The cave hyena (''Crocuta crocuta spelaea''), also known as the Ice Age spotted hyena, was a

paleosubspecies A chronospecies is a species derived from a sequential development pattern that involves continual and uniform changes from an extinct ancestral form on an evolutionary scale. The sequence of alterations eventually produces a population that is p ...

spotted hyena The spotted hyena (''Crocuta crocuta''), also known as the laughing hyena, is a hyena species, currently classed as the sole extant member of the genus ''Crocuta'', native to sub-Saharan Africa. It is listed as being of least concern by the IUC ...

which ranged from the

Iberian Peninsula The Iberian Peninsula (), ** * Aragonese and Occitan: ''Peninsula Iberica'' ** ** * french: Péninsule Ibérique * mwl, Península Eibérica * eu, Iberiar penintsula also known as Iberia, is a peninsula in southwestern Europe, def ...

to eastern

Siberia Siberia ( ; rus, Сибирь, r=Sibir', p=sʲɪˈbʲirʲ, a=Ru-Сибирь.ogg) is an extensive geographical region, constituting all of North Asia, from the Ural Mountains in the west to the Pacific Ocean in the east. It has been a part ...

. It is one of the best known mammals of the

Ice Age An ice age is a long period of reduction in the temperature of Earth's surface and atmosphere, resulting in the presence or expansion of continental and polar ice sheets and alpine glaciers. Earth's climate alternates between ice ages and gre ...

and is well represented in many European bone caves. The cave hyena was a highly specialised animal, with its progressive and regressive features being more developed than in its modern African relative. It preyed on large mammals (primarily

wild horse The wild horse (''Equus ferus'') is a species of the genus ''Equus'', which includes as subspecies the modern domesticated horse (''Equus ferus caballus'') as well as the endangered Przewalski's horse (''Equus ferus przewalskii''). The Europea ...

steppe bison The steppe bisonSeveral literatures address the species as ''primeval bison''. or steppe wisent (''Bison'' ''priscus'')
– Y ...

and

woolly rhinoceros The woolly rhinoceros (''Coelodonta antiquitatis'') is an extinct species of rhinoceros that was common throughout Europe and Asia during the Pleistocene epoch and survived until the end of the last glacial period. The woolly rhinoceros was a me ...

), and was responsible for the accumulation of hundreds of large Pleistocene mammal bones in areas including horizontal

cave A cave or cavern is a natural void in the ground, specifically a space large enough for a human to enter. Caves often form by the weathering of rock and often extend deep underground. The word ''cave'' can refer to smaller openings such as sea ...

sinkhole A sinkhole is a depression or hole in the ground caused by some form of collapse of the surface layer. The term is sometimes used to refer to doline, enclosed depressions that are locally also known as ''vrtače'' and shakeholes, and to openi ...

s, mud pits and muddy areas along rivers. The cause of the cave hyena's extinction is not fully understood, though it could have been due to a combination of factors, including

climate change In common usage, climate change describes global warming—the ongoing increase in global average temperature—and its effects on Earth's climate system. Climate change in a broader sense also includes previous long-term changes to ...

and competition with other predators.

Description and paleoecology

The main difference between the spotted hyena and the cave hyena lies in the different length of the bones of the hind and front limbs. In the cave hyena, the humerus and the femur are longer, indicating an adaptation to environments other than those of the spotted hyena.The former was also a heavier and more robust animal: an almost complete specimen, found from the Los Aprendices cave in northern Spain, was estimated to weigh 103 kilograms. As in the spotted hyena, the females were larger than the males. A study of 16 fossil specimens of Pleistocene Crocuta indicated that the cave hyena was subject to the Bergmann Rule, becoming larger during glacial periods and smaller during interglacial periods. The same study revealed a progressive increase in carnivorous tooth adaptations during glacial periods, indicating that it was an even more active hunter than today's spotted hyena, a behaviour necessitated by the need to feed on calorie rich fresh meat in a freezing environment. Rock paintings in the

Lascaux Lascaux ( , ; french: Grotte de Lascaux , "Lascaux Cave") is a network of caves near the village of Montignac, in the department of Dordogne in southwestern France. Over 600 parietal wall paintings cover the interior walls and ceilings of t ...

and Chauvet Caves indicate that the cave hyena had the characteristic patches and mane of the spotted hyena. It has been proposed that it possessed thicker fur than the spotted hyena as an environmental adaptation.

