Woolly Rhinoceros
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Woolly Rhinoceros

Woolly rhinoceros
Temporal range: 3.6-0.01 Ma
Late Pliocene - Late Pleistocene
Coelodonta antiquitatis .jpg
Woolly rhinoceros skeleton on display
Scientific classification edit
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Perissodactyla
Family: Rhinocerotidae
Genus: +Coelodonta
Species:
C. antiquitatis
Binomial name
Coelodonta antiquitatis
(Blumenbach, 1807)
Synonyms

Rhinoceros tichorhinus (Fischer)[1]

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

The woolly rhinoceros was covered with thick and long hair, which allowed it to survive in the extremely cold, harsh mammoth steppe. It also had a massive hump reaching from its shoulder. It fed mainly on herbaceous plants that grew in the steppe.

Mummified carcasses preserved in permafrost and many bone remains of woolly rhinoceros have been found. Images of woolly rhinoceroses are found among cave paintings in Europe and Asia.

Taxonomy

Wooly rhinoceros remains have been known long before the species was described, and were the basis for some mythical creatures. Native peoples of Siberia believed their horns were the claws of giant birds.[2] A rhinoceros skull was found in Klagenfurt, Germany, in 1335, and was believed to be that of a dragon.[3] In 1590, it was used as the basis for the head on a statue of a lindworm.[4] When a whole skeleton was found in 1663 near Quedlinburg, it was studied by scientist Otto von Guericke, who declared it as the remains of a unicorn.[5]Gotthilf Heinrich von Schubert maintained the belief that the horns were the claws of giant birds, and classified the animal under the name Gryphus antiquitatis, meaning "ancient griffin".[6]

One of the earliest scientific descriptions of an ancient 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 two horns in the permafrost.[7] In 1772, Pallas acquired a head and two legs of a rhinoceros from the locals in Irkutsk,[8] and named the species Rhinoceros lenenesis (after the Lena River).[9] In 1799, Johann Friedrich Blumenbach studied rhinoceros bones from the collection of the University of Göttingen, and proposed the scientific name Rhinoceros antiquitatis.[10] The genus name Coelodonta was chosen in 1831 by geologist Heinrich Georg Bronn, from the Greek words (koilía, "cavity") and (odoús "tooth"), because of the depression in the rhino's molar structure,[11][12] giving the scientific name Coelodonta antiquitatis, "ancient hollow tooth".[13]

Evolution

Woolly rhinoceros by Charles R. Knight

The woolly rhinoceros was the most derived of the genus Coelodonta. The closest extinct relative to the woolly rhinoceros is Elasmotherium, which appeared on the evolutionary arena before the genus Coelodonta. These two lines were divided in the first half of the Miocene. A 1.77 million year old Stephanorhinus hemitoechus rhino mummy may also represent the ancestral stock to Coelodonta.[14][15]

The woolly rhinoceros may have descended from the Eurasian C. tologoijensis or the Tibetan C. thibetana.[14] In 2011, a 3.6-million-year-old woolly rhinoceros fossil, the oldest known, was discovered on the cold Tibetan Plateau.[16]

A study of 40,000- to 70,000-year-old DNA samples showed its closest living relative is the Sumatran rhinoceros.[17]

Description

Structure and appearance

An adult woolly rhinoceros typically measured 3 to 3.8 metres (9.8 to 12.5 ft) from head to tail, with an estimated weight of around 1,800-2,700 kg (4,000-6,000 lb)[18] or 2,000 kg (4,400 lb).[19] It grew up to 2 m (6.6 ft) tall at the shoulder,[18] about the same size as the white rhinoceros.[20] A one-month-old calf was about 120 centimetres (47 in) in length and 72 centimetres (28 in) tall at the shoulder.[21] The two horns were made of keratin, with one long horn reaching forward, and a smaller horn between the eyes.[22] It had thick, long fur, small ears, short, thick legs, and a stocky body.

