Reconstruction of the Gravettian food-web at Předmostí I using multi-isotopic tracking (13C, 15N, 34S) of bone collagen

Quaternary International xxx (2014) 1e18

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Reconstruction of the Gravettian food-web at P
redmostí I using multi-isotopic tracking (13C, 15N, 34S) of bone collagen  Bocherens a, b, *, Dorothe e G. Drucker a, Mietje Germonpre  c, Herve zni
-Galetova  d, e, f, Yuichi I. Naito a, Christoph Wissing a, Martina La ckova g , h , Martin Oliva e Jaroslav Br u
zek €t Tübingen, Ho €lderlinstrasse 12, 72074 Tübingen, Germany Department of Geosciences, Universita €t Tübingen, Ho €lderlinstrasse 12, 72074 Tübingen, Germany Senckenberg Center for Human Evolution and Palaeoenvironment (HEP), Universita Operational Direction “Earth and History of Life”, Royal Belgian Institute of Natural Sciences, Vautierstraat 29, 1000 Brussels, Belgium d 
ckova 15, 306 14 Pilsen, Czech Republic Department of Anthropology, Faculty of Philosophy and Arts, University of West Bohemia, Sedla e Anthropos Institute, Moravian Museum, Zelný trh 6, 659 37 Brno, Czech Republic f  Equipe “Comportements des N eandertaliens et des Hommes anatomiquement modernes replac es dans leur contexte pal eo ecologique”, UMR 7194, D epartement de Pr ehistoire du MNHN, Institut de Pal eontologie Humaine, 1 rue Ren e Panhard, 75 013 Paris, France g A3P-PACEA, UMR 5199 CNRS, Universit e Bordeaux, CS 50023 33615 Pessac Cedex, France h  7, 128 44 Prague 2, Czech Republic Department of Anthropology and Human Genetics, Faculty of Science, Charles University, Vini
cna a

b c

a r t i c l e i n f o

a b s t r a c t

Article history: Available online xxx

The Gravettian site of P
redmostí I in the central Moravian Plain has yielded a rich and diverse large mammal fauna dated around 25e27,000 14C years BP (ca. 29,500e31,500 cal BP). This fauna includes numerous carnivores (cave lion, wolf, brown bear, polar fox, wolverine) and herbivores (reindeer, large bovine, red deer, muskox, horse, woolly rhinoceros, woolly mammoth) whose trophic position could be reconstructed using stable isotopic tracking (d13C, d15N, d34S) of bone collagen (n ¼ 63). Among large canids, two morphotypes, “Pleistocene wolves” and “Palaeolithic dogs”, were considered, and two human bones attributed to the Gravettian assemblage of P
redmostí I were also sampled. The trophic system around the Gravettian settlement of P
redmostí I showed the typical niche partitioning among herbivores and carnivores seen in other mammoth-steppe contexts. The contribution of the analyzed prey species to the diet of the predators, including humans, was evaluated using a Bayesian mixing model (SIAR). Lions included great amounts of reindeer/muskox and possibly bison in their diet, while Pleistocene wolves were more focused on horse and possibly mammoth. Strong reliance on mammoth meat was found for the human of the site, similarly to previously analyzed individuals from other Gravettian sites in Moravia. Interestingly, the large canids interpreted as “Palaeolithic dogs” had a high proportion of reindeer/ muskox in their diet, while consumption of mammoth would be expected from the availability of this prey especially in case of close interaction with humans. The peculiar isotopic composition of the Palaeolithic dogs of P
redmostí I may indicate some control of their dietary intake by Gravettian people, who could have use them more for transportation than hunting purpose. © 2014 Elsevier Ltd and INQUA. All rights reserved.

Keywords: Collagen Dog Food web Gravettian Moravian Plain Stable isotopes

1. Introduction

* Corresponding author. Department of Geosciences, Universit€ at Tübingen, €lderlinstrasse 12, 72074 Tübingen, Germany. Ho E-mail addresses: [email protected] (H. Bocherens), dorothee. [email protected] (D.G. Drucker), mietje.germonpre@naturalsciences. ), [email protected], [email protected] (M. La zni
be (M. Germonpre ckova ), [email protected] (Y.I. Naito), christoph.wissing@uniGaletova tuebingen.de (C. Wissing), [email protected] (J. Br u
zek), moliva@ mzm.cz (M. Oliva).

Stable isotopes of carbon and nitrogen in bone collagen of late Pleistocene large mammals have provided valuable information on the structure of ecosystems in the frame of the so-called “mammoth-steppe”, especially regarding niche partitioning among herbivores and prey selection by their predators, including ancient humans (e.g. Bocherens, 2003; Drucker et al., 2003; Drucker and Henry-Gambier, 2005; Bocherens et al., 2005b, 2011a, b, 2013, 2014; Fox-Dobbs et al., 2008; Yeakel et al., 2013). In contrast to other regions of the mammoth-steppe studied so far,

http://dx.doi.org/10.1016/j.quaint.2014.09.044 1040-6182/© 2014 Elsevier Ltd and INQUA. All rights reserved.

Please cite this article in press as: Bocherens, H., et al., Reconstruction of the Gravettian food-web at P
redmostí I using multi-isotopic tracking (13C, 15N, 34S) of bone collagen, Quaternary International (2014), http://dx.doi.org/10.1016/j.quaint.2014.09.044

2

H. Bocherens et al. / Quaternary International xxx (2014) 1e18

such as south-western France, Belgium, south-western Germany and AlaskaeYukon, the Moravian Plain is characterized by a “mammoth culture”, where mammoth bone strongly dominate the assemblages (Svoboda et al., 2005; Wojtal and Sobczyk, 2005; re et al., 2009; Musil, 2010). So far, only punctual applications Bruge of these isotopic approaches have been performed in the Moravian Plain, the first one including only a handful of faunal samples from
stonice (Ambrose, 1998), another one with only a couple of Dolní Ve human specimens and no associated fauna (Richards et al., 2001), and one more complete isotopic study of faunal remains from the sector G in Milovice but with only three herbivorous and two an, 2001; Drucker et al., in press). Using the carnivorous species (Pe very rich and diverse fauna from P
redmostí I (Musil, 2010), the present study therefore aims at a more complete reconstruction of the trophic system in the Moravian Plain, with a focus on human subsistence and the trophic position of large canids. Indeed, some of these canids are considered to have been domesticated (e.g.,  et al., 2012, 2013; this volume), it is Benecke, 1994; Germonpre therefore of prime importance to test possible dietary differences between domestic and wild large canids, and their similarity to the diet of humans.

2. Material and methods 2.1. Site of P
redmostí I The site of P
redmostí I is located in the central Moravian Plain, in a region very rich in archaeological sites of Gravettian age (Fig. 1). The first archaeological excavations took place in the late 19th century under the direction of J. Wankel, K. Ma
ska, M. K
rí z, and continued during the 20th century under the direction of K.

Absolon, B. Klíma and J. Svoboda (reviews in Svoboda et al., 1994; Oliva, 2007; Velemínsk a and Br u
zek, 2008; d'Errico et al., 2011). P
redmostí I is a very important site for the Gravettian of Central Europe, which yielded an abundant lithic and bone industry (Valoch, 1982), portable art and ornaments (Valoch, 1975; Valoch zni
, 2009), as well as thousands of faunal and La ckov a-Galetova skeletal remains and several human skeletons, unfortunately
almost completely lost during the Second World War (Simek, 1948). Some radiocarbon dating were performed and the results range from 24,340 ± 120 BP to 26,870 ± 250 BP for samples from P
redmostí I (Svoboda, 2008). The central Moravian Plain where P
redmostí I is located has been sometimes considered to be a warm refuge during the late Pleistocene, with signs of arid and humid climatic fluctuations during the time of deposition of the archaeological layers (Musil, 2010). Based on the analysis of tooth cementum, it seems that the site of P
redmostí I was occupied all  Fi
sa kov year long (Nyvltova a, 2013). 2.2. Bone material The studied material was collected during the excavations that took place at the end of the 19th century and was stored in the Moravian Museum in Brno. We tried our best to obtain samples from five different individuals of each large mammal species, by choosing the same anatomical piece for one given species. However, this was not always possible especially in the case of rare species (Table 1). For the abundant species, such as mammoth and large canids (Musil, 2008, 2010), we took more than 5 specimens. The analyzed material from P
redmostí I belongs to the following species (n ¼ 63): reindeer Rangifer tarandus (n ¼ 5), large bovine cf. Bison (n ¼ 5), red deer Cervus elaphus (n ¼ 1), muskox Ovibos

́


v, 7 Jaro
sov, 8 Bor
sice, 9e10 Dolní Ve
stonice and Fig. 1. Map of the Moravian Gravettian. Captions: 1 P
redmostí, 2 Ostrava - Pet
rkovice, 3 Mlade
c III, 4 Brno 2, 5 Napajedla, 6 Spytihne Pavlov, 11 Milovice.