Brain

Intracranial digital casts taken from spotted hyenas and two cave hyena skulls showed that the latter had an encephalic volume of 174-218 cm³, higher than today's spotted hyena which has an average volume of 160 cm³. In cave hyenas, however, the anterior telencephalon occupied only 15.9-16.6% of the total brain volume, in contrast to the spotted hyena,whose anterior telencephalon occupied 24.5%. As previous studies show that there is a correlation between

telencephalon The cerebrum, telencephalon or endbrain is the largest part of the brain containing the cerebral cortex (of the two cerebral hemispheres), as well as several subcortical structures, including the hippocampus, basal ganglia, and olfactory bulb. ...

development and feeding sociability and flexibility in hyenas, it has been proposed, in light of this finding, that the cave hyena didn't demonstrate complex social behaviors or adaptability like the spotted hyena, being instead more similar like brown and striped hyena both known as solitary scavengers.

Diet

The most common prey found in burrows found in Europe are invariably horses : in the Srbsko Chlum-Komin Cave alone in the Czech Republic, horse remains make up 51% of the species present.This predilection for equines is shared with today's spotted hyena. Steppe bison remains are generally rare in hyena burrows and it has been proposed that, except during glacial periods, these were avoided to lessen competition with cave lions and wolves. However, certain sites, such as the cave of San Teodoro, where bison make up 50% of the remains, indicate that certain populations of hyenas specialized in their hunting where mammoths and bears were scarce, whose carcasses were a main source of food in much of Europe. Cervids are rare or absent in the burrows, probably being too fast for hyenas.DIEDRICH, C.G. & ŽÁK, K. 2006
Prey deposits and den sites of the Upper Pleistocene hyena Crocuta crocuta spelaea (Goldfuss, 1823) in horizontal and vertical caves of the Bohemian Karst (Czech Republic)
''Bulletin of Geosciences'' 81(4), 237–276 (25 figures). Czech Geological Survey, Prague. ISSN 1214-1119.Diedrich, C. 2010. “Specialized horse killers in Europe – foetal horse remains in the Late Pleistocene Srbsko Chlum-Komín Cave hyena den in the Bohemian Karst (Czech Republic) and actualistic comparisons to modern African spotted hyenas as zebra hunters.” ''Quaternary International'', vol. 220, no. 1-2, pp. 174-187.

History of discovery and classification

Stamps of Moldova 2016 Crocuta crocuta spelaea (mod)

Although the first full account of the cave hyena was given by

Georges Cuvier Jean Léopold Nicolas Frédéric, Baron Cuvier (; 23 August 1769 – 13 May 1832), known as Georges Cuvier, was a French naturalist and zoologist, sometimes referred to as the "founding father of paleontology". Cuvier was a major figure in na ...

in 1812, skeletal fragments of the cave hyena have been described in scientific literature since the 18th century, though they were frequently misidentified. The first recorded mention of the cave hyena in literature occurs in Kundmann's 1737 tome ''Rariora Naturæ et Artis'', where the author misidentified a hyena's mandibular ramus as that of a calf. In 1774, Esper erroneously described hyena teeth discovered in Gailenreuth as those of a lion, and in 1784, Collini described a cave hyena skull as that of a

seal Seal may refer to any of the following: Common uses * Pinniped, a diverse group of semi-aquatic marine mammals, many of which are commonly called seals, particularly: ** Earless seal, or "true seal" ** Fur seal * Seal (emblem), a device to imp ...

. The past presence of hyenas in

Great Britain Great Britain is an island in the North Atlantic Ocean off the northwest coast of continental Europe. With an area of , it is the largest of the British Isles, the largest European island and the ninth-largest island in the world. It ...

was revealed after

William Buckland William Buckland DD, FRS (12 March 1784 – 14 August 1856) was an English theologian who became Dean of Westminster. He was also a geologist and palaeontologist. Buckland wrote the first full account of a fossil dinosaur, which he named ' ...

's examination of the contents of

Kirkdale Cave Kirkdale Cave is a cave and fossil site located in Kirkdale near Kirkbymoorside in the Vale of Pickering, North Yorkshire, England. The cave was discovered by workmen in 1821, and was found to contain fossilized bones of a variety of mammals ...

, which was discovered to have once been the location of several hyena den sites. Buckland's findings were followed by further discoveries by Clift and Whidbey in

Oreston Oreston ( ), formerly a village on the southern bank of the Cattewater, is now a suburb of Plymouth. It is recorded as ''Horestone'' on the 1591 Spry ''Map of Plimmouth''. Oreston offers many small, local services. It is home to a small cornershop ...

Plymouth Plymouth () is a port city status in the United Kingdom, city and unitary authority in South West England. It is located on the south coast of Devon, approximately south-west of Exeter and south-west of London. It is bordered by Cornwall to ...

