International Journal of Osteoarchaeology

Int. J. Osteoarchaeol. 15: 84–105 (2005)

Published online in Wiley InterScience (www.interscience.wiley.com). DOI: 10.1002/oa.736

Late Mousterian Subsistence and Cave

Use in Dalmatia: the Zooarchaeology of
Mujina Pec¤ina,Croatia
P. MIRACLE*
Department of Archaeology, University of Cambridge, Cambridge CB2 3DZ, UK

ABSTRACT The faunal assemblage from Mujina Pećina provides an initial glimpse of Late Middle
Palaeolithic food procurement, management, and site use, in Dalmatia, Croatia. Radiometric
dates place the entire sequence at about 42 kyr in the middle of OIS 3 (Oxygen Isotope Stage 3)
(Rink et al., 2002). Mujina Pećina is located along a potential migration corridor for hominin
populations moving into Europe from western Asia. The faunal composition shifts from a co-
dominance of red deer and chamois þ ibex in Layer D1 þ D2 that formed during relatively cold
conditions to a clear dominance of wild caprids followed by large bovids and equids in Layer
B þ C that formed during relatively warm conditions. Although non-hominin carnivores played
a significant role in the modification of the faunal assemblages throughout the stratigraphic
sequence, the Late Mousterian faunal assemblage from Mujina Pećina shows the hominins to
be competent hunters within a context of considerable competition from non-hominin
carnivores. These ‘mixed’ hominin-carnivore signatures are pulled apart through a detailed
taphonomic analysis of this well-excavated assemblage. The Mujina Pećina assemblage thus
provides a significant point of reference for a broader-scale study of variability in Mousterian
subsistence practices in their own right as well as within the context of the Initial Upper
Palaeolithic in southeastern Europe. Copyright ß 2005 John Wiley & Sons, Ltd.

Key words: zooarchaeology; Middle Palaeolithic; Neanderthal; Croatia; Europe; hunting;

scavenging

Introduction with the procurement, consumption, and deposi-

Reconstructions of Middle Palaeolithic subsis-
tence practices have figured prominently in
recent debates over the cultural capacities of
‘archaic’ hominins (Burke, 2000a; Gaudzinski,
1999; Marean & Assefa, 1999; Speth & Tchernov,
2001; Stiner 1994). These debates both drew on
and drove research on vertebrate taphonomy
(e.g. Binford, 1981; Brain, 1981). Few palae-
oanthropologists today accept a simple associa-
tion between lithic artefacts and animal remains
as evidence that hominins had anything to do
* Correspondence to: Department of Archaeology, University of
Cambridge, Cambridge CB2 3DZ, UK.
e-mail: ptm21@cam.ac.uk
tion of carcass parts at a given location. Research-
ers venture inferences about Middle Palaeolithic
subsistence only when armed with detailed
taphonomic information about the agents of
accumulation and modification of vertebrate
assemblages. At the forefront has been a debate
over the extent to which Middle Palaeolithic
hominins (assumed to be Neanderthals in Eur-
ope) obtained animal protein from medium- to
large-sized ungulates by scavenging vs. hunting.
Initially these procurement strategies were con-
sidered in opposition to one another; either
Neanderthals scavenged or hunted their prey
(Binford, 1985; Chase, 1988). Second-generation
models posited a ‘scavenging niche’ and exam-
ined the conditions and factors under which

Copyright # 2005 John Wiley & Sons, Ltd. Received 14 March 2003
Revised 30 October 2003
Accepted 25 November 2003
Late Mousterian Subsistence in Dalmatia
Neanderthals occupied it (Stiner, 1994). More
recently, interpretations of Neanderthal scaven-
ging have been questioned on methodological
grounds (Marean & Kim, 1998; Mussi, 1999),
and several lines of evidence suggest that
Neanderthals were both highly dependent on
animal protein and quite adept at acquiring it
(Richards et al., 2000). The consumption side of
subsistence has also been given much needed
attention, and the organization of Neanderthal
food management is in some contexts strikingly
similar to that of Upper Palaeolithic people and
recent hunter-gatherers (Speth & Tchernov, 2001).
Despite considerable progress on all of these
fronts, we still have much to learn about the
range and nature of variability in Middle Palaeo-
lithic subsistence practices. Firstly, if we have
moved beyond a simple hunting/scavenging
dichotomy, we still know little about when/
where/why hominins used a particular tactic.
Figure 1. Location of Mujina Pec¤ina and sites mentioned in the text.
Copyright # 2005 John Wiley & Sons, Ltd.
85
Secondly, detailed studies of food transport,
processing, and the seasonality of procurement
are only starting to provide glimpses of how
Neanderthals managed food resources by provi-
sioning locales and positioning themselves in the
landscape. Finally, the geographic and temporal
coverage of well-studied Middle Palaeolithic
faunal assemblages is spotty at best. Recent
analyses from southern Europe (Boyle, 2000;
Stiner, 1994; Valensi, 2000) and the Caucasus
(Burke, 2000b; Enloe et al., 2000; Hoffecker &
Cleghorn, 2000) are starting to counterbalance
the Perigord bias. The Middle Palaeolithic faunal
assemblage from Mujina Pećina (Figure 1) is thus
of considerable interest because a) it comes from
an understudied part of Europe, b) it was ex-
cavated recently using modern techniques of
recovery and documentation, c) it is associated
with a well-dated, Late Mousterian, archaeologi-
cal context.
Int. J. Osteoarchaeol. 15: 84–105 (2005)
86
Table1. Faunal remains from Levels B,C, D1, D2, and ‘mixed’contexts at Mujina Pe¤cina
Total unid Burned
Level ID # Fragments fragments % ID
B 247 695 17 26.2
C 19 102 0 15.7
D1 91 110 1 45.3
D2 290 703 13 29.2
Mixed 55 44 2 55.6
Total 702 1654 33 29.8
Mujina Pećina faunal assemblage
Mujina Pećina is a small, east-southeast-facing
cave in a broken upland area (entrance at 280 m/
sl) overlooking the Dalmatian coast about 15 km
northwest of the modern city of Split (Karavanić
& Bilich-Kamenjarin, 1997). The cave is about
10 m deep and 8 m wide (ca. 55 m2 within the
dripline); recent excavations by the Department
of Archaeology, Zagreb University and Kaštela
City Museum exposed Middle Palaeolithic depos-
its over approximately 19 m2 between 1995–2000
(Rink et al., 2002). The total depth of deposit is
about 150 cm and is divided into eight major
stratigraphic units (from top to bottom, Levels
A, B, C, D1, D2, E1, E2, E3). Level A is a humus
formed in the surface of Level B. Preliminary
interpretations of the stratigraphy are of deposi-
tion of Levels B-C during warm periods, followed
by deposition of Levels D1-D2 during cold per-
iods, and finally sedimentation of Levels E1-E3
during relatively warm periods (Rink et al., 2002:
944–946). Two areas of localized burning in Level
D2 may represent simple Mousterian hearths. The
lithic assemblages contain small tools similar to
‘Micromousterian’ industries along with ‘typical’
Mousterian tools; byproducts of Levallois tech-
nology are present in Levels D1-D2 (Rink et al.,
2002). Five AMS (Accelerator Mass Spectrome-
try) radiocarbon dates on bone and charcoal from
Levels B-D2 indicate a very rapid deposition of
these sediments about 42 ka (rough calibration
of the mean of five dates, 39,222
2956 BP). Two
ESR (Electron Spin Resonance) dates on tooth
enamel from Level E1 have a mean of 44
5 ka
(LU [linear uptake]), while a single AMS 14C date
on bone from the interface between Levels E1 and
E2 is even older at 45,170
2780/2060 BP (Rink
et al., 2002).
Copyright # 2005 John Wiley & Sons, Ltd.
P. Miracle
% Burned Micro-mammal Bird Amphibian Bone N Bone Wt (g)
2.4 31 11 0 984 1495.4
0.0 0 1 0 122 129.1
0.9 19 9 0 229 1024.8
1.8 44 51 2 1090 2931.6
4.5 0 0 0 99 860
2.0 94 72 2 2524 6441
The Mujina faunal assemblage under consid-
eration is relatively modest, 2524 bones and
teeth1 with a total weight of 6.44 kg from five
major contexts2 (Table 1). Our analyses are
limited to the uppermost metre of the stratigra-
phy and Levels B-D2; remains from underlying
Levels E1-E3 are still under analysis and are not
included here. This sample includes all of the
remains excavated from Levels B, C, D1, D2
between 1995 and 2000. Levels B and D2 have
much larger assemblages than the other levels.
Animal remains were excavated by hand, with all
fragments larger than 2 cm plotted in three
dimensions (Karavanić & Bilich-Kamenjarin,
1997). All sediment was dry-sieved using a
3 mm mesh.
Bones were sorted into ‘identifiable’ and ‘uni-
dentifiable’ fragments by the author. Within the
former were included all fragments that could be
identified to body part and/or taxonomic cate-
gory, as well as all fragments marked with an
individual number (fragments larger than 2 cm
plotted in three dimensions). All fragments larger
than 5 cm were also included in ‘identifiable’. All
identifications were made using the recent com-
parative collections in the Grahame Clark
Zooarchaeology Laboratory at Cambridge Uni-
versity, standard references,3 and the author’s
large comparative data base on Pleistocene faunas
1
The following remains were extracted for absolute dating prior to
detailed analyses by Miracle: square F10, Level C, bone 61; square
G9, Level D1, two bones; square F9, Level D2, Mapa 5, two bones;
square G9, Level D2, two bones.
2
Level D2 includes excavation levels D2 and D2/E. Remains from
mixed or uncertain contexts are grouped under ‘mixed’, which
includes the following levels: S, D?, ‘profile’, ‘B/D profile’, and
‘kost iz niše’.
3
Equus hydruntinus was identified using criteria from Azzaroli (1979)
and Bonifay (1963), while criteria on the cheek teeth (e.g. swelling
of the crown immediately above the enamel-root junction) were
used to distinguish between Bos and Bison (Sala, 1986).
Int. J. Osteoarchaeol. 15: 84–105 (2005)
Late Mousterian Subsistence in Dalmatia
from the Balkans. Many of the ‘identifiable’ frag-
ments were long bone shafts that could at best be
identified to a body size category. Within ‘uni-
dentifiable’ were the remainder of the non-
diagnostic fragments from the sieve.
The bones were for the most part very well
preserved. In most cases, surfaces were very
clean. This was, to a large part, owing to calcite
rinds that had formed on many of the bones. A
number of the bones retained these rinds (NISP
[Number of Identified Specimens] ¼ 155, 22.1%
of identifiable assemblage), while many more
appear to have had rinds that have flaked off.
Calcite rinds were mechanically removed when-
ever possible. Despite these efforts, not all bone
surfaces were visible and frequencies of bone
modification should be treated as minimal esti-
mates. In a few cases, bones showed very heavy
weathering, and some lacked calcite rinds and
may have been more exposed owing to their
position within the cave. The appearance of
faunal remains did vary by level, indicating chan-
ging depositional environments. Material from
Level B was heavily brecciated with cemented
rinds of calcium carbonate and sediment (32% of
NISP retained calcite rinds); beneath these rinds
broken edges were quite rounded, suggesting
abrasion, perhaps from trampling. Several bones
from Level C were stained a mottled black colour,
and none of the specimens were covered by a
calcite rind. Bones so modified were not coded as
burned. Bone is for the most part highly com-
minuted and again gives the overall appearance
of having been rolled. Level D2 contained a
range of patinas. Most of the fragments were a
honey yellow colour, although some were white
to grey, and a few had a reddish brown mineral
stain. Many of the fragments were partially cov-
ered by calcium carbonate (14.8% of NISP
retained calcite rinds), although these rinds were
not as extensive and thick as those in Level B.
Several patterns emerge from these data on
fragments. Firstly, the %ID (percent identifiable)
is a rough measure of fragmentation since more
fragmented bones will tend to preserve fewer
portions/characteristics diagnostic of element
and/or taxon. The %ID is 29.8% for the assem-
blage overall. The very high %ID (55.6%) for the
‘mixed’ level reflects a collection bias; most of
these remains were hand picked without sieving
Copyright # 2005 John Wiley & Sons, Ltd.
87
from a large profile fall that occurred between the
1999 and 2000 field seasons. Among the remain-
ing levels, %ID is relatively high in Level D1 at
45.3% and relatively low in Level C at 15.7%.
Both of these levels have small assemblages
relative to Levels B and D2, and one cannot
rule out the possibility that these deviations
reflect anything more than sample bias. Consid-
ering Levels D2 and B, %ID decreases slightly
from the lower to the upper level (29.2% to
26.2%), suggesting that bone fragmentation was
slightly greater in B compared to D2. Similar
results are obtained if one examines the average
weight per fragment; it is highest in the ‘mixed’
levels at 8.7 g, followed by 4.5 g in Level D1,
2.7 g in Level D2, 1.5 g in Level B, and lowest in
Level C at 1.0 g (calculated from data in Table 1).
The fragmentation of the identifiable remains is
examined in greater detail later in this paper.
Secondly, burned bones are rare among the
unidentified fragments (Table 1). Only 2% of the
unidentified fragments are burned, and burning
frequency does not vary dramatically over time.
The relatively high and low values for Levels C,
D1, and ‘mixed’ could be products of the rela-
tively small samples sizes of these assemblages.
More surprisingly, the frequency of bone burning
in Level D2 is slightly lower than in Level B, even
though only the former level is reported to have
contained hearths. The burning features in Level
D2 appear to have played a negligible role in
food preparation (cooking) and refuse disposal
(cleaning), regardless of whether such bone burn-
ing would have been deliberate or accidental.
Finally, remains of micromammals (smaller than a
hedgehog), are relatively frequent in Level D1
(8.3% of assemblage) compared to Levels B and
D2 (3.2% and 4.0%, respectively), while the
frequency of bird remains decreases from 4.7%
in Level D2 to 1.1% in Level B. Micromammals
are absent from Level C and only a single bird
bone was recovered.4 The contrast in micrommal
frequency suggests that cave-roosting raptors,
owls, and small carnivores were most active at
Mujina during the accumulation of Level D1.
4
These values would certainly increase with systematic flotation and
fine-mesh wet sieving of sediment.
Int. J. Osteoarchaeol. 15: 84–105 (2005)
88 P. Miracle
Table 2. Species representation by level at Mujina Pec¤ina

