JQI=526=Shantha=Venkatachala=BG

Quaternary International 75 (2001) 29}40

The late Middle Paleolithic and earliest Upper Paleolithic in Central

Europe and their relevance for the Out of Africa hypothesis
Michael Bolus*, Nicholas J. Conard
Abteilung Altere Urgeschichte und Quarta( ro( kologie, Institut fu( r Ur- und Fru( hgeschichte und Archa( ologie des Mittelalters der Universita( t Tu( bingen,
Schloss Hohentu( bingen, Burgsteige 11, D-72070 Tu( bingen, Germany

Abstract
The available data from Central Europe is consistent with the hypothesis that Homo sapiens sapiens evolved initially outside Europe
and colonized the di!erent regions of the continent previously occupied by Neanderthals between roughly 50 and 30 kyr BP. Di!erent
transitional industries in Central Europe, which show both Middle and Upper Paleolithic features, appear to root in the local Middle
Paleolithic assemblages. At present it is uncertain which hominids produced the transitional industries. While organic artifacts,
objects of personal ornamentation, and art objects are generally absent in these transitional assemblages, they become abundant in
the Aurignacian, which represents the "rst fully evolved Upper Paleolithic technocomplex spread over larger parts of Europe. Both
radiocarbon and TL dates indicate that the Aurignacian was present in Central Europe, including Germany, as early as about 40 kyr
ago. This means that coexistence of Neanderthals and anatomically modern hominids endured for a time span of some 10 kyr, since
with the appearance of the Aurignacian we "rst encounter the cultural remains of Homo sapiens sapiens with fully developed artistic
and symbolic representations, ornaments, and abundant organic tools, while the last Neanderthals appear to have lived some 30 kyr
ago.  2001 Published by Elsevier Science Ltd. All rights reserved.

1. Introduction
While it is generally accepted that the earliest hominids have their roots in Africa and from there colonized
the low and middle latitudes of the Old World, no consensus exists regarding the evolution and spread of anatomically modern hominids. Scholars continue to debate
whether this evolution took place simultaneously on
di!erent continents or whether Homo sapiens sapiens,
after evolving initially in Africa, replaced existing archaic
populations including the Neanderthals in western
Eurasia (Mellars and Stringer, 1989; Trinkaus, 1989;
BraK uer and Smith, 1992).
This paper cannot present the full story of the
transition from Middle to Upper Paleolithic for the
whole of Central Europe. The cultural manifestations
during the period in question * approximately between
50,000 and 30,000 BP * are too diverse and uncertain in
their interrelations. By focusing on key aspects of the
archaeological record in Germany as well as in Eastern
and Southeastern Central Europe, we can provide
evidence relevant to the Out of Africa hypothesis, and

* Corresponding author.

conclude that this hypothesis is supported by the

archaeological data.

2. The archaeological record in Eastern and Southeastern

Central Europe
In Eastern Central Europe, especially in Moravia,
there are two main transitional industries. The
Bohunician and the Szeletian both root in the local
Middle Paleolithic. The Bohunician has its geographic
center in the region of Brno. The radiocarbon dates for
the important sites of Brno-Bohunice and StraH nska skaH la
(Fig. 1: 7 and 9) lie between 36 and 43 kyr BP and show
relatively high statistical uncertainties ranging from 1.1
to 3.1 kyr (Svoboda et al., 1996: Table 5.2). Noteworthy,
is the abundance of Levallois elements in these lithic
assemblages. This characteristic has led Svoboda et al.
(1996, p. 108) to postulate the existence of cultural a$nities with the Near East and northern and central Asia.
The pattern of lithic reduction in the Bohunician combines Upper Paleolithic concepts such as the reduction
along the edges of cores with the use of the Levallois
concept. An abundance of blades ranges from 20 to 45%
of the larger debitage. The Bohunician appears to have

1040-6182/01/$20.00  2001 Published by Elsevier Science Ltd. All rights reserved.

PII: S 1 0 4 0 - 6 1 8 2 ( 0 0 ) 0 0 0 7 5 - 6

Jqi=526=Shantha=VVC
30

M. Bolus, N.J. Conard / Quaternary International 75 (2001) 29}40

Fig. 1. Major sites with Early Upper Paleolithic industries in Central Europe mentioned in the text. 1, Schwalbenberg; 2, Gei{enkloK sterle and
Sirgenstein (Ach Valley); 3, Vogelherd, Hohlenstein, and Bockstein (Lone Valley); 4, Keilberg-Kirche; 5, Willendorf; 6, Galgenberg-Stratzing;
7, Brno-Bohunice; 8, Vedrovice; 9, StraH nska skaH la; 10, Mladec; 11, Vindija; 12, Velika Pecina; 13, Temnata and 14, Bacho Kiro.

