Woolly rhinoceros

Last updated

Woolly rhinoceros
Temporal range: Middle Pleistocene–Late Pleistocene
Ꞓ
O
S
D
C
P
T
J
K
Pg
N
↓
Coelodonta antiquitatis .jpg
Woolly rhinoceros skeleton
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Perissodactyla
Family: Rhinocerotidae
Genus:† Coelodonta
Species:
†C. antiquitatis
Binomial name
†Coelodonta antiquitatis
(Blumenbach, 1799)
Subspecies [1]
  • C. a. praecursorGuérin, 1980
  • C. a. antiquitatisGuérin, 1980
Synonyms

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

Contents

The woolly rhinoceros (Coelodonta antiquitatis) is an extinct species of rhinoceros that inhabited northern Eurasia during the Pleistocene epoch. 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 range of the woolly rhinoceros 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, with environmental DNA records possibly extending the range of the species around 9,800 years ago. Its closest living relative is the Sumatran rhinoceros (Dicerorhinus sumatrensis).

Taxonomy

Molar tooth showing the cavity the genus was named for Em - Coelodonta antiquitatis - 2.jpg
Molar tooth showing the cavity the genus was named for

Woolly rhinoceros remains have been known long before the species was described and were the basis for some mythical creatures. Native peoples of Siberia believed their horns were the claws of giant birds. [3] A rhinoceros skull was found in Klagenfurt, Austria, in 1335, and was believed to be that of a dragon. [4] In 1590, it was used as the basis for the head on a statue of a lindworm. [5] Gotthilf Heinrich von Schubert maintained the belief that the horns were the claws of giant birds, and classified the animal under the name Gryphus antiquitatis, meaning "griffin of antiquity". [6]

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

Elasmotheriinae

† Elasmotherium

Rhinocerotinae

Black rhinoceros (Diceros bicornis)

White rhinoceros (Ceratotherium simum)

Indian rhinoceros (Rhinoceros unicornis)

Javan rhinoceros (Rhinoceros sondaicus)

Sumatran rhinoceros (Dicerorhinus sumatrensis)

†Woolly rhinoceros (Coelodonta antiquitatis)

† Merck's rhinoceros (Stephanorhinus kirchbergensis)

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

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. [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

Restoration Coelodonta antiquitatis by Benjamin Langlois.jpg
Restoration

An adult woolly rhinoceros typically measured 3.2 to 3.6 metres (10.5 to 11.8 ft) from head to tail, stood 1.45–1.6 metres (4.8–5.2 ft) tall at the shoulder, and weighed up to 1.5–2 metric tons (1.7–2.2 short tons) (with some sources placing the body mass of the species as high as 2.4–2.9 metric tons (5,300–6,400 lb) [21] [22] ) making it comparable in size to the largest living rhinoceros species, the white rhinoceros (Ceratotherium simum). [23] [24] 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. [25] [26] 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. [26] Unlike in modern rhinos, the large nasal horn was often flattened in cross-section, and abrasion patterns on the horn indicate it's possible use in brushing away snow when grazing. [27] 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. [28]

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. [28] 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. [29] Females had two nipples on the udders. [24]

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). [26] 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. [30] [26]

Skull and dentition

Woolly rhinoceros skull Coelodonta antiquitatis Crane.jpg
Woolly rhinoceros skull

The skull had a length between 70 and 90 cm (30 and 35 in). It was longer than those of other rhinoceros, giving the head a deep, downward-facing slanting position, similar to its fossil relative Stephanorhinus hemitoechus and Elasmotherium as well as the white rhinoceros. [31] 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. [26] The teeth of the woolly rhinoceros had thickened enamel and an open internal cavity. [26] Like other rhinos, adults did not have incisors. [32] 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. [33] This adaptation probably evolved as a result of the heavy pressure on the horn and face when the rhinoceros grazed underneath the thick snow. [34] 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. [35] This ossification inspired the junior synonym specific name tichorhinus, from Greek τειχος (teikhos) "wall", ῥις (ῥιν-) (rhis (rhin-)) "nose".

Paleobiology and palaeoecology

Woolly rhinoceros, woolly mammoths, cave lions, and horses in late Pleistocene northern Spain, by Mauricio Anton Ice age fauna of northern Spain - Mauricio Anton.jpg
Woolly rhinoceros, woolly mammoths, 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. [32] The two teats in the female suggest that she raised one calf, or more rarely two, every two to three years. [36] [24]

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. [37] Remains of woolly rhinoceros are frequently found in cave hyena dens with gnaw marks indicating that their remains were consumed by them, [38] which to a large degree likely reflects scavenging of the carcasses of already dead rhinoceroses. [39]

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. [30] They may have also been used to attract mates. [33] 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, [37]   and lower jaws and back ribs show signs of being broken and re-formed, which may have also come from fighting. [40]  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. [34]

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. [26] [31]  It had a wide upper lip like that of the white rhinoceros, which allowed it to easily pluck vegetation directly from the ground. [24] 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. [41] 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. [42] Pollen analysis shows it also ate woody plants (including conifers,  willows  and  alders), [26] along with flowers, [43] forbs and mosses. [44] 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. [45] Dental mesowear measurements further show that the woolly rhinoceros's diet was heavily composed on abrasive grasses. [46]

Growth and pathologies

It is estimated that woolly rhinoceroses could reach around 40 years of age, like their modern relatives. [37] 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). [47]

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. [48]

Habitat and distribution

Range of the woolly rhinoceros, including sites of fossils Woolly rhino area.png
Range of the woolly rhinoceros, including sites of fossils

