Taphonomy and zooarchaeology of the Upper Palaeolithic cave of Dzudzuana, Republic of Georgia

Taphonomy and Zooarchaeologyof the Upper Palaeolithic Cave ofDzudzuana, Republic of Georgia

G. BAR-OZ,a* A. BELFER-COHEN,b T. MESHVELIANI,c

N. DJAKELIc AND O. BAR-YOSEFd

a Zinman Institute of Archaeology, University of Haifa, Mount Carmel 31905, Haifa, Israelb Institute of Archaeology, Hebrew University, Jerusalem 91905, Israelc Georgian State Museum, Department of Archaeology, Tbilisi 380007, Republic of Georgiad Department of Anthropology, Peabody Museum, Harvard University, Cambridge, MA

02138, USA

ABSTRACT We present the results of a detailed taphonomic and zooarchaeological study of the faunalremains from the Upper Palaeolithic layers of Dzudzuana Cave, Republic of Georgia. Thisstudy presents the first carefully analysed Upper Palaeolithic faunal assemblage from thesouthern Caucasus and thus serves as a significant point of reference for inter-regionalstudies of Upper Palaeolithic subsistence in Eurasia. A series of intra-site taphonomiccomparisons are employed to reconstruct the depositional history of the bone assemblageswithin the different occupational phases at the site and to investigate subsistence, meatprocurement and bone-processing strategies. Caucasian tur (Capra caucasica), aurochs(Bos primigenius) and steppe bison (Bison priscus) were the major prey species throughoutthe Upper Palaeolithic. Their frequencies do not change significantly over time, and nor doesbone preservation vary by layer. The assemblage is characterised by significant density-mediated biases, caused by both human bone-processing behaviours and in situ post-burialbone attrition. Bone marrow extraction produced large numbers of unidentified bone frag-ments, many exhibiting green bone fractures. The density and size of bone assemblages andthe extent of fragmentation indicate that Dzudzuana Cave was repeatedly occupied by UpperPalaeolithic foragers over many years. Skeletal part representation and butchery marks fromall stages of carcass processing suggest that prey occasionally underwent field butchery.Intra-site taphonomic comparisons highlight uniform patterns of cultural and economicbehaviours related to food procurement and processing strategies. Copyright � 2007 JohnWiley & Sons, Ltd.

Key words: taphonomy; zooarchaeology; Caucasus; Upper Palaeolithic; subsistence;

hunting; Republic of Georgia

Introduction

This present paper deals with Upper Palaeolithicsubsistence in the southern foothills of theCaucasus Mountains in Western Georgia (i.e.

Imeretia). The region occupies an intermediateposition between Europe and Asia and is boundedby the Caucasus Mountains to the north, theBlack Sea to the west, and the Caspian Sea to theeast. The influence of the Black Sea fostersfavourable climatic conditions that are charac-terised by a mild and warm humid climate withample rainfall (average annual precipitationranges from 800–1600mm), while the massive

International Journal of OsteoarchaeologyInt. J. Osteoarchaeol. 18: 131–151 (2008)Published online 16 July 2007 in Wiley InterScience(www.interscience.wiley.com) DOI: 10.1002/oa.926

* Correspondence to: Zinman Institute of Archaeology, Universityof Haifa, Mount Carmel 31905, Haifa, Israel.e-mail: [email protected]

Copyright # 2007 John Wiley & Sons, Ltd. Received 15 October 2006Revised 28 February 2007Accepted 7 March 2007

ridges of the Caucasus deflect the cold polar airflowing from the Eurasian plains. Southward-flowing streams have cut numerous deep rivervalleys and gorges and formed diverse ecologicalniches (Volodicheva, 2002). These conditionspromote a high degree of floral and faunaldiversity which supported large Upper Pleisto-cene populations (e.g. Vereshchagin, 1967;Liubin, 1989; Lordkipanidze, 1992, 1998). Atthe base of the foothills the landscape ischaracterised by open steppe and grasslands,with gallery forests along the rivers. Broadleafforest is found on the lower slopes, while mixedforest, coniferous forest and alpine meadows arefound at successively higher elevations (Volo-dicheva, 2002). Our excavations and zooarch-aeological investigations in this rich environmentwere aimed at investigating the foraging beha-viours and landscape use of prehistoric peoples(Meshveliani et al., 1999, 2004; Bar-Oz et al., 2004;Bar-Oz & Adler, 2005; Adler et al., 2006).Western Georgia, located between the Cau-

casus range, the Likhi hills and the Black Sea, is aregion known for its wealth of prehistoric sites,most of which are found in the river valleys thatdrain the mountains. Past research established acultural as well as a palaeoenvironmental recordfor the area, commencing with the LowerPalaeolithic up to present times (Vereshchagin,1967; Liubin, 1989, and references therein;Liubin & Bosinski, 1995). These records under-went some changes based on the results of recentinvestigations (Meshviliani et al., 1999, 2004;Adler, 2002; Adler & Tushabramishvili, 2004; andreferences therein). While the archaeologicalremains were reported through the use ofcommon typological terminology, faunal studieswere conducted solely as palaeontological inves-tigations, resulting in lists of presence/absence ofspecies, without any zooarchaeological or tapho-nomic considerations except for some LowerPalaeolithic assemblages (Tappen et al., 2002a,b;Vekua et al., 2002).A Georgian–American–Israeli joint project

centres on the excavations of three sites inWestern Georgia: Dzudzuana Cave and therock-shelters of Ortvale Klde and Kotias Klde.The sites are located within the same ecologicaland geographical setting and provide a culturalsequence from the LateMiddle Palaeolithic to the

Late Mesolithic and Neolithic. The Middle andUpper Palaeolithic bone assemblages of OrtvaleKlde were recently fully published (Bar-Oz &Adler, 2005; Adler et al., 2006) and theexcavations in the Late Upper Palaeolithic andMesolithic site of Kotias Klde are still underway(Meshveliani et al., in press).

The present paper provides a full taphonomicand zooarchaeological documentation of thefaunal remains from the Upper Palaeolithicdeposits of Dzudzuana Cave (1996–2004 exca-vation seasons). Our goals are to reconstruct thedepositional history of the bone assemblageswithin the different occupational phases at thesite, and to investigate their economic structuresin terms of diet breadth and prey procurement,transport and processing. This study presents thefirst carefully analysed Upper Palaeolithic faunalassemblage from the southern Caucasus, and thusserves as a significant point of reference forinter-regional studies of Upper Palaeolithicsubsistence in Eurasia.

The site and its setting

Dzudzuana Cave is located in the Nekressi rivervalley, a tributary of the Kvirila River (Figure 1). Itis situated at approximately 560 m above sealevel, 12 m above the river channel. The cave is alarge, elongated karstic cavity that emerges as atunnel from which a small creek flows on top ofthe deposits. The first series of excavations(1966–1975) was carried out by D. Tushabra-mishvili on behalf of the Georgian State Museum.Excavations were conducted in units of 10 cmover one square metre and the finds werehand-picked without the use of water sieving.The stratigraphy recorded by Tushabramishviliincluded two complexes, ensemble I (Eneolithic)and ensemble II (Upper Palaeolithic), which hesubdivided into eight layers (Liubin, 1989).Faunal remains from these excavations wereidentified by Bendukidze (1978). The boneassemblage was found to include Caucasian tur(Capra caucasica) and the extinct steppe bison(Bison priscus), as well as red deer (Cervus elaphus),aurochs (Bos primigenius), wild boar (Sus scrofa), wildhorse (Equus caballus), wolf (Canis lupus) and pinemarten (Martes martes).

Copyright # 2007 John Wiley & Sons, Ltd. Int. J. Osteoarchaeol. 18: 131–151 (2008)DOI: 10.1002/oa

132 G. Bar-Oz et al.

The new excavations at Dzudzuana Cave werecarried out in ca. 16 m2 at the front (lower area)and inner (upper area) parts of the cave (Figure 2;Meshveliani et al., 1999). Rich lithic and faunalmaterials were systematically collected by usingsmall excavation units of 0.5� 0.5� 0.05 m. Allexcavated sediments were wet-sieved through2mm mesh and the dried sediments werehand-picked for the small bones and lithics. Allbones were saved and processed according totheir spatial and stratigraphic locations. The totaldepth of the Upper Palaeolithic deposits is about3.5 m, divided into four major stratigraphic units:layers B1, B2 and C in the upper area, and layers B,C and D in the lower area. Technological andtypological studies of the lithics have beensummarised and reported alongside a series ofradiocarbon dates obtained from bones andcharcoal samples (Meshveliani et al., 2004). UnitD dates to 32–27 ka BP, Unit C to 23–20 ka BP,

and Unit B to 13–11 ka BP (all dates areuncalibrated).A. Vekua prepared a preliminary report

(Meshveliani et al., 1999) of a small bone sample(NISP¼ 349 identified bones), and recognisedthe presence of the same species as mentionedabove. He encouraged us to conduct a zooarch-aeological and taphonomic study of the entirecollection, the results of which are presentedhere.

