Journal of Taphonomy, 3

Transcription

Journal of Taphonomy, 3
P R O ME T H E US P R E S S / P A L A E O N T O L O G I C A L N E T W O R K F O UN D A T I O N
Journal of Taphonomy
(TERUEL)
2005
VOLUME 3
Available online at www.journaltaphonomy.com
Bar-Oz & Adler
(ISSUE 4)
Taphonomic History of the Middle and Upper
Palaeolithic Faunal Assemblage from Ortvale
Klde, Georgian Republic
Guy Bar-Oz*
Zinman Institute of Archaeology, University of Haifa, Mount Carmel, 31905 Haifa, Israel.
Daniel S. Adler
Department of Anthropology, University of Connecticut, 354 Mansfield Rd,Unit 2176,
Storrs, CT 06269. USA.
Journal of Taphonomy 3 (4) (2005), 185-211.
Manuscript received 9 March 2005, revised manuscript accepted 26 May 2005.
We present the results of a detailed taphonomic and zooarchaeological study of the faunal remains from
the late Middle Palaeolithic (LMP) and early Upper Palaeolithic (EUP) bone assemblage of Ortvale
Klde, Georgian Republic. A series of taphonomic tests and analyses are employed to reconstruct the
depositional history of the bone assemblage and investigate LMP (Neanderthal) and EUP (Modern
human) hunting and subsistence strategies. We identify the maximum number of skeletal elements,
document bone surface modifications, the mode of bone fragmentation, and the demographic structure
of the main hunted ungulate population. The assemblage is characterized by significant densitymediated biases, yet in situ attrition and carnivore damage play a minimal role in assemblage formation.
Data suggest that most bone destruction occurred during site occupation, probably in relation to marrow
consumption as indicated by the mode of bone fragmentation. Caucasian tur (Capra caucasica) is the
major prey species throughout the LMP and EUP, and body part representation, the absence of selective
transport, and butchery marks from all stages of carcass processing suggest that Caucasian tur were
subjected to extensive handling. Analysis of Caucasian tur dental eruption and wear indicates that
prime-age adult individuals dominate the assemblage. The results of this study, the first
zooarchaeological and taphonomic study carried out on a Palaeolithic bone assemblage from the
southern Caucasus, indicates that hunting strategies and meat processing behaviors were not
significantly different between Neanderthals and Modern humans
Keywords: TAPHONOMY, ZOOARCHAEOLOGY, CAUCASUS, NEANDERTHALS, MODERN
HUMANS, SUBSISTENCE, MIDDLE–UPPER PALAEOLITHIC, CAPRA CAUCASICA.
Introduction
Zooarchaeological and taphonomic methods
are now recognized as key elements in any
integrated and meaningful study of
Palaeolithic subsistence. Recognition of the
analytical potential of these methods has led
to a profound transformation in our
Article JTa038. All rights reserved.
* E-mail: guybar@research.haifa.ac.il
185
Taphonomy of Ortvale Klde
understanding of extinct human behavioral
variation through time and across space, yet
the careful application of these methods is
still not commonplace in many parts of the
world. Here we report the results of our
recent work in the southern Caucasus,
specifically the Georgian Republic, where
until
now
zooarchaeological
and
taphonomic studies have never been
attempted on Palaeolithic assemblages. Our
research focuses on two Middle and Upper
Palaeolithic sites in western Georgia
(Ortvale Klde and Dzudzuana Cave) and
our goals have been to analyze newly
excavated faunal assemblages to investigate
the foraging behaviors and subsistence
patterns of local Neanderthal and Modern
human populations. Here we provide the
full taphonomic report of our findings at
one of these sites, Ortvale Klde. A detailed
discussion of the significance of the site in
the larger regional and inter-regional
context and to debates concerning the
Middle–Upper Palaeolithic “transition” is
provided in Adler et al. (in press).
The Middle and Upper Palaeolithic
of the Caucasus is increasingly a focal point
of prehistoric research within Eurasia (e.g.,
Cohen & Stepanchuk, 1999; Golovanova et
al., 1999; Golovanova & Doronichev, 2003;
Hoffecker, 1999, 2002; Hoffecker &
Baryshnikov, 1998; Hoffecker & Cleghorn,
2000; Hoffecker et al., 1991; Liubin, 1977,
1989; Tushabramishvili, 1978). The
geographically isolated nature of the
southern Caucasus and its environmental
and climatic diversity make it an ideal
setting within which to investigate
Palaeolithic lifeways, beginning in the
Pliocene/Pleistocene
(e.g.,
Dmanisi)
through to the late Upper Pleistocene (e.g.,
Ortvale Klde and Dzudzuana Cave), but
large-scale, international research projects
have not been attempted until recently (e.g.,
Adler, 2002; Adler et al., in press; Adler &
Tushabramishvili, 2004; Gabunia et al.,
2000; Meshveliani et al., 2004; Tappen et
al., 2002; Vekua et al., 2002).
The specific goals of this paper are
to: a) provide full taphonomic and
zooarchaeological documentation of the
faunal remains; b) reconstruct the
depositional history of the different
occupational phases of the site and to
investigate their economic structure; and c)
gain insights into the subsistence activities
at the site in terms of food transport and
processing, as evidenced by data on skeletal
part representation and specific modes of
bone modification. Ortvale Klde contains
thick, clearly stratified LMP and EUP layers
from which samples of lithic and faunal
material have been excavated using modern
techniques, thus it is currently the key site
within western Georgia and the southern
Caucasus with which to address these issues
and it is the first to be chronometrically
dated using a variety of radiometric
techniques (Adler, 2002; Adler &
Tushabramishvili, 2004; Adler et al., in
press). All of the data are presented by
archaeological layer (Layers 2–7), and in
certain cases, due to small sample sizes,
quantitative analyses are conducted on
combined LMP or EUP assemblages. The
careful technological and typological study
of the lithic artifacts from Ortvale Klde
indicates that these assemblages can be
treated as two distinct LMP and EUP
archaeological units (Adler, 2002). The
faunal collection from Ortvale Klde is
currently curated at the Georgian State
Museum, Tbilisi, Georgian Republic.
186
Bar-Oz & Adler
Site location and research history
Ortvale Klde is situated in western Georgia
(Imereti region), in the Cherula river valley,
approximately 530 meters above sea level
(m.a.s.l.), and 35 meters (m) above the
gorge (Figure 1). It is a karstic rockshelter
comprised of two chambers opening to the
east (ca. 300 m2) (Figure 2). D.
Tushabramishvili first investigated the
rockshelter between 1973 and 1992, and
excavated roughly 40 m2 in the southern
chamber (Tushabramishvili et al., 1999;
Adler & Tushabramishvili, 2004). Faunal
remains from this excavation were
selectively collected and analyzed by A.
Vekua, Georgian Academy of Sciences,
from a palaeontological perspective that
focused on the taxonomic identification of
teeth and selected long bone epiphyses
(Tushabramishvili et al., 1999). Such an
analytical
procedure
prevents
any
reconstruction of skeletal part representation
and the taphonomic history of the
zooarchaeological assemblage. In addition,
bone surface modifications and the
demographic profiles of the dominant
species were not studied.
Vekua determined that the bone
assemblage (N= 19,372) from Ortvale Klde
was heavily dominated by Caucasian tur
(Capra caucasica: 85% ) in each of the
occupational layers. The extinct steppe
bison (Bison priscus: 6% ) and red deer
(Cervus elaphus: 3%) appear in all layers
while other medium and large mammals
such as aurochs (Bos primigenius), wild
boar (Sus scrofa), and roe deer (Capreolus
capreolus), and carnivores such as the
extinct cave bear (Ursus spelaeus), brown
bear (Ursus arctos), wolf (Canis lupus), and
fox (Vulpes vulpes) are represented in small
proportions
(<1%
in
all
layers)
(Tushabramishvili et al., 1999). Vekua
associated these fauna with a forestmountainous environment. He argued that
the remains of Caucasian tur indicate the
site’s proximity to high altitude, sub-Alpine
habitats while roe and red deer indicate
forest biotopes; the bison was well adapted
to both forests and open habitats.