developed out of the Mousterian and would thus seem to

have been produced by Neanderthals.
The Szeletian, in contrast to the Bohunician, shows
a broader distribution with a far greater number of sites.
Karel Valoch has argued that the Szeletian may have
evolved from earlier central European Micoquian and
Keilmesser industries as typi"ed by the "nds from Kulna
7 and 6a (Valoch, 1993, pp. 57}67). Szeletian assemblages
lack Levallois cores and debitage, but bifacial forms
including leaf points are abundant. Laminar debitage is
less common than in Bohunician assemblages. The bestdated site is Vedrovice V (Fig. 1: 8), which has been
radiocarbon dated to between 35.15$0.65 and 39.5$
1.1 kyr BP (Svoboda et al., 1996, Table 5.3).
Both of these assemblage types include a tool spectrum
that is characterized by both Middle and Upper Paleolithic elements. The Bohunician is particularly dominated by Upper Paleolithic forms, including end
scrapers, carinated scrapers, and burins. Less abundant
within the Bohunician are Middle Paleolithic forms including scrapers and denticulates, and Levallois points.
Although leaf points or Blattspitzen are common in the
Szeletian, they occur only rarely in the Bohunician. Organic artifacts, which are often seen as characteristic of
the Upper Paleolithic, including bone, antler and ivory
tools, ornaments, and artworks are entirely unknown in
both the Bohunician and the Szeletian. While no human
skeletal material has been found in unambiguous association with these assemblages, it is at least plausible that
these assemblages were manufactured by Neanderthals.

Turning to the Aurignacian, the densest concentration

of sites from this period outside France is found in
Moravia, where numerous surface sites are well
documented, but remain undated. In Lower Austria,
strati"ed open-air sites are well known from Willendorf
II, Galgenberg-Stratzing (Fig. 1: 5 and 6), KremsHundssteig, and Langmannersdorf (Hahn, 1977, pp. 104}
109; Neugebauer-Maresch, 1996). Based on the available
chronostratigraphic data, it appears that the Aurignacian
in these parts of Central Europe covers a broad temporal
span between ca. 38 kyr radiocarbon years at Willendorf
II, Kulturschicht 3 (Haesaerts et al., 1996), to around
29 kyr at Milovice (Svoboda et al., 1996, Table 5.4).
Intermediate radiocarbon dates are known from sites
including Pod hradem at 33.3$1.1 kyr BP (Svoboda
et al., 1996, Table 5.4). These Aurignacian assemblages
are dominated by end scrapers, carinated and nosed end
scrapers, and burins; but side scrapers, notches, and denticulates are also present. Unidirectional blade cores usually dominate the assemblages (Hahn, 1977, p. 194) and,
as Bon (1996) has demonstrated for the Aurignacian of
the Grotte des Hye`nes near Brassempouy in France,
a number of the carinated scrapers may well have served
as blade cores. With the appearance of the Aurignacian
in Moravia, the systematic production of bone and antler
tools is documented, among which Mladec points
are noteworthy (Hahn, 1977, pp. 222}225). Additionally,
among others, ivory pendants and perforated teeth are
present and demonstrate the "rst use of ornaments in the
region. Both in terms of the technology and typology of

Jqi=526=SC=VVC
M. Bolus, N.J. Conard / Quaternary International 75 (2001) 29}40

the lithic artifacts and with the presence of numerous

organic artifacts, the Aurignacian seems to appear relatively suddenly in this part of Central Europe, and suggests a cultural development that arises independently of
the earlier Bohunician and Szeletian assemblage types.
When considering the connection between hominid
skeletal morphology and artifact assemblages, the important human fossils from Mladec (Fig. 1: 10) are of only
marginal help since they cannot be attributed with certainty to a particular assemblage type. The reexamination of these fossils by BraK uer and Broeg (1998) did not
identify any evidence for hybridization, as had been argued earlier (e.g. Frayer, 1986, 1992) and suggests that
they can be classi"ed as Homo sapiens sapiens.
In southeastern Europe, assemblages that belong to
the period of the transition from the Middle to the Upper
Paleolithic are also known. These assemblages can be
seen as analogous to the Bohunician in Moravia. In
Bulgaria, Temnata Cave (Fig. 1: 13) provides a transitional industry of Layer VI, Sector TD-II, radiocarbon dated to at least 38.7 kyr (Ginter et al., 1996,
Table 1). The early Aurignacian from a di!erent part of
the same cave has yielded AMS radiocarbon ages in
excess of 40 kyr. The transitional assemblage from Temnata is characterized by a temporal shift from Levallois
cores toward bi-directional Upper Paleolithic cores.
Ginter and colleagues see this assemblage as a continuation of the Moustero-Levalloisian tradition (Ginter et al.,
1996, p. 190). Noteworthy, is an engraved stone from the
transitional assemblage that represents one of the very
few "nds that could perhaps be seen as an indication that
artworks are present before the Aurignacian (CreH mades
et al., 1995). The earliest Aurignacian from Temnata has
to be referred to as the Early Balkan Aurignacian or the
Bachokirian (Ginter et al., 1996, pp. 195}198).
Also in Bulgaria, the cave of Bacho Kiro (Fig. 1: 14)
has played a central role in the discourse about the
origins of the Aurignacian and modern humans in
Europe. Here, Cultural Horizon I of Level 11 has yielded
an early Upper Paleolithic assemblage, which Kozlowski
and colleagues have referred to as the Bachokirian or as
the Early Balkan Aurignacian (Kozlowski, 1982). This
horizon was originally dated to in excess of 43 kyr with
conventional radiocarbon in Groningen (Mook, 1982).
Newer AMS dates from Oxford for Level 11/11a are
younger, with ages of 38.5$1.1, 37.7$1.5 and
34.8$1.2 and 33.8$0.9 kyr BP (Hedges et al., 1994,
pp. 347}348). A clear explanation for these diverse dates
is not yet available. This assemblage lacks bone points
and other organic artifacts of the Aurignacian, and both
side-scrapers and Levallois points are very rare. The
blade production is of Upper Paleolithic character.
Kozlowski and colleagues see no connection between the
Bachokirian assemblage and lower lying Middle Paleolithic assemblages, and emphasize the Upper Paleolithic
nature of this industry (Kozlowski, 1982, p. 163).

