:Woolly rhinoceros

File:Em - Coelodonta antiquitatis - 2.jpg

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.{{cite book | last=Howorth | first=H.H. | title=The Mammoth and the Flood: An Attempt to Confront the Theory of Uniformity with the Facts of Recent Geology | publisher=S. Low, Marston, Searle, & Rivington | year=1887 | url=https://books.google.com/books?id=Jl3PAAAAMAAJ&pg=PA7 | page = 7}} A rhinoceros skull was found in Klagenfurt, Austria, in 1335, and was believed to be that of a dragon.{{cite journal | last=Price | first=Samantha | title=Horns, Tusks, and Flippers: the Evolution of Hoofed Mammals. | journal=Aquatic Mammals | publisher=Aquatic Mammals Journal | volume=33 | issue=2 | date=2007-06-01 | issn=0167-5427 | doi=10.1578/am.33.2.2007.254 | pages=254 | bibcode=2007AqMam..33..254P | s2cid=127893230 | url=https://pdfs.semanticscholar.org/1ecd/7f32bb164d61a4093bf9f8e9530775ff335a.pdf | archive-url=https://web.archive.org/web/20191208204604/https://pdfs.semanticscholar.org/1ecd/7f32bb164d61a4093bf9f8e9530775ff335a.pdf | url-status=dead | archive-date=2019-12-08 }} In 1590, it was used as the basis for the head on a statue of a lindworm.{{cite web | title=RRC: The Klagenfurt Lindwurm | website=RRC | url=http://www.rhinoresourcecenter.com/images/The-Klagenfurt-Lindwurm_i1363386763.php | access-date=2019-12-08}} 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".Schubert, von, G.H., 1823. Die Urwelt und die Fixsterne: eine Zugabe zu den

Ansichten von der Nachtseite der Naturwissenschaft [The Primeval World and the Fixed Stars]. Arnoldischen Buchhandlung, Dresden.

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.{{cite book | last1=Pallas | first1=P.S. | last2=Blagdon | first2=F.W. | title=Travels Through the Southern Provinces of the Russian Empire: In the Years 1793 and 1794 | publisher=John Stockdale, Piccadilly | series=Travels Through the Southern Provinces of the Russian Empire, in the Years 1793 and 1794 | year=1812}} In 1772, Pallas acquired a head and two legs of a rhinoceros from the locals in Irkutsk,{{cite book | last=Owen | first=Richard. | title=A History of British Fossil Mammals, and Birds | publisher=J. Van Voorst | year=1846}} and named the species Rhinoceros lenenesis (after the Lena River). In 1799, Johann Friedrich Blumenbach studied rhinoceros bones from the collection of the University of Göttingen, and proposed the scientific name Rhinoceros antiquitatis.Gehler, Alexander & Reich, Mike & Mol, Dick & Plicht, Hans. (2007). The type material of Coelodonta antiquitatis (Blumenbach) (Mammalia: Perissodactyla: Rhinocerotidae) / Типовой материал Coelodonta antiquitatis (Blumenbach) (Mammalia: Perissodactyla: Rhinocerotidae) / Tipovoj material Coelodonta antiquitatis (Blumenbach) (Mammalia: Perissodactyla: Rhinocerotidae). 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.{{cite book | last=Bronn | first=H.G. | title=Lethaea geognostica oder Abbildungen und Beschreibungen der für die Gebirgs-Formationen bezeichnendsten Versteinerungen | trans-title= Lethaea Geognostica, or pictures and descriptions of the most characteristic fossils of the mountain formations| publisher=Stuttgart | year=1838 | volume=2 | url=https://www.biodiversitylibrary.org/page/39079713#page/1211/ | page=451 | language=de}} This name comes from the Greek words κοιλος (koilos, "hollow") and ὀδούς (odoús "tooth"), from the depression in the rhino's molar structure,{{cite journal|last=van der Made|first=Jan|title=The rhinos from the Middle Pleistocene of Neumark-Nord (Saxony-Anhalt)|journal=Veröffentlichungen des Landesamtes für Archäologie|trans-journal=State Office of Archeology Saxony-Anhalt|volume=62|pages=432–527|url=http://www.rhinoresourcecenter.com/pdf_files/129/1295057876.pdf|language=de, en}}{{cite book | last=von Koenigswald | first=W. | title=Lebendige Eiszeit: Klima und Tierwelt im Wandel | trans-title=Living Ice Age: Climate and wildlife in transition | publisher=Theiss | year=2002 | isbn=978-3-8062-1734-6 | language=de}} giving the scientific name Coelodonta antiquitatis, "hollow-tooth of antiquity".{{cite web | author=RFE/RL | title=Siberia Surrenders Woolly Rhino Mysteries | website=RadioFreeEurope/RadioLiberty | date=2012-12-06 | url=https://www.rferl.org/a/russia-zoology-wolly-rhinoceros-boeskorov/24791250.html | access-date=2019-12-09}}

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.{{Cite journal |last1=Liu |first1=Shanlin |last2=Westbury |first2=Michael V. |last3=Dussex |first3=Nicolas |last4=Mitchell |first4=Kieren J. |last5=Sinding |first5=Mikkel-Holger S. |last6=Heintzman |first6=Peter D. |last7=Duchêne |first7=David A. |last8=Kapp |first8=Joshua D. |last9=von Seth |first9=Johanna |last10=Heiniger |first10=Holly |last11=Sánchez-Barreiro |first11=Fátima |date=24 August 2021 |title=Ancient and modern genomes unravel the evolutionary history of the rhinoceros family |journal=Cell |volume=184 |issue=19 |pages=4874–4885.e16 |doi=10.1016/j.cell.2021.07.032 |issn=0092-8674 |pmid=34433011 |doi-access=free|hdl=10230/48693 |hdl-access=free }}

{{clade|{{clade

|label1=Elasmotheriinae

|1={{extinct}}Elasmotherium

|label2=Rhinocerotinae

|2={{clade

|1={{clade

|1=Black rhinoceros (Diceros bicornis)

|2=White rhinoceros (Ceratotherium simum)

}}

|2={{clade

|label1=

|1={{clade

|1=Indian rhinoceros (Rhinoceros unicornis)

|2=Javan rhinoceros (Rhinoceros sondaicus)

}}

|label2=

|2={{clade

|1=Sumatran rhinoceros (Dicerorhinus sumatrensis)

|2={{clade

|1={{extinct}} Woolly rhinoceros (Coelodonta antiquitatis)

|2={{extinct}} Merck's rhinoceros (Stephanorhinus kirchbergensis)