Level B Level C Level D1 Level D2 Mixed levels Total

Species NISP MNE NISP MNE NISP MNE NISP MNE NISP MNE NISP % MNE % MNI %

Lepus sp. 3 3 2 2 17 10 22 7.9 15 10.0 2 9.1

Rupicapra rupicapra 7 4 6 4 14 7 1 1 28 10.1 16 10.7 5 22.7
Capra ibex 12 6 1 1 13 7 8 4 5 4 39 14.0 23 15.3 2 9.1
Capra/Rupicapra 32 8 2 4 3 13 8 5 3 56 20.1 24 16.0
Cervus elaphus 8 1 1 1 15 3 33 12 6 3 63 22.7 23 15.3 2 9.1
Sus scrofa 2 1 2 0.7 1 0.7 1 4.5
Bison priscus 4 1 4 1.4 1 0.7 1 4.5
Bos primigenius 1 1 1 0.4 1 0.7 1 4.5
Bos/Bison 8 3 3 3 8 7 19 6.8 13 8.7
Equus caballus 1 1 1 0.4 1 0.7 1 4.5
Equus hydruntinus 6 4 1 1 1 1 1 1 9 3.2 7 4.7 1 4.5
Equus sp. 4 1 1 1 1 6 2.2 2 1.3
Small-medium-sized 2 2 2 4 1.4 2 1.3
carnivore
Large-sized carnivore 2 2 0.7 0 0.0
Canis lupus 4 4 1 1 5 1.8 5 3.3 2 9.1
Ursus sp. 12 11 1 1 13 4.7 12 8.0 3 13.6
Meles meles 1 1 3 3 4 1.4 4 2.7 1 4.5
Total ID taxon 102 46 8 6 46 21 103 58 19 13 278 100.0 150 100.0 22 100.0
%Identifiable to taxon 41.3 82.1 42.1 66.7 50.5 75.9 35.5 63.7 34.5 86.7 39.6 62.9
Small-sized animal 1 4 2 5 2
Small-sized ungulate 63 7 4 2 7 1 49 16 10 133 36
Medium-sized ungulate 62 3 7 1 31 4 111 15 24 1 235 47
Large-sized ungulate 18 2 2 16 2 1 38 4
Unidentified 2 4 7 13 0
Grand Total 247 56 19 9 91 28 290 91 55 15 702 239 22

