Cave hyena Temporal range: Middle to Late Pleistocene, | |
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Crocuta crocuta spelaea skeleton from the Muséum de Toulouse. | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Mammalia |
Order: | Carnivora |
Suborder: | Feliformia |
Family: | Hyaenidae |
Genus: | Crocuta |
Species: | |
Subspecies: | †C. c. spelaea |
Trinomial name | |
†Crocuta crocuta spelaea Goldfuss, 1823 |
Cave hyena (Crocuta(crocuta) spelaea and Crocuta(crocuta) ultima) are extinct species or subspecies of hyena known from Eurasia, which ranged from Western Europe to eastern Asia and Siberia during the Pleistocene epoch. It is well represented in many European caves, primarily dating to the Last Glacial Period. It was an apex predator that preyed on large mammals (primarily large ungulates), and was responsible for the accumulation of hundreds of large Pleistocene mammal bones in areas including horizontal caves, sinkholes, mud pits, and muddy areas along rivers.
Often treated as subspecies or populations of the living African spotted hyena (Crocuta crocuta) to which they were closely related and heavily resembled, genetic evidence from the nuclear genome suggests that Eurasian Crocuta populations (including the west Eurasian Crocuta crocuta spelaea and Asian Crocuta crocuta ultima) were highly genetically divergent from African populations (having estimated to have split over 1 million years ago), with evidence suggesting limited interbreeding between Eurasian cave and African spotted hyenas. [1] Some authors have suggested that the two subspecies should be raised to species level as Crocuta spelaea and Crocuta ultima. [2]
Cave hyenas coexisted in Europe alongside both Neanderthals and modern humans. Evidence suggests that they consumed the remains of Neanderthals at least on occasion, with cave hyenas also being recorded in cave art.
The cause of the cave hyena's extinction is not fully understood, though it could have been due to a combination of factors, including human activity, diminished quantities of prey animals, and climate change. [3]
The size of cave hyenas varied depending on environment, with populations inhabiting colder climates having a larger body size than those inhabiting more temperate climates, an example of Bergmann's rule. [4] A 2017 study estimated that on average cave hyenas weighed approximately 88 kilograms (194 lb), around 60% heavier than living spotted hyenas. In comparison to living spotted hyenas, some of the bones of the limbs are more robust (proportionally thicker and shorter), with the ulna being more curved. In the skull, the first and second upper premolars contact each other, while in living spotted hyenas they are separated by a diastema (gap), [5] though collectively the differences between the skeletal anatomy of cave hyenas and living spotted hyenas are "relatively minor". [6] The endocast (brain cavity) also shows differences between from that of the spotted hyena, with the living spotted hyena having a better developed front part of the brain, which may suggest differences in behaviour. [6] Evidence from cave art suggests that cave hyenas had a similar physical appearance to living spotted hyenas. [7]
While "cave hyenas" did use caves, they were not confined to them, [8] being present where caves were absent, and where present only using caves intermittently. [9]
The cave hyena's diet differed little from contemporary African spotted hyenas, [10] with the bones of prey often being cracked open/crushed into order to feed on the interior marrow, as is done by living spotted hyenas. [11]
The diet of the cave hyena is thought to have primarily consisted of large ungulates like wild horse, aurochs, steppe bison, Irish elk/giant deer, wild boar, red deer and reindeer, with larger herbivores like the woolly rhinoceros and woolly mammoth probably being scavenged after death, with their young perhaps sometimes being targeted. [12]
Wild horse are common in Late Pleistocene European cave hyena dens, implying that they were frequent prey, as zebras are for modern African spotted hyenas. [8] [11] In karst cave sites in the Czech Republic which accumulated during the Last Glacial Period, wild horses as well as woolly rhinoceros are the most common remains in cave hyena dens (though this to a significant degree probably reflects the durability of woolly rhinoceros bones making them able to be identified), with other remains including steppe bison, red deer, reindeer, European wild ass, chamois, alpine ibex, and cave bears (who may have been scavenged after dying in the caves). [13] At the Fouvent-Saint-Andoche karst in France, a similar assemblage is found, albeit also including juvenile woolly mammoths, reflecting the similar cold conditions in both regions during the Last Glacial Period. [14] At the Grotta Paglicci den in southern Italy, the assemblage is dominated by European fallow deer, red deer, roe deer and aurochs. [15] At San Teodoro cave in northern Sicily which is a well known hyena den, remains of herbivores likely accumulated in the cave by hyenas include those of aurochs, steppe bison (with aurochs