The new excavations at Ortvale
Klde, directed by Daniel S. Adler
(University of Connecticut) and Nicholas
Tushabramishvili (Georgian State Museum)
between 1996 and 2001, were carried out in
6 m2 in the southern chamber of the
rockshelter (Figure 2) (Adler, 2002; Adler
& Tushabramishvili, 2004). A total of 3,209
EUP and 12,541 LMP faunal specimens
were recovered. All of the excavated
sediments were carefully dry-screened
through 2-millimeter (mm) mesh and then
picked for small bones. All bone fragments
were collected and processed according to
their spatial and stratigraphic location. The
faunal remains discussed here were
recovered from three EUP horizons (Layers
2–4; ca. 19.5–34.5 ka Uncal BP) and three
LMP horizons (Layers 5–7; ca. 36.5–42 ka
Uncal BP) (Figure 3) and thus span the
regional transition from the Middle to
Upper Palaeolithic (Adler et al., in press).
Faunal analysis procedures
The procedures followed in the taphonomic
and zooarchaeological analysis include the
following:
1. Maximum length (to the nearest mm) was
measured for all long bone shaft
fragments that were larger than 10 mm
and did not exhibit new excavation
fractures. Measurements recorded to
0.01 mm were made using a digital
caliper (Sylvac model S225), and were
187
Taphonomy of Ortvale Klde
Figure 1. Map of the Imereti region in western Georgia. Archaeological sites spanning the early Middle Palaeolithic
(Djruchula Cave), Middle Palaeolithic (Ortvale Klde, & Samgle Klde), Upper Palaeolithic (Guargilas Klde, Ortvale
Klde, Dzudzuana Cave, Sareki Klde, Samgle Klde & Samertskhle Klde), and the Neolithic (Samele Klde, Sareki Klde,
& Dzudzuana Cave) are italicizes. Modified after Adler (2002).
Figure 2. Site plan of Ortvale Klde, with excavation grid and the position of Profiles I–XI. Areas and seasons of excavation are shaded. Modified after Adler (2002).
188
Bar-Oz & Adler
including cranial fragments, vertebrae,
long bone articular ends and shafts.
Since the majority of bones were highly
fragmented, the identified bone elements
were documented by describing the
specific element, its side, the portion of
the bone (e.g., proximal-distal epiphysis,
medial shaft), and what part of each
portion was represented (e.g., lateral,
medial). Shaft fragments were coded
according to the presence of specific
zones (i.e., proximal shaft, distal shaft,
mid-shaft) or diagnostic features (e.g.,
foramen, muscle attachments; Stiner,
2004). In most cases identified bone
elements were coded according to their
fraction of completeness (i.e., percentage
of complete bone; percentage of bone
shaft connected to an epiphysis). Each of
the identified vertebrae (cervical,
transferred with an OptoRS232C
compatible interface into a standard
Windows Excel worksheet using
OptoFace (version 1.01) software.
2. The state of burning was recorded for
each of the identified and unidentified
fragments larger than 10 mm. Two
categories of burned bone were
recorded: a) partially or completely
carbonized, and b) calcined.
3. Skeletal element and broad taxonomic
identifications were carried out in the
field using a virtual comparative
collection catalogue of recent specimens.
Finer taxonomic identifications of
closely related species were achieved
later with the assistant of A. Vekua and a
comparative collection.
4. Skeletal elements were identified to the
closest possible taxonomic unit,
Figure 3. Profile X, with Layers 1-7 and 11. Layer 1 is historic, Layers 2–4d are EUP, Layers 5–7 are LMP, and Layer
11 is in situ weathered bedrock. Layers 8–10 (LMP) underlie Layer 7 and are located approximately 2 meters north of
this exposure. See Figure 2 for the location of Profile X. Modified after Adler (2002).
189
Taphonomy of Ortvale Klde
thoracic, lumbar, sacrum, caudal) was
classified according to the portion
represented (e.g., centrum, spinous
process, pre or post-zygapophysis).
Carpal and tarsal bones were
documented according to their percent
completeness.
5. Frequencies of element portions were
used to calculate the minimum number
of skeletal elements (MNE), the
minimum number of animal units
(MAU), and the minimum number of
individuals (MNI) following the
protocol of Klein & Cruz-Uribe (1984)
and Lyman (1994). The number of
identified specimens (NISP) was used as
a basic measure of taxonomic abundance
(Grayson, 1984).
6.Elements were examined during fieldwork
for macroscopic surface modifications
using a low-resolution magnifying lamp
(2.5X). Modifications such as bone
weathering
(Behrensmeyer,
1978),
butchery marks (Binford, 1981),
percussion marks (Blumenschine &
Selvaggio, 1988), and evidence of rodent
gnawing, carnivore punctures, scoring,
and digestion, were recorded (Fisher,
1995; Lyman, 1994).
7. The mode of bone fragmentation was
analyzed for a sample of shaft fragments
with attached epiphyses and isolated
shaft fragments. The morphology of the
fracture angle, fracture outline, and
fracture edge was assessed in order to
determine the stage at which the bones
were broken (i.e., fresh vs. dry; see Villa
& Mahieu, 1991). Shaft circumference
was measured to determine the role of
carnivores and to demonstrate that the
assemblage was fully screened and
collected (Bunn, 1983; Marean et al.,
2004).
8. The age structure of Caucasian tur, the
main hunted species, was analyzed on
the basis of tooth eruption and wear
patterns of the lower deciduous fourth
premolar (dP4) and lower third molar
(M3). The M3 erupts at the age of
approximately 36 months after the dP4
has worn down (based on data for Capra
aegagrus, Evins, 1982; Habermehl,
1985, as cited in Kersten, 1987).
According to Heptner et al. (1989) the
average potential lifespan of Caucasian
tur is about 12 years. Therefore, the
‘life’ of the M3 is estimated at 9 years
(from eruption at 3 years, to no crown
height at 12 years), and the ‘life’ of the
dP4 is estimated at 3 years. Following
Munson & Marean (2003) we present
the mortality profiles as 10% increments
of potential lifespan.
9. The relative abundance of males and
females in the Caucasian tur population
was determined using osteometrical
distinctions of selected post-cranial
elements in comparison to recent sexed
specimens from the St. Petersburg
Zoological Institute and the Humboldt
Zoologische Museum, Berlin.
Species representation, mortality, and sex
ratio
A total of 3234 specimens were identified
from a minimum number of 53 individuals
(Table 1). The LMP and EUP inhabitants of
Ortvale Klde subsisted mainly on Caucasian
tur, which is the most common taxon in all
occupation levels (≥90% of NISP in each
layer), and represents the primary prey
species at the site. Bison contributed less
than 7% to each LMP and EUP assemblage.
Other large ungulates (red and roe deer) are
190
Bar-Oz & Adler
Table 1. Species abundance in the EUP and LMP of Ortvale Klde
2
NISP
MNE
MNI
%NISP
Capra
caucasica
19
13
1
90.5
Bison
priscus
2
2
1
9.5
Ursus
sp.
-
Vulpes
vulpes
-
Cervus
elaphus
-
Capreolus
capreolus
-
3
NISP
MNE
MNI
%NISP
33
17
1
86.8
3
2
1
7.9
1
1
1
2.6
1
1
1
2.6
-
-
38
21
4
4
NISP
MNE
MNI
%NISP
324
132
3
90.0
21
13
1
5.8
12
7
2
3.3
1
1
1
0.3
2
2
1
0.6
-
360
155
8
5
NISP
MNE
MNI
%NISP
191
70
4
92.7
15
10
1
7.3
-
-
-
-
206
80
5
6
NISP
MNE
MNI
%NISP
1408
458
14
95.6
57
23
1
3.9
1
1
1
0.1
-
1
1
1
0.1
5
5
1
0.3
1472
488
18
7
NISP
MNE
MNI
%NISP
1098
384
12
96.6
30
18
1
2.6
1
1
1
0.1
-
2
2
1
0.2
6
5
1
0.5
1137
410
16
Total
NISP
MNE
MNI
%NISP
3073
1074
35
95.0
128
68
6
3.9
15
10
5
0.5
2
2
2
0.1
5
5
3
0.2
11
10
2
0.3
3234
1169
53
Layer
E
U
P
L
M
P
nearly absent from the assemblage (<1% in
each layer). Carnivores (Ursus sp. and
Vulpes vulpes) are also represented in small
proportions, but their relative frequencies in
the EUP (Layer 3 and 4) are higher than in
the LMP (Layer 5–7) (Table 1).
The overwhelming dominance of
Total
21
15
2
Caucasian tur at Ortvale Klde differs from
the pattern found at the neighboring and
contemporaneous EUP site of Dzudzuana
Cave where steppe bison are most common
(Bar-Oz et al., 2004a), and currently
represents the only documented occurrance
of a single ungulate species dominating a
191
Taphonomy of Ortvale Klde
Table 2. Percent mortality as 10% increments of
potential lifespan (12 years) for Caucasian tur
from Layers 6–7 (n= 57) at Ortvale Klde. See
also Figure 4.