Assemblage composition grouped together in a pie chart of faunal fre-

Of the 704 bones and teeth from the identified
assemblage, only 39.8% of NISP was identifiable
to taxon (Table 2). As noted above, most of the
identified assemblage consisted of long bone
shafts identifiable only to body size categories.
Thus only 280 remains (11.1%) were identifiable
to taxon from the assemblage as a whole.
A total of 12 larger mammal species were
identified in the Mujina assemblage. Chamois,
ibex, and red deer are more or less equally
represented, and together make up almost two
thirds of the assemblage identified to taxon
(Table 2). They are followed by hare,5 large
bovids (bison and aurochs), bear, wild half ass,
wolf, badger, wild boar, and horse. Since many of
the caprid, bovid, and equid remains were not
identifiable to species, these remains have been
5
Only European hare has been positively identified from a lower I1
from Level D2 (MP H8D2.50: L ¼ 3.1 mm, B ¼ 2.4 mm). These
dimensions are in the range for Lepus europeaus and out of the range
for Lepus timidus (Koby, 1959).
Copyright # 2005 John Wiley & Sons, Ltd.
quency (Figure 2). Now it becomes clear that
chamois þ ibex dominate the assemblage with
about 43% of NISP, followed by red deer, hare,
large bovids, equids, bear, wolf, badger, and wild
boar (Figure 2).
The issue of Middle Palaeolithic faunal assem-
blage diversity has received considerable atten-
tion over the years because it has been suggested
that targeted procurement of a single species is
indicative of specialized hunting and was not
practised by hominins before the Upper Palaeo-
lithic (Binford, 1968; Mellars, 1973, 1996). Other
authors have argued on comparative (Chase,
1989) and methodological (Grayson & Delpech,
2002) grounds that assemblage diversity does not
change significantly from the Middle Palaeolithic
to Upper Palaeolithic. Evenness (E) is one mea-
sure of diversity; evenness can vary from 0 to 1,
the higher the value, the more even the assem-
blage. We calculated E only on the ungulate
assemblage since these taxa are more likely to
have been the prey of hominins. We used an
Int. J. Osteoarchaeol. 15: 84–105 (2005)
Late Mousterian Subsistence in Dalmatia
Figure 2. Species abundance by %NISP (N ¼ 280) at Mujina Pec¤ina.
approach similar to that described in Grayson
and Delpech (2002); our values for E are directly
comparable to those calculated by Grayson and
Delpech (2002) on Middle Palaeolithic and Early
Upper Palaeolaithic faunal assemblages in south-
western France.6 Considering the assemblage as a
whole, E is fairly high at 0.66 (NISP ¼ 228).
Evenness increases slightly from 0.636 in Layer
D1 þ D2 (NISP ¼ 123) to 0.667 in Layer B þ C
(NISP ¼ 86). These values for E from Mujina
Pećina fall very close to the regression line
calculated by regressing evenness on sample
size for Middle Palaeolithic sites (Grayson &
Delpech, 2002: Figure 2); this slight change in
E may be an artefact of sample size. Mujina
Pećina appears to have a diverse faunal assem-
blage with a relatively even representation of the
major species. Nothing about the assemblage
composition indicates that hominins were target-
ing particular prey to the exclusion of other
species, and hence these data provide no evi-
dence of specialized procurement.
6
Evenness was measured using the formula  pi ln pi/ln S, where p
is the proportion of specimens (NISP) in the ith taxon and S is the
number of non-overlapping taxa. Large bovids (Bos primigenius, Bison
priscus, Bos/Bison) have been treated as a single taxon. Equids (Equus
hydruntinus, Equus caballus, Equus sp.) are also treated as a single taxon.
Specimens identified as Capra/Rupicapra have been allocated to Capra
ibex and Rupicapra rupicapra in proportion to the relative representa-
tion of each species.
Copyright # 2005 John Wiley & Sons, Ltd.
89
The taxonomic composition based on the
minimum number of elements (MNE) and mini-
mum number of individuals (MNI) is broadly
similar to that based on NISP. The MNE was
calculated on the most frequent anatomical fea-
ture preserved from an element for each taxo-
nomic category (often body size class). All
identifiable remains, including those identifiable
only to element, were considered in estimating
the MNE. For example, all long bone shaft
fragments identifiable to element were examined
in the estimation of MNE (in most cases the
MNE is based on the presence of the nutrient
foramen). The MNE of long bones is estimated
relative to the entire bone rather than for each
articular end. The MNE of upper and lower teeth
is calculated relative to complete dentitions, tak-
ing age into consideration. The systematic refit-
ting of the assemblage was not attempted, in part
owing to the high incidence of rounding of
fragment edges. The MNI was calculated for
the aggregate assemblage of all levels combined,
taking side and relative age into consideration.
The relative frequency of less common taxa
increases when quantified by MNE and MNI,
while the relative frequency of more common
taxa decreases (Grayson, 1984). Sample sizes are
too low, particularly for MNI, for anything more
than the most rudimentary discussion of rank
frequency. What emerges from the data on
MNI is that a small number of animals is
Int. J. Osteoarchaeol. 15: 84–105 (2005)
90 P. Miracle
Table 3. Taxonomic frequency by layer at Mujina Pec¤ina

BþC BþC D1 þ D2 D1 þ D2 Total Total

Taxon (%NISP) (%MNE) (%NISP) (%MNE) (%NISP) (%MNE)

Lepus sp. 2.7 5.8 12.8 15.2 7.9 10.0

Capra/Rupicapra 49.1 36.5 38.9 41.8 44.2 42.0
Cervus elaphus 8.2 3.8 32.2 19.0 22.7 15.3
Sus scrofa 1.8 1.9 0.0 0.0 0.7 0.7
Bos/Bison 10.0 11.5 8.7 11.4 8.6 10.0
Equus sp. 9.1 9.6 2.7 3.8 5.8 6.7
Small-sized carnivore 2.7 1.9 3.4 6.3 2.9 4.0
Large-sized carnivore 16.4 28.8 1.3 2.5 7.2 11.3
Total N 110 52 149 79 278 150

represented; only chamois and bear are repre-
sented by more than two individuals. Even if
Levels D to B at Mujina accumulated over a short
period of time, visitations to the cave by homi-
nins or others that produced faunal remains must
have been rare and sporadic.
A few observations about stratigraphic
changes in assemblage composition can be
made from these data, although results are not
firm owing to small sample sizes from individual
levels (e.g. Level C is excluded from this discus-
sion). Remains specifically identified to bison are
restricted to Level D1, while those from aurochs
are restricted to Level D2. Since larger bovid
remains from either bison or aurochs are present
in all levels, it is impossible to assign any strati-
graphic significance to this pattern. Likewise,
equid remains that could come from horse or
Equus hydruntinus are found in all levels. In fact, the
only definite contrast in species composition
comes from wild boar, which is found only in
Level B, and the lack of any carnivores from Level
D1. The latter pattern, however, may reflect the
relatively small sample size from Level D1 (NISP
ID to species ¼ 46).
Thus, owing to small sample sizes from indi-
vidual levels, we have grouped together adjacent
levels that show relatively similar sedimentary
characteristics. Levels B and C are treated
together (Layer B þ C), while Levels D1 and
D2 are treated together (Layer D1 þ D2). As
discussed above, we also group together the
equid (horse, Equus sp. and Equus hydruntinus),
large bovid (Bison, Bos/Bison and Bos), small bovid
(Rupicapra, Rupicapra/Capra and Capra), large car-
nivore (Canis, Ursus, large-sized carnivore), and
Copyright # 2005 John Wiley & Sons, Ltd.
small carnivore (Meles, small-medium-sized carni-
vore) remains (Table 3).
Several interesting changes emerge from these
grouped data. The relative frequency of hare and
red deer decrease significantly from Layer
D1 þ D2 to Layer B þ C, while the relative
frequency of chamois/ibex, equids, and large-
sized carnivores increase dramatically from
Layer D1 þ D2 to Layer B þ C (Table 3,
Figure 3). As noted above, wild boar is present
only in Layer B þ C. The relative frequencies of
Bos/Bison and small-sized carnivores do not
change over time.
Large mammals are notoriously coarse indica-
tors of palaeoecological conditions. Nonetheless,
it has been commonly assumed that red deer and
wild boar are indicative of more temperate and
forested conditions, chamois and ibex of cool,
alpine conditions, and equids of relatively open
biotopes. With the exception of the presence of
wild boar in Layer B þ C, the remaining taxa
indicate more temperate conditions (perhaps
with more shrub/forest cover) in Layer D1 þ D2
changing to cooler (increase in chamois/ibex),
more open (increase in equids) conditions in
Layer B þ C. The confirmed presence of Eur-
opean as opposed to varying hare in Layer
D1 þ D2 supports this interpretation of Layer
D1 þ D2 (Figure 3). Such an interpretation is
contrary to preliminary sedimentological ana-
lyses of cool/dry conditions during the formation
of Layer D1 þ D2 and relatively temperate con-
ditions during the formation of Layer B þ C (Rink
et al., 2002). In this case, we consider the sedi-
ments to be more reliable indicators of local
palaeoclimates. Chamois and ibex are not reliable
Int. J. Osteoarchaeol. 15: 84–105 (2005)
Late Mousterian Subsistence in Dalmatia
Figure 3. Change in faunal composition at Mujina Pec¤ina. Data fromTable 3.
indicators of palaeotemperature (Miracle &
Sturdy, 1991; N. E. H. Phoca-Cosmetatou,
unpublished PhD dissertation, 2001); red deer
thrive in a variety of biotopes, and Equus hydrunti-
nus appears to have favoured relatively temperate
yet open environments. Instead of telling us a
story about changing climates, the large mammal
faunas at Mujina Pećina indicate changes in prey
selection by Neanderthals and/or other bone
collectors.
The second major contrast in species compo-
sition comes in the increase in large carnivores in
Layer B þ C. This increase is almost entirely
accounted for by bear (the few remains identifi-
able to species appear to be cave bear, Ursus
spelaeus). These potentially denning carnivores
increase dramatically in frequency from Layer
D1 þ D2 to Layer B þ C. The frequency of large
carnivores is high enough relative to ungulates in
Layer B þ C to suggest that the former may have
played a significant role in the formation of the
assemblage. Large carnivores may have more
regularly used Mujina Pećina in Layer B þ C
relative to Layer D1 þ D2; the taphonomy of
assemblage formation may have changed signifi-
cantly over time. Thus, even though the Mujina
sediments appear to have accumulated relatively
rapidly (Rink et al., 2002), the fauna indicates
Copyright # 2005 John Wiley & Sons, Ltd.
91
significant contrasts between different deposi-
tional episodes.
Taphonomy
As noted above, many of the animal bones and
teeth appear to have been abraded; broken edges
are commonly rounded. All bone surfaces were
scanned using a 10x magnifying glass; most speci-
mens with suspected cut marks and other unusual
modifications were further examined with a low-
power binocular microscope (7.5–50x magnifica-
tion). Weathering of bone surfaces, through
abrasion, fine-line fractures, spalling of bone
surfaces, etc. is by far the most common type of
modification; over 25% of identified remains are
so modified by ‘light’ or ‘heavy’ weathering
(Table 4).7 Weathering frequencies for remains
identified only to body size are higher than those
for specimens identified to genus/species; this
pattern is not surprising since weathering will
obliterate diagnostic features and move remains
7
‘Light’ weathering is equivalent to Behrensmeyer’s stages 1–2,
while ‘heavy’ weathering is equivalent to stages 3–5 (Behrensmeyer,
1978). Behrensmeyer’s stages are used here simply as a heuristic
device; the causes of ‘weathering’ in a cave environment such as
Mujina Pećina would have been very different from those identified
by Behrensmeyer in open-air contexts in East Africa.
Int. J. Osteoarchaeol. 15: 84–105 (2005)
92 P. Miracle
Table 4. Bone modification by taxon at Mujina Pec¤ina
%NISP