Methods

The bone sample analysed in this study includesonly faunal remains that originated from clean,well defined Upper Palaeolithic contexts. Thecomplete zooarchaeological and taphonomiccoding and analysis procedures used to collectand present the data are detailed in Bar-Oz &

Figure 1. Map showing the location of Dzudzuana Cave, Imereti region, Republic of Georgia.


Taphonomy and Zooarchaeology of Dzudzuana Cave 133

Adler (2005; see also Bar-Oz et al., 2004). Eachexcavation unit was treated separately andrecorded stratigraphically. This procedureenabled us to carry out detailed intra-sitetaphonomic comparisons to determine the mostsignificant agents of assemblage formation(Bar-Oz & Dayan, 2003; Bar-Oz & Munro, 2004).All bones were examined and separated into

two categories: identifiable or unidentifiablefragments. Unidentifiable bone fragments weresorted into three size groups: <20mm,20–50mm, and >50mm in maximum length.The remains of each size group were counted, andfragments >50mm with no recent excavationfractures were measured to their nearest mm. Thestate of burning was recorded for each of thefragments >50mm.Skeletal elements and broad taxonomic identi-

fications were carried out in the field using avirtual comparative collection of recent speci-mens and osteological catalogues (Schmid, 1972;Hillson, 1999). Identifiable bones includedarticular ends and shafts of long bones, teeth,cranial fragments, vertebrae, and all otherrecognisable bone fragments which wereassigned to the most discriminating taxonomiclevel. Finer taxonomic identifications of closely

related species were achieved with the assistanceof A. Vekua and a comparative collection at theGeorgian State Museum. Separation of aurochsand steppe bison specimens was based onmorphological criteria of selected bones (follow-ing Olsen, 1960; Boesneck et al., 1963).

Skeletal elements that could not be assigned tospecies were grouped according to body-sizeclasses. This applies to many of the aurochs andbison remains that were combined in a Bos/Bison-size category. Shaft fragments were codedaccording to the presence of specific zones (i.e.proximal shaft, distal shaft, mid-shaft) or adiagnostic feature (e.g. foramen, muscle attach-ment; following protocol of Stiner, 2004). In mostcases identified specimens were coded accordingto their fraction of completeness (i.e. percentageof complete bone, percentage of bone shaftconnected to the epiphysis). The completenessindex (Marean, 1991) was determined for aselection of bones in order to compare the rate offragmentation and in situ attrition. Frequencies ofelement portions were used to calculate theminimum number of skeletal elements (MNE)and the minimum number of individuals (MNI)(following Klein & Cruz-Uribe, 1984; Lyman,1994). The number of identified specimens(NISP) was used as a basic measure of Taxonomicabundance (Grayson, 1984).

Recorded elements were examined duringfieldwork for macroscopic surface modificationsusing a low-resolution magnifying lens (�2.5).Modifications such as weathering (Behrensmeyer,1978), abrasion (Shipman & Rose, 1988), butch-ery marks (Binford, 1981), percussion marks(Blumenschine & Selvaggio, 1988), and evidenceof rodent gnawing, carnivore punctures, scoringand digestion, were recorded (Lyman, 1994;Fisher, 1995).

The mode of bone fragmentation wasexamined for all shaft fragments connected to aportion of long bone epiphysis to determine thestage at which the bones were broken (i.e.fresh–green vs. old–dry). The morphology of thefracture angle, outline and edge were recordedfollowing Villa and Mahieu (1991). Shaftcircumference was also measured (followingBunn, 1983) to demonstrate that the assemblagewas fully screened and collected (Marean et al.,2004).

Figure 2. Site plan of Dzudzuana Cave: areas of exca-vation are marked.



The age structure of the main hunted specieswas analysed on the basis of tooth eruption andwear of the lower deciduous fourth premolar(dP4) and lower third molar (M3). The faunalcollection of Dzudzuana Cave is curated at theGeorgian State Museum, Tbilisi, Republic ofGeorgia.

The bone assemblage

A total of 29,629 complete and fragmentarybone elements were recovered from the exca-vated areas. Of these, a total of 2214 bonefragments (7%; derived from a minimum numberof 92 individuals) were identified to taxon(including elements that were identified only tobody-size group). The rest of the bone assem-blage comprises unidentified bone fragments(Table 1).The Upper Palaeolithic inhabitants of Dzud-

zuana Cave subsisted mainly on aurochs, steppebison and Caucasian tur, which are the mostcommon taxa in all occupation layers (Figure 3;NISP and MNE of bone elements are detailed inthe Appendix). The fauna do not appear toindicate major contrasts between the differentlayers and no spatial variation in the concen-trations of the different taxa appear between thefront and the rear of the cave. The earliestassemblage (layer D) contains notably higherproportions of Caucasian tur, while the middleassemblage (Layer C) contains higher pro-portions of steppe bison and aurochs (the fewremains identifiable to species appear to includeboth species in equal numbers; Table 2). In layer Bthe ratios of Bos/Bison are similar to those of

Caucasian tur, both in the lower and upper part ofthe cave. Other species, represented in smallfrequencies in all occupational layers at the site,include red deer, wild horse, wild boar, hare (Lepuseuropeus) and carnivores (brown bear [Ursus arctos],wolf, fox [vulpes vulpes] and beech marten). Theirfrequencies do not change over time (Table 2).Although large mammals are rather rough

indicators of palaeoecological conditions, thepresence of Caucasian tur is indicative of coolAlpine conditions, presence of steppe bison andequids indicate open habitats along the Caucasiansteppe, and that of aurochs and red deer, moretemperate and forest conditions. It could be thatthe differences in species abundance observedbetween the layers reflect differences in theseason of site occupation. Caucasian tur livesalong steep rocky slopes at elevations between800–2400 m and it follows a dramatic seasonalmigration that can cover a vertical distance ofmore than 1500 m. In the early spring the herdclimbs high into the mountains, descending intothe upper part of the boreal forest in the laterautumn (Heptner et al., 1989). The steppe bison isnow extinct from the southern parts of theCaucasus, but it used to inhabit both woodlandand grassland landscapes. In the late autumn thisspecies would descend to the open steppe andgrasslands at the base of the foothills in search ofbetter feeding grounds, and in the early springthey climbed to the woodland in the mountainousareas (Heptner et al., 1989). Thus, the highproportion of Caucasian tur in layer Dmay reflecthunting activities that occurred during the latefall or winter, while the high proportion of steppebison in layer C may result from hunting in theearly spring or summer.

Table 1. Frequencies of small (<20mm in maximum dimension), medium (20–50mm) and large unidentified bonefragments (>50mm), and identified fragments in the Upper Palaeolithic layers from the lower and upper areas ofDzudzuana Cave

Layer Unidentified Identified Total

<20 mm 20–50 mm >50 mm

B 4065 3186 596 582Lower area C 5544 3530 575 703

D 66 85 16 114B1 3063 2066 273 457

Upper area B2 1774 1442 176 273C 490 421 47 85

Total 15002 10730 1683 2214 29629



Assemblage completeness anddensity-mediated attrition

The Upper Palaeolithic faunal assemblage ofDzudzuana Cave is highly fragmented; completebones are rare, and the majority of long bones are

shaft fragments (see Appendix). Nevertheless, thebones appear to be moderately well preserved asevidenced by the presence of the whole range ofbone densities, including porous parts such as theunfused proximal humeri. Bone preservation doesnot seem to vary by layer or at different parts of

LOWER AREA

267397

40268

278

70

47 28 40

20

40

60

80

DCB

LAYER

%N

ISP

UPPER AREA

54

10518824

152246

71623

0

20

40

60

80

B1 B2 C

LAYER

%N

ISP

Bos/Bison Capra Other

Figure 3. Relative frequencies of the main hunted species from the mouth (lower area, top) and the inner area (upperarea, bottom) of Dzudzuana Cave. Other species include red deer, wild boar and wild horse (data in Table 2). NISPs aregiven for each layer.

