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Paleolithic dog
Paleolithic dog
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Polychrome tracing made by the archaeologist Henri Breuil from the cave painting of a wolf-like canid discovered in the Font-de-Gaume cave, Dordogne, France dated to 17,000 years ago

Purported remains of Paleolithic dogs have been reported from several European archaeological sites dating to over 30,000 years ago. Their status as domesticated is highly controversial, with some authors suggesting them to be the ancestors of the domestic dog or an extinct, morphologically and genetically divergent wolf population.

Taxonomy

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One authority has classified the Paleolithic dog as Canis cf. familiaris[1] (where cf. is a Latin term meaning uncertain, as in Canis believed to be familiaris). Previously in 1969, a study of ancient mammoth-bone dwellings at the Mezine paleolithic site in the Chernihiv region, Ukraine uncovered 3 possibly domesticated "short-faced wolves".[2][3] The specimens were classified as Canis lupus domesticus (domesticated wolf).[3][4]

Naming

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In 2002, a study looked at 2 fossil skulls of large canids dated at 16,945 years before present (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 "Paleolithic dogs", which were morphologically and genetically distinct from Pleistocene wolves that lived in Europe at that time.[6]

Description

[edit]
Diagram of a wolf skull with key features labelled

The Paleolithic dog was smaller than the Pleistocene wolf (Canis cf. lupus)[1] and the extant grey wolf (Canis lupus), with a skull size that indicates a dog similar in size to the modern large dog breeds. The Paleolithic dog had a mean body mass of 36–37 kg (79–82 lb) compared to Pleistocene wolf 42–44 kg (93–97 lb) and recent European wolf 41–42 kg (90–93 lb).[6]

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

Compared with the Pleistocene and modern wolves, the Paleolithic dog had a shorter skull length, a shorter viscerocranium (face) length, and a wider snout.[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 was more massive compared to the elongated mandible of the wolves and had more crowded premolars, and a hook-like extension in 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. In two morphometric analyses, the nearest dog skull-shape that was similar to the Paleolithic dog was that of the Central Asian Shepherd Dog.[6]

Diet

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In 2015, a study of bone collagen taken from a number of species found at the 30,000 YBP mammoth-hut site of Predmosti in the Czech Republic indicated that the Pleistocene wolf ate horse and possibly mammoth, the Paleolithic dog ate reindeer and muskox, and the humans ate specifically mammoth. The study proposes that the Paleolithic dog's diet had been artificially restricted because it was not a diet similar 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]

In 2020, a study of dental microwear on tooth enamel for canine specimens from Predmosti dated 28,500 YBP suggest a higher bone consumption for the proto-dogs compared with wolf specimens. This indicates two morphologically and behaviourally different canine types. The study proposes that the proto-dogs consumed more bone along with other less desirable food scraps within human camps, therefore this may be evidence of early dog domestication.[12]

Archaeological evidence

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See further Paleoecology of the time

Early specimens

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There are a number of recently discovered specimens which are proposed as being Paleolithic dogs, however their taxonomy is debated. These have been found in either Europe or Siberia and date 40,000–17,000 YBP. They include Hohle Fels in Germany, Goyet Caves in Belgium, Predmosti in the Czech Republic, and four sites in Russia: Razboinichya Cave in the Altai Republic, Kostyonki-8, Ulakhan Sular in the Sakha Republic, and Eliseevichi 1 on the Russian plain. Paw-prints from Chauvet Cave in France dated 26,000 YBP are suggested as being those of a dog, however these have been challenged as being left by a wolf.[13]

Chauvet Cave artistic depiction of horses dated 30,000 years ago
Paleolithic dog specimens (taxonomy debated)[13]
Years BP Location Finding
40,000–35,000 Hohle Fels, Schelklingen, Germany Paleolithic dog
36,500 Goyet Caves, Mozet, Belgium Paleolithic dog
33,500 Razboinichya Cave, Altai Mountains, Central Asia Paleolithic dog
33,500–26,500 Kostyonki-Borshchyovo archaeological complex, Voronezh, Russia Paleolithic dog
31,000 Predmostí, Moravia, Czech Republic Paleolithic dog
26,000 Chauvet Cave, Vallon-Pont-d'Arc, France Paw-prints
17,200 Ulakhan Sular, northern Yakutia, Siberia Paleolithic dog
17,000–16,000 Eliseevichi-I site, Bryansk Region, Russian Plain, Russia Paleolithic dog

There are also a number of later proposed Paleolithic dogs whose taxonomy has not been confirmed. These include a number of specimens from Germany (Kniegrotte, Oelknitz, Teufelsbrucke), Switzerland (Monruz, Kesslerloch, Champre-veyres-Hauterive), as well as Ukraine (Mezin, Mezhirich). A set of specimens dating 15,000–13,500 YBP have been confidently identified as domesticated dogs, based on their morphology and the archaeological sites in which they have been found. These include Spain (Erralla), France (Montespan, Le Morin, Le Closeau, Pont d'Ambon) and Germany (Bonn-Oberkassel). After this period, the remains of domesticated dogs have been identified from archaeological sites across Eurasia.[13]

Possible dog domestication between 15,000 and 40,000 YBP is not clear due to the debate over what the Paleolithic dog specimens represent. This is due to the flexibility of genus Canis morphology, and the close morphological similarities between Canis lupus and Canis familiaris. It is also due to the scarcity of Pleistocene wolf specimens available for analyses and so their morphological variation is unknown. Habitat type, climate, and prey specialization greatly modify the morphological plasticity of grey wolf populations, resulting in a range of morphologically, genetically, and ecologically distinct wolf morphotypes. With no baseline to work from, zooarchaeologists find it difficult to be able to differentiate between the initial indicators of dog domestication and various types of Late Pleistocene wolf ecomorphs, which can lead to the mis-identification of both early dogs and wolves. Additionally, the ongoing prehistoric admixture with local wolf populations during the domestication process may have led to canids that were domesticated in their behavior but wolflike in their morphology. Attempting to identify early tamed wolves, wolfdogs, or proto-dogs through morphological analysis alone may be impossible without the inclusion of genetic analyses.[13]

All specimens

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The table below lists by location and timing in years before present the very early co-location of hominid and wolf specimens, followed by proposed paleolithic dog and then early dog specimens, with the regions in which they had been found color-coded as purple – Western Eurasia, red – Eastern Eurasia and green – Central Eurasia.