}}

}}

}}

}}

}}|style=font-size:100%;line-height:80%}}

Relationships of the woolly rhinoceros based on morphology, excluding African rhinoceros species:{{Cite journal |last=Pandolfi |first=Luca |date=2023-01-19 |title=Reassessing the phylogeny of Quaternary Eurasian Rhinocerotidae |journal=Journal of Quaternary Science |volume=38 |issue=3 |language=en |pages=291–294 |doi=10.1002/jqs.3496 |bibcode=2023JQS....38..291P |issn=0267-8179|doi-access=free |hdl=11563/163194 |hdl-access=free }}{{Clade|{{clade

|label1=Rhinocerotina

|1={{Clade

|1={{Clade

|1=Gaindatherium browni

|2=Lartetotherium sansaniense

}}

|2={{Clade

|1={{Clade

| label1=Nesorhinus

| label2=Rhinoceros

|1={{Clade

|1=Nesorhinus hayasakai

|2=Nesorhinus philippinensis

}}

|2={{Clade

|1=Rhinoceros sondaicus (Javan rhinoceros)

|2=Rhinoceros unicornis (Indian rhinoceros)

}} }}

|2={{Clade

|label1=Dicerorhinus

|1={{Clade

|1=Dicerorhinus gwebinensis

|2={{Clade

|1=Dicerorhinus fusuiensis

|2=Dicerorhinus sumatrensis (Sumatran rhinoceros)

}} }}

|2={{Clade

|1=Dihoplus schleiermacheri

|2={{Clade

|1="Dihoplus" pikermiensis

|2={{Clade

|label1=Coelodonta

|1={{Clade

|1=Coelodonta thibetana

|2={{Clade

|1=Coelodonta nihowanensis

|2={{Clade

|1=Coelodonta tologoijensis

|2=Coelodonta antiquitatis (woolly rhinoceros)

}} }} }}

|2={{Clade

|label1=Pliorhinus

|label2=Stephanorhinus

|1={{Clade

|1=Pliorhinus megarhinus

|2=Pliorhinus miguelcrusafonti

}}

|2={{Clade

|1=Stephanorhinus jeanvireti

|2={{Clade

|1=Stephanorhinus etruscus

|2={{Clade

|1=Stephanorhinus hundsheimensis

|2={{Clade

|1=Stephanorhinus hemitoechus (Narrow-nosed or steppe rhinoceros)

|2=Stephanorhinus kirchbergensis (Merck's or forest rhinoceros)

}} }} }} }} }} }} }} }} }} }} }} }}|style=}}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. The oldest known species of Coelodonta, Coelodonta thibetana is known from the Pliocene of Tibet dating to approximately 3.7 million years ago,{{cite journal |author=Deng, T. |author2=Wang, X. |author3=Fortelius, M. |author4=Li, Q. |author5=Wang, Y. |author6=Tseng, Z.J. |author7=Takeuchi, G.T. |author8=Saylor, J.E. |author9=Säilä, L.K. |author10=Xie, G. |name-list-style=amp |year=2011 |title=Out of Tibet: Pliocene Woolly Rhino Suggests High-Plateau Origin of Ice Age Megaherbivores |url=https://www.science.org/doi/10.1126/science.1206594 |journal=Science |volume=333 |issue=6047 |pages=1285–1288 |doi=10.1126/science.1206594 |pmid=21885780 |bibcode=2011Sci...333.1285D |s2cid=8913866|url-access=subscription }} 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.{{Cite journal |last1=Uzunidis |first1=Antigone |last2=Antoine |first2=Pierre-Olivier |last3=Brugal |first3=Jean-Philip |date=July 2022 |title=A Middle Pleistocene Coelodonta antiquitatis praecursor (Mammalia, Perissodactyla) from Les Rameaux, SW France, and a revised phylogeny of Coelodonta |url=https://linkinghub.elsevier.com/retrieve/pii/S0277379122002256 |journal=Quaternary Science Reviews |language=en |volume=288 |pages=107594 |doi=10.1016/j.quascirev.2022.107594|bibcode=2022QSRv..28807594U }}{{Cite journal |last1=Stefaniak |first1=Krzysztof |last2=Kovalchuk |first2=Oleksandr |last3=Ratajczak-Skrzatek |first3=Urszula |last4=Kropczyk |first4=Aleksandra |last5=Mackiewicz |first5=Paweł |last6=Kłys |first6=Grzegorz |last7=Krajcarz |first7=Magdalena |last8=Krajcarz |first8=Maciej T. |last9=Nadachowski |first9=Adam |last10=Lipecki |first10=Grzegorz |last11=Karbowski |first11=Karol |last12=Ridush |first12=Bogdan |last13=Sabol |first13=Martin |last14=Płonka |first14=Tomasz |date=March 2023 |title=Chronology and distribution of Central and Eastern European Pleistocene rhinoceroses (Perissodactyla, Rhinocerotidae) – A review |url=https://linkinghub.elsevier.com/retrieve/pii/S1040618223000344 |journal=Quaternary International |volume=674-675 |language=en |pages=87–108 |doi=10.1016/j.quaint.2023.02.004|bibcode=2023QuInt.674...87S |s2cid=257337872 |url-access=subscription }} 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. Mitochondrial genomes suggest that the last mitochondrial ancestor of Late Pleistocene woolly rhinoceroses lived around 570,000 years ago.{{Cite journal |last1=Yuan |first1=Junxia |last2=Sun |first2=Guojiang |last3=Xiao |first3=Bo |last4=Hu |first4=Jiaming |last5=Wang |first5=Linying |last6=Taogetongqimuge |last7=Bao |first7=Lei |last8=Hou |first8=Yamei |last9=Song |first9=Shiwen |last10=Jiang |first10=Shan |last11=Wu |first11=Yong |last12=Pan |first12=Dong |last13=Liu |first13=Yang |last14=Westbury |first14=Michael V. |last15=Lai |first15=Xulong |date=2023-09-26 |title=Ancient mitogenomes reveal a high maternal genetic diversity of Pleistocene woolly rhinoceros in Northern China |journal=BMC Ecology and Evolution |language=en |volume=23 |issue=1 |page=56 |doi=10.1186/s12862-023-02168-0 |issn=2730-7182 |pmc=10521388 |pmid=37752413 |doi-access=free }}

File:Coelodonta antiquitatis by Benjamin Langlois.jpg

An adult woolly rhinoceros typically measured {{convert|3.2|to|3.6|m|ft|1}} from head to tail, stood {{convert|1.45|-|1.6|m|ft}} tall at the shoulder, and weighed up to {{convert|1.5|-|2|MT|ST}} (with some sources placing the body mass of the species as high as {{Convert|2.4-2.9|metric ton|lb|abbr=}}{{Cite journal |last1=Mallet |first1=Christophe |last2=Billet |first2=Guillaume |last3=Cornette |first3=Raphaël |last4=Alexandra Houssaye |first4=And |date=2022-11-02 |title=Adaptation to graviportality in Rhinocerotoidea? An investigation through the long bone shape variation in their hindlimb |url=https://academic.oup.com/zoolinnean/article/196/3/1235/6549627 |journal=Zoological Journal of the Linnean Society |language=en |volume=196 |issue=3 |pages=1235–1271 |doi=10.1093/zoolinnean/zlac007 |issn=0024-4082}}{{Cite journal |last1=Ma |first1=Jiao |last2=Wang |first2=Shiqi |last3=Deng |first3=Tao |date=2024-04-11 |title=When the woolly rhinoceroses roamed East Asia: a review of isotopic paleoecology of the genus Coelodonta from the Tibetan Plateau to northern Eurasia |journal=Frontiers in Ecology and Evolution |volume=12 |doi=10.3389/fevo.2024.1377000 |doi-access=free |bibcode=2024FrEEv..1277000M |issn=2296-701X}}) making it comparable in size to the largest living rhinoceros species, the white rhinoceros (Ceratotherium simum).{{cite book