% Lifespan
0–10
10–20
n
10
8
Prime
20–30
30–40
40–50
50–60
60–70
4
10
11
7
6
Old
70–80
80–90
90–100
0
1
0
Age
Young
LMP and EUP site in the southern
Caucasus. Available data on Caucasian tur
life history and its migratory behavior
(Heptner et al., 1989) indicate its abundance
at lower elevations during winter when
herds aggregate for breeding. During spring
and summer herds fission into small
maternal groups of twelve and males
become solitary. It is at this time that
Caucasian tur move to higher elevations to
exploit alpine pastures. Since Ortvale Klde
is situated at an elevation within the lower
limits of this species’ winter migratory
range, we believe that Caucasian tur was
abundant around the site between late fall
and early spring and thus constituted a
seasonally reliable resource for both LMP
and EUP populations.
The mortality profile of the
Caucasian tur population, determined
according to the dental wear height of the
dP4 (n= 19) and M3 (n= 38) from Layers 6
and 7 (LMP) indicates a hunting preference
for prime-age adults (Table 2). The small
sample size from Layers 4 and 5 (EUP–
LMP) did not permit similar analysis. We
follow Stiner (1994, 2002) Steele & Weaver
(2002) and Steele (2004) and present the
ratios of juvenile, prime-adult, and old
Caucasian tur specimens in a triangular plot
(Figure 4). The mortality pattern shows that
Caucasian tur hunting falls within the
'ambush predator' portion of the triangular
diagram, near the median values obtained
by Stiner (1994) for the Middle Palaeolithic
and Late Upper Palaeolithic of Italy (red
deer, roe deer, wild ass, horse, aurochs,
chamois), by Speth & Tchernov (1998) for
the Middle Palaeolithic layers of Kebara
Cave, Israel (gazelle, fallow deer, red deer),
Figure 4. Modified triangular plot (after Steele &
Weaver, 2002) of (1) Caucasian tur mortality patterns in
the combined LMP sample from Layers 6–7 at Ortvale
Klde (Table 2) compared to the median values for (2) the
Middle and (3) Upper Palaeolithic of Italy (Stiner, 1994,
2002), for the mortality patterns of (4) gazelle, (5) fallow
deer, and (6) red deer in the combined Middle Palaeolithic sample of Kebara Cave (Speth & Tchernov, 1998),
and for (7) the combined Middle Palaeolithic sample
from Gabasa 1, Spain (Blasco Sancho, 1995). Bootstrapping of the raw data produces density contours that represent 95% confidence intervals around each data point
(Steele & Weaver, 2002), with assemblages 1, 4, 5, and 7
having a tighter range than assemblages 2, 3, and 6.
192
Bar-Oz & Adler
Table 3. Measurements of sexually dimorphic elements of Caucasian tur from the LMP and EUP of Ortvale
Klde and modern Caucasian tur from the Caucasus. Measurements are based on the breadth of the distal
condyle of the humerus (BT) and its height (HDH), breadth of the astragalus (Bd) and its length (GL1).
Measurements are in millimeters and taken according to von den Driesch (1976).
Humerus BT
LMP
EUP
Modern Male
Modern Female
Range of Variability
43.1–48.6
41.1–46.3
34.0–37.9
Mean
45.0
43.1
36.4
Std. Dev.
2.5
1.8
1.7
N
4
11
5
Humerus
HDH
LMP
EUP
Modern Male
Modern Female
19.3–21.6
17.3–20.9
15.2–17.9
20.5
18.9
16.9
1.1
1.1
1.0
4
11
5
Astragalus
Bd
LMP
EUP
Modern Male
Modern Female
24.9–31.1
24.9–33.0
24.6–27.6
20.8–23.3
27.4
29.3
25.9
22.3
2.3
4.1
1.1
1.0
13
3
11
4
Astragalus
GL1
LMP
EUP
Modern Male
Modern Female
37.0–46.3
39.8–47.5
36.4–40.3
34.1–36.3
41.4
43.9
38.5
35.3
2.5
3.9
1.2
1.1
13
3
11
4
Table 4. Frequencies of 10 mm size classes of Caucasian tur unidentified long bone shaft fragments from
the EUP and LMP of Ortvale Klde. Fragments smaller than 10 mm were not measured.
mm
10–20
20–30
30–40
40–50
50–60
60–70
70–80
80–90
90–100
100–110
110–120
120–130
130–140
140–150
150–160
160–170
Layer 2
64
78
50
21
11
2
0
0
0
0
0
0
0
0
0
0
EUP
Layer 3
43
67
34
31
20
9
5
2
0
0
0
0
0
0
0
0
Layer 4
643
913
489
239
67
28
12
6
1
0
0
0
0
0
0
0
Layer 5
194
581
342
150
58
17
7
1
0
0
0
0
0
0
0
0
LMP
Layer 6
1398
2157
1049
470
173
75
31
9
5
3
1
0
0
0
0
0
Layer 7
641
974
489
216
102
45
15
6
3
1
1
0
0
0
0
0
Total
226
211
2398
1350
5371
2493
Mean Length
28.2
32.3
27.7
28.4
28.1
28.7
Std. Dev.
11.3
12.7
11.2
11.5
11.8
12.7
193
Taphonomy of Ortvale Klde
Table 5. Frequencies of fracture angle, fracture outline, fracture edge, and shaft
circumference of Caucasian tur-sized elements from the EUP and LMP of Ortvale Klde.
Relative frequencies are indicated parenthetically.
and by Blasco Sancho (1995) for data from
Gabasa 1 in Spain (ibex).
The Caucasian tur is sexually
dimorphic, with adult males being larger
and heavier than adult females (the live
weight of adult males is 65–100 kilograms
(kg) and of females 50–60 kg; Heptner et
al., 1989). The difference in weight is
reflected in the breadth and width of the
portions of some elements; astragali and
distal humeri show a particularly high
degree of sexual dimorphism. The latter
were among the most abundant measurable
skeletal elements in the assemblage. Only
adult specimens (i.e., fused epiphyses and
non-porous astragali) were included in the
analysis. Table 3 shows that all Caucasian
tur recovered from the LMP and EUP layers
at Ortvale Klde are larger than recent
specimens of both sexes of Capra
Caucasica and Capra cylindricornis
collected in the Caucasus at the beginning
of the 20th century (5 females and 11 males
from the St. Petersburg Zoological Institute
and the Humboldt Zoologische Museum,
Berlin). It seems reasonable to assume that
body-size diminution of Caucasian tur is
related to processes of post-glacial warming
and desiccation in accordance to
Bergmann’s (1847) rule. A similar pattern
was observed in numerous Late Pleistocene
Levantine mammals (e.g., Bar-Oz et al.,
2004b; Davis, 1981; Kurten, 1965).
Although the archaeological sample from
Ortvale Klde is small, the broad range of
bone measurements shows that both large
and small specimens are present (Table 3;
Appendix 2). Thus, it appears that both
sexes are represented in the assemblage.
194
Bar-Oz & Adler
Assemblage
fragmentation
completeness
and
Intensity of Bone Fragmentation
The intensity of bone fragmentation is
characterized by the size-range of broken
fragments (Lyman, 1994; Lyman &
O’Brien, 1987). Diaphyses fragments of
various sizes constitute the majority of the
unidentified elements. The lengths of
unidentified long bone shaft fragments are
similar for all occupation levels (Table 4).
The frequency distributions of the length of
Caucasian tur specimens is provided by 10
mm size classes and shows that
approximately 80% of the fragments from
each layer fall within the 10–40 mm size
classes (Figure 5; Table 4) (fragments <10
mm were not measured). The similarity
between layers suggests that each faunal
assemblage experienced a similar degree of
fragmentation (Kruskal-Wallis; H= 9.17, P=
0.10).
Density Mediated Attrition
We found a positive and significant
relationship between Caucasian tur bone
survivorship (%MNI; Lyman, 1994) and
bone structural density (based on Lam et al.,
1999; BMD1 density values for Rangifer
tarandus) within Layers 6 and 7 (LMP)
(Figure 6). The small sample from the EUP
(Layers 2–4) and Layer 5 (LMP) did not
enable similar analyses. The results indicate
a pronounced density-mediated bias in
Caucasian tur skeletal part representation. In
contrast,
an
insignificant
negative
relationship, with a low margin of variance,
was observed between Caucasian tur bone
survivorship and the food utility index
(FUI) (Metcalfe & Jones, 1988; based on
the weight of useable tissue of Rangifer
tarandus) (Figure 6). The lack of
relationship between bone abundance and
the FUI, coupled with a significant and
meaningful relationship between bone
abundance and density suggests that bone
survivorship was primarily affected by
selective density-mediated destruction (see
Grayson, 1988; Klein, 1989; Marean &
Assefa, 1999 for discussion). It follows that
the selective transport of high-utility body
parts was not a significant factor in the
formation of the Ortvale Klde assemblage.