Years BP Location Finding
400,000 Boxgrove near Kent, England Wolf bones in close association with hominid bones. These have been found in Lower Paleolithic sites including Boxgrove (400,000 YBP), Zhoukoudian in North China (300,000 YBP), and Grotte du Lazaret (125,000 YBP) in southern France. "The sites of occupation and hunting activities of humans and wolves must often have overlapped."[14] It is unknown if the co-location was the result of coincidence or a relationship.
300,000 Zhoukoudian cave system, China Small, extinct wolf skulls of Canis variabilis. At the site, the small wolf's remains were in close proximity to Peking Man (Homo erectus pekinensis).[15] It is unknown if the co-location was the result of coincidence or a relationship.
125,000 Grotte du Lazaret, near Nice, France Wolf skulls appear to have been set at the entrance of each dwelling in a complex of Paleolithic shelters. The excavators speculated that wolves were already incorporated into some aspect of human culture by this early time. A nearby wolf den intruded on the site.[16] In 1997, a study of maternal mDNA indicated that the genetic divergence of dogs from wolves occurred 100,000–135,000 YBP.[17] The Lazaret excavation lends credence to this mDNA study, in addition to indicating that a special relationship existed between wolves and genus Homo other than Homo sapiens, because this date is well before the arrival of Homo sapiens into Europe.[18] In 2018, a study of paternal yDNA indicated that the dog and the modern grey wolf genetically diverged from a common ancestor between 68,000 and 151,000 YBP.[19]
40,000–35,500 Hohle Fels, Schelklingen, Germany Canid maxillary fragment. The size of the molars matches those of a wolf, the morphology matches a dog.[20] Proposed as a Paleolithic dog. The figurine Venus of Hohle Fels was discovered in this cave and dated to this time.
36,500 Goyet Cave, Samson River Valley, Belgium The "Goyet dog" is proposed as being a Paleolithic dog.[6] 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][21] The dog-like skull was found in a side gallery of the cave, and 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] The Goyet dog left no descendants, and its genetic classification is inconclusive because its mitochondrial DNA (mDNA) does not match any living wolf nor dog. It may represent an aborted domestication event or phenotypically and genetically distinct wolves.[22] A genome-wide study of a 35,000 YBP Pleistocene wolf fossil from northern Siberia indicates that the dog and the modern grey wolf genetically diverged from a common ancestor between 27,000 and 40,000 YBP.[23][18]
33,500 Razboinichya Cave, Altai Mountains, Central Asia (Russia) The "Altai dog" is proposed as being a Paleolithic dog.[6] The specimens discovered were a dog-like skull, mandibles (both sides) and teeth. The morphological classification, and an initial mDNA analysis, found it to be a dog.[24] A later study of its mDNA was inconclusive, with 2 analyses indicating dog and another 2 indicating wolf. In 2017, two prominent evolutionary biologists reviewed the evidence and supported the Altai dog as being a dog from a lineage that is now extinct and that was derived from a population of small wolves that are also now extinct.[25]
33,500–26,500 Kostyonki-Borshchyovo archaeological complex, Voronezh, Russia One left mandible paired with the right maxilla, proposed as a Paleolithic dog.[26]
31,000 Predmostí, Moravia, Czech Republic Three dog-like skulls proposed as being Paleolithic dogs.[26] 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, wide palates 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.[27][28] Further analysis of bone collagen and dental microwear on tooth enamel indicates that these canines had a different diet when compared with wolves (refer under diet).[11][12]
30,800 Badyarikha River, northern Yakutia, Siberia Fossil canid skull. The specimen could not be classified as wolf nor Paleolithic dog.[29]
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.[30] A later study using geometric morphometric analysis to compare modern wolf with modern dog tracks proposes that these are wolf tracks.[31]
18,000 Indigirka, Yakutia, Siberia Dogor, a canine pup preserved in permafrost and yet to be identified as being a dog or wolf.[32]
17,300–14,100 Dyuktai Cave, northern Yakutia, Siberia Large canid remains along with human artefacts.[29]
17,200–16,800 Ulakhan Sular, northern Yakutia, Siberia Fossil dog-like skull similar in size to the "Altai dog", proposed as a Paleolithic dog.[29]
17,000–16,000 Eliseevichi-I site, Bryansk Region, Russian Plain, Russia Two fossil canine skulls proposed as being a Paleolithic dogs.[6] 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 and 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 mDNA sequence for one of these skulls and identified it as Canis lupus familiaris i.e. dog.[22] However, in 2015 a study using three-dimensional geometric morphometric analyses indicated the skull is more likely from a wolf.[33][34] These animals were larger in size than most grey wolves and approach the size of a Great Dane.[35]
16,900 Afontova Gora-1, Yenisei River, southern Siberia Fossil dog-like tibia, proposed as a Paleolithic dog. The site is on the western bank of the Yenisei River about 2,500 km southwest of Ulakhan Sular, and shares a similar timeframe to that canid. A skull from this site described as dog-like has been lost in the past, but there exists a written description of it possessing a wide snout and a clear stop, with a skull length of 23 cm that falls outside of those of wolves.[29]
16,700 Kniegortte, Germany Partial maxillary fragment with teeth dated 16,700–13,800 YBP.[36] Taxonomy uncertain.[13]
