16_Kitagawa et al_TRIS

P ROM E T H E U S P RE S S /P A LAE ON T OL OG IC AL N E T W ORK F OU ND AT IO N
(TERUEL)
2012
Journal of Taphonomy
Available online at www.journaltaphonomy.com
Kitagawa et al.
VOLUME 10 (ISSUE 3-4)
Exploring Cave Use and Exploitation Among
Cave Bears, Carnivores and Hominins in the
Swabian Jura, Germany
Keiko Kitagawa*, Petra Krönneck
Institut für Naturwissenschaftliche Archäologie, Eberhard-Karls-Universität Tübingen
Rümelinstr. 19-24, 72070 Tübingen, Germany
Nicholas J. Conard
Institut für Ur- und Frühgeschichte und Archäologie des Mittelalters
Abteilung Ältere Urgeschichte und Quartärökologie &
Senckenberg Center for Human Evolution and Quaternary Ecology, Schloss Hohentübingen
Eberhard-Karls-Universität Tübingen, 72070 Tübingen, Germany
Susanne C. Münzel
Institut für Naturwissenschaftliche Archäologie, Eberhard-Karls-Universität Tübingen
Rümelinstr. 19-24, 72070 Tübingen, Germany
Journal of Taphonomy 10 (3-4) (2012), 439-461.
Manuscript received 15 March 2012, revised manuscript accepted 15 November 2012.
This study offers an overview of carnivore remains from archaeological contexts and provides evidence
of interaction between carnivores and hominins in the Swabian Jura during the Middle and Upper
Paleolithic (~50,000-27,000 uncal B.P.). First, we present data on the carnivores in the faunal
assemblages from the area, followed by a general comparison of anthropogenic and carnivore
modifications on faunal remains. Further, we describe some archaeological findings that demonstrate
unique ways in which humans interacted with carnivores in the early and middle Upper Paleolithic. This
study documents the pattern of carnivore representation in the zooarchaeological assemblages on a
regional scale. The analyses of faunal assemblages across five cave sites in the two valleys of the Swabian
Jura indicate intensified use of caves by humans, which corresponds to a decreased presence of large and
medium sized carnivores at the beginning of the Upper Paleolithic in addition to the use of carnivore
figures in the cultural repertoire of the Aurignacian and the increased exploitation of carnivores in the
Gravettian period.
Keywords: MIDDLE PALEOLITHIC, AURIGNACIAN, GRAVETTIAN, CAVE USE, CAVE BEARS,
CARNIVORES, SWABIAN JURA
Article JTa145.
JTa124. All rights reserved.
*E-mail:
*E-mail:[email protected]
[email protected]
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Exploring Cave Use and Exploitation in the Swabian Jura
Introduction
Signatures of carnivores prevail in
archaeological sites throughout the
Paleolithic period. While little evidence of
direct interaction between hominins and
carnivores is documented in the archaeological
record until the middle of Upper Paleolithic
(with few exceptions: Blasco et al., 2010;
Münzel & Conard, 2004a), non-human
predators played a crucial role in the
evolution of hominins as they began to
exploit large game for food and entered an
ecological guild of medium and large
carnivores. This change in the adaptation of
hominins triggered debates on hunting and
scavenging as well as the interaction of
carnivores and hominins during the Lower
Paleolithic at eastern African sites
(Blumenschine, 1986; Blumenschine &
Pobiner, 2006; Bunn & Kroll, 1986;
Dominguez-Rodrigo & Barba, 2006) and
led to increased interest in understanding
the taphonomic effects of carnivores to
extrapolate archaeological signals from
faunal assemblages (Brain, 1981; Cleghorn
& Marean, 2007; Selvaggio & Wilder,
2001). Some researchers focused on the
evolution of medium and large carnivores to
infer the nature of competition and
coexistence among different species, which
also has implications for studying the past
adaptation of early Homo (Van
Valkenbergh, 2001; Lewis & Werdelin,
2007). The record of past animal remains,
especially ungulates, provides a direct
evidence of competition for resources,
which shaped the adaptation of hominins
and the nature of coexistence in various
ecological settings.
Hominins and large carnivores also
occupied a similar ecological niche in that
they both used caves as natural shelters (e.g.
Stiner, 1994). Caves provide protection for
carnivores to consume prey for avoiding
competition and rear young, and bears
occupy sheltered area during the period of
hibernation. Further, the majority of
archaeological records derives from the
context of caves or rockshelters due to
better preservation of organic remains. We
infer that carnivores and hominins had
similar need for resources and space ever
since hominins began to regularly exploit
animals for resources and as hominins
dispersed in areas where they encountered
new faunal communities and adapted to
different ecological systems (for a
summary, see Stiner, 2002).
That said, understanding how caves
were occupied and utilized by hominins and
carnivores is often based on isolated cases
of single sites, and with some exceptions,
not many studies have focused on the
regional pattern of cave use by carnivores
and hominins. Mussi (2001) and Straus
(1982) compiled data on a larger scale,
namely in parts of Italy and Cantabria,
Spain to study patterns of carnivore
representation in assemblages spatially and
temporally. Most interactions cannot be
tracked using fossil records alone, but
studying cave use and exploitation of prey
by carnivores and hominins beyond one or
two sites sheds light on the ecological
relationship and trends that speak to
differences in the adaptation of archaic and
modern humans. This study consists of
faunal analyses from caves in the Swabian
Jura during the Middle and Upper
Paleolithic. We sought to observe regional
patterns in the changes of cave use by
considering carnivore abundances, bone
modification, exploitation of carnivores by
humans and lastly, the cultural significance
as symbols during the Upper Paleolithic.