Faunal Transport
Representation
and
Skeletal
Part
Further exploration of bone survivorship
was undertaken through detailed inspection
of Caucasian tur skeletal part profiles.
Figure 7 shows that the representation of
Caucasian tur skeletal elements (%MAU) is
similar in Layers 6 and 7 (LMP) (based on
MNE data from Appendix 1). The body part
profiles show some bias against axial
elements (vertebrae and ribs) and horns, and
a high representation of toe bones.
Statistical comparison of these distributions
indicates no significant differences between
the body part profiles (Kruskal–Wallis; H=
4.98, P= 0.17; based on MNE). The lack of
evidence for selective transport and the low
occurrence of axial units might suggest that
Caucasian tur were subjected to extensive
butchery, either in the field prior to import
to the site or within the site. This is in
accord with Bartram & Marean’s (1999)
observations of Kua hunter-gatherer
treatment of class 3 bovids, equivalent in
size to Caucasian tur, which were
transported either whole in “minimally
disarticulated units” or as extensively
processed carcass parts. According to
195
Taphonomy of Ortvale Klde
50
40
%
30
20
10
10
-2
0
20
-3
0
30
-4
0
40
-5
0
50
-6
0
60
-7
0
70
-8
0
80
-9
0
90
-1
0
10 0
011
11 0
012
12 0
013
13 0
014
14 0
015
15 0
016
16 0
017
0
0
mm
2-UP
3-UP
4-UP
5-MP
6-MP
7-MP
Figure 5. Proportional frequencies of 10 mm size classes of Caucasian tur unidentified long bone shaft fragments
from the EUP and LMP of Ortvale Klde.
A: Layer 6
1.0
B: Layer 7
1.0
y = 1.03x + 0.03
R2 = 0.3617
P<0.001
0.8
y = 1.05x + 0.02
R2 = 0.3265
P<0.01
0.8
0.6
0.6
0.4
0.4
0.2
0.2
0.0
0.0
0
0.1
0.2
0.3
0.4
Bone density
0.5
0.6
0.7
0
0.1
0.2
0.3
0.4
0.5
0.6
0.7
Bone density
D: Layer 7
C: Layer 6
1.0
1.0
y = -0.0011x + 0.5233
R2 = 0.0114
P = 0.37
y = -0.0004x + 0.5159
R2 = 0.0012
P = 0.66
0.8
0.8
0.6
0.6
0.4
0.4
0.2
0.2
0.0
0.0
0
20
40
60
FUI
80
100
0
120
20
40
60
80
100
120
FUI
Figure 6. Relationship between bone density (based on BMD1 values for Rangifer tarandus; Lam et al., 1999) and skeletal part frequency (%MNI) and between food utility (FUI; Metcalfe & Jones, 1988) and skeletal part frequency (%MNI)
for Caucasian tur from Layers 6–7 (LMP) of Ortvale Klde.
196
Bar-Oz & Adler
Figure 7. Skeletal part representation of Caucasian tur from Layers 6–7 (LMP) of Ortvale Klde.
Monahan (1998), the skeletal elements most
frequently transported by the Hadza are
post-cranial axial units, especially the
vertebrae; limbs are transported less often
(see also O’Connell & Hawkes, 1988).
Similar transport patterns were observed
among other contemporary foragers
including the !Kung (Yellen, 1977), the
Nunamiut (Binford, 1978), and the
Kababish nomads (Asher, 1986). In
addition, the bone assemblage from Ortvale
Klde includes a relatively high proportion of
pelves, suggesting that certain axial
elements were transported to the site. At
Hadza base camps, a high frequency of toe
bones in comparison to limb elements was
also observed for class 3 bovids. This
pattern can result from butchering carcasses
at the kill site and carrying the filleted meat
and associated limb bones, hides, and
phalanges back to the site (Monahan, 1998;
O’Connell et al., 1988). Thus it appears that
the inhabitants of Ortvale Klde returned
whole or large portions of Caucasian tur
carcasses to the site rather than a few select
portions of the body. It is tempting to
speculate that the low rate of axial elements
and high frequency of toe bones suggest
that Caucasian tur occasionally underwent
some field butchery before introduction to
the site.…………………………………
Bone damage due to natural taphonomic
agents
In Situ Attrition
Surface modifications on all long-bone
epiphyseal and shaft fragments suggest that
post-depositional agents were not a major
source of bone loss. Ratios of Caucasian tur
197
Taphonomy of Ortvale Klde
100
90
80
9
82 73
41
191
138
70
60
50
40
30
20
1
10
34
8
7 7 11
1 5
5
2 3
0
1
2
10
3
Weathering Stage
L. 2
L. 3
L. 4
L. 5
L. 6
L. 7
Figure 8. Relative frequencies of Caucasian tur long bone shaft fragments per weathering stage from the EUP and
LMP of Ortvale Klde. NISP values are given for each column.
cranial bones to teeth are similar (3/4, 7/10
and 7/11 for Layers 5–7, respectively; based
on MNI for both elements, following Stiner,
1994), suggesting minor (and comparable)
in situ bone loss due to decomposition and/
or advanced physical or chemical
destruction. Similarly, low rates of postburial bone crushing and breakage are
suggested based on Marean's (1991)
completeness index, which investigates the
degree of post-burial attrition using highdensity elements, including the astragalus
and central fourth tarsal. The percentage of
complete astragali (87.5% ) and central
fourth tarsals (80.0% ) in Layer 6 and
astragali (85.3%) and central fourth tarsals
(83.3%) in Layer 7 suggest that the LMP
assemblage of Ortvale Klde did not suffer
considerably from post-depositional decay.
The small sample of tarsal bones from the
EUP layers prevents similar analysis.
Bone Weathering
Caucasian tur long bone shaft fragments
over 50 mm in total length exhibited minor
signs of surface weathering (Figure 8). The
majority of the specimens were attributed to
Behrensmeyer’s (1978) weathering stage 1,
suggesting that bones were buried rapidly
in EUP and LMP contexts and were
exposed to fewer destructive processes
(Andrews, 1995). No surface modifications
typical of bone trampling, for example
shallow parallel scratches (Fiorillo, 1989),
were found in any of the layers.
198
Bar-Oz & Adler
Table 6. Summary of butchery marks on Caucasian tur elements from the EUP and LMP of
Ortvale Klde and activities with which they may be associated (after Binford, 1981).
Abbreviations: vert.= vertebrae; dist.= distal; prox.= proximal; med.= medial; TDF= TarsalDistal Fibula.
199
Taphonomy of Ortvale Klde
of carnivore marks on the inner surface of
Caucasian tur bones (e.g., the occipital part
of the skull) further indicate that carnivores
had access to discarded bones only
following site abandonment by hominins.
We conclude that carnivores were not the
main agents of bone accumulation or
modification of bones at the site, however
we can not reject the hypothesis that
density-mediated bias are partly due to
carnivore ravaging.
Natural experiments show that hyenas
and jackals in the Serengeti typically
scavenge defleshed bones shortly after they
are discarded (Blumenschine, 1988).
Ethnographic studies of the San and Hadza
show a similar pattern in which carnivores
regularly scavenge bones discarded at San
(Bartram et al., 1991) and Hadza (Lupo,
1995) camps. Overall, the low frequency of
carnivore damage likely results from longterm human habitation of the site during
which carnivores could not gain access to
discarded remains (i.e., Bunn, 1993) and/or
the intensive and thorough processing of
faunal remains by hominins, after which
Carnivore Activity
Traces of carnivore activity (e.g., chewing,
gnawing, and scratch marks; Fisher, 1995;
Lyman, 1994) are relatively low (not
exceeding 10%, excluding teeth) among the
identifiable elements and decline in
frequency from Layers 4–7 (Figure 9). In
addition, no signs of digestion were found
on any bones in the assemblage. The nonidentifiable elements (all fragments larger
than 40 mm), comprised mainly of midshafts fragments, show lower rates of
carnivore tooth marks (Figure 9). The low
percentage of carnivore tooth marks on the
mid-shaft portions suggests that carnivores
had only secondary access to bones (see
discussion in Dominguez-Rodrigo, 2002).