Please cite this article in press as: Bocherens, H., et al., Reconstruction of the Gravettian food-web at P
redmostí I using multi-isotopic tracking (13C, 15N, 34S) of bone collagen, Quaternary International (2014), http://dx.doi.org/10.1016/j.quaint.2014.09.044

H. Bocherens et al. / Quaternary International xxx (2014) 1e18

moschatus (n ¼ 3), horse Equus ferus (n ¼ 5), woolly rhinoceros Coelodonta antiquitatis (n ¼ 5), woolly mammoth Mammuthus primigenius (n ¼ 10), brown bear Ursus arctos (n ¼ 4), wolverine Gulo gulo (n ¼ 5), polar fox Alopex lagopus (n ¼ 6), cave lion Panthera spelaea (n ¼ 3), and large canids Canis cf. lupus (n ¼ 11). Following  et al. (2012, this volume) we distinguished amongst Germonpre large canid specimens two morphotypes (i.e. “Pleistocene wolves” versus “Palaeolithic dogs”), in order to clarify whether these two categories can be distinguished by their diet.

Table 1 List of species sampled in P
redmostí I, with the number of specimens, the NMI (Minimum Number of Individuals), and the number of specimens that yielded well preserved collagen. PRDM-53 (98-585-B-Pr 3) is the mandible associated with the  et al. (2012) see Table 3. Palaeolithic dog skull P
redmostí 3 identified in Germonpre Species Woolly mammoth (Mammuthus primigenius) Woolly rhinoceros (Coelodonta antiquitatis) Horse (Equus ferus) large bovine (Bos or Bison, cf. Bison) Muskox (Ovibos moschatus) Red deer (Cervus elaphus) Reindeer (Rangifer tarandus) Cave lion (Panthera spelaea) Brown bear (Ursus priscus) Wolverine (Gulo gulo) Polar fox (Alopex lagopus) Large canid “Pleistocene wolf” (Canis cf. lupus) Large canid “Palaeolithic dog” (Canis cf. familiaris) Human (Homo sapiens) Total

Specimens

MNI Isotopic values

9 4 5 5 3 1 5 3 4 5 6 7 4 2

6 2 3 5 2 1 5 1 1 5 6 7 4 2

5 2 5 5 2 1 4 2 1 3 4 4 2 1

63

49

41

In addition to the faunal samples, we also sampled two human mandibles, P
redmostí 26 (P26) and P
redmostí 30 (P30). The mandibular fragment P26 (Fig. 2) belongs to an adult individual, probably of the female sex (Vl
cek, 2008). This specimen was probably found in 1930 by L. Nov ak (Svoboda, 2008b: p.27), also prior to 1934, the date of publishing Matiegka's monograph (Matiegka, 1934). The P26 specimen is listed in the Catalogue of , 1999). The Fossil Hominids (Vl
cek, 1971; Jelínek and Orvanova other mandibular fragment P30 believed to be part of the Gravettian assemblage of P
redmostí I (Ullrich, 1996) belongs to a young adult individual (Br u
zek et al., 2008). Moreover, one human femur from P
redmostí I (P27) had been originally sampled and yielded collagen, but in the meantime a radiocarbon date proved that this bone was not of Gravettian age, but of the Late glacial period (OxA-

Fig. 2. Lateral view of the human mandible P26 from P
redmostí I sampled for isotopic analysis.

3

27382: 10,675 ± 45 14C BP). Its isotopic values have also been measured but they will not be discussed in the context of the Gravettian assemblage of P
redmostí I, they will only be used as comparison data. 2.3. Methods For each specimen, a small fragment was carefully sawn with a dremel rotating tool equipped with a circular diamond-coated blade, ultrasonicated in acetone and water, rinsed with distilled water, dried and crushed to a powder of 0.7 mm grain size (Bocherens et al., 1997). Then, an aliquot of around 5 mg was used to measure the nitrogen, carbon and sulphur content (%N, %C, %S) of the whole bone, in order to screen out samples with excessive collagen loss or extraneous carbon or sulphur contamination (Bocherens et al., 2005a). For instance, fresh bones contain 4% nitrogen while ancient bones with less than 0.4% nitrogen usually fail to yield good collagen (Bocherens et al., 2005a). The measurements were performed using a Vario EL III elemental analyser using Sulfanilic acid from Merck as internal standard. The mean standard errors were better than of 0.02%, 0.05%, and 0.03% for %C, %N and %S, respectively. To evaluate the amount of non-collagenic carbon in the bones, we subtracted from the whole amount of carbon in the bone the estimated amount of carbon linked to collagen, calculated as %N  2.7, 2.7 being the mass ratio of carbon on nitrogen in collagen (Bocherens et al., 2005a). If the bone only suffered collagen loss without addition of exogenous collagen, it is expected that the non-collagenic carbon equals around 1.4%, which is the carbon amount linked to the carbonate fraction of the bone bioapatite (Bocherens et al., 2005a, 2008). The collagen was purified according to a well established protocol (Bocherens et al., 1997). The elemental and isotopic measurements were performed at the Geochemical unit of the Department of Geosciences at the University of Tübingen (Germany), using an elemental analyser NC 2500 connected to a Thermo Quest Delta þ XL mass spectrometer. The elemental ratios C/N, C/S and N/S were calculated as atomic ratios. The isotopic ratios are expressed using the “d” (delta) value as follows: d13C ¼ [(13C/12C)sample/(13C/12C)reference e 1]  1000‰, d15N ¼ [(15N/14N)sample/(15N/14N)reference e 1]  1000‰, and d34S ¼ [(34S/32S)sample/(34S/32S)reference e 1]  1000‰. The international references are V-PDB for d13C values, atmospheric nitrogen (AIR) for d15N values, and CDT for d34S values. Measurements were normalized to d13C values of USGS24 (d13C ¼ 16.00‰) and to d15N values of IAEA 305A (d15N ¼ 39.80‰). The reproducibility was ±0.1‰ for d13C measurements and ±0.2‰ for d15N measurements, based on multiple analyses of purified collagen from modern bones. Samples were calibrated to d34S values relative to CDT of NBS 123 (d34S ¼ 17.10‰), NBS 127 (d34S ¼ 20.31‰), IAEA-S-1 (d34S ¼ 0.30‰) and IAEA-S-3 (d34S ¼ 21.70‰). The reproducibility is ±0.4‰ for d34S measurements, and the error on amount of S measurement is 5%. The reliability of the isotopic signatures of the extracted collagen was addressed using their chemical composition (%C, %N, % S and C/N, C/S, N/S ratios). These values must be similar to those of collagen extracted from fresh bone to be considered reliable for isotopic measurements and radiocarbon dating. Several studies have shown that collagen with atomic C/N ratios lower than 2.9 or higher than 3.6 is altered or contaminated, and should be discarded, as well as extracts with %N < 5% (e.g., DeNiro, 1985; Ambrose, 1990). For sulphur, %S should be ranging from 0.12 to 0.20% and 100 < N/S < 300 and 300 < C/S < 900 (Nehlich and Richards, 2009; Bocherens et al., 2011b). Cluster analysis is regularly used in modern ecological studies using stable isotopes of animal tissues (e.g. Davenport and Bax,

Please cite this article in press as: Bocherens, H., et al., Reconstruction of the Gravettian food-web at P
redmostí I using multi-isotopic tracking (13C, 15N, 34S) of bone collagen, Quaternary International (2014), http://dx.doi.org/10.1016/j.quaint.2014.09.044

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H. Bocherens et al. / Quaternary International xxx (2014) 1e18

€dt et al., 2002; Riccialdelli et al., 2010; Hobson et al., 2012; Hücksta 2012; Madigan et al., 2012; Miller et al., 2013) but not so far in similar investigations using fossil material. In the present study, the pattern of distribution of individuals from herbivorous species on the one hand and carnivorous species on the other hand based on their d13C and d15N values was determined through a cluster analysis using the Ward's minimum variance method performed on stable carbon and nitrogen isotopic composition, with the software SAS JMP version 10.0, in order to identify trophic groups. The relative contribution of different prey to the average diet of predators, including humans, was simulated using a Bayesian mixing model approach performed in the SIAR package (Parnell et al., 2010), using the R software, version 3.0.2 (R Core Team, 2013). In previous isotopic investigations of the diet of Paleolithic humans, mixing models such as IsoSource (Phillips and Gregg, 2003) were used (e.g. Drucker and Henry-Gambier, 2005; Bocherens et al., 2005b, 2013). The advantages of SIAR are the possibility to incorporate uncertainty in input data, and to yield not only a range of possible dietary proportion, but also to provide their true probability distribution (Parnell et al., 2010). Using data from previous works comparing the d13C and d15N values of the bone collagen of predators and that of their prey in modern and ancient terrestrial ecosystems (Bocherens and Drucker, 2003; Fox-Dobbs et al., 2007), we calculated a Trophic Enrichment Factor (TEF) of þ1.1 ± 0.2‰ and þ3.8 ± 1.1‰ for d13C and d15N values, respectively (Table 2).

