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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 |
The cave hyena (Crocuta crocuta spelaea and Crocuta crocuta ultima), also known as the Ice Age spotted hyena, are paleosubspecies of spotted hyena known from Eurasia, which ranged from the Iberian Peninsula to eastern Siberia. It is one of the best known mammals of the Ice Age and is well represented in many European bone caves. It preyed on large mammals (primarily wild horses, steppe bison and woolly rhinoceros), 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.
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), though the lack of clear separation between mitochondrial genome lineages suggests that the two populations interbred for some time after the initial split. [1] Some authors have suggested that the two subspecies should be raised to species level as Crocutaspelaea and Crocuta ultima. [2]
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 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 kg (227 lb). 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 Bergmann's 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 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.
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 development and feeding sociability and flexibility in hyenas, it has been proposed, in light of this finding, brains were more similar to brown and striped hyena both known as solitary scavengers compared to spotted hyenas. However, the authors still stated that cave hyenas were likely capable of forming social groups, just in smaller numbers due to the reduced size of the anterior brain.
The cave hyena's diet differed little from contemporary African spotted hyenas. [4] The most common prey found in Europe are invariably horses, and in the Srbsko Chlum-Komin Cave alone (in the Czech Republic), horse remains make up 51% of the species present. This predilection for equines distinguishes the cave hyena from today’s spotted hyenas, which are known to target smaller antelope (impala, gazelle, wildebeest) as well as opportunistically scavenge carrion. 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 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. [5] [6] However, exceptions to this pattern also exist; the site of Fouvent-Saint-Andoche represents a hyena den containing remains of red deer, Irish elk, and reindeer. [7] Cave hyenas extensively engaged in cannibalism. [8]
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. [9]
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. [9]
Analysis of the mitochondrial genomes of Eurasian Crocuta specimens shows no clear separation from African lineages. However, analysis of full nuclear genomes 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 Asian populations (the latter often assigned to the separate subspecies C. crocuta ultima) were distinct 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 Asian Crocuta, suggesting gene flow between the two groups after the split between the Asian and European populations. [1]
Crocuta was formerly widely distributed across northern Eurasia during the Middle-Late Pleistocene, with C. c. ultima at times ranging as far southeast as Thailand and Laos in Southeast Asia. [10] [11]
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. [12] The presence of large hyena populations in the Russian Far East may have delayed the human colonisation of North America. [13] There is fossil evidence of humans in Middle Pleistocene Europe butchering and presumably consuming hyenas. [14]
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. [15]
The youngest known specimens of cave hyenas in Europe date to around 31,000 years ago, [16] while the youngest specimens in East Asia date to around 20,000 years ago. [17] Potential causal factors for extinction include decreasing temperatures, competition with other carnivores, including humans for food and living space, and decreased prey abundance. [16] 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]
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.
Wookey Hole Caves are a series of limestone caverns, a show cave and tourist attraction in the village of Wookey Hole on the southern edge of the Mendip Hills near Wells in Somerset, England. The River Axe flows through the cave. It is a Site of Special Scientific Interest (SSSI) for both biological and geological reasons. Wookey Hole cave is a "solutional cave", one that is formed by a process of weathering in which the natural acid in groundwater dissolves the rocks. Some water originates as rain that flows into streams on impervious rocks on the plateau before sinking at the limestone boundary into cave systems such as Swildon's Hole, Eastwater Cavern and St Cuthbert's Swallet; the rest is rain that percolates directly through the limestone. The temperature in the caves is a constant 11 °C (52 °F).
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.
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 known as well.
Panthera spelaea, commonly known as the cave lion is an extinct Panthera species 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 living lions males appear to have lacked manes.
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 Divje Babe flute, also called tidldibab, is a cave bear femur pierced by spaced holes that was unearthed in 1995 during systematic archaeological excavations led by the Institute of Archaeology of the Research Centre of the Slovenian Academy of Sciences and Arts, at the Divje Babe I near Cerkno in northwestern Slovenia. It has been suggested that it was made by Neanderthals as a form of musical instrument, and became known as the Neanderthal flute. The artifact is on prominent public display in the National Museum of Slovenia in Ljubljana as a Neanderthal flute. As such, it would be the world's oldest known musical instrument.
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 bulls on average had a shoulder height of 3.81–4.2 metres (12.5–13.8 ft) and a weight of 10.8–15 tonnes (24,000–33,000 lb). Like modern elephants, the straight-tusked elephant lived in herds, flourishing during interglacial periods, when its range would extend as far north as Great Britain. Skeletons found in association with stone tools and wooden spears suggest they were scavenged and hunted by early humans, including Neanderthals. It is the ancestral species of most dwarf elephants that inhabited islands in the Mediterranean.
Peștera cu Oase is a system of 12 karstic galleries and chambers located near the city Anina, in the Caraș-Severin county, southwestern Romania, where some of the oldest European early modern human (EEMH) remains, between 42,000 and 37,000 years old, have been found.
The 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.
Denisova Cave is a cave in the Bashelaksky Range of the Altai Mountains, Siberian Federal District, Russia. The cave has provided items of great paleoarchaeological and paleontological interest. Bone fragments of the Denisova hominin originate from the cave, including artifacts dated to around 40,000 BP. Remains of a 32,000-year-old prehistoric species of horse have also been found in the cave.
Neanderthals are an extinct group of archaic humans who lived in Eurasia until about 40,000 years ago. The type specimen, Neanderthal 1, was found in 1856 in the Neander Valley in present-day Germany.
The Neanderthals in Gibraltar were among the first to be discovered by modern scientists and have been among the most well studied of their species according to a number of extinction studies which emphasize regional differences, usually claiming the Iberian Peninsula partially acted as a “refuge” for the shrinking Neanderthal populations and the Gibraltar population of Neanderthals as having been one of many dwindling populations of archaic human populations, existing just until around 42,000 years ago. Many other Neanderthal populations went extinct around the same time.
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.
Ursus ingressus is an extinct species of the family Ursidae that lived in Central Europe during the Late Pleistocene. It is named after the Gamssulzen Cave in Austria, where the holotype of this species was found.
Pešturina is a cave in the municipality of Niška Banja in southeast Serbia. It is located southwest of Jelašnica and 20 km (12 mi) southeast of Niš. Artifacts from the Middle and Upper Paleolithic periods were discovered since the archaeological excavations began in 2006. The remains, identified as the Mousterian culture, were dated from 111,000 BP+ 5,000 to 39,000 BP + 3,000, which makes Pešturina one of the latest surviving Neanderthal habitats. The cave has been nicknamed the "Serbian Atapuerca".
The spotted hyena is the most carnivorous member of the Hyaenidae. Unlike its brown and striped cousins, the spotted hyena is primarily a predator rather than a scavenger. One of the earliest studies to demonstrate its hunting abilities was done by Hans Kruuk, a Dutch wildlife ecologist who showed through a 7-year study of hyena populations in Ngorongoro and Serengeti National Park during the 1960s that spotted hyenas hunt as much as lions, and with later studies this has been shown to be the average in all areas of Africa. However spotted hyenas remain mislabeled as scavengers, often even by ecologists and wildlife documentary channels.
The Cave of Pego do Diabo is a small karst cave in a Turonian limestone outcrop located about 250 metres above sea level in the municipality of Loures, about 20 km north of Lisbon in Portugal. The erosion of the outcrop resulted in a set of openings in the rocks wide enough to be exploited by humans. Several archaeological studies have been conducted within the cave, which give insights into the duration of Neanderthal presence on the Iberian Peninsula and to the presence of carnivorous animals during the Paleolithic.