440
Kitagawa et al.
Swabian Jura: Geography, Sites and
Paleoenvironment
The geographical area of Swabian Jura is
the largest karst system in southwestern
Germany (Figure 1). Several caves with
Paleolithic deposits are mainly found in two
valleys of the Swabian Jura, the Ach and
Lone. Sites shown on the map are situated
in these valleys that cut through a dry
upland plateau, which formed during the
Upper Jurassic period and reaches as high
as 1000 m asl, dipping slightly towards the
southeast. Most of the archaeological sites
are found at an elevation of approximately
560-585 m asl (Abel et al., 2002). The Ach
Valley formed as the Danube River, which
runs 5-10 km south in the present,
meandered into the plateau before the Riss
Glacial Stage, and the Lone Valley formed
as a result of a meandering tributary, which
drained into the Danube River. Local flints
(with sources usually within 5-10 km
distance from all caves) were commonly
used for the production of lithic artifacts
(Burkert & Floss, 2006). Ach and Lone
Valleys are similar in their geological
setting, but the Ach Valley is deeply incised
compared to the Lone Valley while the
latter is dominated by open grassland at a
lower elevation, a factor that may explain
the abundance of horses in the Paleolithic
period. Further, the Lone River is fed by the
karst aquifer with no permanent water
source whereas the Ach River flows year
round in the Ach Valley today.
Sites in the Lone Valley, Hohlenstein
-Stadel, Vogelherd and Bockstein and in the
Ach Valley, Hohle Fels and Geißenklösterle
are considered in this study. These sites
Figure 1. Map of the Swabian Jura with major sites included in the study: Hohle Fels and Geißenklösterle in the Ach valley; Bockstein, Vogelherd and Hohlenstein-Stadel in the Lone valley.
441
Exploring Cave Use and Exploitation in the Swabian Jura
have produced one of the richest
archaeological records of the Paleolithic in
Central Europe and revealed an early
emergence of the Aurignacian culture with
an array of innovation and new forms of
symbolic expression including ivory figurines,
music and ornaments (Conard, 2009;
Conard & Malina, 2006; Conard et al.,
2009; Kölbl & Conard, 2003). Große
Grotte, Kogelstein, Sirgenstein and
Brillenhöhle were not included in this study
as these sites either do not have both Middle
and Upper Paleolithic sequences or the
material has not been studied in detail. The
Middle Paleolithic sequences have not been
dated securely with the exception of a
deposit from Geißenklösterle, which dated
to 43,000 BP using TL and ESR dating in
the uppermost level of the Middle
Paleolithic sequence (Richter et al., 2000)
as well as some radiocarbon dates whose
range suggests a minimum age of 50-45,000
cal BP from the Middle Paleolithic horizons
in Hohlenstein-Stadel (Beutelspacher et al.,
2011). The Aurignacian appears to begin
immediately after the end of the Middle
Paleolithic around 43,000 to 41,000 cal BP
(Conard & Bolus, 2003, 2008; Higham et
al., 2012). The deposits in Bockstein, Hohle
Fels and Geißenklösterle also contain
Gravettian sequences, which span the period
of 30-27,000 BP (Moreau, 2009).
Prehistorians and geologists began
archaeological investigations in this area
starting in the mid 19th century and some of
the analyzed faunal assemblages derive from
older excavations that occurred extensively
from the 1930s, namely the three sites from
the Lone Valley. In contrast, the majority of
the faunal material from Hohle Fels and
Geißenklösterle was excavated using more
comprehensive recovery techniques starting
in the 1970s (Hahn, 1988). Despite differences
in the excavation method, all five sites have
cultural sequences that run from the Middle
Paleolithic to Aurignacian and three sites,
Hohle Fels, Geißenklösterle and Bockstein,
also contain Gravettian sequences (another
transitional layer between the Aurignacian
and the Gravettian at Hohle Fels exists, but
it was not considered in the study for
adequate comparison). Geological layers as
well as separate deposits (in the case of
Bockstein) from the Middle Paleolithic,
Aurignacian and the Gravettian are
aggregated to provide adequate sample sizes.
Paleoenvironmental records in the
Swabian Jura allow us a glimpse into the
past climatic shifts, which are particularly
relevant in interpreting the abundance of
fauna with preferred habitats. Previous work
on archaeobotanical remains from archaeological
contexts such as Bockstein indicates steppic
vegetation with some woodland in the
Middle Paleolithic and cooler climate
towards the beginning of the Last Glacial
Maximum (LGM) (Filzer, 1969). In addition,
the nearest pollen core from the alpine
foreland in Germany, Füramoos, reveals mild
climate from roughly 51,000-40,000 BP
followed by a hiatus, which lasted until the
end of the glacial period due to heavy
erosion (Müller et al., 2003).
Some work on mammalian macroand microfauna as well as birds points to a
steppe landscape with woodland area that
persisted from the Middle Paleolithic to the
Aurignacian period (Krönneck, 2012, in
press; Ziegler, in press). Microfaunal data
from Geißenklösterle indicate slightly warmer
conditions during the Aurignacian with a
decrease in tundra and increase in wooded
area (Ziegler, in press). Finally, the Gravettian
period is characterized by a steppe
environment with a decrease in the wooded
areas (Conard et al., 2006; Krönneck, 2012).
