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Woolly rhinoceros

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.

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 was similar in appearance to modern rhinos, with some differences in physique and body structure. It 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. Like all rhinoceroses, it was an exclusively herbivorous animal that fed mainly on herbaceous plants that grew in abundance in the mammoth 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.

EvolutionEdit

 
Woolly rhinoceros by Charles R. Knight

As the last and most derived member of the Pleistocene rhinoceros lineage, the woolly rhinoceros was well adapted to its environment. Stocky limbs and thick woolly pelage made it well suited to the Mammoth steppe environment prevalent across the Palearctic ecozone during the Pleistocene glaciations. Like the vast majority of rhinoceroses, the body plan of the woolly rhinoceros adhered to a conservative morphology, like the first rhinoceroses seen in the late Eocene.

The immediate ancestors of the woolly rhinoceros most likely appeared about 2 million years ago in East Asia, in the northern foothills of the Himalayas. 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. The members of Coelodonta turned out to be more adapted to a variety of conditions compared to elasmotheres. The initial evolution of the genus probably occurred in humid spaces, which explains the absence of remains from the Miocene. The evolution of the woolly rhinoceros began in a frost-free climate, adapting to the cold when the Early Pleistocene climate fluctuated in the area around and north of the Himalayas.[2] Other sources say that the group closest to the woolly rhinoceros was the Early Pleistocene Stephanorhinus, in particular, the species Stephanorhinus hemitoechus.[3] Using the methods of palaeoproteomics, a preserved rhino from Dmanisi known as Stephanorhinus ex gr. etruscus-hundsheimensis was found to be 1.77 million years old. This refers to an earlier line in relation to the related woolly rhinoceros (Coelodonta antiquitatis) and Merck’s rhinoceros (Stephanorhinus kirchbergensis). The genus Coelodonta is descended from the early line of Stephanorhinus.[4]

It is believed that the woolly rhinoceros descended from an earlier member of the genus, Coelodonta tologoijensis. Another rhino belonging to the Middle Pliocene, Coelodonta thibetana, is also mentioned as an ancestor of the woolly rhinoceros.[3] The evolution of the woolly rhinoceros as an independent species occurred at the end of the Early Pleistocene, more than 300 thousand years ago in Central Asia. From here, woolly rhinos migrated north and west into Europe. The woolly rhinoceros became one of the most common inhabitants of the Mammoth steppe, a typical representative of the megafauna.

A study of 40,000- to 70,000-year-old DNA samples showed its closest extant relative is the Sumatran rhinoceros.[5]

DescriptionEdit

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

The appearance of woolly rhinos is known from mummified individuals from Siberia as well as cave paintings.[6] An adult woolly rhinoceros was typically around 3 to 3.8 metres (9.8 to 12.5 ft) in length, with an estimated weight of around 1,800–2,700 kg (4,000–6,000 lb)[7] or 2,000 kg (4,400 lb).[8] The woolly rhinoceros could grow to be 2 m (6.6 ft) tall;[7] the body size was thus comparable to, or slightly larger than, the extant white rhinoceros.[9] Two horns on the skull were made of keratin, the anterior horn being 61 cm (24 in) in length,[10] with a smaller horn between its eyes.[11] It had thick, long fur, small ears, short, thick legs, and a stocky body. Cave paintings suggest a wide dark band between the front and hind legs, but the feature is not universal, and the identification of pictured rhinoceroses as woolly rhinoceros is uncertain.

Compared to other members of its family, the woolly rhinoceros was shorter-legged, with a more elongated head and body. The scruff of the woolly rhinoceros was raised with its powerful hump, which supported its massive front horn. Formed by the animal’s strongly developed muscles, the hump contained a significant amount of fat, a necessary reserve of nutrients to survive in the desolate mammoth steppe.[12] The legs of a woolly rhinoceros, like those of modern rhinos, were three-toed. The woolly rhinoceros had no incisors or canine teeth. Compared with modern rhinoceroses, its teeth were more powerful with thickened enamel.[13] The teeth of the woolly rhinoceros, like other closely related rhinos of the genus Coelodonta, had an open internal cavity.

As the name suggests, woolly rhinoceroses were covered with long hair. Wool is rarely found on preserved carcasses, but the surviving specimens are reddish-brown in color, sometimes with a yellowish tint. There was a thick undercoat, under a layer of long, coarse guard hair that reached its thickest point on the withers and neck. The limbs were covered with shorter hair, keeping snow from attaching.[12] The body's length ended with a 45-50-centimeter tail with a brush of coarse hair at the end.[14][13] Females had two nipples located in the inguinal region.