31

In Croatia and Slovenia there seems to be a relatively

continuous cultural development between 45 and 30 kyr
BP from late Middle Paleolithic to early Upper Paleolithic industries which in most cases cannot be classi"ed
as Aurignacian (Karavanic and Smith, 1998).
The exact stratigraphic positions and cultural attributions, as well as the dates of the hominid remains from
Southwestern Central Europe belonging to this period,
have been questioned (Mellars et al., 1999; Zilha o and
d'Errico, 1999a, b). Among others, this is true for the
Neanderthals from the early Upper Paleolithic level G1
of Vindija cave in Croatia (Fig. 1: 11) which yielded AMS
radiocarbon ages of 33$0.4 kyr BP (Karavanic and
Smith, 1998, Fig. 2) and 28}29 kyr BP (Smith et al., 1999)
respectively. These results suggest the presence of Neanderthals in the region signi"cantly after the appearance of
Aurignacian assemblages and modern Homo sapiens in
other parts of Central Europe. The lithic artifacts from
Vindija are dominated by quartz, with lesser amounts of
graywacke, chert, and radiolarite. The abundance of raw
materials with irregular chipping characteristics contributes to the non-standardized nature of the Vindija lithic
assemblages. Both Middle and Upper Paleolithic tools
are present. The stratigraphic integrity of the site
has been questioned (see Karavanic and Smith, 1998,
pp. 233}236; Zilha o and d'Errico, 1999b, pp. 355}357)
and given the rich bone assemblages from the overlying
layers, there is still uncertainty about whether or not the
Neanderthal remains from Layer G1 are associated with
the worked bone industry.
Apart from the cranial fragment of Homo sapiens
sapiens from Layer j at Velika Pecina cave (Fig. 1: 12;
Malez, 1974, pp. 8}13; Karavanic and Smith, 1998,
pp. 239}241), this deposit yielded only one stone tool.
The lower lying Layer k contained what appears to be
a Middle Paleolithic assemblage. The overlying Layer
i yielded only seven lithic tools, including both Middle
and Upper Paleolithic forms. The latter deposit also
produced three bone points, which have led to the
assumption that this layer should be attributed to
the Upper Paleolithic (Karavanic and Smith, 1998,
pp. 238}239).
It must be stressed that all the dates mentioned so far
are radiocarbon dates, many of them lying outside the
reliable range of the method.

3. The archaeological record in Germany

In Germany, the entirety of the Middle Paleolithic
appears to have been accompanied by the evolution of
Neanderthals out of earlier Homo heidelbergensis populations. The fossil hominids start with Mauer (Wagner and
Beinhauer, 1997), include those from Bilzingsleben
(Mania et al., 1980), Steinheim (Adam, 1988), and
Reilingen (Czarnetzki, 1989, 1991; Condemi, 1996), and

Jqi=526=Shantha=VVC
32

M. Bolus, N.J. Conard / Quaternary International 75 (2001) 29}40

Fig. 2. Schwalbenberg near Remagen, Rhineland/Germany. Stone artifacts. 1}4, side scrapers; 5, 8, splintered pieces; 6, end scraper; 7, 9, pre-forms of
bifacially retouched tools; 10, discoidal core and 11, blade core (modi"ed after App et al., 1995).

"nally reach classical Neanderthals like those from Neandertal itself (Krings et al., 1997). Recent "nds from
Ochtendung, dated to the penultimate glaciation (Berg,
1997; Condemi, 1997), and Warendorf-Neuwarendorf
(Czarnetzki, in press) "t well into this lineage, which
seems to represent a continuous line of evolution. However, there is no direct link between the Neanderthals and

early Homo sapiens sapiens fossils such as the one from

Vogelherd cave (Riek, 1932).
This conclusion is highlighted both by DNA analyses
conducted by a TuK bingen team comparing the Stetten
I fossil from Vogelherd cave with Neanderthals like
Warendorf and Krapina (Scholz et al., 1999; Pusch, pers.
comm.), and by DNA analyses conducted by a team in

