The woolly rhinoceros (Coelodonta antiquitatis), simply known as woolly rhino, is an extinct species of
rhinoceros that inhabited northern Eurasia during the
Pleistoceneepoch. The woolly rhinoceros was a member of the
Pleistocene megafauna. The woolly rhinoceros was covered with long, thick hair that allowed it to survive in the extremely cold, harsh
mammoth steppe. It had a massive hump reaching from its shoulder and fed mainly on
herbaceous plants that grew in the steppe. Mummified carcasses preserved in
permafrost and many bone remains of woolly rhinoceroses have been found. Images of woolly rhinoceroses are found among
cave paintings in Europe and Asia. The species range contracted towards Siberia beginning around 17,000 years ago, with the youngest known records being around 14,000 years old in northeast Siberia, coinciding with the
Bølling–Allerød warming, which likely disrupted its habitat.
Taxonomy
Woolly rhinoceros remains have been known long before the species was described and were the basis for some mythical creatures. Native peoples of Siberia believed their horns were the claws of giant birds.[3] A rhinoceros skull was found in
Klagenfurt, Austria, in 1335, and was believed to be that of a
dragon.[4] In 1590, it was used as the basis for the head on a statue of a
lindworm.[5]Gotthilf Heinrich von Schubert maintained the belief that the horns were the claws of giant birds, and classified the animal under the name Gryphus antiquitatis, meaning "
griffin of antiquity".[6]
One of the earliest scientific descriptions of an ancient rhinoceros species was made in 1769, when the naturalist
Peter Simon Pallas wrote a report on his expeditions to Siberia where he found a skull and two horns in the permafrost.[7] In 1772, Pallas acquired a head and two legs of a rhinoceros from the locals in
Irkutsk,[8] and named the species Rhinoceros lenenesis (after the
Lena River).[9] In 1799,
Johann Friedrich Blumenbach studied rhinoceros bones from the collection of the
University of Göttingen, and proposed the scientific name Rhinoceros antiquitatis.[10] The geologist
Heinrich Georg Bronn moved the species to Coelodonta in 1831 because of its differences in dental formation with members of the Rhinoceros genus.[11] This name comes from the Greek words κοιλος (koilos, "hollow") and ὀδούς (odoús "tooth"), from the depression in the rhino's
molar structure,[12][13] giving the scientific name Coelodonta antiquitatis, "hollow-tooth of antiquity".[14]
Evolution
The woolly rhinoceros was the most recent species of the genus Coelodonta. The closest living relative of Coelodonta is the
Sumatran rhinoceros, and the genus is also closely related to the extinct genus Stephanorhinus. A cladogram showing the relationships of C. antiquitatis to other Late Pleistocene-recent rhinoceros species based on genomic data is given below.[15]
The oldest known species of Coelodonta,
Coelodonta thibetana is known from the
Pliocene of Tibet dating to approximately 3.7 million years ago,[17] 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.[18][19] The woolly rhinoceros is divided into two
chrono-subspecies, with C. a. praecursor from the middle Pleistocene and C. a. antiquitatis from the late Pleistocene.[1] Mitochondrial genomes suggest that the last mitochondrial ancestor of Late Pleistocene woolly rhinoceroses lived around 570,000 years ago.[20]
Description
Structure and appearance
An adult woolly rhinoceros typically measured 3.2 to 3.6 metres (10.5 to 11.8 ft) from head to tail, stood 1.45–1.6 metres (4.8–5.2 ft) tall at the shoulder, and weighed up to 1.5–2 metric tons (1.7–2.2 short tons).[21][22] Both males and females had two horns which were made of
keratin, with one long horn reaching forward and a smaller horn between the eyes.[23][24] The front horn would have measured 1–1.35 metres (3.3–4.4 ft) long for individuals at 25 to 35 years of age, while the second horn would have measured up to 47.5 centimetres (1.56 ft) long.[24] Compared to other rhinoceroses, the woolly rhinoceros had a longer head and body, and shorter legs. Its shoulder was raised with a powerful hump, used to support the animal's massive front horn. The hump also contained a fat reserve to aid survival through the desolate winters of the mammoth steppe.[25]
Frozen specimens indicate that the rhino's long fur coat was reddish-brown, with a thick
undercoat that lay under a layer of long, coarse
guard hair thickest on the withers and neck. Shorter hair covered the limbs, keeping snow from attaching.[25] The body's length ended with a 45-to-50-centimetre (18 to 20 in) tail with a brush of coarse hair at the end.[26] Females had two nipples on the