Table 2. Species abundance in the Upper Palaeolithic layers from the lower and upper areas of Dzudzuana Cave

Layer Lower area Upper area

B C D B1 B2 C

NISP MNI NISP MNI NISP MNI NISP MNI NISP MNI NISP MNI

Bos primigenius 60 2 1 1Bison priscus 23 2 9 1 1 1 4 1 5 1 3 1Bos/Bison 184 5 385 4 39 1 183 4 100 3 54 1Capra 268 9 278 6 70 3 246 7 152 7 24 2Cervus elaphus 31 3 13 1 4 1 3 1 5 1Equus caballus 3 1 14 1 1 1 8 1 7 1Sus scrofa 10 1 1 1 2 1 1 1Lepus europeus 1 1Ursus arctos 2 1 1 1 1Canis lupus 1 1 1 1Vulpes vulpes 1 1 1 1 2 1Martes martes 1 1 2 1 1 1Aves 5 1 2 1Total 582 25 700 14 113 9 457 20 273 18 88 6



the cave. Long bone shaft fragments of varioussizes constitute the majority of the unidentifiedelements. The length of Caucasian tur and Bos/Bison shaft fragments is similar within thedifferent layers (Table 3), with Caucasian turshaft fragments being shorter in total length thanthose of Bos/Bison. The frequency distribution ofthe length of Caucasian tur long bone shaftsshows that over 70% of the fragments from eachlayer fall within the 50–70mm size class(fragments smaller than 50mm were notmeasured). Similarly, the majority (�70%) ofBos/Bison long bone shaft fragments do notexceed 80mm in length. The similarity betweenlayers in the length of Caucasian tur shaftfragments suggests that each of the DzudzuanaCave faunal assemblages experienced a similardegree of fragmentation (Kruskal-Wallis; Cauca-sian tur: H¼ 5.24, P¼ 0.39; Bos/Bison: H¼ 8.56,P¼ 0.13).We compared the completeness of the faunal

assemblages and measured the amount of materiallost due to pre- and post-depositional processes.The probability that skeletal parts will survivevarious taphonomic processes is at least partially afunction of their structural density (e.g. Lyman,1984, 1994; Lam & Pearson, 2004). We foundsignificant and positive relationships betweenCaucasian tur and Bos/Bison bone survivorship

and their structural density (based on Lam et al.,1999: BMD1 density values for Rangifer tarandus).These results indicate a pronounced density-mediated bias in the skeletal part representationof the Upper Palaeolithic assemblage of Dzud-zuana Cave (Figure 4). Similarly we found asignificant and positive correlation between thefragmentation (NISP:MNE) of Caucasian tur andBos/Bison marrow-bearing bones and their bonemarrow index values (based on Bar-Oz & Munro,2007, marrow values for Gazella gazella; Figure 4).This result indicates that long bone shafts withthe highest marrow ‘stores’ (tibia, radius, meta-carpus and metatarsus) are the most fragmented,while elements with the lowest marrow ‘stores’(first and second phalanges) have the lowestfragmentation indices, suggesting that long boneswere intentionally selected and broken formarrow. This observation is supported by themode of bone fragmentation and the presence ofpercussion marks close to fracture edges. Theselective destruction of bones for marrowextraction is also demonstrated by the high rateof fresh bone breakage (discussed below).Conversely, comparisons of Caucasian tur andBos/Bison bone survivorships to food utility index(FUI; based on the weight of useable tissue ofRangifer tarandus; Metcalfe & Jones, 1988) show nopronounced relationship (Figure 4). This suggests

Table 3. Frequencies of 10mm size classes of Caucasian tur and Bos/Bison-sized unidentified long bone shaftfragments from the Upper Palaeolithic layers from the lower and upper areas of Dzudzuana Cave (fragments smallerthan 50mm were not measured)

Length (mm) Lower area Upper area

B C D B C D B1 B2 C B1 B2 C

Capra Bos/Bison Capra Bos/Bison

50–60 68 54 3 70 94 5 47 48 8 45 28 960–70 53 60 1 108 112 4 47 22 3 63 28 870–80 38 35 87 88 2 12 11 2 34 23 480–90 16 8 51 46 10 1 1 14 5 490–100 6 3 32 30 1 1 11 5 4100–110 4 3 26 20 1 1 3110–120 1 1 6 13 1 1 1120–130 10 7130–140 3 4 1 1140–150 1 2150–160 1 3160–170 1 1Total 186 164 4 396 420 12 117 83 14 168 92 34Mean length (mm) 66.8 66.4 58.1 76.5 75.2 62.66 62.7 60.2 61.9 68.2 68.5 73.3SD 13.1 11.8 6.6 19.5 20.2 13.5 9.3 8.6 9.9 12.3 10.2 17.2



that selective transport of high-utility body partswas not a major factor in assemblage formation.The strong relationship between bone survivor-ship, bone density, bone shaft fragmentation andmarrow values on the one hand, and the weakassociation between bone survivorship and theirfood value on the other, indicate that pre-depositional human bone processing behaviourswere a major agent in the formation of the faunalbone assemblage of Dzudzuana Cave.

Figure 5 presents a bivariate scatterplot of thecompleteness index for 11 carpal and tarsal bonesof Caucasian tur and Bos/Bison. The index foreach element was obtained by recording thefraction of the element represented by eachspecimen (i.e. the preserved percentage, follow-ing Marean, 1991). All plotted points fall on ornear the diagonal line, indicating their equalcompleteness. This demonstrates that bonepreservation of the two taxa is similar, and that

Y=1.14X+0.14; r=0.44; P=0.03 Y=0.91X+0.18; r=0.56; P<0.01

0.00

0.20

0.40

0.60

0.80

1.00

0.1 0.2 0.3 0.4 0.5 0.6 0.7

Bone density

Sur

vivo

rshi

p

0.00

0.20

0.40

0.60

0.80

1.00

0.1 0.2 0.3 0.4 0.5 0.6 0.7

Bone density

Sur

vivo

rshi

p

Y=-0.002X+0.41; r=0.24; P=0.43 Y=0.002X+0.40; r=0.07; P=0.78

0

0.2

0.4

0.6

0.8

1

0 20 40 60 80 100

FUI

Sur

vivo

rshi

p

0

0.2

0.4

0.6

0.8

1

0 20 40 60 80 100

FUI

Sur

vivo

rshi

p

Y=-0.05X+1.89; r=0.70; P=0.05 Y=-0.09X+1.80; r=0.83; P=0.01

0

1

2

3

4

5

6

0 10 20 30 40 50

Marrow index

NIS

P/M

NE

0

1

2

3

4

5

6

0 10 20 30 40 50

Marrow index

NIS

P/M

NE

Figure 4. Relationship between bone density and bone survivorship (% MAU) (top), food utility (FUI) and bonesurvivorship (middle), and between marrow index and fragmentation of long bone shafts (NISP/MNE) (bottom) ofCaucasian tur (left) and Bos/Bison (right) from Dzudzuana Cave. Regression equation, Spearman’s r, and P values aregiven for each scatterplot.



both the medium and large-sized animals sufferedfrom analogous density-mediated attrition. Inaddition, the percentage of completeness formost bones is high (>80%, except for thecalcaneus which is only about 60% complete),suggesting that the Dzudzuana Cave assemblagedid not suffer considerably from post-depositional decay.

Bone fragmentation

Analysis of the breakage patterns (fracture angle,fracture outline and fracture edge) of Caucasiantur and Bos/Bison long bone shaft fragments(following fracture typology of Villa & Mahieu,1991) points to the relatively high frequencies ofoblique, V-shaped and jagged bone fractures(Table 4). Since the identification of breakagetypes is somewhat subjective, several specimenswere classified in intermediate categories. Therelatively high frequencies of green bonefractures of both taxa suggest that bones werebroken while still fresh, either prior to depositionor shortly thereafter (Villa & Mahieu, 1991; seealso Enloe, 1993; Outram, 2001), most probablyfor bone marrow extraction. In addition, wefound a small number of percussion marks(Table 5) close to the fracture edges, whichcould also have been made during the marrowextraction process. The high frequency ofgreen long bone fractures is in accordance withprevious findings and demonstrates that a majorshare of bone breakage resulted from pre-depositional human activities. Such bone-processing

0

20

40

60

80

100

0 20 40 60 80 100

Bos/Bison

Cap

ra c

auca

sica

Figure 5. Bivariate scatterplot of completeness index for11 carpal and tarsal bones of Caucasian tur and Bos/Bison from Dzudzuana Cave.