The woolly rhinoceros lived mainly in lowlands, plateaus and river valleys, with dry to arid climates, [49] 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. [49] 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. [50] With its wide distribution, the woolly rhinoceros lived in some areas alongside the other rhinoceroses Stephanorhinus [28] and Elasmotherium . [51]

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. [30] [52] Even during the very warm Eemian interglacial, the range of the woolly rhinoceros extended into temperate regions such as Poland. [53] It had the widest range of any rhinoceros species. [54]

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, [30] 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. [49]

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. [30] 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. [55] Sometimes, only individual teeth or bone fragments are uncovered, which usually came from only one animal. [56] 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. [57] [58]

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. [26] A few sites from the early phase of the Last Glacial Period in the late Middle Paleolithic, such as the Gudenus Cave (Austria) [59] and the open air site of Königsaue (Saxony-Anhalt, Germany), [60] have heavily beaten rhinoceros bones lined with slash marks. This action was done partly to extract the nutritious bone marrow. [61]

Both horns and bones of the rhinoceros were used as raw materials for tools and weapons, as were remains from other animals. [62] In what is now Zwoleń, Poland, a device was made from a battered woolly rhinoceros pelvis. [63] 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. [64] 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. [65]

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. [66]

Ancient art

Cave paintings in Chauvet Cave 16 PanneauDesLions(CentreGauche)RhinocerosEnFuite.jpg
Cave paintings in Chauvet Cave

Many cave paintings from the Upper Paleolithic depict woolly rhinoceroses. The animal's defining features are prominently drawn, complete with the raised back and hump, contrasting with its low-lying head. Two curved lines represent the ears. The animal's horns are drawn with their long curvature, and in some cases, the coat is also indicated. Many paintings show a black band dividing the body. [67]

About 20 Paleolithic drawings of woolly rhinos were known before the discovery of the Chauvet Cave in France. [67] They are dated at over 31,000 years old, probably from the Aurignacian, [30] engraved on cave walls or drawn in red or black. One scene depicts two rhinos fighting each other with their horns. [68] 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. [69] Some images show rhinoceroses struck with spears or arrows, signifying human hunting. [70]

The site of Dolní Věstonice in Moravia, Czech Republic, was found with more than seven hundred statuettes of animals, many of woolly rhinoceroses. [70]

Extinction

Analysis of the nuclear genome suggests that the woolly rhinoceros experienced a population expansion beginning around 30,000 years ago. [71] 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. [30] The woolly rhinoceros was likely intolerant of deep snow, which its short limbs were inefficient in moving through. [72] Population fragmentation is likely to have played a role in its extinction. [73] The presence of large numbers of cervical ribs in specimens from the Netherlands may have been due to inbreeding or harsh environmental conditions. [74] 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. [71] 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. [75] 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, [76] [77] though other authors have argued that this specific environmental DNA record is unlikely to have been reworked. [72] 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. [72]

Frozen specimens

Mummified remains discovered in 1771 Frozen Coelodonta.jpg
Mummified remains discovered in 1771

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. [78] Later in 1877, a Siberian trader recovered a head and one leg from a tributary of the Yana River. [26]

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. [79] Both were sent to the Dzieduszycki Museum, where a detailed description was published in the museum's monograph. [80] Photographs were published in paleontological journals and textbooks, and the first modern paintings of the species were based on the mounted specimen. [81] The rhino is now located in the Lviv National Museum along with the mammoth. [81] Later, in 1929, the Polish Academy of Arts and Sciences sent an expedition to Starunia, finding the mummified remains of three rhinos. [82] 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. [83]

Cast of the mummified Starunia specimen, Natural History Museum, London Wooly Rhino at the Natural History Museum.jpg
Cast of the mummified Starunia specimen, Natural History Museum, 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. [26] [84] 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. [85]

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. [86] [26] Next year in 2008, a nearly complete skeleton came from the Chukochya River. [87] 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. [88] Dental analysis shows that the calf was about seven months old at the time of its death. [89] With its well-intact preservation, scientists proceeded to undergo DNA analysis. [90] [91]

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. [92]

See also

Related Research Articles

<span class="mw-page-title-main">Mammoth</span> Extinct genus of mammals

A mammoth is any species of the extinct elephantid genus Mammuthus. They lived from the late Miocene epoch into the Holocene until about 4,000 years ago, with mammoth species at various times inhabiting Africa, Asia, Europe, and North America. Mammoths are distinguished from living elephants by their spirally twisted tusks and in at least some later species, the development of numerous adaptions to living in cold environments, including a thick layer of fur.

<span class="mw-page-title-main">Beringia</span> Geographical region of Asia and North America currently partly submerged

Beringia is defined today as the land and maritime area bounded on the west by the Lena River in Russia; on the east by the Mackenzie River in Canada; on the north by 72° north latitude in the Chukchi Sea; and on the south by the tip of the Kamchatka Peninsula. It includes the Chukchi Sea, the Bering Sea, the Bering Strait, the Chukchi and Kamchatka Peninsulas in Russia as well as Alaska in the United States and the Yukon in Canada.

<i>Elasmotherium</i> Genus of extinct rhinoceroses

Elasmotherium is an extinct genus of large rhinoceros that lived in Eastern Europe, Central Asia and East Asia during Late Miocene through to the Late Pleistocene, with the youngest reliable dates around 39,000 years ago. It was the last surviving member of Elasmotheriinae, a distinctive group of rhinoceroses separate from the group that contains living rhinoceros (Rhinocerotinae), with the divergence between Elasmotherium

<i>Dicerorhinus</i> Genus of mammals

Dicerorhinus is a genus of the family Rhinocerotidae, consisting of a single extant species, the two-horned Sumatran rhinoceros, and several extinct species. The genus likely originated from the Late Miocene of central Myanmar. Many species previously placed in this genus probably belong elsewhere.

