Paleolithic dog

This article is about the Paleolithic dog. For the dog, see Dog. For the wolf, see Wolf.
Polychrome cave painting of a wolf-like canid, Font-de-Gaume, France.

The Paleolithic dog (Canis c.f. familiaris)[1] is the name given to a number of unique fossil specimens that are proposed to have been a Late Pleistocene – early Holocene European canid[2][1] that was smaller in size compared to the Pleistocene wolf (Canis c.f. lupus)[1] and the extant gray wolf (Canis lupus). They were directly associated with human hunting camps 30,000 years before present (YBP).[1] They are now extinct.

Naming

"Palaeolithic" was the original European spelling however this article uses the US spelling to help maintain consistency with links to related pages.

Mammoth bone dwelling, Mezhirich site

Specimen references used in this article relate to the following institutions:
MAE RAS - Museum of Anthropology and Ethnography (Kunstkamera), Russian Academy of Science, Saint-Petersburg
PM NASU - Palaeontological Museum National Academy of Science of Ukraine, Kiev
RBINS - Royal Belgian Institute of Natural Sciences
ZIN RAS - Zoological Institute of the Russian Academy of Science, Saint-Petersburg

In 1969, a study of ancient mammoth-bone dwellings at the Mezine paleolithic site in the Chernigov region, Ukraine uncovered 3 possibly domesticated "short-faced wolves".[3][4]:p513 The specimens were given the name Canis lupus domesticus (domesticated wolf).[5][4]:p513 In 2002, a study looked at 2 fossil skulls of large canids (references: MAE RAS 447 and ZIN RAS 23781) dated at 13,905 YBP that had been found buried within metres of what was once a mammoth-bone hut at the Upper Paleolithic site of Eliseevichi-1 in the Bryansk region of central Russia, and using an accepted morphologically-based definition of domestication declared them to be "Ice Age dogs".[6] In 2013, a study recalibrated the age to 15,000 BP with the genetic analysis of one skull deposited in GenBank with accession number KF661082 and classified as Canis lupus familiaris (dog).[7] In 2009, another study had looked at these 2 early dog skulls in comparison to other much earlier but morphologically similar fossil skulls that had been found across Europe and concluded that the earlier specimens were "Palaeolithic dogs", which were morphologically and genetically distinct from Pleistocene wolves that lived in Europe at that time.[2]

Taxonomy

The taxonomy is not well defined.[2] In 2015, a study classified the Paleolithic dog as (Canis c.f. familiaris)[1] i.e. Canis believed to be familiaris. A nuclear genome study of recent dogs and wolves has indicated that before the divergence of dogs from wolves there was much more wolf diversity.[8]

Morphology

In comparison to the Pleistocene wolf, they had a short skull length, short snout, a wide palate and braincase, relatively short and massive jaws, and a shorter carnassial length, but these were larger than the modern dog. The mandible of the Paleolithic dog is more massive compared to the elongated mandible of the wolves and presents a high frequency of crowded premolars and of a hook-like extension of the caudal border of the coronoid process. The Paleolithic dog had a mean body mass of 36–37 kg compared to Pleistocene wolves 42–44 kg and recent European wolves 41–42 kg.[2]

Diet

In 2015, a study of bone collagen taken from a number of species found at the 30,000 YBP mammoth-hut site of Predmosti (Czech Republic) indicted that the Eurasian cave lion and the Paleolithic dog ate reindeer and muskox, the Pleistocene wolf ate horse and possibly mammoth, and the humans ate specifically mammoth. The study proposed that the Paleolithic dog's diet had been artificially restricted, else it would have had a similar diet as the Pleistocene wolf. Some remote Arctic tribal people today restrict the diet of their dogs away from what those people prefer to eat. The study also found that when canids were included in the human diet, humans ate wolf rather than dog.[1]

Habitat

The Paleolithic dog's fossil remains have been found across Europe.