Jqi=526=SC=VVC
M. Bolus, N.J. Conard / Quaternary International 75 (2001) 29}40

Munich, using the type specimen itself (Krings et al.,

1997). In both cases the data support the hypothesis that
Neanderthals were not direct ancestors of anatomically
modern hominids.
The latest Middle Paleolithic of Germany is characterized by Blattspitzen assemblages which are, however, less
numerous than in Eastern Central Europe (Bolus and
RuK ck, 2000; Conard and Fischer, in press). These
German assemblages with Blattspitzen do not appear to
have evolved out of the local Middle Paleolithic. Autochthonous transitional industries without or nearly without
Blattspitzen comparable to the Bohunician seem to be
absent in Germany, perhaps with the exception of the
small assemblage from Schwalbenberg in the Rhineland
(Fig. 1: 1), excavated by Joachim Hahn (App et al., 1995).
This site yielded both a discoidal core (Fig. 2: 10) and
a blade core (Fig. 2: 11); the number of blades is small.
Amongst the tools, side-scrapers (Fig. 2: 1}4) predominate, but there is one end-scraper (Fig. 2: 6) as well;
splintered pieces (Fig. 2: 5 and 8) are relatively numerous.
It is noteworthy that splintered pieces also frequently
appear in Blattspitzen assemblages (e.g., Zeitlarn: Heinen
and Beck, 1997, and Vedrovice V: Valoch, 1993). Another
feature linking the Schwalbenberg assemblage with
Blattspitzen assemblages is the morphology of the
end-scraper with heavily retouched edges (Fig. 2: 6)
which was produced out of a thick #ake rather than out
of a blade. Thus, it can be stated that not a single genuine
Upper Paleolithic tool type can be found within the
Schwalbenberg assemblage. Two pre-forms of bifacially
retouched artifacts (Fig. 2: 7 and 9) also provide links to
assemblages with Blattspitzen such as the Szeletian. In
contrast to the Middle Paleolithic tool types and blank
production, the patterns of raw material procurement
resemble those of Upper Paleolithic sites in the Middle
Rhine region. About 60% of the raw material used at the
Schwalbenberg site was obtained from within a distance
of up to 85 km (Floss, 1994, pp. 171}174) while for other
Middle Paleolithic sites of that region the amount of raw
materials from a greater distance is normally less than
3%. This value increases only slightly during the Aurignacian and rises with the Gravettian, reaching its maximum value during the Magdalenian (Floss, 1994).
One of the key sites in southern Germany is
Gei{enkloK sterle cave (Fig. 1: 2) in the Ach Valley near
Blaubeuren. As in most other sites of the region, there is
no continuous stratigraphic sequence from the Middle to
the Upper Paleolithic. Above a Middle Paleolithic horizon, dated by ESR to a mean value of 43.3$4 kyr BP
(Richter et al., 2000), which has, however, so far been
excavated for only a few square meters, there is an archaeologically sterile layer, followed by an Upper Paleolithic sequence. This sequence starts with Aurignacian,
which Joachim Hahn subdivided into two horizons
(Archaeological Horizons III and II: Hahn, 1988) and
is overlain by the Gravettian and Magdalenian.

33

In contrast to Hahn, who addressed the lowest Upper

Paleolithic horizon (Archaeological Horizon III) as
&Protoaurignacien' we refer to this horizon as the Lower
Aurignacian. Unidirectional blade production (Fig. 3: 16)
is well represented in both horizons (Hahn, 1988, pp.
144}147), and we "nd almost the same tool types. There
are, however, more carinated and nosed end scrapers in
the lower horizon (Fig. 3: 1}7) while in the upper (Archaeological Horizon II) there are more simple end scrapers
on blades (Fig. 4: 5, 7 and 9). Burins, especially burins on
truncation, are represented in almost equal numbers in
both horizons (Fig. 3: 8, 11 and 12; Fig. 4: 10). The upper
horizon in turn yielded more blades with lateral retouch
(Fig. 4: 8) and pointed blades (Fig. 4: 19), as well as more
splintered pieces (Fig. 3: 9 and 10; Fig. 4: 13, 14 and 17).
There are only a few Dufour bladelets (Fig. 4: 15) in the
Upper Aurignacian, and they are lacking in the Lower
Aurignacian. Middle Paleolithic type tools are rare in
both horizons.
The Lower Aurignacian yielded two perforated teeth
(Fig. 3: 17 and 18), two ivory pendants (Fig. 3: 14 and 15),
and a pre-form of a bone bead (Fig. 3: 13), while tools
made of bone, antler and ivory appear in considerable
numbers (Fig. 3: 19). Only art objects are absent. They
are in turn present in the Upper Aurignacian (Fig. 6: 13,
15, 16 and 18), representing some of the earliest art
objects known. In the Upper Aurignacian, artifacts made
of organic materials (Fig. 5: 5, 6 and 12) including bone
#utes (Fig. 6: 19), as well as objects of personal ornamentation (Fig. 6: 1}5) are abundant.
While Zilha o and d'Errico * who have not studied the
originals * reject the term Aurignacian or even Protoaurignacian for the lower horizon, which they tend to
label just as non-diagnostic early Upper Paleolithic
(Zilha o and d'Errico, 1999a), we do not hesitate to con"rm Hahn's assessment of the Aurignacian character of
the Level III assemblage. The lack of some tool types in
the Level III assemblage may well be due to the functional context. An analysis of the faunal remains conducted by MuK nzel shows that in the Lower Aurignacian
there are numerous hints for working of organic materials, especially ivory, and the manufacture of organic
tools, even though the number of tools themselves is not
very high (MuK nzel, 1999). In 1977, Hahn already stressed
that di!erences in the frequency of special tool types as
well as the occurrence or non-occurrence of tool types
may be due to functional variability rather than to chronological di!erences (Hahn, 1977, pp. 294}296).
Using the TL signal from burnt #ints, Daniel Richter
dated the Lower Aurignacian of Gei{enkloK sterle cave at
an average of 40.2$1.5 kyr BP, while the Upper Aurignacian produced TL ages at about 37 kyr BP Radiocarbon AMS dates of bone samples from both horizons are
somewhat younger: for Level III a mean age of
38.4$0.85 kyr BP was obtained, for Level II a mean age
of 33.5$0.35 kyr BP (Richter et al., 2000).