.Dawkins, William Boyd; Sanford, W. Ayshford; Reynolds, Sydney Hugh. (1866)
''A monograph of the British pleistocene mammalia''
Palaeontographical Society (Great Britain). pp. 1-6. In his own 1812 account, Cuvier mentioned a number of European localities where cave hyena remains were found, and considered it a different species from the spotted hyena on account of its superior size. He elaborated his view in his ''Ossemens Fossiles'' (1823), noting how the cave hyena's digital extremities were shorter and thicker than those of the spotted hyena. His views were largely accepted throughout the first half of the 19th century, finding support in de Blainville and

Richard Owen Sir Richard Owen (20 July 1804 – 18 December 1892) was an English biologist, comparative anatomist and paleontologist. Owen is generally considered to have been an outstanding naturalist with a remarkable gift for interpreting fossils. Ow ...

among others. Further justifications in separating the two animals included differences in the tubercular portion of the lower

carnassial Carnassials are paired upper and lower teeth modified in such a way as to allow enlarged and often self-sharpening edges to pass by each other in a shearing manner. This adaptation is found in carnivorans, where the carnassials are the modified f ...

. Boyd Dawkins, writing in 1865, was the first to definitely cast doubt over the separation of the spotted and cave hyena, stating that the aforementioned tooth characteristics were consistent with mere individual variation. Writing again in 1877, he further stated after comparing the two animals' skulls that there are no characters of specific value. Analyses of the

mitochondrial A mitochondrion (; ) is an organelle found in the cells of most Eukaryotes, such as animals, plants and fungi. Mitochondria have a double membrane structure and use aerobic respiration to generate adenosine triphosphate (ATP), which is used t ...

cytochrome b Cytochrome b within both molecular and cell biology, is a protein found in the mitochondria of eukaryotic cells. It functions as part of the electron transport chain and is the main subunit of transmembrane cytochrome bc1 and b6f complexes. F ...

genes in both modern African and Pleistocene spotted hyenas demonstrated that the two were the same species, and indicate that spotted hyenas migrated from Africa to Eurasia in three waves around 3, 1, and 0.3 million years ago. Analysis of the cave hyena

genome In the fields of molecular biology and genetics, a genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding ...

also indicates that it is not a separate species but is the Eurasian representative of the Pleistocene spotted hyena.

Relationships with hominids

Interactions

Kills partially processed by

Neanderthal Neanderthals (, also ''Homo neanderthalensis'' and erroneously ''Homo sapiens neanderthalensis''), also written as Neandertals, are an Extinction, extinct species or subspecies of archaic humans who lived in Eurasia until about 40,000 years ag ...

s and then by cave hyenas indicate that hyenas would occasionally steal Neanderthal kills; and cave hyenas and Neanderthals competed for cave sites. Many caves show alternating occupations by hyenas and Neanderthals. The presence of large hyena populations in the

Russian Far East The Russian Far East (russian: Дальний Восток России, r=Dal'niy Vostok Rossii, p=ˈdalʲnʲɪj vɐˈstok rɐˈsʲiɪ) is a region in Northeast Asia. It is the easternmost part of Russia and the Asian continent; and is admin ...

may have delayed the human colonisation of North America.Summerill, Lynette
Gnawed Bones tell Tales
, Summer 2003, ''ASU Research'' There is fossil evidence of humans in Middle Pleistocene Europe butchering and presumably consuming hyenas.

In rock art

The cave hyena is depicted in a few examples of

Upper Palaeolithic The Upper Paleolithic (or Upper Palaeolithic) is the third and last subdivision of the Paleolithic or Old Stone Age. Very broadly, it dates to between 50,000 and 12,000 years ago (the beginning of the Holocene), according to some theories coin ...

rock art in

France France (), officially the French Republic ( ), is a country primarily located in Western Europe. It also comprises of Overseas France, overseas regions and territories in the Americas and the Atlantic Ocean, Atlantic, Pacific Ocean, Pac ...

. A painting from the

Chauvet Cave The Chauvet-Pont-d'Arc Cave (french: Grotte Chauvet-Pont d'Arc, ) in the Ardèche department of southeastern France is a cave that contains some of the best-preserved figurative cave paintings in the world, as well as other evidence of Upper Pale ...

depicts a hyena outlined and represented in profile, with two legs, with its head and front part with well distinguishable spotted coloration pattern. Because of the specimen's steeped profile, it is thought that the painting was originally meant to represent a

cave bear The cave bear (''Ursus spelaeus'') 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. Both the word "cave" and the scientific name ...

