Cave wolf

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Cave wolf
Temporal range: Middle Pleistocene - Late Pleistocene
320,000—12,000 years ago
Canis lupus spelaeus holotype.jpg
The Goldfuss holotype, [1] Berlin's Natural History Museum
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Family: Canidae
Genus: Canis
Species:
Subspecies:
C. l. spelaeus
Trinomial name
Canis lupus spelaeus
Goldfuss, 1823 [2]
Synonyms [3]

C. l. brevis Kuzmina and Sablin, 1994 [4]
C. l. maximus Boudadi-Maligne, 2012 [5]

Contents

The cave wolf (Canis lupus spelaeus) is an extinct glacial mammoth steppe-adapted white wolf that lived during the Middle Pleistocene to the Late Pleistocene. It inhabited Europe, where its remains have been found in many caves. Its habitat included the mammoth steppe grasslands and boreal needle forests. [6] This large wolf was short-legged compared to its body size, [4] yet its leg size is comparable with that of the Arctic wolf C. l. arctos. [6] Mitochondrial DNA analysis shows it to be more closely related to the domestic dog than the modern wolf, indicating possible ancestry. [7] [8]

Taxonomy

The large wolf C. l. spelaeus Georg August Goldfuss, 1823 was first described based on a wolf pup skull found in the Zoolithen Cave located at Gailenreuth, Bavaria, Germany. [2] It is regarded as an ecomorph/chronosubspecies of C. lupus. [9] In Germany, cave wolf populations are known from three caves separated by a few kilometers from each other that are located on the Franconia Karst along the Wiesent and Ahorn River vallies in the Upper Franconia region of the state of Bavaria, Germany. Sophie's Cave sits on the northwest slope of the Ailsbach Valley near Rabenstein Castle in the Ahorntal municipality. Große Teufels Cave (Big Devil's Cave) and the Zoolithen Cave are located nearby. [10] They are also known from Hermann's Cave in the village of Rübeland near the town of Wernigerode, in the district of Harz, Saxony-Anhalt, Germany. [11]

These large wolves have not been studied in any degree of depth, and their relationship to modern wolves has not been clarified using DNA. [12]

Canis lupus bohemica

In 2022, a new species Canis lupus bohemica was taxonomically described after having been discovered in the Bat Cave system located near Srbsko, Central Bohemia, Czech Republic. The Bohemian wolf is an extinct short-legged wolf that first appeared 800,000 years ago (MIS 20, the Rhumian Interglacial of the early Cromerian stage, Middle Pleistocene) and once inhabited what was part of the mammoth steppe. It is proposed as the ancestor of Canis lupus mosbachensis. In comparison, C. etruscus appears to be the ancestor of the Afro-Eurasian jackal. [6]

In Hungary in 1969, a tooth (the premolar of the Maxilla) was found which dated to the Middle Pleistocene, and was assessed as being midway between that of Canis mosbachensis and the cave wolf Canis lupus spelaeus, but leaning towards C.l. spelaeus. [13] During the late Middle Pleistocene around 600,000 years ago, the Bohemian wolf diversified into two wolf lineages that specialized for different environmental and climatic conditions. One was a southern interglacial (warm climate) grey wolf of Europe which was to become the Mosbach wolf, and the other a northern glacial white wolf of Eurasia which was to become C. l. spelaeus. [6]

Canis lupus brevis

The Don wolf Canis lupus brevis Kuzmina and Sablin, 1994 is an extinct wolf whose fossil remains were found at the Kostenki I Late Pleistocene site by the Don River at Kostyonki, Voronezh Oblast, Russia and reported in 1994. Based on the size of its dental rows, the Don wolf was bigger than modern wolves from the tundra or the Middle Russian taiga. The length of its P4 was longer than the row of its molars M1-M2, which is different to the Late Pleistocene wolves from the Caucasus or the Ural Mountains. Its main characteristic was its shorter legs, due to shorter humerus, radius, metacarpals, tibia, and metatarsals. [4] This warm climate grey wolf existed at the same time as C. l. spelaeus. [6] In 2009, a study compared the skull of one of these wolves to those found in Western Europe, and proposed that C. l. brevis of eastern Europe and C. l. spelaeus of western Europe are taxonomic synonyms for the same subspecies. [3] [6] These were both comparable to the remains of a 16,000 YBP wolf skull found on the Taimyr peninsula in far northern Siberia. [3]