Restoration of a woolly rhinoceros

Compared to other rhinoceroses, the woolly rhinoceros had a longer head and body, and shorter legs. Like modern rhinos, it had three toes on each foot. Its shoulder was raised with a powerful hump, used to support the animal's massive front horn. The hump also contained a fat reserve to aid survival through the desolate winters of the mammoth steppe.[23]

The rhino's long fur coat was reddish-brown and sometimes had a yellowish tint. Their thick undercoat lay under a layer of long, coarse guard hair that was thickest on the withers and neck. Shorter hair covered the limbs, keeping snow from attaching.[23] The body's length ended with a 45 to 50 centimetres (18 to 20 in) tail with a brush of coarse hair at the end.[24] Females had two nipples on the udders.[19]

The woolly rhinoceros had several features reducing the body's surface area and minimized heat loss. Its ears had were no longer than 24 centimetres (9.4 in), while those of rhinos in hot climates are about 30 centimetres (12 in) cm.[25] Their tails were also relatively shorter. Also reducing the loss of body heat was its thick skin, ranged from 5 to 15 millimetres (0.20 to 0.59 in), heaviest on the chest and shoulders.[14][25]

Skull and dentition

Woolly rhinoceros skull

The skull had a length between 70 to 90 centimetres (28 to 35 in). It was longer than those of other rhinoceros, giving the head a deep, downward-facing slanting position, similar to its fossil relatives Stephanorhinus hemiotoechus and Elasmotherium as well as the white rhinoceros,.[26] Strong muscles on its long occipital bone formed its neck hock and held the massive skull. Its massive lower jaw measured up to 60 centimetres (24 in) long and to 10 centimetres (3.9 in) high.[25]

The nasal septum of the woolly rhinoceros was ossified, unlike modern rhinos.[27] This adaptation evolved as a result of the heavy pressure on the horn and face when the rhinoceros grazed underneath the thick snow.[28] It was most common in adult males. Unique to this rhino, the nasal bones were fused to the premaxillae, which is not the case in older Coelodonta types or today's rhinoceroses.[29]

The teeth of the woolly rhinoceros had thickened enamel and an open internal cavity.[25] It had no incisors, which in rhinoceroses are usually only in the milk teeth and rudimentarily formed.[30] It had 3 premolars and 3 molars in both jaws. The molars were high-crowned and had a thick coat of cementum. The second premolar was shaped in an especially gracile manner.[26]

Horns

Both males and females had two horns. The preserved wear indicates that they were used for combat, probably including intraspecific combat, as recorded in cave paintings, as well as for pushing aside snow to uncover vegetation during winter.[18] They may have also been used to attract mates.[28]

The front horn reached at least a meter (3.3 ft) long, up to 1.4 metres (4.6 ft),[14][31] and its weight reached 15 kilograms (33 lb). It faced far forward, more than those of modern rhinos.[14] The back horn was shorter, no more than half a meter (1.64 ft), at about 15 centimetres (5.9 in) long from a 14.6 by 8 centimetres (5.7 by 3.1 in) cm base.[25]

Frozen specimens

Cast of the mummified Starunia specimen, Natural History Museum, London

Many rhinoceros remains have been found preserved in the permafrost region. One early discovery came in 1771 in the Vilyuy River in eastern Siberia. A head, two legs and hide were found and sent to the Kunstkamera in Saint Petersburg.[32] Later in 1877, a Siberian trader recovered a head and one leg from a tributary of the Yana River.[25]

In October 1907, miners in Starunia, Ukraine, found a mammoth carcass buried in an ozokerite pit. A month later, a rhinoceros was found 5 metres (16 ft) underneath.[33] Both were sent to the Dzieduszycki Museum, where a detailed description was published in the museum's monograph.[34] This discovery made a significant contribution to the knowledge of the woolly rhinoceros. Photographs were published in paleontological journals and textbooks, and the first modern paintings of the species were based on the mounted specimen.[35] The rhino is now located in the Lviv National Museum along with the mammoth.[35] Later, in 1929, the Polish Academy of Arts and Sciences sent an expedition to Starunia, finding the mummified remains of three rhinos.[36] One had its entire body preserved aside from horns and fur. The specimen was taken to the Aquarium and Natural History Museum in Kraków. A plaster cast was made soon afterwards, which is now held in the Natural History Museum in London.[37]

Skull and rib fragments of a rhinoceros were found in 1972 in Churapcha, between the Lena and Amga rivers. A whole skeleton was found soon afterwards, with preserved skin, fur, and stomach contents.[25][38] In 1976, schoolchildren on a class trip found a 20,000 year old rhinoceros skeleton on the Aldan River's left bank, uncovering a skull with both horns, a spine, ribs and limb bones.[39]

In 2007, a rhinoceros carcass was found in the lower reaches of the Kolyma. Its skull and left half were preserved, along with skin and hair on its limbs. The upward-facing position of the carcass indicates that the animal probably fell into mud and sank.[40][25] Next year in 2008, a nearly complete skeleton came from the Chukochya River.[41] That same year, locals near the Amga discovered mummified rhinoceros remains, and over the next two years, pelvic bones, caudal vertebrae and ribs were excavated along with forelimbs and hind limbs with toes intact.[9]