| isbn = 9780306460920

| url = https://books.google.com/books?id=UZLuF1YXYTcC

| title = Extinctions in Near Time. Causes, Contexts, and Consequences

| publisher = Springer US

| date = June 30, 1999

| access-date = 17 September 2022

| pages = 262

| author = Hans-Dieter Sues, Ross D.E. MacPhee

}}{{Cite journal | last1 = Boeskorov | first1 = G. G. | title = Some specific morphological and ecological features of the fossil woolly rhinoceros (Coelodonta antiquitatis Blumenbach 1799) | doi = 10.1134/S106235901208002X | journal = Biology Bulletin | volume = 39 | issue = 8 | pages = 692–707 | year = 2012 | bibcode = 2012BioBu..39..692B | s2cid = 24868968 }} 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. The front horn would have measured {{convert|1|-|1.35|m|ft}} long for individuals at 25 to 35 years of age, while the second horn would have measured up to {{convert|47.5|cm|ft}} long. 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.{{Cite journal |last1=Shidlovskiy |first1=Fedor K. |last2=Kirillova |first2=Irina V. |last3=Wood |first3=John |date=2012-03-26 |title=Horns of the woolly rhinoceros Coelodonta antiquitatis (Blumenbach, 1799) in the Ice Age Museum collection (Moscow, Russia) |url=https://www.sciencedirect.com/science/article/pii/S1040618211003831 |journal=Quaternary International |series=Mammoths and Their Relatives 1: Biotopes, Evolution and Human Impact V International Conference, Le Puy-en-Velay, 2010 |volume=255 |pages=125–129 |doi=10.1016/j.quaint.2011.06.051 |bibcode=2012QuInt.255..125S |issn=1040-6182|url-access=subscription }} 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.{{cite journal | title=The earliest immigration of woolly rhinoceros (Coelodonta tologoijensis, Rhinocerotidae, Mammalia) into Europe and its adaptive evolution in Palaearctic cold stage mammal faunas | journal=Quaternary Science Reviews | volume=27 | issue=21–22 | date=2008-11-01 | issn=0277-3791 | doi=10.1016/j.quascirev.2008.07.013 | bibcode=2008QSRv...27.1951K |url=http://www.rhinoresourcecenter.com/pdf_files/125/1257355104.pdf| last1=Kahlke | first1=Ralf-Dietrich | last2=Lacombat | first2=Frédéric | pages=1951–1961 }}

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. The body's length ended with a {{convert|45|to|50|cm|in|adj=on}} tail with a brush of coarse hair at the end.Kalandadze N.N., Shapovalov A.V. & Tesakova E.M.— On nomenclatural problems concerning woolly rhinoceros Coelodonta antiquitatis (Blumenbach, 1799) // Researches on paleontology and biostratigraphy of ancient continental deposits (Memories of Professor Vitalii G. Ochev). Eds. M.A. Shishkin & V.P. Tverdokhlebov.— Saratov: «Nauchnaya Kniga» Publishers, 2009. P. 98–111. Females had two nipples on the udders.

The woolly rhinoceros had several features which reduced the body's surface area and minimized heat loss. Its ears were no longer than {{convert|24|cm|in|frac=2|abbr=on}}, while those of rhinos in hot climates are about {{convert|30|cm|in|frac=2|abbr=on}}. Their tails were also relatively shorter. It also had thick skin, ranging from {{convert|5|to|15|mm|in|abbr=on|frac=8}}, heaviest on the chest and shoulders.{{cite journal |last1=Stuart |first1=Anthony J. |last2=Lister |first2=Adrian M. |year=2012 |title=Extinction chronology of the woolly rhinoceros Coelodonta antiquitatis in the context of late Quaternary megafaunal extinctions in northern Eurasia |url=http://www.rhinoresourcecenter.com/pdf_files/135/1351170424.pdf |journal=Quaternary Science Reviews |volume=51 |pages=1–17 |bibcode=2012QSRv...51....1S |doi=10.1016/j.quascirev.2012.06.007 |issn=0277-3791}}

File:Coelodonta antiquitatis Crane.jpg

The skull had a length between {{convert|70|and|90|cm|in|round=5|abbr=on}}. 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. Strong muscles on its long occipital bone formed its neck hock and held the massive skull. Its massive lower jaw measured up to {{convert|60|cm|in|0|abbr=on}} long and {{convert|10|cm|in|0|abbr=on}} high. The teeth of the woolly rhinoceros had thickened enamel and an open internal cavity. Like other rhinos, adults did not have incisors. It had 3 premolars and 3 molars in both jaws. The molars were high-crowned and had a thick coat of cementum.

The nasal septum of the woolly rhinoceros was ossified, unlike modern rhinos. This was most common in adult males. This adaptation probably evolved as a result of the heavy pressure on the horn and face when the rhinoceros grazed underneath the thick snow. 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.{{cite journal | last1=Mazza | first1=Paul | last2=Azzaroli | first2=A. | title=Ethological inferences on Pleistocene rhinoceroses of Europe | journal=Rendiconti Lincei | publisher=Springer Science and Business Media LLC | volume=4 | issue=2 | year=1993 | issn=1120-6349 | doi=10.1007/bf03001424 | pages=127–137| s2cid=86966532 }} This ossification inspired the junior synonym specific name tichorhinus, from Greek τειχος (teikhos) "wall", ῥις (ῥιν-) (rhis (rhin-)) "nose".

File:Ice age fauna of northern Spain - Mauricio Antón.jpgs, 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.{{cite journal | last=Garutt | first=N.V. | title=Dental ontogeny of the woolly rhinoceros Coelodonta antiquitatis (Blumenbach, 1799) | journal=Cranium | volume=11 | issue=1 | url=http://www.rhinoresourcecenter.com/pdf_files/142/1428276199.pdf}} The two teats in the female suggest that she raised one calf, or more rarely two, every two to three years.{{cite web| title=Почему вымер шерстистый носоро| trans-title=Why the woolly rhino died out| website=All-Russian Festival of Science| date=2012-04-22| url=http://www.official.festivalnauki.ru/statya/4577/pochemu-vymer-sherstistyy-nosorog| language=ru| access-date=2019-11-07| archive-date=2014-05-21| archive-url=https://web.archive.org/web/20140521180459/http://www.official.festivalnauki.ru/statya/4577/pochemu-vymer-sherstistyy-nosorog| url-status=dead}}

