thecomplete assemblage. If we assume that prey body size isnot a mediating factor (mean body mass of gazelle is 18kg for females and 25 kg for males) [68] density-mediated bias cannot be attributed to processes of insitu attrition or pre-depositional abiotic factors that areexpected to operate according to bone density regardlessof taxonomic affiliation. Some other process must bepreferentially destroying the low-density cancellousbones of the gazelle skeleton. Because grease is storedin cancellous bone, its extraction should producea pattern of density-mediated attrition, but only inlarge animals with sufficient grease yields, such asungulates. Grease from smaller animals such as haresmay also have been extracted as a byproduct of stewing,but smashing larger bones to extract grease suggestsa more specialized form of grease rendering. Bonesprocessed for grease should also be characterized byhigh fragmentation indices and low survivorship as isthe case in the Epipalaeolithic assemblages. Post-de-positional processes such as trampling and other formsof mechanical loading may fragment cancellous bone,but unlike grease extraction these processes are expectedto act on bones according to their density rather thantheir taxonomy. Our assemblages are dominated bygreen fractures which fit with a hypothesis of grease

processing better than one of post-depositional break-age. Finally, grease-rich vertebrae are also missing fromthe gazelle assemblages. We do not have good evidencefor comminution, but given that bones were not storablein the warm Mediterranean climate, we do not expectEpipalaeolithic grease extraction to resemble the classicethnographic cases in which bones were stored and thencollectively processed for grease [14].

The archaeological record is largely silent on thecharacter of Epipalaeolithic grease extraction. Certainlymilling technology capable of processing bones forgrease was widespread and abundant in the Epipalaeo-lithic [108]. Beyond this, it is quite possible thatEpipalaeolithic humans used boiling technology to cookmeat and other foods. Ethnoarchaeological reportscommonly describe cooking meat and animal bones byadding them to liquids in organic containers (see [14],and references in [43,49]). The contents were heatedeither through the addition of fire-heated stones or bysuspending an organic container over an active hearth[43]. Although, it is common elsewhere in the Mediter-ranean basin, fire cracked rock is rare in the Epipalaeo-lithic record of this region, and although Epipalaeolithicforagers were undoubtedly capable of boiling, there isno clear evidence for it in the Levantine sites. Organiccontainers that would provide equally good evidence forprehistoric cooking activities rarely preserve in thearchaeological record. Still, it is likely that they were

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commonplace in the Upper Palaeolithic and Epipalaeo-lithic periods, especially in light of preserved fibers andrich bone tool kits [9,73,74]. We hypothesize that foodpreparation using milling technology and cooking in hotliquids played a major role in Epipalaeolithic boneassemblage formation, and was a primary source ofdensity-mediated bias in our assemblages.

Our data provide strong evidence that Epipalaeo-lithic foragers intensively exploited gazelle carcasses toextract bone marrow and possibly grease. When thisintensive use began and if it in fact represents anintensification trend, however, is a trickier question toaddress. Unlike other intensification trends that culmi-nate in the Natufian period, bone processing showsgreat stability across the Epipalaeolithic. We requirecomparative data from additional Palaeolithic sites inthe region to comprehensively assess the process ofintensification itself. We believe that grease was beingexploited by Levantine Epipalaeolithic foragers, butwhen it first began is especially relevant for intensifica-tion studies because grease processing (a) is laborintensive and the returns are often insubstantial, and(b) fat may have been a factor limiting humanpopulation size during lean seasons. Like other in-tensification trends, the intensive use of gazelle carcassesin the Epipalaeolithic may be related to pulses ofpopulation growth [96,97] leading up to the transitionto agriculture.

Acknowledgements

We thank Yin Lam, Lee Lyman, Travis Pickering,John Speth, andMary Stiner for their careful reading andthoughtful comments on an earlier draft which greatlyimproved the paper. We also thank Ofer Bar-Yosef, ReidFerring, Nigel Goring-Morris, and Daniel Kaufman forstimulating discussions on the archaeological evidence forgrease production. GBO’s research was supported by theIrene Levi Sala CARE Archaeological Foundation andwas carried out when GBO was a Clore Doctoral Fellowat the Department of Zoology, Tel-Aviv University.NDM’s research was supported by grants from theNational Science Foundation (Dissertation ImprovementGrant SBR-9815083), the Levi Sala CARE Archaeolog-ical Foundation, and doctoral fellowships from the SocialSciences and Humanities Research Council of Canada(SSHRC), and the Department of Anthropology at theUniversity of Arizona (Haury Dissertation Fellowship).

