Content uploaded by Andrei Dorian Soficaru
Author content
All content in this area was uploaded by Andrei Dorian Soficaru on Feb 24, 2021
Content may be subject to copyright.
Content uploaded by João Zilhão
Author content
All content in this area was uploaded by João Zilhão on Mar 05, 2019
Content may be subject to copyright.
Stable Isotope Evidence for Early Modern Human Diet in
Southeastern Europe: Peştera cu Oase, Peştera Muierii and Peştera
Cioclovina Uscată*
ERIK TRINKAUS, ANDREI SOFICARU, ADRIAN DOBOŞ,
SILVIU CONSTANTIN, JOÃO ZILHÃO and MICHAEL RICHARDS
*
Key-words: Radiocarbon dating, Early Upper Paleolithic,
Peştera cu Oase, Peştera Muierii, Peştera Cioclovina Uscată,
Middle Paleolithic, Early Upper Paleolithic.
Abstract: During the process of direct radiocarbon dating of
four Early Upper Paleolithic modern humans from the
Romanian sites of Peştera cu Oase, Peştera Muierii and Peştera
Cioclovina Uscată, carbon and nitrogen stable isotope ratios
(δ13C and δ15N) were measured from their bone collagen.
These individuals have δ13C values similar to other Late
Pleistocene humans. Their δ15N values are well within the
carnivore trophic level range, among the higher of the Middle
Upper Paleolithic values, and significantly greater than those
of preceding Middle Paleolithic and Initial Upper Paleolithic
Neanderthals. These data suggest a shift towards a broader
faunal dietary spectrum among these early modern humans,
despite western and eastern European archeological evidence
and human functional morphology indicating little change in
faunal exploitation from the Middle Paleolithic to the Early
Upper Paleolithic.
Cuvinte cheie: Datări cu radiocarbon, Peştera cu Oase, Peştera
Muierii, Peştera Cioclovina Uscată, Pleistocen superior,
Paleolitic mijlociu, Paleolitic superior.
Rezumat: Cu ocazia datării cu radiocarbon a patru fosile de
oameni moderni de la începutul paleoliticului superior vechi,
descoperite la Peştera cu Oase, Peştera Muierii şi Peştera
Cioclovina Uscată, din colagenul oaselor au fost măsurate şi
proporţiile izotopilor stabili de carbon şi azot (δ13C şi δ15N).
* Erik Trinkaus: Department of Anthropology, Campus
Box 1114, Washington University, Saint Louis MO 63130,
USA; Andrei Soficaru: Institutul de Antropologie “Fr. J.
Rainer,” Eroii Sanitari 8, P.O. Box 35-13, 050471 Bucureşti,
Romania; Adrian Doboş: Institutul de Arheologie „Vasile
Pârvan,” Henri Coandă 11, Sector 1, 010667 Bucureşti,
Romania: Silviu Constantin: Institutul de Speologie ”Emil
Racoviţă”, Str. Frumoasă 31, 010986 Bucureşti, Romania;
João Zilhão: Department of Archaeology and Anthropology,
University of Bristol 43 Woodland Road, Bristol BS8 1UU,
UK; Michael Richards: Department of Human Evolution, Max
Planck Institute for Evolutionary Anthropology, Deutscher
Platz 6, 04103 Leipzig, Germany; Department of Anthropology,
University of British Columbia, Vancouver BC V6T 1Z1, Canada.
Valorile δ13C sunt similare cu cele ale populaţiilor din
Pleistocenul superior, în vreme ce valorile δ15N indică o dietă
perponderent carnivoră. Aportul de carne în cazurile studiate
este printre cele mai ridicate din Paleoliticul superior mijlociu,
şi în acelaşi timp mult mai însemnat decât al neandertalilor din
Paleoliticul mijlociu şi de la începutul Paleoliticului superior
vechi. Aceste date arată că primii oamenii moderni din această
regiune au inclus în dietă o gamă mai variată de animale, deşi
datele arheologice şi morfologia funcţională a scheletelor
umane din vestul şi estul Europei indică schimbări minore în
exploatarea faunei pentru Paleoliticul mijlociu şi Paleoliticul
superior vechi.
Introduction
Insight into the dietary spectra of Late Pleistocene
humans, at least in Europe, is primarily based on
the associated faunal remains, especially for assem-
blages in which the primary accumulating agent
appears to have been humans, organic preservation
is good, and the excavation has been sufficient to
permit detailed zooarcheological analysis. However,
not all sites have sufficient preservation, many
were excavated in the past with less rigorous
methods (including industrial exploitation of the
site), and/or represent complex palimpsests of
human and carnivore activities variably combined
with geological disturbances of the sediments. As a
result, stable isotope analysis of human (and
faunal) remains from the Late Pleistocene has been
increasingly undertaken to provide further insight
into Late Pleistocene human dietary spectra,
especially for well-dated (and in many cases,
directly-dated) human remains.
These latter conditions variably apply to three
of the European early modern human samples,
those from the Peştera cu Oase, Peştera Muierii and
Peştera Cioclovina Uscată, all in southwestern
Romania (Rainer and Simionescu 1942; Gheorghiu
MATERIALE ŞI CERCETĂRI ARHEOLOGICE (serie nouă) V, 2009, p. 5–14
Erik Trinkaus, Andrei Soficaru, Adrian Doboş, Silviu Constantin, João Zilhão and Michael Richards
6
and Haas 1954; Trinkaus et al. 2003; Soficaru et al.
2006, 2007; Rougier et al. 2007). These samples
represent a substantial portion of the more
complete remains for early modern humans ≥29 ka
14C BP; only the Moravian Mladeč sample is
larger, the western European remains (such as
those from Brassempouy, La Quina-Aval and Les
Rois) consist principally of partial mandibles and
isolated teeth (Martin 1936; Gambier and Houët
1993; Henry-Gambier et al. 2004; Teschler-Nicola
2006), and further eastern European remains (from
Kostenki) consist of undescribed and/or undiagnostic
teeth and postcrania (Sinitsyn 2004; Anikovich
et al. 2007). The Romanian human remains are,
however, approached in time by the (probably)
early Gravettian remains from Cro-Magnon and
Paviland (Henry-Gambier 2002; Jacobi and Higham
2008).