A number of external features of the woolly rhinoceros indicate its adaptability to long-term severe frosts. Its ears were relatively much smaller than those of rhinoceros species in tropical climates; the preserved ears of woolly rhinos have a length of no more than 24 cm, while modern rhinos living in hot climates have a length of about 30 cm.[13] Their tails were also relatively much shorter. These adaptations reduced the body's total surface area from which heat loss occurs, as is common to animals in cold climates. The skin of the woolly rhinoceros was very thick, also reducing the loss of body heat. It ranged from 5 to 15 mm in thickness, heaviest on the chest and shoulders.[3][13]

Its shape was known only from prehistoric cave drawings until a completely preserved specimen (missing only the fur and hooves) was discovered in a tar pit in Starunia, Ukraine. The specimen, an adult female, is now on display in the Polish Academy of Sciences' Museum of Natural History in Kraków. Several frozen specimens have also been found in Siberia, the latest in 2015.[15]

HornsEdit

The woolly rhinoceros had two horns, found on both males and females. Like all rhinoceroses, it used its horns for defensive purposes and to attract mates.[16] However, the horns of the woolly rhinoceros served another purpose in raking away snow. This was made possible by their unique shape, flattened along the edges and curved backwards.

The front horn reached a considerable size. Its length reached a meter or more, up to 1.4 m,[3][17] and its weight reached 15 kg. The front horn faced forward, to a much greater extent than those of modern rhinos.[3] One specimen found in the Kolyma basin had a front horn with a length of 84.5 cm around the outer edge, along with a base 22.9 cm long and 12.3 cm wide. As typical for the species, the middle of the horn reached a much thinner length, at only 23 mm. The second horn was significantly shorter, no more than half a meter, at about 15 cm long from a 14.6 × 8 cm base.[13]

The nasal septum of the woolly rhinoceros was ossified, which is not observed in modern rhinos.[18] This appears to be another adaptation to increased pressure on the horn and the entire face when feeding.[19] This feature was most prevalent among adult males.

Found horns of woolly rhinos bear traces of friction caused by contact with snow. Their abrasions suggest that the rhino often moved its head from side to side as it dug snow with its horn, extracting vegetation beneath.[7]

DistributionEdit

 
Range of the woolly rhinoceros, including sites of fossils

By the end of the Riss glaciation (about 130 thousand years ago), the area of the woolly rhinoceros occupied a huge space, which included almost all of Eurasia north of the tropical zone. The rhinoceros inhabited most of Europe, the Russian Plain, the south of western and eastern Siberia, and the Mongolian Plateau, ranging to extremes of 72° to 33° north latitude. Findings of woolly rhinos occur even in the New Siberian Islands.[3][20] In 2011, a 3.6-million-year-old woolly rhinoceros fossil, the oldest known, was discovered on the cold Tibetan Plateau, suggesting that it existed there during a period of general climate warmth around the earth. It is believed they migrated from there to northern Asia and Europe when the Ice Age began.[21]

During Greenland Stadial 2 (the Last Glacial Maximum[22]) the North Sea retreated northward, as sea levels were up to 125 metres (410 ft) lower than today. The woolly rhinoceros roamed the exposed Doggerland and much of Northern Europe and was common in the cold, arid desert that later became southern England[23] and the North Sea. Its geographical range expanded and contracted with the alternating cold and warm cycles, forcing populations to migrate as glaciers receded. The woolly rhinoceros co-existed with woolly mammoths and several other extinct larger mammals of the Pleistocene megafauna. A close relative, Elasmotherium, had a more southerly range.

The rhinoceros was apparently absent in Japan[3] and in Ireland since its bones were not found there.[7] In the northern parts of central Siberia, it was also not common. The lack of woolly rhinoceros remains in North America suggests that none were found there, and represents a certain mystery to science. It remains unclear why woolly rhinoceroses did not cross the Bering land bridge to reach this continent, while other large animals such as the mammoth and the steppe bison did (especially since rhinos were found in the Chukotka Peninsula, the easternmost part of Asia).[3]

Likely, the rhino did not migrate to North America due to strong food competition from other large ungulates in Beringia, where the food supply was limited. Glaciated mountain ranges in Alaska would have posed a physical barrier, while vegetation in the north Yukon may have consisted more of bushes and sedge swamp rather than the grass they preferred.[24] Rhinos do not form herds, suggesting a low potential for migration, compared to other Pleistocene mammals such as mammoths, bison, and horses. Individual rhinoceros visits to the North American continent are not ruled out, but it most likely never became a long-term habitat.[20]

Behavior and habitatEdit

 
Restoration of a woolly rhinoceros

ReproductionEdit

Little is known about the reproduction of the woolly rhinoceros. Estimates by comparison with modern species suggest that woolly rhinos formed pairs once every 3-4 years for the short time needed for mating. During this period, the males entered into battle with each other as they competed for mating with a female rhinoceros. The presence of only two teats in the female suggests that she usually gave birth to one, or more rarely, two calves.[25][26] Pregnancy lasted about a year and a half. The calf remained with its mother for several months to two years, before searching for its own individual territory. This implies that the natural reproduction of woolly rhinos was very slow – during 20–25 years of fertility, the female could produce only 6–8 calves.