Table

4.Frequenciesoffracture

angle,fracture

outline,fracture

edge,andshaftcircumferenceofCaucasianturandBos/Bison-sizedelements

from

theUpper

Palaeolithic

layers

ofDzudzuanaCave(relativefrequenciesare

givenin

parentheses)

Layer

Fracture

angle

Fracture

outline

Fracture

edge

Shaftcircumference

Obliq

ue

(fresh)

Right

(dry)

Interm

ediate

Vshaped

(fresh)

Transverse

(dry)

Interm

ediate

Jagged

(fresh)

Smoothed

(dry)

Interm

ediate

Less

than1/2

More

than1/2

Complete

Bos/Bison

B127(44%)91(31%)

73(25%)

130(43%)

69(23%)

102(34%)

132(44%)

90(30%)

76(26%)

282(93%)

19(6%)

1(0%)

B1

29(52%)18(32%)

9(16%)

33(58%)

16(28%)

8(14%)

40(68%)

12(20%)

7(12%)

47(81%)

8(14%)

3(5%)

B2

50(48%)36(34%)

19(18%)

39(40%)

27(28%)

31(32%)

65(63%)

28(27%)

10(10%)

103(98%)

1(1%)

1(1%)

C28(54%)16(31%)

8(15%)

32(60%)

9(17%)

12(23%)

30(59%)

7(14%)

14(27%)

52(100%)

00

Capra

caucasicaB

52(43%)32(26%)

37(31%)

60(47%)

48(37%)

21(16%)

66(50%)

34(26%)

31(24%)

126(96%)

5(4%)

0

B1

19(32%)26(44%)

14(24%)

25(39%)

27(42%)

12(19%)

40(67%)

15(25%)

5(8%)

47(78%)

10(17%)

3(5%)

B2

14(47%)

9(30%)

7(23%)

20(63%)

10(31%)

2(6%)

14(42%)

7(21%)

12(36%)

33(100%)

00

C16(52%)12(39%)

3(10%)

17(52%)

12(36%)

4(12%)

25(81%)

4(13%)

2(6%)

29(94%)

2(6%)

0



behaviours produced high frequencies of fresh/green fractures and probably account for the highnumber of small, unidentified shaft fragmentswithin each layer. Still, the Dzudzuana Caveassemblage contains relatively high proportionsof dry bone fractures (>20% in most assem-blages). Thus, it appears that after their depo-sition in the cave the remains were subjected tofurther breakage and modification by attritionalprocesses such as sediment compaction andtrampling. The shaft circumference data(Table 5) indicate that the majority of long boneshaft fragments are represented by less than halfof the complete circumference (>80% in alllayers), as would be expected from assemblagesthat were carefully screened and collected(Marean et al., 2004)

Specific attritional processes

We searched for signs of specific attritionalprocesses that may have been responsible forbone loss and fragmentation. Analysis of allepiphyses with attached shaft fragments indi-cated some loss of bone due to physical erosion,as evidenced by the presence of abraded andbleached bones. In our analysis we recordedabraded bones as those having rounded orsmoothed fracture surfaces (Shipman & Rose,1988). Bleached bones were coded as being

chalky in texture, creamy-white in colour, andfriable with a powdery surface. On average,>20% of the Dzudzuana Cave bones appear tohave been abraded, with rounded broken edges.The ratios appear to be similar among the layersfrom both the lower and upper areas of the cave.In addition, on average more than 10% of bonesfrom all layers and from both areas of excavationwere bleached (Table 5). Interestingly, Layer C,identified at the mouth of the cave, contains onlya small number of bleached bones (5 specimens),with two of these derived from the upper levels ofthe layer. Bone decomposition within both theinner part and mouth of the cave was most likelyproduced by the creek that flows from the innerpart of the cave. Indeed, our results indicate thatthe majority of post-depositional bone destruc-tion was caused by post-burial physical erosion.However, fluvial sorting of skeletal elements hasnot occurred and the represented elements arefrom opposite transport groups (Voorhies, 1969).While ribs and vertebrae are the most transport-able elements and so could have been washedaway from the excavated area, skulls and teeth,which are considered low transportable elements,are also underrepresented. Thus, it appears thatloss of bones owing to fluvial transport did notaffect the Dzudzuana Cave bone assemblage in asubstantial way. This is also in accordance withthe lithics and abundant flint chips and flakes thatare distributed in different sizes in all layers andareas of the cave.

Table 5. Frequencies of bone modifications from the Upper Palaeolithic layers from the lower and upper areas ofDzudzuana Cave

Capra caucasica Bos/Bison Total

B B1 B2 C B B1 B2 C

Total NISP (excluding teeth) 153 180 109 219 223 143 76 379 1482Abrasion 39 37 27 34 59 23 20 57 296% 25.5 20.6 24.8 15.5 26.5 16.1 26.3 15.0 20.0Bleached 21 32 18 4 17 20 13 1 126% 13.7 17.8 16.5 1.8 7.6 14.0 17.1 0.3 8.5Carnivore impacts 4 2 2 2 4 3 2 9 28% 2.6 1.1 1.8 0.9 1.8 2.1 2.6 2.4 1.9Rodent impacts 5 1 2 3 8 0 2 5 26% 3.3 0.6 1.8 1.4 3.6 0.0 2.6 1.3 1.8Impact fractures 2 7 4 5 4 8 4 4 38% 1.3 3.9 3.7 2.3 1.8 5.6 5.3 1.1 2.6Butchery marks 2 8 3 5 5 3 0 14 40% 1.3 4.4 2.8 2.3 2.2 2.1 0.0 3.7 2.7Burned 10 9 2 7 12 3 0 9 52% 6.5 5.0 1.8 3.2 5.4 2.1 0.0 2.4 3.5



Many of the bones from Dzudzuana Cavedisplay surface cracks, evidence of exfoliation,and moderate signs of surface weathering(Figure 6). The majority of bones conform toweathering stages 2–3 of Behrensmeyer’s (1978)six weathering stages. No pronounced differenceis observed between the different occupationallayers (Kruskal-Wallis; Caucasian tur: H¼ 4.15,P¼ 0.25; Bos/Bison: H¼ 4.50, P¼ 0.21). Theseresults indicate that the majority of bones weresubjected to a moderate degree of subaerialweathering (stages 2–3). Thus, bones from alllayers were probably exposed for several yearsprior to burial but were buried under largelysimilar sedimentological conditions.Additional bone surface modifications were

somewhat difficult to assess because of the friablecondition of the bones and the fact that numerousbone surfaces were dissolved by erosion. Sincenot all bone surfaces were visible, the frequenciesof bone modifications should be treated asminimal estimates. Certain bone modificationswere observed on the surface of the bones. A fewspecimens (n¼ 28; 2% of total NISP, excluding

teeth) exhibit unambiguous carnivore score marks(e.g. chewing, gnawing and scratch marks;Table 5). The non-identifiable elements (frag-ments >50mm), which comprise mainly mid-shaft fragments, also showed lower rates ofcarnivore tooth-marks (not exceeding 2% in alllayers). The low percentage of carnivore tooth-marks on the mid-shaft portions suggests thatcarnivores had only secondary access to thebones. This further attests that the DzudzuanaCave bone assemblage was not significantlyaltered or modified by carnivore ravaging.Similarly, rodent gnawing is evident only in asmall proportion of the bones from each layer(not exceeding 4%, Table 5) and does not appearto be a major contributing factor to modification.