, but was modified as a hyena. In

, a red and black rock painting of a hyena is present in the part of the cave known as the Diverticule axial, and is depicted in profile, with four limbs, showing an animal with a steep back. The body and the long neck have spots, including the flanks. An image on a cave in Ariège shows an incompletely outlined and deeply engraved figure, representing a part of an elongated neck, smoothly passing into part of the animal's forelimb on the proximal side. Its head is in profile, with a possibly re-engraved muzzle. The ear is typical of the spotted hyena, as it is rounded. An image in the Le Gabillou Cave in

Dordogne Dordogne ( , or ; ; oc, Dordonha ) is a large rural department in Southwestern France, with its prefecture in Périgueux. Located in the Nouvelle-Aquitaine region roughly half-way between the Loire Valley and the Pyrenees, it is named ...

shows a deeply engraved zoomorphic figure with a head in frontal view and an elongated neck with part of the forelimb in profile. It has large round eyes and short, rounded ears which are set far from each other. It has a broad, line-like mouth that evokes a

smile A smile is a facial expression formed primarily by flexing the muscles at the sides of the mouth. Some smiles include a contraction of the muscles at the corner of the eyes, an action known as a Duchenne smile. Among humans, a smile expresses ...

. Though originally thought to represent a composite or zoomorphic hybrid, it is probable it is a spotted hyena based on its broad muzzle and long neck. The relative scarcity of hyena depictions in Paleolithic rock art has been theorised to be due to the animal's lower rank in the

animal worship The term Animal worship (or zoolatry) is an umbrella term designating religious or ritual practices involving animals. This includes the worship of animal deities or animal sacrifice. An animal 'cult' is formed when a species is taken to represe ...

hierarchy; the cave hyena's appearance was likely unappealing to Ice Age hunters, and it was not sought after as prey. Also, it was not a serious rival like the cave lion or bear, and it lacked the impressiveness of the

mammoth A mammoth is any species of the extinct elephantid genus ''Mammuthus'', one of the many genera that make up the order of trunked mammals called proboscideans. The various species of mammoth were commonly equipped with long, curved tusks an ...

woolly rhino The woolly rhinoceros (''Coelodonta antiquitatis'') is an extinct species of rhinoceros that was common throughout Europe and Asia during the Pleistocene epoch and survived until the end of the last glacial period. The woolly rhinoceros was a me ...

.Spassov N.; Stoytchev T. 2004
The presence of cave hyaena (Crocuta crocuta spelaea) in the Upper Palaeolithic rock art of Europe
. ''Historia naturalis bulgarica'', 16: 159-166.

Extinction

The ultimate cause of the cave hyena's extinction is still poorly understood. While

has been forwarded as a possible reason, it is insufficient to explain the animal's complete extinction; although the extremely cold conditions following the

Last Glacial Maximum The Last Glacial Maximum (LGM), also referred to as the Late Glacial Maximum, was the most recent time during the Last Glacial Period that ice sheets were at their greatest extent. Ice sheets covered much of Northern North America, Northern Eu ...

(LGM) diminished favourable hyena habitat in

Northern Europe The northern region of Europe has several definitions. A restrictive definition may describe Northern Europe as being roughly north of the southern coast of the Baltic Sea, which is about 54°N, or may be based on other geographical factors ...

, and separated cave hyena populations from their African kin, there were still habitable localities in

Southern Southern may refer to: Businesses * China Southern Airlines, airline based in Guangzhou, China * Southern Airways, defunct US airline * Southern Air, air cargo transportation company based in Norwalk, Connecticut, US * Southern Airways Express, M ...

and

Central Europe Central Europe is an area of Europe between Western Europe and Eastern Europe, based on a common historical, social and cultural identity. The Thirty Years' War (1618–1648) between Catholicism and Protestantism significantly shaped the a ...

at that time, and the animal survived many other cold periods during the Pleistocene.Varela, S., Lobo, J. M., Rodríguez, J. and Batra, P. (2010)
Were the Late Pleistocene climatic changes responsible for the disappearance of the European spotted hyena populations?
''Quaternary Science Reviews'', 29: 2027-2035. In the

, climate change was ruled out as the sole cause of the cave hyena's extinction, for although the LGM resulted in a mass extinction of several hyena prey species, the

red deer The red deer (''Cervus elaphus'') is one of the largest deer species. A male red deer is called a stag or hart, and a female is called a hind. The red deer inhabits most of Europe, the Caucasus Mountains region, Anatolia, Iran, and parts of ...