Canis lupus maximus

The large wolf Canis lupus maximus Boudadi-Maligne, 2012 was a subspecies larger than all other known fossil and extant wolves from Western Europe. The fossilized remains of this Late Pleistocene subspecies were found across a wide area of south-western France at: Jaurens cave, Nespouls, Corrèze dated 31,000 YBP; Maldidier cave, La Roque-Gageac, Dordogne dated 22,500 YBP; and Gral pit-fall, Sauliac-sur-Célé, Lot dated 16,000 YBP. The wolf's long bones are 10 percent longer than those of extant European wolves and 20 percent longer than its probable ancestor, C. l. lunellensis. The teeth are robust, the posterior denticules on the lower premolars p2, p3, p4 and upper P2 and P3 are highly developed, and the diameter of the lower carnassial (m1) were larger than any known European wolf. [5]

Wolf body size in Europe has followed a steady increase from their first appearance up to the peak of the Last Glacial Maximum. The size of these wolves are thought to be an adaptation to a cold environment (Bergmann's rule) and plentiful game as their remains have been found in association with reindeer fossils. [5]

In 2022, a paleontologist proposed that C. l. maximus was a taxonomic synonym for C. l. spelaeus. [6]

Italian wolves

A 2014 study found wolves in Late Pleistocene Italy were comparable in tooth morphology – and therefore in size – with C. l. maximus from France. These wolves were found near Avetrana, Taranto and near Buco del Frate, Brescia and Pocala cave in Friuli Venezia Giulia. [14]

Description

The short-legged C. l. spelaeus was a glacial mammoth steppe-adapted white wolf. It first appeared 320,000 years ago (MIS 8, Middle Pleistocene), and disappeared along with the mammoth steppe and cave bear fauna at the end of the Late Pleistocene, somewhere between 24,000-12,000 years ago (MIS 2). Compared with the modern day Arctic wolf C. l. arctos and the Siberian Tundra wolf C. l. albus, its legs are shorter than the Tundra wolf but similar to the Arctic wolf and may be its ancestor. [6]

During MIS 8-3, its habitat included the mammoth steppe grasslands and boreal needle forests. It spread across all of Europe through the middle-high elevated mountains, and followed mountain glaciers down to the Mediterranean. This wolf specialized in feeding on cave bear carcasses found inside of caves, and did more damage to its teeth in doing so than any other wolf subspecies. [6]

Its bone proportions are close to those of the Canadian Arctic-boreal mountain-adapted timber wolf and a little larger than those of the modern European wolf. Some postcranial bones have similarly large proportions to those from the Sophie's and Große Teufels Caves, [10] where the bone sizes are closer to those of Scandinavian Arctic and Canadian Columbian wolf subspecies than to those of the smaller European wolves. Bone sizes are one-eighth larger than European wolves, and this wolf was a specialized Late Pleistocene wolf ecomorph. [15]

Diagram of a wolf skull with key features labelled Wolf cranium labelled.jpg
Diagram of a wolf skull with key features labelled

During the Late Pleistocene, C. l. spelaeus exhibited a changed skeletal development due to the harsh climatic and environmental conditions, and a preference for taking larger prey. This resulted in a larger and more robust wolf with a shortened rostrum, a pronounced development of the temporalis muscles, and proportionally enlarged and wider premolars and carnassials. These features were specialised adaptations for the processing of fast-freezing carcasses associated with the hunting and scavenging of larger prey. Some populations of C. l. spelaeus showed an increase in tooth breakage when compared with the extant C. lupus because they were habitual bone crackers. The large dimensions, robust build and cranio-dental adaptations formed at this time enabled these wolves to hunt larger prey, making the adult Steppe Bison Bison priscus a more attainable target than for their C. lupus relatives. [8]