Faunal transport and skeletalpart representation

Additional investigation of skeletal completenessis determined through detailed analysis of skeletalpart profiles. Compression of individual elements

Figure 6. Relative frequencies of Caucasian tur (top) and Bos/Bison (bottom) long bone shaft fragments per weatheringstage from Dzudzuana Cave. NISP values are given for each column.



into nine anatomical regions (horn, head, neck,axial skeleton, upper forelimb, lower forelimb,upper hindlimb, lower hindlimb and toes;following Stiner, 1994, 2002) allows us tohighlight skeletal biases that may have resultedfrom prey transport.Figures 7 and 8 show the representation of

Caucasian tur and Bison/Bos skeletal elements(%MAU) in the different Upper Palaeolithiclayers at Dzudzuana Cave. The representation ofskeletal elements is similar for both the medium

and large ungulate taxa in all layers. All skeletalpart profiles are biased against the vertebralcolumn and neck, but are relatively rich in upperfront and hind limbs of both species. Heads(horns and skulls), mandibles and toes also appearin low ratios in all layers. The bone assemblagealso exhibits low representation of pelves andscapulae, a pattern which is most pronounced inthe large-sized prey. The differential representa-tion of skeletal parts, and in particular the lowrepresentation of heads, toes and axial skeleton

Layer B

020406080

100

Horn

Skul

l

Man

dible

Cervic

al

Thora

cic

Lumbar

Scapu

la

Humer

us

Radiu

sUln

a

Meta

carp

us

Carpa

ls

Pelv

is

Fem

urTib

ia

Patella

Tarsa

ls

Meta

tarsa

lToe

s

%M

AU

Layer B1

02040

6080

100

Horn

Skull

Man

dible

Cervic

al

Thora

cic

Lumbar

Scap

ula

Humer

us

Radiu

sUln

a

Meta

carp

us

Carpa

ls

Pelvis

Fem

urTib

ia

Patel

la

Tarsa

ls

Meta

tarsa

lToes

%M

AU

Layer B2

020406080

100

Horn

Skull

Man

dible

Cervic

al

Thorac

ic

Lumbar

Scapu

la

Humer

us

Radiu

sUln

a

Meta

carp

us

Carpa

ls

Pelvis

Femur

Tibia

Patella

Tarsa

ls

Meta

tarsa

lToe

s

%M

AU

Layer C

02040

6080

100

Horn

Skul

l

Man

dible

Cervic

al

Thora

cic

Lumbar

Scap

ula

Humer

us

Radiu

sUln

a

Meta

carp

us

Carpa

ls

Pelvis

Femur

Tibia

Patella

Tarsa

ls

Meta

tarsa

lToe

s

%M

AU

Figure 7. Skeletal part representation of Caucasian tur from Dzudzuana Cave (based on MNE, see Appendix).



(including pelves and scapulae) might suggestthat both large and medium-sized prey weresubject to extensive field butchery at the kill-siteprior to import to the cave.

Butchery marks and burning

A small number of butchery marks and burnedbones were found among the Caucasian tur and

Bos/Bison remains. Burned specimens wereobserved in all of the archaeological horizonsand appear to be randomly distributed. However,the frequencies of burned bones are too low toanalyse by anatomical unit. Only 3.5% of theidentified specimens (excluding teeth) wereburned (Table 5), and only 2.7% bear butcherymarks (Table 6). Most of the cut marks were madeduring the process of dismemberment (65% inBos/Bison and 62% in tur, following Binford,1981). The small number of cut-marked bones is

Layer B

020406080

100

Horn

Skul

l

Man

dible

Cervic

al

Thora

cic

Lumba

r

Scapu

la

Humer

us

Radiu

sUln

a

Meta

carp

us

Carpa

ls

Pelv

is

Femur

Tibia

Patella

Tarsa

ls

Meta

tarsa

lToe

s

%M

AU

Layer B1

0

20

4060

80

100

Horn

Skul

l

Man

dible

Cervic

al

Thorac

ic

Lumbar

Scap

ula

Humer

us

Radiu

sUln

a

Meta

carp

us

Carpa

ls

Pelv

is

Femur

Tibia

Patella

Tarsa

ls

Meta

tarsa

lToe

s

%M

AU

Layer B2

020406080

100

Horn

Skul

l

Man

dible

Cervic

al

Thorac

ic

Lumbar

Scap

ula

Humer

us

Radiu

sUlna

Meta

carp

us

Carpa

ls

Pelvis

Femur

Tibia

Patel

la

Tarsa

ls

Meta

tarsa

lToe

s

%M

AU

Layer C

02040

6080

100

Horn

Skul

l

Man

dible

Cervic

al

Thorac

ic

Lumbar

Scap

ula

Humer

us

Radiu

sUln

a

Meta

carp

us

Carpa

ls

Pelvis

Fem

urTib

ia

Patel

la

Tarsa

ls

Meta

tarsa

lToes

%M

AU

Figure 8. Skeletal part representation of Bos/Bison from Dzudzuana Cave (based on MNE, see Appendix).



most probably related to the relatively poorpreservation of the bone surfaces.

Age and season at death

Age at death was inferred on the basis ofepiphyseal fusion and dental eruption and wear.While reconstruction of the age profile, and inparticular the presence of neonatal individuals,enables us to determine season of death, thesample size of the different layers at DzudzuanaCave is not sufficient to conduct detailed analysesof mortality profiles. We clustered our specimens

into four major age categories: neonatal, juven-iles, adult and old adult (Table 7; followingMiracle, 2005). ‘Neonatal’ includes bones that,according to their size and texture, come fromnear-born fetuses or recently-born young. ‘Juven-ile’ is defined based on unfused epiphyses ofseveral skeletal elements whose fusion age isabout 24 months (radii, metapodia, femora, tibiaeand calcanea; Davis, 1983) and deciduousdentition. The ‘Adult’ category consists ofpermanent teeth and fused bones, while the‘Old adults’ category is marked by teeth withheavy wear.

Table 6. Summary of butcherymarks onCaucasian tur and Bos/Bison bone elements from the Upper Palaeolithic layersof Dzudzuana Cave, and activities with which they may be associated (after Binford, 1981)

Layer Dismembering n Filleting n Skinning n Other n

B Bos/Bison Scapula gle-fos. 1 Radius prox-shaft 1Radius prox. 1 Tibia prox-shaft 1Rib shaft 3

Caucasian tur Femur dis-shaft 2 Mandible sym. 1Sus Scrofa Metapod prox. 2

Calcaneus prox. 2B1 Bos/Bison Carpal 1 Tibia prox-shaft 1 Femur med-shaft 2

Caucasian tur Humerus dis. 2 Scapula spine 1Femur prox. 1 Metapod med-shaft 2Tibia dis. 1Astragallus 1Navicular-cuboid 1

B2 Bos/Bison Tibia dis. 1Caucasian tur Radius prox. 2 Rib shaft 1

C Bos/Bison Toracic ver. sp.pro. 1 Caudal ver. cent. 1Humerus dis. 2 Humerus dis-shaft 1 Radius med-shaft 1Ulna dis. 2 Femur dis-shaft 1Metacarpus prox. 1Astragallus 3Metapod dis-shaft 1

Caucasian tur Humerus dis. 1 Humerus dis-shaft 1 Phalanx 1 prox. 1Radius prox. 1 Radius prox-shaft 1Metacarpus prox. 1

Abbreviations: ver., vertebrae; dis., distal; prox., proximal; med., medial; sym., symphasis; gle-fos., glenoid fossa;sp.pro., spinous process; cent., centrum.

Table 7. Summary of ageing from teeth and bones of Caucasian tur and Bos/Bison remains from the Upper Palaeolithiclayers of Dzudzuana Cave

% NISP Aged NISP

Fetal/neonatal Juvenile Adult Old adult

B Caucasian tur 4 21 75 0 48Bos/Bison 0 28 68 4 25

C Caucasian tur 0 27 73 0 33Bos/Bison 3 14 78 5 37



The resulting mortality profiles are similar forthe two major archaeological layers (layers B andC) for both the Caucasian tur and Bos/Bison taxa.The small samples from layers B1 and B2 did notallow a similar analysis. The mortality profiles ofDzudzuana Cave are dominated by adult indi-viduals. In addition, the Caucasian tur from layerB comprise two newborn individuals (twocomplete medial-shaft humeri). The size of thosebones suggests that they are from calves less thana month old (Amorosi, 1989). Similarly, a singleunworn dP4 of a Bos/Bison, from layer C, is from acalf only a few months old (Grant, 1982).Assuming that the season of birth of tur andBos/Bison in the Caucasus region is in the latespring to early summer (May–June; Vereshcha-gin, 1967; Heptner et al., 1989), these remainswould have come from animals killed during latesummer and possibly early autumn. Thesefindings suggest that some of the huntingencounters and the occupation of DzudzuanaCave took place in the summer and possibly theautumn.