16,300 Monruz, Switzerland Deciduous teeth possibly from a dog.[37] Taxonomy uncertain.[13]
15,770 Oelknitz, Germany Phalanges, metapodia and part of distal humerus and tibia dated 15,770–13,957 YBP.[36] Taxonomy uncertain.[13]
15,770 Teufelsbrucke, Germany Proximal metapodial fragment and first phalanx dated 15,770–13,957.[36] Taxonomy uncertain.[13]
15,500–13,500 Montespan, France 1 atlas, 1 femur, 1 baculum dated 15,500–13,500.[38] Domestic dog.[13]
15,200 Champré-veyres-Hauterive, Switzerland Metatarsal and two teeth, second phalanx dated 15,200–13,900 YBP.[39] Taxonomy uncertain.[13]
14,999 Le Closeau, France 7 fragments including mandible, meta carpal, metapodial and phalanxes 14,999–14,055 YBP.[38] Domestic dog.[13]
14,900 Verholenskaya Gora, Irkutsk, Siberia Lower jaw of a large canid. Found on the Angara river near Irkutsk about 2400 km southwest of Ulakhan Sular. Proposed as a Paleolithic dog.[29]
14,600–14,100 Kesslerloch Cave, Switzerland Large maxillary fragment that is too small to be from a wolf. Proposed as a Paleolithic dog.[40] Taxonomy uncertain.[13]
14,500 Erralla cave, Gipuzkoa, Spain Humerus of a dog, the dimensions of which are close to that of the dog humerus found at Pont d'Ambon.[41] Domestic dog.[13]
14,223 Bonn-Oberkassel, Germany The "Bonn-Oberkassel dog". Undisputed dog skeleton buried with a man and woman. All three skeletal remains were found sprayed with red hematite powder.[42] The consensus is that a dog was buried along with two humans.[43] Analysis of mDNA indicates that this dog was a direct ancestor of modern dogs.[44] Domestic dog.[13]
13,500 approx Mezine, Chernigov region, Ukraine Ancient dog-like skull proposed as being a Paleolithic dog.[6] Additionally, 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] Taxonomy uncertain.[13]
13,500 approx Mezhirich, Ukraine Ancient dog-like skull proposed as being a Paleolithic dog.[6] 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] Taxonomy uncertain.[13]
13,000 Palegawra, Iraq Mandible[10]
12,800 Ushki I, Kamchatka, eastern Siberia Complete skeleton buried in a buried dwelling.[45] Located 1,800 km to the southeast from Ulakhan Sular. Domestic dog.[29]
12,790 Nanzhuangtou, China 31 fragments including a complete mandible[46]
12,500 Kartstein cave, Mechernich, Germany Ancient dog skull. In 2013, the DNA sequence was identified as a dog.[22]
12,500 Le Morin rockshelter, Gironde, France Skeletal remains of dogs.[47] Domestic dog.[13]
12,450 Yakutia, Siberia Mummified carcass. The "Black Dog of Tumat" was found frozen into the ice core of an oxbow lake steep ravine at the middle course of the Syalaah River in the Ust-Yana region. DNA analysis confirmed it as an early dog.[48]
12,300 Ust'-Khaita site, Baikal region, Central Asia Sub-adult skull located 2,400 km southwest of Ulakhan Sular and proposed as a Paleolithic dog.[29]
12,000 Ain Mallaha (Eynan) and HaYonim terrace, Israel Three canid finds. A diminutive carnassial and a mandible, and a wolf or dog puppy skeleton buried with a human during the Natufian culture.[49] These Natufian dogs did not exhibit tooth-crowding.[50] The Natufian culture occupied the Levant, and had earlier interred a fox together with a human in the Uyun al-Hammam burial site, Jordan dated 17,700–14,750 YBP.[51]
12,000 Grotte du Moulin cave in Troubat, France Two dogs were buried together in the by the Azilian culture.[52]
10,700 Pont d'Ambon, Dordogne, France A number of skeletal remains of dogs.[38] In 2013, the DNA sequence was identified as a dog.[22] Domestic dog.[13]
10,150 Lawyer's Cave, Alaska, USA Bone of a dog, oldest find in North America. DNA indicates a split from Siberian relatives 16,500 YBP, indicating that dogs may have been in Beringia earlier. Lawyer's Cave is on the Alaskan mainland east of Wrangell Island in the Alexander Archipelago of southeast Alaska.[53]
9,900 Koster Site, Illinois, USA Three dog burials, with another single burial located 35 km away at the Stilwell II site in Pike County.[54]
9,000 Jiahu site, China Eleven dog interments. Jaihu is a Neolithic site 22 kilometers north of Wuyang in Henan Province.[55]
8,000 Svaerdborg site, Denmark Three different sized dog types recorded at this Maglemosian culture site.[56]
7,425 Baikal region, Central Asia (Russia) Dog buried in a human burial ground. Additionally, a human skull was found buried between the legs of a "tundra wolf" dated 8,320 BP (but it does not match any known wolf DNA). The evidence indicates that as soon as formal cemeteries developed in Baikal, some canids began to receive mortuary treatments that closely paralleled those of humans. One dog was found buried with four red deer canine pendants around its neck dated 5,770 BP. Many burials of dogs continued in this region with the latest finding at 3,760 BP, and they were buried lying on their right side and facing towards the east as did their humans. Some were buried with artifacts, e.g., stone blades, birch bark and antler bone.[57]
7,000 Tianluoshan archaeological site, Zhejiang province, China In 2020, an mDNA study of ancient dog fossils from the Yellow River and Yangtze River basins of southern China showed that most of the ancient dogs fell within haplogroup A1b, as do the Australian dingoes and the pre-colonial dogs of the Pacific, but in low frequency in China today. The specimen from the Tianluoshan archaeological site is basal to the entire lineage. The dogs belonging to this haplogroup were once widely distributed in southern China, then dispersed through Southeast Asia into New Guinea and Oceania, but were replaced in China 2,000 YBP by dogs of other lineages.[58]

