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Extinct species of rhinoceros of northern Eurasia

Woolly rhinoceros Temporal range: 3.6–0.01 Ma PreꞒ Ꞓ O South D C P T J Thousand Pg N ↓ Late Pliocene – Late Pleistocene
Woolly rhinoceros skeleton
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Form:	Mammalia
Order:	Perissodactyla
Family:	Rhinocerotidae
Genus:	† Coelodonta
Species:	† C. antiquitatis
Binomial name
† Coelodonta antiquitatis (Blumenbach, 1799)
Synonyms
Rhinoceros lenenesis Pallas Rhinoceros antiquitatis Blumenbach Rhinoceros tichorhinus Fischer. [1]

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 concluding glacial period. 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 accept been institute. Images of woolly rhinoceroses are found among cave paintings in Europe and Asia.

Taxonomy

[edit]

Woolly rhinoceros remains have been known long before the species was described, and were the ground for some mythical creatures. Native peoples of Siberia believed their horns were the claws of giant birds.^[2]
A rhino skull was found in Klagenfurt, Austria, in 1335, and was believed to exist that of a dragon.^[3]
In 1590, it was used as the basis for the head on a statue of a lindworm.^[4]
Gotthilf Heinrich von Schubert maintained the belief that the horns were the claws of behemothic birds, and classified the brute under the name
Gryphus antiquitatis, meaning "griffin of anquity".^[v]

Ane of the earliest scientific descriptions of an aboriginal rhinoceros species was made in 1769, when the naturalist Peter Simon Pallas wrote a report on his expeditions to Siberia where he found a skull and 2 horns in the permafrost.^[6]
In 1772, Pallas acquired a head and two legs of a rhino from the locals in Irkutsk,^[7] and named the species
Rhinoceros lenenesis
(after the Lena River).^[8]
In 1799, Johann Friedrich Blumenbach studied rhinoceros bones from the collection of the University of Göttingen, and proposed the scientific proper noun
Rhinoceros antiquitatis.^[ix]
The geologist Heinrich Georg Bronn moved the species to
Coelodonta
in 1831 considering of its differences in dental germination with members of the
Rhino
genus.^[10]
This name comes from the Greek words κοιλος (koilos, "hollow") and ὀδούς (odoús
"tooth"), from the low in the rhino's molar construction,^[11]
[12]
giving the scientific proper noun
Coelodonta antiquitatis, "hollow-tooth of artifact".^[13]

Evolution

[edit]

The woolly rhinoceros was the most derived of the genus
Coelodonta. The closest extinct relative to the woolly rhino isElasmotherium. These ii lines were divided in the beginning one-half of the Miocene. A one.77 million year erstwhile
Stephanorhinus
rhinoceros mummy may also stand for a sis grouping to
Coelodonta.^{[fourteen]
[fifteen]}

The woolly rhino may have descended from the Eurasian Coelodonta tologoijensis or the Tibetan Coelodonta thibetana''.^[xiv]
In 2011, a 3.half dozen-million-year-one-time woolly rhinoceros fossil, the oldest known, was discovered on the cold Tibetan Plateau.^[16]

A study of xl,000- to seventy,000-twelvemonth-old Dna samples showed its closest living relative is the Sumatran rhino.^[17]

Description

[edit]

Construction and appearance

[edit]

An developed woolly rhinoceros typically measured 3 to 3.viii metres (x to
12+
i⁄2
 ft) from head to tail, with an estimated weight of effectually 2,000–2,700 kg (4,400–6,000 lb)^[18]
or 3,000 kg (6,600 lb).^{[19]
[twenty]}
Information technology grew up to 2 m (vi+
1⁄2
 ft) tall at the shoulder and 4.6 m (15 ft) in length,^[21]
well-nigh the same size equally the white rhinoceros. A one-month-old calf was about 120 cm (three ft xi in) in length and 72 cm (2 ft 4 in) tall at the shoulder.^[22]
The 2 horns were made of keratin, with one long horn reaching forward, and a smaller horn betwixt the eyes.^[23]
Compared to other rhinoceroses, the woolly rhinoceros had a longer head and body, and shorter legs. Its shoulder was raised with a powerful hump, used to back up the animate being's massive front horn. The hump likewise independent a fat reserve to aid survival through the desolate winters of the mammoth steppe.^[24]