Tooth marked Digested Cut Impact scars Burned Light Heavy Total
Taxon weathering weathering

Rupicapra/Capra 5.7 4.1 0.8 1.6 0.8 11.4 2.4 123

Cervus elaphus 1.6 9.5 1.6 6.3 1.6 12.7 14.3 63
Sus scrofa 0.0 0.0 0.0 0.0 0.0 0.0 0.0 2
Bos/Bison 4.2 8.3 4.2 4.2 0.0 0.0 4.2 24
Equus sp. 6.3 18.8 0.0 0.0 0.0 0.0 0.0 16
Small-medium-sized carnivore 0.0 0.0 0.0 0.0 0.0 25.0 0.0 4
Large-sized carnivore 0.0 0.0 0.0 0.0 0.0 0.0 0.0 2
Canis lupus 0.0 0.0 0.0 0.0 0.0 0.0 0.0 5
Ursus sp. 15.4 0.0 0.0 0.0 0.0 30.8 0.0 13
Meles meles 0.0 0.0 0.0 0.0 0.0 0.0 0.0 4
Lepus sp. 4.5 4.5 0.0 0.0 0.0 4.5 0.0 22
Small-sized animal 0.0 0.0 0.0 0.0 0.0 20.0 0.0 5
Small-sized ungulate 10.5 6.8 5.3 4.5 0.8 29.3 4.5 133
Medium-sized ungulate 3.8 3.0 3.4 4.3 0.9 24.7 9.4 235
large-sized ungulate 0.0 0.0 7.9 2.6 5.3 28.9 13.2 38
Unidentified 0.0 30.8 0.0 0.0 15.4 7.7 15.4 13
Grand Total 5.1 5.3 3.0 3.4 1.3 19.7 6.8 702

Table 5. Bone modification by layer at Mujina Pec¤ina

%NISP

Tooth marked Digested Cut Impact scars Burned Light Heavy Total
Layer weathering weathering

BþC 5.3 3.0 1.9 1.5 0.8 24.8 6.4 266

D1 þ D2 5.5 7.3 3.7 4.5 1.8 15.5 7.1 381
Mixed 1.8 1.8 3.6 5.5 0.0 23.6 7.3 55
Total 5.1 5.3 3.0 3.4 1.3 19.7 6.8 702

from relatively more to less identifiable cate-
gories. There is a second pattern, however, of
weathering frequency increasing with body size.
Smaller-sized remains would have been incorpo-
rated in the sediment more rapidly than larger-
sized remains, while the latter were more likely to
be displaced, trampled, or otherwise abraded.
Comparing different levels (Table 5), weathering
frequency appears to increase somewhat from
Layer D1 þ D2 to Layer B þ C.
The weathering data corroborate previously
mentioned evidence of more bone fragmentation
in Layer B þ C relative to Layer D1 þ D2. The
%ID and average weight per fragment, however,
are very rough measures of fragmentation. A
more direct measure of bone fragmentation is
fragment length. This was measured to the near-
est millimetre on all identifiable specimens for
which the greatest linear dimension was not
affected by recent breaks. Long bone shafts not
identifiable to element and smaller than 5 cm
were not included with the identifiable speci-
mens, which partially truncates the lower end of
the size range. Thus these data are not directly
comparable to length measures from other faunal
assemblages sorted using criteria different from
those outlined above. They are, however, extre-
mely useful for intra-assemblage comparisons. To
control for the original size of bones (big bones
tend to produce larger fragments than small
bones), comparisons are made using only the
diaphyseal fragments of major long bones8 and
different body sizes are treated separately. For
reasons of small sample sizes, the major long
bones are treated as a single group. Results are
8
We include under ‘major long bone’ the humerus, radius, metacar-
pal, femur, tibia, and metatarsal, as well as indeterminate long bone
shafts that must come from one of the aforementioned elements.