The development of young animals was similar to that of modern rhinoceros species. Development and change of milk teeth in the woolly rhinoceros correlates with the same data on calves of the white and black rhinoceros.[27] However, early life stages of the woolly rhinoceros are poorly studied due to the absence of preserved bodies of nursing calves.[27]

DietEdit

 
Frozen head, leg and horn from Siberia, 1849

Controversy has long surrounded the precise dietary preference of Coelodonta as past investigations have found both grazing and browsing modes of life to be plausible. The palaeodiet of the woolly rhinoceros has been reconstructed using several lines of evidence. Climatic reconstructions indicate the preferred environment to have been cold and arid Mammoth steppe, with large herbivores forming an important part of the feedback cycle. Pollen analysis shows a prevalence of grasses and sedges within a more complicated vegetation mosaic.[28]

Numerous anatomical features suggest that the woolly rhinoceros had a diet largely consisting of grasses, as well as shrubs and branches.[13] These include a long, slanted head with its downward-facing posture, high-crowned teeth with their high cement content and the graceful form of the second premolar.[29] Horizontal abrasion of the teeth has been observed, which is caused by the silica contained in grasses. Coniferswillows and alders were eaten.[24] Recent isotope studies on horns indicate 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.[30]

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.

Comparisons with extant perissodactyls confirm that Coelodonta was a hindgut fermentor with a single stomach, and as such would have grazed upon cellulose-rich, protein-poor fodder. This method of digestion would have required a large throughput of food and thus links the large mouthful size to the low nutritive content of the chosen grasses and sedges.[31]

Recent evidence suggests that woolly rhinos lived in the Arctic during the Last Glacial Maximum consumed approximately equal volumes of forbs, such as Artemisia, and graminoids.[32]

LifestyleEdit

Whether the woolly rhinoceros was a social animal is not known, but is relatively unlikely, due to the fact that modern rhinoceros species are predominantly solitary. Bull rhinoceroses live territorially, defending their territory against potential competitors including other rhinoceroses. Some skulls of woolly rhinos show injuries that indicate battles with other rhinos. Piercings occur on the skull, often on the parietal bone, but also on the orbit or the maxilla. Broken and re-deformed lower jaws are preserved, as well as broken and partly healed back ribs, which may have also resulted from fighting.[33] 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.

ExtinctionEdit

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

Many species of Pleistocene megafauna, like the woolly rhinoceros, became extinct around the same time period. Human hunting is often cited as one cause.[34] 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).[35] 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 (Shapiro). This suggests that climate change was not the only factor contributing to the rhinoceros's extinction (Naish). 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.

Recent radiocarbon dating indicates that populations survived as recently as 8,000 BC in western Siberia. 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.).

Fossil specimensEdit

 
Mounted woolly rhinoceros skeleton in the Ruhr Museum in Essen, Germany

Woolly rhinoceroses are among the most common fossil finds of Pleistocene species. Their mostly isolated bones and teeth are mainly found from gravel, sand and clay pits, that indicate former rivers or lake shores. Other finds come from caves. In addition, rhinoceros remains have been recovered from shelf seas, such as the North and Baltic, which were dry during periods of glaciation.[33] More than 50 sites are known in Poland alone, and at least 20 in Iberia.[36][37] Bones have also been located from the Italian Peninsula.[38] In China there are more than 70 places where rhinoceros remains have been discovered.[39][40]

The woolly rhinoceros is also known from many finds in Germany, including more than 30 known localities in Westphalia alone. [33] However, articulated skeletal remains or even complete skeletons are extremely rare. One of the earliest skeletons found comes from Thuringia, and is now exhibited in the Museum of Natural History in Gera. Another was salvaged in a gravel and sand pit near Petershagen, containing 66 skeletal elements including skulls; it is now in the Natural History Museum in Bielefeld.[41]