Discussion

The taphonomic history of Dzudzuana Caveindicates that humans were the primary agents ofaccumulation, modification and destruction of thebone assemblage from each occupational level.The cave occupation was apparently associatedwith the hunting of large mammals, mainlyaurochs, steppe bison and Caucasian tur, thefrequencies of which do not change profoundlyover time. Taphonomic analysis reveals that theassemblage is characterised by significant density-mediated biases, caused by both human bone-processing behaviours and in situ post-burial boneattrition. Bone preservation does not vary bylevel, and little evidence was found to indicatechanges in the depositional environments indifferent areas of the cave.Although we can not reject the hypothesis that

density-mediated bias is partially caused by in situattrition, we have found that the main agent ofbone destruction is human subsistence behaviour,in particular the processing of bone marrow. Suchan activity produced large numbers of unidenti-fied bone fragments of various sizes, most of

which exhibit green bone fractures. The sys-tematic breakage of fresh bones indicates thatfood resources were not wasted by the site’sinhabitants. The presence of cut-marks from allstages of butchery suggests that at least somefaunal processing took place at or near the site.Furthermore, the abundance of prey body partsand butchery marks, coupled with the diversity oflithic artefacts (Meshveliani et al., 2004), indicatethat multiple activities were carried out at the site.The low rates of axial units and high representa-tion of upper fore- and hindlimbs for both Bos/Bison and Caucasian tur suggest that carcassesoccasionally underwent field butchery and dis-memberment at kill sites before transport to thecave for consumption. Unfortunately, the com-bined post-depositional impacts (weathering,trampling and physical erosion) on the faunalremains prevent us from developing detailedinferences about the role of selective transport ofprey to the site.While radiometric dates of Dzudzuana Cave

show gaps in the Upper Palaeolithic record, as inthe nearby late Middle Palaeolithic and earlyUpper Palaeolithic site of Ortvale Klde (Adleret al., 2006), these are not evident in thetaphonomic analysis of the excavated assem-blages. The faunal record does not trackenvironmental changes and can be consideredat best only a coarse-grained indicator of long-term palaeoecological changes within the region.We believe that the Dzudzuana Cave boneassemblage was created and accumulated as partof the same larger economic system, as indicatedby the stability of carcass processing, marrowextraction and butchering behaviours. These datahighlight uniform patterns of cultural andeconomic behaviours related to food procure-ment and processing strategies. Intra-site tapho-nomic comparisons provide multiple lines ofevidence for intensive occupations throughoutthe Upper Palaeolithic. The density and size ofbone assemblages, the high percentage of freshbone fractures and the extent of fragmentation allindicate that Dzudzuana Cave was repeatedlyoccupied by Upper Palaeolithic foragers overmany years. The absence of carnivore activitiessuggests that bones were not available ordesirable to carnivores following site abandon-ment.



The faunal assemblage from Dzudzuana Caveprovides detailed insight into Upper Palaeolithicfood procurement and processing strategies in thesouthern Caucasus. However, the potentialimportance of such insight is difficult to realisegiven the paucity of similar contemporaneousassemblages in the region with which to compareour findings. Comparison with the Palaeolithiclayers at Ortvale Klde reveal similar speciesrepresentation, with Caucasian tur being the mostabundant prey species in each of the late Middleand early Upper Palaeolithic layers (Bar-Oz &Adler, 2005; Adler et al., 2006). Conversely, in theUpper Palaeolithic layers of Dzudzuana Cave,the proportions of steppe bison and aurochsincrease at the expense of the Caucasian tur. Itseems reasonable to conclude that the differencesin the taxonomic abundance between the twosites reflect differences in the season of occu-pation. While zooarchaeological data suggestthat Ortvale Klde was occupied during lateautumn to early spring, it seems that DzudzuanaCave was occupied primarily during the latespring to early autumn.Results of the detailed intra-site taphonomic

analysis at Dzudzuana Cave show no majordifferences in hunting and food-processingstrategies from those of the late Middle Palaeo-lithic and early Upper Palaeolithic layers ofOrtvale Klde. Although the Upper Palaeolithicpopulations in the southern Caucasus weremodern humans (Adler et al., 2006), zooarchaeo-logical analyses were unable to document newtrends in food procurement strategies during thetransition from the Middle to the UpperPaleolithic. Analysis of Caucasian tur and Bos/Bison age classes demonstrates that the UpperPalaeolithic hunters of Dzudzuana Cave prefer-entially targeted prime-age adult individuals. Thedominance of prime-age adults may be indicativeof intercept hunting, rather than some form ofencounter hunting. The complex valley andravine topography of the southern Caucasuswould have been well suited to stalking andambushing specific individuals of particular preyspecies.The zooarchaeological data from Dzudzuana

Cave and Ortvale Klde is of interest as the lithicindustries tell a different story. While the MiddlePalaeolithic lithic assemblages on both sides of

the Caucasus range are different, the UpperPalaeolithic tool kits indicate that the mountainsdid not serve as a geographical barrier, suggestingthat the Upper Palaeolithic newcomers to theCaucasus were more mobile and could exploitmuch larger territories than their Middle Palaeo-lithic predecessors (Adler, 2002; Meshvelianiet al., 2004; Adler et al., 2006). Thus the results ofthis study support our previous argument thathunting behaviours are poor proxies for majorbehavioural differences between Neanderthalsand Modern humans (see detailed discussion inAdler et al., 2006). Both Middle and UpperPalaeolithic lifeways were largely determined bywhat food resources were available on a local andseasonal basis (Bar-Yosef, 2004).

Acknowledgements

We thank the American School for PrehistoricResearch (Peabody Museum, Harvard Univer-sity) for the financial support of this project.We are grateful to our colleagues at the GeorgiaState Museum, D. Lordkipanidze, A. Vekua andN. Tushabramishvili, for their collaboration andassistance during our research in the Republic ofGeorgia. Bar-Oz thanks the American School ofPrehistoric Research, Harvard University, for theMacCurdy post-doctoral fellowship duringwhich part of the zooarchaeological analyseswere conducted. He also thanks RichardMeadowfrom Harvard University for important adviceand support. Thanks are due to Reuven Yeshurunfor his help with laboratory analysis during field-work in 2005–2006, and Dan Adler, ReuvenYeshurun and two anonymous referees for theircomments on an earlier draft of the text.

References

Adler DS. 2002. Late Middle Palaeolithic Patterns of LithicReduction, Mobility, and Land Use in the SouthernCaucasus. Unpublished PhD thesis, Department ofAnthropology, Harvard University, Cambridge,MA.

Adler DS, Tushabramishvili N. 2004. Middle Palaeo-lithic patterns of settlement and subsistence in thesouthern Caucasus. In Middle Palaeolithic Settlement



Dynamics, Conard N (ed.). Publications in Prehis-tory, Kerns Verlag: Tubingen; 91–132.

Adler DS, Bar-Oz G, Belfer-Cohen A, Bar-Yosef O.2006. Ahead of the game: Middle and UpperPalaeolithic hunting behaviors in the southern Cau-casus. Current Anthropology 47: 89–118.

Amorosi T. 1989. A Postcranial Guide to Domestic Neo-Natal and Juvenile Mammals – The Identification and Agingof Old World Species, BAR International Series 533:Oxford.

Bar-Oz G, Adler DS. 2005. Taphonomic history of theMiddle and Upper Palaeolithic faunal assemblagefrom Ortvale Klde, Georgian Republic. Journal ofTaphonomy 3: 185–211.

Bar-Oz G, Dayan T. 2003. Testing the use of multi-variate intersite taphonomic comparisons: the fau-nal analysis of Hefzibah in its Epipalaeolithiccultural context. Journal of Archaeological Science 30:885–900.

Bar-Oz G, Munro ND. 2004. Beyond cautionary tales:a multidimensional taphonomic approach for iden-tifying sub-patterns in ungulate body-part data.Journal of Taphonomy 2: 201–220.

Bar-Oz G, Munro ND. 2007. Gazelle bone marrowyields and Epipalaeolithic animal exploitation strat-egies in the southern Levant. Journal of ArchaeologicalScience 34: 946–956.