The oldest of these remains, from the Peştera cu
Oase (Anina, Caraş-Severin) at ~35 ka 14C BP,
were water-transported surface finds within a
karstic system that served primarily as a hibernation
locale for large cave bear (Ursus spelaeus) with
fossil and taphonomic evidence for some wolf
(Canis lupus) activity (Zilhão et al. 2007). There
are no archeological remains or other evidence of
Pleistocene human activity within the cave system,
and the human fossils appear to be secondary
intrusions into the system. The isotopic data (Table 1)
derive from the direct 14C dating of the Oase 1
mandible (Trinkaus et al. 2003); repeated attempts
to directly date the Oase 2 cranium provided only a
minimum age for the specimen due to poor
collagen preservation (Rougier et al. 2007).
The next oldest specimens, from the Peştera
Muierii (Baia de Fier, Gorj; aka Peştera Muierilor)
at ~29-30 ka 14C BP, were found during
excavations in the Galeria Musteriană (Gheorghiu
and Haas 1954), probably secondarily mixed with
Middle Paleolithic lithic remains (Soficaru et al.
2006). Although there is evidence for an earlier
Upper Paleolithic occupation of the adjacent
Galeria Principală and especially Middle Paleolithic
activity in both galleries, the site contains an
abundance of U. spelaeus remains, combined with
those of other carnivores and herbivores. Given the
use of the cave system as a carnivore den, partial
excavation recovery of faunal remains, and un-
certainties regarding the association of the human
remains with specific archeological assemblages,
the only dietary data available for the Muierii
humans derives from their stable isotope values,
also obtained as part of the 14C dating process.
The youngest specimen, slightly more recent
than the Muierii remains at ~28,500 14C BP, is the
isolated human neurocranium from Peştera Cioclovina
Uscată (Boşorod, Hunedoara) (Rainer and Simionescu
1942; Soficaru et al. 2007). The cranium was found
during phosphate mining, and its original position
in the cave is unknown. Although Middle Paleolithic
and Early Upper Paleolithic assemblages have been
found in the cave (Dobrescu 2008), the cave served
principally as a hibernation den for U. spelaeus,
and remains of other species are rare. The human
fossil is therefore without stratigraphic, paleonto-
logical or archeological context (cf., Rainer and
Simionescu 1942; Păunescu 2001; Soficaru et al.
2007), and both its age and any dietary inferences
must be based on its bone chemistry.
Stable Isotopes and Diet
The measurements of the stable isotope ratios of
carbon (13C/12C, the δ
13C value) and nitrogen
(15N/14N, the δ15N value) in bone collagen extracted
from human and animal hard tissue has become a
well established method for determining past diets
(Sealy 2001; Lee-Thorp 2008). Through a number
of lines of research, including empirical observations
and controlled experiments on living organisms, it
has been determined that the carbon and nitrogen
isotope ratios in mammal bone collagen reflect the
isotope ratios of dietary protein consumed over the
last years of the life of that mammal (Wild et al.
2000).
Nitrogen in bone can only come from dietary
protein, so mammal bone collagen δ15N values
indicate the main sources of dietary protein in a
mammal’s long-term diet, and it is enriched by
approximately 2-4‰ over dietary protein δ15N
values (Schoeninger and DeNiro 1984; Jenkins
et al. 2001). In temperate environments such as
Europe, strict herbivores tend to have low δ15N
values, generally <7‰, whereas dedicated carnivores
have higher δ15N values, generally >8‰, and
omnivores have intermediate values (cf., Bocherens
2002).
There is variation between sites, and ideally one
should evaluate human δ15N values with respect to
values from animals of known diet from the same
levels of the same sites, something that is not
always possible (e.g., Richards et al. 2000;
Bocherens et al. 2005; Beauval et al. 2006). Such
Stable Isotope Evidence for Early Modern Human Diet in Southeastern Europe
7
correction is partly feasible for Peştera cu Oase
(Richards et al. 2008a) and, to a lesser extent, for
Peştera Muierii (Doboş et al. 2009). However,
dietarily significant differences in δ15N values,
given within population variation and within
dietary category variation, should mitigate intersite
variation in values.
Carbon isotope ratios (δ13C) do not change, or
change only slightly, between trophic levels, and
they therefore reflect variation in plant carbon
isotopes (C3 versus C4 plants, with the former
having lower or more negative δ13C values), access
to marine resources (which have higher δ13C
values), and possibly climate (van Klinken et al.
1994, Richards and Hedges 2003). In Late
Pleistocene Europe, C4 plants were not present, so
δ13C variation should reflect minor paleoecological
differences and, to some extent, access to marine
resources (Richards 2000; Pettitt et al. 2003).
Materials and Methods
The stable isotope data for the Oase, Muierii
and Cioclovina human remains was generated
through the direct AMS radiocarbon dating of the
specimens, principally by the Oxford Radiocarbon
Accelerator Unit (ORAU); in addition three
corroborative AMS dates derive from the Centrum
voor Isotopen Onderzoek in Groningen and the
Laboratoriet för 14C-datering i Lund (Table 1). The
dates and available chemistry have been published
previously (Trinkaus et al. 2003; Soficaru et al.
2006, 2007); all of the samples have δ13C values
and C:N ratios (DeNiro 1985) within acceptable
ranges; the initial ORAU Oase 1 date was a
minimum age, so further dating was done through
Groningen to produce a combined age of ~35 ka
14C BP. The ORAU dates were all run using
ultrafiltration (Brown et al. 1988; Higham et al.
2006a).