Frozen specimensEdit

 
A mummified carcass of woolly rhinoceros in Starunia, 1929

Preserved carcasses from the woolly rhinoceros have been found in the permafrost region, giving clear information about the animal's soft tissue anatomy. Around half a dozen are known, mostly from Yakutia. These are far less common than those of the mammoth.[42] One of the first known frozen carcasses was discovered in 1771 in the Vilyuy River.[43] Later in 1877, a Siberian trader discovered mummified rhino remains from a tributary of the Yana River from which a head and one leg were recovered.[13] One finding from 1972 came from Churapcha, between the Lena and the Amga rivers. Only individual skull and rib fragments were initially found, but further investigations that year found a whole skeleton with preserved skin, fur, and stomach contents.[13][44] A short time later, in 1976, a skull, horn, vertebrae and other rhinoceros bones were found by schoolchildren during a class trip on the left bank of the Aldan River.[45]

In 2007, at the lower reaches of the Kolyma, the skeleton of a rhinoceros was found with the skull and left part of the body preserved, along with skin and hair on its limbs.[46][13] Next year in 2008, findings from the Chukochya River yielded a nearly complete skeleton with a skull and two horns. [47] That same year, partly mummified rhinoceros remains were discovered by local residents near the Amga and were excavated over the years of 2009 and 2010. Here, pelvic bones, forelimbs and hind limbs were presented complete with fingers and toes, caudal vertebrae and ribs. [48]

In September 2014, at an unnamed tributary of Semjuljach River in Yakutia, Abyysky District, a mummified young rhinoceros was discovered by Alexander “Sasha” Banderov and Simeon Ivanov. Parts of the body not covered by permafrost had already been thawed and consumed by animals. The intact remains in permafrost included the head with its fur and horns, and soft tissue parts were also recovered. The two excavators who found it handed over their discovery to the Yakutia Academy of Sciences, who named it “Sasha” after one of its discoverers.[49] After their first assessment, the remains are well preserved. According to dental analysis, the calf is estimated to have been seven months old at the time of its death. However, it was the same size as an eighteen-month old modern rhinoceros, due to the immense size of the extinct animal.[50] Due to the state of preservation, scientists hope to obtain usable DNA.[49][51]

Outside of Yakutia and the permafrost area, four individuals have been found in an ozokerite pit in Starunia. The first carcass, with a complete body and skull and two intact horns, was found in 1907. Another mummy was found in 1929, completely preserved aside from its horns. Two partially preserved remains of other rhinoceroses were located nearby, with their soft tissue well preserved, but having lost fur coverage. The second complete carcass is now exhibited in the Aquarium and Natural History Museum in Kraków.[52][53]

Relationship with humansEdit

 
Reconstruction of woolly rhinoceros with human hunter

The rhinoceros appears to not have been one of the most frequent hunting targets by ancient people. Remains and images of woolly rhinoceros are associated with only 11% of the known sites of the Paleolithic tribes of Siberia.[3]

However, there is clear evidence that prehistoric people hunted the woolly rhino. In Western Europe, particularly France, most rhinoceros remains are found at sites frequented by Paleolithic humans. One of the corpses of a rhino, found in 1907 in western Ukraine, had injuries sustained from a human. Traces of a wound with a sharp object marked the shoulder and thigh, and a preserved spear was found near the carcass. However, this rhino was not killed by hunters; it died having fallen into a hole filled with ozokerite, due to which it was well preserved.[13] Meat and fat of the rhinoceros was eaten, and its horns and bones were used for a variety of crafts. On the banks of the Yana River, half-meter spear throwers made from the horn of a woolly rhinoceros about 27 thousand years ago were found.

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.[54]

Ancient artEdit

 
Chauvet cave art depicting a woolly rhinoceros

The woolly rhinoceros is often represented among cave paintings from the Upper Paleolithic. Depictions of the rhinoceros demonstrate the characteristic back line with the high neck hump, as shown by the mummified carcasses, along with the low posture of the head. Ears are indicated by two arcuate lines or a black band dividing the body. The animal’s long horns are prominently featured. In some cases, the coat is also indicated. These illustrations give a deep insight into the appearance of this Pleistocene species, in accompaniment to its scientific studies.[11]

Among the earliest illustrations are from the Chauvet Cave in France, probably from the Aurignacian, with an age of more than 31,000 years.[3] 47 drawings are located from the cave, making around a quarter of all of its animal drawings. They are alternately drawn in red or black, while other depictions were made as engravings. Of particular note is a scene of two rhinos facing each other, fighting with their horns.[55]

Other noteworthy depictions are those from the Rouffignac and Lascaux caves. One drawing from Font-de-Gaume shows a noticeably higher head posture, and red-pigmented drawings of woolly rhinos appear in the Kapova Cave in the Ural Mountains, which belong to the late Upper Palaeolithic.[56]

A woolly rhinoceros statuette was found in Dolní Věstonice in Moravia.[57]

See alsoEdit

Elasmotherium, another woolly rhinoceros species

ReferencesEdit

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