Bar-Oz G, Adler DS, Vekua A, Meshveliani T, Tush-abramishvili N, Belfer-Cohen A, Bar-Yosef O. 2004.Faunal exploitation patterns along the southernslopes of the Caucasus during the Late Middleand Early Upper Palaeolithic. In Colonisation,Migration, and Marginal Areas: A ZooarchaeologicalApproach, Mondini S, Munoz M, Wickler S (eds).Oxbow Books: Oxford; 46–54.

Bar-Yosef O. 2004. Eat what is there: hunting andgathering in the world of Neanderthals and theirneighbors. International Journal of Osteoarchaeology 14:333–342.

Behrensmeyer AK. 1978. Taphonomic and ecologicalinformation from bone weathering. Paleobiology 4:150–162.

Bendukidze O. 1978. Mammal Fauna of the Dzud-zuana Paleolithic Settlement in Western Georgia.Cbornik Pechereh Gruzia (The Caves of Georgia) 7: 72–79.

Binford LR. 1981. Bones: Ancient Men and Modern Myths.Academic Press: New York.

Blumenschine RJ, Selvaggio M. 1988. Percussionmarks on bone surface as a new diagnostic of humanbehaviour. Nature 333: 763–765.

Boesneck J, Jequier JP, Stampfli HR. 1963. SeebergBurgaschisee-Sud, Teil 3: Die Tierreste. Verlag Stampfli& Cie Bern: Bern; 118–159.

Bunn HT. 1983. Evidence on the diet and subsistencepatterns of Plio-Pleistocene hominids at Kobi Fora,Kenya, and Oldovai Gorge, Tanzania. In Animals andArchaeology: 1. Hunters and Their Prey, Clutton-Brock J,Grigson C (eds). BAR International Series 163:Oxford; 21–30.

Davis SJM. 1983. The age profiles of gazelles predatedby ancient man in Israel: possible evidence for ashift from seasonality to sedentism in the Natufian.Paleorient 9: 55–62.

Enloe JG. 1993. Ethnoarchaeology of marrow crack-ing: implications for the recognition of prehistoricsubsistence organization. In From Bones to Behavior:Ethnoarchaeological and Experimental Contributions to theInterpretation of Faunal Remains, Hudson J (ed.).Southern Illinois University, Center for Archaeo-logical Investigations: Occasional paper No. 21:Illinois; 82–97.

Fisher JW. 1995. Bone surface modifications inzooarchaeology. Journal of Archaeological Method andTheory 2: 7–68.

Grant A. 1982. The use of tooth wear as a guide to theage of domestic ungulates. In Ageing and Sexing AnimalBones from Archaeological Sites, Wilson B, Grigson C,Payne S (eds). BAR International Series 109:Oxford; 91–108.

Grayson DK. 1984. Quantitative Zooarchaeology. Aca-demic Press: New York.

Heptner VG, Nasimovich AA, Bannikov AG. 1989.Mammals of the Soviet Union, Vol. 1: Ungulates. E.J. Brill:Leiden.

Hillson S. 1999. Mammal Bones and Teeth: an IntroductoryGuide to Methods of Identification. Institute of Archae-ology, University College London: London.

Klein RG, Cruz-Uribe K. 1984. The Analysis of AnimalBones from Archaeological Sites. University of ChicagoPress: Chicago.

Lam YM, Pearson OM. 2004. The fallibility of bonedensity values and their use in archaeologicalanalyses. Journal of Taphonomy 2: 99–115.

Lam YM, Chen X, Pearson OM. 1999. Intertaxonomicvariability in patterns of bone density and thedifferential representation of bovid, cervid, andequid elements in the archaeological record.American Antiquity 64: 343–362.

Liubin VP. 1989. Paleolit Kavkaza (The Paleolithic ofthe Caucasus). In Paleolit Kavkaza i Severnoj Azii,Boriskovsky PI (ed.). Nauka: Moscow; 9–142.

Liubin VP, Bosinski G. 1995. The earliest occupationof the Caucasus region. In The Earliest Occupation ofEurope, Roebroeks W, Kolfschoten V (eds). Univer-sity of Leiden Press: Leiden; 207–254.

Lordkipanidze DO. 1992. Osobenosti Vzaimodeistvia Per-vobytnogo Obshchestva I Prirodnoi Sredi Vuslovi’akh



Gornikh Teritorii (The Interactions of Primitive Societies andthe Environments of Mountain Territories). UnpublishedPhD thesis, University of Moscow.

Lordkipanidze DO. 1998. Pleistocene settlement ofthe Transcaucasus by hominids. In Anatolian Prehis-tory at the Crossroads of TwoWorlds Vol. 1, Otte M (ed.).ERAUL 85: Liege; 15–28.

Lyman RL. 1984. Bone density and differential survi-vorship of fossil classes. Journal of AnthropologicalArchaeology 3: 259–299.

Lyman RL. 1994. Vertebrate Taphonomy. CambridgeUniversity Press: Cambridge.

Marean CW. 1991. Measuring the postdepositionaldestruction of bone in archaeological assemblages.Journal of Archaeological Science 18: 677–694.

Marean CW, Domınguez-Rodrigo M, Pickering TR.2004. Skeletal element equifinality in zooarchaeol-ogy begins with method: the evolution andstatus of the ‘‘shaft critique’’. Journal of Taphonomy2: 69–98.

Meshveliani T, Bar-Yosef O, Belfer-Cohen A, DjakeliN, Kraus A, Lordkipanidze DO, Tvalchrelidze M,Vekua A. 1999. Excavations at Dzudzuana Cave,Western Georgia (1996-1998): preliminary results.Prehistoire Europeenne 15: 79–86.

Meshveliani T, Bar-Yosef O, Belfer-Cohen A. 2004.The Upper Paleolithic of Western Georgia. In TheEarly Upper Paleolithic beyond Western Europe, Brantig-ham PJ, Kuhn SL, Kerry KW (eds). University ofCalifornia Press: Berkeley, CA; 129–143.

Meshveliani T, Bar-Oz G, Bar-Yosef O, Belfer-CohenA, Boaretto E, Djakeli N, Matskevich Z. In press.Mesolithic hunters at Kotias Klde, western Georgia.Paleorient.

Metcalfe D, Jones KT. 1988. A reconsideration ofanimal body part utility indices. American Antiquity53: 486–504.

Miracle P. 2005. Late Mousterian subsistence and caveuse in Dalmatia: the zooarchaeology of MujinaPecina, Croatia. International Journal of Osteoarchaeol-ogy 15: 84–105.

Olsen SJ. 1960. Post-cranial skeletal characters of Bison andBos. Papers of the Peabody Museum of Archaeologyand Ethnology, Vol XXXV, No. 4: Cambridge.

Outram AK. 2001. A new approach to identifyingbone marrow and grease exploitation: why the‘‘intermediate’’ fragments should not be ignored.Journal of Archaeological Science 28: 401–410.

Schmid E. 1972. Atlas of Animal Bones for Prehistorians,Archaeologists and Quaternary Geologists. Elsevier Pub-lishing Company: Amsterdam.

Shipman P, Rose J. 1988. Bone tools: an experimentalapproach. In Scanning Electron Microscopy in Archae-ology, Olsen SL (ed.). British ArchaeologicalReports: Oxford; 303–335.

Stiner MC. 1994.Honor among Thieves: a ZooarchaeologicalStudy of Neanderthal Ecology. Princeton UniversityPress: Princeton.

Stiner MC. 2002. On in situ attrition and vertebratebody part Profiles. Journal of Archaeological Science 29:979–991.

Stiner MC. 2004. A comparison of photon densito-metry and computed tomography parameters ofbone density in ungulate body part profiles. Journalof Taphonomy 2: 117–145.

Tappen M, Adler DS, Ferring CR, Gabunia M, VekuaA, Swisher CC III. 2002a. Akhalkalaki: the taph-onomy of an early Pleistocene locality in the Repub-lic of Georgia. Journal of Archaeological Science 29:1367–1391.

Tappen M, Ferring CR, Lordkipanidze DO, Vekua A,Kiladze G. 2002b. Preliminary observations on thevertebrate taphonomy of the Dmanisi locality in theRepublic of Georgia. In Current Topics on Taphonomyand Fossilization, De Renzi M, Alonso MVP,Belinchon M, Penalve E, Montoya P, Marquez-Aliaga A (eds). Ajuntament de Valencia: Valencia;161–170.