Oase 1 sample derives from the posteroinferior
mandibular corpus. The Muierii 1 sample is from
the zygomatic bone, and the Muierii 2 one is from
the squamous temporal. The Cioclovina 1 sample is
from a detached portion of the inferior occipital
bone. The relevant faunal remains from Oase, the
wolf (C. lupus), red deer (Cervus elaphus) and ibex
(Capra ibex) (Figure 2) bracket the Oase
1 mandible in time (Richards et al. 2008a). The
moose (Alces alces) sample from Muierii (an M1)
is radiometrically the same age as the human
remains, but the cave lion (Panthera spelaea) is
older (Soficaru et al. 2006; Doboş et al., 2009).
The other fauna providing stable isotopes from
Oase and Muierii, and all of the fauna providing
such data from Cioclovina, are cave bears
(Richards et al. 2008a; Doboş et al. 2009). Cave
bears, as with modern brown bears (Mowat and
Heard 2006), appear to have potentially broad
dietary spectra (Richards et al. 2008a) and
therefore cannot be used as a baseline for
evaluating the human isotopic results.
Comparative Late Pleistocene human stable
isotope values derive from published analyses of
European late archaic (Neandertal) and early
modern (Early and Middle Upper Paleolithic)
humans, many of which were generated in the
context of direct AMS radiocarbon dating of the
human remains (Table 2). Eight of the Neandertals
with stable isotope data are Late Pleistocene (OIS 4
and 3) Middle Paleolithic in context; the other three
Neandertals (Saint-Césaire 1, Spy 572a and Vindija
208) providing data are from the Initial Upper
Paleolithic (Lévêque et al. 1993; Janković et al.
2006; Semal et al. 2009). In cases in which
specimens have been redated (e.g., Higham et al.
2006b; Jacobi and Higham 2008), values from the
more recent analyses are employed. Variation in
values from repeated measures of the same
specimens and from different sample preparation
protocols tend to be modest, close to ±1.0‰.
Most of the human and faunal data derive from
fully mature bone. However, four of the specimens
are immature (the Middle Upper Paleolithic
Kostenki 4 and Sunghir 2 and 3 late juveniles and
the Neandertal Engis 2 infant) and one (the Jonzac
1 Neandertal) is a premolar root. In addition,
among the comparative faunal remains, the Peştera
Muieri A. alces data are from the root of an M1. It
appears that nursing immature individuals should
be considered to be a trophic level higher than
adults due to lactation. As a result, bone or dentin
formed postnatally and prior to weaning may have
δ15N values a few per mil higher than that of the
mother, depending on the duration of nursing and
the degree to which the diet of the infant is
supplemented with other foods (Jenkins et al. 2001;
Fuller et al. 2006; Jay et al. 2008). This should not
pose a problem for the bone samples from the older
juvenile Kostenki and Sunghir remains, nor for the
Jonzac premolar given late juvenile root formation
for human premolars (Smith 1991). However, the
Engis 2 δ15N value of 12.6‰ (Bocherens et al.
Erik Trinkaus, Andrei Soficaru, Adrian Doboş, Silviu Constantin, João Zilhão and Michael Richards
8
2001) may be elevated relative to its population
average, and the A. alces δ
15N value of 7.3‰
(Doboş et al. 2009) may be similarly elevated
given early postnatal formation of cervid first
molar roots (Brown and Chapman 1991).
Given the non-parametric nature of the stable
isotope distributions, sample comparisons are made
using Wilcoxon P-values. Significance levels are
adjusted using a sequentially reductive multiple
comparison corrections within sets of tests
(Proschan and Waclawiw 2000).
Results
Among Late Pleistocene humans, there is a shift
in average δ13C and δ15N values from the Nean-
dertals to the Middle Upper Paleolithic modern
humans (Fig. 1). The latter sample has significantly
higher (less negative) δ13C values (P = 0.001). This
is due in part to the high values for Arene Candide
1 and La Rochette 1 and the low values for
Feldhofer 1 and 2. The Arene Candide 1 δ13C value,
as well as the slightly lower Paviland 1 one, are
apparently due to the consumption of significant
amounts of maritime resources (Richards 2000;
Pettitt et al. 2003). It is not clear why Feldhofer 1
and 2 have very low δ13C values, but they are close
to those of cervids from the same site (Richards
and Schmitz 2008).
In δ15N values, there is a modest but non-
significant shift between the adults of the
Neandertal and Middle Upper Paleolithic samples
(P = 0.129), despite the low values for the
Feldhofer specimens; including the value for the
Engis 2 provides less of a difference (P = 0.236).
The δ15N values for the Initial Upper Paleolithic
Saint-Césaire 1, Spy 572a and Vindija 208 (11.4‰,
11.0‰ and 10.3‰ respectively) fall among the
Middle Paleolithic Neandertal values.
The δ15N values for Peştera cu Oase and Peştera
Muierii carnivores and herbivores are within
expected values for these dietary groups, with the
wolves and the cave lion being ~8‰ up to ~12‰
and the herbivores being below 7.5‰ (Fig. 2);
reducing the A. alces value slightly, given the
young age of its M1 formation, would place it
closer to the other herbivores. The four Romanian
early modern humans are in the middle of the
faunal range in δ13C values, but they are at the top
of the range and above the carnivores in δ15N.
When considered among other Late Pleistocene
humans (Fig. 1), these Early Upper Paleolithic
humans are among the highest of the Middle Upper
Paleolithic humans, being exceeded only by Barma
Grande 6 and Kostenki 4 in their δ15N values. The
very high δ15N Early Upper Paleolithic value is
provided by Kostenki 8, an undescribed tibia and
fibula dated to ~32.6 ka 14C BP (Richards et al.
2001; Sinitsyn 2004). At the same time, these four
human δ15N values are above those of all of the
mature Neandertals, but matched by the Engis
2 infant’s value. As a group they are significantly
(P = 0.006) higher than those of the adult
Neandertals (P = 0.003 with Kostenki 8; P = 0.005
with both Kostenki 8 and Engis 2 included).