seeming to predominate over bison), European wild ass, wild boar, red deer, and the endemic dwarf elephant Palaeoloxodon mnaidriensis . [16] At Kirkdale Cave in Yorkshire, northern England which dates to the Last Interglacial when Europe had a temperate climate similar to modern times, the assemblage includes juvenile straight-tusked elephant, Irish elk, red deer, European fallow deer, bison, and the narrow-nosed rhinoceros. [17] At the Manot Cave site in Israel, the bone assemblage accumulated by cave hyenas is predominantly Persian fallow deer, as well as to a lesser extent goats, aurochs and equines. [18] At Wezmeh Cave in the Zagros Mountains of western Iran, dating to the Last Glacial Period, remains include mouflon, wild goat, red deer, aurochs, wild boar, gazelles, and the narrow-nosed rhinoceros. [19] Cave hyenas also appear to have extensively engaged in cannibalism, demonstrated as numerous sites. [20]
Cave hyenas likely came into conflict with cave lions (which despite their name, probably only rarely if ever used caves) over carcasses, with remains of cave lions found in European cave deposits possibly being the result of being brought into the cave by cave hyenas. A significant number of these cave lion remains do not bear any evidence of consumption, which may suggest that like living spotted hyenas, they did not generally consume the remains of lions after killing them. [13]
Although the first full account of the cave hyena was given by Georges Cuvier 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. The past presence of hyenas in Great Britain was revealed after William Buckland's examination of the contents of Kirkdale Cave, 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, Plymouth. [21]
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 among others. Further justifications in separating the two animals included differences in the tubercular portion of the lower carnassial. 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. [21]
Analysis of the mitochondrial genomes of Eurasian Crocuta specimens shows no clear separation from African lineages. However, an analysis of full nuclear genomes of both European and East Asian cave hyenas published in 2020 suggests that African and Eurasian Crocuta populations were largely separate, having estimated to have diverged from each other around 2.5 million years ago, closely corresponding to the age of the earliest Crocuta specimens in Eurasia, which are around 2 million years old from China. The nuclear genome results also suggested that the European and East Asian populations (often assigned to the separate subspecies C. crocuta ultima) were strongly genetically divergent from each other, but were more closely related to each other overall than to African Crocuta populations. Analysis of the nuclear genome suggests that there had been interbreeding between these populations for some time after the split, which likely explains the discordance between the nuclear and mitochondrial genome results, with the mitochondrial genomes of African and European Crocuta more closely related to each other than to East Asian Crocuta, suggesting gene flow between the two groups after the split between the East Asian and European populations. [1] Some authors have suggested that the two subspecies should be raised to species level as Crocutaspelaea and Crocuta ultima. [2] A 2024 study of a cave hyena genome from Sicily found that as with the 2020 study, there was strong genetic separation between Eurasian cave and African spotted hyenas, but unlike the 2020 study, there was no robust support for a basal split between East Asian/Siberian and European cave hyenas, with the Sicilian cave hyena found to be the earliest diverging cave hyena lineage, with less interbreeding with African hyenas than other European cave hyenas. [22]
Crocuta first appeared outside of Africa in Asia during the Early Pleistocene around 2 million years ago, [1] before arriving in Europe at the beginning of the Middle Pleistocene around 800,000 years ago. [23] Crocuta was widely distributed across northern Eurasia during the Middle-Late Pleistocene, spanning from the Iberian Peninsula, Britain and Ireland in the West, across southern Siberia, Mongolia and northern China to the Pacific Coast of the Russian Far East. [12] C. c. ultima at times ranged as far southeast as Taiwan, as well as Thailand and Laos in Southeast Asia, [24] [25] while C. c. spelaea ranged into the Middle East, as far south as the Judaean Desert [26] and as far east as the Zagros Mountains of western Iran. [12] The northernmost records are from the banks of the Vilyuy River in Northeast Siberia, with indirect evidence of feeding on woolly rhinoceros carcasses suggesting cave hyenas may have reached the far northeast of Siberia near the Arctic circle. [9]
Kills partially processed by Neanderthals 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. [27] There is fossil evidence of humans in Middle Pleistocene Europe butchering and presumably consuming hyenas. [28] At a number of cave hyena den sites, the remains of Neanderthals have been found showing evidence of having been gnawed on by the hyenas, [29] [30] this may be the result of cave hyenas scavenging Neanderthal burials, though some of these remains may also be the result of cave hyenas attacking and killing Neanderthals. [31]