In September 2014, a mummified young rhinoceros was discovered by Alexander "Sasha" Banderov and Simeon Ivanov at a tributary of the Semyulyakh River in the Abyysky District in Yakutia, Russia. Its head and horns, fur, and soft tissues were recovered. Some parts had been thawed and eaten, since they were not covered by permafrost. The body was handed over to the Yakutia Academy of Sciences, where it was named "Sasha" after one of its discoverers.[42] Dental analysis shows that the calf was about seven months old at the time of its death. [43] With its well-intact preservation, scientists hope to obtain usable DNA.[42][44]

Distribution

Range of the woolly rhinoceros, including sites of fossils

By the end of the Riss glaciation (about 130 thousand years ago), the woolly rhinoceros lived all across 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 as far north as the New Siberian Islands.[14][45] It had the widest range of all rhinoceros species.[46]

It seemingly did not cross the Bering land bridge during the last ice age (which connected Asia to North America), with its easterly-most occurrence at the Chukotka Peninsula,[14] probably due to the low grass density and suitable habitat in the Yukon, competition with other large herbivores on the frigid land bridge, and vast glaciers creating physical barriers.[47] Even if some arrived in North America, it was probably not very common.[45]

Paleobiology

The woolly rhinoceros had a similar life history to modern rhinos. Studies on milk teeth show that individuals developed similarly to both the white and black rhinoceros.[30] The two teats in the female suggest that she usually gave birth to one, or more rarely, two calves.[48][49] It could reach up to about 40 years of age, similar to today's rhinos.[50] With their massive horns and size, adult woolly rhinos had few predators, but young animals could be attacked by animals such as hyenas and cave lions. A skull was found with trauma indicating an attack from a feline, but the animal survived to adulthood.[50]

Like its modern relatives, bull woolly rhinos were territorial and defended themselves from competitors, particularly during the rutting season. Fossil skulls indicate damage from the front horns of other rhinos.[50] Lower jaws and back ribs show signs of being broken and re-formed, which may have also come from fighting.[51] The relative frequency of combat injuries compared to recent rhinos is attributed to rapidly changing climatic conditions during the last glacial period, with the woolly rhinoceros facing increased stress in competition with other large and medium-sized herbivores.[52]

Diet

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.[25][26] It had a wide upper lip like that of the white rhinoceros, which easily allowed it to plucking vegetation directly from the ground.[18]

Pollen analysis shows that the rhinoceros ate a variety of plants aside from grasses.[53]Coniferswillows and alders were eaten,[47] as were forbs and mosses.[54] Isotope studies on horns show that it had a seasonal diet; different areas of horn growth suggest that the woolly rhinoceros mainly grazed in summer, while it browsed for shrubs and branches in the winter.[55]

A strain vector biomechanical investigation of the skull, mandible and teeth of a well-preserved last cold stage individual recovered from Whitemoor Haye, Staffordshire, revealed musculature and dental characteristics that support a grazing feeding preference. In particular, the enlargement of the temporalis and neck muscles is consistent with that required to resist the large tugging forces generated when taking large mouthfuls of fodder from the ground. The presence of a large diastema supports this theory.[56]

Comparisons with living perissodactyls confirm that the woolly rhinoceros was a hindgut fermentor with a single stomach, consuming cellulose-rich, protein-poor fodder. It had to consume a heavy amount of food to account for the low nutritive content of its diet.[57] Woolly rhinos living in the Arctic during the Last Glacial Maximum consumed approximately equal volumes of forbs, such as Artemisia, and graminoids.[58]

Habitat

Woolly rhinoceros, woolly mammoths, cave lions, and Equus lambei herd in late Pleistocene northern Spain, by Mauricio Antón

The woolly rhinoceros lived mainly in lowlands, plateaus and river valleys, with dry to arid climates,[47] and migrated to higher elevations in favourable climate phases. It avoided mountain ranges, due to heavy snow and steep terrain that the animal could not easily cross.