Copyright # 2005 John Wiley & Sons, Ltd. Int. J. Osteoarchaeol. 15: 84–105 (2005)
Late Mousterian Subsistence in Dalmatia
Table 6. Comparison of long bone shaft lengths (in mm) between Mujina Pec¤ina Layers B þ C and D1 þD2. Only diaphyses of
major long bones (humerus, radius, metacarpal, femur, tibia, metatarsal) not shortened by recent breaks were measured. ‘Small
ungulate’ includes remains identified to small ungulate and chamois/ibex.‘Medium ungulate’ includes remains identified to medium
ungulate and red deer.The 95% confidence intervals were calculated using t0.05, sample SD and sample size
N min
Small ungulate
Layer B þ C 41 23
Layer D1 þ D2 44 26
Medium ungulate
Layer B þ C 37 23
Layer D1 þ D2 114 27
Large ungulate
Layer B þ C 9 22
Layer D1 þ D2 8 36
tabulated in Table 6. Long bone shafts are much
larger in Layer D1 þ D2 than in Layer B þ C;
average length decreases over time by 13.6 mm in
small ungulates (t-test: t ¼ 3.26, d.f. ¼ 67, p ¼
0.002) and by 22.4 mm in medium ungulates (t-
test: t ¼ 7.82, d.f. ¼ 133, p < 0.001). The size
decrease in large ungulate shaft size is not statis-
tically significant. Unexpectedly, long bone
shafts of small ungulates are slightly larger than
those of medium ungulates in Layer B þ C,
although this difference is not statistically sig-
nificant. The fragmentation of animal bones
increased significantly over time at Mujina
Pećina. The cave was clearly a dynamic sedimen-
tary environment during the Pleistocene.
Turning to other kinds of bone modification,
tooth marks (including punctures, tooth scores,
crenulated edges, and gouged cancellous bone)
and evidence of etching from gastric acids (often
with thin and/or feathered, broken edges and rare
punctures) are present on over 10% of the assem-
Figure 4. Bos/Bison lower deciduous third premolar, digested
(Level D2, square G9).
Copyright # 2005 John Wiley & Sons, Ltd.
93
Length in mm
max average (95% CI) SD
88 43.9 (39.7–48.0) 13.1
151 57.5 (40.1–64.8) 24.1
79 41.7 (38.1–45.4) 11.0
164 64.1 (59.6–68.6) 23.6
106 50.2 (31.0–69.5) 25.5
101 59.6 (41.9–77.4) 21.8
blage (Figure 4). All of the main species show
evidence of tooth marks and/or digestion. Sample
sizes are too small for a detailed analysis, but
there is an interesting pattern of more digestion
relative to gnawing in larger-sized animals (e.g.
equids, large bovids, red deer) and the reverse
pattern in smaller-sized animals (chamois/ibex,
hare, small ungulate sized). The frequency of
carnivore modification decreases from 12.8% to
8.3% of NISP from Layer D1 þ D2 to Layer
B þ C, even though the frequency of carnivore
remains shows the opposite trend. As discussed
above, the increase in large carnivore frequency
in Layer B þ C comes primarily from bear, and as
discussed below these remains come mostly from
neonate/juvenile individuals. That is, bear sows
may have been using the site occasionally as a
nursery during the accumulation of Layer B þ C.
The concurrent reduction in hominin activity
would have also reduced the amount of scaveng-
able food waste created.
Only 1.3% of NISP were burned, and only
3.0% of NISP bear cut marks (Tables 4 and 5,
Figure 5). A specimen was coded as bearing an
‘impact scar’ when there was a negative flake scar
emanating from loading on the cortical surface of
a fractured edge. Our usage generally corre-
sponds to Capaldo and Blumenschine’s (1994:
744) definition of a ‘normal notch.’ Impact scars
at Mujina were relatively wide with acute
release angles. Visually they resemble percussion
notches described by Capaldo and Blumenschine
(1994). We did not, however, measure the size
and shape of these notches. In the absence of
more detailed data about notch morphology and
Int. J. Osteoarchaeol. 15: 84–105 (2005)
94
Figure 5. Large bovid femur, distal shaft, with cut marks, calcium
carbonate rinds, and impact scars (Level D2: F8D2.22).
size, we cannot unambiguously attribute all of
them to hominin activity. Nonetheless, the
majority were probably produced by hominins
cracking open long bones to extract marrow.
Impact scars are present on 3.4% of NISP. The
statistic on burning is very similar to that of the
unidentified fragments (Table 1). Of the six
taxonomic groups with NISP > 10 (chamois/
ibex, red deer, large bovid, equid, bear, hare),
only remains of red deer, and chamois/ibex were
burned (Table 4). As with weathering, there
seems to be a strong pattern with body size—
burning frequency increases with larger body
size; this pattern is also visible in the remains
identified only to body size—weathering fre-
quency increases from small ungulate to large
ungulate-sized remains. Cut marks are found on
chamois/ibex, red deer, and large bovid remains,
along with bone shafts identified to different
body sizes. The latter marks are all on long
bone diaphyses; these mid-shaft cut marks are
most likely from defleshing/meat consumption.
Remains from these taxa also bear impact scars.
Taken together, the burning and cut mark data
provide clear evidence of human modification for
only chamois/ibex, red deer and large bovids.
Copyright # 2005 John Wiley & Sons, Ltd.
P. Miracle
Significantly, none of the carnivore remains
appear to have been modified by hominins.
More interestingly, the same appears also to be
true of equids and hare. The only conclusive
evidence of modification of equid and hare
remains comes from carnivores. Viewed tempo-
rally, frequencies of burning and cut marks
decrease significantly (approximately by half)
from Layer D1 þ D2 to Layer B þ C.
How can we further untangle the actions of
hominins and other carnivores at Mujina Pećina?
One approach is to focus on the mid-shaft por-
tions of long bones. A series of actualistic experi-
ments show that mid-shafts of long bones
frequently bear tooth marks when carnivores
have the first access to meat and/or marrow in
limbs, while these same portions are generally
ignored by carnivores when humans have already
processed long bone mid-shafts for marrow (e.g.
Blumenschine, 1988; Marean & Spencer, 1991;
Marean & Bertino, 1994; Capaldo, 1998; Sel-
vaggio, 1998). Hominins, on the other hand,
frequently leave damage marks from defleshing
limbs (cut marks) or breaking open long bones
for marrow (percussion marks, including impact
scars). In situations where only hominins or
carnivores acted on an assemblage, the expecta-
tions are fairly straightforward. In the former
case, there should be frequent percussion and
cut marks and no tooth marks or digested bones.
In the latter case, there should be frequent tooth
marks and digested bones, but no cut marks or
percussion marks. What about assemblages, how-
ever, where tooth, percussion, and/or cut marks
are all present? This issue was initially investi-
gated by examining bones marked by teeth and
cuts; did tooth marks overlap cut marks or vice
versa (Shipman,1986)? In recent years, the most
significant advance of the actualistic experiments
mentioned above has been to generate quantita-
tive predictions about the relative frequency of
percussion, tooth, and cut marks when both
hominins and carnivores have had access to
remains. Three basic scenarios have been mod-
elled and examined with actualistic data
(Capaldo, 1998; Selvaggio, 1998; Marean et al.,
2000): hominids followed by carnivores (carni-
vores ravage hominid food waste), carnivores
followed by hominids (hominids scavenge
carniovre kills), and carnivores followed by
Int. J. Osteoarchaeol. 15: 84–105 (2005)
Late Mousterian Subsistence in Dalmatia 95
Table 7. The frequency of long bone mid-shafts with tooth and With the exception of the small ungulates from
cut marks from Mujina Pec¤ina Layers B þ C and D1 þD2 com-
pared to experimental data, with 95% confidence intervals Layer D1 þ D2 (32.4% tooth marked), the fre-
listed in parentheses. Fragments with calcite rinds are not in- quency of tooth marks on long bone mid-shafts
cluded. Confidencepffiintervals for percentages calculated using (0.0–10.0%) is lower than the experimentally
the formula 1.96* ( p)*(100-p)/N where p is the percentage
and N is the sample size (Wonnacott & Wonnacott,1990: 273) scavenged assemblages (11.2% is the lowest
95% confidence interval), while within some of
Tooth marked Cut marked the 95% confidence intervals for carnivore-
N % (95% CI) % (95% CI)
ravaged assemblages (Table 7). Cut-mark fre-
Layer B þ C quencies, on the other hand, give equivocal
Small ungulate 30 10.0 (0.0–20.7) 6.7 (0.0–15.6) results. Cut mark frequencies on large ungulates
Medium ungulate 27 7.4 (0.0–17.3) 3.7 (0.0–10.8) in Layer D1 þ D2 (33.3%) and on small ungu-
Large ungulate 6 0.0 0.0
Layer D1 þ D2 lates in Layer B þ C (6.7%) are above those for
Small ungulate 34 32.4 (16.6–48.1) 5.9 (0.0–13.8) scavenged assemblages (6.0% is the highest 95%
Medium ungulate 90 5.6 (0.8–10.3) 5.6 (0.8–10.3) confidence interval). The remaining assemblages
Large ungulate 9 0.0 33.3 (2.5–64.1)
Experimental from Mujina fit in one or both of the experimen-
Hominid to carnivore tally-scavenged assemblages and show cut-mark
Blumenschine 10.5 (0.0–21.5) frequencies well below that reported for carni-
Capaldo2 15.4 (12.3–18.5) 13.5 (10.6–16.4)
Marean1 7.1 (4.1–9.8) vore-ravaged assemblages (10.6% is the lowest
Carnivore to hominid3 22.0 (13.0–31.0) 2.4 (0.0–5.2) 95% confidence interval). In summary, the fre-
Carnivore to hominid 23.2 (11.2–35.2) 3.2 (1.0–6.0)
to carnivore3
quency of cut marks fall within the expected
ranges of scavenged assemblages (hominids sec-
1
Data as reported by Marean et al. (2000: Table 3). ond), while the frequency of tooth marks falls
2
Data from Capaldo (1998: Table 13A), 95% confidence intervals within the expected ranges of ravaged assem-
calculated using t0.05, sample SD and sample size.
3
Data from Selvaggio (1998: Table 6).
blages (hominids first).
This kind of direct comparison between
archaeological and experimental/actualistic as-
hominids followed by carnivores (carnivores semblages is potentially problematic for several
ravage remains from hominid-scavenged food reasons. Firstly, the Mujina Pećina assemblages
waste). In all three cases, one expects there to have been significantly affected by taphonomic
be percussion, tooth, and cut marks. processes other than carnivore gnawing. For
The frequency of long bone mid-shafts with example, surface modifications at Mujina may
tooth marks and cut marks from Mujina Pećina is have been obliterated by weathering; about
presented in Table 7.9 We compare frequencies 25% of identifiable remains from Mujina show
among small-, medium-, and large-sized ungu- signs of weathering (Table 5). Assuming that
lates from Layer B þ C and Layer D1 þ D2. Long remains with tooth marks and cut marks were
bone mid-shafts are more frequently modified in not preferentially weathered, one can compen-
Layer D1 þ D2 than in Layer B þ C, regardless of sate for the effects of weathering by simply
body size and agent of modification. We have dividing all frequencies by 0.75. This transforma-
excluded all specimens with calcite rinds. Results tion increases slightly the frequency of modifica-
from the long bone mid-shafts confirm those tions, but has little effect on where different
obtained for the assemblage as a whole (Tables assemblages fall with respect to the carnivore-
4 and 5). To aid further interpretation, we com- ravaging and hominin-scavenging scenarios
pare our results to experimental studies that described above. Something more than bone
modelled the three basic scenarios discussed weathering would appear to be at work. Sec-
above. Tooth-mark data give interesting results. ondly, our comparison makes strong analogies
not only between agents (i.e. carnivores and
9
As noted earlier, our criteria for identifying and coding impact scars hominins) and their traces (tooth or cut mark),
cover only a subset of the marks coded as percussion marks but also between contexts of consumption/bone
Blumenshine et al. (1996). Hence we do not attempt to compare
impact scar frequencies to actualistic data on frequency of percus- modification and the frequency with which such
sion marks. traces are produced. The first analogy between
Copyright # 2005 John Wiley & Sons, Ltd. Int. J. Osteoarchaeol. 15: 84–105 (2005)
96
agents and effects is relatively unproblematic,
and most researchers are confident about distin-
guishing between different taphonomic agents
based on bone surface modifications. The second
analogy is more problematic. Are butchery meth-
ods today, whether those of professionals wield-
ing metal implements or amateurs (e.g. Euro-
American academics) wielding stone tools appro-
priate analogues for Middle Palaeolithic homi-
nins (in a European context, presumably
Neanderthals)? Past hominins clearly used stone
tools in butchery. They may, however, have
relied more heavily on their physical strength
to disarticulate carcasses after initial cuts and/or
break open bones after the surface had been
cracked with a hammerstone. Our point here is
not to propose a specific model of Middle
Palaeolithic butchery, but to raise the issue that
the relationship between butchery practices and
trace frequency is mediated by many variables,
only some of which are potentially under experi-
mental control. That said, the experimental and
actualistic approaches taken by Blumenschine,
Marean, Capaldo, Selvaggio, and others are
clearly the way forward. Finally, the experiments
were based on feeding captive hyenas butchered
carcass parts (Marean & Spencer, 1991; Marean &
Bertino, 1994), while actualistic research was
conducted on the effects of hyenas and other
scavengers on butchered remains laid out in East
African savannah environments (Blumenshine,
1988; Capaldo, 1998; Selvaggio, 1998). Mujina
Pećina differs from these experimental and actua-
listic situations in the following (among other)
ways: it lacks direct evidence of the presence of
hyenas, the ecology of Pleistocene Dalmatia is
quite different from modern-day East Africa, and
the cave is spatially constrained relative to the
open-air contexts of the actualistic studies.
Although remains of hyenas are not present at
Mujina Pećina, several remains appear to have
been modified by hyenas (discussed below), and
their remains and/or coprolites are present at
other Pleistocene caves in the region such as
Pećina u Brini (Malez, 1975a), Crvena Stijena
(Malez, 1975b), and Parska Golobina (Rakovec,
1961) (for an overview see Miracle, 1991). We
cannot control for differences in the ecology of
Pleistocene Dalmatia and East Africa today, and
the effects of any differences remain unknown.
Copyright # 2005 John Wiley & Sons, Ltd.
P. Miracle
As for contrasts in the contexts of consumption
(e.g. cave vs. open air), this is precisely one of the
variables one would hope to better understand
through the comparisons made above. Marean
et al. (2000) productively used these experimen-
tal/actualistic data to interpret the taphonomy of
Die Kelders 1, like Mujina Pećina, a Pleistocene
cave from a Mediterranean-type environment
with evidence of both hominin and carnivore
activities. We conclude that our comparisons
between the Mujina Pećina faunal assemblages
and these experimental/actualistic assemblages
are sound as long as the potential points of
divergence are kept in mind.
Another way to use the experimental data is to
compare the frequency of hominid modifications
relative to carnivore modifications among the
different scenarios. In all of the experimentally
ravaged assemblages, the frequency of cut marks
was only slightly less frequent than tooth marks.
In the scavenged assemblages, in contrast, the
frequency of tooth marks was greater than the
frequency of cut marks by at least an order of
magnitude.
Using these criteria, we suggest that the large
and medium ungulate assemblages from Layer
D1 þ D2 were accumulated by hominins and
then ravaged by carnivores. The small ungulate
assemblage from Layer D1 þ D2 better fits a
scavenging model. In Layer B þ C, we lack data
to interpret the large ungulate assemblage, while
the small and medium ungulates were probably
accumulated by hominins and then ravaged by
carnivores. Hominins would appear to have been
hunting large- and medium-ungulates, while
small ungulates were scavenged, particularly in
Layer D1 þ D2, as opportunities arose. Taking
into account the 95% confidence intervals for
percentages from the Mujina assemblages
(Table 7), then it becomes apparent that the
small ungulates from Layer D1 þ D2 could fit
both a carnivore-ravaged and hominid-scavenged
model. Thus the long bone mid-shafts indicate
that there is relatively little firm evidence for
hominids scavenging from carnivores, while
there is abundant evidence for carnivores scaven-
ging from hominids (ravaging assemblages).
Two specimens warrant further discussion.
The first is a shaft fragment from Level D2
(E7D2.5) with three small holes that pierce the
Int. J. Osteoarchaeol. 15: 84–105 (2005)
Late Mousterian Subsistence in Dalmatia
Figure 6. Long bone shaft fragment with three holes (Level D2:
E7D2.5).The specimen was probably partially digested and regur-
gitated by a hyena.
cortical bone (Figure 6). The largest hole
(ca. 3.5 mm diameter) is broken by an old break
of the shaft, while the other two holes (ca. 2 mm
and 1.5 mm diameter, respectively) are complete.
All three holes are circular in shape. There is
relatively little crushing of the bone surface
around the edge of the holes. The specimen
resembles bone fragments that have been par-
tially digested and regurgitated by spotted hye-
nas (D’Errico & Villa, 1997: Figure 13). It is even
more similar to bone fragments from the Pleisto-
cene hyena den of Bois Roche (D’Errico & Villa,
1997: Figure 16). The largest hole may represent
a nutrient foramen that has been enlarged by a
hyena’s gastric acid.
The second specimen is a long bone shaft
(F7D.13) that appears to have been chipped
along several of its broken edges (Figure 7).
These flake scars start from the cortical surface
and extend in towards the medullary cavity. We
doubt that this edge damage might be the result
of abrasion since other surfaces appear fresh. On
the other hand, these flake scars are too small to
be from deliberate retouch. Apparently hyena-
modified bones from the hyena den of Bois Roche
again provide a close analogue (Villa & Bartram,
1996). Together, these two specimens provide
strong indirect evidence of the hyena scaven-
ging/ravaging of the Mujina faunal assemblage.
Based on this evidence of bone modification,
we suggest the following interpretation of the
taphonomy of Layers B-D. During the accumula-
tion of Layer D1 þ D2, which based on sedimen-
tological evidence was a time of relatively cold
climate, hominins hunted red deer and large
bovids and perhaps scavenged chamois and
Copyright # 2005 John Wiley & Sons, Ltd.
97
Figure 7. Long bone shaft fragment with chipping along its edge
(Level D: F7D.13). The surface modification was probably caused
by hyena chewing.
ibex. Hyenas and wolves were also active at the
site; they visited it, probably following on homi-
nin occupations, to scavenge bones and other
food waste. These carnivores may have also at
times hunted hare. Mujina was also used occa-
sionally as a latrine by scavenging carnivores, but
there is no evidence from juvenile teeth and
bones of the site being used as a nursery den or
lair. The climate ameliorated somewhat in Layer
B þ C, and the appearance of wild boar indicates
denser thickets and/or moisture near to Mujina
Pećina. Hominins continued to visit the site, but
with less frequency than before. Despite the
climatic amelioration, they focused more on
mountain caprids than during earlier occupations.
Bear were using the cave, perhaps as a hiberna-
tion den by bear sows with dependent young
and/or neonates, at times when hominins were
less active in the area. Equid remains were
brought to the site, probably by wolves and
perhaps hyenas. Bone fragmentation and weath-
ering increase too, perhaps from the periodic
Int. J. Osteoarchaeol. 15: 84–105 (2005)
98 P. Miracle