and several other herbivore species survived and would still have adequately sustained hyena populations.Varela, S., Lobo, J. M. and Rodríguez, J. (2010)
Are herbivores and spotted hyena extinctions at the end of the Pleistocene related?
''Zona Arqueologica'', 13: 76-91. In

Western Europe Western Europe is the western region of Europe. The region's countries and territories vary depending on context. The concept of "the West" appeared in Europe in juxtaposition to "the East" and originally applied to the ancient Mediterranean ...

at least, the cave hyena's extinction coincided with a decline in

grassland A grassland is an area where the vegetation is dominated by grasses ( Poaceae). However, sedge ( Cyperaceae) and rush ( Juncaceae) can also be found along with variable proportions of legumes, like clover, and other herbs. Grasslands occur na ...

s 12,500 years ago. Europe experienced a massive loss of lowland habitats favoured by cave hyenas, and a corresponding increase in

mixed woodland Temperate broadleaf and mixed forest is a temperate climate terrestrial habitat type defined by the World Wide Fund for Nature, with broadleaf tree ecoregions, and with conifer and broadleaf tree mixed coniferous forest ecoregions. These fo ...

s. Cave hyenas, under these circumstances, would have been outcompeted by

wolves The wolf (''Canis lupus''; : wolves), 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, and gray wolves, as popularly un ...

and

human Humans (''Homo sapiens'') are the most abundant and widespread species of primate, characterized by bipedalism and exceptional cognitive skills due to a large and complex brain. This has enabled the development of advanced tools, cultu ...

s which were as much at home in forests as in open lands, and in highlands as in lowlands. Cave hyena populations began to shrink after roughly 20,000 years ago, completely disappearing from Western Europe between 14 and 11,000 years ago, and earlier in some areas.C. Stiner, Mary (2004
Comparative ecology and taphonomy of spotted hyenas, humans, and wolves in Pleistocene Italy
''Revue de Paléobiologie, Genève''. (December 2004) 23 (2) : 771-785. ISSN 0253-6730

Gallery

File:British Pleistocene Mammalia (1866) Cave Hyena Skull 1.png, Cranium from Wookey Hole, now in Taunton Museum File:British Pleistocene Mammalia (1866) Cave Hyena Skull 2.png, Skull from Wookey Hole, now in Taunton Museum File:British Pleistocene Mammalia (1866) Cave Hyena Manus 1.png, Anterior and posterior views of the right forefoot, from Tor Bryan Caves near Torquay, now kept in British Museum File:British Pleistocene Mammalia (1866) Cave Hyena Pes 1.png, Anterior and posterior views of the right hind foot, from Tor Bryan Caves near Torquay, now kept in British Museum. File:British Pleistocene Mammalia (1866) Cave Hyena Dentition 1.png, Permanent dentition of a Pleistocene cave hyena from Tor Bryan Caves near Torquay, now kept in British Museum. File:British Pleistocene Mammalia (1866) Cave Hyena Dentition 2.png, Permanent dentition, from Tor Bryan Caves near Torquay (now kept in British Museum), Creswell Caves in Derbyshire (now kept in Manchester's Owen College Museum), Kirkdale Cave and Wookey Hole (now kept in Oxford Museum). File:British Pleistocene Mammalia (1866) Cave Hyena Jaws & Cranium.png, Jaws and cranium from Kent's Hole, Torquay (now in British Museum) and Wookey Hole (now in Taunton Museum). File:British Pleistocene Mammalia (1866) Cave Hyena Vertebrae.png, Vertebrae from Wookey Hole (now in Taunton Museum). File:British Pleistocene Mammalia (1866) Cave Hyena Vertebrae 2.png, Vertebrae from Wookey Hole (now in Taunton Museum). File:British Pleistocene Mammalia (1866) Cave Hyena Vertebrae 3.png, Vertebrae from Wookey Hole (now in Taunton Museum). File:British Pleistocene Mammalia (1866) Cave Hyena Pelvis.png, Pelvis from Wookey Hole (now in Taunton Museum). File:British Pleistocene Mammalia (1866) Cave Hyena Scapula.png, Scapula from Creswell Caves, Derbyshire (now in Owens College Museum, Manchester).

References

External links

* Cuvier, Georges
Note sur les ossemens fossiles d’hyènes
''Nouveau Bulletin des Sciences'', Tome 1, 1807-1809 (pp. 149–150). {{Taxonbar, from=Q1139935 Pleistocene mammals of Asia Pleistocene carnivorans Prehistoric hyenas