Woolly rhinoceros

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. A rhinoceros skull was found in Klagenfurt, Austria, in 1335, and was believed to be that of a dragon. In 1590, it was used as the basis for the head on a statue of a lindworm. 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". 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. In 1772, Pallas acquired a head and two legs of a rhinoceros from the locals in Irkutsk, and named the species Rhinoceros lenenesis (after the Lena River). 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. This name comes from the Greek words κοιλος (koilos, "hollow") and ὀδούς (odoús "tooth"), from the depression in the rhino's molar structure, giving the scientific name Coelodonta antiquitatis, "hollow-tooth of antiquity". 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. |style=font-size:100%;line-height:80%}} Relationships of the woolly rhinoceros based on morphology, excluding African rhinoceros species:The ancestors of Coelodonta are suggested to have diverged from those of the Sumatran rhinoceros around 9.4 million years ago, with Coelodonta diverging from Stephanorhinus around 5.5 million years ago. with the genus being present in Siberia, Mongolia, and China during the Early Pleistocene. The woolly rhinoceros first appeared during the early 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. 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. ==Description==

Size and general morphology An adult woolly rhinoceros typically measured from head to tail, stood tall at the shoulder, and weighed up to (with some sources estimate the species' maximum shoulder height at and its weight at ) making it comparable in size to the largest living rhinoceros species, the white rhinoceros (Ceratotherium simum). Compared to other rhinoceroses, the woolly rhinoceros had a longer head and body, and shorter legs. Skull and dentition The skull had a length between . 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. Unlike in modern rhinos, the large nasal horn was often flattened in cross-section, and abrasion patterns on the horn indicate its possible use in brushing away snow when grazing. The nasal septum of the woolly rhinoceros was ossified, unlike modern rhinos. This was most common in adult males. This ossification inspired the junior synonym specific name tichorhinus, from Greek τειχος (teikhos) "wall", ῥις (ῥιν-) (rhis (rhin-)) "nose". External appearance Frozen specimens indicate that the woolly rhino's long fur coat was brown in adults and light brown in juveniles, 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. Females had two nipples on the udders. Beneath the skin, the body was covered in a layer of subcutaneous fat, ranging from thick on the chest, to on the lower jaw and the posterior part of the back. ==Paleobiology and palaeoecology==

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. The two teats in the female suggest that she raised one calf, or more rarely two, every two to three years. which to a large degree likely reflects scavenging of the carcasses of already dead rhinoceroses. A piece of juvenile woolly rhinoceros skin with blond fur (possibly representing that of a calf) was recovered from the stomach content of two frozen juvenile female Pleistocene wolves, which was inferred to be part of the wolves' last meal. 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.  and lower jaws and back ribs show signs of being broken and re-formed, which may have also come from fighting. 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. 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. It had a wide upper lip like that of the white rhinoceros, which allowed it to easily pluck vegetation directly from the ground. 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. Woolly rhinos living in the Arctic during the Last Glacial Maximum consumed approximately equal volumes of forbs, such as Artemisia, and graminoids. Pollen analysis shows it also ate woody plants (including conifers, willows and alders), forbs and mosses. Dental mesowear measurements further show that the woolly rhinoceros's diet was heavily composed on abrasive grasses. Growth and pathologies An adult female discovered in Yakutia suggests that it could have had a lifespan of about 40 years or more, similar to that of its modern relatives. C. antiquitatis individuals of old age display extensive wear and loss of their anterior premolars as a result of tooth abrasion from their intensive grazing lifestyle. Habitat and distribution The woolly rhinoceros lived mainly in lowlands, plateaus and river valleys, with dry to arid climates, With its wide distribution, the woolly rhinoceros lived in some areas alongside the other rhinoceroses Stephanorhinus and Elasmotherium. By the end of the Riss glaciation about 130,000 years ago, 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 as far north as the New Siberian Islands. Even during the very warm Eemian interglacial, the range of the woolly rhinoceros extended into temperate regions such as Poland. It had the widest range of any rhinoceros species. 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, ==Relationship with humans==

Hunting Paired δ13C and δ15N measurements of fossils from Les Pradelles indicate that woolly rhinoceroses made up a significant proportion of the diet of the local Neanderthal population. Woolly rhinoceroses shared their habitat with modern 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. Sometimes, only individual teeth or bone fragments are uncovered, which usually came from only one animal. 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. 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 spear was found near the carcass. and the open air site of Königsaue (Saxony-Anhalt, Germany), have heavily beaten rhinoceros bones lined with slash marks. This action was done partly to extract the nutritious bone marrow. Both horns and bones of the rhinoceros were used as raw materials for tools and weapons, as were remains from other animals. In what is now Zwoleń, Poland, a device was made from a battered woolly rhinoceros pelvis. 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. 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. Ancient art 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. About 20 Paleolithic drawings of woolly rhinos were known before the discovery of the Chauvet Cave in France. 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. Some images show rhinoceroses struck with spears or arrows, signifying human hunting. 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. ==Extinction==

Analysis of the nuclear genome suggests that the woolly rhinoceros experienced a population expansion beginning around 30,000 years ago. Population fragmentation is likely to have played a role in its extinction. The presence of large numbers of abnormal cervical ribs in specimens from the North Sea, much higher than that found in living rhinoceroses, may have been the result of inbreeding due to low population size or harsh environmental conditions. 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. A specimen of 14,400-year old woolly rhinoceros muscle tissue found preserved in a permafrost-frozen juvenile wolf's stomach in the Tumat region of far north-northeastern Siberia, which represents among the youngest specimens of the species, had a similar level of genetic diversity to the 18,500-year old specimen, implying that the woolly rhinoceros population in the region was stable up until a likely rapid decline during the Bølling–Allerød warming. A Holocene survival of the species has been suggested by the finding of environmental DNA of the woolly rhinoceros in sediments of the Kolyma region of Northeast Siberia dating to 9,800 ± 200 years ago. 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, though other authors have argued that this specific environmental DNA record is unlikely to have been reworked. == Frozen specimens ==

Many rhinoceros remains have been found preserved in the permafrost region. In 1771, a head, two legs and hide were found in the Vilyuy River in eastern Siberia and sent to the Kunstkamera in Saint Petersburg. Later in 1877, a Siberian trader recovered a head and one leg from a tributary of the Yana River. Both were sent to the Dzieduszycki Museum, where a detailed description was published in the museum's monograph. Photographs were published in paleontological journals and textbooks, and the first modern paintings of the species were based on the mounted specimen. Later, in 1929, the Polish Academy of Arts and Sciences sent an expedition to Starunia, finding the mummified remains of three rhinos. One specimen, missing only its horns and fur, 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. 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. 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. In 2007, a partial rhinoceros carcass was found in the lower reaches of the Kolyma river. Its upward-facing position indicates that the animal probably fell into mud and sank. That same year, locals near the Amga discovered mummified rhinoceros remains, and over the next two years, pelvic bones, tail vertebrae and ribs were excavated along with forelimbs and hind limbs with toes intact. 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. Dental analysis shows that the calf was about seven months old at the time of its death. With its well-intact preservation, scientists proceeded to undergo DNA analysis. 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. ==See also==