442
Kitagawa et al.
Geological and micromorphological
analyses from sites of Geißenklösterle and
Hohle Fels offer a different reconstruction
of the environment with some intersite
variability (Conard et al., 2006; Goldberg
and Conard, in press; Miller, 2009). In
Geißenklösterle, micromorphological samples
taken from the transition between the
Middle Paleolithic and the Aurignacian are
marked by an episode of erosion, indicating
an abrupt shift from a mild condition with
warm and wet climate to a colder and drier
climate. While the general trend is comparable
at Hohle Fels, the change appears to be gradual
with mild climate lasting through to the
beginning of the Aurignacian, which is then
followed by a cooler climate, a climatic shift
that occurred presumably after the arrival of
modern humans. In the Gravettian, there is a
clear indication of colder and drier climate
leading towards the beginning of the LGM.
In all, the paleoenvironmental record
shows that the steppe environment mostly
prevailed in the Swabian Jura with some
woodland and riverine forests. To date, an
abrupt shift in the climate between the
Middle Paleolithic and the Aurignacian is
documented only by the micromorphological
study of Geißenklösterle, while other lines of
evidence mostly document relatively stable
conditions during the transitional period,
followed by some climatic fluctuations
within the Aurignacian period. From the
end of the Aurignacian, the cooler climate
prevailed and persisted throughout the
Gravettian period until ~27,000 uncal. B.P.
Previous Work on Fauna of the Swabian Jura
Zooarchaeological research of the Swabian
Jura largely focused on two themes: the
subsistence pattern of Neanderthals and
modern humans (Krönneck et al., 2004;
Münzel, in press; Münzel & Conard, 2004a;
Niven, 2006) and the paleoecology of cave
bears and their possible interactions with
hominins (Münzel & Conard, 2004b; Münzel
et al., 2011). Horses (Equus ferus) and reindeer
(Rangifer tarandus) were extensively exploited
in the Swabian Jura during the Paleolithic
period. Horses were a preferred taxon in the
Lone Valley and reindeer remains are more
common in the Ach Valley. Bovids are also
present across sites, but there is a greater
proportion of ibex (Capra ibex) at the sites
in the Ach Valley, which probably served as an
ideal habitat with high reliefs for these
bovids (Münzel, in press). Further, Vogelherd
holds one of the richest assemblages of
mammoth (Mammuthus primigenius) remains
in Central Europe although the degree of its
exploitation for subsistence purpose is not
clearly demonstrated (Niven, 2006). Across
sites, many mammoth remains are found in a
form of ivory fragments, which was commonly
used as raw material for the production of
artifacts (Conard et al., 2006; Niven, 2007).
Other ungulates found in the Swabian Jura
include woolly rhinoceros (Coelodonta
antiquitatis), aurochs/bison (Bos/Bison),
giant deer (Megaloceros giganteus), red
deer (Cervus elaphus), chamois (Rupicapra
rupicapra) and wild boars (Sus scrofa).
In some sites such as Vogelherd,
repetitive cutmarks and selective transport
of body parts point to the significant role of
ungulates and proboscideans in humans’
diet during the Aurignacian period (Niven,
2006, 2007). However, the selection of prey
by hominins does not largely alter during
the Paleolithic. While the archaeological
record reveals differences in the intensity
and settlement pattern of Neanderthals and
early modern humans, the choice of prey
remained unchanged (Conard et al., in press).
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Exploring Cave Use and Exploitation in the Swabian Jura
Such continuity in the selection of prey
most likely reflects the abundance of
respective ungulates that inhabited or migrated
through the Swabian Jura, with horses
preferring the open land of the Lone Valley
and reindeer, which were exploited during the
seasons of migration (Müller-Beck, 1988;
Niven, 2006).
In recent years, some studies
explored the possible use of small game by
Neanderthals and modern humans (Conard
et al., in press; Krönneck, in press; Münzel,
in press). While preservational bias against
small bodied animals must be investigated
in greater detail, preliminary results suggest
a gradual increase in the abundance and the
diversity of prey including small mammals
such as hares, birds and fish from the
Middle Paleolithic to the Gravettian period.
Such patterns may reflect changing subsistence
practices that resulted from increased
human population or longer residential
times at caves in the Aurignacian period
(Conard 2011; Conard et al., 2012), but the
pattern of prey choice of the Neanderthals
and early modern humans are comparable,
documenting no clear shift in the economic
behavior of the hominins in the Swabian
Jura.
Other faunal studies concentrated on
cave bears that are most commonly found in
the Ach Valley, including at sites discussed here
and Große Grotte (Weinstock, 1999) and the
sites of Hohlenstein-Stadel and Hohlenstein
-Bärenhöhle in the Lone Valley (Lehmann,
1960). Genetic analyses demonstrated the
existence of two genetically different cave
bears (Ursus spelaeus and Ursus ingressus)
with the latter replacing the former around
31,270 ± 325 cal BP and Ursus ingressus
becoming extinct around 29 760 ± 530 cal.