<span class="mw-page-title-main">Steppe bison</span> Extinct species of mammal

The steppe bison or steppe wisent is an extinct species of bison. It was widely distributed across the mammoth steppe, ranging from Western Europe to eastern Beringia in North America during the Late Pleistocene. It is ancestral to all North American bison, including ultimately modern American bison. Three chronological subspecies, Bison priscus priscus, Bison priscus mediator, and Bison priscus gigas, have been suggested.

<i>Rhinoceros</i> (genus) Genus of mammals

Rhinoceros is a genus comprising one-horned rhinoceroses. This scientific name was proposed by Swedish taxonomist Carl Linnaeus in 1758. The genus contains two species, the Indian rhinoceros and the Javan rhinoceros. Although both members are threatened, the Javan rhinoceros is one of the most endangered large mammals in the world with only 60 individuals surviving in Java (Indonesia). The word 'rhinoceros' is of Greek origin meaning "nose-horn".

<i>Panthera spelaea</i> Extinct species of lion

Panthera spelaea, commonly known as the cave lion, is an extinct Panthera species that was native to Eurasia and northwest North America during the Pleistocene epoch. Genetic analysis of ancient DNA has revealed that while closely related, it was a distinct species genetically isolated from the modern lion, with the genetic divergence between the two species estimated at around 500,000 years ago. The earliest fossils of the P. spelaea lineage in Eurasia date to around 700,000 years ago. It is closely related and probably ancestral to the American lion. The species ranged from Western Europe to eastern Beringia in North America, and was a prominent member of the mammoth steppe fauna, and an important apex predator across its range. It became extinct about 13,000 years ago. It closely resembled living lions with a coat of yellowish-grey fur though unlike extant lions, males appear to have lacked manes.

<i>Coelodonta</i> Extinct genus of rhinoceros

Coelodonta is an extinct genus of Eurasian rhinoceroses that lived from about 3.7 million years to 14,000 years ago, in the Pliocene and the Pleistocene epochs. It is best known from the type species, the woolly rhinoceros, which ranged throughout northern Eurasia during the Pleistocene. The earliest known species, Coelodonta thibetana, lived in Tibet during the Pliocene, with the genus spreading to the rest of Eurasia during the Pleistocene.

<span class="mw-page-title-main">Mammoth steppe</span> Prehistoric biome

The mammoth steppe, also known as steppe-tundra, was once the Earth's most extensive biome. During glacial periods in the later Pleistocene it stretched east-to-west, from the Iberian Peninsula in the west of Europe, then across Eurasia and through Beringia and into the Yukon in northwest Canada; from north-to-south, the steppe reached from the Arctic southward to southern Europe, Central Asia and northern China. The mammoth steppe was cold and dry, and relatively featureless, though climate, topography, and geography varied considerably throughout. Certain areas of the biome—such as coastal areas—had wetter and milder climates than others. Some areas featured rivers which, through erosion, naturally created gorges, gulleys, or small glens. The continual glacial recession and advancement over millennia contributed more to the formation of larger valleys and different geographical features. Overall, however, the steppe is known to be flat and expansive grassland. The vegetation was dominated by palatable, high-productivity grasses, herbs and willow shrubs.

<span class="mw-page-title-main">Steppe mammoth</span> Extinct species of mammoth

Mammuthus trogontherii, sometimes called the steppe mammoth, is an extinct species of mammoth that ranged over most of northern Eurasia during the Early and Middle Pleistocene, approximately 1.7 million to 200,000 years ago. The evolution of the steppe mammoth marked the initial adaptation of the mammoth lineage towards cold environments, with the species probably being covered in a layer of fur. One of the largest mammoth species, it evolved in East Asia during the Early Pleistocene, around 1.8 million years ago, before migrating into North America around 1.3 million years ago, and into Europe during the Early/Middle Pleistocene transition, around 1 to 0.7 million years ago. It was the ancestor of the woolly mammoth and Columbian mammoth of the later Pleistocene.

<span class="mw-page-title-main">Cave hyena</span> Extinct subspecies of carnivore

Cave hyena are extinct species or subspecies of hyena known from Eurasia, which ranged from Western Europe to eastern Asia and Siberia during the Pleistocene epoch. It is well represented in many European caves, primarily dating to the Last Glacial Period. It was an apex predator that preyed on large mammals, and was responsible for the accumulation of hundreds of large Pleistocene mammal bones in areas including horizontal caves, sinkholes, mud pits, and muddy areas along rivers.

<span class="mw-page-title-main">Woolly mammoth</span> Extinct species of mammoth

The woolly mammoth is an extinct species of mammoth that lived from the Middle Pleistocene until its extinction in the Holocene epoch. It was one of the last in a line of mammoth species, beginning with the African Mammuthus subplanifrons in the early Pliocene. The woolly mammoth began to diverge from the steppe mammoth about 800,000 years ago in Siberia. Its closest extant relative is the Asian elephant. The Columbian mammoth lived alongside the woolly mammoth in North America, and DNA studies show that the two hybridised with each other. Mammoth remains had long been known in Asia before they became known to Europeans. The origin of these remains was long a matter of debate and often explained as being remains of legendary creatures. The mammoth was identified as an extinct species of elephant by Georges Cuvier in 1796.