The
low
frequency
of
carnivore
modifications indicates that the assemblage
from Ortvale Klde was not accumulated or
significantly altered by carnivores. This
result is in accordance with the high
proportion of fragments with maximum
circumferences less than half of the original
skeletal element (Table 5). Four examples
Table 7. Identified and unidentified burned bones of Caucasian tur and bison from the EUP and LMP of
Ortvale Klde. Relative frequencies are indicated parenthetically.
Unidentified Bone Fragments
Identified Bone Fragments
Capra caucasica
Bison priscus
Capra caucasica
Bison priscus
2
0
0
7/214 (3.3)
0
3
0
0
4/211 (1.9)
0
4
2/140 (1.4)
0
278/2312 (12.0)
2/57 (3.5)
5
4/177 (2.3)
1/11 (9.1)
142/1535 (9.2)
0
6
14/1235 (1.1)
0
576/5639 (10.2)
13/156 (8.3)
7
20/905 (2.2)
1/24 (4.2)
323/2493 (13.0)
8/71 (11.3)
Layer
EUP
LMP
200
Bar-Oz & Adler
10
14/144
9
8
19/123
%NISP
7
64/957
38/705
6
5
4
7/357
3
9/825
2
1
10/433
6/259
2/303
2/237
0
L. 2
L. 3
L. 4
ID
L. 5
L. 6
L. 7
UNID
%NISP
Figure 9. Relative frequencies of carnivore modifications on Caucasian tur identifiable epiphyseal elements (ID) and
unidentified long bone shaft fragments (UNID). NISP values are given for each column.
90
80
70
60
50
40
30
20
10
0
57
58
49
9
8
13
14
12
17
Fresh
Dry
Both
Fracture Angle: Oblique/Right/Both
Fracture Outline: V-shaped/Intermediate/Transverse
Fracture Edge: Jagged/Smooth/Both
Figure 10. Relative frequencies of fracture angle, fracture outline and fracture edge of first phalanges of Caucasian tur
from the LMP of Ortvale Klde. NISP values are given for each column.
201
Taphonomy of Ortvale Klde
10
17/205
15/205
8
25/348
4/61
%NISP
6
9/147
4
6/182
2
5/309
0
Head
Axial
Skeleton
Upper Limb
Lower Limb
Front Limb
Hind Limb
Toes
Layers 6 and 7
Figure 11. Caucasian tur bones from Layer 6 and 7 (LMP) of Ortvale Klde, with butchery marks arranged by anatomical
unit (based on NISP, excluding isolated teeth). NISP values are given for each column.
50
419
40
356
30
20
143
10
0
65
26
4
10-20
20-30
30-40
40-50
50-60
60-70
10
7
70-80
80-90
mm
Figure 12. Proportional frequencies of 10 mm size classes of unidentified burned long bone shaft fragments from the
combined LMP assemblage of Ortvale Klde. NISP values are given for each column (n= 1030).
202
Bar-Oz & Adler
little remained for carnivores to ravage
(Lupo, 1995). Our results are in accordance
with natural experiments and actualistic
studies with captive spotted hyena (Hyaena
hyaena) that show few carnivore tooth
marks on assemblages processed by humans
(Marean & Spencer, 1991; Marean et al.,
1992). Decreasing rates of carnivoredamaged identifiable bone fragments in the
LMP layers at Ortvale Klde may reflect
extended and intensive use of the site by
Neanderthals.
Bone damage due to human subsistence
behaviors
Bone Fragmentation
Analysis of the breakage patterns (fracture
angle, fracture outline, and fracture edge) of
long bone shaft fragments (Villa & Mahieu,
1991) points to the prevalence of oblique,
V-shaped, and jagged bone fractures (Table
5), indicating that the majority of bones
from each layer were broken while fresh.
These data demonstrate that the faunal
assemblage results from intensive bone
processing, possibly related to the extraction
of bone marrow. Such bone processing
behaviors produce high frequencies of
fresh/green fractures and likely account for
the high number of small, unidentified shaft
fragments within each layer. The presence
of percussion fractures (Blumenschine &
Selvaggio, 1988) close to the fracture edge
of long bone shaft fragments (<1% in all
layers: 1/138, 1/177, 5/1297, 7/929 in
Layers 4–7, respectively) and average
fragment length (Table 4; Figure 5) further
support this interpretation. In addition, the
high rate of fresh breakage among the first
phalanges (Figure 10) and the low relative
frequency of complete bones (22.0% of
MNE; following Munro & Bar-Oz, 2005)
suggests that these elements were also
systematically split open for marrow
extraction.
Butchery Marks
Caucasian tur remains preserve evidence for
butchery marks from all stages of carcass
processing (n= 53, combined EUP and LMP
assemblage):
dismemberment (62.3% ),
filleting (18.9% ), and skinning (3.8% )
(Table 6). Butchery marks for which a
related activity could not be determined
using Binford’s cut marks typology (1981)
were included in the “other” category
(15.1% ). Only three unidentified shaft
fragments from the small bison assemblage
bore cut marks.
While the proportion of butchery
marks on Caucasian tur is low for all
archaeological layers (<3%), the presence of
cut marks from all stages of the butchering
process suggests that the full range of
butchering activities occurred within each
layer. Overall, the percentage of butchery
marks does not exceed 20% of the NISP for
any bone element, and is usually below 10%
of the NISP for identifiable elements. The
most obvious similarity in butchery patterns
from each assemblage is the location of cut
marks on and around the major limb joints.
Figure 11 shows the proportional
distribution of cut marks by anatomical unit
for Caucasian tur and indicates higher
frequencies of cut marks on the head and
limb bones, and fewer cut marks on axial
elements and foot bones. However, no
difference is observed between the less
meaty lower limbs and the meatier upper
limbs. These data indicate that Caucasian
203
Taphonomy of Ortvale Klde
tur
carcasses
underwent
thorough
dismemberment and preparation following
arrival on site, and that carcass processing
included both low and high utility parts.
Burning
Burned specimens were observed in all of
the archaeological horizons and appear
randomly distributed. Layers 2 and 3 (EUP)
are characterized by low rates of burning
(2–3%) while Layer 4 (EUP) and the Layers
5–7 (LMP) exhibit higher rates of burning
(<13%; Table 7). The low frequency of
burned bones in Layers 2 and 3 (EUP) is
similar to the rate of burning among the
lithics, suggesting more ephemeral use of
the site during later phases of the EUP. The
frequencies of burned bones among the
identified Caucasian tur and bison remains
are too small to analyze by anatomical unit.
Burning was most common on small
fragments (n= 1030, mean= 25 mm, sd= 11
mm) in the unidentified fraction from the
combined LMP sample (Figure 12). As
argued by Speth (in Bar-Yosef et al., 1992)
the relationship between burning and
specimen size support the view that burning
is related to food preparation; if bones were
accidentally burned by later activities, larger
bones with bigger surface areas should
display high rates of burning (Stiner et al.,
1995).
Discussion
The LMP and EUP occupants of Ortvale
Klde subsisted primarily on prime-aged
Caucasian tur. A hunting pattern focusing
on prime age adults has been identified at
numerous LMP sites across Eurasia and
appears to be a characteristic of this period.
The taphonomic analysis of Ortvale Klde
suggests minor loss of bone due to various
post-depositional processes. High rates of
fragmentation and green bone fractures
indicate that most of the bone destruction
occurred while the site was occupied,
probably as a result of processing bones for
marrow. The presence of cut marks from all
stages of butchery, the absence of selective
transport, and the body part data suggest
that processing of Caucasian tur carcasses
was carried out in the vicinity of the site.
The low rate of axial elements and high
frequency of toe bones might suggest that
Caucasian tur also occasionally underwent
field butchery before introduction to the
site. Since the inhabitants of Ortvale Klde
likely returned whole or large portions of
prime-age adult Caucasian tur carcasses to
the site rather than a few select portions of
the body, we conclude that the net
nutritional gains must have out-weighed
transport costs. Heptner et al. (1989)
estimate that recent adult Caucasian tur can
yield 40–55 kg of dressed meat, and that
individuals caught before the rut
(November), when they are fattest, yield up
to 60% of total weight in meat, 4% in edible
viscera (e.g., liver, kidney, etc.), 19% in
stomach and intestines, and 17% in heads,
legs, and hide.
The formation of the faunal
assemblages in each EUP and LMP layer at
Ortvale Klde is heavily influenced by
human
subsistence
behaviors.
