Paleolithic dog

Watercolor tracing made by archaeologist Henri Breuil from a cave painting of a wolf-like canid, Font-de-Gaume, France dated 19,000 years ago.

The Paleolithic dog was a Late Pleistocene canine. They were directly associated with human hunting camps in Europe over 30,000 years before present (YBP) and it is proposed that they were domesticated. They are also proposed to be either a proto-dog and the ancestor of the domestic dog or an extinct morphologically and genetically divergent wolf population.

Taxonomy

A study has classified the Paleolithic dog as Canis c.f. familiaris[1] where c.f. is used in Latin to mean confer, uncertain (Canis believed to be familiaris). Previously 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".[2][3]:p513 The specimens were given the name Canis lupus domesticus (domesticated wolf).[3]:p513[4]

Naming

In 2002, a study looked at 2 fossil skulls of large canids dated at 16,945 YBP that had been found buried 2 metres and 7 metres from 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".[5] In 2009, another study 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.[6]

Description

Wolf mandible diagram showing the names and positions of the teeth.

The Paleolithic dog was smaller than the Pleistocene wolf (Canis c.f. lupus)[1] and the extant gray wolf (Canis lupus), with a skull size that indicated a dog similar in size to the modern large dog breeds. The Paleolithic dog had a mean body mass of 36–37 kg compared to Pleistocene wolf 42–44 kg and recent European wolf 41–42 kg.[6]

The earliest sign of domestication in dogs was thought to be the neotonization of skull morphology[7][8][9] and the shortening of snout length that results in tooth crowding, reduction in tooth size, and a reduction in the number of teeth,[7][10] which has been attributed to the strong selection for reduced aggression.[7][8]

The Paleolithic dog had a skull length shorter, a viscerocranium (face) length shorter, and a snout wider than Pleistocene and modern wolves.[6] It had a wider palate and wider braincase,[6][9] relatively short and massive jaws, and a shorter carnassial length but these were larger than the modern dog and closer to those of the wolf. The mandible of the Paleolithic dog is more massive compared to the elongated mandible of the wolves and has more crowded premolars, and of a hook-like extension of the caudal border of the coronoid process of the mandible. The snout width was greater than those of both the Pleistocene and modern wolves, and implies well-developed carnassials driven by powerful jaws.[6]

In two of the morphometric analyses, the nearest dog skull-shape that was similar to the Paleolithic dog was that of the Central Asian Shepherd Dog.

See further: Altai dog – 33,000 YBP (below)

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 Pleistocene wolf ate horse and possibly mammoth, the Paleolithic dog ate reindeer and muskox, and the humans ate specifically mammoth. The study proposed that the Paleolithic dog's diet had been artificially restricted as it was not a similar diet to the Pleistocene wolf. Some remote Arctic tribal people today restrict the diet of their dogs away from what those people prefer to eat.[1] An analysis of a specimen from the Eliseevichi-1 site on the Russian plain also revealed that the Paleolithic dog ate reindeer.[11]

Archaeological evidence

The Paleolithic dog's fossil remains have been found across Europe and with one instance in Central Asia.