BP in this part of Central Europe (Münzel et
al., 2011). Stable isotopic analyses confirmed
their herbivorous diets showing distinct
values of d15N and d13C from coeval
carnivorous brown bears (Bocherens et al.,
1994; Münzel et al., 2011). Co-occurrence
of cave bear remains and hominin
occupations has allowed researchers to
explore the nature of interaction among
humans and cave bears and propose
possible cause(s) for the extinction of cave
bears during the Gravettian period (Münzel
& Conard, 2004b; Stiller et al., 2010).
Data and Results
In addition to the faunal analysis of
Bockstein, Vogelherd and Geißenklösterle,
the analysis of material from Hohle Fels and
the taphonomic study from HohlensteinStadel are currently in progress. Approximately
half of the material from each cultural layer
at Hohle Fels has been analyzed by S.
Münzel and the previous work by C. Gamble
on Hohlenstein-Stadel (1979, 1999) gives us
a general understanding of the species’ relative
abundance and diversity. All five sites consist
of caves of different sizes, which have been
occupied during the Middle and early Upper
Paleolithic. Further, faunal data from the
Gravettian layers of Bockstein, Hohle Fels
and Geißenklösterle are reported here. Data
in this study consist of species abundance with
a focus on carnivores, bone modification by
carnivores and humans, as well as anthropogenic
modification of carnivore remains.
Taxonomic Data
Intersite variability characterizes the faunal
spectrum of the Swabian Jura. The relative
abundances of fauna categorized under
ungulate, large carnivores, small carnivores,
cave bear and small mammals, notably
444
Kitagawa et al.
hares (Table 1-3, Figure 2), show that while
some sites are dominated by ungulates,
some sites are represented by high
concentrations of cave bear. Such contrast
points to the difference in the site use by
animals throughout the Paleolithic period.
Ungulates, horses and reindeer,
dominate throughout the occupational
sequences of Vogelherd and Bockstein. The
assemblage size of Bockstein from the
Aurignacian and Gravettian is small in
comparison to the Middle Paleolithic, but
high %NISP of ungulates and relatively low
%NISP of large carnivores points to a
continuing trend in the exploitation of prey
that were transported and butchered in the
caves by hominins. Vogelherd served an
ideal site to observe prey movement due to
its elevation, allowing for a greater visibility
over a spot where two valleys merge
(Niven, 2006). When we consider the
taxonomic data and the relative abundance
of ungulates, these sites appear to have
served as locales for transport and
processing of animal carcasses with some
degree of temporal continuity.
Cave bears are taxonomically placed
in the order of Carnivora, but have a
different ecological niche as other carnivores,
because of their herbivorous diet as indicated
by the tooth morphology (Kurtén, 1976;
Rabeder & Hofreiter, 2004) and isotopic
studies (Bocherens et al., 1994). In this study,
we include cave bears in the analysis as they
directly competed with hominins for the use of
shelters. Cave bears dominate the assemblages
of Geißenklösterle, Hohle Fels, and
Hohlenstein-Stadel. With the exception of
Hohlenstein-Stadel, %NISP of cave bear at
these sites decreases from the Middle
Paleolithic to the Aurignacian period, but
cave bear continues to be a dominant taxon
throughout the sequences, including the
Gravettian period in the Ach Valley. We can
conclude that sites were often occupied in the
winter season by cave bears and their remains
accumulated as they died of natural causes
and predation. Further, many examples of
fetal remains indicate the active use of the
sites as dens, as is especially well documented
in Geißenklösterle and Hohle Fels. The
abundance of cave bears in association with
artifacts suggests that the sites were visited
by various animals, which included hominins
but with long intervals between the occupations
that allowed other animals access to the
caves. Factors such as the frequency of visits
by animals and conditions of the cave
including the size and locality probably
played a role in the preference of cave bears
to use the caves as shelters for the winter.
These favorable conditions for some sites
probably remained consistent throughout the
Paleolithic period, as the dominance of cave
bears at certain sites does not change over
different cultural phases.
Large and middle sized carnivores
include lion (Panthera leo spelaea), hyena
(Crocuta crocuta spelaea), wolf (Canis
lupus), brown bear (Ursus arctos) as well as
lynx (Lynx lynx). They are found in varying
abundances across the sites (Table 1-3,
Figure 3). A general temporal pattern points
to a decrease in the NISP and %NISP of
large/middle carnivores and an increase of
ungulates, small carnivores and hares, from
the Middle Paleolithic to the Gravettian
with the exception of Hohlenstein-Stadel.
Such a trend signifies the decreased use of
caves by carnivores on a regional scale.
While a link between the trend and its
probable cause, increased human activities,
remains tentative, there is an inverse
correlation between intensified use of caves
in the Upper Paleolithic and decreasing
NISP of large/middle carnivores at sites
445
Exploring Cave Use and Exploitation in the Swabian Jura
Table 1. NISP of mammalian fauna in the Middle Paleolithic from the five sites of the Swabian Jura.
Taxa
Panthera leo spelaea
Crocuta crocuta spelaea
Canis lupus
Ursus arctos
Vulpes/Alopex
Ursus spelaeus
Mammuthus primigenius
Coelodonta antiquitatis
Bos/Bison
Equus sp.