Frozen specimens point that the rhinoceros'southward long fur glaze was crimson-dark-brown, with a thick undercoat that lay under a layer of long, coarse guard hair thickest on the withers and cervix. Shorter hair covered the limbs, keeping snowfall from attaching.^[24]
The body's length ended with a 45-to-50-centimetre (xviii to 20 in) tail with a brush of coarse hair at the end.^[25]
Females had two nipples on the udders.^[20]

The woolly rhino had several features reducing the body's surface area and minimized heat loss. Its ears were no longer than 24 cm (ix+
1⁄ii
 in), while those of rhinos in hot climates are about thirty cm (12 in).^[26]
Their tails were also relatively shorter. It too had thick pare, ranging from 5 to fifteen mm (
1⁄4

to

v⁄viii
 in), heaviest on the chest and shoulders.^[14]
[26]

Skull and dentition

[edit]

The skull had a length betwixt seventy to xc cm (xxx to 35 in). It was longer than those of other rhino, giving the head a deep, downwards-facing slanting position, like to its fossil relatives
Stephanorhinus hemiotoechus
and
Elasmotherium
as well every bit the white rhino.^[27]
Stiff muscles on its long occipital os formed its neck hock and held the massive skull. Its massive lower jaw measured up to 60 cm (24 in) long and 10 cm (iv in) high.^[26]

The nasal septum of the woolly rhino was ossified, different modern rhinos. This was nearly mutual in adult males.^[28]
This accommodation probably evolved as a result of the heavy pressure on the horn and face when the rhino grazed underneath the thick snow.^[29]
Unique to this rhinoceros, the nasal bones were fused to the premaxillae, which is not the case in older
Coelodonta
types or today's rhinoceroses.^[30]
This ossification inspired the junior synonym specific name
tichorhinus, from Greek τειχος (teikhos) "wall", ῥις (ῥιν-) (rhis (rhin-)) "nose".

The teeth of the woolly rhinoceros had thickened enamel and an open up internal cavity.^[26]
Like other rhinos, adults did not accept incisors.^[31]
It had 3 premolars and 3 molars in both jaws. The molars were high-crowned and had a thick coat of cementum.^[eleven]

Horns

[edit]

Both sexes had ii horns. The front horn reached at least one metre (3 ft iii in) long, up to ane.4 metres (4 ft vii in),^{[fourteen]
[32]}
and its weight reached 15 kilograms (33 lb). It faced far forwards, more than those of modern rhinos.^[fourteen]
The back horn was shorter.^[26]

Paleobiology

[edit]

The woolly rhinoceros had a similar life history to mod rhinos. Studies on milk teeth show that individuals developed similarly to both the white and blackness rhinoceros.^[31]
The two teats in the female propose that she raised ane calf, or more than rarely two, every ii to iii years.^[33]
[20]
If similar to modern rhinos, calves lived with their mother for around three years before searching for their own private territory, reaching sexual maturity within five years.^[34]
[35]
Woolly rhinoceroses could accomplish around twoscore years of historic period, similar their modern relatives.^[36]

With their massive horns and size, adults had few predators, only young individuals could be attacked by animals such as hyenas and cave lions. A skull was institute with trauma indicating an assail from a feline, but the animal survived to adulthood.^[36]