See further Paleoecology of the time
Years BP Location Finding
36,000 Goyet Cave, Samson River Valley, Belgium Dog-like skull was found in a side gallery of the cave, together with remains from mammoth, lynx, red deer and large canids. The Goyet skull is very similar in shape to that of the Eliseevichi-I dog skulls (16,900 YBP) and to the Epigravettian Mezin 5490 and Mezhirich dog skulls (13,500 BP), which are about 18,000 years younger.[6][12] Palaeolithic artifacts in this system of caves date from the Mousterian, Aurignacian, Gravettian, and Magdalenian, which indicates recurrent occupations of the cave from the Pleniglacial until the Late Glacial.[6] No descendants, genetic classification of species is inconclusive.[13] Likely represents an extinct morphologically and genetically divergent wolf population.[14]
33,500 Razboinichya Cave, Altai Mountains, Central Asia (Russia) Dog-like skull, mandibles (both sides) and teeth. Specimen is unlike ancient and modern wolves but similar to the skulls of fully domesticated prehistoric dogs from Greenland (about 1,000 YBP) and the putative dogs from Eliseevichi-I site in central Russia.[15] No modern descendants, the morphological classification is that of a dog, the genetic classification of the species is inconclusive as it is not clear if it is a dog or a wolf - of 4 different types of tests two rated it as a wolf and two as a dog.[13]
32,000–22,000 Predmostí, Moravia, Czech Republic Three dog-like skulls. Predmostí is a Gravettian site. The skulls were found in the human burial zone and identified as Palaeolithic dogs, characterized by – compared to wolves – short skulls, short snouts, widepalates and braincases, and even-sized carnassials. Wolf skulls were also found at the site. One dog had been buried with a bone placed carefully in its mouth. The presence of dogs buried with humans at this Gravettian site corroborates the hypothesis that domestication began long before the Late Glacial.[16][17]
26,000 Chauvet Cave, Vallon-Pont-d'Arc, Ardèche region, France 50-metre trail of footprints made by a boy of about ten years of age alongside those of a large canid. The size and position of the canid's shortened middle toe in relation to its pads indicates a dog rather than a wolf. The footprints have been dated by soot deposited from the torch the child was carrying. The cave is famous for its cave paintings.[18] A later study using geometric morphometric analysis to compare modern wolf with modern dog tracks suggested that these were wolf tracks.[19]
22,000 Kostyonki-8, Voronezh, Russia One left mandible paired with the right maxilla. Declared a Paleolithic dog.[20]
16,945 Eliseevichi-I site, Bryansk Region, Russian Plain, Russia Two fossil canine skulls. In 2002, a study looked at the fossil skulls of two large canids that had been found buried 2 metres and 7 metres from what was once a mammoth-bone hut at this Upper Paleolithic site, and using an accepted morphologically based definition of domestication declared them to be "Ice Age dogs". The carbon dating gave a calendar-year age estimate that ranged between 16,945-13,905 YBP.[5] The Eliseevichi-1 skull is very similar in shape to the Goyet skull (36,000 BP), the Mezine dog skull (13,500 BP) and Mezhirich dog skull (13,500 BP).[6] In 2013, a study looked at the mitochondrial DNAsequence for one of these skulls and identified it as Canis lupus familiaris i.e. dog.[13] However, in 2015 a study using three-dimensional geometric morphometric analyses indicated the skull is more likely from a wolf.[14][21]
16,700 Kniegortte, Germany Partial maxillary fragment with teeth dated 16,700-13,800 YBP.[22]
15,770 Oelknitz, Germany Phalanges, metapodia and part of distal humerus and tibia dated 15,770-13,957 YBP.[22]
15,770 Teufelsbrucke, Germany Proximal metapodial fragment and first phalanx dated 15,770 -13,957.[22]
15,500 Montespan, France 1 atlas, 1 femur, 1 baculum dated 15,500-13,500.[23]
15,200 Hauterive-Champréveyres, Switzerland Metatarsal and two teeth, second phalanx dated 15,200-13,900 YBP.[24]
14,999 Le Closeau, France 7 fragments including mandible, meta carpal, metapodial and phalanxes 14,999-14,055 YBP.[23]
13,500 approx Mezine, Ukraine Ancient dog-like skull as well as ancient wolf specimens found at the site. Dated to the Epigravettian period (17,000–10,000 BP). The Mezine skull is very similar in shape to the Goyet skull (36,000 YBP), Eliseevichi-1 dog skulls (16,900 YBP) and Mezhirich dog skull (13,500 YBP). The Epigravettian Mezine site is well known for its round mammoth bone dwelling.[6]
13,500 approx Mezhirich, Ukraine Ancient dog-like skull. Dated to the Epigravettian period (17,000–10,000 YBP). The Mezhirich skull is very similar in shape to the Goyet skull (36,000 YBP), the Eliseevichi-1 dog skulls (15,000 YBP) and Mezine dog skull (13,500 YBP). The Epigravettian Mezhirich site had four mammoth bone dwellings present.[6]

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.[13]

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 analysis to look for evidence of the presence of Paleolithic dogs that were separate from Pleistocene wolves. Reference groups included the Eliseevichi-1 prehistoric dogs, recent dogs and wolves. 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 dated at 36,000 YBP 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 and Mezhirich, Ukraine skulls were also identified as being Paleolithic dogs. Collagen analysis indicated that the Paleolithic dogs associated with human hunter-gatherer 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.[6][12]