<i>Coelodonta tologoijensis</i> Extinct species of mammal

Coelodonta tologoijensis is an extinct species of rhinoceros belonging to the genus Coelodonta, related to the woolly rhinoceros. It is known from fossils found in Siberia and Mongolia, dating from the Early Pleistocene to Middle Pleistocene. One skull found in the Kyffhauser hills near the town of Bad Frankenhausen, Germany, dating to approximately 450,000 years was formerly assigned to the species by researchers, which would have made it the earliest known member of Coelodonta in Europe, However, a 2022 study refuted the assignment of the Bad Frankenhausen skull to C. tologoijensis, interpreting it as the skull of the woolly rhinoceros instead, meaning that the species is currently confined to Asia.

<i>Stephanorhinus</i> Extinct genus of rhinoceros

Stephanorhinus is an extinct genus of two-horned rhinoceros native to Eurasia and North Africa that lived during the Late Pliocene to Late Pleistocene. Species of Stephanorhinus were the predominant and often only species of rhinoceros in much of temperate Eurasia, especially Europe, for most of the Pleistocene. The last two species of Stephanorhinus – Merck's rhinoceros and the narrow-nosed rhinoceros – went extinct during the last glacial period.

<span class="mw-page-title-main">Narrow-nosed rhinoceros</span> Extinct species of rhinoceros

The narrow-nosed rhinoceros, also known as the steppe rhinoceros is an extinct species of rhinoceros belonging to the genus Stephanorhinus that lived in western Eurasia, including Europe, as well as North Africa during the Pleistocene. It first appeared in Europe around 500,000 years ago during the Middle Pleistocene and survived there until at least 34,000 years Before Present. It was native to temperate and Mediterranean environments, where it fed on low growing plants and to a lesser extent woody plants. Evidence has been found that it was exploited for food by archaic humans, including Neanderthals.

<i>Stephanorhinus kirchbergensis</i> Extinct species of rhinoceros native to Eurasia during the Pleistocene

Stephanorhinus kirchbergensis, also known as Merck's rhinoceros is an extinct species of rhinoceros belonging to the genus Stephanorhinus from the Early-Middle to Late Pleistocene of Eurasia. Its range spanned from Western Europe to Eastern Asia. Among the last members of the genus, it co-existed alongside Stephanorhinus hemitoechus in the western part of its range.

Dream Cave is a natural limestone cavern located near Wirksworth in Derbyshire, England. It was discovered by lead miners in 1822 and was found to contain the almost complete skeletal remains of a woolly rhinoceros and other large mammal bones. These remains were acquired by the geologist William Buckland and are now housed in Oxford Museum.

<i>Dihoplus</i> Extinct genus of rhinoceros

Dihoplus is an extinct genus of rhinoceros that lived in Eurasia from the Late Miocene to Pliocene.

The Yana Rhinoceros Horn Site is an Upper Palaeolithic archaeological site located near the lower Yana River in northeastern Siberia, Russia, north of the Arctic Circle in the far west of Beringia. It was discovered in 2001, after thawing and erosion exposed animal bones and artifacts. The site features a well-preserved cultural layer due to the cold conditions, and includes hundreds of animal bones and ivory pieces as well as numerous artifacts, which are indicative of sustained settlement and a relatively high level of technological development. With an estimated age of around 32,000 calibrated years before present, the site provides the earliest archaeological evidence for human settlement in this region, or anywhere north of the Arctic Circle, where people survived extreme conditions and hunted a wide range of fauna before the onset of the Last Glacial Maximum. The Yana site is perhaps the earliest unambiguous evidence of mammoth hunting by humans.

