Palaeolithic dog skulls at the Gravettian Předmostí site, the Czech Republic PDF

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Palaeolithic dog skulls at the Gravettian Předmostí site, the Czech Republic Mietje Germonpré

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Journal of Archaeological Science 39 (2012) 184e202

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Journal of Archaeological Science journal homepage: http://www.elsevier.com/locate/jas

Palaeolithic dog skulls at the Gravettian P
redmostí site, the Czech Republic Mietje Germonpré a, *, Martina Lázni
cková-Galetová b, Mikhail V. Sablin c a

Department of Palaeontology, Royal Belgian Institute of Natural Sciences, Vautierstraat 29, 1000 Brussel, Belgium Moravian Museum Anthropos Institute, Zelnytrh 6, 65937 Brno, The Czech Republic c Zoological Institute RAS, Universitetskaya nab. 1, 199034 Saint-Petersburg, Russia b

a r t i c l e i n f o

a b s t r a c t

Article history: Received 10 June 2011 Received in revised form 16 September 2011 Accepted 17 September 2011

Whether or not the wolf was domesticated during the early Upper Palaeolithic remains a controversial issue. We carried out detailed analyses of the skull material from the Gravettian P
redmostí site, Czech Republic, to investigate the issue. Three complete skulls from P
redmostí were identified as Palaeolithic dogs, characterized by short skull lengths, short snouts, and wide palates and braincases relative to wolves. One complete skull could be assigned to the group of Pleistocene wolves. Three other skulls could not be assigned to a reference group; these might be remains from hybrids or captive wolves. Modifications by humans of the skull and canine remains from the large canids of P
redmostí indicate a specific relationship between humans and large canids. Ó 2011 Elsevier Ltd. All rights reserved.

Keywords: Gravettian P
redmostí Dog (Canis familiaris) Wolf (Canis lupus) Domestication Skull Canine

1. Introduction P
redmostí, in the Czech Republic, is a very well-known Gravettian site. Excavated since the 1880s, it is famous for its large human skeletal assemblage composed of several dozen individuals. Unfortunately most of these human remains were destroyed in a fire during the Second World War (Velemínská and Br u
zek, 2008). Furthermore, P
redmostí is renowned for its huge amount of mammoth remains, from more than 1000 individuals (Musil, 2008). It is also an exceptional canid site, delivering more than 4000 bones of this taxon (Pokornỷ, 1951). As early as 1894, Ma
ska mentioned in his diary the finding of a complete canid skeleton, possibly from a Palaeolithic dog (Ma
ska, 2008, p. 185). That the dog originated in the Late Glacial is a view that is widely accepted among archaeologists (Morey, 2010 and references herein). However, a recent study has shown that the domestication of the wolf might have begun much earlier, during the Aurignacian (Germonpré et al., 2009). On the assumption that the hypothesis of an Aurignacian origin is correct, there is an hiatus between the oldest Aurignacian Goyet dog and the putatively second oldest dogs, the Epigravettian Eliseevichi dogs (Sablin and Khlopachev,

* Corresponding author. Tel.: þ32 2 627 44 64; fax: þ32 2 627 41 13. E-mail addresses: [email protected] (M. Germonpré), [email protected] (M. Lázni
cková-Galetová), [email protected] (M.V. Sablin). 0305-4403/$ e see front matter Ó 2011 Elsevier Ltd. All rights reserved. doi:10.1016/j.jas.2011.09.022

2002). Recently, Benecke (1994) argued that the P
redmostí site provides some indications of wolf domestication, using as a basis an analysis of the lower jaws that were found at the site. In addition, several papers on DNA analysis of recent dogs have suggested that the dog originated earlier than the Late Glacial (Vilà et al., 1997; Lindblad-Toh et al., 2005). In this context, the Gravettian P
redmostí site is critical for confirming that the domestication of the wolf started long before the end of the Last Glacial. Therefore, we studied the P
redmostí large canid skulls to determine whether some skulls could be identified as Palaeolithic dogs, thereby corroborating the hypothesis of an early origin for the dog. For this study, we also included unidentified large canid skulls from the Gravettian (Avdeevo) and the early Upper Paleolithic (Kostenki 17). 2. Sites 2.1. P
redmostí P
redmostí is located in the Moravian Corridor, Czech Republic (Fig. 1). This open air site is situated on a gentle south-oriented slope in the valley of the Be
cva. Originally, it extended to the north, west and south of the Skalka rock, a limestone outcrop and occurred in loess deposits that were exploited from the 19th century but have since been mined away, resulting in an almost complete destruction of the site (Oliva, 1997; Svoboda, 2008a,b; Svoboda et al., 1994). The first archaeological investigations of the