See further Paleoecology of the time

Relationship to the domestic dog

The Upper Paleolithic

Upper Paleolithic (40–10 ka)

Châtelperronian (45-40 ka)
Baradostian (36 ka)
Aurignacian (32–26 ka)
Gravettian (28–22 ka)
Solutrean (21–17 ka)
Magdalenian (18–10 ka)
Hamburg (15 ka)
Ahrensburg (13 ka)
Swiderian (10 ka)
↓ Mesolithic
↓ Stone Age

In 2013, a major Mitochondrial DNA study has found that divergence times from wolf to dog implies a European origin of the domestic dog dating 18,800-32,100 years ago, which supports the hypothesis that dog domestication preceded the emergence of agriculture and occurred in the context of European hunter-gatherer cultures.[7]

In 2009, a study proposed that there was a low frequency of recognized dog skulls in Upper Paleolithic sites because existing specimens had not yet been recognized as dogs. The study looked at the 2 Eliseevichi-1 dog skulls in comparison to much earlier Late Pleistocene but morphologically similar fossil skulls that had been found across Europe, and proposed the much earlier specimens were Paleolithic dogs that were morphologically and genetically distinct from the Pleistocene wolves living in Europe at that time. The study looked at 117 skulls of recent and fossil large canids. Several skulls of fossil large canids from sites in Belgium, the Ukraine and Russia were examined using multivariate techniques to look for possible evidence of the presence of Paleolithic dogs that were separate from Pleistocene wolves. Reference groups constituted of the Eliseevichi-1 prehistoric dogs, and recent wolves and dogs. The osteometric analysis of the skulls indicated that the Paleolithic dogs fell outside the skull ranges of the Pleistocene wolf group and the modern wolf group, and were closer related to those of the Eliseevichi-1 prehistoric dog group. The fossil large canid from Goyet, Belgium (RBINS 2860), dated at 31,700 BP was clearly different from the recent wolves, resembling most closely the Eliseevichi 1 prehistoric dogs and suggesting that dog domestication had already started during the Aurignacian. The two Epigravettian Mezine, Ukraine (PM NASU 5490) and Mezhirich, Ukraine (PM NASU 4493) skulls are also identified as being Paleolithic dogs. Collagen analysis indicated that the Paleolithic dogs associated with human camp-sites (Eliseevichi 1, Mezine and Mezhirich) had been specifically eating reindeer, while other predator species in those locations and times had eaten a range of prey.[2]

Further studies have looked at wolf-like fossils from Paleolithic hunter-gatherer sites across Europe and propose to have identified Paleolithic dogs at Predmosti (Czech Republic 26,000-27,000 YBP), Kosteki-8 (Russia 23,000-27,700 YBP), Kosteki-1 (Russia 22,000-24,000 BP), Kosteki-17 (Russia Upper Paleolithic) and Verholenskaya (Russia late glacial).[9] In the human burial zone at the Predmosti site, 3 Paleolithic skulls were found that resemble those of a Siberian husky but they were larger and heavier than the modern husky. For one skull, "a large bone fragment is present between the upper and lower incisors that extends several centimetres into the mouth cavity. The size, thickness and shape of the fragment suggest that it could be a fragment of a bone of a large mammal, probably from a mammoth. The position of the bone fragment in the mouth and the articulated state of the lower jaw with the skull indicate that this mammoth bone fragment was inserted artificially into the mouth of the dog post-mortem." The morphology of some wolf-like fossils was such that they could not be assigned to either the Pleistocene wolf nor Paleolithic dog groups.[10]

It has been proposed that, based on the genetic evidence of the timeline and European location, the archaeological evidence of the Paleolithic dog remains being found at known European hunting camp-sites, their morphology, and collagen analysis that indicated their diet had been restricted compared to wolves, that the Paleolithic dog was domesticated. It has also been hypothesized that the Paleolithic dog may have provided the stock from which early dogs arose, or alternatively that they are a type of wolf that is not known to science.[2]

There has been ongoing debate in the scientific press about what the fossil remains of the Paleolithic dog might be, with some commenters declaring them as either wolves or a unique form of wolf. These include a first article proposing the Paleolithic dog,[2] its refutation,[11] a counter to the refutation,[12] a second article,[10] its refutation,[13] a third article that includes a counter to the refutation,[9] its refutation,[14] a counter to the refutation,[15] another refutation,[16] and another article in support based on bone collagen analysis.[1]

As the ancestor of the dog has not been positively identified by scientists, this debate continues.

Two domestication events?