Megaloceros giganteus
Cervus elaphus
Rangifer tarandus
Capra ibex
Lepus sp.
others
total NISP
Geißenklösterle
Hohle Fels
Hohlenstein-Stadel
Bockstein
Vogelherd
4
7
19
_
26
586
3
8
_
21
7
2
53
20
8
10
1
1
5
3
4
531
_
2
_
10
_
1
7
3
1
_
80
476
240
_
161
4066
61
125
156
1557
14
203
102
_
4
56
7
48
40
1
18
55
59
72
82
678
_
9
130
2
7
21
2
10
8
2
_
25
16
57
49
342
7
11
12
_
2
1
774
569
7301
1229
544
Table 2. NISP of mammalian fauna in the Aurignacian from the five sites of the Swabian Jura.
Taxa
Panthera leo spelaea
Crocuta crocuta spelaea
Canis lupus
Ursus arctos
Vulpes/Alopex
Ursus spelaeus
Mammuthus primigenius
Coelodonta antiquitatis
Bos/Bison
Equus sp.
Cervus elaphus
Rangifer tarandus
Capra ibex
Rupicapra rupicapra
Lepus sp.
others
total NISP
Geißenklösterle
Hohle Fels
Hohlenstein-Stadel
Bockstein
Vogelherd
1
13
91
9
5
25
3
34
1018
21
4
_
126
9
226
27
1
52
2
1562
15
44
157
_
97
1935
3
14
_
50
4
82
_
_
22
5
2428
6
5
_
6
28
6
_
8
72
1
32
_
_
8
1
173
4
17
38
2
20
120
1127
124
61
1423
19
1633
_
2
27
14
4631
159
2972
220
62
1
483
10
493
104
35
209
11
4864
446
Kitagawa et al.
Table 3. NISP of mammalian fauna in the Gravettian from the three sites of the Swabian Jura.
Taxa
Panthera leo spelaea
Crocuta crocuta spelaea
Canis lupus
Ursus arctos
Vulpes/Alopex
Mustela sp.
Ursus spelaeus
Mammuthus primigenius
Coelodonta antiquitatis
Equus sp.
Cervus elaphus
Rangifer tarandus
Capra ibex
Rupicapra rupicapra
Lepus sp.
others
total NISP
Geißenklösterle
1
2
14
1
109
1
1418
51
4
116
3
183
43
7
240
1
2194
(Conard 2011; Conard et al., 2006, 2012).
Interestingly, this trend is not observed for
cave bears, as NISP decreases but the %
NISP remains consistent over the Middle
Paleolithic to the Gravettian period.
Among the large and middle sized
carnivores in the Middle Paleolithic, the
most common species is hyenas, followed
by wolves (Figure 3). Besides the sites
discussed in text, the site of Kogelstein in
the Ach Valley, now a collapsed cave, has
yielded evidence of denning by hyenas
intermittent with hominin occupation
(Böttcher et al., 2000). Among the sites, the
overall abundance of carnivores indicates
that there is a decrease in the abundance of
hyenas during Aurignacian period, while
wolves become the most abundant predator,
a trend that continues to the Gravettian
period. We tentatively propose that the local
population of hyenas decreased in abundance
Hohle Fels
6
2
34
5
73
1
2268
40
1
336
3
309
49
2
474
2
3605
Bockstein
7
9
8
26
3
13
51
4
62
11
6
200
from the Middle Paleolithic to the
Aurignacian and Gravettian and the wolf
population remained unaltered during these
cultural phases. Lion comprises a small
proportion of the entire assemblages
without any apparent temporal trends or
spatial patterns. Large and middle sized
felids were often depicted in the figurines
from the Aurignacian period as discussed
below, but they did not utilize the cave for
shelter or to transport prey as intensively as
hyenas and wolves in this region. Further,
brown bears are also scarce in the
assemblages. Even accounting for
fragmentary remains and juvenile bones,
which may be misidentified as cave bears,
few brown bears are identified, showing
that brown bears, which coexisted and
competed for sheltered space with cave
bears, seldom visited the Swabian Jura
caves during the Paleolithic period.
447
Exploring Cave Use and Exploitation in the Swabian Jura
Figure 2a. Proportion of ungulates, middle/large and small carnivores, cave bears and hares in the
Middle Paleolithic.
Figure 2b. Proportion of ungulates, middle/large and small carnivores, cave bears and hares in the
Aurignacian.
Figure 2c. Proportion of ungulates, middle/large and small carnivores, cave bears and hares in the
Gravettian.
448
Kitagawa et al.
The Middle Paleolithic assemblage
in Hohlenstein-Stadel represents one of the
richest assemblages with large and middle
sized carnivores. Besides cave bears, hyenas,
wolves and lions exist in greater proportions
than in other deposits. Hints of Neanderthal
occupation, including artifacts such as lithics,
burnt bones and charcoal (Beck, 1999;
Beutelspacher et al., 2011) as well as a femur
of Neanderthal (Wetzel, 1961) exist, but
scant evidence of anthropogenic modification
on fauna suggests sparse use of the cave by
the archaic humans. Conversely, abundant
carnivore remains attest to greater input of
bones by non-human predators that can
account for the presence of ungulate remains.
Findings of coprolites and regurgitated
bones (identified by marks of acid etching,
thinning of the edges and polishing of the
bone surface) demonstrate an active use of
the cave by hyenas in addition to wolves. C.
Diedrich (2011) suggests that the predation
of cave bears in winter by hyenas and other
large carnivores occurred frequently during
the late Pleistocene. Cave bears may have
been a prey of choice in addition to ungulates
during winter when they were dormant or
may have been scavenged after their natural
deaths, although this idea should be tested
with detailed taphonomic studies. Thus, it
appears that the assemblage of HohlensteinStadel represents a palimpsest of hominin,
cave bear and carnivore activities, which is
unique in the Swabian Jura for the high
occurrences of large and middle sized
carnivores.