Discussion
The modest shift in δ15N values between the
Neandertal and Middle Upper Paleolithic samples,
in combination with the generally higher more
recent δ13C values, has been attributed to a broader
food spectrum and a greater reliance on aquatic
resources with their longer food chains among
some of the Middle Upper Paleolithic populations
(Richards et al. 2001). The growing evidence for
some degree of maritime resource exploitation
among Neandertals (Stiner, 1994; Stringer et al.
2008), however, suggests that any dietary contrasts
may be more subtle than originally suggested
and/or more regionally variable.
The addition of data from the earliest modern
humans in Europe, now represented isotopically by
those from Oase, Muierii and Cioclovina, as well
as the undescribed Kostenki 8 specimen, extends
any isotopic contrasts further back in the Upper
Paleolithic. The significant increase in δ15N values
for these early modern humans relative to the
Neandertals implies an increase in dietary protein
from animals and/or in aquatic resources with long
food chains relative to the Neandertals. Yet, any
such differences are likely to be in degree, given
the proximity of most of the isotopic values
between the Neandertals and these early modern
humans, all of them well within the “carnivore”
range.
At the same time, the available archeological
and human paleontological evidence does not imply
a major dietary difference between Neandertals and
Early Upper Paleolithic modern humans. There is
archeological evidence for improved weaponry in
the Aurignacian (Knecht, 1993; Liolios, 2006;
Bolus and Conard, 2006), and similar evidence for
effective spears is known from the Galeria
Stable Isotope Evidence for Early Modern Human Diet in Southeastern Europe
9
Principală in the Peştera Muierii (Gheorghiu et al.
1954; Păunescu 2000) and occurs in other
Romanian Aurignacian assemblages (Chirica et al.
1996; Dobrescu 2008). Yet, Aurignacian faunal
assemblages, to the extent that they have been
appropriately analyzed zooarcheologically (albeit
not in southeastern Europe [cf., Dobrescu 2008]),
show little difference with earlier Middle Paleolithic
or Initial Upper Paleolithic ones (Grayson and
Delpech 2002, 2003, 2008; Morin 2008). Organic
residue analysis of eastern European Middle Paleo-
lithic and Early Upper Paleolithic lithics (Hardy et
al. 2001) shows similar patterns of exploitation in
both groups of a broad spectrum of plant and
animal resources. In addition, functional analyses
of the few known Early Upper Paleolithic human
upper limb remains (including the Muierii 1 scapula)
show little of the anatomical change associated
with habitual spear throwing (Trinkaus et al. 2006;
Trinkaus 2008; cf., Churchill and Rhodes 2006).
It also remains to be assessed whether some
degree of the isotopic contrast may be related to
variation in geographic and/or climatic milieu. The
Neandertal data derive from central and western
Europe, whereas the Early Upper Paleolithic data
are all eastern European (Tables 1 and 2). Yet, the
Middle Upper Paleolithic human sample derives
from sites across Europe, and the few herbivore
and carnivore isotopic values for Peştera cu Oase
and Peştera Muierii are similar to those for western
European Late Pleistocene mammals.
Clearly, more sampling of both Early Upper
Paleolithic and preceding humans in diverse
ecozones of Europe is needed to resolve this
isotopic contrast, in addition to zooarcheological
data from across Europe and including species
other than ungulates.
Conclusion
The addition of carbon and nitrogen stable
isotopic data for four Early Upper Paleolithic
modern humans from Romania adds significantly
to our dietary record for the European transitional
period from Middle Paleolithic to Middle Upper
Paleolithic and from Neandertals to early modern
humans. A significant increase in δ15N values
among the more recent humans implies greater
carnivory and/or a broader proteinaceous dietary
spectrum for them, a trend apparently present into
the Middle Upper Paleolithic. However, current
archeological analyses suggest little dietary change
from the Middle Paleolithic through the Early
Upper Paleolithic, turning this isotopic insight into
hypotheses to be, hopefully, tested against
additional isotopic and faunal data.
Acknowledgments
The excavation and/or analysis of the Oase,
Muierii and Cioclovina human remains has been
supported by National Science Foundation (USA)
grants BCS-0409194 and BCS-0509072, Wenner-
Gren Foundation grants 7111 and 7290, the Leakey
Foundation, the Instituto Português de Arqueologia,
the Institut royal des Sciences naturelles de
Belgique, the Romanian National Council for
Academic Research (CNCSIS 1258/2005), and the
Romanian Ministry for Education and Research
(CEEX 627/2005). I. Povara, C. Petrea, M. Fifor,
F. Ridiche and A. Popescu provided access to the
Muierii human remains, and T. Neagu provided
access to the Cioclovina fossil. A. Lister and
M. Breda helped identify the Alces molar. To all of
them we are grateful.
Table 1
Direct radiocarbon ages and stable isotopes for the Oase, Muierii and Cioclovina human remains. For detailed chemistry, see
Trinkaus et al. (2003) and Soficaru et al. (2006, 2007). The two dates for Oase 1 provide a combined age of 34,950 +990, -890 B.P.
(Trinkaus et al. 2003). δ13C values are measured relative to the VPDB (Vienna Pee Dee Belemnite) standard, and δ15N values are
measured relative to the AIR (Ambient Inhalable Reservoir) standard
Specimen 14C age δ13C (‰) δ15N (‰)
Oase 1 >35,200 (OxA-11711) -18.7 13.3
34,290, +970, -870 (GrA-22810) -19.0 --
Muierii 1 30,150 ± 800 (LuA-5228) -20.0 --
29,930 ± 170 (OxA-15529) -19.3 12.3
Muierii 2 29,110 ± 190 (OxA-16252) -19.1 12.4
Cioclovina 1 29,000 ± 700 (LuA-5229) -- --
28,510 ± 170 (OxA-15527) -19.6 12.7
Erik Trinkaus, Andrei Soficaru, Adrian Doboş, Silviu Constantin, João Zilhão and Michael Richards
10
Table 2.