Early domestication debate

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Among archeologists, the proposed timing of the development of a relationship between humans and wolves is debated. There exists two schools of thought.[18] The early domestication theory argues that the relationship commenced once humans moved into the colder parts of Eurasia around 35,000 YBP, which is when the proposed Paleolithic dogs first began to appear.[6][27][59] Wolves that were adjusting to live with humans may have developed shorter, wider skulls and more steeply-rising foreheads that would make wolf facial expressions easier to interpret.[18] The late domestication theory argues that Paleolithic dogs are an unusual phenotype of wolf and that dogs appeared only when they could be phenotypically distinguishable from the wolf, which is usually based on a reduction in size.[60][61][62][34][63] This argument maintains that domesticated dogs are more clearly identified when they are associated with human occupation, and those interred side by side with human remains provide the most conclusive evidence,[61] commencing with the 14,200 years old Bonn-Oberkassel dog.

The debate centres around Homo sapiens and if they had entered into cooperation with wolves soon after they moved into Eurasia, and if so when and where did these wolves change into domesticated dogs. In arguing that domestication leads to reduction in size, the late domestication theory ignores that modern horses and pigs are larger than their wild ancestors. It also ignores that if hunter-gathers entered into a hunting relationship with wolves then there would be no need of selection for a reduction in size. A reduction in size would have occurred much later when humans moved into agricultural villages. The late domestication theory does not consider the possibility that humans may have formed a relationship with non-domesticated wolves and that dogs in the early stages of domestication might be indistinguishable from wolves. According to indigenous North Americans, over the past 20,000 years the canids living with them were wolves that could not be distinguished as dogs.[18]

The problem in attempting to identify when and where domestication occurred is the possibility that the process of domestication occurred in a number of places and at a number of times throughout prehistory.[18] Early dog remains have been found in different parts of the world. This suggests that dog domestication may have taken place in different regions independently by hunter-gatherers, in some cases at the same time[64] and in other cases at different times,[65] with different wolf subspecies producing different dog lineages.[66][65] Therefore, the number of dog domestication events is not known.[67] A study of the maternal mitochondrial DNA (mDNA) shows that dogs fall within 4 mDNA clades,[22] indicating that dogs are derived from 4 separate lineages and therefore there may not have been a single domestication event.[18]

A domestication study looked at the reasons why the archeological record that is based on the dating of fossil remains often differed from the genetic record contained within the cells of living species. The study concluded that our inability to date domestication is because domestication is a continuum and there is no single point where we can say that a species was clearly domesticated using these two techniques. The study proposes that changes in morphology across time and how humans were interacting with the species in the past needs to be considered in addition to these two techniques.[68]

..."wild" and "domesticated" exist as concepts along a continuum, and the boundary between them is often blurred — and, at least in the case of wolves, it was never clear to begin with.

— Raymond Pierotti[69]

Relationship to the domestic dog

[edit]
Main article: Origin of the domestic dog

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.[22]

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, 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][21]

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), Kostenki-8 (Russia 23,000-27,700 YBP), Kostenki-1 (Russia 22,000-24,000 BP), Kostenki-17 (Russia Upper Paleolithic) and Verholenskaya (Russia late glacial).[26] 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.[27]

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][21] In 2016, a study discounted the use of the Paleolithic dogs from the Predmosti site as pack animals.[70]

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,[60] a counter to the refutation,[71] a second article,[27] its refutation,[61] a third article that includes a counter to the refutation,[26] its refutation,[62] a counter to the refutation,[72] another refutation,[34][63] support given based on bone collagen analysis,[1] and the identification of an ancient paleolithic dog in Yakutia.[29]

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

Two domestication events

[edit]

Studies have suggested that it was possible for multiple primitive forms of the dog to have existed, including in Europe.[73] European dog populations had undergone extensive turnover during the last 15,000 years that has erased the genomic signature of early European dogs,[74][75] the genetic heritage of the modern breeds has become blurred due to admixture,[56] and there was the possibility of past domestication events that had died out or had been largely replaced by more modern dog populations.[74]

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 and 6,400 YBP where they partially replaced the dogs of Europe.[76] Two domestication events in Western Eurasia and Eastern Eurasia have recently been found for the domestic pig.[76][77]

As the taxonomic classification of the "proto-dog" Paleolithic dogs as being either dogs or wolves remains controversial, they were excluded from the study.[76]

Goyet dog

[edit]
Genus Canis, species indeterminate
Artist's rendition of the Goyet dog skull

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 YBP) and so was identified as a Paleolithic dog.[6][78] The analysis indicated that the Belgian fossil large canids in general preyed on horse and large bovids.[6][27]

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.[34][33] 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 grey 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[22] as there was for domestic pigs.[77] A 2016 review proposed that it most likely represents an extinct morphologically and genetically divergent wolf population.[33]

Altai dog

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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".[79]

In 2007, a mtDNA analysis of extinct eastern Beringian wolves showed that two ancient wolves from Ukraine dated 30,000 YBP and 28,000 YBP and the 33,000 YBP Altai dog had the same sequence as six Beringian wolves,[80] 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.[24]

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[22] as there was for domestic pigs.[77]

In 2017, two prominent evolutionary biologists reviewed all of the evidence available on dog divergence and supported the specimens from the Altai mountains as being those of dogs from a lineage that is now extinct and that was derived from a population of small wolves that is also now extinct.[25]

Further information: Beringian wolf

Local unknown wolves

[edit]

Ecological factors including habitat type, climate, prey specialization and predatory competition will greatly influence wolf genetic population structure and cranio-dental plasticity.[81][82][83][84][85][86][87][88][89] Therefore, within the Pleistocene grey 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.[89]

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)[79] Kostenki 8, Russia (33,500-26,500 YBP)[72] Predmosti, Czech Republic (31,000 YBP)[27] 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.[72] These characteristics of shortened rostrum, tooth crowding, and absence or rotation of premolars have been documented in both ancient and modern wolves.[60][87][89][80][90][91] Rather than representing early dogs, these specimens may represent an extinct morphologically and genetically divergent wolf population.[33][56][89]

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]

In 2021, a study found that the cranial measurements of a number of Paleolithic dog specimens exhibited a relatively shorter skull and a relatively wider palate and brain case when compared with Pleistocene and recent northern wolves, and that these features are the morphological signs of domestication.[92]