Therefore, all the analyzed bones have incorporated some exogenous carbon during the diagenetic processes in P
redmostí I, possibly from carbonate concretions (Musil, 2010). The amount of sulphur in bone (%S) ranges from 0.11% to 0.43% with an average value of 0.20 ± 0.06% when all analyzed samples are considered (Table 3). There is no statistically significant correlation between %Cncoll and %S (%S ¼ (0.0034  %Cncoll) þ 0.1837, r2 ¼ 0.007). Even mammoth bones (%S ¼ 0.21 ± 0.11%) do not exhibit higher %S than any other species, which range from 0.16 ± 0.02% (polar fox) to 0.27 ± 0.05% (reindeer). 3.2. Chemical and isotopic composition of collagen The range of %C, %N and C/N values was large in the collagen extracted from the bones from P
redmostí I (Table 3). Among the 63 sampled specimens, 22 had chemical characteristics outside the range of fresh collagen, i.e. %N < 6% or C/N > 3.6 (Fig. 5) and therefore will not be considered in the following discussion. Among the 24 collagen that could be analyzed for sulphur isotopes, 14 yielded %S that were lower than 0.13%, which is the lowest amount found in collagen extracted with the same method from fresh bones (Bocherens et al., 2011b). Most of these collagen also exhibit C/S higher than 900, and some of them also C/N higher than 300, these values being the upper limits for collagen considered to yield biogenic isotopic composition (Nehlich and Richards, 2009; Bocherens et al., 2011b).

Table 2 Summmary of the trophic enrichment factors between predator collagen and prey collagen used to calculate the Trophic Enrichment Factor (TEF) used for SIAR. Predator-prey

Tissue

Age

Origin

Dd13C

Wolf-prey Lynx-prey Wolf-prey Wolf-deer Coyote-deer Wolf-caribou Carnivore-herbivore Wolf-herbivores Wolf-herbivores Lynx-rabbit Modern average 1sd Archaeological average 1sd Modern þ archaeological average 1sd

Collecoll Collecoll Collecoll Collecoll Collecoll Collecoll Collecoll Collecoll Collecoll Collecoll

Modern Modern Modern Modern Modern Modern Modern Ca 20 kyr BP Ca 15 kyr BP Ca 12 kyr BP

Bialowieza Forest (Poland) Bialowieza Forest (Poland) Isle Royale (Canada) Ontario (Canada) Ontario (Canada) Alaska Africa Les Jamblancs (France) St-Germain-la-Riviere (France) Pont d'Ambon (France)

1.0 1.1 1.3 e e 1.2 e 1.3 0.9 0.8 1.2 0.1 1.0 0.3 1.1 0.2

SD

Dd15N 3.5 4.0 4.6 2.9 2.7 2.4 4.8 5.5 4.4 2.8 3.6 0.9 4.2 1.4 3.8 1.1

SD

0.7 0.9 0.9

Reference Bocherens and Drucker (2003) Bocherens and Drucker (2003) Fox-Dobbs et al. (2007) Schwarcz 1991 Schwarcz (1991) Szepanski et al. (1999) Ambrose and DeNiro (1986) Bocherens and Drucker (2003) Bocherens and Drucker (2003) Bocherens and Drucker (2003)

3. Results

3.3. Isotopic results of bone collagen according to species

3.1. Chemical composition of the bones

Globally the d13C and the d15N values range from 21.8 to 18.3‰ and from 1.3 to 12.3‰, respectively (Table 3). The herbivorous species present d13C ranging from 21.8 to 18.7‰, while the predatory species present d13C ranging from 19.8 to 18.3‰. The d15N values of herbivorous species range from 1.3 to 9.8‰, while those of predatory species range from 6.2 to 10.7‰. The carnivorous species exhibit d15N higher than those of herbivorous species except mammoth, but the d13C values of the mammoths are lower than those of any carnivores, and there is therefore no overlap between herbivorous and carnivorous species. The human mandible P26, the only Gravettian human bone to provide reliable results, presents a d13C value of 19.4‰ and a d15N value of 12.6‰, the highest d15N value measured in the whole assemblage. In contrast, the human femur of Lateglacial age presents lower d13C and d15N values (d13C ¼ 20.1‰; d15N ¼ 9.7‰). The d34S values for collagen with reliable chemical composition range from 1.3 to

Nitrogen content in bones (%N) from P
redmostí I exhibited a wide range of values, from 0.24% to 3.02% (Table 3). Among the samples analyzed, eight had %N values lower than 0.4% and therefore have lost more than 90% of their original collagen. These samples that have suffered from heavy collagen loss are scattered in different species and there is no obvious difference among the species regarding this chemical parameter (Fig. 3). The non-collagenic carbon (%Cncoll) in the bones also exhibited a wide range of variation, ranging from 2.5 to 8.5% (Table 3, Fig. 4). Among species, there is a clear trend for bones of larger species, such as mammoth and woolly rhinoceros, to have higher %Cncoll than species of smaller size (Fig. 4). In any case, even the lowest % Cncoll values are clearly higher than those expected for bones loosing their collagen without addition of exogenous carbon (1.4%).

Please cite this article in press as: Bocherens, H., et al., Reconstruction of the Gravettian food-web at P
redmostí I using multi-isotopic tracking (13C, 15N, 34S) of bone collagen, Quaternary International (2014), http://dx.doi.org/10.1016/j.quaint.2014.09.044

d13C

d15N

%S

N/S

d34S

3.4 3.6 3.5 3.5

18.9 19.3 18.8 18.9

2.8 2.7 2.6 3.9

0.14

738

218

0.44

0.13

797

228

0.76

2.4 13.6 13.2 0.8 9.0

4.1 3.3 3.4 7.2 3.8

(19.3) 18.6 18.6 (20.3) (19.8)

(6.4) 9.4 8.1

0.15 0.14

681 702

206 209

1.88 0.33

(8.6)

37.0 34.9 15.6 31.9 2.8 37.4 37.6

13.2 11.4 4.5 10.5 0.6 13.5 12.8

3.3 3.6 4.0 3.6 5.6 3.2 3.4

19.8 19.3 (19.0) 19.4 (22.0) 18.9 21.0

9.0 8.8 (7.5) 7.4 (4.4) 9.0 5.6

0.10

908

255

(0.02)

0.12 0.11

825 945

254 276

1.28 (0.09)

68.7

36.1

12.4

3.4

20.8

5.2

0.07

1343

397

(1.69)

0.20

60.4

38.3

13.1

3.4

20.4

5.1

0.09

1081

318

(6.39)

3.1 3.0 4.2 2.7 3.2 3.7 3.3 3.1 3.0 5.4 7.3 3.0 2.5 3.3 3.1 2.8 2.9 2.7 3.3 3.3 4.3 8.1

0.26 0.23 0.16 0.18 0.15 0.14 0.18 0.15 0.34 0.27 0.16 0.21 0.16 0.16 0.13 0.18 0.19 0.19 0.15 0.17 0.14 0.13

41.8 67.6 17.3 39.5 17.6 12.2 25.5 28.5 25.5 31.0 34.1 58.6 129.8 59.0 9.0 50.6 52.7 29.4 12.8 12.0 9.8 10.9

35.7 41.1 25.5 39.0 32.4 28.5 28.9 40.1 3.3 2.2 27.9 41.1 41.6 37.5 34.9 38.4 40.3 37.5 34.6 36.8 16.0 10.2

12.3 14.5 7.7 13.6 11.3 8.1 10.3 14.6 0.6 0.2 9.1 14.5 15.1 12.9 12.0 13.1 14.5 12.3 11.4 12.6 4.5 2.3

3.4 3.3 3.8 3.4 3.3 4.1 3.3 3.2 6.6 11.9 3.6 3.3 3.2 3.4 3.4 3.4 3.2 3.6 3.5 3.4 4.1 5.2

20.8 20.7 (19.8) 19.5 19.8 (19.0) 19.6 19.7 (22.6) (22.1) 20.5 20.3 20.7 19.6 19.6 20.2 20.3 20.9 19.1 18.7 (19.6) (21.3)

3.7 5.0 (8.1) 6.4 8.0 (6.2) 7.0 6.2 (4.5)

0.08 0.12

1160 928

342 281

(2.64) (1.02)

0.15 0.14

706 603

211 181

0.98 6.26

0.14

780

243

6.34

0.12 0.13 0.11

926 858 947

280 267 280

(1.74) 2.83 (2.16)

0.12 0.11 0.14

868 948 696

253 293 196

3.67 (1.72) 0.19

13.5

7.5

0.43

20.2

22.0

7.1

3.6

21.5

8.0

1.1

8.7

5.8

0.15

60.5

36.5

12.8

3.3

21.1

7.7

1.3

11.3

7.9

0.23

4.8

7.1

1.4

5.8

(23.3)

(5.9)

1.1

9.6

6.7

0.20

29.4

9.1

2.9

3.6

(21.3)

(6.2)

Lab no

Species

Piece

Side

Remark

%Nbone

%Cbone

%Cncoll

%Sbone

Yield (mg g1)

%C

%N

PRDM-1 PRDM-2 PRDM-3 PRDM-4 PRDM-5 PRDM-6 PRDM-7 PRDM-8 PRDM-9 PRDM-10 PRDM-11 MPU-32 PRDM-12 PRDM-13 PRDM-14 PRDM-15 PRDM-16 PRDM-17