Comparative Middle and Initial Upper Paleolithic Neandertal and
Early and Middle Upper Paleolithic modern human stable isotope data from Europe
δ
13C (‰) δ15N (‰) Source
Middle Paleolithic Neandertals
Engis 2 (child), Belgium –19.6 12.6 Bocherens et al. 2001
Feldhofer 1, Germany –21.6 7.9 Richards and Schmitz 2008
Feldhofer 2, Germany –21.5 9.0 Richards and Schmitz 2008
Jonzac 1, France –20.7 10.6 Richards et al. 2008b
Les Pradelles 64801, France –19.1 11.6 Bocherens et al. 2005
Les Pradelles M300, France –19.1 11.5 Bocherens et al. 2005
Les Pradelles M400, France –19.5 11.4 Bocherens et al. 2005
Rochers-de-Villeneuve 1, France –19.0 11.6 Beauval et al. 2006
Initial Upper Paleolithic Neandertals
Saint-Césaire 1, France –19.8 11.4 Bocherens et al. 2005
Spy 572a, Belgium –19.8 11.0 Bocherens et al. 2001
Vindija 208, Croatia –20.2 10.3 Higham et al. 2006b
Early Upper Paleolithic
Kostenki 8, Russia –18.2 15.3 Richards et al. 2001
Middle Upper Paleolithic
Arene Candide 1, Italy –17.6 12.4 Pettitt et al. 2003
Barma Grande 6, Italy –19.7 12.9 Formicola et al. 2004
Brno-Francouzská 2, Czech Rep. –19.0 12.3 Richards et al. 2001
Dolní Vĕstonice 35, Czech Rep. –18.8 12.3 Richards et al. 2001
Eel Point 1, UK –19.0 10.9 Schulting et al. 2005
Kostenki 4, Russia –19.1 13.1 Richards et al. 2001
Paviland 1, UK –18.2 10.35 Jacobi and Higham 2008
La Rochette 1, France –17.1 11.2 Orschiedt 2002, per. comm.
Sunghir 1, Russia –19.2 11.3 Richards et al. 2001
Sunghir 2, Russia –19.0 11.2 Richards et al. 2001
Sunghir 3, Russia –18.9 11.3 Richards et al. 2001
REFERENCES
Anikovich et al. 2007 – M. V. Anikovich, A. A. Sinitsyn, J. F.
Hoffecker, V. T. Holliday, V. V. Popov, S. N. Lisitsyn,
S. L. Forman, G. M. Levkovskaya, G. A. Pospelova,
I. E. Kuz’mina, N. D. Burova, P. Goldberg, R. I. Macphail,
B. Giaccio, N. D. Praslov, Early Upper Paleolithic in
eastern Europe and implications for the dispersal of
modern humans, Science 315, 2007, p. 223-226.
Beauval et al. 2006 – C. Beauval, F. Lacrampe-Cuyaubère,
B. Maureille, E. Trinkaus, Direct radiocarbon dating and
stable isotopes of the Neandertal femur from Les Rochers-
de-Villeneuve (Lussac-les-Châteaux, Vienne), Bulletins et
Mémoires de la Société d’Anthropologie de Paris 18, 2006,
p. 35-42.
Bocherens 2002 – H. Bocherens, Alimentation des ours et
signatures isotopiques, ERAUL 100, 2002, p. 41-49.
Bocherens et al. 2001 – H. Bocherens, D. Billiou, A. Mariotti,
M. Toussaint, M. Patou-Mathis, D. Bonjean, M. Otte,
M., New isotopic evidence for dietary habits of Neandertals
from Belgium, Journal of Human Evolution 40, 2001,
p. 497-505.
Bocherens et al. 2005 – H. Bocherens, D. Drucker, D. Billiou,
M. Patou-Mathis, B. Vandermeersch, Isotopic evidence for
diet and subsistence pattern of the Saint-Césaire I
Neanderthal: review and use of a multi-source mixing
model. Journal of Human Evolution 49, 2005, p. 71-87.
Bolus şi Conard 2006 – M. Bolus, N.J. Conard., Zur Zeit-
stelling von Geschossspitzen aus organischen Materialien
im späten Mittelpaläolithikum und Aurignacien, ArchKorr
36, 2006, p. 1-15.
Stable Isotope Evidence for Early Modern Human Diet in Southeastern Europe
11
Brown et al. 1988 – T. A. Brown, D. E. Nelson, J. S. Vogel,
J. R. Southon, Improved collagen extraction by modified
Longin method, Radiocarbon 30, 1988, p. 171-177.
Brown şi Chapman 1991 – W.A.B. Brown, N. G. Chapman,
Age assessment of fallow deer (Dama dama): from a
scoring scheme based on radiographs of developing
permanent molariform teeth, Journal of Zoology 224,
1991, p. 367-379.
Chirica et al. 1996 – V. Chirica, I. Borziac, N. Chetraru,
Gisements du paléolithique superieur ancien entre le
Dniestr et la Tissa, Iaşi, 1996.
Churchill, Rhodes 2006 – S. E. Churchill, J. A. Rhodes How
strong were the Neandertals? Leverage and muscularity at
the shoulder and elbow in Mousterian foragers, Periodicum
Biologorum 108, 2006, p. 457-470.
DeNiro 1985 – M. J. DeNiro, Post-mortem preservation and
alteration of in vivo bone collagen isotope ratios in
relation to paleodietary reconstruction, Nature 317, 1985,
p. 806-809.
Doboş et al. 2009 – A. Doboş, A. Soficaru, A. Popescu,
E. Trinkaus, Radiocarbon dating and faunal stable isotopes
for the Galeria Principală, Peştera Muierii, Baia de Fier,
Gorj County, Romania, MCA 5, 2009, p. 15–20.
Dobrescu 2008 – R. Dobrescu, Aurignacianul din Transilvania,
Bucureşti, 2008.
Formicola et al. 2004 – V. Formicola, P. B. Pettitt, A. del
Lucchese, A direct AMS radiocarbon date on the Barma
Grande 6 Upper Paleolithic skeleton, Current Anthropology
45, 2004, p. 114-118.