With their massive horns and size, adults had few predators, but young individuals could have been attacked by cave hyenas and cave lions. A skull was found with trauma indicating an attack from a feline, but the animal survived to adulthood. Remains of woolly rhinoceros are frequently found in cave hyena dens with gnaw marks indicating that their remains were consumed by them,Diedrich, C.G. & ŽÁK, K. 2006. [http://www.geology.cz/bulletin/fulltext/bullgeosci200604237.pdf Prey deposits and den sites of the Upper Pleistocene hyena Crocuta crocuta spelaea (Goldfuss, 1823) in horizontal and vertical caves of the Bohemian Karst (Czech Republic)]. Bulletin of Geosciences 81(4), 237–276 (25 figures). Czech Geological Survey, Prague. ISSN 1214-1119. which to a large degree likely reflects scavenging of the carcasses of already dead rhinoceroses.{{Cite journal |last=Diedrich |first=C.G. |date=June 2012 |title=Late Pleistocene Crocuta crocuta spelaea (Goldfuss, 1823) clans as prezewalski horse hunters and woolly rhinoceros scavengers at the open air commuting den and contemporary Neanderthal camp site Westeregeln (central Germany) |url=https://linkinghub.elsevier.com/retrieve/pii/S0305440312000258 |journal=Journal of Archaeological Science |language=en |volume=39 |issue=6 |pages=1749–1767 |doi=10.1016/j.jas.2012.01.013|bibcode=2012JArSc..39.1749D |url-access=subscription }}

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. They may have also been used to attract mates. 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,

{{cite journal|last=Garutt|first=N.V.|title=Traumatic skull damages in the woolly rhinoceros, Coelodonta antiquitatis Blumenbach, 1799|journal=Cranium|year=1997|volume=14|issue=1|pages=37–46|url=http://www.rhinoresourcecenter.com/pdf_files/142/1428276474.pdf}}  and lower jaws and back ribs show signs of being broken and re-formed, which may have also come from fighting.{{cite journal|last=Diedrich|first=Cajus|date=2008|title=A skeleton of an injured Coelodonta antiquitatis (Blumenbach, 1807) from the Upper Pleistocene of north-western Germany|journal=Cranium|volume=25|issue=1|pages=1–16|url=http://www.rhinoresourcecenter.com/pdf_files/142/1428542866.pdf}} 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.{{cite thesis|last=Garutt|first=N.V.|date=1998|title=Woolly Rhinoceros: Morphology, Systematics and Geological Significance|type=Doctoral dissertation abstract|publisher=Russian Mining Institute|url=http://www.rhinoresourcecenter.com/pdf_files/139/1393195316.pdf}}

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.{{cite book |first1=Jan |last1=van der Made |first2=René |last2=Grube |title=The rhinoceroses from Neumark-Nord and their nutrition |editor-first=Harald |editor-last=Meller |publisher=Halle/Saale |year=2010 |pages=382–394 |language=de, en |url=http://www.rhinoresourcecenter.com/ref_files/1295058899.pdf}} It had a wide upper lip like that of the white rhinoceros, which allowed it to easily pluck vegetation directly from the ground. 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.{{cite journal |last1=Schreve |first1=Danielle |last2=Howard |first2=Andy |last3=Currant |first3=Andrew |last4=Brooks |first4=Stephen |last5=Buteux |first5=Simon |last6=Coope |first6=Russell |last7=Crocker |first7=Barnaby |last8=Field |first8=Michael |last9=Greenwood |first9=Malcolm |last10=Greig |first10=James |last11=Toms |first11=Phillip |date=2012-12-18 |title=A Middle Devensian woolly rhinoceros (Coelodonta antiquitatis) from Whitemoor Haye Quarry, Staffordshire (UK): palaeoenvironmental context and significance |url=http://www.rhinoresourcecenter.com/pdf_files/136/1360867011.pdf |journal=Journal of Quaternary Science |publisher=Wiley |volume=28 |issue=2 |pages=118–130 |doi=10.1002/jqs.2594 |issn=0267-8179 |s2cid=54874495}} 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.{{cite journal |vauthors=Willerslev E, Davison J, Moora M, Zobel M, Coissac E, Edwards ME, Lorenzen ED, Vestergård M, Gussarova G, Haile J, Craine J, Gielly L, Boessenkool S, Epp LS, Pearman PB, Cheddadi R, Murray D, Bråthen KA, Yoccoz N, Binney H, Cruaud C, Wincker P, Goslar T, Alsos IG, Bellemain E, Brysting AK, Elven R, Sønstebø JH, Murton J, Sher A, Rasmussen M, Rønn R, Mourier T, Cooper A, Austin J, Möller P, Froese D, Zazula G, Pompanon F, Rioux D, Niderkorn V, Tikhonov A, Savvinov G, Roberts RG, MacPhee RD, Gilbert MT, Kjær KH, Orlando L, Brochmann C, Taberlet P |year=2014 |title=Fifty thousand years of Arctic vegetation and megafaunal diet |url=https://eprints.soton.ac.uk/377215/3/Willerslev_Fig2.pdf |journal=Nature |volume=506 |issue=7486 |pages=47–51 |bibcode=2014Natur.506...47W |doi=10.1038/nature12921 |pmid=24499916 |s2cid=4461741}} Pollen analysis shows it also ate woody plants (including conifers, willows and alders), along with flowers,{{cite web | title=Woolly Mammoths and Rhinos Ate Flowers | website=livescience.com | date=2014-02-05 | url=https://www.livescience.com/43131-woolly-mammoths-ate-flowers.html | access-date=2019-11-07}} forbs and mosses. 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.Tiunov, Alexei & Kirillova, Irina. (2010). Stable isotope (C-13/C-12 and N-15/N-14) composition of the woolly rhinoceros Coelodonta antiquitatis horn suggests seasonal changes in the diet. Rapid communications in mass spectrometry : RCM. 24. 3146-50. 10.1002/rcm.4755. Dental mesowear measurements further show that the woolly rhinoceros's diet was heavily composed on abrasive grasses.{{Cite journal |last1=Diana |first1=Pushkina |last2=Juha |first2=Saarinen |last3=Reinhard |first3=Ziegler |last4=Hervé |first4=Bocherens |date=1 January 2020 |title=Stable isotopic and mesowear reconstructions of paleodiet and habitat of the Middle and Late Pleistocene mammals in south-western Germany |url=https://www.sciencedirect.com/science/article/pii/S0277379119301325 |journal=Quaternary Science Reviews |language=en |volume=227 |pages=106026 |doi=10.1016/j.quascirev.2019.106026 |bibcode=2020QSRv..22706026D |access-date=30 September 2024 |via=Elsevier Science Direct}}