Adaptation

Cave wolf illustration Prehistorische dierenresten uit Noord-Brabant (1998) fig. 19 colorized.png
Cave wolf illustration

Their dens have been identified, with the Zoolithen Cave supporting a large population and has yielded more than 380 bones as well as several skulls (including a holotype). Sophie's Cave has demonstrated the first "Early Late Pleistocene wolf den", with intensive faecal places and the first European record of half-digested cave bear bones found within the faecal areas in the cave. It demonstrates that wolves seem not to have used this cave as a cub-raising den, but that they were cave dwellers that fed on cave bear carcasses, similar to but less so than cave hyenas, but more so than cave lions. The abundant faeces seem to play a role in the "orientation" for trail tracking, similar to modern wolves, and less as den marking. The high abundance in a limited area of the Bear's Passage of the cave might be the result of periodical short-term den use of smaller cave areas. Wolves were scavenging on the bears that hibernated and died there, and therefore a simultaneous use as both a wolf and a cave bear den cannot be expected. Remains of a skeleton of at least one high adult wolf also might have been the result of a battle within the cave with the bears, the same as in the lion taphonomic record. [12]

The ecology of the early to middle Late Pleistocene wolves on the mammoth steppe and the boreal forests is not known, nor is whether they used caves as dens. [12]

All of the top predators in Europe commenced going extinct with the loss of the pleistocene megafauna when conditions became colder during the peak of the Last Glacial Maximum around 23,000 years ago. The last cave wolves used the side branches of the main caves to protect their pups from the cold climate. [15] During this time the cave wolf was replaced by a smaller wolf-type, which then disappeared along with the reindeer, to finally be replaced by the Holocene warm-period European wolf Canis lupus lupus . [12]

Relationship with the dog

Mitochondrial DNA (mDNA) passes along the maternal line and can date back thousands of years. [16] Therefore, phylogenetic analysis of mDNA sequences within a species provides a history of maternal lineages that can be represented as a phylogenetic tree. [17] [18]

In 2013, a study analysed the complete and partial mitochondrial genome sequences of 18 fossil canids dated from 1,000 to 36,000 YBP from the Old and New Worlds, and compared these with the complete mitochondrial genome sequences from modern wolves and dogs. Phylogenetic analysis showed that modern dog mDNA haplotypes resolve into four monophyletic clades with strong statistical support, and these have been designated by researchers as clades A-D. [7] [19] [20]

Based on the specimens used in this study, clade A included 64% of the dogs sampled and these were sister to a 14,500 YBP wolf sequence from the Kessleroch cave near Thayngen in the canton of Schaffhausen, Switzerland, with a most recent common ancestor estimated to 32,100 YBP. This group of dogs matched three fossil pre-Columbian New World dogs dated between 1,000 and 8,500 YBP, which supported the hypothesis that pre-Columbian dogs in the New World share ancestry with modern dogs and that they likely arrived with the first humans to the New World. [7]

Clade C included 12% of the dogs sampled and these were sister to two ancient dogs from the Bonn-Oberkassel cave (14,700 YBP) and the Kartstein cave (12,500 YBP) near Mechernich in Germany, with a common recent ancestor estimated to 16,000–24,000 YBP. Clade D contained sequences from 2 Scandinavian breeds (Jamthund, Norwegian Elkhound) and were sister to another 14,500 YBP wolf sequence also from the Kesserloch cave, with a common recent ancestor estimated to 18,300 YBP. Its branch is phylogenetically rooted in the same sequence as the "Altai dog" (not a direct ancestor). The data from this study indicated a European origin for dogs that was estimated at 18,800–32,100 years ago based on the genetic relationship of 78% of the sampled dogs with ancient canid specimens found in Europe. [21] [7] The data supports the hypothesis that dog domestication preceded the emergence of agriculture [19] and was initiated close to the Last Glacial Maximum when hunter-gatherers preyed on megafauna. [7] [22]