Woolly rhinos may accept used their horns for combat, probably including intraspecific combat as recorded in cave paintings, as well every bit for moving snow to uncover vegetation during winter.^[14]
They may have as well been used to attract mates.^[28]
Balderdash woolly rhinos were probably territorial like their modern counterparts, defending themselves from competitors, particularly during the rutting flavor. Fossil skulls signal damage from the forepart horns of other rhinos,^[36]  and lower jaws and back ribs show signs of being broken and re-formed, which may have likewise come up from fighting.^[37] The credible frequency of intraspecific combat, compared to recent rhinos, was probable a result of rapid climatic modify during the last glacial period, when the animal faced increased stress from competition with other large herbivores.^[29]

Diet

[edit]

Woolly rhinoceroses mostly fed on grasses and sedges that grew in the mammoth steppe. Its long, slanted head with a downward-facing posture, and tooth structure all helped it graze on vegetation.^[26]
[27] It had a broad upper lip like that of the white rhinoceros, which allowed it to easily pluck vegetation directly from the ground.^[20]
Pollen analysis shows information technology also ate woody plants (including conifers, willows and alders),^[26]
forth with flowers,^[38]
forbs and mosses.^[39]
Isotope studies on horns show that the woolly rhino had a seasonal diet; different areas of horn growth suggest that it mainly grazed in summer, while it browsed for shrubs and branches in the wintertime.^[forty]

A strain vector biomechanical investigation of the skull, mandible and teeth of a well-preserved concluding cold stage private recovered from Whitemoor Haye, Staffordshire, revealed musculature and dental characteristics that support a grazing feeding preference. In particular, the enlargement of the temporalis and cervix muscles is consistent with that required to resist the large tugging forces generated when taking big mouthfuls of provender from the footing. The presence of a large diastema supports this theory.^[41]

Comparisons with living perissodactyls ostend that the woolly rhinoceros was a hindgut fermentor with a single stomach, consuming cellulose-rich, protein-poor fodder. It had to swallow a heavy corporeality of food to account for the low nutritive content of its diet. Woolly rhinos living in the Arctic during the Terminal Glacial Maximum consumed approximately equal volumes of forbs, such every bit
Artemisia, and graminoids.^[42]

Habitat and distribution

[edit]

The woolly rhinoceros lived mainly in lowlands, plateaus and river valleys, with dry to arid climates,^[43]
and migrated to college elevations in favourable climate phases. It avoided mountain ranges, due to heavy snow and steep terrain that the animal could not easily cross.^[43]
The rhino'south chief habitat was the mammoth steppe, a large, open up mural covered with wide ranges of grass and bushes. The woolly rhinoceros lived alongside other large herbivores, such as the woolly mammoth, giant deer, reindeer, saiga antelope and bison – an array of animals known as the
Mammuthus-Coelodonta
Faunal Circuitous.^[44]
With its wide distribution, the woolly rhinoceros lived in some areas alongside the other rhinoceroses
Stephanorhinus
^[24]
and
Elasmotherium.^[45]

By the end of the Riss glaciation about 130,000 years agone, the woolly rhinoceros lived throughout northern Eurasia, spanning most of Europe, the Russian Plain, Siberia, and the Mongolian Plateau, ranging to extremes of 72° to 33°N. Fossils have been found every bit far northward as the New Siberian Islands.^[14]
[46]
It had the widest range of whatever rhino species.^[47]

It seemingly did not cross the Bering land span during the last water ice age (which connected Asia to Northward America), with its easterly-most occurrence at the Chukotka Peninsula,^[14]
probably due to the depression grass density and lack of suitable habitat in the Yukon, competition with other large herbivores on the frigid country bridge, and vast glaciers creating physical barriers.^[43]
Even if some arrived in Due north America, this was probably uncommon.^[46]

Human relationship with humans

[edit]

Hunting

[edit]

Woolly rhinoceroses shared their habitat with humans, but direct prove that they interacted is relatively rare. Only xi% of the known sites of prehistoric Siberian tribes have remains or images of the beast.^[14]
Many rhinoceros remains are found in caves (such as the Kůlna Cave in Central Europe), which were non the natural habitat of either rhinos or humans, and large predators such as hyenas may have carried rhinoceros parts there.^[48]
Sometimes, only individual teeth or bone fragments are uncovered, which normally came from but 1 animal.^[49]
Nigh rhinoceros remains in Western Europe are establish in the same places where homo remains or artifacts were found, but this may have occurred naturally.^[l]
[51]