Rangifer tarandus Rangifer tarandus Rangifer tarandus Rangifer tarandus Rangifer tarandus Panthera spelaea Panthera spelaea Panthera spelaea Ursus arctos Ursus arctos Ursus arctos Ursus arctos Gulo gulo Gulo gulo Gulo gulo Gulo gulo Gulo gulo Equus ferus

Humerus (distal end) Humerus (distal end) Humerus (distal end) Humerus (distal end) Humerus (distal end) Mandible Femur (distal end) Humerus (distal end) Mandible Mandible Mandible Mandible Femur (distal end) Femur (distal end) Femur (distal end) Femur (distal end) Femur (distal end) Femur (distal end)

R R R R R R R? L? L L L

5 s individuals 5 s individuals 5 s individuals 5 s individuals 5 s individuals May not be s individuals May not be s individuals May not be s individuals 3 s individuals 3 s individuals 3 s individuals

1.1 0.5 2.2 0.9 0.3 0.9 1.0 1.5 0.3 0.8 0.3 1.3 0.5 0.5 0.5 0.9 1.5 1.1

8.1 8.2 9.0 7.2 6.0 5.7 6.0 6.8 5.6 8.1 5.3 7.1 6.8 6.3 5.5 8.1 7.8 6.2

5.2 6.8 3.1 4.7 5.1 3.4 3.3 2.8 4.7 5.8 4.6 3.6 5.3 4.9 4.1 5.6 3.8 3.1

0.31 0.29 0.32 0.22 0.22 0.27 0.18 0.21 0.20 0.18 0.16 0.16 0.14 0.14 0.16 0.30 0.17 0.28

41.2 11.5 62.1 16.5 n.a. 3.8 38.8 53.3 23.4 39.3 n.a. 31.9 29.8 21.3 15.4 14.5 56.3 30.2

37.7 19.5 40.1 35.7

13.0 6.3 13.4 11.9

8.3 38.6 37.9 5.1 29.2

PRDM-18

Equus ferus

Femur (distal end)

1.3

8.7

5.2

0.15

PRDM-19

Equus ferus

Femur (distal end)

1.6

8.2

3.8

PRDM-20 PRDM-21 PRDM-22 PRDM-23 PRDM-24 PRDM-25 PRDM-26 PRDM-27 PRDM-28 PRDM-29 PRDM-30 PRDM-31 PRDM-32 PRDM-33 PRDM-34 PRDM-35 PRDM-36 PRDM-37 PRDM-38 PRDM-39 PRDM-40 PRDM-41

Equus ferus Equus ferus Alopex lagopus Alopex lagopus Alopex lagopus Alopex lagopus Alopex lagopus Alopex lagopus Coelodonta antiquitatis Coelodonta antiquitatis Coelodonta antiquitatis Coelodonta antiquitatis bovini cf. Bison bovini cf. Bison bovini cf. Bison bovini cf. Bison bovini cf. Bison Cervus elaphus Ovibos moschatus Ovibos moschatus Ovibos moschatus Mammuthus primigenius

Humerus (distal end) Humerus (distal end) Femur (diaphysis) Femur (diaphysis) Femur (diaphysis) Femur (diaphysis) Femur (diaphysis) Femur (diaphysis) Mandible Mandible Femur (prox. End) Tibia Metapodium (dist.) Metapodium (dist.) Metapodium (diaph.) Metapodium Metapodium Mandible Calcaneum Calcaneum Femur (distal end) Astragalus

1.1 2.1 0.8 1.6 0.9 0.9 0.9 1.1 1.4 1.6 1.0 2.2 3.0 1.6 0.4 1.3 1.2 1.3 0.6 0.5 0.4 0.7

6.1 8.6 6.3 6.9 5.5 6.0 5.7 6.1 6.9 9.8 10.0 8.8 10.7 7.6 4.1 6.2 6.0 6.1 4.8 4.5 5.3 10.1

PRDM-42

Mammuthus primigenius

Astragalus

2.2

PRDM-43

Mammuthus primigenius

Astragalus

PRDM-44

Mammuthus primigenius

Astragalus

PRDM-45

Mammuthus primigenius

Astragalus

R R L L L L R L R

R

5 s individuals 5 s individuals 5 s individuals 5 s individuals 5 s individuals 3 s individuals, same side as next 3 s individuals, same side as previous 3 s individuals, opposite side but s morphology May not be s individuals may not be s individuals 2 s individuals 2 s individuals 4 s individuals 4 s individuals 4 s individuals 4 s individuals 2 s individuals, 1931, bigger 2 s individuals, 1928, smaller Exc. 1926 Exc. 1934 Bigger Smaller

Complete but fragile Piece taken on mandible, not teeth 2 s individuals 2 s individuals Epiphyse not fused 4 s individuals, big 1926, s from last 2 4 s individuals, big 1924, s from last 2 4 s individuals, medium 1926, s from last 2 4 s individuals, medium 1924, s from last 2

C/N

3.9 5.0 3.3 3.3 4.2 6.8 5.3 2.7 1.6 1.3 (2.7) (7.4)

C/S

H. Bocherens et al. / Quaternary International xxx (2014) 1e18

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Table 3 List of analyzed material, chemical and isotopic results. Figures in italics indicate values outside the range for sufficiently well preserved collagen. Isotopic values in brackets are considered unreliable due to anomalous chemical composition and were not discussed any further.

(continued on next page) 5

3.15 560 0.18

167

(1.28) 216 743

9.9 (5.8) 8.9 10.7 (10.0) 12.6 9.7 8.7 0.9 6.9 10.5 4.2 10.6 13.2 26.8 4.2 21.2 31.0 16.5 32.5 38.1

3.6 5.2 3.6 3.4 4.6 3.6 3.4

19.4 (20.8) 18.7 19.5 (20.3) 19.4 20.1

0.11

190 9.0 10.2 31.1

3.6

19.5

0.10 (10.4) 7.8 8.9 (19.3) 18.7 18.3 3.7 3.4 3.3 6.5 11.1 13.4 20.4 32.1 37.5

9.2 9.7 45.9 n.a. 16.1 n.a. n.a. 19.9 10.5 21.5 23.9 39.7 41.6 26.2 0.18 0.18 0.19 0.29 0.16 0.12 0.11 0.20 0.32 0.17 0.16 0.18 n.d. 0.23 3.8 3.0 3.4 3.9 5.1 3.5 4.9 3.5 4.1 3.6 2.6 7.6 n.d. 4.5 5.5 5.5 7.1 4.6 6.3 4.3 5.7 6.7 6.6 5.7 5.7 9.5 n.d. 7.53 0.6 0.9 1.4 0.2 0.5 0.3 0.3 1.2 0.9 0.8 1.1 0.7 n.d. 1.1

8.2 21.8 3.5 8.4 25.2 0.0 n.d. n.d. n.d. n.d.

675

965

627 0.16 9.8 20.7 n.d. n.d.

n.d.

n.d.

21.9

38.6

13.9

3.2

9.6 20.9 n.d. n.d.

n.d.

n.d.

8.7

23.3

7.6

3.6

7.3 21.2 3.6 9.8 30.1 20.9 0.15 8.5 9.9

Not totally sure if s individual from next Not totally sure if s individual from previous Sample taken from one growth layer in ivory Sample taken from one growth layer in ivory Sample taken from one growth layer in ivory 97-584-B-93 97-584-F-Pr9 98-585-B-Pr3 98-585-B(3) 97-583-E-P(3) 97-583-F-94 97-584-A-16 97-584B-Pr1928(2) 97-584-F-99 97-584-F-Pr1924(1) 97-584-F P30 P26 P
r 27 (OxA-27382: 10,675 ± 4514C BP)

0.5

0.12

297

3.53 194

d34S N/S C/S %S

d15N d13C C/N %N %C Yield (mg g1) %Sbone %Cncoll %Cbone %Nbone

Canis lupus “dog” Canis lupus “dog” Canis lupus “dog” Canis lupus “dog” Canis lupus “wolf” Canis lupus “wolf” Canis lupus “wolf” Canis lupus “wolf” Canis lupus “wolf” Canis lupus “wolf” Canis lupus “wolf” Homo sapiens Homo sapiens Homo sapiens PRDM-48 PRDM-49 PRDM-53 PRDM-54 PRDM-55 PRDM-56 PRDM-57 PRDM-58 PRDM-60 PRDM-61 PRDM-63 PRDM-69 PRDM-70 PRDM-68

Mandible Mandible Mandible Mandible Mandible Mandible Mandible Mandible Mandible Mandible Mandible Mandible Mandible Femur

Mammuthus primigenius TSK-II 7

Tusk

Mammuthus primigenius TSK-II 3

Tusk

Mammuthus primigenius TSK-II 1

Tusk

Mammuthus primigenius PRDM-46

Astragalus

Species Lab no

Table 3 (continued )

Piece

Side

Remark

3.20

H. Bocherens et al. / Quaternary International xxx (2014) 1e18

(2.51)