Fuller et al. 2006 – B.T Fuller, J. L. Fuller, D. A. Harris,
R.E.M. Hedges, Detection of breastfeeding and weaning in
modern human infants with carbon and nitrogen stable
isotope ratios, American Journal of Physical Anthropology
129, 2006, p. 279–293.
Gambier, Houët 1993 – D. Gambier, F. Houët, Hominid
Remains An Up-Date: France Upper Palaeolithic,
Anthropologie et Préhistoire/Supplément 6, 1993, p. 1-120.
Gheorghiu, Haas 1954 – A. Gheorghiu, N. Haas, Date privind
omul primitiv de la Baia de Fier. Consideraţii paleo-
antropologice, AAR-Sesiunea Secţiunii de Ştiinţe
Medicale a Academiei R.P.R. 41, 1954, p. 641-63.
Gheorghiu et al. 1954 – A. Gheorghiu, C.S. Nicolăescu-
Plopşor, N. Haas, E. Comşa, C. Preda, G. Bombiţă,
G. Enea, F. Gheorghiu, S. Iofcea, D. Nicolăescu-Plopşor,
A. Neagoe, R. Silveanu, I. Surdu I., Raport preliminar
asupra cercetărilor de paleontologie umană de la Baia de
Fier (Reg. Craiova) din 1951, Probleme de Antropologie
1, 1954, p. 73-86.
Grayson, Delpech 2002 – D. K. Grayson, F. Delpech, Specialized
Early Upper Palaeolithic hunters in southwestern France?
JArS 29, 2002, p. 1439–1449.
Grayson, Delpech 2003 – D. K. Grayson, F. Delpech,
Ungulates and the Middle-to-Upper Paleolithic transition
at Grotte XVI (Dordogne, France), JArS 30, 2003,
p. 1633–1648.
Grayson, Delpech 2008 – D. K. Grayson, F. Delpech, The large
mammals of Roc de Combe (Lot, France): The Châtel-
perronian and Aurignacian assemblages, Journal of
Anthropological Archaeology 27, 2008, p. 338–362.
Hardy et al. 2001 – B. L.Hardy, M. Kay, A. E. Marks,
K. Monigal, Stone tool function at the paleolithic sites of
Starosele and Buran Kaya III, Crimea: Behavioral
implications, Proceedings of the National Academy of
Sciences USA 98, 2001, p. 10972–10977.
Henry-Gambier 2002 – D. Henry-Gambier, Les fossiles de
Cro-Magnon (Les Eyzies-de-Tayac, Dordogne): nouvelles
données sur leur position chronologique et leur attribution
culturelle, Bulletins et Mémoires de la Société d’Anthro-
pologie de Paris 14, 2002, p.89–112.
Henry-Gambier et al. 2004 – D. Henry-Gambier, B. Maureille,
R. White, Vestiges humains des niveaux de l’Aurignacien
ancien du site de Brassempouy (Landes), Bulletins et
Mémoires de la Société d’Anthropologie de Paris 16, 2004,
p. 49–87.
Higham et al. 2006a – T. F. G. Higham, R. M. Jacobi,
C. Bronk Ramsey, AMS radiocarbon dating of ancient
bone using ultrafiltration, Radiocarbon 48, 2006, p. 179–195.
Higham et al. 2006b – T.F. F. Higham, C. Bronk Ramsey,
I. Karavanić, F. H. Smith, E. Trinkaus, Revised direct
radiocarbon dating of the Vindija G1 Upper Paleolithic
Neandertals, Proceedings of the National Academy of
Sciences USA 103, 2006, p. 553-557.
Jacobi, Higham 2008 – R. M. Jacobi, T. F. G. Higham, The
‘‘Red Lady’’ ages gracefully: new ultrafiltration AMS
determinations from Paviland, Journal of Human
Evolution 55, 2008, p. 898–907.
Janković et al. 2006 – I. Janković, I. Karavanić, J. C. M. Ahern,
D. Brajković, J. M. Lenardić, F. H. Smith, Vindija Cave
and the modern human peopling of Europe, Collegium
Antropologicum 30, 2006, p. 315-324.
Jay et al. 2008 – M. Jay, B. T. Fuller, M. P. Richards, C. J. Knüsel,
S. S. King, Iron Age breastfeeding practices in Britain:
Isotopic evidence from Wetwang Slack, East Yorkshire,
American Journal of Physical Anthropology 136, 2008,
p. 327–337.
Jenkins et al. 2001 – S. G. Jenkins, S. T. Partridge, T. R.
Stephenson, S. D. Farley, C. T Robbins, Nitrogen and
carbon isotope fractionation between mothers, neonates,
and nursing offspring, Oecologia 129, 2001, p. 336–341.
Knecht 1993 – H. Knecht, Splits and wedges: The techniques
and technology of early Aurignacian antler working, in
H. Knecht, A. Pike-Tay, R. White (eds.) Before Lascaux:
The Complex Record of the Early Upper Paleolithic, 1993,
p. 137-162.
Lee-Thorp 2008 – J. A. Lee-Thorp, On isotopes and old bones,
Archaeometry 50, 2008, p. 925-950.
Lévêque et al. 1993 – F. Lévêque, A. M. Backer, M. Guilbaud
(eds.) Context of a Late Neandertal. Implications of
Multidisciplinary Research for the Transition to Upper
Paleolithic Adaptations at Saint-Césaire, Charente-Maritime,
France, 1993.
Liolios D. 2006, Reflections on the role of bone tools in the
definition of the Early Aurignacian, in O. Bar-Yosef,
J. Zilhão (eds.) Towards a Definition of the Aurignacian,
Trabalhos de Arqueologia 45, 2006, p. 37-51.
Martin 1936 – H. Martin, Nouvelles consatations faites dans la
station aurignacienne de La Quina (Charente), BSPF 31,
1936, p. 177-202.
Erik Trinkaus, Andrei Soficaru, Adrian Doboş, Silviu Constantin, João Zilhão and Michael Richards
12
Morin 2008 – E. Morin, Evidence for declines in human
population densities during the early Upper Paleolithic in
western Europe, Proceedings of the National Academy of
Sciences USA 105, 2008, p. 48-53.