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M. Germonpré et al. / Journal of Archaeological Science 39 (2012) 184e202

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Desna

Moscow

NORTH SEA

Danube

EA

NS

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Fig. 1. Map showing the most important sites discussed in the text. 1, P
redmostí; 2, Avdeevo; 3, Kostenki 17; 4, Goyet; 5, Dolní Vĕstonice; 6, Eliseevichi; 7, Mezin; 8, Mezhirich.

site were undertaken by Wankel in 1880 and were followed by a long series of archaeological excavations, including those of Ma
ska, until the beginning of the 20th century. Excavations of the remaining parts of the site were conducted between 1943 and 1992, and resumed in the beginning of the 21st century. In 2006, a museum pavilion that was constructed to protect the last remnants of the cultural layer, was opened to the public (Svoboda, 2008b). Velemínská and Br u
zek (2008) give an extensive review of the excavation history. Two Gravettian cultural layers have been recognized, of which the lower one delivered most of the human and animal material. The Gravettian occupations took place during the warmer episodes of the Pleniglacial that preceded the last glacial maximum. The main Gravettian horizon can be considered to be composed of separate settlement units that are probably not contemporaneous but were occupied in the period 26,000e27,000 y BP (Svoboda et al., 1994). P
redmostí yielded a rather small number of portable art objects, including two representations of human females engraved on mammoth bones (d’Errico et al., 2011). The human skeletal remains were identified as anatomically modern humans (Velemínská and Br u
zek, 2008). The funerary assemblage indicates a long-term use of the burial area, located to the west of the Skalka rock, with consecutive interments. Most of the adult assemblage is composed of young people. The funerary assemblage contains many children u
zek and Velemínská, 2008). Mammoth scapulae covered at (Br least three human skeletons at the margins of the burial area (Svoboda, 2008b). Nývltová-Fi
sáková (2007) claimed that the Gravettian people probably lived at P
redmostí year round, basing this conclusion on the increment structures of animal teeth. The mammal assemblage is dominated by the mammoth (73.8% of the Minimum Number of Individuals). Remains from more than 1000 mammoths were discovered (Oliva, 1997; Musil, 2008). The age profile is dominated by young females aged from 20 to 30 years. This suggests a selective mortality, which is probably due to a selective hunting of inexperienced female mammoths (Oliva, 1997). Mammoth meat was probably the staple food for the P
redmostí people (Absolon and Klíma, 1977; Oliva, 1997). Remains from other large herbivores such as horse, bison and reindeer, are present in low frequencies (Pokornỷ, 1951). Large canids are the second most abundant group in the Minimum Number of Individuals (MNI), representing 7.6% of the

individual mammals present. The total number of identified specimens (NISP) is 4143 and the MNI is 103 (Pokornỷ, 1951). Ma
ska, when excavating the northern zone in 1894, discovered seven or eight complete wolf skeletons, but unfortunately, their skulls were broken (Absolon and Klíma, 1977; Svoboda, 2008b). Furthermore, in the same year, Ma
ska found within the human burial zone a complete skeleton of a dog or small wolf (Ma
ska, 2008, p. 185). This configuration is somewhat reminiscent of the finding of a dog skeleton in a Middle Holocene hunteregatherer cemetery at the site of Shamanka, Siberia (Losey et al., 2011). Unfortunately, the exact spatial distribution of each skeletal element of the P
redmostí material is not known, so we have no information on the exact position in which the analysed skulls were found within the human burial zone. Also in the same year, Ma
ska excavated three canid mandibulae. He concluded in his notes that one was probably from a dog (Absolon and Klíma, 1977). Ma
ska’s conjecture that dogs were present at P
redmostí was not taken up by later researchers. Pokornỷ (1951) distinguished two size groups: Canis lupus major and Canis lupus minor. Musil (2000) followed Pokornỷ’s (1951) subdivision of the large canid material and explained the size difference in terms of the sexual dimorphism of wolves. Benecke (1994) analysed the lower jaws. He discovered that tooth crowding occurs frequently in this material, much more than in recent wolves (Benecke, 1994, table 2: 21.3% vs 4.2%). He suggests that, given that tooth crowding can be present in captive wolves, at least a part of the P
redmostí large canids lived under conditions created by humans. Benecke (1994) also discovered that polyodonty in the P
redmostí lower jaws occurs much more commonly than in wild wolf populations. According to Benecke (1994), this could be an indication of a first genetic isolation of the P
redmostí canids from the Pleniglacial wild wolf population and therefore a first step in the domestication of the wolf. 2.2. Avdeevo Avdeevo is a Gravettian open air site located in the Russian Plain, in the Sejm valley near the city of Kursk and is dated between 20,000 and 21,000 BP (Fig. 1). It is well known for its human female and animal figurines, including images of wolves. The fauna consists especially of mammoth bones. The presence of a large