It is estimated that woolly rhinoceroses could reach around 40 years of age, like their modern relatives. 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 {{convert|1.2|m|ft}} in length and {{convert|72|cm|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 {{convert|1.3|m|ft}}. At 7 to 10 years of age, woolly rhinos became young adults with a shoulder height of {{convert|1.4|-|1.5|m|ft}}. By more than 14 years of age, woolly rhinos became fully mature, old adults with a shoulder height of {{convert|1.6|m|ft}}.{{cite journal | last=Shpansky | first=A.V. | title=Juvenile remains of the "woolly rhinoceros" Coelodonta antiquitatis (Blumenbach 1799) (Mammalia, Rhinocerotidae) from the Tomsk Priob'e area (southeast Western Siberia) | journal=Quaternary International | publisher=Elsevier BV | volume=333 | year=2014 | issn=1040-6182 | doi=10.1016/j.quaint.2014.01.047 | bibcode=2014QuInt.333...86S | pages=86–99 | url=http://www.rhinoresourcecenter.com/pdf_files/140/1400554721.pdf}}

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.{{Cite journal |last=Diedrich |first=Cajus G. |date=1 February 2023 |title=Extinct Eurasian rhinoceros Coelodonta and Stephanorhinus dental pathologies and tooth change modus |url=https://linkinghub.elsevier.com/retrieve/pii/S0277379122005534 |journal=Quaternary Science Reviews |language=en |volume=301 |pages=107922 |bibcode=2023QSRv..30107922D |doi=10.1016/j.quascirev.2022.107922 |access-date=8 May 2024 |via=Elsevier Science Direct|url-access=subscription }}

File:Woolly rhino area.png

The woolly rhinoceros lived mainly in lowlands, plateaus and river valleys, with dry to arid climates, 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.{{cite journal | title=The origin of Eurasian Mammoth Faunas (Mammuthus–Coelodonta Faunal Complex) | journal=Quaternary Science Reviews | volume=96 | date=2014-07-15 | issn=0277-3791 | doi=10.1016/j.quascirev.2013.01.012 | bibcode=2014QSRv...96...32K | pages=32–49 | url=http://www.rhinoresourcecenter.com/pdf_files/140/1404659577.pdf| last1=Kahlke | first1=Ralf-Dietrich }} With its wide distribution, the woolly rhinoceros lived in some areas alongside the other rhinoceroses Stephanorhinus and Elasmotherium.{{cite journal | last=Pushkina | first=Diana | title=The Pleistocene easternmost distribution in Eurasia of the species associated with the Eemian Palaeoloxodon antiquus assemblage | journal=Mammal Review | publisher=Wiley | volume=37 | issue=3 | year=2007 | issn=0305-1838 | doi=10.1111/j.1365-2907.2007.00109.x | pages=224–245| bibcode=2007MamRv..37..224P }}

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.Garutt, N. V., & Boeskorov, G. G. (2001). Woolly rhinoceroses: On the history of the genus. Mamont i ego okruzhenie, 200, 157-167. Even during the very warm Eemian interglacial, the range of the woolly rhinoceros extended into temperate regions such as Poland.{{Cite journal |last1=Stefaniak |first1=Krzysztof |last2=Kovalchuk |first2=Oleksandr |last3=Marciszak |first3=Adrian |last4=Sobczyk |first4=Artur |last5=Socha |first5=Paweł |date=28 September 2023 |title=Environmental conditions across Poland during the Eemian Interglacial reconstructed from vertebrate remains |url=https://geojournals.pgi.gov.pl/agp/article/view/33877 |journal=Acta Geologica Polonica |language=en |volume=73 |issue=3 |pages=379–410 |doi=10.24425/agp.2023.145621 |access-date=17 September 2024|doi-access=free }} It had the widest range of any rhinoceros species.Prothero, D.R., Guérin, C. & Manning, E. 1989. The History of Rhinocerotoidea. In: The Evolution of Perissodactyls (eds. Prothero, D. R. & Schoch, R.M.). Oxford University Press, New York, 321-340.

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, 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.{{cite journal | last=Boeskorov | first=G. | title=Woolly rhino (Coelodonta antiquitatis) distribution in Northeast Asia | journal=Deinsea | volume=8 | issue=1 | url=http://www.rhinoresourcecenter.com/pdf_files/136/1360793475.pdf}}

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. 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.{{cite journal|last=Diedrich|first=Cajus|title=Eingeschleppte und benagte Knochenreste von Coelodonta antiquitatis (Blumenbach 1807) aus dem oberpleistozänen Fleckenhyänenhorst Perick-Höhlen im Nordsauerland (NW Deutschland) und Beitrag zur Taphonomie von Wollnashornknochen in Westfalen| journal=Mitteilungen der Höhlen und Karstforscher|year=2008|pages=100–117|language=de|url=http://www.rhinoresourcecenter.com/pdf_files/131/1312136882.pdf}} Sometimes, only individual teeth or bone fragments are uncovered, which usually came from only one animal.Bratlund, B. (2005). Comments on a cut-marked woolly rhino mandible from Zwolen. In: R. Schild (ed.), The killing fields of Zwolén. A Middle Paleolithic kill-butchery-site in Central Poland. Warsaw, Institute of Archaeology and Ethnology, Polish Academy of Sciences: 217-221. 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.{{cite journal|last1=Baumann|first1=Willfried|last2=Mania|first2=Dietrich|title=Ein mittelpaläolithisches Rentierlager bei Salzgitter-Lebenstedt.|trans-title=A Middle Paleolithic reindeer site near Salzgitter-Lebenstedt.|journal=Die paläolithischen Neufunde von Markkleeberg bei Leipzig (Mit Beiträgen von L. Eißmann und V. Toepfer). Veröffentlichungen des Landesmuseums für Vorgeschichte Dresden.|volume=Band 16|location=Berlin|year=1983|language=de}}{{cite journal|last=Guérin|first=Claude|title=Coelodonta antiquitatis praecursor (Rhinocerotidae) du Pléistocène moyen final de l'aven de Romain-la-Roche (Doubs, France)|trans-title=The Late Middle Pleistocene Coelodonta antiquitatis praecursor (Rhinocerotidae) from the sinkhole of Romain-la-Roche (Doubs, France)|year=2010|language=fr}}

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. A few sites from the early phase of the Last Glacial Period in the late Middle Paleolithic, such as the Gudenus Cave (Austria){{cite journal|last=Döppes|first=Doris|title=Die jungpleistozäne Säugetierfauna der Gudenushöhle (Niederösterreich).|trans-title=The Late Pleistocene mammal fauna of the Gudenus Cave (Lower Austria)|journal=Wissenschaftliche Mitteilungen des Niederösterreichischen Landesmuseums|trans-journal=Scientific communications of the Lower Austria Museum|volume=10|year=1997|pages=17–32|language=de}} and the open air site of Königsaue (Saxony-Anhalt, Germany),{{cite journal|last1=Mania|first1=Dietrich|last2=Toepfer|first2=V.|title=Königsaue. Gliederung, Ökologie und mittelpaläolithische Funde der letzten Eiszeit. Veröffentlichungen des Landesmuseums für Vorgeschichte Halle/Saale|volume=26|year=1973|language=de}} have heavily beaten rhinoceros bones lined with slash marks. This action was done partly to extract the nutritious bone marrow.{{cite book | last1=Ashton | first1=N. | last2=Lewis | first2=S. | last3=Stringer | first3=C. | title=The Ancient Human Occupation of Britain | publisher=Elsevier Science | series=ISSN | year=2010 | isbn=978-0-444-53598-6 | url=https://books.google.com/books?id=-ykpHxlzBf4C&pg=PA173}}