6

6.3‰, with most values between 0 and 3.7‰. There are not enough results to perform statistical tests on the differences in d34S values among species, but some species, such as reindeer and cave lion, seem to have lower d34S values than others, such as large bovines, mammoths, canids and polar foxes. The pattern of distribution of individuals from herbivorous species on the one hand and carnivorous species on the other hand based on their d13C and d15N values was determined through a cluster analysis. The specimens from herbivorous species cluster in four groups, one of them (cluster H4) includes only mammoths, and all the mammoths, another one (cluster H1) includes all reindeer, all muskoxen and some large bovines, and the rest of the specimens are within the two remaining clusters, one cluster (H2) including herbivores with relatively low d15N values, i.e. the red deer, one large bovine, one horse and one rhinoceros, and one cluster (H3) with herbivores with relatively higher d15N values, i.e. the remaining horses and one rhinoceros. Interestingly, the large bovines are the most widespread in this cluster analysis, since the specimens from this taxon are spread in 3 of the 4 clusters. Among the carnivores, one cluster (C4) well separated from the rest of the specimens includes all dogs and cave lions, together with one wolf and one wolverine. The rest of the carnivores can be separated into three additional clusters, one of them (C3) including the human sample and two wolves, another one (C1) with most polar foxes and one wolverine, and finally a third one (C2) including members of different species such as polar fox, brown bear, wolverine and wolf. The wolverines are scattered among the three of the four clusters (C1, C2 and C4), as well as the wolves (C2, C3 and C4). In contrast, both cave lions and both Palaeolithic dogs are included into the same cluster, C4. Among these carnivores, the only predators included into the same cluster as the human specimen (C3) are two wolves (Fig. 7 and Fig. 8). The differences in d13C and d15N values among species were tested statistically, using a non-parametric Wilcoxon test with the software JMP 10. In the statistical analysis were taken into account only those species with at least three specimens, therefore brown bear, red deer, musk ox, woolly rhinoceros and human, were not included in the test. The results of the statistical tests are given in Table 4. Among the main possible prey species, horse, large bovine, reindeer, and mammoth, each is significantly different from all three others for d13C and/or d15N values (Table 4). The one red deer exhibits d13C and d15N values within the range of those of reindeer, the woolly rhinoceros d13C and d15N values are close to those of the large bovine, and the muskox presents similar d13C values than reindeer but slightly lower d15N values. Among the predators, wolf has the highest average d15N value (9.6 ± 0.87‰) and polar fox the lowest (6.9 ± 0.80‰) but statistical comparisons could not be performed with other predators due to the small number of samples with collagen for most species. The human specimen exhibits the highest d15N value of all the measured specimens (12.6‰). 3.4. Proportions of consumed prey calculated using SIAR For each predator, the possible proportions of prey species as well as those of herbivores that belong to different clusters were calculated using SIAR, first considering each predator species as a group, then testing each individual separately (Fig. 9, Fig. S5 to S9). To test the sensitivity of our calculations to possible uncertainties in Trophic Enrichment Factor (TEF), we simulated the effect of changing the TEF for nitrogen from 3.6 to 4.2 and the TEF for carbon from 0.8 to 1.4 (Fig. S1 to S4). This sensitivity analysis shows that the proportion of mammoths in human diet was not affected by changes in the TEF of nitrogen, while the change in carbon TEF could result in ~20% change (median) in the proportion of mammoth. Using the cluster of the individuals that belong to

Please cite this article in press as: Bocherens, H., et al., Reconstruction of the Gravettian food-web at P
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Table 4 Results of non-parametric Wilcoxon statistical tests for d13C and d15N values for species with at least three specimens with reliable collagen.

Canis lupus Gulo gulo Alopex lagopus Rangifer tarandus Bos or Bison Equus ferus M. primigenius

n

d13C/d15N

Wolverine

Polar fox

Reindeer

Bovine

Horse

Mammoth

4 3 4 4 5 5 6

Av ± sd (19.2 ± (19.2 ± (19.6 ± (19.0 ± (20.1 ± (20.8 ± (21.2 ±

Gulo gulo (NS/NS)

Alopex lagopus (NS/0.0304) (NS/NS)

Rangifer tarandus (NS/0.0304) (NS/0.0518) (0.0294/0.0304)

Bos or Bison (0.0189/0.0195) (0.0358/0.0358) (NS/NS) (0.0189/NS)

Equus ferus (0.0189/0.0200) (0.0358/0.0369) (0.0195/0.0200) (0.0189/0.0373) (0.0269/NS)

M. primigenius (0.0139/NS) (0.0282/NS) (0.0142/NS) (0.0139/0.0142) (0.0102/0.0080) (NS/0.0081)

0.39/9.6 0.23/8.4 0.12/6.9 0.25/3.0 0.40/4.6 0.20/4.9 0.40/8.4

± ± ± ± ± ± ±

0.87) 0.88) 0.80) 0.63) 1.48) 0.7) 0.74)

Values in bold stand for statistically significant differences.

Fig. 3. Nitrogen content in bones from P
redmostí I.

Fig. 4. Non-collagenous carbon content in bones from P
redmostí I. The dotted line (1.4%) represents the amount of carbon of the carbonate fraction.

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H. Bocherens et al. / Quaternary International xxx (2014) 1e18

Fig. 5. Cluster analysis of specimens from herbivorous species based on their d13C and d15N values and sorted according to their d15N values.

herbivorous species allowed us to include specimens of species with too few individuals to be included in the dietary analysis based on prey species. The main results of these analyses are described below. For cave lions, the contribution of each prey was relatively even, with a median around 14e20% though a slightly higher contribution of reindeer and muskoxen could be possible (Fig. 9). Among clusters, the cluster H1 (made up essentially of these two species) showed the highest contribution (33%). One cave lion (PRDM-7) included less muskoxen than the other cave lion (PRDM-8) (Fig. S5). Wolverines as a group may have included a bit of every type of prey in their diet with possibly less reindeer and muskoxen as

opposed to cave lions (Fig. 9). However, contributions of the clusters H3 (essentially horses) and H4 (mammoths) were negatively correlated (r ¼ 0.55), meaning that if the contribution of one type of prey is increasing, the contribution of the second one must decrease. Two individuals (PRDM-12 and 16) were similar, with high mammoth contribution, and a lower contribution of the other prey species, while another individual (PRDM-14) was more focused on large bovine, rhino, horse, and less on the other prey species (Fig. S6). Palaeolithic dogs were similar to cave lions, they included in their diet more reindeer and muskoxen than any other herbivore prey (Fig. 9). Contributions of the clusters H1 (essentially reindeer and muskoxen) and H3 (essentially horses) were negatively correlated (r ¼ 0.52). One Palaeolithic dog (PRDM-49) included more reindeer and muskoxen, while the other Palaeolithic dog (PRDM-53) included more reindeer, large bovine, and muskoxen (Fig. S7). Pleistocene wolf included relatively high amount of mammoth (median ¼ 28%), but much less reindeer and muskoxen (Fig. 9).

Fig. 6. d13C and d15N values of herbivorous species with grouping of the values according to species and to the cluster defined through the cluster analysis (Fig. 5).

Fig. 7. Cluster analysis of specimens from omnivorous and carnivorous species based on their d13C and d15N values and sorted according to their d15N values.

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collagen samples that could be analyzed had a chemical composition indicating extensive diagenetic alteration for sulphur and therefore the d34S values of these samples could not be used in the paleobiological discussion. However, even with a limited dataset, some grouping occured when considering d34S values, especially between cave lions and reindeer, and similar d34S values were found in mammoths and wolves (Fig. 11). Some polar foxes exhibited higher d34S values than the rest of the samples, which may indicate different foraging areas or an access to different food items than the rest of the animals. These sulfur isotopic values are still too limited to provide independent evidence of the trophic link between these predators and their prey, but they do not contradict the conclusions of the carbon and nitrogen isotopic compositions and they may indicate differences in foraging territories. Interestingly, the human femur dated to a more recent period, coeval to the Younger Dryas, exhibit similar d34S values than the terrestrial fauna from the Gravettian times, and Neolithic human individuals from the Moravian Plain are also in a similar range of d34S values (Zvelebil et al., 2012). 4. Discussion 4.1. Diagenetic conditions in P
redmostí I 13

15

Fig. 8. d C and d N values of omnivorous and carnivorous species with grouping of the values according to species and to the cluster defined through the cluster analysis (Fig. 7).