The
taphonomic history of each assemblage, as
reflected
by
patterns
of
species
representation, mortality profile, and the
degree of bone fragmentation are similar.
The mild effects of weathering, and the
relatively low frequency of carnivore
damage,
coupled
with
the
high
204
Bar-Oz & Adler
completeness of cranial and tarsal elements,
and the low proportion of dry bone
fractures, confirm the hypothesis that much
of the bone-density mediated destruction is
related to human agency. Our taphonomic
analysis
also
reveals
considerable
similarities between the EUP and LMP
assemblages in terms of human food
transport, consumption, and processing
behaviors. Similar patterns are now
routinely documented at many Middle and
Upper Palaeolithic sites across Eurasia (e.g.,
Burke,
2000
and papers
therein;
Gaudzinski, 1996; Grayson & Delpech,
2002, 2003; Jaubert et al., 1990; Marean &
Assefa, 1999; Roebroeks, 2001; Speth &
Tchernov, 2001; Stiner, 1994, 2002; Stiner
& Kuhn, 1992; Villa et al., 2004) Given
these results, we believe that the
zooarchaeological data from Ortvale Klde
provide no evidence for shifts in human
hunting strategies and meat processing
behaviors between the LMP (Neanderthals)
and the EUP (Modern humans) (see also
Bar-Yosef, 2004). Hence LMP and EUP
faunal assemblages are poor proxies for
behavioral
differences
between
Neanderthals and Modern humans, a
conclusion that has important implications
for debates surrounding the Middle–Upper
Palaeolithic transition and which domains
of human behavior should be considered
“modern.”
Acknowledgements
We are grateful to our colleagues at the
Georgia State Museum: D. Lordkipanidze,
A. Vekua, N. Tushabramishvili, T.
Meshveliani and N. Jakeli for their
collaboration and assistance during our
many years of research in the Georgian
Republic. We also thank Ofer Bar-Yosef
and Richard Meadow from Harvard
University and Anna Belfer-Cohen from the
Hebrew University for important advice,
support and many profitable discussions and
encouragements
throughout
the
development of this research. G. Bar-Oz
thanks the American School of Prehistoric
Research, Harvard University, for a postdoctoral fellowship during which much of
the zooarchaeological analyses were
conducted, to Leonid Vishniatsky for his
hospitality during his visit to St. Petersburg,
and the Humbolt University, Berlin, and the
Zoological Institute, St. Petersburg, for
access to their collection of modern tur
specimens. D. S. Adler thanks the WennerGren Foundation for Anthropological
Research (Grants 6881 and 7059), the L. S.
B. Leakey Foundation, the American
School of Prehistoric Research at Harvard
University, and the Davis Center for
Russian Studies at Harvard University for
their generous financial support for
fieldwork and analyses. Finally, thanks are
due to N. Munro for providing thoughtful
comments on an earlier draft of this text.
References
Adler, D. S. (2002). Late Middle Palaeolithic Patterns
of Lithic Reduction, Mobility, and Land Use in the
Southern
Caucasus.
Ph.D.
Dissertation,
Department of Anthropology, Harvard University,
Cambridge.
Adler, D. S., Bar-Oz, G., Belfer-Cohen, A. & BarYosef, O. (2005). Ahead of the game: Middle and
Upper Palaeolithic hunting behaviors in the
southern Caucasus. Current Anthropology, in
Press.
Adler, D. S. & Tushabramishvili, N. (2004). Middle
Palaeolithic patterns of settlement and subsistence
in the southern Caucasus. In (Conard, N. ed.)
Middle Palaeolithic Settlement Dynamics.
Tübingen: Publications in Prehistory, Kerns
205
Taphonomy of Ortvale Klde
Verlag, pp. 91–132.
Andrews, P. (1995). Experiments in taphonomy.
Journal of Archaeological Science, 22: 147-153.
Asher, M. (1986). A Desert Dies. London: Viking.
Bar-Oz, G., Adler, D. S., Vekua, A., Meshveliani, T.,
Tushabramishvili, N., Belfer-Cohen, A. & BarYosef, O. (2004a). Faunal exploitation patterns
along the southern slopes of the Caucasus during
the Late Middle and Early Upper Palaeolithic. In
(Mondini, M., Munoz, S. & Wickler, S., eds.)
Colonisation, Migration, and Marginal Areas: A
Zooarchaeological Approach. Oxford: Oxbow
Books, pp. 46–54.
Bar-Oz, G., Dayan, T., Weinstein-Evron, M. and
Kaufman, D. (2004b). The Natufian economy at
el-Wad Terrace with special reference to gazelle
exploitation patterns. Journal of Archaeological
Science, 31: 217–231.
Bartram, L. E. & Marean, C. W. (1999). Explaining the
“Klasies Pattern”: Kua ethnoarchaeology, the Die
Kelders Middle Stone Age archaeofauna, long
bone fragmentation and carnivore ravaging.
Journal of Archaeological Science, 26: 9–20.
Bartram, L. E., Kroll, E. M. & Bunn, H. T. (1991).
Variability in camp structure and bone food refuse
patterning at Kua San camps. In (Kroll, E. M. &
Price, T. D., eds.) The interpretation of
archaeological spatial patterning. New York:
Plenum Press, pp. 77–148.
Bar-Yosef, O. (2004). Eat what is there: hunting and
gathering in the world of Neanderthals and their
neighbors.
International
Journal
of
Osteoarchaeology, 14: 333–342.
Bar-Yosef, O., Vandermeersch, B., Arensburg, B.,
Belfer-Cohen, A., Goldberg, P., Laville, H.,
Meignen, L., Rak, Y., Speth, J. D., Tchernov, E.
Tillier, A. M. and Weiner, S. (1992). The
excavations in Kebara Cave, Mount Carmel.
Current Anthropology, 33: 497–550.
Behrensmeyer, A. K. (1978). Taphonomic &
ecological information from bone weathering.
Paleobiology, 4: 150–162.
Bergmann, C. (1847). Ueber die verheltnisse der
warmerkonomie de thiere zu iherer grosse.
Gottingen Studien, 3: 595–708.
Binford, L. R. (1978). Nunamiut ethnoarchaeology.
Academic Press, New York.
Binford, L. R. (1981). Bones: ancient men and modern
myths. Academic Press, New York.
Blasco Sancho, M. F. (1995). Hombres, fieras y presas.
Estudio arqueozoológico y tafonómico del
yacimiento del Paleolítico Medio Cueva de
Gabasa 1(Huesca). Monografías Arqueológicas
38, Zaragoza.
Blumenschine, R. J. (1988). An experimental model of
the timing of hominid and carnivore influence on
archaeological bone assemblages. Journal of
Archaeological Science, 15: 483–502.
Blumenschine, R. J. & Selvaggio, M. (1988).
Percussion marks on bone surface as a new
diagnostic of human behavior. Nature, 333: 763–
765.
Bunn, H. T. (1983). Evidence on the diet & subsistence
patterns of Plio-Pleistocene hominids at Koobi
Fora, Kenya, & Olduvai Gorge, Tanzania. In
(Clutton-Brock, J. & Grigson, C., eds.) Animals &
archaeology: 1. hunters & their prey. Oxford:
British Archaeological Report, pp. 21–30.
Bunn, H. T. (1993). Bone assemblages at base camps:
a further consideration of carcass transport and
bone destruction by the Hadza. In (Hudson, J., ed.)
From bones to behavior: ethnoarchaeological and
experimental contributions to the interpretation of
faunal remains. Center for archaeological
investigations. Occasional paper No. 21, Illinois:
Southern Illinois University, pp. 156–168.
Burke, A. (Editor) (2000). Hunting in the Middle
Palaeolithic.
International
Journal
of
Osteoarchaeology, 10: 281–285.
Cohen, V. & Stephanchuk, V. N. (1999). Late Middle
and Early Upper Paleolithic evidence from the east
European Plain and Caucasus: a new look at
variability, interactions, and transitions. Journal of
World Prehistory, 13(3): 265-–319.
Davis, S. J. M. (1981). The effect of temperature
change and domestication on the body size of late
Pleistocene to Holocene mammals of Israel.
Paleobiology, 7: 101–114.
Domínguez-Rodrigo, M. (2002).
Hunting and
scavenging by early humans: the state of the
debate. Journal of World Prehistory, 16: 1–54.
Driesch, A. von den. (1976). A guide to a measurement
of animal bones from archaeological sites.
Peabody Museum Bulletins 1, Peabody Museum
of Archaeology and Ethnology, Cambridge.