References

  1. 1 2 Uzunidis, A.; Antoine, P.-O.; Brugal, J.-P. (2022). "A Middle Pleistocene Coelodonta antiquitatis praecursor Guérin (1980) (Mammalia, Perissodactyla) from Les Rameaux, SW France, and a revised phylogeny of Coelodonta Bronn, 1831" (PDF). Quaternary Science Reviews. 288. 107594. Bibcode:2022QSRv..28807594U. doi:10.1016/j.quascirev.2022.107594.
  2. ↑ Pei Wen-Chung (1956). "Quaternary mammalian fossils from Hsintsai, South-Eastern part of Honan". Acta Palaeontologica Sinica. Archived from the original on 21 March 2018.
  3. ↑ Howorth, H.H. (1887). The Mammoth and the Flood: An Attempt to Confront the Theory of Uniformity with the Facts of Recent Geology. S. Low, Marston, Searle, & Rivington. p. 7.
  4. ↑ Price, Samantha (2007-06-01). "Horns, Tusks, and Flippers: the Evolution of Hoofed Mammals" (PDF). Aquatic Mammals. 33 (2). Aquatic Mammals Journal: 254. doi:10.1578/am.33.2.2007.254. ISSN   0167-5427. S2CID   127893230. Archived from the original (PDF) on 2019-12-08.
  5. ↑ "RRC: The Klagenfurt Lindwurm". RRC. Retrieved 2019-12-08.
  6. ↑ 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.
  7. ↑ 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.
  8. ↑ Owen, Richard. (1846). A History of British Fossil Mammals, and Birds. J. Van Voorst.
  9. 1 2 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.
  10. ↑ 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).
  11. ↑ Bronn, H.G. (1838). Lethaea geognostica oder Abbildungen und Beschreibungen der für die Gebirgs-Formationen bezeichnendsten Versteinerungen [Lethaea Geognostica, or pictures and descriptions of the most characteristic fossils of the mountain formations] (in German). Vol. 2. Stuttgart. p. 451.
  12. 1 2 van der Made, Jan. "The rhinos from the Middle Pleistocene of Neumark-Nord (Saxony-Anhalt)" (PDF). Veröffentlichungen des Landesamtes für Archäologie[State Office of Archeology Saxony-Anhalt] (in German and English). 62: 432–527.
  13. ↑ von Koenigswald, W. (2002). Lebendige Eiszeit: Klima und Tierwelt im Wandel[Living Ice Age: Climate and wildlife in transition] (in German). Theiss. ISBN   978-3-8062-1734-6.
  14. ↑ RFE/RL (2012-12-06). "Siberia Surrenders Woolly Rhino Mysteries". RadioFreeEurope/RadioLiberty. Retrieved 2019-12-09.
  15. 1 2 Liu, Shanlin; Westbury, Michael V.; Dussex, Nicolas; Mitchell, Kieren J.; Sinding, Mikkel-Holger S.; Heintzman, Peter D.; Duchêne, David A.; Kapp, Joshua D.; von Seth, Johanna; Heiniger, Holly; Sánchez-Barreiro, Fátima (24 August 2021). "Ancient and modern genomes unravel the evolutionary history of the rhinoceros family". Cell. 184 (19): 4874–4885.e16. doi: 10.1016/j.cell.2021.07.032 . hdl: 10230/48693 . ISSN   0092-8674. PMID   34433011.
  16. ↑ Pandolfi, Luca (2023-01-19). "Reassessing the phylogeny of Quaternary Eurasian Rhinocerotidae". Journal of Quaternary Science. 38 (3): 291–294. Bibcode:2023JQS....38..291P. doi: 10.1002/jqs.3496 . hdl: 11563/163194 . ISSN   0267-8179.
  17. ↑ Deng, T.; Wang, X.; Fortelius, M.; Li, Q.; Wang, Y.; Tseng, Z.J.; Takeuchi, G.T.; Saylor, J.E.; Säilä, L.K. & Xie, G. (2011). "Out of Tibet: Pliocene Woolly Rhino Suggests High-Plateau Origin of Ice Age Megaherbivores". Science. 333 (6047): 1285–1288. Bibcode:2011Sci...333.1285D. doi:10.1126/science.1206594. PMID   21885780. S2CID   8913866.
  18. ↑ Uzunidis, Antigone; Antoine, Pierre-Olivier; Brugal, Jean-Philip (July 2022). "A Middle Pleistocene Coelodonta antiquitatis praecursor (Mammalia, Perissodactyla) from Les Rameaux, SW France, and a revised phylogeny of Coelodonta". Quaternary Science Reviews. 288: 107594. Bibcode:2022QSRv..28807594U. doi:10.1016/j.quascirev.2022.107594.
  19. ↑ Stefaniak, Krzysztof; Kovalchuk, Oleksandr; Ratajczak-Skrzatek, Urszula; Kropczyk, Aleksandra; Mackiewicz, Paweł; Kłys, Grzegorz; Krajcarz, Magdalena; Krajcarz, Maciej T.; Nadachowski, Adam; Lipecki, Grzegorz; Karbowski, Karol; Ridush, Bogdan; Sabol, Martin; Płonka, Tomasz (March 2023). "Chronology and distribution of Central and Eastern European Pleistocene rhinoceroses (Perissodactyla, Rhinocerotidae) – A review". Quaternary International. 674–675: 87–108. Bibcode:2023QuInt.674...87S. doi:10.1016/j.quaint.2023.02.004. S2CID   257337872.
  20. ↑ Yuan, Junxia; Sun, Guojiang; Xiao, Bo; Hu, Jiaming; Wang, Linying; Taogetongqimuge; Bao, Lei; Hou, Yamei; Song, Shiwen; Jiang, Shan; Wu, Yong; Pan, Dong; Liu, Yang; Westbury, Michael V.; Lai, Xulong (2023-09-26). "Ancient mitogenomes reveal a high maternal genetic diversity of Pleistocene woolly rhinoceros in Northern China". BMC Ecology and Evolution. 23 (1): 56. doi: 10.1186/s12862-023-02168-0 . ISSN   2730-7182. PMC   10521388 . PMID   37752413.