Contributions of the clusters H3 (essentially horses) and H4 (mammoths) were negatively correlated (r ¼ 0.59). Three wolves (PRDM-55, 58 and 63) were rather similar, with mammoths as the most prominent prey (median ¼ 27e36%), and may have included few reindeer and muskoxen (median < 8%). A fourth wolf (PRDM61) was clearly different, with reindeer (median ¼ 21%) and large bovine (median ¼ 17%) and muskoxen (median ¼ 18%) having higher contributions relative to those of the other individuals (Fig. S8). The polar fox relied more on horse and rhino (Fig. 9). The cluster H2 showed the highest contribution (median ¼ 43%). The H2 cluster contribution was negatively correlated with those of H3 (r ¼ 0.55) and H4 (r ¼ 0.61). PRDM-24 individual were different from the others in that reliance on mammoth was higher (median ¼ 25%) than those of the other polar foxes (median ¼ 7e13%) (Fig. S9). Brown bear was similar to wolf in terms of prey consumption with high amount of mammoth (median ¼ 33%) and low muskoxen contribution (median ¼ 4%), as expected from its similar isotopic values (Fig. 9). Contributions of the clusters H4 and H3 were negatively correlated (r ¼ 0.54). The human specimen relied heavily on mammoths (median ¼ 56%) (Fig. 10). The mammoth contribution was negatively correlated with that of large bovine (r ¼ 0.64). Contributions of the cluster H4 and H3 were also negatively correlated (r ¼ 0.71). If canids were included in the spectrum of possible prey, then wolf may have represented 13% (median) of the consumed prey, decreasing the amount of mammoth (median ¼ 41%). Contributions of these two species were negatively correlated (r ¼ 0.49). This relationship was also illustrated by the negative correlation between the cluster H4 and C4 (r ¼ 0.50). 3.5. Sulphur isotopic composition Unfortunately, several collagen extracted from the bones could not be analyzed for sulphur isotopes due to too small quantities, lower than the 5 mg necessary for this analysis. Moreover, several

The diagenetic alteration looks very variable among the bones from P
redmostí I. The loss of collagen is very variable and it does not seem to be related to the species to which the bone belonged. In contrast, the amount of exogenous carbon is higher in bone of the largest species, mammoth and woolly rhinoceros, than in bones from smaller species. This could be linked to slower burial for large bones than for small ones, and therefore a longer weathering process for these large bones. Mammoth bones were used as building material of settlement (Oliva, 1988, 2009), and this special use could be part of the reason why these bones were containing more exogenous carbon than the bones of other species, not involved in this process. The sulphur amounts in the bones are similar to those of rock-shelter sites, higher than those of cave sites and much lower than those of bones from fluvial sediments (Bocherens et al., 2011b). Moreover, the pattern of sulphur amount in bones does not differ significantly according to the species or the size of the bones. Altogether, the diagenetic status of the Gravettian bones from P
redmostí I is quite variable and requires caution before interpreting the obtained results as reflecting palaeobiological features. However, when the proper reliability criteria are taken into account, it is possible to screen out the specimens that can be considered further in the study. 4.2. Niche partitioning among herbivores and distinction of possible prey The d13C and d15N values of the different herbivorous species exhibit a similar distribution in the carbon/nitrogen isospace than in other areas of the mammoth steppe ecosystem. The relatively high d13C values of reindeer have been reported in previous studies in France (Fizet et al., 1995; Drucker et al., 2000, 2003; Bocherens, 2003; Drucker and Henry-Gambier, 2005; Bocherens et al., 2005b, 2011a, b; Richards et al., 2008), in Belgium (Bocherens et al.,  et al., 2009), in Germany (Stevens 2001, 2011a, b, 2013; Germonpre et al., 2009), as well as in Siberia (Iacumin et al., 2000) and in AlaskaeYukon (Fox-Dobbs et al., 2008). This feature has been linked to the consumption of lichen by this ungulate species (Fizet et al., 1995; Bocherens, 2003; Drucker et al., 2003, 2012), a type of plant that has systematically higher d13C values than vascular plants (e.g. Barnett, 1994; Fizet et al., 1995). Even if the difference is not statistically significant because of the low number of specimen

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Fig. 9. Prey contributions in the diet of the predators from P
redmostí I calculated using SIAR represented as Violin-plots. These plots show the proportional prey contributions to predators' diets by species or by cluster. The mammoth data include both bone and ivory collagen data. Black boxes and whiskers show the median with 1st and 3rd quartiles and ranges with 1.5 times length of the interquartile range above the 3rd quartile or below the 1st quartile, respectively. The shaded area indicates the Kernel density plot of the probability density of prey proportions.

analyzed, muskoxen also exhibit relatively high d13C values compared to other herbivores, as it has been reported in late  et al., 2009). This could indicate Pleistocene Belgium (Germonpre that during the late Pleistocene of Europe, muskoxen used lichen in the same way as modern reindeer, although modern muskoxen do not compete with reindeer for lichen in winter but rather select sedges and mosses (e.g. Klein, 1992; Ihl and Barboza, 1997; Larter and Nagy, 2004; Skarin, 2004), and moss does not exhibit the 13 C-enrichment seen in lichen compared to vascular plants (e.g. Teeri, 1981; Galimov, 2000). The low d15N values of muskox compared to all other herbivorous species is probably linked to the consumption of leaves rather than grass, since shrubs and trees

have systematically lower d15N values due to their association with mycorrhizae (e.g. Schulze et al., 1994; Michelsen et al., 1998; Kristensen et al., 2011). Among the remaining herbivorous species, large bovines (Bos/ Bison) present d13C values intermediate between those of reindeer, muskoxen and red deer, which are higher than 19‰, and those of horse, woolly rhinoceros and mammoth, which are lower than 20‰ on average, with a larger spread of values between individuals than in other herbivorous species. Such a pattern is commonly found in other mammoth steppe fauna studies performed in France (Drucker et al., 2003), in Belgium (Bocherens et al., 2001, 2011a, b), in Germany (Stevens et al., 2009), in Siberia

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Fig. 10. Prey contributions in the diet of the human from P
redmostí I (P26) calculated using SIAR, shown as Violin-plots. These plots show the proportional prey contributions to human diet. Graphs A to D indicate the proportions when only herbivorous species are included as possible prey (A), with herbivorous, omnivorous, and carnivorous species (B), according to clusters among herbivorous species (C), according to cluster among herbivorous, omnivorous and carnivorous species (D). Mammoth data include both bone and ivory collagen data.

(Iacumin et al., 2000) and in AlaskaeYukon (Fox-Dobbs et al., 2008). In P
redmostí I, the isotopic compositions of the large bovines are more scattered than those of other herbivorous species, suggesting some heterogeneity in their ecology. This could be due to the failure to recognize specimens belonging to Bison or to Bos, which may have had different ecological habits (Musil, 2010), but previous studies where the isotopic composition of both taxa in the same sites were measured did not show any significant differences (e.g. Bocherens et al., 2005b, 2011a, b). The fact that the large bovines from P
redmostí I cluster with different other herbivorous species could reflect individual ecological habits of the analyzed

Fig. 11. d13C and d34S values of collagen from herbivores and carnivores from P
redmostí I.

specimens, and could indicate that individuals from large bovines were dwelling in the different habitats available around the prehistoric site. Moreover, differences also occur in d15N values among these herbivorous species. Especially mammoths exhibit d15N values higher than those of all other herbivorous species. The mammoth specimen with the lowest d15N value (7.3‰) is still higher than any representative of all the other herbivorous species. These differences are interpreted as reflecting different plant selection and possibly differences in the digestive physiology (Bocherens, 2003). The consumption of plants with higher d15N values by mammoths in comparison with other herbivores seems to be the main reason for this pattern (Bocherens, 2003). By comparison with recent plants in boreal and arctic environments, grass and graminoids would be good candidates (e.g. Michelsen et al., 1998). Interestingly, forbs have also been suggested based on ancient DNA analysis of mammoth coprolites and stomach content (Willerslev et al., 2014) and indeed forbs also exhibit relatively higher d15N values in comparison to shrubs for instance (Michelsen et al., 1998; Miller and Bowman, 2002; Yi and Yang, 2006). The results obtained on the herbivorous species from P
redmostí I confirm the ecological stability of the mammoth steppe across its geographical expansion and shows that the Gravettian people were part of this ecosystem. Deviations from this isotopic pattern among large herbivores could indicate shifts in the ecosystem functioning, due to climate fluctuations or human impact (Drucker et al., in press; this volume). Mammoths analyzed in P
redmostí I exhibit significant variations of their d15N values, with higher d15N values in ivory compared to bone, and variations within a single tusk. Such patterns can be detected in previous works with similar set of isotopic values (e.g. Richards et al., 2012). Moreover, variations in d15N values of mammoth individuals reaching 3‰ have been described within tusks (e.g. Gohman et al., 2010), as well as hair (Iacumin et al., 2006), reflecting seasonal changes of diet composition. In addition, d15N values of juvenile mammoth tissues are higher than

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H. Bocherens et al. / Quaternary International xxx (2014) 1e18

those of adults, due to the influence of milk consumption, which is one trophic level higher than adult diet (Rountrey et al., 2007; Bocherens et al., 2013). To calculate an average d15N value of mammoth meat as available to predators including humans, we chose to average all values, from bone and ivory, to incorporate isotopic variations occurring in mammoth tissues, especially young specimens that are often found in the bone assemblage (Oliva, 1997). The cluster analysis of herbivores allowed us to define 4 groups including members of one or several species (Figs. 5 and 6). These clusters seem to correspond to different ecological preferences related to choice of plant food. The cluster H1, including all reindeer, all muskoxen and some large bovines includes specimens with d13C values higher than 20‰. Such a pattern has been linked to the consumption of lichen (e.g. Fizet et al., 1995; Bocherens, 2003). The members of this cluster also exhibit d15N values in the lower range of values for herbivores, a feature that is consistent with the consumption of shrubs rather than grass (Bocherens, 2003). The three other clusters are mostly distinct due to their d15N values, with cluster H2 having the lowest and H4 the highest d15N values, H3 having intermediate d15N values (Fig. 5). This pattern looks similar to the pattern of d15N values in vascular plants in modern boreal and arctic environments, where shrubs have the lowest d15N values and grass and forbs the highest. Therefore the isotopic differences between clusters seem to be linked to ecological preferences of individuals. In some cases, all specimens from one species belong to the same cluster, such as reindeer, muskoxen and mammoths, but in other cases, specimens of one taxon are scattered among different clusters, suggesting different ecological preferences within individuals of this taxon. Due to the isotopic distinctiveness of different herbivorous species or clusters of individuals with similar ecological preferences, it is possible to use d13C and d15N values as tracers of prey consumption. Depending on the relative proportion of meat coming from each potential prey, the different predators will incorporate a mixture of C and N atoms with different isotopic composition, and their own tissues will integrate the isotopic composition of their average diet. The context is therefore favorable to the determination of prey consumption by different predators in P
redmostí I.