Appendix A. Number of identified specimens (NISP)

and minimum number of elements (MNE) of gazelle

elements and bone portions from the five

Epipalaeolithic sites

Bones that contained shaft and articular end frag-ments were included in the MNE calculations for eachcategory (Table A.1).

Table A.1

NISP and MNE of gazelle bone elements at the Epipalaeolithic sites of Northern Israel

NHV KEB HEF G-KEB HAYC EN HAYC LN EWT LN

NISP MNE NISP MNE NISP MNE NISP MNE NISP MNE

Head

Horn 50 7 84 10 29 7 62 20 20 4

Skull 235 138 124 61 66 11 62 9 66 23

Mandible 173 119 87 50 66 15 47 7 35 14

Teeth upper 429 50 26 251 96 17 67 9 105 16

Teeth lower 103 947 564 52 101 13 73 11 107 16

Body

Vert atlas 5 2 8 5 16 4 7 3 2 2

Vert: axis 19 4 8 6 14 3 4 2 7 2

Vert: cervical 115 19 63 12 64 20 31 4 38 11

Vert: thoracic 81 28 36 12 71 20 26 9 30 16

Vert: lumbar 260 34 120 24 97 13 36 11 92 25

Vert: caudal 10 6 4 1 7 6 2 2 6 5

Sacrum 0 0 0 0 5 2 1 1 0 0

Sternum 0 0 0 0 3 1 8 1 2 1

Rib 154 68 100 11 218 48 147 54 74 23

Forelimb

Scapula-glen.fos 89 52 56 31 9 9 14 11 21 21

Scapula-blade 36 8 26 7 18 4 17 4 25 6

Humerus-prox 47 15 26 9 11 4 9 5 12 7

Humerus-dist 220 119 111 64 50 36 44 36 42 27

Humerus-shaft 54 5 24 3 40 20 46 27 25 3

Radius-prox 193 71 125 48 25 14 38 22 35 10

Radius-dist 115 82 63 33 15 9 13 7 11 7

(continued on next page)

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Table A.1 (continued)

NHV KEB HEF G-KEB HAYC EN HAYC LN EWT LN

NISP MNE NISP MNE NISP MNE NISP MNE NISP MNE

Radius-shaft 22 3 22 3 44 13 48 18 14 2

Ulna-prox 91 44 36 15 22 17 30 24 16 7

Metacarpus-prox 125 45 58 28 26 16 25 14 13 7

Metacarpus-shaft 35 9 24 3 37 12 37 12 5 2

Carpal-cuneiform 184 172 113 109 22 22 5 5 30 30

Carpal-lunate 150 137 59 57 14 12 6 6 17 13

Carpal-magnum 193 173 105 102 8 8 7 6 34 27

Carpal-scaphoid 260 250 160 149 27 27 12 12 30 25

Carpal-unciform 195 185 105 101 13 13 7 7 22 22

Hindlimb

Pelvis-ilium 31 20 37 23 14 11 12 10 17 11

Pelvis-isch 69 15 44 8 14 11 7 4 28 4

Pelvis-pubis 112 14 36 12 5 5 12 12 8 7

Pelvis-acetabulum 211 150 117 73 24 12 27 10 52 37

Femur-prox 146 95 96 89 27 18 9 3 32 28

Femur-dist 99 23 52 18 23 9 16 8 36 9

Femur-shaft 45 4 15 2 43 11 18 6 20 1

Tibia-prox 11 4 7 2 30 15 12 8 3 1

Tibia-dist 229 119 123 50 49 34 24 10 43 26

Tibia-shaft 62 9 34 4 94 23 43 7 18 2

Patella 142 125 109 87 18 18 13 13 33 28

Astragalus 538 403 249 166 32 31 29 24 64 51

Calcaneum 368 190 176 79 50 36 31 24 30 13

Navicular cuboid 165 121 91 54 23 19 19 17 24 17

Ext.Cmid. cuneiform 299 290 163 160 8 8 2 2 44 41

Lateral malleolus 141 137 83 82 7 7 3 3 20 20

Metatarsus-prox 167 57 69 23 40 23 25 15 28 9

Metatarsus-shaft 50 5 14 2 72 19 49 13 5 2

Toes

Phalanx 1 1611 940 647 339 234 120 151 89 211 99

Phalanx 2 1308 888 507 293 145 103 122 92 158 93

Phalanx 3 695 632 243 213 92 81 74 65 61 50

Seasamoid 393 385 214 208 37 37 19 19 69 69

Metapod-dist 1561 605 701 509 149 71 132 57 166 57

Metapod -shaft 62 9 42 5 130 28 99 28 17 2

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