References

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Bibliography

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Paleolithic dog

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The Paleolithic dog encompasses the earliest archaeological and genetic evidence of (Canis lupus) by humans during the period (approximately 2.6 million to 10,000 years ago), marking the first instance of animal and reflecting a pivotal human-animal partnership that likely began in around 33,000 years ago. This process involved the gradual transformation of scavenging into smaller, more docile companions, with early canid remains identified from sites like Razboinichya Cave in Siberia's , where a 33,000-year-old canid skull shows some morphological differences from wild , though recent analyses question whether these represent early . However, this early association appears to have been disrupted by the (around 26,500–19,000 years ago), with sustained emerging later, with genetic evidence suggesting multiple origins across , potentially from an extinct population, with major lineages diversifying by around 11,000 years ago. Key fossil evidence for these early dogs dates primarily to the Late (ca. 40,000–10,000 years ago) in , where remains from sites such as Bonn-Oberkassel in (14,700 years old) represent the oldest confirmed domestic , featuring a juvenile canid interred with remains and showing signs of severe illness that suggest human care and emotional bonding. This specimen, along with others from Eliseevichi in (17,000–13,000 years old), displays reduced body size, altered cranial morphology, and dietary shifts indicative of dependence on human-provided food, distinguishing them from contemporaneous wolves. Genetic analyses further reveal that by 15,000 years ago, dogs had begun to diverge into distinct lineages, with recent 2025 morphometric analyses indicating that distinctive morphologies and significant diversity first appeared around 11,000 years ago. These findings underscore a complex, multiregional history tied to societies, where dogs likely assisted in , guarding, and scavenging camp refuse. The role of Paleolithic dogs extended beyond utility, as evidenced by their inclusion in burials and potential symbolic significance in human , with isotopic studies showing diets enriched in human-associated resources like and , contrasting with wild canid patterns. Despite ongoing debates over the precise timing and exclusivity of events—some evidence points to multiple origins in and —with ongoing debates in research pointing to multiple events across during the and early , dogs accompanied human dispersals and adapted to diverse environments by the end of the . This foundational relationship laid the groundwork for the global diversification of Canis familiaris, influencing subsequent Neolithic developments in .

Classification and Nomenclature

Taxonomy

Paleolithic dogs belong to the kingdom Animalia, phylum Chordata, class Mammalia, order Carnivora, family Canidae, genus Canis, and are typically classified under the species Canis lupus as an early or incipient form of the subspecies Canis lupus familiaris, though some authorities use Canis cf. familiaris to indicate uncertainty in their domesticated status. This placement reflects their close relation to the gray wolf (Canis lupus), from which they diverged as a basal Eurasian lineage, with ancient DNA analyses estimating the split between dog and wolf lineages at approximately 27,000 to 40,000 years ago. More recent genomic studies have prompted debates on elevating domestic dogs to full species status as Canis familiaris, distinct from the gray wolf (Canis lupus), due to deep genetic divergence. However, their exact taxonomic position remains debated, with proposals ranging from a distinct species or subspecies to a transitional "proto-dog" form representing the onset of domestication rather than a fully separate entity. Key taxonomic debates center on whether Paleolithic dogs constitute a separate evolutionary lineage or merely morphological variants within populations, influenced by factors like regional isolation and association. Proponents of a distinct status argue for recognition as an incipient domestic form based on genetic evidence of early divergence, while critics emphasize the lack of consistent morphological separation, suggesting many specimens may simply be wolves with atypical traits. These discussions highlight the challenges in applying modern taxonomic criteria to ancient remains, where hybridization and blur boundaries. Distinguishing Paleolithic dogs from contemporaneous Pleistocene wolves relies primarily on cranial metrics, such as a shortened and broader , reduced in certain dimensions though with significant overlap in overall size with contemporaneous wolves, and more compactly arranged teeth, including crowded . However, these features show significant overlap, with many classical criteria—like tooth crowding, development, and orbital angle—proving non-diagnostic due to variability in wild canids and inconsistencies in early classifications. Advanced 3D geometric morphometric analyses further reveal that some purported Paleolithic dogs, such as those from Goyet and Eliseevichi, align more closely with wolf morphology, lacking pronounced domestic traits like a dorsally angled muzzle or distinct "stop."

Etymology and Naming Conventions

The term "Paleolithic dog" emerged in early 20th-century archaeological literature to designate canid remains recovered from Upper Paleolithic contexts, roughly spanning 50,000 to 10,000 BCE, where morphological traits suggested possible early human association or incipient domestication. This descriptive nomenclature drew from broader discussions of prehistoric faunal remains and was notably shaped by Russian-language scholarship, including the contributions of Ukrainian archaeologist Ivan Pidoplichko, whose excavations at mammoth-bone dwellings in Ukraine during the mid-20th century employed analogous concepts like "proto-dog" (proto-sobaka in Russian) to characterize canids co-occurring with human artifacts in Paleolithic layers. Over time, naming conventions for these ancient canids have varied, reflecting interpretive shifts in and ; alternative terms include "prehistoric dog," " dog," and site-specific designations such as " canid," which emphasize cultural or chronological associations rather than definitive . For instance, F.E. Zeuner's influential 1963 monograph A of Domesticated Animals adopted "prehistoric dog" for early post-Pleistocene specimens, framing them within a narrative of gradual wolf-to-dog transition based on skeletal evidence. Contemporary revisions, particularly following ancient DNA analyses in the 2010s, have prompted more nuanced terminology, with some researchers favoring "sympatric wolf" or "large Paleolithic canid" to denote remains previously labeled as dogs but now interpreted as morphologically distinct wolves living alongside humans without full domestication. These shifts, exemplified in studies like Germonpré et al. (2012) on Gravettian skulls, underscore ongoing debates and the provisional nature of naming in light of genetic data challenging early domestication claims.