The cave hyena is depicted in a few examples of Upper Palaeolithic rock art in France. A painting from the Chauvet Cave 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, but was modified as a hyena. In Lascaux, 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 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. 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 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 or woolly rhino. [7]
A 2014 study concluded that the youngest well-dated remains of cave hyenas in Europe date to around 31,000 years ago. [12] A later 2020 study concluded that cave hyenas may have persisted as late as 7,000 years ago in the southern Iberian Peninsula based on radiocarbon dating of likely hyena coprolites found in caves in the region, but suggested that the dates should be considered with caution due to potential issues with contamination. [32] A 2021 study found the youngest specimens in East Asia date to around 20,000 years ago. [33] Potential causal factors for extinction include decreasing temperatures, competition with other carnivores, including humans for food and living space, and decreased prey abundance. [12] Evidence suggests that climate change alone cannot account for the cave hyena's extinction in Europe and that other factors, such as human activity and decreasing prey abundance, are necessary to explain it. [3]
The aurochs is an extinct species of bovine, considered to be the wild ancestor of modern domestic cattle. With a shoulder height of up to 180 cm (71 in) in bulls and 155 cm (61 in) in cows, it was one of the largest herbivores in the Holocene; it had massive elongated and broad horns that reached 80 cm (31 in) in length.
Hyenas or hyaenas are feliform carnivoran mammals belonging to the family Hyaenidae. With just four extant species, it is the fifth-smallest family in the order Carnivora and one of the smallest in the class Mammalia. Despite their low diversity, hyenas are unique and vital components of most African ecosystems.
The woolly rhinoceros is an extinct species of rhinoceros that inhabited northern Eurasia during the Pleistocene epoch. The woolly rhinoceros was a member of the Pleistocene megafauna. The woolly rhinoceros was covered with long, thick hair that allowed it to survive in the extremely cold, harsh mammoth steppe. It had a massive hump reaching from its shoulder and fed mainly on herbaceous plants that grew in the steppe. Mummified carcasses preserved in permafrost and many bone remains of woolly rhinoceroses have been found. Images of woolly rhinoceroses are found among cave paintings in Europe and Asia. The range of the woolly rhinoceros contracted towards Siberia beginning around 17,000 years ago, with the youngest known records being around 14,000 years old in northeast Siberia, coinciding with the Bølling–Allerød warming, which likely disrupted its habitat, with environmental DNA records possibly extending the range of the species around 9,800 years ago. Its closest living relative is the Sumatran rhinoceros.
The spotted hyena, 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 IUCN due to its widespread range and large numbers estimated between 27,000 and 47,000 individuals. The species is, however, experiencing declines outside of protected areas due to habitat loss and poaching. Populations of Crocuta, usually considered a subspecies of Crocuta crocuta, known as cave hyenas, roamed across Eurasia for at least one million years until the end of the Late Pleistocene. The spotted hyena is the largest extant member of the Hyaenidae, and is further physically distinguished from other species by its vaguely bear-like build, rounded ears, less prominent mane, spotted pelt, more dual-purposed dentition, fewer nipples, and pseudo-penis. It is the only placental mammalian species where females have a pseudo-penis and lack an external vaginal opening.
Crocuta is a genus of hyena containing the largest extant member of the family, the spotted hyena (Crocuta crocuta). Several fossil species are also known, with the Pleistocene Eurasian cave hyenas either being regarded as distinct species or subspecies of the spotted hyena.
Panthera spelaea, commonly known as the cave lion, is an extinct Panthera species that was native to Eurasia and northwest North America during the Pleistocene epoch. Genetic analysis of ancient DNA has revealed that while closely related, it was a distinct species genetically isolated from the modern lion, with the genetic divergence between the two species estimated at around 500,000 years ago. The earliest fossils of the P. spelaea lineage in Eurasia date to around 700,000 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, and an important apex predator across its range. It became extinct about 13,000 years ago. It closely resembled living lions with a coat of yellowish-grey fur though unlike extant lions, males appear to have lacked manes.