udders.[22]
The woolly rhinoceros had several features which reduced the body's surface area and minimized heat loss. Its ears were no longer than 24 cm (9+1⁄2 in), while those of rhinos in hot climates are about 30 cm (12 in).[24] Their tails were also relatively shorter. It also had thick skin, ranging from 5 to 15 mm (1⁄4 to 5⁄8 in), heaviest on the chest and shoulders.[27][24]
Skull and dentition
The skull had a length between 70 and 90 cm (30 and 35 in). It was longer than those of other rhinoceros, giving the head a deep, downward-facing slanting position, similar to its fossil relative Stephanorhinus hemitoechus and Elasmotherium as well as the white rhinoceros.[28] Strong muscles on its long
occipital bone formed its neck hock and held the massive skull. Its massive lower jaw measured up to 60 cm (24 in) long and 10 cm (4 in) high.[24] The teeth of the woolly rhinoceros had thickened enamel and an open internal cavity.[24] Like other rhinos, adults did not have
incisors.[29] It had 3
premolars and 3
molars in both jaws. The molars were high-crowned and had a thick coat of
cementum.[12]
The nasal septum of the woolly rhinoceros was
ossified, unlike modern rhinos. This was most common in adult males.[30] This adaptation probably evolved as a result of the heavy pressure on the horn and face when the rhinoceros grazed underneath the thick snow.[31] 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.[32] This ossification inspired the
junior synonymspecific nametichorhinus, from Greek τειχος (teikhos) "wall", ῥις (ῥιν-) (rhis (rhin-)) "nose".
Paleobiology
The woolly rhinoceros had a similar life history to modern rhinos. Studies on milk teeth show that individuals developed similarly to both the white and
black rhinoceros.[29] The two teats in the female suggest that she raised one calf, or more rarely two, every two to three years.[33][22]
With their massive horns and size, adults had few predators, but young individuals could be attacked by animals such as
hyenas and
cave lions. A skull was found with trauma indicating an attack from a feline, but the animal survived to adulthood.[34]
Woolly rhinos may have used their horns for combat, probably including
intraspecific combat as recorded in cave paintings, as well as for moving snow to uncover vegetation during winter.[27] They may have also been used to attract mates.[30] Bull woolly rhinos were probably territorial like their modern counterparts, defending themselves from competitors, particularly during the
rutting season. Fossil skulls indicate damage from the front horns of other rhinos,[34] and lower jaws and back ribs show signs of being broken and re-formed, which may have also come from fighting.[35] The apparent frequency of intraspecific combat, compared to recent rhinos, was likely a result of rapid climatic change during the last glacial period, when the animal faced increased stress from competition with other large herbivores.[31]
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.[24][28] It had a wide upper lip like that of the white rhinoceros, which allowed it to easily pluck vegetation directly from the ground.[22]Pollen analysis shows it also ate woody plants (including
conifers,
willows and
alders),[24] along with flowers,[36]forbs and
mosses.[37] Isotope studies on horns show that the woolly rhinoceros had a seasonal diet; different areas of horn growth suggest that it mainly
grazed in summer, while it
browsed for shrubs and branches in the winter.[38]
A strain vector
biomechanical investigation of the
skull,
mandible and
teeth of a well-preserved last cold stage individual recovered from
Whitemoor Haye,
Staffordshire, revealed
musculature and dental characteristics that support a grazing feeding preference. In particular, the enlargement of the
temporalis and neck muscles is consistent with that required to resist the large tugging forces generated when taking large mouthfuls of
fodder from the ground. The presence of a large
diastema supports this theory.[39]
Comparisons with living
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.[40]
Growth
It is estimated that woolly rhinoceroses could reach around 40 years of age, like their modern relatives.[34] In 2014, Shpansky analysed the growth of woolly rhinoceros from its early life stages based on several lower jaw fragments and limb bones. A one-month-old calf was about 1.2 metres (3.9 ft) in length and 72 centimetres (2.36 ft) tall at the shoulder. The most intensive growth in woolly rhinos occurred during the juvenile stage around 3 to 4 years of age with a shoulder height of 1.3 metres (4.3 ft). At 7 to 10 years of age, woolly rhinos became young adults with a shoulder height of 1.4–1.5 metres (4.6–4.9 ft). By more than 14 years of age, woolly rhinos became fully mature, old adults with a shoulder height of 1.6 metres (5.2 ft).[41]