This genetic analysis indicates that C. l. spelaeus possessed mitochondrial lineages which cannot be found among the modern C. lupus, and therefore they are now extinct. It also indicates that the domestic dog C. l. familiaris descended from these extinct lineages. [8]

Related Research Articles

<span class="mw-page-title-main">Canidae</span> Family of mammals

Canidae is a biological family of dog-like carnivorans, colloquially referred to as dogs, and constitutes a clade. A member of this family is also called a canid. The family includes three subfamilies: the Caninae, the extinct Borophaginae and Hesperocyoninae. The Caninae are known as canines, and include domestic dogs, wolves, coyotes, foxes, jackals and other species.

Subspecies of <i>Canis lupus</i>

There are 38 subspecies of Canis lupus listed in the taxonomic authority Mammal Species of the World. These subspecies were named over the past 250 years, and since their naming, a number of them have gone extinct. The nominate subspecies is the Eurasian wolf.

<span class="mw-page-title-main">Wolf</span> Type of canine

The wolf, also known as the gray wolf or grey wolf, is a large canine native to Eurasia and North America. More than thirty subspecies of Canis lupus have been recognized, including the dog and dingo, though gray wolves, as popularly understood, only comprise naturally-occurring wild subspecies. The wolf is the largest extant member of the family Canidae, and is further distinguished from other Canis species by its less pointed ears and muzzle, as well as a shorter torso and a longer tail. The wolf is nonetheless related closely enough to smaller Canis species, such as the coyote and the golden jackal, to produce fertile hybrids with them. The wolf's fur is usually mottled white, brown, gray, and black, although subspecies in the arctic region may be nearly all white.

<span class="mw-page-title-main">Cave bear</span> Extinct species of carnivore

The cave bear is a prehistoric species of bear that lived in Europe and Asia during the Pleistocene and became extinct about 24,000 years ago during the Last Glacial Maximum.

<span class="mw-page-title-main">Dire wolf</span> Extinct species of the genus Aenocyon from North America

The dire wolf is an extinct canine. The dire wolf lived in the Americas during the Late Pleistocene and Early Holocene epochs. The species was named in 1858, four years after the first specimen had been found. Two subspecies are recognized: Aenocyon dirus guildayi and Aenocyon dirus dirus. The largest collection of its fossils has been obtained from the Rancho La Brea Tar Pits in Los Angeles.

<i>Canis</i> Genus of carnivores

Canis is a genus of the Caninae which includes multiple extant species, such as wolves, dogs, coyotes, and golden jackals. Species of this genus are distinguished by their moderate to large size, their massive, well-developed skulls and dentition, long legs, and comparatively short ears and tails.

<i>Panthera spelaea</i> Extinct species of lion

Panthera spelaea, also known as the cave lion or steppe lion, is an extinct Panthera species that most likely evolved in Europe after the third Cromerian interglacial stage, less than 600,000 years ago. Genetic analysis of ancient DNA has revealed that while closely related, it was a distinct species genetically isolated from the modern lion occurring in Africa and Asia, with the genetic divergence between the two species variously estimated between 1.9 million and 600,000 years ago. It is closely related and probably ancestral to the American lion. The species ranged from Western Europe to eastern Beringia in North America, and was a prominent member of the mammoth steppe fauna. It became extinct about 13,000 years ago.

<span class="mw-page-title-main">Hokkaido wolf</span> Extinct subspecies of the gray wolf

The Hokkaido wolf, also known as the Ezo wolf and in Russia as the Sakhalin wolf, is an extinct subspecies of gray wolf that once inhabited coastal northeast Asia. Its nearest relatives were the wolves of North America rather than Asia. It was exterminated in Hokkaido during the Meiji Restoration period, when American-style agricultural reforms incorporated the use of strychnine-laced baits to kill livestock predators. Some taxonomists believe that it survived up until 1945 on the island of Sakhalin. It was one of two subspecies that were once found in the Japanese archipelago, the other being the Japanese wolf.