Jqi=526=Shantha=VVC
34

M. Bolus, N.J. Conard / Quaternary International 75 (2001) 29}40

Fig. 3. Gei{enkloK sterle Cave, Southern Germany. Lower Aurignacian}Archaeological Horizon III. 1}3, carinated end scrapers; 4}7, nosed end
scrapers; 8, 11, 12, burins; 9, 10, splintered pieces; 13, pre-form of a bone bead; 14, 15, ivory pendants; 16, blade core with re"tted blades; 17, 18,
perforated teeth and 19, ivory projectile point (modi"ed after Hahn, 1988).

Only one other German Aurignacian assemblage has

been dated this early: the one from Keilberg-Kirche near
Regensburg in Bavaria (Fig. 1: 4), which yielded conventional charcoal dates ranging between 37.5$1.45 and
38.6$1.2 kyr BP (Uthmeier, 1996).
In addition to Gei{enkloK sterle, Aurignacian art objects
and objects of personal ornamentation are well known
from the Lone Valley (Fig. 1: 3), at the cave sites of

Vogelherd (Fig. 6: 14 and 17; Riek, 1934; Hahn, 1977, pp.

87}91), Hohlenstein-Stadel (Fig. 6: 7 and 8; Hahn, 1977,
pp. 86}87), and Bockstein (Fig. 6: 9 and 12; Hahn, 1977,
pp. 82}85), excavated by Gustav Riek and Robert Wetzel
in the 1930s, and from the Ach Valley (Fig. 1: 2) at the
cave site of Sirgenstein (Fig. 6: 6), excavated by Robert
Rudolf Schmidt in 1906 (Hahn, 1977, pp. 92}94). In these
caves the ornaments and art objects were associated with

Jqi=526=SC=VVC
M. Bolus, N.J. Conard / Quaternary International 75 (2001) 29}40

35

Fig. 4. Stone artifacts from the Upper Aurignacian/Archaeological Horizon II of Gei{enkloK sterle Cave (7}10, 13}15, 17, 19) and artifacts from
comparable cave sites in Southwestern Germany: Sirgenstein IV (1, 4), Sirgenstein V (2), Vogelherd V (3, 5, 6, 16, 18), and Bockstein-ToK rle VII (11, 12).
1}3, carinated end scrapers; 4, 6, nosed end scrapers; 5, 9, end scrapers; 7, burin combined with end scraper; 8, blade with retouched edges; 10}12, 16,
burins; 13, 14, 17, splintered pieces; 15, Dufour bladelet; 18, 19, pointed blades (modi"ed after Hahn, 1977, 1988).

assemblages corresponding to the Upper Aurignacian of

Gei{enkloK sterle (Fig. 4: 1}6, 11,12, 16 and 18; Fig. 5: 1}4,
7}11).
The dates for the assemblages from Vogelherd and
Stadel caves range from about 30}32 kyr BP, suggesting
that they postdate the Upper Aurignacian from Gei{en-

kloK sterle. It must be stressed, however, that in contrast to

the other four sites mentioned, Gei{enkloK sterle cave has
been recently excavated with modern standards of documentation and a more systematic sampling, allowing the
TL and C dates from Gei{enkloK sterle to be regarded as
much more reliable. We are just getting new dates for the

Jqi=526=Shantha=VVC
36

M. Bolus, N.J. Conard / Quaternary International 75 (2001) 29}40

Fig. 5. Organic artifacts from the Upper Aurignacian/Archaeological Horizon II of Gei{enkloK sterle Cave (5, 6, 12) and artifacts from comparable cave
sites in Southwestern Germany: Vogelherd V (1}4, 7, 9}11) and BocksteinhoK hle (8). 1}6, 8, projectile points with split base; 7, cave bear tooth used as
retoucher; 9, 10, bone points with massive base; 11, smoothing tool; 12, ba( ton perceH (modi"ed after Hahn, 1977, 1988).

other sites, using bones de"nitely worked by man, and

this will give us the possibility of checking the dates
obtained earlier.
While early Upper Paleolithic hominid remains are
absent in Gei{enkloK sterle and only a single tooth was
recovered from the Stadel Aurignacian, Vogelherd cave
yielded the very well-preserved specimen Stetten I
(Fig. 7), which was found at the base of the Aurignacian
sequence (Fig. 8) and thus represents one of the earliest
"nds of Homo sapiens sapiens in Europe. This demonstrates that anatomically modern hominids are respon-

sible for the fully developed Aurignacian, while no fossil

evidence is at present available for the earliest
Aurignacian from Gei{enkloK sterle.

4. Conclusions
Let us now look at the archaeological evidence with
regard to the Out of Africa hypothesis. Neither in Eastern or Southeastern Central Europe, nor in Germany can
an autochthonous evolution of anatomically modern hu-

Jqi=526=SC=VVC
M. Bolus, N.J. Conard / Quaternary International 75 (2001) 29}40

37

Fig. 6. Ornamental objects, art objects, and a musical instrument from the Upper Aurignacian/Archaeological Horizon II of Gei{enkloK sterle Cave
(1}5, 13, 15, 16, 18, 19) and from comparable cave sites in Southwestern Germany: Sirgenstein IV (6), Hohlenstein-Stadel VI}V (7, 8), Bockstein-ToK rle
IV}VI (9}12), and Vogelherd V (14, 17). 1}6, pendants; 7, 8, perforated teeth; 9}11, stone beads; 12, stone pendant; 13}16, ivory "gurines; 17, 18,
ornamented objects and 19, bone #ute (modi"ed after Hahn, 1977, 1986, 1988; Hahn and MuK nzel, 1995).