Further studies later looked at wolf-like fossils from Paleolithic hunter-gatherer sites across Europe and proposed 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).[20] 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.[16]

It has been proposed that based on the genetic evidence of modern dogs being traced to the ancient wolves of Europe, the archaeological evidence of the Paleolithic dog remains being found at known European hunting camp-sites, their morphology, and collagen analysis that indicated that their diet had been artificially restricted compared to nearby 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.[6][12] In 2016, a study discounted the use of the Paleolithic dogs from the Predmosti site as pack animals.[25]

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,[6] its refutation,[26] a counter to the refutation,[27] a second article,[16] its refutation,[28] a third article that includes a counter to the refutation,[20] its refutation,[29] a counter to the refutation,[30] another refutation,[21] support based on bone collagen analysis,[1] and the identification of an ancient paleolithic dog in Yakutia.[31]

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

Two domestication events

Chauvet cave artistic depiction of horses (refer #Archaeological evidence 26,000 YBP)

Studies have suggested that it was possible for multiple primitive forms of the dog to have existed, including in Europe.[32] European dog populations had undergone extensive turnover during the last 15,000 years that has erased the genomic signature of early European dogs,[33][34] the genetic heritage of the modern breeds has become blurred due to admixture,[35] and there was the possibility of past domestication events that had died out or had been largely replaced by more modern dog populations.[33]

In 2016, a study proposed that dogs may have been domesticated separately in both Eastern and Western Eurasia from two genetically distinct and now extinct wolf populations. East Eurasian dogs then made their way with migrating people to Western Europe between 14,000-6,400 YBP where they partially replaced the dogs of Europe.[36] Two domestication events in Western Eurasia and Eastern Eurasia has recently been found for the domestic pig.[36][37]

As the taxonomic classification of the "proto-dog" Paleolithic dogs as being either dogs or wolves remains controversial, they were excluded from the study.[36]

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.[12] 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.[6][38] The analysis indicated that the Belgian fossil large canids in general preyed on horse and large bovids.[6][16]

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 Genbank accession number KF661079, and with Belgium 30,000 YBP KF661080 and 26,000 years YBP KF661078 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. However, in 2015 three-dimensional geometric morphometric analyses indicated this, and the Eliseevichi-1 dog, is more likely from a wolf.[21][14] 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[13] as there was for domestic pigs.[37] A 2016 review proposed that it most likely represents an extinct morphologically and genetically divergent wolf population.[14]

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".[39]

In 2007, a mtDNA analysis of extinct eastern Beringian wolves showed that two ancient wolves from the Ukraine dated 30,000 YBP and 28,000 YBP and the 33,000 YBP Altai dog had the same sequence as six Beringian wolves,[40] indicating a common maternal ancestor. In 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.[15]

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 GenBank accession number JX173682. Of four tests, 2 tests showed its sequence to fall within the wolf clade and 2 tests within the dog clade. 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[13] as there was for domestic pigs.[37]

Local unknown wolves

Ecological factors including habitat type, climate, prey specialization and predatory competition will greatly influence wolf genetic population structure and cranio-dental plasticity.[41][42][43][44][45][46][47][48][49] 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.[49]

There are a small number of Canis remains that have been found at Goyet Cave, Belgium (36,500 YBP)[6] Razboinichya Cave, Russia (33,500 YBP)[39] Kostenki 8, Russia (33,500-26,500 YBP)[30] Predmosti, Czech Republic (31,000 YBP)[16] and Eliseevichi-1, Russia (17,000 YBP).[5] Based on cranial morphometric study of the characteristics thought to be associated with the domestication process, these have been proposed as early Paleolithic dogs.[30] These characteristics of shortened rostrum, tooth crowding, and absence or rotation of premolars have been documented in both ancient and modern wolves.[26][47][49][40][50][51] Rather than representing early dogs, these specimens may represent an extinct morphologically and genetically divergent wolf population.[14][35][49]

However, regardless of it eventually proving to be either a proto-dog or an unknown species of wolf, the original proposal was that the "Paleolithic dog" was domesticated.[6]

References

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