<span class="mw-page-title-main">Italian wolf</span> Subspecies of carnivore

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<span class="mw-page-title-main">Great Plains wolf</span> Subspecies of gray wolf

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<span class="mw-page-title-main">Domestication of the dog</span> Process which created the domestic dog

The domestication of the dog was the process which led to the domestic dog. This included the dog's genetic divergence from the wolf, its domestication, and the emergence of the first dogs. Genetic studies suggest that all ancient and modern dogs share a common ancestry and descended from an ancient, now-extinct wolf population – or closely related wolf populations – which was distinct from the modern wolf lineage. The dog's similarity to the grey wolf is the result of substantial dog-into-wolf gene flow, with the modern grey wolf being the dog's nearest living relative. An extinct Late Pleistocene wolf may have been the ancestor of the dog.

<span class="mw-page-title-main">Japanese wolf</span> Extinct subspecies of the gray wolf

The Japanese wolf, also known as the Honshū wolf, is an extinct subspecies of the gray wolf that was once endemic to the islands of Honshū, Shikoku and Kyūshū in the Japanese archipelago.

<span class="mw-page-title-main">Himalayan wolf</span> Subspecies of mammal

The Himalayan wolf is a canine of debated taxonomy. It is distinguished by its genetic markers, with mitochondrial DNA indicating that it is genetically basal to the Holarctic grey wolf, genetically the same wolf as the Tibetan and Mongolian wolf, and has an association with the African wolf. No striking morphological differences are seen between the wolves from the Himalayas and those from Tibet. The Himalayan wolf lineage can be found living in Ladakh in the Himalayas, the Tibetan Plateau, and the mountains of Central Asia predominantly above 4,000 m (13,000 ft) in elevation because it has adapted to a low-oxygen environment, compared with other wolves that are found only at lower elevations.

<span class="mw-page-title-main">Armbruster's wolf</span> Extinct species of carnivore

Armbruster's wolf is an extinct species that was endemic to North America and lived during the Irvingtonian stage of the Pleistocene epoch, spanning from 1.9 Mya—250,000 years BP. It is notable because it is proposed as the ancestor of one of the most famous prehistoric carnivores in North America, the dire wolf, which replaced it.

<i>Canis lupus dingo</i> Subspecies of canine

In the taxonomic treatment presented in the third (2005) edition of Mammal Species of the World, Canis lupus dingo is a taxonomic rank that includes both the dingo that is native to Australia and the New Guinea singing dog that is native to the New Guinea Highlands. It also includes some extinct dogs that were once found in coastal Papua New Guinea and the island of Java in the Indonesian Archipelago. In this treatment it is a subspecies of Canis lupus, the wolf, although other treatments consider the dog as a full species, with the dingo and its relatives either as a subspecies of the dog, a species in its own right, or simply as an unnamed variant or genetic clade within the larger population of dogs. The genetic evidence indicates that the dingo clade originated from East Asian domestic dogs and was introduced through the Malay Archipelago into Australia, with a common ancestry between the Australian dingo and the New Guinea singing dog. The New Guinea singing dog is genetically closer to those dingoes that live in southeastern Australia than to those that live in the northwest.

<span class="mw-page-title-main">Beringian wolf</span> Extinct type of wolf that lived during the Ice Age in Alaska, Yukon, and northern British Columbia

The Beringian wolf is an extinct population of wolf that lived during the Ice Age. It inhabited what is now modern-day Alaska, Yukon, and northern British Columbia. Some of these wolves survived well into the Holocene. The Beringian wolf is an ecomorph of the gray wolf and has been comprehensively studied using a range of scientific techniques, yielding new information on their prey species and feeding behaviors. It has been determined that these wolves are morphologically distinct from modern North American wolves and genetically basal to most modern and extinct wolves. The Beringian wolf has not been assigned a subspecies classification and its relationship with the extinct European cave wolf is not clear.