Habitat and distribution
The woolly rhinoceros lived mainly in
lowlands,
plateaus and river valleys, with dry to arid climates,[42] and migrated to higher elevations in favourable climate phases. It avoided mountain ranges, due to heavy snow and steep terrain that the animal could not easily cross.[42] The rhino's main habitat was the mammoth steppe, a large, open landscape covered with wide ranges of grass and bushes. The woolly rhinoceros lived alongside other large herbivores, such as the
woolly mammoth,
giant deer,
reindeer,
saiga antelope and
bison – an assortment of animals known as the Mammuthus-Coelodonta Faunal Complex.[43] With its wide distribution, the woolly rhinoceros lived in some areas alongside the other rhinoceroses Stephanorhinus[25] and Elasmotherium.[44]
By the end of the
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.[27][45] It had the widest range of any rhinoceros species.[46]
It seemingly did not cross the
Bering land bridge during the last ice age (which connected Asia to North America), with its easterly-most occurrence at the
Chukotka Peninsula,[27] probably due to the low grass density and lack of suitable habitat in the
Yukon combined with competition from other large herbivores on the frigid land bridge.[42]
Relationship with humans
Hunting
Woolly rhinoceroses shared their habitat with humans, but direct evidence that they interacted is relatively rare. Only 11% of the known sites of prehistoric Siberian tribes have remains or images of the animal.[27] Many rhinoceros remains are found in caves (such as the
Kůlna Cave in Central Europe), which were not the natural habitat of either rhinos or humans, and large predators such as hyenas may have carried rhinoceros parts there.[47] Sometimes, only individual teeth or bone fragments are uncovered, which usually came from only one animal.[48] Most rhinoceros remains in Western Europe are found in the same places where human remains or artifacts were found, but this may have occurred naturally.[49][50]
Signs that early humans hunted or scavenged the rhinoceros come from markings on the animal's bones. One specimen had injuries caused by human weaponry, with traces of a wound from a sharp object marking the shoulder and thigh, and a preserved
spear was found near the carcass.[24] A few sites from the early phase of the
Last Glacial Period in the late
Middle Paleolithic, such as the
Gudenus Cave (Austria)[51] and the open air site of
Königsaue (
Saxony-Anhalt, Germany),[52] have heavily beaten rhinoceros bones lined with slash marks. This action was done partly to extract the nutritious
bone marrow.[53]
Both horns and bones of the rhinoceros were used as raw materials for tools and weapons, as were remains from other animals.[54] In what is now
Zwoleń, Poland, a device was made from a battered woolly rhinoceros pelvis.[55] 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.[56] A 13,300-year-old spear found on
Bolshoy Lyakhovsky Island has a tip made of rhinoceros horn, the furthest north a human artifact has ever been found.[57]
The
Pinhole Cave Man is a late Paleolithic figure of a man engraved on a rib bone of a woolly rhinoceros, found at
Creswell Crags in England.[58]
Ancient art
Many cave paintings from the
Upper Paleolithic depict woolly rhinoceroses. The animal's defining features are prominently drawn, complete with the raised back and hump, contrasting with its low-lying head. Two curved lines represent the ears. The animal's horns are drawn with their long curvature, and in some cases, the coat is also indicated. Many paintings show a black band dividing the body.[59]
About 20 Paleolithic drawings of woolly rhinos were known before the discovery of the
Chauvet Cave in France.[59] They are dated at over 31,000 years old, probably from the
Aurignacian,[27]engraved on cave walls or drawn in red or black. One scene depicts two rhinos fighting each other with their horns.[60] 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.[61] Some images show rhinoceroses struck with spears or arrows, signifying human hunting.[62]
The site of
Dolní Věstonice in
Moravia, Czech Republic, was found with more than seven hundred
statuettes of animals, many of woolly rhinoceroses.[62]