Mowat, Heard 2006 – G. Mowat, D. C. Heard, Major
components of grizzly bear diet across North America,
Canadian Journal of Zoology 84, 2006, p. 473–489.
Orschiedt 2002 – J. Orschiedt, Datation d’un vestige humain
provenant de La Rochette (Saint Léon-sur-Vézère,
Dordogne) par la méthode de carbone 14 en spectrométrie
de masse, Paléo 14, 2002, p. 239-240.
Păunescu 2000 – A. Păunescu, Paleoliticul şi mezoliticul din
spaţiul cuprins între Carpaţi şi Dunăre, Bucureşti, 2000.
Păunescu 2001 – A. Păunescu, Paleoliticul şi Mezoliticul din
spaţiul Transilvan, Bucureşti, 2001.
Pettitt et al. 2003 – P. B. Pettitt, M. P. Richards, R. Maggi,
V. Formicola, The Gravettian burial known as the Prince
(‘Il Principe’): new evidence for his age and diet,
Antiquity 295, 2003, p. 15-19.
Proschan, Waclawiw 2000 – M. A. Proschan, M. A. Waclawiw,
Practical guidelines for multiplicity adjustment in clinical
trials, Controlled Clinical Trials 21, 2000, p. 527–539.
Rainer, Simionescu 1942 – F. Rainer, I. Simionescu, Sur le
premier crâne d’homme Paléolithique trouvé en Roumanie,
AAA-MSS Seria III/17, 1942, p. 489-503.
Richards 2000 – M. P. Richards, Human and faunal stable
isotope analyses from Goat’s Hole and Foxhole Caves,
Gower in Aldhouse-Green, S. (ed.) Paviland Cave and the
‘Red Lady,’ Bristol, 2000, p. 71-75.
Richards, Hedges 2003 – M. P. Richards, R. E. M. Hedges,
Bone collagen d13C and d15N values of fauna from
northwest Europe reflect palaeoclimatic variation over the
last 40,000 years, Palaeogeography, Palaeoclimatology,
Palaeoecology 193, 2003, p. 261-267
Richards et al. 2000 – M. P. Richards, P. B. Pettitt, E. Trinkaus,
F. H. Smith, M. Paunović, I. Karavanić, Neanderthal diet
at Vindija and Neanderthal predation: The evidence from
stable isotopes, Proceedings of the National Academy of
Science USA 97, 2000, p. 7663-7666.
Richards et al. 2001 – M. P. Richards, P. B. Pettitt, M. C. Stiner,
E. Trinkaus, Stable isotope evidence for increasing dietary
breadth in the European mid-Upper Paleolithic,
Proceedings of the National Academy of Science USA 98,
2001, p. 6528-6532.
Richards, Schmitz 2008 – M. P. Richards, R. W. Schmitz,
Isotopic evidence for the diet of the Neanderthal type
specimen, Antiquity 82, 2008, p. 553–559.
Richards et al. 2008a – M. P. Richards, M. Pacher, M. Stiller,
J. Quilès, M. Hofreiter, S. Constantin, J. Zilhão, E. Trinkaus,
Isotopic evidence for omnivory among European cave
bears: Late Pleistocene Ursus spelaeus from the Peştera cu
Oase, Romania, Proceedings of the National Academy of
Sciences USA 105, 2008, p. 600-604.
Richards et al. 2008b – M. P. Richards, G. Taylor, T. Steele,
S. P. McPherron, M. Soressi, J. Jaubert, J. Orschiedt, J. B.
Mallye, W. Rendu, J. J. Hublin, Isotopic dietary analysis
of a Neanderthal and associated fauna from the site of
Jonzac (Charente-Maritime), France, Journal of Human
Evolution 55, 2008, p. 179-185.
Rougier et al. 2007 – H. Rougier, Ş. Milota, R. Rodrigo, M.
Gherase, L. Sarcină, O. Moldovan, J. Zilhão, S. Constantin,
R. G. Franciscus, C. P. E. Zollikofer, M. Ponce-de-León,
E. Trinkaus, Peştera cu Oase 2 and the cranial morphology of
early modern Europeans, Proceedings of the National
Academy of Sciences USA 104, 2007, p. 1164-1170.
Schoeninger, DeNiro 1984 – M. Schoeninger, M. DeNiro,
Nitrogen and carbon isotopic composition of bone
collagen from marine and terrestrial animals, Geochimica
et Cosmochimica Acta 48, 1984, p. 625-639.
Schulting et al. 2005 – R. J. Schulting, E. Trinkaus, T.
Higham, R. Hedges, M. Richards, B. Cardy, A Mid-Upper
Palaeolithic human humerus from Eel Point, South Wales,
UK, Journal of Human Evolution 48, p. 493-505.
Sealy 2001 – Sealy, J., Body tissue chemistry and palaeodiet in
D. R. Brothwell, A. M. Pollard, (eds.) Handbook of
Archaeological Sciences. New York, 2001, p. 269-279.
Semal et al. 2008 – P. Semal, H. Rougier, I. Crevecoeur, C.
Jungels, D. Flas, A. Hauzeur, B. Maureille, M. Germonpré,
H. Bocherens, S. Pirson, L. Cammaert, N. De Clerck, A.
Hambucken, T. Higham, M. Toussaint, J. van der Plicht,
New data on the late Neandertals: Direct dating of the
Belgian Spy fossils, American Journal of Physical
Anthropology 138, p. 421–428.
Sinitsyn 2004 – A. A. Sinitsyn, Les sépultures de Kostenki:
Chronologie, attribution culturelle, rite funéraire. Études
et Recherches Archéologiques de l’Université de Liège
106, 2004, p. 237-244.
Smith 1991 – B. H. Smith, Standards of human tooth formation
and dental age assessment. In Kelley, M.A, Larsen, C.S.
(eds.) Advances in Dental Anthropology. New-York, 1991.
p. 143-168.
