Paraceratherium - Knowledge

2197: 129: 2470: 1836: 1926: 1687: 2393: 1323: 1827: 1167: 941: 751: 1158: 1020: 4554: 2097: 158: 2257:. Some Russian authors suggested that the tusks were probably used for breaking twigs, stripping bark and bending high branches and that, because species from the early Oligocene had larger tusks than later ones, they probably had a more bark than leaf based diet. Since the species involved are now known to have been contemporaneous, and the differences in tusks are now thought to be sexually dimorphic, the latter idea is not accepted today. Herds of 2018: 1818: 2278: 1748: 742: 2083:—similar to the running rhinoceroses from which they descended. Some foot bones were almost 50 centimetres (20 in) long. The thigh bones typically measured 1.5 m (4.9 ft), a size only exceeded by those of some elephants and dinosaurs. The thigh bones were pillar-like and much thicker and more robust than those of other rhinoceroses, and the three 1129:, based on remains too fragmentary to identify properly. By analysing alleged differences between named genera and species, Lucas and Sobus found that these most likely represented variation within populations, and that most features were indistinguishable between specimens, as had been pointed out in the 1930s. The fact that the single skull assigned to 713:, and Chinese scientists worked in isolation from each other for much of the 20th century and published research mainly in their respective languages. Scientists from different parts of the world tried to compare their finds to get a more complete picture of these animals, but were hindered by politics and wars. The opposing taxonomic tendencies of " 2497:. Prothero and the zoologist Pavel V. Putshkov have considered these causes unlikely since these animals managed to survive regardless of these issues for millions of years under the harsh conditions of their environment, and were not much larger than the biggest proboscideans, extinct as well as extant, which faced similar challenges. 2196: 128: 1766:

because this species is known from the most complete remains. Estimates have been based on skull, teeth, and limb bone measurements, but the known bone elements are represented by individuals of different sizes, so all skeletal reconstructions are composite extrapolations, resulting in several weight

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of Mongolia represents an arid desert basin, and the environment is thought to have had few tall trees and limited brush cover, as the fauna consisted mainly of animals that fed from tree tops or close to the ground. A study of fossil pollen showed that much of China was woody shrubland, with plants

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Granger and Gregory argued that the large incisors were used for defence or for loosening shrubs by moving the neck downwards, thereby acting as picks and levers. Tapirs use their proboscis to wrap around branches while stripping off bark with the front teeth; this ability would have been helpful to

1980:

differed from the other species in that the nasal notch was deeper, with the bottom placed above the middle of molar M2, a proportionally higher occipital condyle compared to the occipital surface's height, short muzzle bones and diastema in front of the cheek teeth, and a high zygomatic arch with a

2516:, which would have been their most likely competitors. While cautioning that the true cause of their extinction will never be known for certain, Prothero found it to be more than a coincidence that paraceratheres disappeared just as large predators and other large herbivores entered Asia during the 2131:

would not have been able to run and move quickly, but they would have been able to cross large distances, which would be necessary in an environment with a scarcity of food. They may therefore have had large home ranges and have been migratory. Prothero suggests that animals as big as indricotheres

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as separate genera. In 2016, the Chinese researchers Haibing Wang and colleagues used the name Paraceratheriidae for the family and Paraceratheriine for the subfamily, and placed them outside of Hyracodontidae. Deng and colleagues confirmed previous studies with their 2021 analysis, suggesting that

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had been able to migrate back north to Central Asia during this time when that area had become tropical (it was arid during the early Oligocene). This implies the Tibetan region was not yet a high-elevation plateau that could act as a barrier, and large animals may therefore have been able to move

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There are no indications of the colour and skin texture of the animal because no skin impressions or mummies are known. Most life restorations show the creature's skin as thick, folded, grey, and hairless, based on modern rhinoceroses. Because hair retains body heat, modern large mammals such as

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from the Jiaozigou Formation of the Linxia Basin (to which the name refers) of northwestern China. A multitude of other species and genus names—mostly based on differences in size, snout shape, and front tooth arrangement—have been coined for various indricothere remains. Fossils attributable to

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specimens known by then, using the proportions of a modern rhinoceros as a guide. The result was too squat and compact, and Osborn had a more slender version drawn later the same year. Some later life restorations have made the animal too slender, with little regard to the underlying skeleton.