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number of remains from arctic fox and wolf suggests that the inhabitants of the site hunted for fur. Furthermore, perforated canines from these two canids were recovered in substantial numbers. In addition, numerous decorated metapodial bones from large canids and imitations of these bones carved out of mammoth ivory have been discovered (Gvozdover, 1995). A complete skull of a large canid (GIN RAS 911) was analysed by Germonpré et al. (2009) and is restudied here.

2.3. Kostenki 17 Kostenki 17 is one of several Kostenki-Borshchevo sites located on the second terrace along the west bank of the River Don, near Voronezh on the Russian Plain (Fig. 1). Culture Level II is found below a layer of volcanic ash, the Campanian Ignimbrite Y tephra, dated to 41,000e38,500 BP (Holliday et al., 2007). This level yielded an artifact assemblage that has been assigned to the early Upper

Table 1 List of the complete skulls of the large canids used in this study; RBINS, Royal Belgian Institute of Natural Sciences, Brussels; Forestia, Parc animalier, Theux, Belgium; GIN RAS, Geological Institute Russian Academy of Science, Moscow; MAE RAS Museum of Anthropology and Ethnography (Kunstkamera), Russian Academy of Science, Saint-Petersburg; MZM, Moravian Museum, Anthropos Institute, Brno; PM NASU, Palaeontological Museum National Academy of Science of Ukraine, Kiev; ULB, Université libre de Bruxelles, Belgium; ZIN RAS, Zoological Institute of the Russian Academy of Science, Saint-Petersburg; Herd., Herding; Guard., Guarding; D., Dog.

Unknown cases Type/site P
redmostí Kostenki 17 36233 Avdeevo 911 F1 wolf-Husky hybrid Zoo wolves 3356, 14730 Zoo wolf Han-sur-Lesse Zoo wolf Thor, Belgium Female Russian wolves

Age

Institute

Gravettian Early Upper Palaeolithic Gravettian Recent Recent Recent Recent Recent

MZM ZIN RAS GIN RAS ZIN RAS RBINS ULB Forestia ZIN RAS

Reference

n

Total

7 1 Germonpré et al. (2009) Germonpré et al. (2009)

1 1 2 1 1 6 20

Reference groups Recent Eurasian wolves (RW) Recent Eurasian wolves

Recent

ZIN RAS, RBINS

Germonpré et al. (2009)

35 35

Pleistocene Eurasian wolves (PlW) Pleistocene Eurasian wolves

Pleistocene

RBINS, PM NASU, ZIN RAS

Germonpré et al. (2009)

4 4

Palaeolithic dog (PalD) Site Goyet Eliseevichi 447

Aurignacian Epigravettian

RBINS MAE RAS

Eliseevichi 23781

Epigravettian

ZIN RAS

Mezin 5490

Epigravettian

PM NASU

Mezhirich 4493

Epigravettian

PM NASU

Germonpré et al. (2009) Germonpré et al. (2009), Sablin and Khlopachev (2002) Germonpré et al. (2009), Sablin and Khlopachev (2002) Benecke (1987), Germonpré et al. (2009), Pidoplichko (1998) Benecke (1987), Germonpré et al. (2009), Pidoplichko (1998)

1 1 1 1 1 5

Prehistoric small Dog (PrsD) Site Saint-Thibaud Senckenberg Bedburg

Epipalaeolithic Mesolithic Mesolithic

Chaix (2000) Degerbøl (1961) Street (1989)