Both horns and bones of the rhinoceros were used as raw materials for tools and weapons, as were remains from other animals.Gaudzinski, S. 1999a. The faunal record of the Lower and Middle Palaeolithic of Europe: remarks on human interference. In The Middle Palaeolithic Occupation of Europe. (ed. W. Roebroeks and C. Gamble) Leiden: University of Leiden, pp. 215 - 233. In what is now Zwoleń, Poland, a device was made from a battered woolly rhinoceros pelvis.{{cite book | last=Schild | first=R. | title=The Killing Fields of Zwolén: A Middle Paleolithic Kill-butchery-site in Central Poland | publisher=Institute of Archaeology and Ethnology, Polish Academy of Sciences | year=2005 | isbn=978-83-89499-23-3 | pages=2017–216}} 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.{{cite journal | last1=Nikolskiy | first1=Pavel | last2=Pitulko | first2=Vladimir | title=Evidence from the Yana Palaeolithic site, Arctic Siberia, yields clues to the riddle of mammoth hunting | journal=Journal of Archaeological Science | publisher=Elsevier BV | volume=40 | issue=12 | year=2013 | issn=0305-4403 | doi=10.1016/j.jas.2013.05.020 | pages=4189–4197| bibcode=2013JArSc..40.4189N }} 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.{{cite news | title=13,300 year old spear made of woolly rhinoceros horn found on Arctic island | website=Siberian Times | url=https://siberiantimes.com/science/others/features/13300-year-old-spear-made-of-woolly-rhinoceros-horn-found-on-arctic-island/}}

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.

File:16 PanneauDesLions(CentreGauche)RhinocérosEnFuite.jpg]]

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.{{cite book | last1=Bahn | first1=P.G. | last2=Vertut | first2=J. | title=Journey Through the Ice Age | publisher=University of California Press | year=1997 | isbn=978-0-520-22900-6 | page=153}}

About 20 Paleolithic drawings of woolly rhinos were known before the discovery of the Chauvet Cave in France. They are dated at over 31,000 years old, probably from the Aurignacian, engraved on cave walls or drawn in red or black. One scene depicts two rhinos fighting each other with their horns.{{cite web | last=Foundation | first=Bradshaw | title=Fighting Rhino & Four Horses - The Cave Art Paintings of the Chauvet Cave | website=Bradshaw Foundation | url=http://www.bradshawfoundation.com/chauvet/fighting_rhino_four_horses.php | access-date=2019-10-28}} 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.{{cite web|title=Kapova Cave|website=Don's Maps|url=https://www.donsmaps.com/kapova.html|access-date=24 Aug 2019}} Some images show rhinoceroses struck with spears or arrows, signifying human hunting.

The site of Dolní Věstonice in Moravia, Czech Republic, was found with more than seven hundred statuettes of animals, many of woolly rhinoceroses.{{cite book | last=Guthrie | first=R.D. | title=The Nature of Paleolithic Art | publisher=University of Chicago Press | year=2005 | isbn=978-0-226-31126-5 | url=https://books.google.com/books?id=3u6JNwMyMCEC}}

{{Broader|Late Pleistocene extinctions}}

Analysis of the nuclear genome suggests that the woolly rhinoceros experienced a population expansion beginning around 30,000 years ago. 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 and 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. The woolly rhinoceros was likely intolerant of deep snow, which its short limbs were inefficient in moving through.{{Cite journal |last1=Fordham |first1=Damien A. |last2=Brown |first2=Stuart C. |last3=Canteri |first3=Elisabetta |last4=Austin |first4=Jeremy J. |last5=Lomolino |first5=Mark V. |last6=Haythorne |first6=Sean |last7=Armstrong |first7=Edward |last8=Bocherens |first8=Hervé |last9=Manica |first9=Andrea |last10=Rey-Iglesia |first10=Alba |last11=Rahbek |first11=Carsten |last12=Nogués-Bravo |first12=David |last13=Lorenzen |first13=Eline D. |date=2024-06-11 |title=52,000 years of woolly rhinoceros population dynamics reveal extinction mechanisms |journal=Proceedings of the National Academy of Sciences |language=en |volume=121 |issue=24 |pages=e2316419121 |doi=10.1073/pnas.2316419121 |pmid=38830089 |pmc=11181021 |bibcode=2024PNAS..12116419F |issn=0027-8424}} Population fragmentation is likely to have played a role in its extinction.{{Cite journal |last1=Rey-Iglesia |first1=Alba |last2=Lister |first2=Adrian M. |last3=Stuart |first3=Anthony J. |last4=Bocherens |first4=Hervé |last5=Szpak |first5=Paul |last6=Willerslev |first6=Eske |last7=Lorenzen |first7=Eline D. |date=July 2021 |title=Late Pleistocene paleoecology and phylogeography of woolly rhinoceroses |journal=Quaternary Science Reviews |language=en |volume=263 |pages=106993 |doi=10.1016/j.quascirev.2021.106993 |bibcode=2021QSRv..26306993R |doi-access=free}} The presence of large numbers of cervical ribs in specimens from the Netherlands may have been due to inbreeding or harsh environmental conditions.{{Cite journal |last1=van der Geer |first1=Alexandra A.E. |last2=Galis |first2=Frietson |date=29 August 2017 |title=High incidence of cervical ribs indicates vulnerable condition in Late Pleistocene woolly rhinoceroses |journal=PeerJ |language=en |volume=5 |pages=e3684 |doi=10.7717/peerj.3684 |doi-access=free |issn=2167-8359 |pmc=5580387 |pmid=28875067 }} 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.{{Cite journal |last1=Lord |first1=Edana |last2=Dussex |first2=Nicolas |last3=Kierczak |first3=Marcin |last4=Díez-del-Molino |first4=David |last5=Ryder |first5=Oliver A. |last6=Stanton |first6=David W.G. |last7=Gilbert |first7=M. Thomas P. |last8=Sánchez-Barreiro |first8=Fátima |last9=Zhang |first9=Guojie |last10=Sinding |first10=Mikkel-Holger S. |last11=Lorenzen |first11=Eline D. |last12=Willerslev |first12=Eske |last13=Protopopov |first13=Albert |last14=Shidlovskiy |first14=Fedor |last15=Fedorov |first15=Sergey |date=October 2020 |title=Pre-extinction Demographic Stability and Genomic Signatures of Adaptation in the Woolly Rhinoceros |url=https://linkinghub.elsevier.com/retrieve/pii/S096098222031071X |journal=Current Biology |language=en |volume=30 |issue=19 |pages=3871–3879.e7 |doi=10.1016/j.cub.2020.07.046|pmid=32795436 |bibcode=2020CBio...30E3871L |hdl=10037/20986 |s2cid=221114305 |hdl-access=free }} 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.{{Cite journal|last1=Wang|first1=Yucheng|last2=Pedersen|first2=Mikkel Winther|last3=Alsos|first3=Inger Greve|last4=De Sanctis|first4=Bianca|last5=Racimo|first5=Fernando|last6=Prohaska|first6=Ana|last7=Coissac|first7=Eric|last8=Owens|first8=Hannah Lois|last9=Merkel|first9=Marie Kristine Føreid|last10=Fernandez-Guerra|first10=Antonio|last11=Rouillard|first11=Alexandra|date=2021-10-20|title=Late Quaternary dynamics of Arctic biota from ancient environmental genomics|journal=Nature|volume=600|issue=7887|language=en|pages=86–92|doi=10.1038/s41586-021-04016-x|pmid=34671161|pmc=8636272|bibcode=2021Natur.600...86W|s2cid=239051880|issn=1476-4687}} 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,{{Cite journal |last1=Miller |first1=Joshua H. |last2=Simpson |first2=Carl |date=2022-12-01 |title=When did mammoths go extinct? |journal=Nature |language=en |volume=612 |issue=7938 |pages=E1–E3 |doi=10.1038/s41586-022-05416-3 |issn=0028-0836 |pmc=9712083 |pmid=36450914|bibcode=2022Natur.612E...1M }}{{Cite report |url=https://elifesciences.org/reviewed-preprints/89992v1 |title=Mitochondrial genomes of Pleistocene megafauna retrieved from recent sediment layers of two Siberian lakes |last1=Seeber |first1=Pa |last2=Batke |first2=L |date=2023-09-01 |publisher=elife |doi=10.7554/elife.89992.1 |last3=Dvornikov |first3=Y |last4=Schmidt |first4=A |last5=Wang |first5=Y |last6=Stoof-Leichsenring |first6=Kr |last7=Moon |first7=Kl |last8=Shapiro |first8=B |last9=Epp |first9=Ls |doi-access=free }} though other authors have argued that this specific environmental DNA record is unlikely to have been reworked. Low level human hunting may have played a decisive role in the extinction by reducing the ability of woolly rhinoceros populations to colonise newly suitable habitat, thereby exacerbating the population fragmentation brought on by environmental change.