<span class="mw-page-title-main">Pleistocene wolf</span> Extinct lineage of the grey wolf

The Pleistocene wolf, also referred to as the Late Pleistocene wolf, is an extinct lineage or ecomorph of the grey wolf. It was a Late Pleistocene 129 Ka – early Holocene 11 Ka hypercarnivore. While comparable in size to a large modern grey wolf, it possessed a shorter, broader palate with large carnassial teeth relative to its overall skull size, allowing it to prey and scavenge on Pleistocene megafauna. Such an adaptation is an example of phenotypic plasticity. It was once distributed across the northern Holarctic. Phylogenetic evidence indicates that despite being much smaller than this prehistoric wolf, the Japanese wolf, which went extinct in the early 20th century, was of a Pleistocene wolf lineage, thus extending its survival to several millennia after its previous estimated extinction around 7,500 years ago.

<span class="mw-page-title-main">Paleolithic dog</span> Late Pleistocene canine

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

<span class="mw-page-title-main">Evolution of the wolf</span>

The evolution of the wolf occurred over a geologic time scale of at least 300 thousand years. The grey wolf Canis lupus is a highly adaptable species that is able to exist in a range of environments and which possesses a wide distribution across the Holarctic. Studies of modern grey wolves have identified distinct sub-populations that live in close proximity to each other. This variation in sub-populations is closely linked to differences in habitat – precipitation, temperature, vegetation, and prey specialization – which affect cranio-dental plasticity.

<i>Canis mosbachensis</i> Extinct species of carnivore

Canis mosbachensis is an extinct wolf that once inhabited Europe 600,000—420,000 years ago. The Mosbach wolf was a short-legged carcass feeder adapted for scavenging megafauna on the mammoth steppe. The Mosbach wolf is proposed as the ancestor of the grey wolf Canis lupus but some mammalogists have assigned it as the subspecies Canis lupus mosbachensis.