trampling of the site by bears as well as bone
consumption by hyenas.
Element representation
Different elements of the skeleton are grouped
into carcass units owing to small sample sizes
(Table 8). The elements included in each carcass
unit are indicated on the table, and the MNE is
standardized by dividing MNE by the frequency
with which elements are found in a complete
skeleton. Data are provided for chamois þ ibex
(combined), red deer, Bos þ Bison, equids, hare,
bear, small ungulate-sized animals, and medium
ungulate-sized animals. Data are provided for the
Mujina Pećina assemblage in its entirety, again
owing to small sample sizes.
All of the major taxa are represented by both
teeth and bones. There is significant variation in
the relative frequency of teeth to bones; cha-
mois þ ibex, large bovid, and equid assemblages
are dominated by teeth (%NISP ¼ 50–66.7%), in
contrast to red deer and bear assemblages with
relatively few teeth (%NISP ¼ 7.7–17.5%). The
taphonomic significance of these differences is
not clear as red deer and bear show relatively
high frequencies of weathering and tooth/diges-
tion marks (Table 4); the high frequency of bones
relative to teeth is not simply a reflection of less
bone destruction.
We compare carcass unit representation, stan-
dardized by the relative frequency of elements
per carcass unit, for chamois þ ibex þ small ungu-
late and red deer þ medium ungulate in Figure 8.
Inclusion of remains identifiable only to body
size, brings into the analysis parts of the axial
skeleton not easily identifiable to species (e.g.
small vertebral and rib fragments) as well as long
bone shafts that could be identified to element on
the basis of nutrient foramina and other anato-
mical landmarks. Our approach to identification
thus addresses some of the biases created by the
preferential destruction of long bone ends rela-
tive to mid-shafts (Marean & Frey, 1997), and in
most cases limb bone MNE was calculated on
shaft fragments instead of articular ends. The
pattern of carcass representation is strikingly
similar between these different-sized ungulates.
Upper and lower teeth are present in similar
frequencies, suggesting that complete heads
were introduced to the site. Likewise, antler/
horn and bony parts of the head are present in
similar frequencies. Considering the axial skele-
ton in general, frequencies decrease dramatically
from the head to the tail, with a particularly large
decrease from the head to the neck (Figure 8).
Within the limbs, carcass unit frequencies

Table 8. Carcass unit frequency of major taxa at Mujina Pec¤ina

Chamois þ Ibex Red deer Large bovid Equid Hare Bear Small ungulate Medium ungulate

Carcass unit NISP MNE NISP MNE NISP MNE NISP MNE NISP MNE NISP MNE NISP MNE NISP MNE

Antler/horn 8 5 15 3
Head 12 8 3 3 1 1 2 2 1 7 2 8 2
Upper teeth 25 12 6 2 7 3 3 2
Lower teeth 30 13 3 3 8 7 2 2 8 3
Teeth 15 1 2 1 3 1 1 1 1 3 3 1 1
Neck 1 1 13 7 26 4
Pelvis 1 1
Upper front 8 7 1 1 1 1 9 9 10 6 8 6
Lower front 5 5 9 3 1 1 1 2 2
Upper hind 5 4 1 2 2 1 1 3 2 1 1 13 7 16 9
Lower hind 4 2 20 4 3 3
Feet 3 3 2 2 1 1 1 1 1 1
Other 23 4 5 1 6 1 2 1 1 86 176 2
Total 123 60 63 20 24 15 16 10 22 15 13 12 133 26 235 24

Carcass unitdefinition (# bones): Antler/horn (2); head ¼ bony parts of head (4
maxilla and mandible); upper teeth ¼ upper premolars
(permanent and deciduous) and molars (2); lower teeth ¼ lower premolars (permanent and deciduous) and molars (2); teeth ¼ other
teeth (2); neck ¼ hyoid, atlas, axis, cervical vertebra (8); back ¼ thoracic vertebra, lumbar vertebra, ribs (45); pelvis ¼ sacrum, innominate
(3); upper front ¼ scapula, humerus, radius, ulna (8); lower front ¼ carpals, metacarpals (12); upper hind ¼ femur, patella, tibia (6); lower
hind ¼ tarsals, metatarsals (12); feet ¼ phalanges (24); other ¼accessory phalanges and metapodials, sesamoids, etc.