1 1 1 3

Recent archaic dog (RDa) Breed Chow Chow (Northern Breed) Husky (Northern Breed)

Recent Recent

RBINS ZIN RAS

Germonpré et al. (2009) Germonpré et al. (2009)

3 15 18

Recent other dog (RDo) Breed Malinois (Herding Dog) German Shepherd (Herd. Dog) Rottweiler (Guardian Dog) Great Dane (Guardian Dog) Mastiff (Guarding Dog) Tibetian Mastiff (Guard. Dog) Doberman Pinscher (Guard. D.) Irish Wolfhound (Hound) Grand total

Recent Recent Recent Recent Recent Recent Recent Recent

RBINS RBINS RBINS RBINS, ZIN RAS RBINS RBINS RBINS RBINS

Germonpré Germonpré Germonpré Germonpré Germonpré Germonpré Germonpré Germonpré

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(2009) (2009) (2009) (2009) (2009) (2009) (2009) (2009)

4 13 2 6 2 1 4 2 34 119

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Palaeolithic. Bone points, awls, and some worked ivory were recovered (Praslov and Rogachev, 1982). The fauna is dominated by remains of wolf (70%) (Sablin, 2007). A complete large canid skull (ZIN RAS 36233) of Culture Level II is included in this study. 3. Material and methods 3.1. Skulls Here, we compare European Pleistocene canid skulls with recent and fossil Eurasian grey wolves, and recent and fossil dogs. All reference groups contain only adult skulls. Morphological and genetic studies have shown that the sole ancestor of the dog is the grey wolf (Benecke, 1987; Morey, 1992, 2010; Pang et al., 2009; Savolainen et al., 2002; Vilà et al., 1997; vonHoldt et al., 2010). Our Pleistocene wolf group is limited to four skulls: one from Belgium (Trou des Nutons), two from the Ukraine (Mezin, Mezhirich) and one from Siberia (Anabar) (Germonpré et al., 2009). Studies on the genetics of recent Eurasian wolves revealed that they can be genetically split in two main clades: haplogroups 1 and 2. These haplogroups do not show any clear allopatric distribution but cooccur in Eurasia (Pilot et al., 2010). Furthermore, analysis of ancient DNA has shown that Pleistocene European wolves, including the Trou des Nutons wolf used here, all belong to haplogroup 2 (Pilot et al., 2010; Stiller et al., 2006), which indicates a continuation to the present day of at least a part of the fossil European wolf populations. Given that all the ancient haplotypes were closely related to contemporary ones, Pilot et al. (2010) conclude that ancient and contemporary wolves represent the same population in different time periods. Their conclusion lends credence to the view that the recent wolf group we use as reference are representative of the Eurasian Pleistocene wolves. Except for one Belgian specimen, which dates from the 1850s, the group of Recent Eurasian wolves consists only of specimens from Russia, from different populations from the Russian Plain to Kamchatka, to yield a natural range of size variation in this reference set. These skulls were collected from the middle of the 19th century to the end of the 20th century. Wolf-dog hybrids have been described, especially from wolf populations from Italy (Verardi et al., 2006) and from the Baltic regions (Andersone et al., 2002). According to Randi (2011), F1-hybrids do not readily backcross into wolf populations and introgression in nature is counteracted by behavioural or

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selective constraints. Although we cannot totally eliminate the possibility that our reference set contains wolf-dog hybrids, we believe that the probability is low. Hybridization is likely to be most frequent near human settlements, where wolf density is low and dogs are common (Wayne and Vilà, 2003). However, we took samples from a large geographic area, where the density of human settlements is low, so the probability that hybrids were taken will be low. Male wolves are larger, on average, than females for both cranial and dental characters (Dayan et al., 1992). Unfortunately, sex is known for only a limited number of our reference wolf skulls: six female specimens and 17 male specimens. In addition to the Pleistocene Wolf and the Recent Eurasian Wolf groups, the comparative sample consists of four further reference groups, all of which comprised of dogs: the Palaeolithic Dog, the Prehistoric Dog, the Recent Archaic Dog and the Recent Other Dog group. The amount of shape variation of the skull among dogs far exceeds that in grey wolves (Drake and Klingenberg, 2010). The Recent Archaic Dog group includes two northern breeds. The Recent Other Dog group consists of the following functional groups as defined by the American Kennel Club (2006): Herding Dog (two breeds), Gua...