File:Frozen Coelodonta.jpg

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.{{cite web | title=Three Centuries of Hunting for Ice Age Mummies and the Prospect of De-Extinction | website=Capeia | date=2018-05-18 | url=https://beta.capeia.com/paleobiology/2018/05/18/three-centuries-of-hunting-for-ice-age-mummies-and-the-prospect-of-de-extinction | access-date=2019-11-02 | archive-date=2019-11-07 | archive-url=https://web.archive.org/web/20191107171523/https://beta.capeia.com/paleobiology/2018/05/18/three-centuries-of-hunting-for-ice-age-mummies-and-the-prospect-of-de-extinction | url-status=dead }} Later in 1877, a Siberian trader recovered a head and one leg from a tributary of the Yana River.

In October 1907, miners in Starunia, Russian Empire, found a mammoth carcass buried in an ozokerite pit. A month later, a rhinoceros was found {{convert|5|m|ft|round=5}} underneath.{{cite journal|last1=Kotarba|first1=Maciej J.|last2=Dzieniewicz|first2=Marek|last3=Móscicki|first3=Wlodzimierz J.|last4=Sechman|first4=Henryk|title=Unique Quaternary environment for discoveries of woolly rhinoceroses in Starunia, fore-Carpathian region, Ukraine: Geochemical and geoelectric studies|journal=Geology|volume=36|issue=7|date=2008-07-01|issn=0091-7613|doi=10.1130/G24654A.1|bibcode=2008Geo....36..567K|pages=567–570}} Both were sent to the Dzieduszycki Museum, where a detailed description was published in the museum's monograph.Bayger, J.A., Hoyer, H., Kiernik, E., Kulczyński, W., Łomnicki, M., Łomnicki J., Mierzejewski, W., Niezabitowski, E., Raciborski, W., Szafer, W. & Schille, F., 1914. Wykopaliska staruńskie. Słoń mamut (Elephas primigenius Blum.) i nosorożec włochaty (Rhinoceros antiquitatis Blum. s. tichorhinus Fisch.) wraz z współczesną florą i fauną. Muzeum im. Dzieduszyckich we Lwowie, 15, 386 pp + atlas (67 tab.). (In Polish). Photographs were published in paleontological journals and textbooks, and the first modern paintings of the species were based on the mounted specimen. The rhino is now located in the Lviv National Museum along with the mammoth.{{cite journal|last1=Chornobay|first1=Yuriy M.|last2=Drygant|first2=Daniel M.|title=The Starunia collections in the Natural History Museum of the National Academy of Sciences of Ukraine in Lviv|journal= Geotourism/Geoturystyka|volume=3|issue=18|year=2009|issn=1731-0830|doi=10.7494/geotour.2009.18.3.45|pages=45–50|url=http://www.rhinoresourcecenter.com/pdf_files/142/1422710219.pdf|doi-access=free}} Later, in 1929, the Polish Academy of Arts and Sciences sent an expedition to Starunia, finding the mummified remains of three rhinos.{{cite journal | last=Kucha | first=Henryk | title=The Starunia collections in the Institute of Systematics and Evolution of Animals, Polish Academy of Sciences in Kraków | journal=Geotourism/Geoturystyka | publisher=AGHU University of Science and Technology Press | volume=18 | issue=1 | year=2009 | issn=1731-0830 | doi=10.7494/geotour.2009.18.71 | page=71| s2cid=128734734 | doi-access=free }} 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.Nowak, J., Panow, E., Tokarski, J., Szafer, W. & Stach, J. 1930. The second woolly rhinoceros (Coelodonta antiquitatis Blum.) from Starunia, Poland (Geology, Mineralogy, Flora and Fauna). Classe des Sciences Mathématiques et Naturelles, Série B: Sciences Naturelles, Supplément 1-47.

File:Wooly Rhino at the Natural History Museum.jpg

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.Lazarev, P.A., Boeskorov, G.G., Tomskaya, A.I., Garutt, N.V., Vasil’ev, E.M., and Kasparov, A.K., Mlekopitayushchie antropogena Yakutii (Mammals of the Anthropogene in Yakutia), Yakutsk: Yakut. Nauch. Tsentr Akad. Nauk SSSR, 1998. 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.{{cite news | title=Woolly Rhino Fossils Found | website=The New York Times | date=1976-10-10 | url=https://www.nytimes.com/1976/10/10/archives/woolly-rhino-fossils-found.html}}

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.{{cite journal | last1=Boeskorov | first1=G. G. | last2=Lazarev | first2=P. A. | last3=Bakulina | first3=N. T. | last4=Shchelchkova | first4=M. V. | last5=Davydov | first5=S. P. | last6=Solomonov | first6=N. G. | title=Preliminary study of a mummified woolly rhinoceros from the lower reaches of the Kolyma River | journal=Doklady Biological Sciences | volume=424 | issue=1 | year=2009 | issn=0012-4966 | doi=10.1134/s0012496609010165 | pmid=19341085 | pages=53–56 | s2cid=2674199 | url=http://www.rhinoresourcecenter.com/pdf_files/129/1297725357.pdf?view}} Next year in 2008, a nearly complete skeleton came from the Chukochya River.{{cite journal | last1=Kirillova | first1=Irina V. | last2=Shidlovskiy | first2=Fedor K. | title=Estimation of individual age and season of death in woolly rhinoceros, Coelodonta antiquitatis (Blumenbach, 1799), from Sakha-Yakutia, Russia | journal=Quaternary Science Reviews | volume=29 | issue=23–24 | date=2010-11-01 | issn=0277-3791 | doi=10.1016/j.quascirev.2010.06.036 | bibcode=2010QSRv...29.3106K | pages=3106–3114}} 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.Lazarev, P.A., Grigoriev, S.E., Plotnikov, V.V., 2010. Woolly rhinoceroses from Yakutia//evolution of life on the Earth. In: Proceedings of the IV International

Symposium. TML-Press, Tomsk, pp. 555e558.

