Woolly rhinoceros

Source: Wikipedia, the free encyclopedia.

Woolly rhinoceros
Temporal range: Middle Pleistocene–Late Pleistocene
Woolly rhinoceros skeleton
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Perissodactyla
Family: Rhinocerotidae
Genus: Coelodonta
Species:
C. antiquitatis
Binomial name
Coelodonta antiquitatis
(Blumenbach, 1799)
Subspecies[1]
  • C. a. praecursor Guérin, 1980
  • C. a. antiquitatis Guérin, 1980
Synonyms

Rhinoceros lenenesis Pallas
Rhinoceros antiquitatis Blumenbach
Rhinoceros tichorhinus Fischer.[2]

The woolly rhinoceros (Coelodonta antiquitatis), simply known as woolly rhino, is an extinct species of

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 species range 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.

Taxonomy

Molar tooth showing the cavity the genus was named for

Woolly 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.[3] A rhinoceros skull was found in Klagenfurt, Austria, in 1335, and was believed to be that of a dragon.[4] In 1590, it was used as the basis for the head on a statue of a lindworm.[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 "griffin of antiquity".[6]

One of the earliest scientific descriptions of an ancient rhinoceros species was made in 1769, when the naturalist

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 geologist Heinrich Georg Bronn moved the species to Coelodonta in 1831 because of its differences in dental formation with members of the Rhinoceros genus.[11] This name comes from the Greek words κοιλος (koilos, "hollow") and ὀδούς (odoús "tooth"), from the depression in the rhino's molar structure,[12][13] giving the scientific name Coelodonta antiquitatis, "hollow-tooth of antiquity".[14]

Evolution

The woolly rhinoceros was the most recent species of the genus Coelodonta. The closest living relative of Coelodonta is the Sumatran rhinoceros, and the genus is also closely related to the extinct genus Stephanorhinus. A cladogram showing the relationships of C. antiquitatis to other Late Pleistocene-recent rhinoceros species based on genomic data is given below.[15]

Elasmotheriinae

Elasmotherium

Rhinocerotinae

Black rhinoceros (Diceros bicornis)

White rhinoceros (Ceratotherium simum)

Indian rhinoceros (Rhinoceros unicornis)

Javan rhinoceros (Rhinoceros sondaicus)

Sumatran rhinoceros (Dicerorhinus sumatrensis)

Woolly rhinoceros (Coelodonta antiquitatis)

Merck's rhinoceros
(Stephanorhinus kirchbergensis)

Relationships of the woolly rhinoceros based on morphology, excluding African rhinoceros species:[16]

Hyrachyus eximius

Tapirus terrestris
(South American tapir)

Rhinocerotidae

Trigonias osborni

Ronzotherium filholi

Aceratheriini
Teleoceratina
Rhinocerotina
Nesorhinus

Nesorhinus hayasakai

Nesorhinus philippinensis

Rhinoceros

Rhinoceros sondaicus
(Javan rhinoceros)

Rhinoceros unicornis
(Indian rhinoceros)

Dicerorhinus

Dicerorhinus gwebinensis

Dicerorhinus fusuiensis

Dicerorhinus sumatrensis
(Sumatran rhinoceros)

Dihoplus schleiermacheri

"Dihoplus" pikermiensis

Coelodonta

Coelodonta thibetana

Coelodonta nihowanensis

Coelodonta tologoijensis

Coelodonta antiquitatis antiquitatis (woolly rhinoceros)

Coelodonta antiquitatis praecursor (woolly rhinoceros)

Pliorhinus

Pliorhinus megarhinus

Pliorhinus miguelcrusafonti

Stephanorhinus

Stephanorhinus jeanvireti

Stephanorhinus etruscus

Stephanorhinus hundsheimensis

Stephanorhinus hemitoechus
(Narrow-nosed or steppe rhinoceros)

Stephanorhinus kirchbergensis (Merck's or forest rhinoceros)

The oldest known species of Coelodonta,

Middle Pleistocene in China, and the oldest remains of the species in Europe, which represents the only species of Coelodonta to have been present in the region, date to approximately 450,000 years ago.[18][19] The woolly rhinoceros is divided into two chrono-subspecies, with C. a. praecursor from the middle Pleistocene and C. a. antiquitatis from the late Pleistocene.[1] Mitochondrial genomes suggest that the last mitochondrial ancestor of Late Pleistocene woolly rhinoceroses lived around 570,000 years ago.[20]