Signs that early humans hunted or scavenged the rhinoceros come from markings on the animal'southward bones. One specimen had injuries caused by human weaponry, with traces of a wound from a abrupt object mark the shoulder and thigh, and a preserved spear was found near the carcass.^[26]
A few sites from the early phase of the Last Glacial Period in the belatedly Heart Paleolithic, such every bit the Gudenus Cave (Austria)^[52]
and the open air site of Königsaue (Saxony-Anhalt, Germany),^[53]
have heavily beaten rhinoceros bones lined with slash marks. This action was done partly to excerpt the nutritious bone marrow.^[54]

Both horns and bones of the rhinoceros were used as raw materials for tools and weapons, equally were remains from other animals.^[55]
In what is now Zwoleń, Poland, a device was made from a battered woolly rhino pelvis.^[56]
One-half-meter spear throwers, made from a woolly rhinoceros horn about 27,000 years ago, came from the Yana Rhinoceros Horn Site on the banks of the Yana River.^[57]
A 13,300 year-erstwhile spear found on Bolshoy Lyakhovsky Island has a tip made of rhinoceros horn, the furthest north a human artifact has ever been constitute.^[58]

The Pinhole Cave Man is a tardily Paleolithic figure of a man engraved on a rib bone of a woolly rhino, found at Creswell Crags in England.^[59]

Ancient art

[edit]

Many cave paintings from the Upper Paleolithic depict woolly rhinoceroses. The animate being's defining features are prominently fatigued, complete with the raised back and hump, contrasting with its depression-lying caput. Two curved lines represent the ears. The beast's horns are drawn with their long curvature, and in some cases the coat is as well indicated. Many paintings bear witness a blackness band dividing the body.^[lx]

Virtually 20 Paleolithic drawings of woolly rhinos were known before the discovery of the Chauvet Cave in France.^[sixty]
They are dated at over 31,000 years quondam, probably from the Aurignacian,^[14]
engraved on cave walls or fatigued in red or black. One scene depicts 2 rhinos fighting each other with their horns.^[61]
Other illustrations are establish in the Rouffignac and Lascaux caves. One drawing from Font-de-Gaume shows a noticeably higher caput posture, and others were drawn in red pigments in the Kapova Cave in the Ural Mountains.^[62]
Some images show rhinoceroses struck with spears or arrows, signifying human hunting.^[63]

The site of Dolní Věstonice in Moravia, Czech Republic, was plant with more than than seven hundred statuettes of animals, many of woolly rhinoceroses.^[63]

Extinction

[edit]

Many species of Pleistocene megafauna, like the woolly rhinoceros, became extinct effectually the same time period. Human hunting is often cited every bit one cause. Other theories for the cause of the extinctions are climate modify associated with the receding Ice age and the hyperdisease hypothesis (q.v. Quaternary extinction event).^[64]
One of the more widely accustomed theories states that, although the woolly rhinoceros was specialized for cold weather, information technology was capable of surviving in warmer climates. This suggests that climate change was not the only factor contributing to the rhinoceros'south extinction. Other cold-adapted species, such as reindeer, muskox and wisent, survived this menstruum of climatic change and many others like it, supporting the 'overkill' hypothesis for the woolly rhino. However, more contempo studies indicate that the woolly rhinoceros may have been specialized to the mammoth steppe, and the paludification and dramatic shrinkage of this habitat due to increased preciptation post-obit the Last Glacial Maximum may have caused its range to slowly contract to whatever refugia were left for the steppe, and ultimately become extinct.^[65]

