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:
C. lupus
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] Genetic evidence suggests that Late Pleistocene European wolves shared high genetic connectivity with contemporary wolves from Siberia, with continual gene flow from Siberian wolves into European wolves over the course of the Late Pleistocene. Modern European wolves derive most of their ancestry from Siberian wolf populations that expanded into Europe during and after the Last Glacial Maximum, but retain a minor fraction of their ancestry from earlier Late Pleistocene European wolves. [7]

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. [8] 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. [9] 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. [10]

These large wolves have not been studied in any degree of depth, and their relationship to modern wolves has not been clarified using DNA. [11]

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. [12] 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. [13]

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, [9] 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. [14]

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. [15]

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. [11]

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. [11]

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. [14] 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 . [11]

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. [19] [20] [21]

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. [19]

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. [22] [19] The data supports the hypothesis that dog domestication preceded the emergence of agriculture [20] and was initiated close to the Last Glacial Maximum when hunter-gatherers preyed on megafauna. [19] [23]

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. [15]

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, and 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">Beringia</span> Geographic region of Asia and North America currently partly submerged

Beringia is defined today as the land and maritime area bounded on the west by the Lena River in Russia; on the east by the Mackenzie River in Canada; on the north by 72° north latitude in the Chukchi Sea; and on the south by the tip of the Kamchatka Peninsula. It includes the Chukchi Sea, the Bering Sea, the Bering Strait, the Chukchi and Kamchatka Peninsulas in Russia as well as Alaska in the United States and the Yukon in Canada.

<span class="mw-page-title-main">Dire wolf</span> Extinct species of canine mammal

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.

<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

The Italian wolf, also known as the Apennine wolf, is a subspecies of the grey wolf native to the Italian Peninsula. It inhabits the Apennine Mountains and the Western Alps, though it is undergoing expansion towards the north and east. As of 2022 the wolf population within Italy is estimated to be 3,307 individuals. Although not universally recognised as a distinct subspecies, it nonetheless possesses a unique mtDNA haplotype and a distinct skull morphology.

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

The Great Plains wolf, also known as the buffalo wolf or loafer, is a subspecies of gray wolf that once extended throughout the Great Plains, from southern Manitoba and Saskatchewan in Canada southward to northern Texas in the United States. The subspecies was thought to be extinct in 1926, until studies declared that its descendants were found in Minnesota, Wisconsin and Michigan. They were described as a large, light-colored wolf but with black and white varying between individual wolves, with some all white or all black. The Native Americans of North Dakota told of how only three Great Plains wolves could bring down any sized bison.

<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>Xenocyon</i> Extinct subgenus of carnivores

Xenocyon is an extinct group of canids, either considered a distinct genus or a subgenus of Canis. The group includes Canis (Xenocyon) africanus, Canis (Xenocyon) antonii and Canis (Xenocyon) falconeri that gave rise to Canis (Xenocyon) lycanoides. The hypercarnivorous Xenocyon is thought to be closely related and possibly ancestral to modern dhole and the African wild dog, as well as the insular Sardinian dhole.

<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

During the Pleistocene, wolves were widely distributed across the Northern Hemisphere. Some Pleistocene wolves, such as Beringian wolves and those from Japan, exhibited large body size in comparison to modern gray wolf populations. Genetic analysis of the remains of Late Pleistocene wolves suggest that across their range populations of wolves maintained considerable gene flow between each other and thus there was limited genetic divergence between them. Modern wolves mostly draw their ancestry from some Siberian populations of Late Pleistocene gray wolves, which largely replaced other gray wolf populations after the Last Glacial Maximum.

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

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.

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

It is widely agreed that the evolutionary lineage of the grey wolf can be traced back 2 million years to the Early Pleistocene species Canis etruscus, and its successor the Middle Pleistocene Canis mosbachensis. 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 inhabited Europe from the late Early Pleistocene to the Middle Pleistocene, around 1.4 million to 400,000 years ago. Canis mosbachensis is widely considered to have descended from the earlier Canis etruscus, and to be the ancestor of the living grey wolf with some considering it as a subspecies of the wolf as Canis lupus mosbachensis. The morphological distinction between C. mosbachensis and C. lupus has historically been vague, and attribution of fossils to C. mosbachensis or to C. lupus around the transition time between the two species is ambiguous.

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 .
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  14. 1 2 Diedrich 2015, pp. 137
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Bibliography

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