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.{{cite news | last=Liesowska | first=Anna | title=Meet Sasha - the world's only baby woolly rhino| newspaper= The Siberian Times| url=https://siberiantimes.com/science/casestudy/news/n0129-meet-sasha-the-worlds-only-baby-woolly-rhino/}} Dental analysis shows that the calf was about seven months old at the time of its death.{{cite news | last=Liesowska | first=Anna | title=Lifelike again after 34,000 years, the world's only baby woolly rhino | website=Siberian Times | url=https://siberiantimes.com/other/others/features/lifelike-again-after-34000-years-the-worlds-only-baby-woolly-rhino/}} With its well-intact preservation, scientists proceeded to undergo DNA analysis.{{cite news | last=Liesowska | first=Anna | title=Sasha, the world's only baby woolly rhino, is 34,000 years old, say scientists | website=Siberian Times | date=2019-12-23 | url=https://siberiantimes.com/home/born-in-siberia/sasha-the-worlds-only-baby-woolly-rhino-is-34000-years-old-say-scientists/}}{{cite journal | last1=Chernova | first1=O. F. | last2=Protopopov | first2=A. V. | last3=Perfilova | first3=T. V. | last4=Kirillova | first4=I. V. | last5=Boeskorov | first5=G. G. | title=Hair microstructure of the first time found calf of woolly rhinoceros Coelodonta antiquitatis | journal=Doklady Biological Sciences | volume=471 | issue=1 | year=2016 | issn=0012-4966 | doi=10.1134/s0012496616060090 | pmid=28058607 | pages=291–295 | s2cid=18015793 | url=http://www.rhinoresourcecenter.com/pdf_files/148/1487482154.pdf}}

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.{{cite web|url=https://www.bbc.co.uk/news/world-europe-55490037 |title=Woolly rhino from Ice Age unearthed in Russian Arctic |date=30 December 2020 |work=BBC News |access-date=31 December 2020}}

• Elasmotherium, another Pleistocene Eurasian rhinoceros
• Narrow-nosed rhinoceros, temperate adapted rhinoceros species native to Europe, the Middle East and North Africa during Middle-Late Pleistocene
• Stephanorhinus kirchbergensis also known as Merck's rhinoceros, temperate adapted rhinoceros species native to Europe and Asia during Middle-Late Pleistocene

{{Portal bar|Paleontology}}

{{Reflist|2|refs=

{{cite book | url = https://books.google.com/books?id=8ikKezOKRB4C | title = Frozen Fauna of the Mammoth Steppe | isbn = 9780226311234 | last1 = Dale Guthrie | first1 = R. | year = 1990 | publisher = University of Chicago Press }}

{{cite journal | title= The morphology and paleobiological significance of the horns of Coelodonta antiquitatis (Mammalia: Rhinocerotidae) | last1= Fortelius | first1= Mikael | journal= Journal of Vertebrate Paleontology | volume= 3 | issue= 2 | year= 1983 | pages= 125–135 | issn = 0272-4634 | doi = 10.1080/02724634.1983.10011964 | bibcode= 1983JVPal...3..125F }}

{{cite web | url = https://www.britishmuseum.org/research/search_the_collection_database/search_object_details.aspx?objectId=1528116&partId=1 | title = engraved bone/antler | publisher = British Museum }}

{{cite journal | title=Woolly rhino discovery in the lower Kolyma River | journal=Quaternary Science Reviews | volume=30 | issue=17–18 | date=2011-08-01 | issn=0277-3791 | doi=10.1016/j.quascirev.2011.02.010 | bibcode=2011QSRv...30.2262B | pages=2262–2272 | url=http://www.rhinoresourcecenter.com/pdf_files/131/1313208126.pdf | last1=Boeskorov | first1=Gennady G. | last2=Lazarev | first2=Peter A. | last3=Sher | first3=Andrei V. | last4=Davydov | first4=Sergei P. | last5=Bakulina | first5=Nadezhda T. | last6=Shchelchkova | first6=Marina V. | last7=Binladen | first7=Jonas | last8=Willerslev | first8=Eske | last9=Buigues | first9=Bernard | last10=Tikhonov | first10=Alexey N. }}

{{cite journal | last1=Shidlovskiy | first1=Fedor K. | last2=Kirillova | first2=Irina V. | last3=Wood | first3=John | title=Horns of the woolly rhinoceros Coelodonta antiquitatis (Blumenbach, 1799) in the Ice Age Museum collection (Moscow, Russia) | journal=Quaternary International | publisher=Elsevier BV | volume=255 | year=2012 | issn=1040-6182 | doi=10.1016/j.quaint.2011.06.051 | bibcode=2012QuInt.255..125S | pages=125–129 | url=http://www.rhinoresourcecenter.com/pdf_files/131/1313208377.pdf}}

}}

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

{{Commons category|Coelodonta antiquitatis}}

{{Wikispecies|Coelodonta antiquitatis}}

• [https://web.archive.org/web/20160314031316/http://www.zwoje-scrolls.com/zwoje40/text19p.htm More pictures of the fully preserved tar pit wholly rhinoceros that was found in Poland (text in Polish)]
• [https://web.archive.org/web/20080807172800/http://worldmuseumofman.org/woollyrhinocerosf3.htm Fossil skull of a woolly rhinoceros from Belgium]
• [https://web.archive.org/web/20080517111716/http://worldmuseumofman.org/woollyrhinocerosf4.htm Fossil skull of a woolly rhinoceros from Germany]
• [https://web.archive.org/web/20111111011208/http://www.rhinos-irf.org/woolly/ International Rhino Foundation: Woolly Rhino]

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Category:Pleistocene extinctions

Category:Prehistoric mammals of Europe

Category:Pliocene rhinoceroses

Category:Piacenzian first appearances

Category:Pleistocene rhinoceroses

Category:Fossil taxa described in 1831

Category:Fossil taxa described in 1799

Category:Taxa named by Johann Friedrich Blumenbach

Category:Coelodonta

Category:Species that are or were threatened by habitat loss