Small sized carnivores include artic
foxes (Alopex lagopus) and red foxes (Vulpes
vulpes) as well as occasional findings of
wolverine (Gulo gulo), wild cat (Felis
silvestris), marten (Martes sp.), badger (Meles
meles) and mustelids. Besides foxes, other
small carnivores are not represented in large
numbers for adequate comparison. In addition
to their high occurrences in the Middle
Paleolithic at Hohlenstein-Stadel on the
basis of NISP, foxes are more commonly
represented in the Aurignacian and in the
Gravettian. While the number of large and
middle sized carnivores decreases, the NISP
and %NISP of foxes increase over time.
Some researchers suggest an increasing
exploitation of foxes for use of pelts starting
in the Upper Paleolithic and beyond (Charles,
1997), but to date, few anthropogenically
modified remains of small carnivores in the
Swabian Jura have been identified. Since
small animals usually bear fewer cutmarks
and anthropogenic modifications, the increase
in the number of foxes over time tentatively
points to human involvement in the
accumulations of foxes at these sites, but
will require further taphonomic assessment.
Modification by carnivores and hominins:
implication for cave use
The modification of the bones by carnivores
and hominins reflects the degree in which
both agents were involved in the transport
and accumulation of animal remains in the
caves. The percentage of total specimens for
carnivore and anthropogenic modifications is
represented in Figure 4 and 5. Carnivore
modification includes evidence of gnawing,
scoring, pits and punctures. Regurgitated bones
and occasional teeth, a common form of
modification by wolves or hyenas, were
identified at the sites as well. Anthropogenic
modification compiled for this study refers to
cutmarks, percussion marks, burning, artifacts
and organic debitages.
We observe an increase in
anthropogenic modification and decrease in
carnivore modification consistently for the
major sites from the Middle Paleolithic to
449
Exploring Cave Use and Exploitation in the Swabian Jura
Figure 3a. %NISP of carnivores and non-carnivores in the Middle Paleolithic.
Figure 3b. %NISP of carnivores and non-carnivores in the Aurignacian.
Figure 3c. %NISP of carnivores and non-carnivores in the Gravettian.
450
Kitagawa et al.
the Aurignacian, although Hohlenstein-Stadel
will be examined further to assess this
pattern. Gravettian assemblages have also
produced a large number of human modified
remains and relatively few remains modified
by carnivores across all three sites. Different
forms of anthropogenic modifications,
including butchering and processing of
animal remains and the use of organic remains
for fuel and artifact production, increase
significantly from the Middle Paleolithic to
the Aurignacian period. To date, few remains
have yielded evidence of both anthropogenic
and carnivore traces, which points to the
primary agent responsible for the transport
of the remains into the caves. Few bone
fragments at Bockstein from the Middle
Paleolithic show cutmarks superimposed by
carnivore puncture marks (Krönneck, 2012).
Taxonomic representation of fauna in the
Swabian Jura differs among sites and shows
no regional trend, but the taphonomic study
reveals a consistent increase of human
traces and a decrease in the input of animal
remains by carnivores across sites from the
Middle and to the early Upper Paleolithic.
In addition to fauna, this trend is also
supported by micromorphological studies
from Geißenklösterle and Hohle Fels, which
show that fragments of coprolites are identified
in higher frequencies in the Middle
Paleolithic than in the Aurignacian (Miller,
2009). Further, there is an increased artifact
density as well as clearer visibility of human
activities in the sediments of the deposit
when the Middle Paleolithic is compared to
the Aurignacian (Conard 2011; Conard et
al., 2012). To securely determine whether
such trend reflects an actual signal of hominin
behavior, we need a better understanding of
the duration of Middle Paleolithic occupation
by dating the lower cultural sequences and
detailed taphonomic analyses. Nonetheless,
such increase of human input point to shifts
in the use of caves in the region by
Neanderthals and modern humans.
Direct human exploitation and cultural
significance of carnivores
Economic Use: Exploitation and Tools
Some carnivore bones and teeth bear unique
evidence of anthropogenic modifications
(Table 4). Evidence of cutmarks, burning
and use of carnivores as artifacts remains
relatively low during the Middle Paleolithic
and the Aurignacian, but increase during the
Gravettian period, a trend which is most
visible at Hohle Fels. One of the evident
examples is a flint projectile embedded in a
cave bear vertebra, dating to 28,000 uncal. BP
(Münzel & Conard 2004b). This specimen
alone cannot speak to the degree in which
cave bears were exploited systematically,
but it demonstrates that humans preyed on
cave bears during hibernation on occasion.
Exploitation of cave bear during winter
season is also demonstrated by a canine of
circa one year old cave bear with cutmarks
(Figure 6). Further, evidence of cutmarks
documents different steps of processing
carcasses including skinning (i.e. cutmarks
on skull and metapodials), disarticulation
(i.e. cutmarks on head of axis) and
defleshing (i.e. cutmarks on pelvis).
Anthropogenic modification of cave bear
remains is also found at other sites of the
Swabian Jura although it is relatively low in
number (Table 4, Figure 7a, 7b). S. Münzel
argues for a possible link between human
predation on cave bears and their extinction,
which occurred locally around the end of
the Gravettian (Münzel & Conard, 2004b;
Münzel et al., 2011).