mans from the existing Neanderthal populations be demonstrated. An in situ development of Upper Paleolithic
features out of the local Middle Paleolithic industries
appears to be possible regarding the archaeological record in Central Europe. To what extent processes of

acculturation took place and in which direction a presumed acculturation was oriented remains subject to
controversial discussions. In any case, it is remarkable
that essential features of fully developed Upper Paleolithic assemblages are lacking in the very places where

Jqi=526=Shantha=VVC
38

M. Bolus, N.J. Conard / Quaternary International 75 (2001) 29}40

Fig. 7. Vogelherd Cave, Southwestern Germany. Homo sapiens sapiens

specimen &Stetten I' found at the base of the Aurignacian sequence/Layer V (modi"ed after Czarnetzki, 1983; Fig. 132).

objects as well as art objects, had been initiated by

contacts between Neanderthals and anatomically modern hominids. This seems especially true for Western
Europe, but may also hold for Southwestern Germany.
In this context, the western European Cha( telperronian
should be mentioned. On the basis of lithic technology
and various organic artifacts as well as objects of personal ornamentation, this appears to be Upper Paleolithic, but was obviously manufactured by Neanderthals,
as Neanderthal remains associated with Cha( telperronian
in Arcy-sur-Cure and St. CeH saire, both in France, reveal
(Ba$er, 1999). At present, it cannot be "nally stated
whether the Cha( telperronian is a genuine Neanderthal
creation (Zilha o and d'Errico, 1999a, b), or if it is rather
the result of acculturation processes.
Due to gaps in the fossil record, it remains uncertain
who was responsible for the transitional industries and
for the earliest Upper Paleolithic assemblages in Central
Europe. It should be kept in mind that between 40 and
30 kyr BP there seems to have been a coexistence of
Neanderthals and anatomically modern hominids in
Europe. Examples of very late Neanderthals, for instance,
come from Zafarraya and CariguK ela in Spain, Columbeira, Figueira Brava, and Salemas in Portugal (Straus,
1996), and Vindija Layer G, in Croatia (Karavanic and
Smith, 1998, pp. 239}241) while early Homo sapiens
sapiens specimens have been found in Vogelherd in
Southwestern Germany (Riek, 1932).
In summary, it can be stated that the archaeological
record in Central Europe, including Germany, appears to
support the replacement model rather than the
multiregional model, thus providing evidence that Homo
sapiens sapiens originated outside Europe.

Acknowledgements
We would like to thank Jean-Michel Geneste
(Bordeaux/France), Paul Haesaerts (Brussels/Belgium),
Jakov Radovcic (Zagreb/Croatia), Jack Rink (Hamilton/
Canada), Fred H. Smith (DeKalb, Illinois/USA), and
Nicolas Teyssandier (Paris-Nanterre/France) for providing helpful information.

Fig. 8. Vogelherd Cave, Southwestern Germany. Stratigraphic sequence at the southwestern entrance (modi"ed after Riek, 1934). I,
Holocene "nds; II, Magdalenian; III, Magdalenian; IV, Aurignacian;
V, Aurignacian; VI, "nd horizon of the specimen &Stetten I' and VII,
Middle Paleolithic.

early anatomically modern hominids are known in Africa

and the Near East. It may be discussed if this variety of
features, associated with `cultural modernitya including
a broad spectrum of artifacts made of bone, antler and
ivory and a pronounced production of varied decorative

References
Adam, K.D., 1988. Der Urmensch von Steinheim an der Murr und seine
Umwelt. Ein Lebensbild aus der Zeit vor einer viertel Million
Jahren. Jahrbuch des RGZM 35, 3}23.
App, V., Au!ermann, B., Hahn, J., Pasda, C., Stephan, E., 1995. Die
altsteinzeitliche Fundstelle auf dem Schwalbenberg bei Remagen.
(With contributions by M. Baales, E. Bibus, W. RaK hle, R. RottlaK nder, W. Schoch, and K.-H. Steppan). Berichte zur ArchaK ologie an
Mittelrhein und Mosel 4, 11}136.
Ba$er, D., 1999. Les derniers NeH andertaliens. Le Cha( telperronien,
Paris.