Analysis of the nuclear genome suggests that the woolly rhinoceros experienced a population expansion beginning around 30,000 years ago.[63] The end of the last glacial period shows a progressive contraction of the range of the woolly rhinoceros, with the species disappearing from Europe during the interval between 17-15,000 years ago, with its youngest confirmed records being from the
Urals, dating to 14,200 years ago, and northeast Siberia, dating to around 14,000 years ago. The youngest records of the species coincide with the onset of the
Bølling–Allerød warming, which likely resulted in increased
precipitation (including snowfall), which transformed the woolly rhinoceros' preferred low-growing grass and herb habitat into one dominated by shrubs and trees.[27]Population fragmentation is likely to have played a role in its extinction,[64] as evidenced by the incidences of genetic deformities such as unusually high occurrences of cervical ribs.[65] A genetic study of the woolly rhinoceros remains in northeast Siberia, dating to around 18,500 years ago, a few thousand years before its extinction, found that the population size was stable and relatively large, despite long-term co-existence with humans in the region.[63] A
Holocene survival of the species has been suggested by the finding of
environmental DNA of the woolly rhinoceros in sediments dating to 9,800 ± 200 years ago.[66] 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.[67][68]
Fossil specimens
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.[69] Later in 1877, a Siberian trader recovered a head and one leg from a tributary of the
Yana River.[24]
In October 1907, miners in
Starunia, Russian Empire, found a mammoth carcass buried in an
ozokerite pit. A month later, a rhinoceros was found 5 metres (15 ft) underneath.[70] Both were sent to the
Dzieduszycki Museum, where a detailed description was published in the museum's
monograph.[71] Photographs were published in paleontological journals and textbooks, and the first modern paintings of the species were based on the mounted specimen.[72] The rhino is now located in the
Lviv National Museum along with the mammoth.[72] Later, in 1929, the
Polish Academy of Arts and Sciences sent an expedition to Starunia, finding the mummified remains of three rhinos.[73] 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.[74]
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.[24][75] 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.[76]
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.[77][24] Next year in 2008, a nearly complete skeleton came from the
Chukochya River.[78] 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.[9]
In September 2014, a mummified young rhinoceros was discovered by two hunters, Alexander “Sasha” Banderov and Simeon Ivanov, at a tributary of the Semyulyakh River in the
Abyysky District in
Yakutia, Russia. Its head and horns, fur, and soft tissues were recovered. Some parts had been thawed and eaten since they were not covered by permafrost. The body was handed over to the Yakutia Academy of Sciences, where it was named “Sasha” after one of its discoverers.[79] Dental analysis shows that the calf was about seven months old at the time of its death.[80] With its well-intact preservation, scientists proceeded to undergo DNA analysis.[81][82]
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.[83]
See also
Elasmotherium, another Pleistocene Eurasian rhinoceros
^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.
^Pallas, P.S.; Blagdon, F.W. (1812). Travels Through the Southern Provinces of the Russian Empire: In the Years 1793 and 1794. Travels Through the Southern Provinces of the Russian Empire, in the Years 1793 and 1794. John Stockdale, Piccadilly.
^Owen, Richard. (1846). A History of British Fossil Mammals, and Birds. J. Van Voorst.
^
abLazarev, 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.
^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).
^von Koenigswald, W. (2002). Lebendige Eiszeit: Klima und Tierwelt im Wandel [Living Ice Age: Climate and wildlife in transition] (in German). Theiss.
ISBN978-3-8062-1734-6.
^
abcdBoeskorov, G. G. (2012). "Some specific morphological and ecological features of the fossil woolly rhinoceros (Coelodonta antiquitatis Blumenbach 1799)". Biology Bulletin. 39 (8): 692–707.
doi:
10.1134/S106235901208002X.
S2CID24868968.
^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.
^Mazza, Paul; Azzaroli, A. (1993). "Ethological inferences on Pleistocene rhinoceroses of Europe". Rendiconti Lincei. 4 (2). Springer Science and Business Media LLC: 127–137.
doi:
10.1007/bf03001424.