Description

Structure and appearance

Restoration

An adult woolly rhinoceros typically measured 3.2 to 3.6 metres (10.5 to 11.8 ft) from head to tail, stood 1.45–1.6 metres (4.8–5.2 ft) tall at the shoulder, and weighed up to 1.5–2 metric tons (1.7–2.2 short tons).[21][22] Both males and females had two horns which were made of keratin, with one long horn reaching forward and a smaller horn between the eyes.[23][24] The front horn would have measured 1–1.35 metres (3.3–4.4 ft) long for individuals at 25 to 35 years of age, while the second horn would have measured up to 47.5 centimetres (1.56 ft) long.[24] 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 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.[25]

Frozen specimens indicate that the rhino's long fur coat was reddish-brown, with a thick undercoat that lay under a layer of long, coarse guard hair thickest on the withers and neck. Shorter hair covered the limbs, keeping snow from attaching.[25] The body's length ended with a 45-to-50-centimetre (18 to 20 in) tail with a brush of coarse hair at the end.[26] Females had two nipples on the udders.[22]

The woolly rhinoceros had several features which reduced the body's surface area and minimized heat loss. Its ears were no longer than 24 cm (9+12 in), while those of rhinos in hot climates are about 30 cm (12 in).[24] Their tails were also relatively shorter. It also had thick skin, ranging from 5 to 15 mm (14 to 58 in), heaviest on the chest and shoulders.[27][24]

Skull and dentition

Woolly rhinoceros skull

The skull had a length between 70 and 90 cm (30 and 35 in). It was longer than those of other rhinoceros, giving the head a deep, downward-facing slanting position, similar to its fossil relative

Stephanorhinus hemitoechus and Elasmotherium as well as the white rhinoceros.[28] Strong muscles on its long occipital bone formed its neck hock and held the massive skull. Its massive lower jaw measured up to 60 cm (24 in) long and 10 cm (4 in) high.[24] The teeth of the woolly rhinoceros had thickened enamel and an open internal cavity.[24] Like other rhinos, adults did not have incisors.[29] It had 3 premolars and 3 molars in both jaws. The molars were high-crowned and had a thick coat of cementum.[12]

The nasal septum of the woolly rhinoceros was

tichorhinus, from Greek τειχος (teikhos) "wall", ῥις (ῥιν-) (rhis (rhin-)) "nose".

Paleobiology

cave lions, and horses in late Pleistocene northern Spain, by Mauricio Antón

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.[29] The two teats in the female suggest that she raised one calf, or more rarely two, every two to three years.[33][22]

With their massive horns and size, adults had few predators, but young individuals 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.[34]

Woolly rhinos may have used their horns for combat, probably including

intraspecific combat as recorded in cave paintings, as well as for moving snow to uncover vegetation during winter.[27] They may have also been used to attract mates.[30] Bull woolly rhinos were probably territorial like their modern counterparts, defending themselves from competitors, particularly during the rutting season. Fossil skulls indicate damage from the front horns of other rhinos,[34]  and lower jaws and back ribs show signs of being broken and re-formed, which may have also come from fighting.[35] The apparent frequency of intraspecific combat, compared to recent rhinos, was likely a result of rapid climatic change during the last glacial period, when the animal faced increased stress from competition with other large herbivores.[31]

Charging woolly rhinoceros

Diet

Woolly rhinoceroses mostly fed on grasses and

Pollen analysis shows it also ate woody plants (including coniferswillows and alders),[24] along with flowers,[36] forbs and mosses.[37] Isotope studies on horns show that the woolly rhinoceros 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 winter.[38]

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.[39]

Comparisons with living

Artemisia, and graminoids.[40]