Evins, M. A. (1982). The fauna from Shanidar Cave:
Mousterian wild goat exploitation in northeastern
Iraq. Paléorient, 8: 37–58.
Fiorillo, A. R. (1989). An experimental study of
trampling: implication for the fossil record. In
(Bonnichsen, R. & Sorg, M. H., eds.) Bone
modification. Orono: University of Maine Center
for the Study of the First Americans, pp. 61–71.
Fisher, J. W. (1995). Bone surface modifications in
zooarchaeology. Journal of Archaeological
Method and Theory, 2: 7–68.
Gaudzinski, S. (1996). On bovid assemblages and their
consequences for for the knowledge of subsistence
206
Bar-Oz & Adler
patterns in the Middle Palaeolithic. Proceedings of
the London Prehistoric Societies, 62: 19-39.
Gabunia, L. K., Vekua, A. K., Lordkipanidze, D.,
Swisher III, C. C., Ferring, R., Justus, A.,
Nioradze, M., Tvalchrelidze, M., Anton, S. C.,
Bosinski, G., Jöris, O., de Lumley, M-A.,
Majsuradze, G. & Mouskhelishvili, A. (2000).
Earliest Pleistocene Cranial Remains from
Dmanisi, Republic of Georgia: Taxonomy,
Geological Setting, and Age. Science, 288: 1019–
1025.
Golovanova, L. V. & Doronichev, V. B. (2003). The
Middle Paleolithic of the Caucasus. Journal of
World Prehistory, 17: 71–140.
Golovanova, L. V., Hoffecker, J. F., Kharitonov, V. M.
& Romanova, G. P. (1999). Mezmaiskaya Cave: a
Neanderthal occupation in the northern Caucasus.
Current Anthropology, 40(1): 77–86.
Grayson, D. K. (1984). Quantitative zooarchaeology.
Academic Press, New York.
Grayson, D. K. (1988). Danger Cave, Last Supper
Cave, and Hanging Rockshelter: the faunas.
Anthropological papers of the American Museum
of Natural History, 66: 1–130.
Grayson, D. K. & Delpech, F. (2002). Specialized early
Upper Palaeolithic hunters in southwestern
France? Journal of Archaeological Science, 29:
1439–1449.
Grayson, D. K. & Delpech, F. (2003). Ungulates and
the Middle-to-Upper Paleolithic transition at
Grotte XVI (Dordogne, France). Journal of
Archaeological Science, 30: 1633–1648.
Habermehl, K.-H. 1985. Altersbestimmung bei Wildund Pelztieren, 2nd Auflage. Verlag Paul Parey,
Berlin and Hamburg.
Heptner, V. G., Nasimovich, A. A. & Bannikov, A.
G. (1989). Mammals of the Soviet Union, vol. 1:
ungulates. E. J. Brill, Leiden.
Hoffecker, J. F. 1999. Neanderthals and Modern
Humans in eastern Europe. Evolutionary
Anthropology, 7(4): 129–141.
Hoffecker, J. F. (2002). Desolate landscapes: Ice-Age
settlement in eastern Europe. Rutgers University
Press, New Brunswick.
Hoffecker, J. F. & Baryshnikov, G. (1998).
Neanderthal ecology in the northwestern
Caucasus: faunal remains from the Borisovskoe
Gorge site. In (Saunders J. J., Styles, B. W. &
Baryshnikov, G., eds.) Quaternary paleozoology
in the northern hemisphere. Springfield: Illinois
State Museum, pp. 187–211.
Hoffecker, J. F., Baryshnikov, G. & Potapova, O.
(1991). Vertebrate remains from the Mousterian
site of Il'skaya I (northern Caucasus, U.S.S.R.):
new analysis and interpretations. Journal of
Archaeological Science, 18: 113–147.
Hoffecker, J. F. & Cleghorn, N. (2000). Mousterian
hunting patterns in the northwestern Caucasus and
the ecology of the Neanderthals. International
Journal of Osteoarchaeology, 10: 368–378.
Jaubert, J. M., Lorblanchet, H., Laville, R., SlotMoller, A. & Brugal, J. P. (1990). Les chasseurs
d'aurochs de la Borde: un site de Paleolithique
moyen. Maison de Sciences de l'Homme, Vol. 1,
Documents d'Archaeologie Francaise. Paris.
Kersten, A. M. P. (1987). Age and sex composition of
Epipalaeolithic fallow deer and wild goat from
Ksar 'Akil. Palaeohistoria, 29: 119–131.
Klein, R. G. (1989). Why does skeletal part
representation differ between smaller and larger
bovids at Klasies River Mouth and other
archaeological sites? Journal of Archaeological
Science, 16: 363–381.
Klein, R. G. and Cruz-Uribe, K. (1984). The analysis
of animal bones from archaeological sites.
University Chicago Press, Chicago.
Kurten, B. (1965). Canivora of the Palestine caves.
Acta Zoologica Fennica, 107: 1–74.
Lam, Y. M., Chen, X. & Pearson, O. M. (1999).
Intertaxonomic variability in patterns of bone
density and the differential representation of bovid,
cervid, and equid elements in the archaeological
record. American Antiquity, 64: 343–362.
Liubin, V. P. (1977). Mustierskie kul'turi Kavkaza
(Mousterian cultures of the Caucasus). Nauka,
Leningrad.
Liubin, V. P. (1989). Paleolit Kavkaza (Paleolithic of
the Caucasus). In (Boriskovski, P. I., ed.) Paleolit
Kavkaza i Sredney Azii (The Palaeolithic of the
Caucasus and northern Asia). Leningrad: Nauka,
pp. 9–144.
Lupo, K. D. (1995). Hadza bone assemblages and
hyena attrition: an ethnographic example of the
influence of cooking and mode of discard on the
intensity of scavenger ravaging. Journal of
Anthropological Archaeology, 14: 288–314.
Lyman, R. L. (1994). Vertebrate taphonomy.
Cambridge University Press, Cambridge.
Lyman, R. L. & O’Brien, (1987). Plow-zone
archaeology: fragmentation and identifiability.
Journal of Field Archaeology, 14: 493–498.
Marean, C. W. (1991). Measuring the postdepositional
destruction of bone in archaeological assemblages.
Journal of Archaeological Science, 18: 677–694.
Marean, C. W. & Assefa, Z. (1999). Zooarchaeological
evidence for the faunal exploitation behavior of
Neandertals and Early Modern Humans.
Evolutionary Anthropology, 8: 22–37.
207
Taphonomy of Ortvale Klde
Marean, C. W. & Spencer, L. M. (1991). Impact of
carnivore ravaging on zooarchaeological measures
of element abundant. American Antiquity, 56: 645–
658.
Marean, C. W., Spencer, L. M., Blumenschine, R. J.
and Capaldo, S. (1992). Captive hyena bone choice
and destruction, the Schlepp Effect, and Olduvai
archaeofaunas. Journal of Archaeological Science,
19: 101–121.
Marean, C. W., Domínguez-Rodrigo, M. & Pickering,
T. R. (2004). Skeletal element equifinality in
zooarchaeology begins with method: the evolution
and status of the “shaft critique”. Journal of
Taphonomy, 2: 69–98.
Meshveliani, T., Bar-Yosef, O. & Belfer-Cohen, A.
(2004). The Upper Paleolithic of Western Georgia.
In Brantigham P. J., Kuhn, S. L. & Kerry, K. W.,
eds.) The Early Upper Paleolithic beyond western
Europe. Berkeley: University of California Press,
pp. 129–143.
Metcalfe, D. & Jones, K. T. (1988). A reconsideration
of animal body part utility indices. American
Antiquity, 53: 486–504.
Monahan, C. M. (1998). The Hadza carcass transport
debate
revisited
and
its
archaeological
implications. Journal of Archaeological Science,
25: 405–24.
Munro, N. D. & Bar-Oz, G. (2005). Gazelle bone fat
processing in the Levantine Epipalaeolithic.
Journal of Archaeological Science, 32: 223–239.
Munson, P. J. & Marean, C. W. (2003). Adults only? A
reconsideration of Middle Palaeolithic ‘prime
dominated’ reindeer hunting at Salzgitter
Lebenstedt. Journal of Human Evolution, 44: 263–
273.
O’Connell, J. F. & Hawkes, K. (1988). Hadza hunting,
butchering, and bone transport and their
archaeological
implications.
Journal
of
Anthropological Research, 44: 113–161.
O’Connell, J. F., Hawkes, K. & Blurton Jones, N.
(1988). Hadza hunting, butchering, and bone
transport and their archaeological implications.