It is possible that multiple primitive forms of the dog existed, including in Europe.[17] European dog populations have undergone extensive turnover during the last 15,000 years that has erased the genomic signature of early European dogs,[18][19] the genetic heritage of the modern breeds has become blurred due to admixture,[20] and studies of today's dogs cannot exclude the possibility of past domestication events that died out or were overwhelmed by more modern dog populations.[19]

Goyet dog – 36,000 YBP

Genus Canis, species indeterminate

In 2009, a study looked at 117 skulls of recent and fossil large canids. None of the 10 canid skulls from the Belgian caves of Goyet, Trou du Frontel, Trou de Nutons, and Trou de Chaleux could be classified, so the team took as their basic assumption that all of these canid samples were wolves.[21] The DNA sequence of seven of the skulls indicated seven unique haplotypes that represented ancient wolf lineages lost until now. The osteometric analysis of the skulls showed that one large canid fossil from Goyet was clearly different from recent wolves, resembling most closely the Eliseevichi-1 dogs (15,000 years YBP) and so was identified as a Paleolithic dog (see below).[2][22]

In November 2013, a DNA study sequenced three haplotypes from the ancient Belgium canids (the Goyet dog - Belgium 36,000 YBP cataloged as Canis species, and with Belgium 30,000 YBP and 26,000 years YBP cataloged as Canis lupus) and found they formed the most diverging group. Although the cranial morphology of the Goyet dog has been interpreted as dog-like, its mitochondrial DNA relation to other canids places it as an ancient sister-group to all modern dogs and wolves rather than a direct ancestor. Belgium 26,000 YBP has been found to be uniquely large but was found not to be related to the Beringian wolf. This Belgium canid clade may represent a phenotypically distinct and not previously recognized population of gray wolf, or the Goyet dog may represent an aborted domestication episode. If so, there may have been originally more than one ancient domestication event for dogs[7] as there was for domestic pigs.[23]

Altai dog – 33,000 YBP

Genus Canis, species indeterminate
33,000-year-old skull of a dog-like canid found in the Altai Mountains. It has no direct descendants today.

In 2011, a study looked at the well-preserved 33,000-year-old skull and left mandible of a dog-like canid that was excavated from Razboinichya Cave in the Altai Mountains of southern Siberia (Central Asia). The morphology was compared to the skulls and mandibles of large Pleistocene wolves from Predmosti, Czech Republic, dated 31,000 YBP, modern wolves from Europe and North America, and prehistoric Greenland dogs from the Thule period (1,000 YBP or later) to represent large-sized but unimproved fully domestic dogs. "The Razboinichya Cave cranium is virtually identical in size and shape to prehistoric Greenland dogs" and not the ancient nor modern wolves. However, the lower carnassial tooth fell within the lower range of values for prehistoric wolves and was only slightly smaller than modern European wolves, and the upper carnassial tooth fell within the range of modern wolves. "We conclude, therefore, that this specimen may represent a dog in the very early stages of domestication, i.e. an incipient dog, rather than an aberrant wolf... The Razboinichya Cave specimen appears to be an incipient dog...and probably represents wolf domestication disrupted by the climatic and cultural changes associated with the Last Glacial Maximum".[24]

In March 2013, a DNA study of the Altai dog deposited the sequence in GenBank with a classification of Canis lupus familiaris (dog). "The analyses revealed that the unique haplotype of the Altai dog is more closely related to modern dogs and prehistoric New World canids than it is to contemporary wolves... This preliminary analysis affirms the conclusion that the Altai specimen is likely an ancient dog with shallow divergence from ancient wolves. These results suggest a more ancient history of the dog outside of the Middle East or East Asia." The haplotype groups closest to the Altai dog included such diverse breeds as the Tibetan mastiff, Newfoundland, Chinese crested, cocker spaniel and Siberian husky.[25]

In November 2013, a study looked at 18 fossil canids and compared these with the complete mitochondrial genome sequences from 49 modern wolves and 77 modern dogs. A more comprehensive analysis of the complete mDNA found that the phylogenetic position of the Altai dog as being either dog or wolf was inconclusive and cataloged its sequence as Canis species. All tests showed it to fall equally in both the wolf and dog clades. The sequence strongly suggests a position at the root of a clade uniting two ancient wolf genomes, two modern wolves, as well as two dogs of Scandinavian origin. However, the study does not support its recent common ancestry with the great majority of modern dogs. The study suggests that it may represent an aborted domestication episode. If so, there may have been originally more than one ancient domestication event for dogs[7] as there was for domestic pigs.[23]

See also Origin of the domestic dog
See also Megafaunal wolf

Local, unknown wolves?