Copyright # 2005 John Wiley & Sons, Ltd. Int. J. Osteoarchaeol. 15: 84–105 (2005)
Late Mousterian Subsistence in Dalmatia
Figure 8. Carcass part frequency (corrected MNE) of chamois þ ibex þ small ungulate and red deer þ medium ungulate at Mujina
Pec¤ina.
progressively drop from upper limbs to feet. From
a food utility perspective, we have a similar
representation of relatively high-ranked (upper
limbs) and low-ranked (antler/horn) carcass units.
Further discussion of these issues requires more
detailed data on element frequency and bone
modification by element. With the exception of
chamois þ ibex þ small ungulate, data sets are
too small to warrant such analyses. Therefore
we limit discussion to these smaller ungulates
(Table 9).
Chamois þ ibex þ small ungulate element re-
presentation is not significantly correlated with
sheep meat weight. On the other hand, there is a
very strong correlation between element repre-
sentation and sheep marrow volume (Figure 9,
Spearman’s r ¼ 0.722, t ¼ 3.296, 0.008, 10 d.f.).
This pattern suggests preferential selection of
marrow-bearing bones for transport and proces-
sing. However, there is also a strong correlation
between element representation and caribou
bone mineral density (Figure 10, Spearman’s
r ¼ 0.642, t ¼ 3.741, p ¼ 0.001, 20 d.f.). These
latter data suggest that any patterning in element
frequency can be explained by differential
destruction of bones mediated by bone mineral
density. While these two patterns are not neces-
sarily contradictory, we think that the bias
towards long bones (relatively high marrow
yield) reflects in the first instance the destruction
of lower-density articular ends and axial bones.
This strong pattern of density-mediated destruc-
tion suggests that whole carcasses were brought
Copyright # 2005 John Wiley & Sons, Ltd.
99
to the site followed by significant bone destruc-
tion. Modifications to chamois þ ibex þ small
ungulate elements show interesting patterns
(Table 9). Firstly, impact scars are found only
on long bone shafts. These fracture scars were
most likely created by hominins breaking open
long bone shafts for marrow extraction. Cut
marks are also limited solely to long bone shafts.
The location and morphology of these marks
suggests defleshing; dismemberment, apparently
was achieved without leaving marks. Evidence of
carnivore gnawing and digestion, in contrast, is
found on a wide range of elements, including
long bone shafts, articular ends, small bones
(carpals, phalanges), and bones of the axial ske-
leton. We suggest that the most likely scenario
for the accumulation of these remains is that
hominins hunted chamois and ibex and trans-
ported their carcasses back to the cave. Carcasses
were minimally processed before consumption,
with hominin damage mostly arising from
defleshing meaty limbs.10 After the hominins
left the site, there was sufficient food value left
in the bone waste to attract carnivores (hyena
and/or wolf) that then ravaged the assemblage
and used the site as a latrine. These scavengers
either destroyed or removed axial elements (neck
and pelvis) and long bone articular ends (Marean
et al., 1992). The relatively high frequency of
10
The available data suggest a similar interpretation of the red
deer+medium ungulate assemblage, although sample sizes are not
sufficiently large to support such a detailed analysis.
Int. J. Osteoarchaeol. 15: 84–105 (2005)
100 P. Miracle
Table 9. Element frequency and bone modification in chamois þ ibex þ small ungulate. Bone mineral density data from Lam et al.
(1999); whole-bone (WB) sheep utility data from Binford (1978) as modified by Miracle (2002)

%NISP marked Bone mineral density WB Sheep Utility

Element NISP MNE MAU Cut Impact scar Tooth Digested Site CT Caribou1 Meat wt. Marrow vol.

UP Teeth 25 12 6.00 4.0

LOW Teeth 30 13 6.50
Teeth 18 1 0.50 5.6
HC 8 5 2.50 12.5 1
SK 13 7 3.50 179
MD 6 3 1.50 DN3 1.07 1026 5
AT 0 0 0.00 AT3 0.49 320.34
AX 0 0 0.00 AX1 0.62 320.34
CV 2 2 0.40 CE1 0.45 951.24
TV 2 2 0.15 50.0 TH2 0.53 1469.62
LV 1 0 0.00 LU3 0.51 665.94
PEL 0 0 0.00 IL2 1.02 1303.75 3
RIB 10 5 0.19 20.0 RI3 0.96 1622.8
STM 0 0 0.00 1807.01
SC 1 0 0.00 SP2 1.04 769.66 2
HUM 6 5 2.50 16.7 HU3 1.12 629.71 19
RC 11 8 2.00 18.2 9.1 RA3 1.09 433.07 14
CAR 2 2 0.17 50.0 UNICF 0.72 258.32
MC 3 3 1.50 33.3 33.3 33.3 MC3 1.1 258.32 8
FEM 5 2 1.00 20.0 FE4 1.15 1353.2 24
TIB 11 7 3.50 9.1 9.1 18.2 9.1 TI3 1.13 869.08 24
TAR 1 1 0.17 NC2 0.62 869.08
AST 0 0 0.00 AS2 0.7 869.08
CAL 1 1 0.50 CA2 0.8 869.08 1
MT 3 1 0.50 33.3 33.3 MR3 1.1 479.64 10
P1 1 1 0.13 P12 0.92 255.93 2
P2 1 1 0.13 100.0 P22 0.72 255.93 1
P3 2 2 0.25 50.0 P31 0.48 255.93
MP 4 0 50.0 25.0
LB shafts 79 0 7.6 5.1 7.6 8.9
Other 10 2 10.0
Total 256 86 3.1 3.1 8.6 5.1

1
BMD2 from Lam et al. (1999).

Figure 9. Sheep marrow volume vs. MAU (Minimal Animal Unit) Figure10. Log-normal plot of caribou BMD (bone mineral density).
chamois þ ibex þ small ungulate at Mujina Pec¤ina. vs. MAU chamois þ ibex þ small ungulate at Mujina Pec¤ina.

Copyright # 2005 John Wiley & Sons, Ltd. Int. J. Osteoarchaeol. 15: 84–105 (2005)
Late Mousterian Subsistence in Dalmatia
tooth marks on small ungulate long bone mid-
shafts relative to other taxa (Table 7) may reflect
the greater food value of the small ungulates
relative to other parts of the assemblage. Thus,
the tooth- and cut- mark data may be more
informative about processing techniques than
procurement strategies. These scavenging carni-
vores also transported to Mujina remains of equid
and hare carcasses; the few neonatal bear remains
from Layer B þ C might represent hibernation
deaths or carnivore food remains.
Age and season at death
Age at death was inferred on the basis of bone
growth, epiphyseal fusion, and dental eruption
and wear. While remains were aged as precisely
as possible, and as discussed below some data are
precise enough to also indicate season of death,
they were not sufficiently numerous to warrant a
detailed analysis of mortality profiles. Instead,
we use four major age categories: neonatal,
juvenile, adult, old adult (Table 10). ‘Neonatal’
includes bones that based on size and texture
come from late-term fetuses and/or recently born
young. ‘Juvenile’ includes all other remains with
unfused epiphyses and/or deciduous dentitions
with wear on occlusal surfaces. ‘Adult’ includes
dentitions that have replaced deciduous with
permanent teeth as well as late-fusing bones
with fused epiphyses, while late-erupting cheek
teeth (premolars and third molars) with heavy
wear were assigned to ‘old adult’. Although
sample sizes are very small, the results are strik-
ing. The small bear assemblage is dominated by
Table10. Summarizedageingdatafromteethandbonesfor the
major taxa at Mujina Pec¤ina
%NISP
Taxa Fetal/neonatal Juvenile Adult Old adult NISP
Chamois þ ibex 0.0 20.3 74.3 5.4
Red deer 0.0 13.3 66.7 20.0
Large bovid 10.5 73.7 10.5 5.3
Equid 0.0 57.1 42.9 0.0
Bear 50.0 41.7 8.3 0.0
Hare 0.0 50.0 50.0 0.0
Copyright # 2005 John Wiley & Sons, Ltd.
101
remains of very young or juvenile cubs. These
very young cubs may have died during hiberna-
tion or have been killed by males, either at
Mujina itself or scavenged and transported to
Mujina from a nearby lair. The small hare assem-
blage contains equal numbers of juvenile and
adult remains. The large bovid and equid assem-
blages are dominated by remains from juvenile
animals; the former includes a few fetal/neonatal
bones and a large number of juvenile remains,
while the latter includes a more even representa-
tion of juvenile and adult remains. The large
bovid data, in particular, are suggestive of
attritional mortality. The red deer and cha-
mois þ ibex assemblages, in contrast, are domi-
nated by remains from adult animals. This
dominance of prime-aged adults, in conjunction
with other evidence of carcass processing, sug-
gests hunting by Mujina’s hominids. With
regards to chamois þ ibex, our interpretation of
mortality data contrast with that suggested by
modifications to long bone mid-shafts.
Only a few specimens (N ¼ 10) can be aged
with sufficient precision to be useful indicators
of season of death. These data are presented in
Table 11. Two bovid dentitions from Level D,
including a maxilla with deciduous teeth identi-
fied as Bison priscus, are from calves only four to
seven months old. Assuming a season of birth in
late spring/early summer (May-June), these
remains would have come from an animal
killed during autumn/early winter (Wegrzyn &
Serwatka, 1984). The majority of seasonality
indicators from Level B are fetal/neonatal bear
bones (Table 11). The very small size of all of
these bones indicates mother/cub deaths during
winter hibernation (Stiner, 1998). The other
seasonality indicators are from spring (fetal large
bovid tibia) and autumn (ibex mandible). Homi-
nins were definitely coming to Mujina during
the autumn in Level B. If they were also respon-
sible for the large bovid remains, which seems
Aged likely given the evidence of bone modification,
then they also had occasion to visit the site
during the spring in Level B and during the
74
15 autumn in Level D. There is no evidence of
19 hominins at Mujina during the summer or win-
7 ter, while bears were definitely at the site and/or
12
8 were brought to the site by scavengers during
the winter.
Int. J. Osteoarchaeol. 15: 84–105 (2005)
102 P. Miracle