Jqi=526=SC=VVC
M. Bolus, N.J. Conard / Quaternary International 75 (2001) 29}40
Berg, A.V., 1997. Die SchaK delkalotte eines Neandertalers aus dem
Wannenvulkan bei Ochtendung, Kreis Mayen-Koblenz. Berichte
zur ArchaK ologie an Mittelrhein und Mosel 5, 11}22.
Bolus, M., RuK ck, O., 2000. Eine Blattspitze aus Wittislingen, Lkr.
Dillingen a. d. Donau (Bayern). Zur suK dwestlichen Verbreitungsgrenze spaK tmittelpalaK olithischer Blattspitzeninventare. ArchaK ologisches Korrespondenzblatt 30, 1}8.
Bon, F., 1996. L'industrie lithique aurignacienne de la couche 2A de la
grotte des Hye`nes a` Brassempouy (Landes). In: Delporte, H.,
Clottes, J. (Eds.), PyreH neH es preH historiques * arts et socieH teH s, 118
Congre`s National des socieH teH s historiques et scienti"ques, Pau,
1993, pp. 439}455.
BraK uer, G., Broeg, H., 1998. On the degree of Neandertal * modern
continuity in the earliest Upper Palaeolithic crania from the Czech
Republic: evidence from non-metrical features. In: Omoto, K.,
Tobias, P.V. (Eds.), The Origins and Past of Modern Humans *
Towards Reconciliation. World Scienti"c, Singapore, pp. 106}125.
BraK uer, G., Smith, F.H. (Eds.), 1992. Continuity or Replacement. Controversies in Homo sapiens Evolution. Balkema, Rotterdam.
Conard, N.J., Fischer, B. Are there recognizable cultural entities in the
German Middle Paleolithic? In: Ronen, A., Weinstein-Evron, M.,
(Eds.), The Yabrudian and Micoquian in the Levant and in Europe.
BAR International Series, Oxford, in press.
Condemi, S., 1996. Does the human fossil specimen from Reilingen
(Germany) belong to the Homo erectus or to the Neanderthal
lineage? Anthropologie 34(1) (Brno), 69}77.
Condemi, S., 1997. Preliminary study of the Calotte of the Ochtendung
cranium. Berichte zur ArchaK ologie an Mittelrhein und Mosel 5,
23}28.
CreH mades, M., Laville, H., Sirakov, N., Kozlowski, J.K., 1995.
Une pierre graveH e de 50,000 ans BP dans les Balkans. PaleH o 7,
201}209.
Czarnetzki, A., 1983. Zur Entwicklung des Menschen in SuK dwestdeutschland. In: MuK ller-Beck, H. (Ed.), Urgeschichte in BadenWuK rttemberg. Stuttgart, pp. 217}240.
Czarnetzki, A., 1989. Ein archaischer Hominidencalvariarest aus einer
Kiesgrube in Reilingen, Rhein-Neckar-Kreis. QuartaK r 39/40,
191}201.
Czarnetzki, A., 1991. Nouvelle deH couverte d'un fragment de cra( ne d'un
hominideH archamK que dans le sud-ouest de l'Allemagne (rapport preH liminaire). L'Anthropologie 95, 103}111.
Czarnetzki, A. Das Fragment eines Os parietale des klassischen Neandertalers aus Warendorf-Neuwarendorf. Ausgrabungen und Funde
in Westfalen-Lippe 10, in press.
Floss, H., 1994. Rohmaterialversorgung im PalaK olithikum des Mittelrheingebietes. Monographein des RGZM Z1, Dr. R. Habert, Bonn.
Frayer, D.W., 1986. Cranial variation at Mladec and the relationship
between Mousterian and Upper Paleolithic hominids. In: Novotny,
V.V., MizerovaH , A. (Eds.), Fossil Man. New Facts}New Ideas.
Anthropos, Brno, 23, pp. 243}255.
Frayer, D.W., 1992. Evolution at the European edge: Neanderthal and
Upper Paleolithic relationships. PreH histoire EuropeH enne 2, 9}69.
Ginter, B., Kozlowski, J.K., Laville, H., Sirakov, N., Hedges, R.E.M.,
1996. Transition in the Balkans: news from the Temnata Cave,
Bulgaria. In: Carbonell, E., Vaquero, M. (Eds.), The Last Neandertals, the First Anatomically Modern Humans: a Tale About the
Human Diversity. Cultural Change and Human Evolution:
The Crisis At 40 ka BP. Universitat Rovira i Virgili, Tarragona,
pp. 169}200.
Haesaerts, P., Damblon, F., Bachner, M., Trnka, G., 1996. Revised
stratigraphy and chronology of the Willendorf II sequence, Lower
Austria. Archaeologia Austriaca 80, 25}42.
Hahn, J., 1977. Aurignacien, das aK ltere JungpalaK olithikum in Mittelund Osteuropa. KoK ln/Wien.
Hahn, J., 1986. Kraft und Aggression. Die Botschaft der Eiszeitkunst im
Aurignacien SuK ddeutschlands? Archaeologica Venatoria Band 7,
TuK bingen.