  21. ↑ Mallet, Christophe; Billet, Guillaume; Cornette, Raphaël; Alexandra Houssaye, And (2022-11-02). "Adaptation to graviportality in Rhinocerotoidea? An investigation through the long bone shape variation in their hindlimb". Zoological Journal of the Linnean Society. 196 (3): 1235–1271. doi:10.1093/zoolinnean/zlac007. ISSN   0024-4082.
  22. ↑ Ma, Jiao; Wang, Shiqi; Deng, Tao (2024-04-11). "When the woolly rhinoceroses roamed East Asia: a review of isotopic paleoecology of the genus Coelodonta from the Tibetan Plateau to northern Eurasia". Frontiers in Ecology and Evolution. 12. doi: 10.3389/fevo.2024.1377000 . ISSN   2296-701X.
  23. ↑ Hans-Dieter Sues, Ross D.E. MacPhee (June 30, 1999). Extinctions in Near Time. Causes, Contexts, and Consequences. Springer US. p. 262. ISBN   9780306460920 . Retrieved 17 September 2022.
  24. 1 2 3 4 Boeskorov, G. G. (2012). "Some specific morphological and ecological features of the fossil woolly rhinoceros (Coelodonta antiquitatis Blumenbach 1799)". Biology Bulletin. 39 (8): 692–707. Bibcode:2012BioBu..39..692B. doi:10.1134/S106235901208002X. S2CID   24868968.
  25. ↑ Fortelius, Mikael (1983). "The morphology and paleobiological significance of the horns of Coelodonta antiquitatis (Mammalia: Rhinocerotidae)". Journal of Vertebrate Paleontology. 3 (2): 125–135. Bibcode:1983JVPal...3..125F. doi:10.1080/02724634.1983.10011964. ISSN   0272-4634.
  26. 1 2 3 4 5 6 7 8 9 10 11 12 Boeskorov, Gennady G.; Lazarev, Peter A.; Sher, Andrei V.; Davydov, Sergei P.; Bakulina, Nadezhda T.; Shchelchkova, Marina V.; Binladen, Jonas; Willerslev, Eske; Buigues, Bernard; Tikhonov, Alexey N. (2011-08-01). "Woolly rhino discovery in the lower Kolyma River" (PDF). Quaternary Science Reviews. 30 (17–18): 2262–2272. Bibcode:2011QSRv...30.2262B. doi:10.1016/j.quascirev.2011.02.010. ISSN   0277-3791.
  27. ↑ Shidlovskiy, Fedor K.; Kirillova, Irina V.; Wood, John (2012-03-26). "Horns of the woolly rhinoceros Coelodonta antiquitatis (Blumenbach, 1799) in the Ice Age Museum collection (Moscow, Russia)". Quaternary International. Mammoths and Their Relatives 1: Biotopes, Evolution and Human Impact V International Conference, Le Puy-en-Velay, 2010. 255: 125–129. Bibcode:2012QuInt.255..125S. doi:10.1016/j.quaint.2011.06.051. ISSN   1040-6182.
  28. 1 2 3 Kahlke, Ralf-Dietrich; Lacombat, Frédéric (2008-11-01). "The earliest immigration of woolly rhinoceros (Coelodonta tologoijensis, Rhinocerotidae, Mammalia) into Europe and its adaptive evolution in Palaearctic cold stage mammal faunas" (PDF). Quaternary Science Reviews. 27 (21–22): 1951–1961. Bibcode:2008QSRv...27.1951K. doi:10.1016/j.quascirev.2008.07.013. ISSN   0277-3791.
  29. ↑ 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.
  30. 1 2 3 4 5 6 7 Stuart, Anthony J.; Lister, Adrian M. (2012). "Extinction chronology of the woolly rhinoceros Coelodonta antiquitatis in the context of late Quaternary megafaunal extinctions in northern Eurasia" (PDF). Quaternary Science Reviews. 51: 1–17. Bibcode:2012QSRv...51....1S. doi:10.1016/j.quascirev.2012.06.007. ISSN   0277-3791.
  31. 1 2 van der Made, Jan; Grube, René (2010). Meller, Harald (ed.). The rhinoceroses from Neumark-Nord and their nutrition (PDF) (in German and English). Halle/Saale. pp. 382–394.
  32. 1 2 Garutt, N.V. "Dental ontogeny of the woolly rhinoceros Coelodonta antiquitatis (Blumenbach, 1799)" (PDF). Cranium. 11 (1).
  33. 1 2 Shidlovskiy, Fedor K.; Kirillova, Irina V.; Wood, John (2012). "Horns of the woolly rhinoceros Coelodonta antiquitatis (Blumenbach, 1799) in the Ice Age Museum collection (Moscow, Russia)" (PDF). Quaternary International. 255. Elsevier BV: 125–129. Bibcode:2012QuInt.255..125S. doi:10.1016/j.quaint.2011.06.051. ISSN   1040-6182.
  34. 1 2 Garutt, N.V. (1998). Woolly Rhinoceros: Morphology, Systematics and Geological Significance (PDF) (Doctoral dissertation abstract). Russian Mining Institute.
  35. ↑ 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. ISSN   1120-6349. S2CID   86966532.
  36. ↑ "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.
  37. 1 2 3 Garutt, N.V. (1997). "Traumatic skull damages in the woolly rhinoceros, Coelodonta antiquitatis Blumenbach, 1799" (PDF). Cranium. 14 (1): 37–46.
  38. ↑ Diedrich, C.G. & ŽÁK, K. 2006. 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.
  39. ↑ Diedrich, C.G. (June 2012). "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)". Journal of Archaeological Science. 39 (6): 1749–1767. Bibcode:2012JArSc..39.1749D. doi:10.1016/j.jas.2012.01.013.
  40. ↑ Diedrich, Cajus (2008). "A skeleton of an injured Coelodonta antiquitatis (Blumenbach, 1807) from the Upper Pleistocene of north-western Germany" (PDF). Cranium. 25 (1): 1–16.