The rhino's main habitat was the mammoth steppe, a large, open landscape 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 assortment of animals known as the Mammuthus-Coelodonta Faunal Complex.[59] With its wide distribution, the woolly rhinoceros lived in some areas alongside the other rhinoceroses Stephanorhinus[23] and Elasmotherium.[60]

Extinction

Many species of Pleistocene megafauna, like the woolly rhinoceros, became extinct around the same time period. Human hunting is often cited as one cause.[61] Other theories for the cause of the extinctions are climate change associated with the receding Ice age and the hyperdisease hypothesis (q.v. Quaternary extinction event).[62] One of the more widely accepted theories states that, although the woolly rhinoceros was specialized for cold weather, it was capable of surviving in warmer climates. This suggests that climate change was not the only factor contributing to the rhinoceros's extinction. Other cold-adapted species, such as reindeer, muskox and wisent, survived this period of climatic change and many others like it, supporting the 'overkill' hypothesis for the woolly rhino.

Radiocarbon dating indicates that populations survived as recently as 8,000 BC in western Siberia.[63] However, the accuracy of this date is uncertain, as several radiocarbon plateaus exist around this time. The extinction does not coincide with the end of the last ice age, but does coincide with a minor yet severe climatic reversal that lasted for about 1,000–1,250 years, the Younger Dryas (GS1--Greenland Stadial 1), characterized by glacial readvances and severe cooling globally, a brief interlude in the continuing warming subsequent to the termination of the last major ice age (GS2), thought to have been due to a shutdown of the thermohaline circulation in the ocean due to huge influxes of cold fresh water from the preceding sustained glacial melting during the warmer Interstadial (GI1--Greenland Interstadial 1: ca. 16,000-11,450 14C years B.P.).

Relationship with humans

Reconstruction of woolly rhinoceros with human hunter

Hunting

Woolly rhinoceroses shared their habitat with humans, but direct evidence that they interacted is relatively rare. Only 11% of the known sites of prehistoric Siberian tribes have remains or images of the animal.[14] Many rhinoceros remains are found in caves (such as the K?lna Cave in Central Europe), which were not the natural habitat of either rhinos or humans, and large predators such as hyenas may have carried rhinoceros parts there.[64] Sometimes, only individual teeth or bone fragments are uncovered, which were usually came from only one animal.[65] Most rhinoceros remains in Western Europe are found in the same places where human remains or artifacts were found, but this may have occurred naturally.[66][67]

Signs that early humans hunted or scavenged the rhinceros come from markings on the animal's bones. One specimen had injuries caused by human weaponry, with traces of a wound from a sharp object markings from a the shoulder and thigh, and a preserved spear was found near the carcass.[25] A few sites from the early phase of the Last Glacial Period in the late Middle Paleolithic, such as the Gudenus Cave (Austria)[68] and the open air site of Königsaue (Saxony-Anhalt, Germany),[69] have heavily beaten rhinoceros bones lined with slash marks. This action was done partly to extract the nutritious bone marrow.[70]

Both horns and bones of the rhinoceros were used as raw materials for tools and weapons, as were remains from other animals.[71] In what is now Zwole?, Poland, a device was made from a battered woolly rhinoceros pelvis.[72] Half-meter spear throwers, made from a woolly rhinoceros horn about 27 thousand years ago, came from the banks of the Yana River.[73] A 13,300 year-old spear found on Bolshoy Lyakhovsky Island has a tip made of rhinoceros horn, the furthest north a human artifact has ever been found.[74]

The Pinhole Cave Man is a late Paleolithic figure of a man engraved on a rib bone of a woolly rhinoceros, found at Creswell Crags in England.[75]

Ancient art

Chauvet cave art depicting a woolly rhinoceros

Many cave paintings from the Upper Paleolithic depict woolly rhinoceroses. The animal's defining features are prominently drawn, complete with the raised back and hump, contrasting with its low-lying head. Two curved lines represent the ears. The animal's horns are drawn with their long curvature, and in some cases the coat is also indicated. Many paintings show a black band dividing the body.[27][76]

About 20 Paleolithic drawings of woolly rhinos were known before the discovery of the Chauvet Cave in France, yielding about 60 rhinoceros images.[27] They are dated at over 31,000 years old, probably from the Aurignacian,[14]engraved on cave walls or drawn in red or black. One scene depicts two rhinos fighting each other with their horns.[77] Other illustrations are found in the Rouffignac and Lascaux caves. One drawing from Font-de-Gaume shows a noticeably higher head posture, and others were drawn in red pigments in the Kapova Cave in the Ural Mountains.[78] Some images show rhinoceroses struck with spears or arrows, signifying human hunting.[79]

A woolly rhinoceros statuette was found in Dolní V?stonice in Moravia.[79]

See also

Elasmotherium, another Pleistocene Eurasian rhinoceros

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