Growth

It is estimated that woolly rhinoceroses could reach around 40 years of age, like their modern relatives.[34] In 2014, Shpansky analysed the growth of woolly rhinoceros from its early life stages based on several lower jaw fragments and limb bones. A one-month-old calf was about 1.2 metres (3.9 ft) in length and 72 centimetres (2.36 ft) tall at the shoulder. The most intensive growth in woolly rhinos occurred during the juvenile stage around 3 to 4 years of age with a shoulder height of 1.3 metres (4.3 ft). At 7 to 10 years of age, woolly rhinos became young adults with a shoulder height of 1.4–1.5 metres (4.6–4.9 ft). By more than 14 years of age, woolly rhinos became fully mature, old adults with a shoulder height of 1.6 metres (5.2 ft).[41]

Habitat and distribution

Range of the woolly rhinoceros, including sites of fossils

The woolly rhinoceros lived mainly in

lowlands, plateaus and river valleys, with dry to arid climates,[42] 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.[42] 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.[43] With its wide distribution, the woolly rhinoceros lived in some areas alongside the other rhinoceroses Stephanorhinus[25] and Elasmotherium.[44]

By the end of 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.[27][45] It had the widest range of any rhinoceros species.[46]

It seemingly did not cross the

Chukotka Peninsula,[27] probably due to the low grass density and lack of suitable habitat in the Yukon combined with competition from other large herbivores on the frigid land bridge.[42]

Relationship with humans

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.[27] Many rhinoceros remains are found in caves (such as the Kůlna 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.[47] Sometimes, only individual teeth or bone fragments are uncovered, which usually came from only one animal.[48] 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.[49][50]

Signs that early humans hunted or scavenged the rhinoceros 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 marking the shoulder and thigh, and a preserved

Gudenus Cave (Austria)[51] and the open air site of Königsaue (Saxony-Anhalt, Germany),[52] have heavily beaten rhinoceros bones lined with slash marks. This action was done partly to extract the nutritious bone marrow.[53]

Both horns and bones of the rhinoceros were used as raw materials for tools and weapons, as were remains from other animals.

Yana River.[56] 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.[57]

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.[58]

Ancient art

Cave paintings in Chauvet Cave

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.[59]

About 20 Paleolithic drawings of woolly rhinos were known before the discovery of the

Kapova Cave in the Ural Mountains.[61] Some images show rhinoceroses struck with spears or arrows, signifying human hunting.[62]

The site of

statuettes of animals, many of woolly rhinoceroses.[62]

Extinction

Analysis of the nuclear genome suggests that the woolly rhinoceros experienced a population expansion beginning around 30,000 years ago.[63] The end of the last glacial period shows a progressive contraction of the range of the woolly rhinoceros, with the species disappearing from Europe during the interval between 17-15,000 years ago, with its youngest confirmed records being from the Urals, dating to 14,200 years ago, and northeast Siberia, dating to around 14,000 years ago. The youngest records of the species coincide with the onset of the Bølling–Allerød warming, which likely resulted in increased precipitation (including snowfall), which transformed the woolly rhinoceros' preferred low-growing grass and herb habitat into one dominated by shrubs and trees.[27] Population fragmentation is likely to have played a role in its extinction.[64] A genetic study of the woolly rhinoceros remains in northeast Siberia, dating to around 18,500 years ago, a few thousand years before its extinction, found that the population size was stable and relatively large, despite long-term co-existence with humans in the region.[63] A Holocene survival of the species has been suggested by the finding of environmental DNA of the woolly rhinoceros in sediments dating to 9,800 ± 200 years ago.[65] However, it has been demonstrated that ancient DNA in permafrost can be reworked into sediment layers dating to well after the extinction of the originating species.[66][67]

Fossil specimens

Frozen specimens

Mummified remains discovered in 1771

Many rhinoceros remains have been found preserved in the

Yana River.[24]

In October 1907, miners in

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.[73]

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

Skull and rib fragments of a rhinoceros were found in 1972 in

Aldan River's left bank, uncovering a skull with both horns, a spine, ribs and limb bones.[75]

In 2007, a partial rhinoceros carcass was found in the lower reaches of the

tail 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 two hunters, 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.[78] Dental analysis shows that the calf was about seven months old at the time of its death.[79] With its well-intact preservation, scientists proceeded to undergo DNA analysis.[80][81]

In August 2020, a rhinoceros was found, after being revealed by melting permafrost, close to the site of the 2014 discovery. The rhino was between three and four years old and it is thought that the cause of death was drowning. It is one 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 normally decompose quickly.[82]

See also

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External links