Vekua A, Lordkipanidze DO, Rightmire PG, Agusti J,Ferring CR, Maisuradze G, Mouskhelishvili A,Nioradze M, Ponce de Leon M, Tappen M, Tval-chrelidze M, Zollikofer C. 2002. A new skull ofearly Homo from Dmanisi, Georgia. Science 297:85–89.

Vereshchagin NK. 1967. The Mammals of the Caucasus: AHistory of the Evolution of the Fauna. Israel Program forScientific Translations: Jerusalem.

Villa P, Mahieu E. 1991. Breakage patterns of humanlong bones. Journal of Human Evolution 21: 27–48.

Volodicheva N. 2002. The Caucasus. In The PhysicalGeography of Northern Eurasia, Shahgedanova M (ed.).Oxford University Press: Oxford; 350–376.

Voorhies M. 1969. Taphonomy and Population Dynamics ofan Early Pliocene Vertebrate Fauna, Knox Country,Nebraska, University of Wyoming Contributionsto Geology Special Paper No. 1: Laramie.



Appendix

NISP and MNE of Bos/Bison and Caucasian tur bone elements in the UP layers of Dzudzuana Cave(bone counts for identified Bos and Bison elements are given in parentheses).

B C D

Capracaucasica

Bos/Bison Capracaucasica


Bos/Bison

NISP MNE NISP MNE NISP MNE NISP MNE NISP MNE NISP MNE

(a) Lower area

HeadHorn 2 1 2 1 3 1 2 1Sk.frag.-occipital 1 1 2 [0/1] 2 [0/1]Sk.frag.-parietal 1 1Sk.frag.-petrosum 2 2 1 1 1 1 1 1Man. Cond.þ frag. 7 2 12 [1/0] 3 [1/0] 2 2 7 3 1 1Teeth 113 9 32 [12/8] 5 [2/2] 75 6 66 4 18 3 13 3

BodyVer: Cervical 3 2 4 1 3 1 4 2 2 1Ver: Thoracic 1 1 1 1 5 2Ver: Lumbar 3 1 4 1 2 1 11 3 1 1 1 1Ver: caudal 1 1 2 1SacrumSternumRib frag. 30 47 18 36 1 1 1

ForelimbScapula-gle.fos 2 2 3 1 1 1 2 2Scapula-shold.bl. 6 2 1 1 2 2 3 1 2 1Humerus-prox 2 1 3 2Humerus-dis 9 [7/0] 2 [1/0] 4 2 1 1Humerus-shaft 14 3 23 [0/1] 5 [0/1] 17 4 25 [0/1] 4 [0/1]Radius-prox 1 1 1 1 6 [0/1] 4 [0/1] 1 1Radius-dis 1 1 1 1Radius-shaft 18 3 12 2 8 3 8 2 2 1 2 1Ulna-prox 1 1Ulna-distal 2 1 2 [1/0] 2 [1/0] 4 3 5 3Ulna-shaft 2 1 2 1 8 2 1 1Metacarpus-prox 1 1 6 [4/0] 3 [2/0] 9 5 8 3 3 1 1 1Metacarpus-disMetacarpus-shaft 1 [1/0] 1 [1/0] 1 1 1 1Carpal-c 2 2 1 [0/1] 1 [0/1]Carpal-l 3 3Carpal-m 1 1 2 2 2 2 1 1Carpal-u 1 1 1 1 2 2Carpal-s 2 2 1 1

HindlimbAcetabulum-il 2 1 3 2Acetabulum-isch 1 1 2 1 1 1 1 1Acetabulum-pub 1 1 1 1Other pelvic frag. 1 [0/1] 1 [0/1] 1 1Femur-prox 3 1 5 2Femur-dis 1 1 3 1 6 2Femur-shaft 22 4 19 [1/0] 2 [1/0] 17 2 26 [0/3] 4 [0/1] 7 2 1 1Tibia-prox 3 1Tibia-dis 5 2 10 [0/1] 4 [0/1]Tibia-shaft 4 1 31 [2/2] 3 [1/1] 7 2 26 [0/2] 4 [0/1] 1 1 2 1

(Continues)



Appendix (Continued)

B C D

Capracaucasica



Bos/Bison


Patella 1 1 1 1 2 1Astragalus 1 1 4 3 2 2Calcaneum 2 1 4 2 3 2Coboid 1 1 1 1 4 3 3 2Metatarsus-prox 2 1 1 [1/1] 1 1 5 3 1 1Metatarsus-dis 1 1Metatarsus-shaft 1 1 1 [1/1] 1 1 1 1 1 1Tarsal-c 1 1 3 3Tarsal-tdf 1 1 1 1 5 4 1 1

ToesPhalanx 1 6 5 11 [2/1] 4 [2/1] 20 7 12 5 10 5 3 1Phalanx 2 6 4 7 [2/0] 4 [2/0] 14 7 24 9 4 2 1 1Phalanx 3 1 1 2 [1/0] 2 [1/0] 2 1 5 3 3 3 1 1Seasamoid 2 2 6 6 8 7Metapod cond. 1 1 1 1 3 1 10 7 4 4 3 [0/1] 2 [0/1]Metapod -shaft 3 1 10 [1/3] 2 [1/1] 20 3 20 3 4 1 3 1NISP 268 267 278 394 70 40

B1 B2 C

Capracaucasica



Bos/Bison


(b) Upper areaHeadHorn 1 1Sk.frag.-occipitalSk.frag.-parietalSk.frag.-petrosum 2 2 1 1Man. Cond.þ frag. 7 2 3 2 2 1 2 1 1 1 2 1Teeth 63 7 48 [0/4] 4 [0/1] 45 7 28 [0/4] 3 [0/1] 6 1 7 [0/1] 1

BodyVer: Cervical 2 1 4 1 2 1Ver: Thoracic 1 1Ver: Lumbar 3 1 1 1 3 2Ver: caudalSacrumSternumRib frag. 20 15 16 7 1 1 1

ForelimbScapula-gle.fos 1 1 2 2 1 1Scapula-shold.bl. 1 1 2 1 1 1 2 1Humerus-prox 1 1 1 1Humerus-dis 1 1 5 2 1 1 1 1Humerus-shaft 19 6 15 4 6 2 10 3 2 1 5 2Radius-prox 2 2 1 [1/0] 1 [1/0] 1 1 2 1Radius-dis 2 2 1 1Radius-shaft 21 5 6 2 19 6 3 1 1 1 4 2Ulna-proxUlna-distal 1 1 3 2 1 1Ulna-shaft

(Continues)



Appendix (Continued)

B1 B2 C

Capracaucasica



Bos/Bison


Metacarpus-prox 3 2 2 1 2 1Metacarpus-dis 2 1Metacarpus-shaft 2 1 9 2 5 2 2 1 2 1 1 1Carpal-c 1 1Carpal-l 1 1Carpal-m 1 1Carpal-u 1 1 2 2Carpal-s 2 2

HindlimbAcetabulum-il 2 2Acetabulum-isch 2 1 2 1 1 1 1 1 1 1Acetabulum-pub 1 1Other pelvic frag. 1 1 2 1 1 1Femur-prox 1 1 2 [0/1] 2 [0/1] 1 1Femur-dis 2 1 4 1 1 1 1 1Femur-shaft 17 4 12 3 14 2 13 3 6 2Tibia-prox 2 1Tibia-dis 2 2 1 1 1 1 2 1Tibia-shaft 16 4 20 5 7 2 13 2 1 1 9 2Patella 1 1 1 1Astragalus 4 3 1 1 1 1Calcaneum 3 2 2 1 2 2 1 1Coboid 1 1 2 1 1 1Metatarsus-prox 1 1 1 1 2 1 1 1Metatarsus-dis 1 1Metatarsus-shaft 3 1 2 1 1 1 2 1Tarsal-c 1 1 2 1Tarsal-tdf 1 1

ToesPhalanx 1 7 3 6 2 1 1 4 2 3 [0/1] 2Phalanx 2 6 3 10 4 4 3 2 2 2 2Phalanx 3 1 1 1 1 3 2Seasamoid 1 1 1 1Metapod cond. 8 5 3 1 2 1 4 3 2 2Metapod -shaft 11 1 2 1 3 1 3 1 1 1 2 1NISP 246 183 152 24 57



Taphonomy and zooarchaeology of the Upper Palaeolithic cave of Dzudzuana, Republic of Georgia

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Transcript of Taphonomy and zooarchaeology of the Upper Palaeolithic cave of Dzudzuana, Republic of Georgia