  41. ↑ Schreve, Danielle; Howard, Andy; Currant, Andrew; Brooks, Stephen; Buteux, Simon; Coope, Russell; Crocker, Barnaby; Field, Michael; Greenwood, Malcolm; Greig, James; Toms, Phillip (2012-12-18). "A Middle Devensian woolly rhinoceros (Coelodonta antiquitatis) from Whitemoor Haye Quarry, Staffordshire (UK): palaeoenvironmental context and significance" (PDF). Journal of Quaternary Science. 28 (2). Wiley: 118–130. doi:10.1002/jqs.2594. ISSN   0267-8179. S2CID   54874495.
  42. ↑ 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 (2014). "Fifty thousand years of Arctic vegetation and megafaunal diet" (PDF). Nature. 506 (7486): 47–51. Bibcode:2014Natur.506...47W. doi:10.1038/nature12921. PMID   24499916. S2CID   4461741.
  43. ↑ "Woolly Mammoths and Rhinos Ate Flowers". livescience.com. 2014-02-05. Retrieved 2019-11-07.
  44. ↑ Dale Guthrie, R. (1990). Frozen Fauna of the Mammoth Steppe. University of Chicago Press. ISBN   9780226311234.
  45. ↑ 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.
  46. ↑ Diana, Pushkina; Juha, Saarinen; Reinhard, Ziegler; Hervé, Bocherens (1 January 2020). "Stable isotopic and mesowear reconstructions of paleodiet and habitat of the Middle and Late Pleistocene mammals in south-western Germany". Quaternary Science Reviews . 227: 106026. Bibcode:2020QSRv..22706026D. doi:10.1016/j.quascirev.2019.106026 . Retrieved 30 September 2024– via Elsevier Science Direct.
  47. ↑ Shpansky, A.V. (2014). "Juvenile remains of the "woolly rhinoceros" Coelodonta antiquitatis (Blumenbach 1799) (Mammalia, Rhinocerotidae) from the Tomsk Priob'e area (southeast Western Siberia)" (PDF). Quaternary International. 333. Elsevier BV: 86–99. Bibcode:2014QuInt.333...86S. doi:10.1016/j.quaint.2014.01.047. ISSN   1040-6182.
  48. ↑ Diedrich, Cajus G. (1 February 2023). "Extinct Eurasian rhinoceros Coelodonta and Stephanorhinus dental pathologies and tooth change modus". Quaternary Science Reviews . 301: 107922. Bibcode:2023QSRv..30107922D. doi:10.1016/j.quascirev.2022.107922 . Retrieved 8 May 2024– via Elsevier Science Direct.
  49. 1 2 3 Boeskorov, G. "Woolly rhino (Coelodonta antiquitatis) distribution in Northeast Asia" (PDF). Deinsea. 8 (1).
  50. ↑ Kahlke, Ralf-Dietrich (2014-07-15). "The origin of Eurasian Mammoth Faunas (Mammuthus–Coelodonta Faunal Complex)" (PDF). Quaternary Science Reviews. 96: 32–49. Bibcode:2014QSRv...96...32K. doi:10.1016/j.quascirev.2013.01.012. ISSN   0277-3791.
  51. ↑ 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. ISSN   0305-1838.
  52. ↑ Garutt, N. V., & Boeskorov, G. G. (2001). Woolly rhinoceroses: On the history of the genus. Mamont i ego okruzhenie, 200, 157-167.
  53. ↑ Stefaniak, Krzysztof; Kovalchuk, Oleksandr; Marciszak, Adrian; Sobczyk, Artur; Socha, Paweł (28 September 2023). "Environmental conditions across Poland during the Eemian Interglacial reconstructed from vertebrate remains". Acta Geologica Polonica . 73 (3): 379–410. doi:10.24425/agp.2023.145621 . Retrieved 17 September 2024.
  54. ↑ 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.
  55. ↑ Diedrich, Cajus (2008). "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" (PDF). Mitteilungen der Höhlen und Karstforscher (in German): 100–117.
  56. ↑ 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.
  57. ↑ 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.
  58. ↑ 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)
  59. ↑ 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.
  60. ↑ 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)
  61. ↑ Ashton, N.; Lewis, S.; Stringer, C. (2010). The Ancient Human Occupation of Britain. ISSN. Elsevier Science. ISBN   978-0-444-53598-6.
  62. ↑ 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.
  63. ↑ 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. ISBN   978-83-89499-23-3.
  64. ↑ 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. ISSN   0305-4403.
  65. ↑ "13,300 year old spear made of woolly rhinoceros horn found on Arctic island". Siberian Times.
  66. ↑ "engraved bone/antler". British Museum.
  67. 1 2 Bahn, P.G.; Vertut, J. (1997). Journey Through the Ice Age. University of California Press. p. 153. ISBN   978-0-520-22900-6.
  68. ↑ Foundation, Bradshaw. "Fighting Rhino & Four Horses - The Cave Art Paintings of the Chauvet Cave". Bradshaw Foundation. Retrieved 2019-10-28.
  69. ↑ "Kapova Cave". Don's Maps. Retrieved 24 Aug 2019.
  70. 1 2 Guthrie, R.D. (2005). The Nature of Paleolithic Art. University of Chicago Press. ISBN   978-0-226-31126-5.
  71. 1 2 Lord, Edana; Dussex, Nicolas; Kierczak, Marcin; Díez-del-Molino, David; Ryder, Oliver A.; Stanton, David W.G.; Gilbert, M. Thomas P.; Sánchez-Barreiro, Fátima; Zhang, Guojie; Sinding, Mikkel-Holger S.; Lorenzen, Eline D.; Willerslev, Eske; Protopopov, Albert; Shidlovskiy, Fedor; Fedorov, Sergey (October 2020). "Pre-extinction Demographic Stability and Genomic Signatures of Adaptation in the Woolly Rhinoceros". Current Biology. 30 (19): 3871–3879.e7. Bibcode:2020CBio...30E3871L. doi:10.1016/j.cub.2020.07.046. hdl: 10037/20986 . PMID   32795436. S2CID   221114305.