than dog, and that even if the contribution of mammoth by the human decreases, this megaherbivore remains clearly the most consumed prey species (Fig. 12). The analyzed wolverines from P
redmostí I are variable in their diet, as reflected by their scattering in the cluster analysis. When we consider the members of this species as a group, their diet may include on average similar amount of each available prey, but different individuals exhibit different amounts of each prey in their diet, indicating some individual preferences since bone collagen averages food consumed during long time periods. Evidence that some wolverine ate significant amounts of mammoth indicates the possibility to access carcasses and scavenge them, since this relatively small carnivore could not kill a mammoth by itself. Moreover, although recent wolverines are known to be able to kill large ungulates such as reindeer, they rely essentially on scavenging (e.g. van Dijk et al., 2008; Dalerum et al., 2009). This scavenging of large carcasses by wolverine could have been favored by the absence of cave hyenas, which were already extirpated from central Europe around 28,000 years ago (Stuart and Lister, 2014). The relatively high amount of mammoth in the diet of the brown bear may also be the result of scavenging, as modern brown bears are known to scavenge ungulate carcasses killed by other predators, especially for prey species of large size (e.g. Mattson, 1997; Vulla et al., 2009). In contrast, Palaeolithic dog and cave lion relied more on reindeer and muskoxen than on mammoth and other herbivores (Figs. 12 and 13). The results of sulphur isotopic analysis are consistent with this conclusion. For cave lion, this result is consistent with previous isotopic investigations of this predator in Belgium and southewest Germany between around 40,000 and 12,000 years ago that concluded that most analyzed cave lions were preferentially feeding on reindeer (Bocherens et al., 2011a). In the sector G of Milovice, a Gravettian site younger in age than P
redmostí I, a similar isotopic pattern was obtained for cave lion an, 2001; Drucker et al., in press), suggesting that this strong (Pe trophic link between cave lion and reindeer was probably a general feature in the Moravian Plain during the Gravettian.

4.3. Prey preference of predators The results of the cluster analysis of carnivores already suggested that some predators had more similar prey choices than others (Figs. 7 and 8). This is for instance the case for Palaeolithic dogs and cave lions, while humans are clustering with two Pleistocene wolves. This is not what was predicted under the assumption that dogs were fed by humans and fed with the same kind of food that the humans themselves used to eat (e.g. Guiry, 2012). However, there are many occurrences where such a dog feeding pattern is not applied (see discussion below). The results of the SIAR indicate that for Palaeolithic dogs and cave lions, reindeer and muskoxen were among the most consumed prey. In contrast, the human specimen and the two wolves that cluster with it have mammoth as primary prey. Large bovines are the second most important prey kind, especially when canids are not included in the human diet. Both prey for the human are strongly negatively correlated, so if one prey increases in the diet, the other one has to decrease. Canids were tested using SIAR in the human diet since  some cut-marks were found on large canid bones (Germonpre et al., 2012) and to evaluate if the consumption of these carnivores by humans could explain the high d15N values of humans rather than mammoth consumption. The results show that, when canids are included in the range of prey, it is mostly wolf rather

Fig. 12. Percentage of prey contribution (as median value) for the different predators in P
redmostí I, with two situations in the case of the human: one were only herbivorous species were consumed and another one including canids (Palaeolithic dog and Pleistocene wolf) as possible prey.

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redmostí I using multi-isotopic tracking (13C, 15N, 34S) of bone collagen, Quaternary International (2014), http://dx.doi.org/10.1016/j.quaint.2014.09.044

H. Bocherens et al. / Quaternary International xxx (2014) 1e18

13

Fig. 13. Schematic representation of the trophic relationships of herbivores and their predators in P
redmostí I. (a) represents the situation assuming that humans did not consume wolf meat, (b) represents the situation considering that wolf meat was consumed in addition to the meat of herbivorous species.

4.4. Mammoth as food resource Mammoth meat was consumed in high quantities by humans, as well as by some predators and scavengers, including some wolves, some wolverines, one polar fox (PRDM-24) and the analyzed brown bear (Figs. 12 and 13). One possible explanation for such a pattern is that mammoths suffered from high mortality in Moravia and were scavenged by at least some of the predators mentioned above, especially wolverines, polar fox and brown bears that were unlikely to actively hunt the mammoths. The cause of this mortality of mammoth is not straightforward to establish. Natural mortality is unlikely since mammoths, like modern elephants, are long-lived animals with a low reproduction rate and a low mortality rate (Wittemyer et al., 2013). Mortality could be due to accidents related to a difficult landscape that would lead to accidental mortality of the mammoths (Bosch, 2012), or to active hunting (Svoboda et al., 2005). Hunting is the favored hypothesis in view of the age profile of the mammoth, dominated by young female cows (Oliva, 1997). The results of the present isotopic analyses are fully consistent with mammoth meat being the staple food of the P
redmostí people, as suggested by several authors (Absolon and  and Br Klíma, 1977; Oliva, 1997; Velemínska u
zek, 2008; Br u
zek et al., 2008). Since the collagen reflects an average of the protein fraction of the diet of an individual during several years, it means that this amount of mammoth meat was regularly consumed and that the large accumulation of mammoth remains was not only an artefact due to the long period of deposition. It means that the bones of mammoths were not just collected from old carcasses and only used as construction material, but that mammoth meat was also a staple food of Gravettian humans in Moravia. In addition to humans, some scavenging predators probably also took advantage of this anthropogenic subsidy in the form of mammoth carcasses, among them brown bears, wolverines, polar fox and Pleistocene wolves, species within which some individuals could rely heavily on mammoth meat. Furthermore, it must be underlined here that the Pleistocene wolves, thus the canids with elongated mandibles and well-spaced premolars, ate more mammoth than the Palaeolithic dogs, that is the morphotype characterized by a shorter muzzle and crowded premolars. It has been suggested that the canids that utilized the mammoths present on Gravettian sites were the ones that developed dog-like features (e.g. Fladerer, 2001; Crockford and Kuzmin, 2012). However, the present study suggests

that the presence of dog-like features in large canids is not related to a diet rich in mammoth meat. In order to confirm this trend, more individuals from both morphotypes need to be analyzed. 4.5. Implications for the status of canids The domestication of dogs is expected to have modified their diet in comparison to wild wolves. One expectation is that dog's diet would be more similar to human diet, since humans could supply their dogs with a fraction of their own food (e.g. Waguespack, 2002), leading to similar isotopic composition in dogs and humans (e.g. Guiry, 2012). This is documented with isotopes in different cases (Losey et al., 2011), also when the wild canid is not a wolf but a fox (Rick et al., 2011), but many such studies did not actually test the difference between wolves and dogs (e.g. Guiry, 2012). In fact, some cases show that wolves and dogs have similar d13C and d15N values, which are different from those of humans, as in the Mesolithic sites of the Iron Gates, in the Danube valley (Bori c et al., 2004). Furthermore, in several hunter-gatherer societies, it has been observed that humans feed their dogs with items different from their own preferred food. In some cases this food must be kept away from dogs or the dogs receive what their owners do not appreciate much. For the Tareumiut (Point Barrow, Alaska), the whale formed the keystone of their economic, social and religious live. Whale, seals and caribou were their preferred food. The whales were communally hunted and the meat was distributed between participating crews according to well-defined regulations and ceremony. Their dogs could not eat any part of the whale, as this was considered offensive to the whale, although they could be fed the viscera from a freshly killed whale. Walrus meat, a meat unpopular with this people, was given to the dogs. The dogs received further parts of game not destined for human consumption such as entrails. They also were fed fish, blubber and reindeer droppings (Spencer, 1959). The Koyukon (Alaska) use mainly moose and caribou for food; the black and brown bears are also significant in their subsistence economy. The latter animals possess potent spirits, which are easily provoked and must be properly treated. The Koyukon dogs are mostly fed fish; bear meat is strictly tabooed for dogs (Nelson, 1983). The diet of the Evenki from Northern Transbaikal consists mainly of deer meat and fish. Their dogs are fed with specially cooked dog food composed of old meat, hooves