Upperdp4, very light wearon mesial cusps

Conclusion

with occlusal surface just above alveolus

Measurements (mm) and wear stages

Lower dp4 ¼ stage 4, M1 ¼erupting

The faunal assemblage from Mujina Pećina thus
provides an initial glimpse of Late Middle Palaeo-

Lower dp4, very light wear

lithic food procurement and management, as well
as site use, in Dalmatia. The significance of these
GL ¼115, SD ¼17.1

initial results is considerably heightened by the

paucity of other assemblages in the region (Figure
1). The nearest Middle Palaeolithic faunal assem-
SD ¼ 10.2
SD ¼ 13.9

SD ¼ 5.8

SD ¼ 7.6
SD ¼ 7.6
blages to the southeast are from Crvena stijena in
Montenegro (Malez, 1975b) or Asprochaliko in
northwestern Greece (Bailey et al., 1983), while
the closest, well-excavated assemblages to the
northwest are from Divje Babe in Slovenia (Turk,
Source of information

Habermehl (1975: 65)

Couturier (1962: 330)

Serwatka (1984: 118)

Serwatka (1984:118) 1997) or Riparo di Fumane and Riparo Tagliente

in northeastern Italy (Mussi, 2001).11 We cur-
Wegrzyn and

Wegrzyn and

rently lack an appropriate data base for compar-

isons within the region, although revision of old
assemblages like Krapina (Miracle, in prepara-
tion) and Veternica (Miracle & Brajković, 1992)
in the Hrvatsko Zagorje at least provides a start.
We thus have to limit discussion to widely
September-January
September-January
October-November
Spring mid-March

divergent spatial and analytical scales—the small

Autumn/winter
Autumn/winter
Seaon of death

scale of human activities at the site on the one

to mid-May

hand, vs. the large scale of Neanderthal strategies

Autumn

in Mediterranean Europe on the other hand.

Winter
Winter

Winter
Winter

Winter
Winter

At the local scale of the site, the larger mam-

mal remains do not track the environmental
changes indicated by preliminary observations
(1.5 months before birth)

of the sediments. Radiometric dates place the

Fetus 220^230 days

entire sequence in the middle of OIS 3 (Oxygen

Fetal/neonate
Fetal/neonate

Fetal/neonate
Fetal/neonate
Fetal/neonate

Isotope Stage 3) (Rink et al., 2002). The precision

4^5 months

4^7 months
4^7 months

of these dates does not allow a closer correlation

with specific stadials/interstadials within OIS 3.
Fetal
Age

The faunal composition shifts from a co-domi-

Table11. Seasonality indicators from Mujina Pec¤ina

nance of red deer and chamois þ ibex in Layer

D1 þ D2 that formed during relatively cold con-
alveolus for erupting M1
Mandible with dp3 -M1

ditions to a clear dominance of wild caprids

Radius, proximal end
Radius, proximal end
Humerus, distal shaft
Humerus, distal end
Humerus, shaft with

Bison priscus Maxilla with dp2^4,

followed by large bovids and equids in Layer

nutrient foramen

B þ C that formed during relatively warm condi-

Radius, shaft
Tibia, shaft

Lower dp4

tions. Non-hominin carnivores played a signifi-

Element

cant role in the modification of the faunal

assemblages throughout the stratigraphic
sequence. Carnivore activities on site, however,
changed over time. In Layer D1 þ D2 carnivores
Capra ibex
Bos/Bison

Bos/Bison
Ursus sp.
Ursus sp.
Ursus sp.

Ursus sp.
Ursus sp.
Ursus sp.

scavenged hominin food refuse, as shown by the

Species

11
The final publication of Crvena Stijena provides little more than a
species list (Malez, 1975b), while we have only a preliminary report
Level

of species composition without taphonomic analyses from Aspro-

D2
D1

chaliko (Bailey et al., 1983).

B

B
B
B

B
B
B

Copyright # 2005 John Wiley & Sons, Ltd. Int. J. Osteoarchaeol. 15: 84–105 (2005)
Late Mousterian Subsistence in Dalmatia
relatively high frequency of digested bones (all
from ungulates), without using the cave as a den.
In Layer B þ C, in contrast, bears were occasion-
ally using the site as a den during the winter;
other carnivores only occasionally scavenged
food refuse, whether created by hominins or
bears. Hominin meat processing and/or con-
sumption is thus more prevalent at Mujina during
the colder conditions of Layer D1 þ D2; bears
succeeded in hibernating at Mujina only during
the warmer conditions of Layer B þ C. This tem-
poral pattern suggests that hominins controlled
access/use of the cave; they had priority of access
at a time when one might expect greater competi-
tion for shelter (e.g. relatively cold conditions).
The Late Mousterian faunal assemblage from
Mujina Pećina shows the hominins to be compe-
tent hunters within a context of considerable
competition from non-hominin carnivores. Our
pattern of red deer and caprid hunting at ca.
42 kyr fits Stiner’s (1994) suggestion of a shift in
hominin subsistence niches from scavenging to
hunting of medium- and large-sized ungulates
about 55 kyr in Latium, Italy. A simple case of
red deer and caprid hunting at Mujina even seems
banal in the context of accumulating evidence of
systematic hunting by Mousterian (and earlier?)
hominins in other contexts (Gaudzinski, 1999;
Marean & Assefa 1999). As we have repeatedly
stressed above, patterns in the faunal data are
much more subtle than ‘hunting vs. scavenging’ or
‘hominins vs. carnivores’. These ‘mixed’ hominin-
carnivore signatures can be pulled apart through
detailed taphonomic analyses of well-excavated
assemblages. Our analyses thus highlight some of
the dangers of calculating species diversity or
meat weight from faunal composition without
taking into account detailed taphonomic analyses.
One of the surprises of this study is the extent to
which hominin behavioural information remains
accessible despite the considerable modification
of the assemblage by carnivores, weathering, and
other taphonomic agents. We have touched on
the potential impact of environmental changes
and competition from non-hominin carnivores on
faunal patterning at Mujina Pećina. The location
the site and the age of its deposits make competi-
tion with other hominins an intriguing possibility.
The site is located along a potential migration
corridor for hominin populations moving into
Copyright # 2005 John Wiley & Sons, Ltd.
103
Europe from western Asia. Similarly there is
potential overlap between Mujina’s radiometric
dates and those from Early Upper Palaeolithic
contexts in Europe. The Mujina Pećina assem-
blage thus provides a significant point of reference
for a broader-scale study of variability in Mous-
terian subsistence practices in their own right as
well as within the context of the Initial Upper
Palaeolithic in southeastern Europe.
Acknowledgements
Ivor Karavanić provided me with the opportunity
to study the Mujina faunal assemblage. He also
provided invaluable information about excava-
tions at Mujina and the context of the faunal
assemblage. The manuscript has benefited from
the comments of Katie Boyle, Terry Hopkinson,
and Mike Petraglia. Two anonymous reviewers
helped tighten the taphonomic arguments, and
one in particular offered very useful suggestions
for the interpretation of some of the illustrated
specimens. Any errors that remain are my own.
Research visits to Croatia have been supported,
in part, by the University of Cambridge and
St John’s College.
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