451
Exploring Cave Use and Exploitation in the Swabian Jura
Figure 4. %NISP of specimens with carnivore modification.
Figure 5. %NISP of specimens with anthropogenic modification (Note: ^ small burnt
material not collected).
452
Kitagawa et al.
In addition, we observe uses of lions
during the Aurignacian and the Gravettian
period. Two forms of exploitation are
documented. Retouchers made from lion
canines are recovered from the sites of
Hohle Fels and Vogelherd (Figure 8). In
contrast, cutmarks on shafts of long bones, a
possible indication of skinning and defleshing,
were recovered from the transitional layer
between the Aurignacian and the Gravettian
period and from the Gravettian at Hohle
Fels (Figure 9). While these examples are few
in number, they nonetheless demonstrate
cases of exploitation that on the one hand
focused on the use of large carnivore
remains as raw material for the production
of tools, and on the other hand, processing
of the remains for meat and/or skin. It also
appears that canines from large carnivores
(including cave bear) were often used as
retouchers in the Aurignacian period in the
Swabian Jura (Taute, 1965).
Cultural significance of carnivores in the
Swabian Jura
The cultural significance of carnivores
emerged in the Aurignacian and persisted in
the Gravettian through different art forms.
Ivory played a crucial role as a medium for
symbolic expression of the humans in the
Swabian Jura during this period (Conard,
2003; Conard et al., 2006, 2009). We recognize
abundant depictions of carnivores and at times,
depictions of mixed human and carnivore
traits. Some of the well-known ivory figures
feature lions, which are highly concentrated
in Vogelherd (Figure 10), in addition to a bear
figurine from Geißenklösterle and a possible
hyena or lion from Vogelherd. Further, the
largest ivory figurine known as Lionman
(Löwenmensch) from the site of Hohlenstein
-Stadel has both anthropomorphic features
of a man’s body with a lion head (Figure 11)
(Wehrberger, 1994). Two additional figurines
from Geißenklösterle and Hohle Fels depict
a similar therianthropic form with felid
features. Here, we postulate that large felids
in particular had a crucial role in the
iconography and system of beliefs of the
Aurignacian culture in the Swabian Jura
(Conard, 2003). Aside from the faunal data,
ivory figurines from this period demonstrate
a unique ‘use’ of carnivores as motifs in the
cultural repertoire of the Aurignacian period.
During the Gravettian period, carnivore
teeth are increasingly utilized for personal
ornaments. The production and use of personal
ornaments emerge in Europe at the beginning
of the early Upper Paleolithic (Vanhaeren &
d’Errico, 2006). Common raw material
includes ivory, animal teeth and mollusks. In
the Swabian Jura, there is a clear preference
of ivory during the Aurignacian, although
some perforated fox teeth are recovered as
well (Floss, 2007). In the Gravettian, we see
an increased use of animal teeth as raw
material for ornament production (Kölbl &
Conard, 2003; Pacher, 2005). This is well
documented from the assemblages of the Ach
Valley. While ivory beads are still recovered
in the Gravettian, perforated teeth increase
in number and become more diversified at
Hohle Fels and Geißenklösterle. Carnivore
teeth include canines and incisors from
hyena, wolves, brown bear and cave bears
(both deciduous and permanent teeth)
(Figure 12). While the exact meaning of
these symbolic expressions remains open for
interpretation, carnivores played a significant
role in the cultures of the Aurignacian and
Gravettian peoples. In addition, the way in
which carnivores were employed in the
cultural repertoire shifts from the use of
carnivores as motifs to the use of carnivore
remains themselves for ornaments. We can
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Exploring Cave Use and Exploitation in the Swabian Jura
Table 4. Number of anthropogenic modification on carnivore remains across sites over the Middle
Paleolithic, Aurignacian and Gravettian (Note: numbers in parentheses indicate tools, organic debitages
and tooth pendants out of the total number of modified remains).
Geißenklösterle
Canis lupus
Ursus arctos
Vulpes/Alopex
Ursus spelaeus
Middle Paleolithic
1
1
6
Aurignacian
1
Gravettian
1
2(2)
10 (1)
7
17 (2)
indet. Carnivore
1
8
13
26
4 (4)
5
1 (1)
Hohle Fels
Panthera leo spelaea
Crocuta crocuta spelaea
Canis lupus
1
4
8 (5)
1
3 (1)
5 (4)
3 (2)
5
40 (3)
164 (14)
6
54
184
Ursus arctos
Vulpes/Alopex
Ursus spelaeus
Hohlenstein-Stadel
Vulpes/Alopex
4 (4)
Ursus spelaeus
Ursus sp.
1
1
6
Bockstein
Crocuta crocuta spelaea
Ursus spelaeus
1
2
2 (2)
3
2
Vogelherd
Panthera leo spelaea
1 (1)
1 (1)
Ursus arctos
Vulpes/Alopex
1
3
454
Kitagawa et al.
thus infer that there was a change in the
iconography and cultural system, which was
manifested through motifs and artifacts from
carnivore remains over the Aurignacian to
the Gravettian in the Swabian Jura.
Figure 6. Canine of a juvenile cave bear from
the Gravettian in Hohle Fels with cut marks.