39

Hahn, J., 1988. Die Gei{enkloK sterle-HoK hle im Achtal bei Blaubeuren I.
Fundhorizontbildung und Besiedlung im MittelpalaK olithikum und
im Aurignacien. (With contributions by H. Gollnisch, A. Scheer, N.
Symens, R. Whallon, and J. Wei{haupt). Karl Theiss, Stuttgart.
Hahn, J., MuK nzel, S., 1995. Knochen#oK ten aus dem Aurignacien des
Gei{enkloK sterle bei Blaubeuren, Alb-Donau-Kreis. Fundberichte
aus Baden-WuK rttemberg 20, 1}12.
Hedges, R.E.M., Housley, R.A., Bronk Ramsey, C., van Klinken, G.J.,
1994. Radiocarbon dates from the Oxford AMS system: Archaeometry datelist 18. Archaeometry 36, 337}374.
Heinen, M., Beck, D., 1997. Ausgrabungen auf dem Szeletien-Fundplatz
Zeitlarn, Lkr. Regensburg. BeitraK ge zur ArchaK ologie in der Oberpfalz 1, 71}89.
Karavanic, I., Smith, F.H., 1998. The Middle/Upper Paleolithic interface and the relationship of Neanderthals and early modern humans
in the Hrvatsko Zagorje, Croatia. Journal of Human Evolution 34,
223}248.
Kozlowski, J.K. (Ed.), 1982. Excavation in the Bacho Kiro Cave (Bulgaria).
Final Report. Panstwowe Wydawnictwo Naukowe, Warszawa.
Krings, M., Stone, A., Schmitz, R.W., Krainitzki, H., Stoneking, M.,
PaK aK bo, S., 1997. Neandertal DNA sequences and the origin of
modern humans. Cell 90, 19}30.
Malez, M., 1974. Noviji rezultati istraz\ ivanja paleolitika u Velikoj
Pecini, Veternici i S[ andalji. Neue Ergebnisse der PalaK olithikumForschungen in Velika Pecina, Veternica und S[ andalja (Kroatien).
Arheolocki radovi i rasprave 7, 7}44.
Mania, D., Toepfer, V., Vlcek, E., 1980. Bilzingsleben I. Homo erectus
* seine Kultur und seine Umwelt. Berlin.
Mellars, P., Otte, M., Straus, L.G., Zilha o, J., d'Errico, F., 1999. The
Neanderthal Problem Continued. Current Anthropology 40, 341}364.
Mellars, P., Stringer, C. (Eds.), 1989. The Human Revolution. Behavioural and Biological Perspectives on the Origins of Modern Humans. Princeton University Press, Princeton, NJ.
Mook, W.G., 1982. Radiocarbon Dating. In: Kozlowski, J.K. (Ed.),
Excavations in the Bacho Kiro Cave (Bulgaria). Final Report,
Warszawa, p. 168.
MuK nzel, S., 1999. Die Gro{saK ugerfauna aus dem Gei{enkloK sterle. Unpublished "nal report for Deutsche Forschungsgemeinschaft.
Neugebauer-Maresch, C., 1996. Zu Stratigraphie und Datierung der
Aurignacien-Station am Galgenberg von Stratzing/Krems-Rehberg, NOG . In: Svoboda, J. (Ed.), Paleolithic in the Middle Danube
Region. Anniversary volume to Bohuslav Klima, Brno, ArcheologickyH uH stav AV, pp. 67}80.
Richter, D., Waiblinger, J., Rink, W.J., Wagner, G.A., 2000. Thermoluminescence, electron spin resonance and C-dating of the Late
Middle and Early Upper Palaeolithic site of Gei{enkloK sterle Cave in
Southern Germany. Journal of Archaeological Science 27, 71}89.
Riek, G., 1932. PalaK olithische Station mit Tierplastiken und menschlichen Skelettresten bei Stetten ob Lontal. Germania 16, 1}8.
Riek, G., 1934. Die EiszeitjaK gerstation am Vogelherd im Lonetal I. Die
Kulturen. TuK bingen.
Scholz, M., Bachmann, L., Nicholson, G.L., Bachmann, J., Hengst, S.,
Giddings, I., RuK scho!-Thale, B., Klostermann, J., Protsch von
Zieten, R.R., Czarnetzki, A., Pusch, C.M., 1999. Fossil}DNA based
classi"cation of the most northern Neanderthal man. In: Central
and Eastern Europe from 50,000}30,000 BP. Abstracts for the
International Workshop in the Neanderthal Museum, March
18}21, 1999, p. 72. Neandertal Museum, Neandertal, Germany.
Smith, F.H., Trinkaus, E., Pettitt, P.B., Karavanic, I., Paunovic, M.,
1999. Direct radiocarbon dates from Vindija G1 and Velika Pecina
Late Pleistocene Hominid remains. Proceedings of the National
Academy of Sciences 96 (22), 12281}12286.
Straus, L.G., 1996. Continuity or rupture; convergence or invasion;
adaptation or catastrophe; mosaic or monolith: views on the Middle
to Upper Paleolithic transition in Iberia. In: Carbonell, E., Vaquero,
M. (Eds.), The last Neandertals, the First Anatomically Modern
Humans: a Tale About the Human Diversity. Cultural Change and

Jqi=526=Shantha=VVC
40

M. Bolus, N.J. Conard / Quaternary International 75 (2001) 29}40

Human Evolution: the Crisis at 40 ka BP. Universitat Rovira i Virgili, Tarragona, pp. 203}218.
Svoboba, J., Lozek, V., Vlcek, E., 1996. Hunters between East and West.
The Paleolithic of Moravia. Plenum Press, New York.
Trinkaus, E. (Ed.), 1989. The Emergence of Modern Humans. Biocultural Adaptations in the Later Pleistocene. Cambridge University
Press, Cambridge.
Uthmeier, T., 1996. Ein bemerkenswert fruK hes Inventar des Aurignacien
von der Freilandfundstelle `Keilberg-Kirchea bei Regensburg. ArchaK ologisches Korrespondenzblatt 26, 233}248.
Valoch, K., 1993. Vedrovice V, eine Siedlung des Szeletien in
SuK dmaK hren. (With contributions by A. Kozi, W.G. Mook, E. Op-

ravil, J. v. d. Plicht, L. SmolmH kovaH , and Z. Weber). QuartaK r 43/44,

7}93.
Wagner, G.A., Beinhauer, K.W. (Eds.), 1997. Homo heidelbergensis von
Mauer. Das Auftreten des Menschen in Europa. Heidelberg.
UniversitaK tsverlag, Winter, 3165.
Zilha o, J., d'Errico, F., 1999a. The chronology and taphonomy of
the earliest Aurignacian and its implications for the understanding of Neandertal extinction. Journal of World Prehistory 13,
1}68.
Zilha o, J., d'Errico, F., 1999b. Reply. In: Mellars, P. et al. (Eds.), The
Neanderthal Problem Continued. Current Anthropology 40,
355}360.