Soficaru et al. 2006 – A. Soficaru, A. Doboş, E. Trinkaus,
Early modern humans from the Peştera Muierii, Baia de
Fier, Romania, Proceedings of the National Academy of
Sciences USA 103, 2006, p. 17196-17201.
Soficaru et al. 2007 – A. Soficaru, C. Petrea, A. Doboş, E.
Trinkaus, The human cranium from the eştera Cioclovina
Uscată, Romania: Context, age, taphonomy, morphology
and paleopathology, Current Anthropology 48, 2007, p.
611-619.
Stiner 1994 – M. C. Stiner, Honor Among Thieves, Princeton,
Princeton University Press, 1994.
Stringer et al. 2008 – C. B. Stringer, J. C. Finlayson, R. N. E.
Barton, Y. Fernández-Jalvo, I. Cáceres, R. C. Sabin, E. J.
Rhodes, A. P. Currant, J. Rodríguez-Vidal, F. Giles-
Pacheco, J. A. Riquelme-Cantal, Neanderthal exploitation
of marine mammals in Gibraltar, Proceedings of the
National Academy of Sciences USA 105, 2008, p. 14319–
14324.
Teschler-Nicola 2006 – M. Teschler-Nicola, Early Modern
Humans at the Moravian Gate, Vienna, Spinger Verlag,
2006.
Trinkaus 2008 – E. Trinkaus, Behavioral implications of the
Muierii 1 early modern human scapula. Annuaire
Roumain d’Anthropologie 45, p. 27–41.
Trinkaus et al. 2003 – E. Trinkaus, O. Moldovan, Ş. Milota, A.
Bîlgăr, L. Sarcina, S. Athreya, S.E. Bailey, R. Rodrigo, M.
Gherase, T. Higham, C. Bronk Ramsey, J. van der Plicht,
Stable Isotope Evidence for Early Modern Human Diet in Southeastern Europe
13
An early modern human from the Peştera cu Oase,
Romania, Proceedings of the National Academy of
Sciences USA 100, 2003, p. 11231-11236.
Trinkaus et al. 2006 – E. Trinkaus, F. H. Smith, T. C. Stockton,
L. L. Shackelford, The human postcranial remains from
Mladeč, in M. Teschler-Nicola (ed.) Early Modern
Humans at the Moravian Gate: The Mladeč Caves and
their Remains, Vienna, 2006, Springer Verlag, p. 385-445.
van Klinken et al. 1994 – G. J. van Klinken, H. van der Plicht,
R. Hedges, Bone 13C/12C ratios reflect (palaeo-) climatic
variations, Geophysical Research Letters 21, 1994, p. 445-
448.
Wild et al. 2000 –, E. M. Wild, , K. A. Arlamovsky, R. Golser,
W. Kutschera, A. Priller, S. Puchegger, W. Rom, P. Steier,
W. Vycudilik, 14C dating with the bomb peak: An
application to forensic medicine, Nuclear Instruments and
Methods in Physics Research 172B, 2000, p. 944-950.
Zilhão et al. 2007 – J. Zilhão, E. Trinkaus, S. Constantin,
Ş. Milota, M. Gherase, L. Sarcină, A. Danciu, H. Rougier,
J. Quilès, R. Rodrigo, The Peştera cu Oase people,
Europe’s earliest modern humans in P. Mellars, K. Boyle,
O. Bar-Yosef, C. Stringer (eds.) Rethinking the Human
Revolution, McDonald Institute of Archaeology
Monographs, Cambridge UK, 2007, p. 249-262.
CAPTIONS TO THE FIGURES
Fig. 1. Human stable isotope values for Late Pleistocene
European humans. Black circles: Early Upper Paleolithic
humans; gray diamonds: Neandertals; gray triangles:
Middle Upper Paleolithic humans. The δ15N for Engis 2
(the highest Neandertal value at 12.6‰) may be elevated
given the young age of the individual at death. The high
EUP δ15N outlier is Kostenki 8, and the low Neandertal
values are Feldhofer 1 and 2.
Fig. 2. Human stable isotopes (black circles) from the Peştera
cu Oase (O1), Peştera Muierii (M1, M2) and Peştera
Cioclovina (C1), plotted against carnivores (gray
diamonds) and herbivores (gray triangles) from the first
two sites. The human isotopic values are in Table 1, and
the faunal values are in Richards et al. (2008) and Doboş et
al. (2008) respectively for Peştera cu Oase and Peştera
Muierii. The high herbivore δ15N value is the Muierii
A. alces (7.3‰), which is probably elevated since it
derives from an M1 root. The Oase herbivores consist of
Cervus elaphus and Capra ibex, the Oase carnivores are
Canis lupus, and the Muierii carnivore is Panthera
spelaea.
Erik Trinkaus, Andrei Soficaru, Adrian Doboş, Silviu Constantin, João Zilhão and Michael Richards
14
Fig. 1. Human stable isotope values for Late Pleistocene European humans. Black circles: Early Upper Paleolithic humans;
gray diamonds: Neandertals; gray triangles: Middle Upper Paleolithic humans. The δ15N for Engis 2 (the highest Neandertal value at
12.6‰) may be elevated given the young age of the individual at death. The high EUP δ15N outlier is Kostenki 8,
and the low Neandertal values are Feldhofer 1 and 2.
Fig. 2. Human stable isotopes (black circles) from the Peştera cu Oase (O1), Peştera Muierii (M1, M2) and Peştera Cioclovina (C1),
plotted against carnivores (gray diamonds) and herbivores (gray triangles) from the first two sites. The human isotopic values are in
Table 1, and the faunal values are in Richards et al. (2008) and Doboş et al. (2008) respectively for Peştera cu Oase and
Peştera Muierii. The high herbivore δ15N value is the Muierii A. alces (7.3‰), which is probably elevated since it derives
from an M1 root. The Oase herbivores consist of Cervus elaphus and Capra ibex, the Oase carnivores are Canis lupus,
and the Muierii carnivore is Panthera spelaea.