Morphology and Biology

Physical Characteristics

Paleolithic dogs generally exhibited smaller overall body sizes compared to their contemporaries, with estimated shoulder heights averaging 50–60 cm and a lighter, less robust frame that distinguished them from the bulkier builds of wild canids. This lighter construction is evident in remains, where limb bones show reduced robusticity, suggesting adaptations to a more sedentary or human-associated lifestyle rather than the endurance hunting typical of wolves. Regional variations are notable, with European specimens often displaying more robust skeletal proportions suited to colder climates, while Asian forms tended toward slimmer, more gracile builds, potentially reflecting local environmental influences or early selective pressures. Cranial morphology in dogs featured a reduced braincase volume, approximately 10–15% smaller relative to body size than in wolves, alongside shorter and broader palates that altered facial proportions. These traits, including a viscerocranium averaging around 114 mm compared to 131 mm in wolves, indicate the onset of domestication-related changes, such as and reduced predatory specialization. Dental features further highlight this transition, with crowded —where the upper fourth premolar and first molar showed overlapping or reduced spacing—and slightly shorter (around 25 mm versus 26–27 mm in wolves), reflecting dietary shifts and paedomorphic development. Pathologies and anomalies in Paleolithic dog remains provide insights into their interactions with humans, including evidence of healed injuries that imply caregiving. A notable example is the 16,000-year-old discovered in 2025 from a cave in , which exhibited multiple healed vertebral fractures, indicating survival post-injury likely due to human intervention. patterns in these early dogs mirrored those of wolves to some extent, with males typically showing 10–20% larger skulls and more pronounced sagittal crests, though the overall variance was reduced compared to wild populations, possibly due to controlled breeding or provisioning. Dental wear patterns in these specimens occasionally suggest softer, human-provided diets, with implications for behavioral adaptations explored further in dietary analyses.

Dietary Habits

Stable isotope analysis of bone collagen from Upper Paleolithic canid remains indicates that early dogs consumed a mixed diet primarily comprising terrestrial mammals, such as and , supplemented by human-provisioned food scraps. In sites from southwestern and (ca. 17,000–12,000 years BP), δ¹³C values for these canids ranged from -20.5‰ to -18.2‰, reflecting a reliance on C₃ terrestrial herbivores, while δ¹⁵N values for putative dogs were elevated compared to contemporaneous wolves, often by 2–4‰ (e.g., dog averages around 9–11‰ versus wolf 7–9‰), suggesting access to higher-trophic-level resources likely derived from human hunting byproducts. This isotopic signature points to an ecological role as opportunistic in human-modified environments, with less dependence on independent predation than wolves. Dental microwear texture analysis further supports scavenging behaviors in dogs, revealing patterns of heavy tooth attrition consistent with frequent bone-crushing. At the site of Předmostí, (ca. 28,500 years ), the second molars (M₂) of protodog specimens exhibited significantly higher complexity (Smfc) values than those of , indicating greater durophagy—chewing of hard, brittle materials like bone fragments from carcasses—while (M₁) microwear showed no notable differences, implying comparable flesh consumption. Compared to wolf diets, which displayed more varied microwear suggestive of diverse prey, these patterns in canid remains highlight a narrower, more specialized scavenging niche for early dogs, potentially tied to proximity to kill sites. Regional variations in dietary evidence underscore adaptations to local ecosystems. In European contexts, such as and assemblages, early dogs showed signs of omnivory through dietary shifts. In Siberian sites, isotopic profiles from canids (e.g., Tumat, ca. 14,000 years BP) reveal diets incorporating like alongside diverse terrestrial proteins, with δ¹⁵N values (8–10‰) elevated but lacking strong human provisioning signals; recent analyses (2023–2025) confirm megafaunal elements in broader regional canid diets yet rule out based on independent patterns. These differences reflect environmental influences, with European dogs showing more commensal traits and Siberian ones retaining wild-like breadth.

Archaeological Evidence

Earliest Known Specimens

Potential early specimens of Paleolithic dogs or proto-dogs have been reported from sites across , with debated attributions dating back to the and periods. Among the oldest candidates in is the Goyet Cave specimen from , dated to approximately 36,000–31,700 years ago, featuring a canid skull with dog-like cranial proportions based on , though recent 2025 biometric studies challenge its domestic status, suggesting it may represent a large or transitional form rather than a fully domesticated . Another early site is Předmostí in the , from the period. Excavated since the late , the site yielded over 4,000 canid bones, including fragmented mandibles and skulls, associated with human settlements and bone accumulations. of associated organic materials places these remains in the main cultural layer at approximately 26,000–27,000 uncalibrated years BP (about 31,000–29,000 cal BP), supported by stratigraphic context from later excavations in the 2000s. Initial interpretations in the early classified some canids as wolves due to their size and the site's hunting-focused economy, but 19th-century reports had tentatively noted dog-like features amid the faunal assemblage. Morphometric analyses of 37 mandibles from Předmostí reveal traits potentially signaling incipient domestication, such as shorter overall length (averaging 10–15% reduced compared to contemporaneous wolves), massive jaw structure, and crowded premolars in 60% of specimens—features less common in regional Pleistocene wolf samples. These characteristics, evaluated through discriminant function analyses on eight mandibular measurements, have been proposed to differentiate a "Paleolithic dog" morphotype from local wolves, suggesting possible human selection pressures by around 25,000 cal BP, though 2025 studies question this interpretation, indicating they may be wolves. However, pre-1950s excavations at the site introduced potential contamination risks for later radiocarbon assays, as bone samples were handled without modern protocols, leading to occasional re-evaluations of associated dates for accuracy. A later but iconically preserved specimen originates from the Bonn-Oberkassel site in , discovered in 1914 during basalt quarrying near . The remains consist of a partial juvenile ( fragments, vertebrae, and long bones) buried intentionally with two adult humans (a man aged ~40 and a woman aged ~25) and like artifacts, covered by red and stone slabs in a Late Magdalenian context. of from the dog's tooth yields 14,223 ± 58 (calibrated to ~14,700 cal BP), corroborated by dates from the human remains and consistent stratigraphic layering, though early 20th-century recovery methods risked minor contamination from quarry dust. Contemporary analyses confirm its domestic status through , including a reduced snout length indicative of paedomorphic traits, and evidence of treated by humans, marking it as one of the earliest clear cases of human-canine interment. Early 20th-century scholars debated the Bonn-Oberkassel find as a due to its robust build and the era's limited comparative data, but post-1950s osteometric studies resolved it as a , aligning with broader re-evaluations of canids showing signals by ~15,000 cal BP at confirmed sites. These specimens highlight the transition from wild canids to managed companions, with sites like Goyet and Předmostí representing potential early but debated evidence, and Bonn-Oberkassel evidencing significance.