References

  1. Kempe, Stephan; Döppes, Doris (2009). "Cave Bear, Cave Lion and Cave Hyena Skulls from the Public Collection at the Humboldt Museum in Berlin". Acta Carsologica. 38 (2–3). doi: 10.3986/ac.v38i2-3.126 .
  2. 1 2 Goldfuss, G. A. (1823). "5-Ueber den Hölenwolf (Canis spelaeus) (About the Cave wolf)". Osteologische Beiträge zur Kenntniss verschiedener Säugethiere der Vorwelt (Osteological contributions to different knowledge Beast of the ancients). Vol. 3. Nova Acta Physico-Medica Academiea Caesarae Leopoldino-Carolinae Naturae Curiosorum. pp. 451–455.
  3. 1 2 3 Baryshnikov, Gennady F.; Mol, Dick; Tikhonov, Alexei N (2009). "Finding of the Late Pleistocene carnivores in Taimyr Peninsula (Russia, Siberia) with paleoecological context". Russian Journal of Theriology. 8 (2): 107–113. doi:10.15298/rusjtheriol.08.2.04 . Retrieved September 7, 2022.
  4. 1 2 3 Kuzmina, I. E.; Sablin, M. V. (1994). "Wolf Canis lupus L. from the Late Paleolithic sites Kostenki on the Don River". Trudy Zoologicheskogo Instituta (in Russian) (256): 44–58. See the last page for the summary in English
  5. 1 2 3 Boudadi-Maligne, Myriam (2012). "Une nouvelle sous-espèce de loup (Canis lupus maximus nov. Subsp.) dans le Pléistocène supérieur d'Europe occidentale [A new subspecies of wolf (Canis lupus maximus nov. subsp.) from the upper Pleistocene of Western Europe]". Comptes Rendus Palevol. 11 (7): 475. doi:10.1016/j.crpv.2012.04.003.
  6. 1 2 3 4 5 6 7 8 9 Cajus G. Dietrich (2022). "Eurasian Grey and White wolf ancestors—800,000 years evolution, adaptation, pathologies and European dog origins". Acta Zoologica. 105 (1). Wiley: 2–37. doi:10.1111/azo.12451. S2CID   254340729.
  7. 1 2 3 4 5 Thalmann O, Shapiro B, Cui P, Schuenemann VJ, Sawyer SK, Greenfield DL, Germonpre MB, Sablin MV, Lopez-Giraldez F, Domingo-Roura X, Napierala H, Uerpmann HP, Loponte DM, Acosta AA, Giemsch L, Schmitz RW, Worthington B, Buikstra JE, Druzhkova A, Graphodatsky AS, Ovodov ND, Wahlberg N, Freedman AH, Schweizer RM, Koepfli KP, Leonard JA, Meyer M, Krause J, Paabo S, Green RE, Wayne RK (2013). "Complete Mitochondrial Genomes of Ancient Canids Suggest a European Origin of Domestic Dogs". Science. 342 (6160): 871–874. Bibcode:2013Sci...342..871T. doi:10.1126/science.1243650. PMID   24233726. S2CID   1526260.
  8. 1 2 3 Marciszak, Adrian (2023). "History of Polish Canidae (Carnivora, Mammalia) and Their Biochronological Implications on the Eurasian Background". Genes . 14 (3). MDPI: 539. doi: 10.3390/genes14030539 . PMC   10048199 . PMID   36980812.
  9. Marciszak, Adrian (2017). "Large mammals (carnivores, artiodactyls) from Solna Jama Cave (Bystrzyckie Mts, Southwestern Poland) in the context of faunal changes in the postglacial period of Central Europe". Palaeontologia Electronica . 20.2.3A: 1–37. doi: 10.26879/581 .
  10. 1 2 Diedrich 2015, pp. 96–99
  11. Diedrich 2017, pp. 108–118
  12. 1 2 3 4 Diedrich, C. G. (2013). "Extinctions of Late Ice Age Cave Bears as a Result of Climate/Habitat Change and Large Carnivore Lion/Hyena/Wolf Predation Stress in Europe". ISRN Zoology. 2013: 1–25. doi: 10.1155/2013/138319 .
  13. Jánossy, Dénes (2012). "Vertebrate faunas of the Middle Pleistocene of Hungary". Pleistocene Vertebrate Faunas of Hungary. Elsevier Science. p. 102. ISBN   978-0444556356.
  14. Berte, E.; Pandolfi, L. (2014). "Canis lupus (Mammalia, Canidae) from the Late Pleistocene deposit of Avetrana (Taranto, Southern Italy)". Rivista Italiana di Paleontoligia e Stratigrafia. 120 (3): 367–379.
  15. 1 2 Diedrich 2015, pp. 137
  16. Arora, Devender; Singh, Ajeet; Sharma, Vikrant; Bhaduria, Harvendra Singh; Patel, Ram Bahadur (2015). "Hgs Db: Haplogroups Database to understand migration and molecular risk assessment". Bioinformation. 11 (6): 272–275. doi:10.6026/97320630011272. PMC   4512000 . PMID   26229286.
  17. Avise, J. C. (1994). Molecular Markers, Natural History, and Evolution. Chapman & Hall. pp. 109–110. ISBN   978-0-412-03781-8.
  18. Robert K. Wayne; Jennifer A. Leonard; Carles Vila (2006). "Chapter 19:Genetic Analysis of Dog Domestication". In Melinda A. Zeder (ed.). Documenting Domestication:New Genetic and Archaeological Paradigms. University of California Press. pp. 279–295. ISBN   978-0-520-24638-6.
  19. 1 2 Vila, C. (1997). "Multiple and ancient origins of the domestic dog". Science. 276 (5319): 1687–1689. doi:10.1126/science.276.5319.1687. PMID   9180076.
  20. Bjornerfeldt, S (2006). "Relaxation of selective constraint on dog mitochondrial DNA followed domestication". Genome Research. 16 (8): 990–994. doi:10.1101/gr.5117706. PMC   1524871 . PMID   16809672.
  21. Miklosi, Adam (2018). "1-Evolution & Ecology". The Dog: A Natural History. Princeton University Press. pp. 13–39. ISBN   978-0-691-17693-2.
  22. Shipman, P. (2015). The Invaders:How humans and their dogs drove Neanderthals to extinction. Harvard University Press. p. 149. ISBN   9780674736764.

Bibliography