ISSN1120-6349.
S2CID86966532.
^"Почему вымер шерстистый носоро" [Why the woolly rhino died out]. All-Russian Festival of Science (in Russian). 2012-04-22. Archived from
the original on 2014-05-21. Retrieved 2019-11-07.
^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.
^Pushkina, Diana (2007). "The Pleistocene easternmost distribution in Eurasia of the species associated with the Eemian Palaeoloxodon antiquus assemblage". Mammal Review. 37 (3). Wiley: 224–245.
doi:
10.1111/j.1365-2907.2007.00109.x.
ISSN0305-1838.
^Garutt, N. V., & Boeskorov, G. G. (2001). Woolly rhinoceroses: On the history of the genus. Mamont i ego okruzhenie, 200, 157-167.
^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.
^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.
^Baumann, Willfried; Mania, Dietrich (1983). "Ein mittelpaläolithisches Rentierlager bei Salzgitter-Lebenstedt" [A Middle Paleolithic reindeer site near Salzgitter-Lebenstedt.]. Die paläolithischen Neufunde von Markkleeberg bei Leipzig (Mit Beiträgen von L. Eißmann und V. Toepfer). Veröffentlichungen des Landesmuseums für Vorgeschichte Dresden. (in German). Band 16. Berlin.
^Guérin, Claude (2010). "Coelodonta antiquitatis praecursor (Rhinocerotidae) du Pléistocène moyen final de l'aven de Romain-la-Roche (Doubs, France)" [The Late Middle Pleistocene Coelodonta antiquitatis praecursor (Rhinocerotidae) from the sinkhole of Romain-la-Roche (Doubs, France)] (in French). {{
cite journal}}: Cite journal requires |journal= (
help)
^Döppes, Doris (1997). "Die jungpleistozäne Säugetierfauna der Gudenushöhle (Niederösterreich)" [The Late Pleistocene mammal fauna of the Gudenus Cave (Lower Austria)]. Wissenschaftliche Mitteilungen des Niederösterreichischen Landesmuseums [Scientific communications of the Lower Austria Museum] (in German). 10: 17–32.
^Mania, Dietrich; Toepfer, V. (1973). "Königsaue. Gliederung, Ökologie und mittelpaläolithische Funde der letzten Eiszeit. Veröffentlichungen des Landesmuseums für Vorgeschichte Halle/Saale" (in German). 26. {{
cite journal}}: Cite journal requires |journal= (
help)
^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.
^Schild, R. (2005). The Killing Fields of Zwolén: A Middle Paleolithic Kill-butchery-site in Central Poland. Institute of Archaeology and Ethnology, Polish Academy of Sciences. pp. 2017–216.
ISBN978-83-89499-23-3.
^Nikolskiy, Pavel; Pitulko, Vladimir (2013). "Evidence from the Yana Palaeolithic site, Arctic Siberia, yields clues to the riddle of mammoth hunting". Journal of Archaeological Science. 40 (12). Elsevier BV: 4189–4197.
Bibcode:
2013JArSc..40.4189N.
doi:
10.1016/j.jas.2013.05.020.
ISSN0305-4403.
^Kotarba, Maciej J.; Dzieniewicz, Marek; Móscicki, Wlodzimierz J.; Sechman, Henryk (2008-07-01). "Unique Quaternary environment for discoveries of woolly rhinoceroses in Starunia, fore-Carpathian region, Ukraine: Geochemical and geoelectric studies". Geology. 36 (7): 567–570.
Bibcode:
2008Geo....36..567K.
doi:
10.1130/G24654A.1.
ISSN0091-7613.
^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).
^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.
^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.
^Kirillova, Irina V.; Shidlovskiy, Fedor K. (2010-11-01). "Estimation of individual age and season of death in woolly rhinoceros, Coelodonta antiquitatis (Blumenbach, 1799), from Sakha-Yakutia, Russia". Quaternary Science Reviews. 29 (23–24): 3106–3114.
Bibcode:
2010QSRv...29.3106K.
doi:
10.1016/j.quascirev.2010.06.036.
ISSN0277-3791.