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H. Bocherens et al. / Quaternary International xxx (2014) 1e18

with tendons and sinews. The dogs receive also intestines, reindeer blood and periosteum (Abe, 2005). The staple food of the Maritime Chukchee is meat from sea mammals and fish. Their dogs are fed mainly with intestines and blubber of sea mammals, they receive only little meat and not much fish (Bogoras, 1904). Based on the short review above, it can be concluded that it is not because an animal is staple food for humans that dogs will automatically receive its meat. Restrictions can be present for several reasons. It cannot be excluded that also at P
redmostí a taboo existed so that the living dogs could not have access to mammoth meat. Such differences between the diet of humans and that of their dogs have been documented in archaeological sites. In several instances, dogs from archaeological sites have isotopic values significantly different from those of coeval humans (e.g. Chu, 1998; Losey et al., 2013; Tsutaya et al., 2014). At P
redmostí, the dogs and the human have different isotopic composition, reflecting different proportions of prey consumption. It will be interesting to analyze the stable isotopic composition of palaeolithic dogs from other sites to see if such a dietary pattern was widespread in space and time or if the Gravettian of P
redmostí was a special case maybe linked to the diversity and abundance of available food resources. In the case of P
redmostí, we can provide some insights on the use of dogs based on the dietary reconstruction. One of the first possible use of dog is helping in hunting, for instance in immobilizing large herbivores (e.g. Fiedel, 2005; Shipman, 2012). In such a case, it is difficult to imagine that these dogs would not have access to a part of the meat of the prey. The P
redmostí dogs do not seem to have consumed high amounts of mammoth meat, in contrast to their human owners, the median of the mammoth meat contribution in the Palaeolithic dog diet is around 15%, compared to 41e56% for humans (Fig.12). If they were helping in hunting mammoths, they could have received a minimum amount of mammoth tissues at the kill site, but if such hunting events did not occur often, they must have been prevented access to mammoth meat the rest of the time. Most of the time, the Palaeolithic dogs were probably fed by humans a kind of meat that themselves did not eat much, especially reindeer, and this has several implications.

First of all, it means that the dogs were not allowed to move freely in the settlement, and they were probably tied up. One has to remember that these early dogs were as big as wolves, and probably as dangerous. Also, ethnographic data show that the arctic and subarctic peoples that use their dogs for transport usually keep them tied up, in contrast with those northern people who utilize their dogs as hunting aids: those dogs are free ranging (Abe, 2005:  et al. (2012) suggested that the dog-like canids of p.42). Germonpre P
redmostí played an important role in helping to solve the transportation problems of the Gravettian peoples, and the conclusion about their diet is consistent with this hypothesis. 4.6. Implications for human subsistence pattern in the Gravettian In P
redmostí, we have for the first time a stable isotopic investigation that combines human and coveal fauna, including a large diversity of herbivorous and carnivorous mammal species. This allows us not only to reconstruct the possible diet of the Gravettian humans, but also to replace them into their trophic system. The pattern that emerges is that of an ecosystem with a taxonomical and ecological diversity of herbivores that were preyed upon by numerous predatory species. Some species exhibit restricted ecological preferences, such as mammoth, reindeer, muskox and cave lion, while others are more flexible with individuals having different preferences, such as large bovines and wolverines. The humans were clearly trophically connected to the mammoths, but with no detectable impact on the ecology of the mammoths, indicating that despite the important consumption of mammoths by humans and some scavengers, the mammoth population was apparently not profoundly affected by this consumption. Younger sites of the Gravettian in the Moravian Plain have yielded some isotopic data that can be compared to those of P
redmostí, although they are less complete. Humans from Dolní
stonice and Brno 2-Francouzska  can be compared with isotopic Ve data from Milovice G fauna, since all these findings are dated around 22e23 kyr 14C BP (Richards et al., 2001). The pattern is very similar to that of P
redmostí, with each species represented in both

Fig. 14. d13C and d15N values of Gravettian humans, compared to those of herbivores of the same period.

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H. Bocherens et al. / Quaternary International xxx (2014) 1e18

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Table 5 List of Gravettian humans with d13C and d15N values. Site

Country

d13C

d15N

Labcode

14

sd

Reference

P
redmostí I 2 Brno-Francouzska
stonice 35 Dolní Ve Muerii 1 Muerii 2 Cioclovina 1 Sunghir 3 Sunghir 2 Sunghir 1 Paviland 1 Paviland 1 Paviland 1 Eel Point 1 La Rochette 1 Arene Candide IP

Czech Republic Czech Republic Czech Republic Romania Romania Romania Russia Russia Russia UK UK UK UK France Italy

19.4 19 18.8 19.3 19.1 19.6 19.6 19 19.5 18.4 18.4 18.4 19.7 17.1 17.6

12.2 12.3 12.3 12.3 12.4 12.7 11 11.2 10.7 10.4 9.3 10.2 11.4 11.2 12.4

OxA-8293 OxA-8292 OxA-15529 OxA-16252 OxA-15527 OxA-9038 OxA-9037 OxA-9036 OxA-16412 OxA- 8025 OxA-16413 OxA-14164 OxA-11053 OxA-10700

23,680 22,840 29,930 29,110 28,510 24,100 23,830 22,930 28,870 25,840 29,490 24,470 23,630 23,440

±200 þ990/904 ±170 ±190 ±170 ±240 ±220 ±200 ±180 ±280 ±210 ±110 ±130 ±190

This paper Pettitt and Trinkaus (2000), Richards et al. (2001) Richards et al. (2001) Soficaru et al. (2006), Trinkaus et al. (2009) Soficaru et al. (2006), Trinkaus et al. (2009) Soficaru et al. (2007), Trinkaus et al. (2009) Dobrovolskaya et al. (2012) Richards et al. (2001) Dobrovolskaya et al. (2012) Jacobi and Higham (2008) Richards et al. (2001) Jacobi and Higham (2008) Schulting et al. (2005) Richards (2009) Pettitt et al. (2003)

sites, P
redmostí and Milovice G, having similar isotopic composition, indicating a similar niche partitioning and a similar prey choice for the predators. Especially the human specimens had also a high amount of mammoth meat in their average diet, suggesting the stability of the ecosystem of the Moravian Plain during the Gravettian period. The only indication of a possible change in Milovice G is the isotopic nitrogen composition of one of the horses that is similar to those of the mammoths, and this could indicate that the ecological niche occupied by the mammoth may have been less densely occupied due a demographic decline of the proboscideans (Drucker et al., in press). In Europe, there is evidence for a large isotopic diversity of Gravettian humans, with d13C values ranging from 19.7 to 17.1‰ and d15N values ranging from 9.3 to 12.4‰ (Table 5). Due to the relatively stable pattern of isotopic variation in the main prey species available to prehistoric populations at the European scale, it is possible to link this isotopic variability with a diversity of subsistence strategies that can be compared to that of the Moravian Plain (Fig. 14). Some Gravettian humans from Romania exhibit d13C and d15N values similar to those of the Moravian specimens (Soficaru et al., 2006; Trinkaus et al., 2009), which could reflect a similar high amount of mammoth meat in their diet. In contrast, several human individuals from Russia (Sunghir: Richards et al., 2001; Dobrovolskaya et al., 2012), and Great-Britain (Paviland, Eel Point: Schulting et al., 2005; Jacobi and Higham, 2008) have lower d15N values than their Moravian and Romanian counterparts, possible indicating a diet lower in mammoth meat consumption but still with a protein source dominated by terrestrial ungulates. Some individuals deviate from this pattern, such as one individual from Italy (Arene Candide: Pettitt et al., 2003) and one from southwestern France (La Rochette: Richards, 2009). Their less negative d13C values and higher d15N values point to the integration of some marine food resources, either directly from the sea, as in Arene Candide, or through the migration in rivers of anadromous salmon, as in La Rochette. It seems therefore that the Gravettian populations were diverse in their food procurement depending on the different environmental context they lived in (Musil, 2011). 5. Conclusion The present study has shown that, despite variable state of preservation of bone collagen, it has been possible to perform a detailed reconstruction of the trophic relationships in P
redmostí I, including humans and dogs. The isotopic values of large herbivores are consistent with a typical mammoth steppe ecosystem with no sign of deviation from the pattern observed from western Europe to

C B.P.

Alaska during this period. The abundance of mammoth had an impact on the diet of several predators, such as wolves, wolverines and brown bears, while others such as cave lion and Palaeolithic dogs relied more on reindeer and muskoxen. The Gravettian humans relied heavily on mammoth meat, even if the occasional consumption of canid meat cannot be ruled out. The different diet of the Palaeolithic dogs suggests that they were mostly used for transportation and not primarily for helping in hunting, but this trend should be confirmed by analyzing more individuals of Palaeolithic dogs from P
redmostí. Acknowledgements This work is dedicated to the memory of our dear colleague Prof. Karel Valoch (1920e2013). Thanks are due to S. Ali, T. Himpel, C. Must, B. Steinhilber and H. Taubald for assistance in the sample preparation and isotopic measurements. We are thankful to Martina Pacher for her help with the mandible of brown bear from P
redmostí (MPU-32). Appendix A. Supplementary data Supplementary data related to this article can be found at http:// dx.doi.org/10.1016/j.quaint.2014.09.044. References Abe, Y., 2005. Hunting and Butchering Patterns of the Evenki in Northern Transbaikalia, Russia (Ph.D. dissertation). Department of Anthropology, Stony Brook University. €hren. In: Fontes Absolon, K., Klíma, B., 1977. P
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