Radiocarbon dating indicates that populations survived every bit recently as nigh 10,000 BC in western Siberia.^[66]
However, the accurateness of this engagement is uncertain, as several radiocarbon plateaus be around this time. A Holocene survival of the species in supported by eDNA studies showing that the woolly rhinoceros persisted in northeastern Siberia until 9.8 ± 0.2 ka.^[65]

Fossil specimens

[edit]

Frozen specimens

[edit]

Many rhinoceros remains have been plant preserved in the permafrost region. In 1771, a head, two legs and hide were establish in the Vilyuy River in eastern Siberia and sent to the Kunstkamera in Saint Petersburg.^[67]
Afterwards in 1877, a Siberian trader recovered a head and one leg from a tributary of the Yana River.^[26]

In October 1907, miners in Starunia, Russian Empire, found a mammoth carcass cached in an ozokerite pit. A month later, a rhino was constitute v metres (15 ft) underneath.^[68]
Both were sent to the Dzieduszycki Museum, where a detailed clarification was published in the museum'southward monograph.^[69]
Photographs were published in paleontological journals and textbooks, and the get-go modern paintings of the species were based on the mounted specimen.^[70]
The rhino is now located in the Lviv National Museum along with the mammoth.^[70]
Later, in 1929, the Polish University of Arts and Sciences sent an expedition to Starunia, finding the mummified remains of 3 rhinos.^[71]
One specimen, missing simply its horns and fur, was taken to the Aquarium and Natural History Museum in Kraków. A plaster bandage was made soon afterwards, which is now held in the Natural History Museum in London.^[72]

Skull and rib fragments of a rhinoceros were found in 1972 in Churapcha, between the Lena and Amga rivers. A whole skeleton was plant soon afterwards, with preserved skin, fur, and stomach contents.^[26]
[73]
In 1976, schoolchildren on a course trip found a xx,000 year old rhinoceros skeleton on the Aldan River's left depository financial institution, uncovering a skull with both horns, a spine, ribs and limb bones.^[74]

In 2007, a fractional rhinoceros carcass was institute in the lower reaches of the Kolyma river. Its upward-facing position indicates that the animate being probably fell into mud and sank.^[75]
[26]
Next yr in 2008, a nearly complete skeleton came from the Chukochya River.^[76]
That same twelvemonth, locals near the Amga discovered mummified rhinoceros remains, and over the side by side two years, pelvic bones, tail vertebrae and ribs were excavated along with forelimbs and hind limbs with toes intact.^[eight]

In September 2014, a mummified young rhinoceros was discovered by two hunters, Alexander “Sasha” Banderov and Simeon Ivanov, at a tributary of the Semyulyakh River in the Abyysky District in Yakutia, Russian federation. Its caput and horns, fur, and soft tissues were recovered. Some parts had been thawed and eaten, since they were not covered past permafrost. The torso was handed over to the Yakutia Academy of Sciences, where information technology was named “Sasha” after ane of its discoverers.^[77]
Dental assay shows that the calf was well-nigh seven months sometime at the fourth dimension of its death.^[78]
With its well-intact preservation, scientists proceeded to undergo Dna analysis.^[79]
[80]

In August 2020, a rhinoceros was found, afterwards being revealed by melting permafrost, close to site of the 2014 discovery. The rhinoceros was between three and four years former and it is idea that the cause of death was drowning. Information technology is ane of the best preserved animals recovered from the region, having most of its internal organs intact. The discovery was also notable for the preservation of a small nasal horn, a rarity as these unremarkably decompose quickly.^[81]

See likewise

[edit]

• Elasmotherium, another Pleistocene Eurasian rhinoceros

References

[edit]

• Parker, Steve.
  Dinosaurus: The Complete Guide to Dinosaurs. Firefly Books Inc, 2003. Pg. 422.

External links

[edit]

• More pictures of the fully preserved tar pit wholly rhinoceros that was found in Poland (text in Polish)
• Fossil skull of a woolly rhinoceros from Belgium
• Fossil skull of a woolly rhinoceros from Germany
• International Rhino Foundation: Woolly Rhino