The cut mark on ca. one year old individual
indicates hunting during hibernation, since cubs
were born during winter time (Photo: H. Jensen
©University of Tübingen).
Conclusion
The faunal record of the sites in the Ach and
Lone Valleys in the Swabian Jura reveals a
mixed signal of continuity and change
(Münzel & Conard, 2004a). On the one hand,
the proportion of species abundance at each site
is fairly consistent throughout the Paleolithic
sequence even after the arrival of modern
humans. The cave bears continued to inhabit
the area for shelter until the end of the
Gravettian and the diversity of species does
not change over time, documenting no
evidence of resource intensification or large
turnover in the faunal community. On the
other hand, there is a clear decrease in the
large/middle sized carnivores and an increase
of small carnivores over time. Further, the
frequency of anthropogenic and carnivore
modification demonstrates diminishing role
of carnivore in the accumulation of fauna in
the caves diachronically, which coincides
with the intensified use of the sites by humans,
reflecting clear anthropogenic signal in the
faunal assemblages (Conard 2011; Conard
et al., 2012). This trend is most apparent
between the Middle Paleolithic and the
Aurignacian period.
Many faunal assemblages from the
Middle and early Upper Paleolithic are
characterized by variability and document
no significant shifts in the subsistence practices
that reflect unique and ‘modern’ behavior of
Upper Paleolithic humans (Adler et al.,
2006; Grayson & Delpech, 2003; Hoffecker,
2009; Morin, 2008; Stiner et al., 2000).
Differences in the settlement patterns and
population density between Neanderthals
and modern humans in the Swabian Jura
have been argued previously (Conard et al.,
2006; Conard, 2011; Conard et al., 2012)
and greater presence of humans in caves of
the Swabian Jura is supported by these data.
Such a hypothesis has been previously
suggested by Kurtén (1976) and applied by
Grayson and Delpech’s study (2003) with
respect to cave bears and their extinction.
Thus, studying faunal assemblages from the
region has demonstrated likely changes in the
demography of archaic and modern humans
as well as their ecological relationship with
the carnivores and cave bears that occupied
the Swabian Jura.
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Exploring Cave Use and Exploitation in the Swabian Jura
Figure 7a. Cut marks on a bear rib from the
Aurignacian in Hohlenstein-Stadel (Photo: K.
Kitagawa).
Figure 7b. Close-up view of the cut marks on a bear
rib from the Aurignacian in Hohlenstein-Stadel
(Photo: K. Kitagawa).
Figure 8. Retoucher made from a canine of
Panthera leo spelaea from the Aurignacian in
Hohle Fels (Photo: S. Münzel).
Figure 9 (right). Tibia of Panthera leo spelaea
from the Aurignacian/Gravettian transitional layers
in Hohle Fels with cut marks (close-up view)
(Photo: S. Münzel).
456
Kitagawa et al.
Figure 10. Ivory sculpture of a lion
from the Aurignacian in Vogelherd
(Photo: Hilde Jensen ©University of
Tübingen).
Figure 11. Ivory sculpture of the ‘Löwenmensch’, a
therianthropic creature with human and lion traits from the
Aurignacian in Hohlenstein-Stadel (Photo: Thomas Stephan
©Ulmer Museum).
Figure 12. Tooth pendant of a milk canine
of a cave bear from the Gravettian in
Geißenklösterle (Photo: Hilde Jensen
©University of Tübingen).
457
Exploring Cave Use and Exploitation in the Swabian Jura
The Gravettian assemblages indicate
increased and diversified exploitation of
carnivores by humans. The evidence is well
documented in the Ach Valley, where a large
number of cave bears were clearly exploited for
subsistence and skin. In addition, carnivores
had cultural significance as their remains
were used for pendants and served as motifs in
figurines. Carnivore remains with anthropogenic
modifications provide evidence for the
changing relationship between carnivores and
hominins from that of competition to
predation and active exploitation. Although
opportunistic in nature, we begin to see that
animals such as non-human predators and
cave bears became incorporated in the
economic practices of Paleolithic people.
Carnivore remains in many
zooarchaeological assemblages often contribute
to our understanding of taphonomic history
and the processes affecting site formation.
The occurrences of carnivores and cave
bears on a regional scale also provide insight
into the changing use of caves and population
dynamics of animals on the Paleolithic
landscape. The sites of the Swabian Jura
further demonstrate the diversified exploitation
of animals that included the use of carnivores
and cave bears for cultural and economic
purposes. Faunal assemblages from caves
which include carnivores can serve as an
opportune case study to explore hominin
interaction with animals that go beyond a
predator-prey relationship.
Acknowledgments
We thank all of the organizations that have
funded this work including the Deutsche
Forschungsgemeinschaft, the Heidelberger
Akademie der Wissenschaften, the Universität
Tübingen, Heidelberger Cement, the Voith
Corporation, Alb-Donau-Kreis, Kreis
Heidenheim, the Gesellschaft für Urgeschichte,
the Museumsgesellschaft Schelklingen, the
Landesamt für Denkmalpflege BadenWürttemberg, the cities of Blaubeuren and
Niederstotzingen, and the Förderverein
Eiszeitkunst im Lonetal. Further, we thank
the organizers of the symposium in Salou
and the editors of this volume. We would
also like to thank our colleages in Institut
für Naturwissenschaftliche Archäologie, and
Institut für Ur- und Frühgeschichte und
Archäologie des Mittelalters for their
continual support and help.
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