Ecological factors including habitat type, climate, prey specialization and predatory competition will greatly influence their genetic population structure and cranio-dental plasticity.[26][27][28][29][30][31][32][33][34] Therefore, within the Pleistocene gray wolf population the variations between local environments would have encouraged a range of wolf ecotypes that were genetically, morphologically and ecologically distinct from one another.[34]

There are a small number of Canis remains that have been found at Goyet Cave, Belgium (36,500 YBP)[2] Razboinichya Cave, Russia (33,500 YBP)[24] Kostenki 8, Russia (33,500-26,500 YBP)[15] Predmosti, Czech Republic (31,000 YBP)[10] and Eliseevichi 1, Russia (17,000 YBP).[6] Based on cranial morphometric study of the characteristics thought to be associated with the domestication process, these have been proposed as early Paleolithic dogs.[15] These characteristics of shortened rostrum, tooth crowding, and absence or rotation of premolars have been documented in both ancient and modern wolves.[35][36][32][37][11][34] Rather than representing early dogs, these specimens may represent "a morphologically distinct local, now extinct, population of wolves".[20][34]

References

  1. 1 2 3 4 5 6 7 Bocherens, H (2015). "Reconstruction of the Gravettian food-web at Předmostí I using multi-isotopic tracking (13C, 15N, 34S) of bone collagen". Quaternary International. 359-360: 211–228. doi:10.1016/j.quaint.2014.09.044.
  2. 1 2 3 4 5 6 7 8 9 Germonpré, Mietje; Sablin, Mikhail V.; Stevens, Rhiannon E.; Hedges, Robert E.M.; Hofreiter, Michael; Stiller, Mathias; Despre´s, Viviane R. (2009). "Fossil dogs and wolves from Palaeolithic sites in Belgium, the Ukraine and Russia: osteometry, ancient DNA and stable isotopes". Journal of Archaeological Science 36 (2): 473–490. doi:10.1016/j.jas.2008.09.033.
  3. ↑ Pidoplichko, I. G. (1969) Late Palaeolithic dwellings of Mammoth bones in the Ukraine. Academy of Sciences of the Ukraine. Institute of Zoology. Nauka, Doumka, Kiev. (in Russian)
  4. 1 2 Olsen, S.J. (2000). "8". In Kipple, K.F. and Ornelas, K.C. The Cambridge World History of Food. Cambridge University Press.
  5. ↑ Pidoplichko I.G. (1998) Upper Paleolithic Dwellings of Mammoth Bones in the Ukraine. Oxford: BAR International Series 712
  6. 1 2 Sablin, Mikhail V.; Khlopachev, Gennady A. (2002). "The Earliest Ice Age Dogs:Evidence from Eliseevichi I" (PDF). Wenner-Gren Foundation for Anthropological Research. Retrieved 10 January 2015.
  7. 1 2 3 4 Thalmann, O.; Shapiro, B.; Cui, P.; Schuenemann, V.J.; Sawyer, S.K.; Greenfield, D.L.; Germonpré, M.B.; Sablin, M.V.; López-Giráldez, F.; Domingo-Roura, X.; Napierala, H.; Uerpmann, H-P.; Loponte, D.M.; Acosta, A.A.; Giemsch, L.; Schmitz, R.W.; Worthington, B.; Buikstra, J.E.; Druzhkova, A.S.; Graphodatsky, A.S.; Ovodov, N.D.; Wahlberg, N.; Freedman, A.H.; Schweizer, R.M.; Koepfli, K.-P.; Leonard, J.A.; Meyer, M.; Krause, J.; Pääbo, S.; Green, R.E.; Wayne, Robert K. (15 November 2013). "Complete Mitochondrial Genomes of Ancient Canids Suggest a European Origin of Domestic Dogs". Science (AAAS) 342 (6160): 871–874. doi:10.1126/science.1243650. PMID 24233726. Retrieved 24 December 2014. refer Supplementary material Page 27 Table S1