  72. 1 2 3 Fordham, Damien A.; Brown, Stuart C.; Canteri, Elisabetta; Austin, Jeremy J.; Lomolino, Mark V.; Haythorne, Sean; Armstrong, Edward; Bocherens, Hervé; Manica, Andrea; Rey-Iglesia, Alba; Rahbek, Carsten; Nogués-Bravo, David; Lorenzen, Eline D. (2024-06-11). "52,000 years of woolly rhinoceros population dynamics reveal extinction mechanisms". Proceedings of the National Academy of Sciences. 121 (24): e2316419121. Bibcode:2024PNAS..12116419F. doi:10.1073/pnas.2316419121. ISSN   0027-8424. PMC   11181021 . PMID   38830089.
  73. ↑ Rey-Iglesia, Alba; Lister, Adrian M.; Stuart, Anthony J.; Bocherens, Hervé; Szpak, Paul; Willerslev, Eske; Lorenzen, Eline D. (July 2021). "Late Pleistocene paleoecology and phylogeography of woolly rhinoceroses". Quaternary Science Reviews. 263: 106993. Bibcode:2021QSRv..26306993R. doi: 10.1016/j.quascirev.2021.106993 .
  74. ↑ van der Geer, Alexandra A.E.; Galis, Frietson (29 August 2017). "High incidence of cervical ribs indicates vulnerable condition in Late Pleistocene woolly rhinoceroses". PeerJ . 5: e3684. doi: 10.7717/peerj.3684 . ISSN   2167-8359. PMC   5580387 . PMID   28875067.
  75. ↑ Wang, Yucheng; Pedersen, Mikkel Winther; Alsos, Inger Greve; De Sanctis, Bianca; Racimo, Fernando; Prohaska, Ana; Coissac, Eric; Owens, Hannah Lois; Merkel, Marie Kristine Føreid; Fernandez-Guerra, Antonio; Rouillard, Alexandra (2021-10-20). "Late Quaternary dynamics of Arctic biota from ancient environmental genomics". Nature. 600 (7887): 86–92. Bibcode:2021Natur.600...86W. doi:10.1038/s41586-021-04016-x. ISSN   1476-4687. PMC   8636272 . PMID   34671161. S2CID   239051880.
  76. ↑ Miller, Joshua H.; Simpson, Carl (2022-12-01). "When did mammoths go extinct?". Nature. 612 (7938): E1–E3. Bibcode:2022Natur.612E...1M. doi:10.1038/s41586-022-05416-3. ISSN   0028-0836. PMC   9712083 . PMID   36450914.
  77. ↑ Seeber, Pa; Batke, L; Dvornikov, Y; Schmidt, A; Wang, Y; Stoof-Leichsenring, Kr; Moon, Kl; Shapiro, B; Epp, Ls (2023-09-01). Mitochondrial genomes of Pleistocene megafauna retrieved from recent sediment layers of two Siberian lakes (Report). elife. doi: 10.7554/elife.89992.1 .
  78. ↑ "Three Centuries of Hunting for Ice Age Mummies and the Prospect of De-Extinction". Capeia. 2018-05-18. Archived from the original on 2019-11-07. Retrieved 2019-11-02.
  79. ↑ 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. ISSN   0091-7613.
  80. ↑ 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).
  81. 1 2 Chornobay, Yuriy M.; Drygant, Daniel M. (2009). "The Starunia collections in the Natural History Museum of the National Academy of Sciences of Ukraine in Lviv" (PDF). Geotourism/Geoturystyka. 3 (18): 45–50. doi: 10.7494/geotour.2009.18.3.45 . ISSN   1731-0830.
  82. ↑ Kucha, Henryk (2009). "The Starunia collections in the Institute of Systematics and Evolution of Animals, Polish Academy of Sciences in Kraków". Geotourism/Geoturystyka. 18 (1). AGHU University of Science and Technology Press: 71. doi: 10.7494/geotour.2009.18.71 . ISSN   1731-0830. S2CID   128734734.
  83. ↑ 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.
  84. ↑ 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.
  85. ↑ "Woolly Rhino Fossils Found". The New York Times. 1976-10-10.
  86. ↑ Boeskorov, G. G.; Lazarev, P. A.; Bakulina, N. T.; Shchelchkova, M. V.; Davydov, S. P.; Solomonov, N. G. (2009). "Preliminary study of a mummified woolly rhinoceros from the lower reaches of the Kolyma River" (PDF). Doklady Biological Sciences. 424 (1): 53–56. doi:10.1134/s0012496609010165. ISSN   0012-4966. PMID   19341085. S2CID   2674199.
  87. ↑ 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. ISSN   0277-3791.
  88. ↑ Liesowska, Anna. "Meet Sasha - the world's only baby woolly rhino". The Siberian Times.
  89. ↑ Liesowska, Anna. "Lifelike again after 34,000 years, the world's only baby woolly rhino". Siberian Times.
  90. ↑ Liesowska, Anna (2019-12-23). "Sasha, the world's only baby woolly rhino, is 34,000 years old, say scientists". Siberian Times.
  91. ↑ Chernova, O. F.; Protopopov, A. V.; Perfilova, T. V.; Kirillova, I. V.; Boeskorov, G. G. (2016). "Hair microstructure of the first time found calf of woolly rhinoceros Coelodonta antiquitatis" (PDF). Doklady Biological Sciences. 471 (1): 291–295. doi:10.1134/s0012496616060090. ISSN   0012-4966. PMID   28058607. S2CID   18015793.
  92. ↑ "Woolly rhino from Ice Age unearthed in Russian Arctic". BBC News. 30 December 2020. Retrieved 31 December 2020.
This page is based on this Wikipedia article
Text is available under the CC BY-SA 4.0 license; additional terms may apply.
Images, videos and audio are available under their respective licenses.