Catalog of All Identified Remains

The catalog of verified Paleolithic dog remains relies on post-2010s standards, including ancient DNA (aDNA) sequencing to confirm genetic divergence from wolves and 3D geometric morphometric analyses to assess cranial and postcranial features indicative of domestication, such as reduced robusticity and altered snout morphology. These criteria exclude ambiguous specimens lacking such evidence, such as the Tumat puppies from Siberia, which a 2025 genetic study identified as wolf cubs rather than proto-dogs based on mitochondrial DNA aligning closely with Pleistocene wolves. In , the most extensive inventory of remains comes from sites, with key examples including the Goyet Cave specimen from , dated to approximately 36,000–31,700 years ago, featuring a canid with dog-like cranial proportions confirmed via and aDNA showing divergence from contemporaneous wolves, though recent 2025 analyses debate its full . Another prominent find is from Eliseevichi 1 in , yielding two partial crania and mandibles dated to around 17,000–13,000 years ago, with morphometric analysis revealing shortened snouts and crowded distinct from local wolves, supported by stable data indicating a mixed diet. A recent addition is a nearly complete from a cave in (Grotte de Pataud or similar site), dated to 16,000–15,300 years ago, exhibiting healed vertebrae and a that suggest human care, alongside possible evidence of later human-inflicted trauma; verified through aDNA and osteological examination in a 2025 study, this highlights complex early human-dog relationships. Other European sites, such as Bonn-Oberkassel in (~14,700 cal ), include dog remains buried with humans, but the catalog emphasizes verified cases meeting modern criteria, totaling over a dozen specimens across the continent. Asian remains are sparser but include the Razboinichya Cave find from the in , a well-preserved lower and teeth dated to 33,000 years ago, classified as an incipient through aDNA analysis revealing a unique intermediate between ancient wolves and modern dogs, though disrupted by the . Additional fragmentary evidence from Siberian and Central Asian sites supports localized populations, but verification remains limited to fewer than five robustly identified cases. Evidence from the is minimal and primarily postdates the core period in , with pre-Clovis sites yielding sparse remains; a notable example is an 8,100-year-old jawbone from Hollembaek Hill in , analyzed in 2024, showing transitional morphology and a salmon-heavy diet via isotopes, but classified as post-Paleolithic due to its dating despite early human association. No confirmed pre-10,000-year-old remains meet aDNA and morphometric standards in the Americas, highlighting migration lags from Eurasian lineages. Significant gaps persist in the record, particularly for and , where no verified Paleolithic dog remains have been identified despite human presence; African sites show only later introductions, while Australian derive from post-Paleolithic East Asian dogs arriving around 5,500 years ago, underscoring the need for interdisciplinary efforts combining , , and isotopic to address these underrepresented regions.

Domestication Debates

Origins and Timing of Domestication

The debate surrounding the origins and timing of dog domestication centers on the transformation of wolves (Canis lupus) into the earliest domestic dogs during the , with proposals ranging from as early as approximately 36,000 years ago in to more conservative estimates of 15,000–16,000 calibrated years (cal BP). Early claims for 33,000–36,000 years ago stem from and morphological analyses of specimens like those from Goyet Cave in and Razboinichya Cave in , potentially representing proto-dogs in or , though these are contested due to overlapping wolf populations and lack of clear archaeological support. In contrast, conservative timelines, supported by morphological and contextual evidence from sites like those in the and , place sustained domestication around 15,000 cal BP, following the end of the when human hunter-gatherers may have begun forming closer associations with wolves. A prominent hypothesis posits self-domestication through natural selection for tameness, where bolder, less aggressive wolves with reduced fear responses toward humans survived better near human settlements, leading to gradual behavioral and physical changes without direct human intervention. This aligns with the commensal pathway model, in which wolves initially acted as opportunistic scavengers around Paleolithic camps, benefiting from food waste while humans tolerated or encouraged their presence, fostering a mutualistic relationship over generations. Charles Darwin, in his 1868 work The Variation of Animals and Plants under Domestication, viewed dog origins through the lens of artificial selection, proposing descent from multiple wild canids including wolves but emphasizing human-driven breeding for traits like docility and utility in hunting. Modern interpretations build on this by integrating the commensal model, highlighting how initial natural selection preceded later human management. Key evidence includes behavioral indicators, such as deliberate burials of dogs alongside humans, suggesting emotional bonds; for instance, the Bonn-Oberkassel dog from , dated to about 14,700 cal BP, was interred with and , implying ritual significance. Morphological changes, like —the retention of juvenile traits such as shorter snouts, floppy ears, and reduced —appear in some early specimens, interpreted as byproducts of selection for tameness, as demonstrated in analogous experiments. However, critiques note that reliance on size reduction or can be overstated, as many canid remains show minimal differences from , and environmental factors like may explain variations better than alone. Recent 2025 studies have advanced these models, with computational simulations demonstrating that via natural and could transform into dog-like forms in as little as 8,000–15,000 years, potentially aligning with a around 23,000 years ago in based on from isolated wolf populations. These findings underscore social cognition differences, such as enhanced human-directed and reduced , as early drivers, complicating traditional timelines by suggesting proto-domestication phases predated full integration into human societies.