The mammoth steppe, also known as steppe-tundra, was once the Earth's most extensive biome. During glacial periods in the later Pleistocene it stretched east-to-west, from the Iberian Peninsula in the west of Europe, across Eurasia to North America, through Beringia and northwest Canada; from north-to-south, the steppe reached from the Arctic southward to southern Europe, Central Asia and northern China. The mammoth steppe was cold and dry, and relatively featureless, though climate, topography, and geography varied considerably throughout. Certain areas of the biome—such as coastal areas—had wetter and milder climates than others. Some areas featured rivers which, through erosion, naturally created gorges, gulleys, or small glens. The continual glacial recession and advancement over millennia contributed more to the formation of larger valleys and different geographical features. Overall, however, the steppe is known to be flat and expansive grassland. The vegetation was dominated by palatable, high-productivity grasses, herbs and willow shrubs.
Pachycrocuta is an extinct genus of prehistoric hyenas. The largest and most well-researched species is Pachycrocuta brevirostris, colloquially known as the giant short-faced hyena as it stood about 90–100 cm (35–39 in) at the shoulder and it is estimated to have averaged 110 kg (240 lb) in weight, approaching the size of a lioness, making it the largest known hyena. Pachycrocuta first appeared during the late Miocene. By 800,000 years ago, it became locally extinct in Europe, with it surviving in East Asia until at least 500,000 years ago, and possibly later elsewhere in Asia.
The European wild ass or hydruntine is an extinct equine from the Middle Pleistocene to Late Holocene of Europe and West Asia, and possibly North Africa. It is a member of the subgenus Asinus, and closely related to the living Asiatic wild ass. The specific epithet, hydruntinus, means from Otranto.
The straight-tusked elephant is an extinct species of elephant that inhabited Europe and Western Asia during the Middle and Late Pleistocene. One of the largest known elephant species, mature fully grown bulls on average had a shoulder height of 4 metres (13 ft) and a weight of 13 tonnes (29,000 lb). Straight-tusked elephants likely lived very similarly to modern elephants, with herds of adult females and juveniles and solitary adult males. The species was primarily associated with temperate and Mediterranean woodland and forest habitats, flourishing during interglacial periods, when its range would extend across Europe as far north as Great Britain and eastwards into Russia, while persisting in southern Europe during glacial periods. Skeletons found in association with stone tools and wooden spears suggest they were scavenged and hunted by early humans, including Homo heidelbergensis and their Neanderthal successors.
Panthera fossilis is an extinct species of cat belonging to the genus Panthera, known from remains found in Eurasia spanning the Middle Pleistocene and possibly into the Early Pleistocene.
Wezmeh Cave is an archaeological site near Islamabad Gharb, western Iran, around 470 km (290 mi) southwest of the capital Tehran. The site was discovered in 1999 and excavated in 2001 by a team of Iranian archaeologists under the leadership of Dr. Kamyar Abdi. Wezmeh cave was re-excavated by a team under direction of Fereidoun Biglari in 2019.
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Stephanorhinus is an extinct genus of two-horned rhinoceros native to Eurasia and North Africa that lived during the Late Pliocene to Late Pleistocene. Species of Stephanorhinus were the predominant and often only species of rhinoceros in much of temperate Eurasia, especially Europe, for most of the Pleistocene. The last two species of Stephanorhinus – Merck's rhinoceros and the narrow-nosed rhinoceros – went extinct during the last glacial period.
The narrow-nosed rhinoceros, also known as the steppe rhinoceros is an extinct species of rhinoceros belonging to the genus Stephanorhinus that lived in western Eurasia, including Europe, as well as North Africa during the Pleistocene. It first appeared in Europe around 500,000 years ago during the Middle Pleistocene and survived there until at least 34,000 years Before Present. It was native to temperate and Mediterranean environments, where it fed on low growing plants and to a lesser extent woody plants. Evidence has been found that it was exploited for food by archaic humans, including Neanderthals.
Stephanorhinus kirchbergensis, also known as Merck's rhinoceros is an extinct species of rhinoceros belonging to the genus Stephanorhinus from the Early-Middle to Late Pleistocene of Eurasia. Its range spanned from Western Europe to Eastern Asia. Among the last members of the genus, it co-existed alongside Stephanorhinus hemitoechus in the western part of its range.
Panthera pardus spelaea, also known as the European Ice Age leopard or the cave leopard, is a fossil leopard subspecies which roamed Europe in the Late Pleistocene and possibly the Holocene.
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