  8. ↑ Freedman, Adam H.; Gronau, Ilan; Schweizer, Rena M.; Ortega-Del Vecchyo, Diego; Han, Eunjung; Silva, Pedro M.; Galaverni, Marco; Fan, Zhenxin; Marx, Peter; Lorente-Galdos, Belen; Beale, Holly; Ramirez, Oscar; Hormozdiari, Farhad; Alkan, Can; Vilà, Carles; Squire, Kevin; Geffen, Eli; Kusak, Josip; Boyko, Adam R.; Parker, Heidi G.; Lee, Clarence; Tadigotla, Vasisht; Siepel, Adam; Bustamante, Carlos D.; Harkins, Timothy T.; Nelson, Stanley F.; Ostrander, Elaine A.; Marques-Bonet, Tomas; Wayne, Robert K.; Novembre, John (16 January 2014). "Genome Sequencing Highlights Genes Under Selection and the Dynamic Early History of Dogs". PLOS Genetics (PLOS Org) 10 (1): e1004016. doi:10.1371/journal.pgen.1004016. PMC 3894170. PMID 24453982. Retrieved December 8, 2014.
  9. 1 2 Germonpré, Mietje; Laznickova-Galetova, Martina; Losey, Robert J.; Raikkonen, Jannikke; Sablin, Mikhail V. (2014). "Large canids at the Gravettian Predmostí site, the Czech Republic:The mandible". Quaternary International xxx: 1–19. doi:10.1016/j.quaint.2014.07.012.
  10. 1 2 3 Germonpré, Mietje; Laznickova-Galetova, Martina; Sablin, Mikhail V. (2012). "Palaeolithic dog skulls at the Gravettian Predmosti site, the Czech Republic". Journal of Archaeological Science 39: 184–202. doi:10.1016/j.jas.2011.09.022.
  11. 1 2 Crockford SJ, Kuzmin YV (2012) Comments on Germonpre et al., Journal of Archaeological Science 36, 2009 "Fossil dogs and wolves from Palaeolithic sites in Belgium, the Ukraine and Russia: osteometry, ancient DNA and stable isotopes"
  12. ↑ Germonpré, Mietje; Sablin, MV; Despres, V; Hofreiter, M; Laznickova-Galetova, M; et al. (2013). "Palaeolithic dogs and the early domestication of the wolf: a reply to the comments of Crockford and Kuzmin (2012)". Journal of Archaeological Science 40: 786–792. doi:10.1016/j.jas.2012.06.016.
  13. ↑ Boudadi-Maligne, Myriam; Escarguel, Gilles (2014). "A biometric re-evaluation of recent claims for Early Upper Palaeolithic wolf domestication in Eurasia". Journal of Archaeological Science (Elsevier Ltd) 45: 80–89. doi:10.1016/j.jas.2014.02.006.
  14. ↑ Morley, Darcy F. (2014). "In search of Paleolithic dogs: a quest with mixed results". Journal of Archaeological Science 52: 300–307. doi:10.1016/j.jas.2014.08.015.
  15. 1 2 3 Germonpré, Mietje; Sablin, Mikhail V.; Laznickova-Galetova, Martina; Despre´s, Viviane R.; Stevens, Rhiannon E.; Stiller, Mathias; Hofreiter, Michael (2015). "Palaeolithic dogs and Pleistocene wolves revisited: a reply to Morey (2014)". Journal of Archaeological Science 54: 210–216. doi:10.1016/j.jas.2014.11.035.
  16. ↑ Drake, Abby Grace; Coquerelle, Michael; Colombeau, Guillaume (5 February 2015). "3D morphometric analysis of fossil canid skulls contradicts the suggested domestication of dogs during the late Paleolithic". Scientific Reports 5 (2899): 8299. doi:10.1038/srep08299. PMID 25654325.
  17. ↑ Wang, G (2015). "Out of southern East Asia: the natural history of domestic dogs across the world". Cell Research. doi:10.1038/cr.2015.147.
  18. ↑ Malmström, Helena; Vilà, Carles; Gilbert, M; Storå, Jan; Willerslev, Eske; Holmlund, Gunilla; Götherström, Anders (2008). "Barking up the wrong tree: Modern northern European dogs fail to explain their origin". BMC Evolutionary Biology 8: 71. doi:10.1186/1471-2148-8-71. PMC 2288593. PMID 18307773.
  19. 1 2 Shannon, L (2015). "Genetic structure in village dogs reveals a Central Asian domestication origin". Proceedings of the National Academy of Sciences: 201516215. doi:10.1073/pnas.1516215112.
  20. 1 2 Larson G (2012). "Rethinking dog domestication by integrating genetics, archeology, and biogeography". PNAS 109 (23): 8878–8883. doi:10.1073/pnas.1203005109.
  21. ↑ Shipman, P. (2011). The Animal Connection: A New Perspective on What Makes Us Human. W W Norton & Co New York. p. 218.