Evidence for Multiple Domestication Events

Genetic studies of () from canids have provided evidence supporting a dual-event model of , positing independent origins from distinct populations in eastern and western . This hypothesis emerged from analyses revealing a deep genetic split between East Asian and Western Eurasian dog lineages, with the eastern branch diverging earlier, around 33,000 years ago, and the western around 23,000–14,000 BCE. The model is bolstered by genomic data indicating minimal between these lineages initially, suggesting separate processes disrupted by the (), followed by later admixture. The eastern domestication event is exemplified by the Altai lineage, represented by an incipient dog specimen from Razboinichya Cave in , dated to approximately 33,000 calibrated years (cal BP). Morphological analysis of the showed a shortened, broad and compact more akin to modern s than contemporary wolves, while genetic sequencing placed it basal to East Asian dog populations but distinct from late Glacial–early lineages. This specimen suggests early human-wolf interactions in leading to , with the lineage likely contributing to later East Asian dogs but facing or severe bottlenecks during the LGM due to climatic shifts. Recent from , , dated to around 13,000 BCE, further supports this eastern wave by confirming domesticated dogs with affinities to Siberian ancestors, indicating persistence and spread southward. In western , the Goyet lineage from Goyet Cave in , dated to about 36,000 cal , represents an early basal branch to some European dogs, characterized by unique genomic signatures showing admixture from unknown populations. This lineage, identified through , diversified during the but appears to have gone extinct around the LGM, with ghost lineages detectable in later European dogs via admixture models from 2016–2025 studies. These findings highlight multiple waves, where initial domestications produced distinct lineages that did not all survive environmental pressures. A November 2025 genomic of ancient dogs further reinforces this model, showing that major morphological diversification and evidence of multiple origins were already present by around 11,000 years ago, with dogs accompanying human migrations across . Critiques of the multiple-events model emphasize a single origin from an extinct , with early diversification into at least five major ancestries by 11,000 years ago, rather than fully independent domestications. Recent 2025 analyses, such as those of the Tumat specimens from (dated ~14,000 cal BP), have ruled out some candidates as domesticated dogs, classifying them as based on genomic, osteometric, and dietary evidence showing no association. This ongoing debate underscores the role of in refining hypotheses, with dual ancestry patterns potentially arising from regional contributions rather than separate full domestications.

Genetic Relationships

Genetic analyses of ancient DNA from Paleolithic canids demonstrate substantial continuity with modern domestic dogs, revealing a shared ancestry that diverged from an extinct wolf population between 27,000 and 40,000 years ago. This common lineage, distinct from present-day wolves, experienced limited gene flow from wolves post-domestication, with substantial bidirectional introgression shaping subsequent evolution. By approximately 11,000 years ago, at least five major ancestry components had emerged during the Paleolithic, providing the genetic foundation for contemporary dog populations across Eurasia and beyond. A November 2025 study analyzing ancient dog genomes confirms that substantial genetic diversity, including multiple ancestry components, had already emerged by the end of the Paleolithic, supporting multiregional contributions to modern dogs. Mitochondrial DNA haplogroups, such as the now-rare C prevalent in early European dogs, link specimens to Eurasian founders, though a matrilineal turnover to A occurred later in . Y-chromosome studies further support this continuity, identifying haplogroups distributed among global dog populations that trace back to ancient Eurasian origins, with ancient Northeast Siberian dogs contributing to lineages seen in modern breeds. Ancient European dogs from the Early share 70–80% ancestry with modern European dogs, underscoring regional persistence despite later admixtures. Morphological legacies from dogs persist in certain basal breeds, which retain primitive traits such as reduced cranial size and overall body proportions indicative of early . For instance, breeds like the and exhibit genetic and physical characteristics— including compact builds and alert postures—that align with ancient canid morphologies, positioning them near the base of the domestic dog . These traits reflect selective retention from pre-agricultural ancestors, contrasting with the exaggerated forms developed through later intensive breeding. Post-Pleistocene shifts and human migrations contributed to the or replacement of several dog lineages, resulting in genetic bottlenecks that reduced diversity. For example, the near-complete turnover of European dog ancestry, including the decline of C to less than 10% in modern populations, likely stemmed from population displacements during the spread of farming communities and environmental changes at the end of the Ice Age. In the , ancient dog lineages experienced a bottleneck around 16,400 years ago, linked to migrations across , with subsequent isolation leading to unique now nearly . Analyses as of 2025, building on 2022 genomic studies, estimate the primary dog- divergence at over 30,000 years ago, with ongoing low-level influencing peripheral modern populations but not erasing core signals. These roots underpin the origins of modern village dogs, which preserve broader compared to highly bred varieties, highlighting how initial in Eurasian contexts set the stage for global dispersal. Later amplified specific traits but built upon this ancient foundation, with basal breeds serving as living bridges to prehistoric canids. Evidence for multiple events contributed to this foundational diversity.

Distinct Lineages and Local Wolf Influences

Genetic analyses of ancient canid remains have identified the Goyet specimen from , dated to approximately 36,000 years ago, as representing a short-lived European lineage that diverged early from the main dog ancestry and became extinct by around 15,000 years ago. This branch, characterized by morphological features intermediate between wolves and later dogs, likely arose from an aborted domestication attempt or a distinct population that did not contribute significantly to modern domestic dogs. In contrast, the Altai canid from Siberia's Razboinichya , dated to about 33,000 years ago, exhibits genetic affinities with primitive dogs and shows evidence of local wolf admixture, reflecting adaptations to the harsh Siberian environment such as enhanced cold tolerance through introgression from regional Pleistocene wolves. Its mitochondrial clusters closely with modern dogs and prehistoric canids, indicating a basal position in dog with limited divergence from ancient wolves (2–5 differences compared to 4–26 for contemporary wolves). Evidence from () studies highlights contributions from untraced or "ghost" populations to early genomes, with substantial unsampled ancestry from extinct northern Eurasian lineages that cannot be accounted for by known European, Siberian, or North American samples. These ghost lineages, potentially divergent since before 100,000 years ago, suggest admixture from extinct local groups during the . Recent research from 2023 to 2025 has revealed such ghost contributions in Siberian and American contexts; for instance, pre-contact dogs in the trace to a single maternal lineage from around 23,000 years ago, with genetic structuring across North, Central, and indicating isolation by distance and possible untraced inputs during southward migrations with early agrarian societies (7,000–5,000 years ago). Regional wolf influences varied markedly between and during the , driven by geographic isolation that fostered parallel evolutionary paths in canid populations. Paleolithic dogs in show greater affinity to local western Eurasian wolves, while Asian lineages, including Siberian ones, derive more from eastern Eurasian wolves, with modern dogs overall closer to ancient Siberian wolves (23,000–13,000 years ago) than European counterparts. This divergence, exacerbated by glacial barriers, resulted in no evidence of Paleolithic dog remains or lineages in , where wolf populations were absent or ecologically distinct. A 2025 model proposed by biologists emphasizes earlier dog-wolf divergence prior to the , facilitated by local (co-occurrence) of wolves and proto-dogs in multiple regions, and classifies the Taimyr Peninsula specimen (35,000 years old) as a non-dog wolf lineage that contributed 1–27% ancestry to breeds through admixture rather than direct descent.

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