Journal of Anthropological Research, 44: 113–
161.
Roebroeks, W. (2001). Hominid behavior and the
earliest occupation of Europe: an exploration.
Journal of Human Evolution, 41: 437-461.
Speth, J. D. & Tchernov, E. (1998). The role of hunting
and scavenging in Neanderthal procurement
strategies: new evidence from Kebara Cave
(Israel). In (Akazawa T., Aoki, K. & Bar-Yosef,
O., eds.) Neanderthals and Modern Human in
western Asia. New York: Plenum Press, pp. 223–
240.
Speth, J. D. & Tchernov, E. (2001). Neandertal hunting
and meat processing in the Near East, evidence
from Kebara Cave (Israel). In (Stanford, C. B. &
Bunn H. T., eds.) Meat eating and human
evolution. Oxford and New York: The Human
Evolution Series, Oxford University Press, pp. 52–
72.
Steele, T. E. (2004). Variation in mortality profiles of
red deer (Cervus elaphus) in Middle Palaeolithic
assemblages from western Europe. International
Journal of Osteoarchaeology, 14: 307–320.
Steele, T. E. & Weaver, T. D. (2002). The Modified
Triangular Graph: A Refined Method for
Comparing Mortality Profiles in Archaeological
Samples. Journal of Archaeological Science, 29:
317–322.
Stiner, M. C. (1994). Honor among thieves: a
zooarchaeological study of Neanderthal ecology.
Princeton University Press, Princeton.
Stiner, M. C. (2002). Carnivory, coevolution, and the
geographic spread of the genus Homo. Journal of
Archaeological Research, 10: 1–63.
Stiner, M. C. & Kuhn, S. L. (1992). Subsistence,
technology, and adaptive variation in Middle
Paleolithic Italy. American Anthropologist, 94:
306–339.
Stiner, M. C., Kuhn, S. L., Weiner, S. & Bar-Yosef, O.
(1995). Differential burning, recrystallization, and
fragmentation of archaeological bone. Journal of
Archaeological Science, 22: 223–237.
Tappen, M., Adler, D. S., Ferring, C. R., Gabunia, M.,
Vekua, A. & Swisher III, C. C. (2002).
Akhalkalaki: the taphonomy of an early
Pleistocene locality in the Republic of Georgia.
Journal of Archaeological Science, 29: 1367–
1391.
Tushabramashvili,
D.
M.
(Editor)
(1978).
Arkheologicheskie panyatniki Tsutskhvatskogo
mnogoetajnogo
peshernogo
kompleska
(Archaeological sites of the Tsutskhvati Cave
Complex). Metsniereba, Tbilisi.
Tushabramashvili, N., Lordkipanidze, D., Vekua, A.,
Tvalcherlidze, M., Muskhelishvili, A. & Adler, D.
S. (1999). The Middle Palaeolithic rockshelter of
Ortvale Klde, Imereti Region, the Georgian
Republic. Préhistoire Europeenne, 15: 65–77.
Vekua, A. K., Lordkipanidze, D., Rightmire, G. P.,
Agusti, J., Ferring, R., Maisuradze, G.,
Mouskhelishvili, A., Nioradze, M., Ponce de Leon,
M., Tappen, M., Tvalchrelidze, M. & Zollikofer,
C. (2002). A New Skull of Early Homo from
Dmanisi, Georgia. Science, 297 (5): 85–89.
Villa, P., Castel, J, C., Beauval, C., Bourdillat, V. &
Goldberg, P. (2004). Human and carnivore sites in
208
Bar-Oz & Adler
the European Middle and Upper Paleolithic:
similarities and differences in bone modification
and fragmentation. Revue de Paleobiologie,
Geneve, 23: 705-730.
Villa, P. & Mahieu, E. (1991). Breakage patterns of
human long bones. Journal of Human Evolution,
21: 27–48.
Yellen, J. E. (1977). Cultural patterning in faunal
remains: evidence from the Kung! Busmen. In
(Ingerssol, D., Yellen, J. E. & Macdonald, W.,
eds.) Experimental archaeology. New York:
Columbia University Press, pp. 271–331.
Appendix 1. Number of identified specimens (NISP) and minimum number of elements (MNE) of Caucasian
tur elements and bone portions in Layer 4 (EUP) and Layers 5–7 (LMP) at Ortvale Klde.
BODY PART
HEAD
Horn/Antler
Skull fragment-Frontal
Skull fragment-Occipital
Skull fragment-Parietal
Skull fragment-Petrosum
Mandible fragment- Condyle
Teeth
BODY
Vertebrae: Cervical
Vertebrae: Thoracic
Vertebrae: Lumbar
Vertebrae: Caudal
Sacrum
Sternum
Rib fragment
FORELIMB
Scapula-glenoid fossa
Scapula-shoulder blade
Humerus-proximal
Humerus-distal
Humerus-shaft
Radius-proximal
Radius-distal
Radius-shaft
Ulna-proximal
Ulna-distal
Ulna-shaft
Metacarpus-proximal
Metacarpus-distal
Metacarpus-shaft
Carpal-Cuneiform
Carpal-Lunate
Carpal-Magnum
Carpal-Scaphoid
Carpal-Unciform
EUP
Layer 4
NISP
MNE
Layer 5
NISP
MNE
LMP
Layer 6
NISP
MNE
Layer 7
NISP
MNE
1
2
1
4
2
56
1
1
1
3
1
3
1
2
8
54
1
1
3
4
5
5
7
4
10
22
319
2
2
2
1
5
7
10
1
1
4
3
5
19
223
1
1
2
1
3
7
11
6
9
19
16
1
3
4
5
14
3
14
5
2
1
1
1
57
26
95
3
1
7
86
11
7
23
3
1
3
32
54
30
71
1
2
3
72
7
9
17
1
2
1
22
1
3
4
8
3
4
8
2
1
3
1
2
1
1
1
1
2
2
1
3
1
1
1
6
1
1
5
1
-
1
1
1
2
1
1
2
1
-
11
8
11
42
10
8
7
24
2
10
2
13
2
1
4
1
1
1
3
5
3
3
3
4
4
4
3
1
4
1
6
2
1
2
1
1
1
3
5
4
11
8
21
8
3
19
7
1
10
1
1
3
3
7
1
5
3
1
5
2
3
2
1
2
3
1
3
1
1
3
3
6
1
5
209
Taphonomy of Ortvale Klde
BODY PART
HIND LIMB
Acetabulum-ilium
Acetabulum-ischium
Acetabulum-pubis
Other pelvic fragment
Femur-proximal
Femur-distal
Femur-shaft
Tibia-proximal
Tibia-distal
Tibia-shaft
Patella
Astragalus
Calcaneum
Central fourth tarsal
Metatarsus-proximal
Metatarsus-distal
Metatarsus-shaft
Tarsal-Cuneiform
Tarsal-Distal Fibula
TOES
Phalanx 1
Phalanx 2
Phalanx 3
Seasamoid
Metapod condyle
Metapod-shaft
TOTAL
MNI
EUP
Layer 4
NISP
MNE
Layer 5
NISP
MNE
LMP
Layer 6
NISP
MNE
Layer 7
NISP
MNE
2
4
1
1
1
13
1
6
3
3
1
5
3
1
2
1
1
1
2
1
2
2
1
1
2
3
2
1
1
1
2
1
2
1
4
1
3
1
1
1
1
1
1
1
1
1
1
1
1
1
6
12
14
1
11
15
42
5
17
25
3
8
14
10
25
2
3
4
3
5
7
1
7
3
3
2
8
4
2
6
4
9
13
2
3
4
5
10
6
1
10
7
21
2
8
23
3
7
17
6
18
1
2
4
2
2
5
5
1
6
2
2
1
3
3
3
6
5
5
8
1
1
3
2
40
28
7
36
9
7
324
16
11
6
36
4
1
132
23
15
2
5
6
2
191
14
10
1
5
3
1
70
139
100
36
36
58
14
1408
67
47
27
36
32
2
458
102
86
26
44
50
30
1098
51
40
21
44
28
5
384
3
4
210
14
12
Bar-Oz & Adler
Appendix 2. Caucasian tur bone measurements from the EUP and LMP at Ortvale Klde and modern
Caucasian tur from the Caucasus. The measurements are based on the breadth of the distal condyle of
the humerus (BT) and its height (HDH), breadth of the astragalus (Bd) and its length (GL1). Measurements are in millimeters and taken according to von den Driesch (1976)
211
Taphonomy of Ortvale Klde
212