  22. ↑ Royal Belgium Institute of Natural Sciences. "Goyet skull photo".
  23. 1 2 Frantz, L. (2015). "Evidence of long-term gene flow and selection during domestication from analyses of Eurasian wild and domestic pig genomes". Nature Genetics 47: 1141–1148. doi:10.1038/ng.3394.
  24. 1 2 Ovodov, N. (2011). "A 33,000-year-old incipient dog from the Altai Mountains of Siberia: Evidence of the earliest domestication disrupted by the Last Glacial Maximum". PLoS ONE 6 (7): e22821. doi:10.1371/journal.pone.0022821. PMC 3145761. PMID 21829526.
  25. ↑ Druzhkova, A. (2013). "Ancient DNA analysis affirms the canid from Altai as a primitive dog". PLoS ONE 8 (3): e57754. doi:10.1371/journal.pone.0057754. PMC 3590291. PMID 23483925.
  26. ↑ Carmichael, L. E.; Nagy, J. A.; Larter, N. C.; Strobeck, C. (2001). "Prey specialization may influence patterns of gene flow in wolves of the Canadian Northwest". Molecular Ecology 10 (12): 2787. doi:10.1046/j.0962-1083.2001.01408.x. PMID 11903892.
  27. ↑ Carmichael, L.E., 2006. Ecological Genetics of Northern Wolves and Arctic Foxes. Ph.D. Dissertation. University of Alberta.
  28. ↑ Geffen, ELI; Anderson, Marti J.; Wayne, Robert K. (2004). "Climate and habitat barriers to dispersal in the highly mobile grey wolf". Molecular Ecology 13 (8): 2481. doi:10.1111/j.1365-294X.2004.02244.x. PMID 15245420.
  29. ↑ Pilot, Malgorzata; Jedrzejewski, Wlodzimierz; Branicki, Wojciech; Sidorovich, Vadim E.; Jedrzejewska, Bogumila; Stachura, Krystyna; Funk, Stephan M. (2006). "Ecological factors influence population genetic structure of European grey wolves". Molecular Ecology 15 (14): 4533. doi:10.1111/j.1365-294X.2006.03110.x. PMID 17107481.
  30. ↑ Musiani, Marco; Leonard, Jennifer A.; Cluff, H. Dean; Gates, C. Cormack; Mariani, Stefano; Paquet, Paul C.; Vilà, Carles; Wayne, Robert K. (2007). "Differentiation of tundra/taiga and boreal coniferous forest wolves: Genetics, coat colour and association with migratory caribou". Molecular Ecology 16 (19): 4149. doi:10.1111/j.1365-294X.2007.03458.x. PMID 17725575.
  31. ↑ Hofreiter, Michael; Barnes, Ian (2010). "Diversity lost: Are all Holarctic large mammal species just relict populations?". BMC Biology 8: 46. doi:10.1186/1741-7007-8-46. PMC 2858106. PMID 20409351.
  32. 1 2 Flower, Lucy O.H.; Schreve, Danielle C. (2014). "An investigation of palaeodietary variability in European Pleistocene canids". Quaternary Science Reviews 96: 188. doi:10.1016/j.quascirev.2014.04.015.
  33. ↑ Leonard, Jennifer (2014). "Ecology drives evolution in grey wolves" (PDF). Evolution Ecology Research.
  34. 1 2 3 4 Perri, Angela (2016). "A wolf in dog's clothing: Initial dog domestication and Pleistocene wolf variation". Journal of Archaeological Science 68: 1. doi:10.1016/j.jas.2016.02.003.
  35. ↑ Leonard, J. (2007). "Megafaunal extinctions and the disappearance of a specialized wolf ecomorph". Current Biology 17 (13): 1146–50. doi:10.1016/j.cub.2007.05.072. PMID 17583509.
  36. ↑ Boudadi-Maligne, M., 2010. Les Canis pleistocenes du sud de la France: approche biosystematique, evolutive et biochronologique. Ph.D. dissertation. Universit�e Bordeaux 1
  37. ↑ Dimitrijević, V.; Vuković, S. (2015). "Was the Dog Locally Domesticated in the Danube Gorges? Morphometric Study of Dog Cranial Remains from Four Mesolithic-Early Neolithic Archaeological Sites by Comparison with Contemporary Wolves". International Journal of Osteoarchaeology 25: 1. doi:10.1002/oa.2260.
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