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finds implies that they inhabited a continuous landmass with a similar environment across it, but this is contradicted by palaeogeographic maps that show this area had various marine barriers, so the genus was successful in being widely distributed despite this. The fauna which coexisted with

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were long and formed a long "hump" along the back, where neck muscles and nuchal ligaments for holding up the skull were attached. The ribs were similar to those of modern rhinoceroses, but the ribcage would have looked smaller in proportion to the long legs and large bodies, because modern

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would need very large home ranges or territories of at least 1,000 square kilometres (250,000 acres) and that, because of a scarcity of resources, there would have been little room in Asia for many populations or a multitude of nearly identical species and genera. This principle is called

602:. The shoulder height was about 4.8 metres (15.7 feet), and the length about 7.4 metres (24.3 feet). Its weight is estimated to have been about 15 to 20 tonnes (33,000 to 44,000 lb). The long neck supported a skull that was about 1.3 metres (4.3 ft) long. It had large, tusk-like 2009:. The premolars only partially formed the pi pattern. Each molar was the size of a human fist; among mammals they were only exceeded in size by proboscideans, though they were small relative to the size of the skull. The lower cheek teeth were L-shaped, which is typical of rhinoceroses. 1290:. A 2004 paper by Deng and colleagues also recognised three distinct genera. Some western writers have similarly used names otherwise considered invalid since the 1989 revision, but without providing detailed analysis and justification. Deng and colleagues recognised six 2512:, and as their numbers dwindled, they would have become more vulnerable to other threats. Prothero has pointed out that gomphotheres are not known to have generally coexisted with paraceratheres, and there are no known co-occurrences between paraceratheres and the large 1761:

Early estimates of 30 tonnes (66,000 lb) are now considered exaggerated; it may have been in the range of 15 to 20 tonnes (33,000 to 44,000 lb) at maximum, and as low as 11 tonnes (24,000 lb) on average. Calculations have mainly been based on fossils of

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may be large mammals such as elephants, rhinoceroses and hippopotamuses. To aid in thermoregulation, these animals cool down during the day by resting in the shade or by wallowing in water and mud. They also forage and move mainly at night. Because of its large size,

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that have ever existed, but its precise size is unclear because of the lack of complete specimens. Its total body length was estimated as 8.7 m (28.5 ft) from front to back by Granger and Gregory in 1936, and 7.4 m (24.3 ft) by the palaeontologist

1988:

were reduced to a single pair of incisors in either jaw, which were large and conical, and have been described as tusks. The upper incisors pointed downwards; the lower ones were shorter and pointed forwards. Among known rhinoceroses, this arrangement is unique to

2151:. Adult individuals would be too large for any land predators to attack, but the young would have been vulnerable. Bite marks on bones from the Bugti beds indicate that even adults may have been preyed on by 10-to-11-metre (33 to 36 ft)-long crocodiles, 2074:

with those of elephants and sauropod dinosaurs with their likewise graviportal (heavy and slow moving) builds. Unlike such animals, which tend to lengthen the upper limb bones while shortening, fusing and compressing the lower limb, hand, and foot bones,

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had a long forehead, which was smooth and lacked the roughened area that serves as attachment point for the horns of other rhinoceroses. The bones above the nasal region are long and the nasal incision goes far into the skull. This indicates that

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on the sides were much reduced, as this robustness diminished their importance. The limbs were held in a column-like posture instead of bent, as in smaller animals, which reduced the need for large limb muscles. The front limbs had three toes.

1259:, the subfamily name Indricotheriinae is still in use because genus name synonymy does not affect the names of higher level taxa that are derived from these. Members of the subfamily are therefore still commonly referred to as indricotheres. 634:, eating mainly leaves, soft plants, and shrubs. It lived in habitats ranging from arid deserts with a few scattered trees to subtropical forests. The reasons for the animal's extinction are unknown, but various factors have been proposed. 1379:

forms. Most species did not have horns. Rhinoceros fossils are identified as such mainly by characteristics of their teeth, which is the part of the animals most likely to be preserved. The upper molars of most rhinoceroses have a

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Sander, P. M.; Christian, A.; Clauss, M.; Fechner, R.; Gee, C. T.; Griebeler, E. M.; Gunga, H. C.; Hummel, J. R.; Mallison, H.; Perry, S. F.; Preuschoft, H.; Rauhut, O. W. M.; Remes, K.; Tütken, T.; Wings, O.; Witzel, U. (2011).

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appears to have had large, strong neck muscles, which allowed it to sweep its head strongly downwards while foraging from branches. The upper profile of the skull was arched, a distinguishing feature of the genus. One skull of

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and its relatives became extinct after surviving for about 11 million years are unknown, but it is unlikely that there was a single cause. Theories include that their large size was related to the now outdated concept of

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may have lived in small herds, perhaps consisting of females and their calves, which they protected from predators. It has been proposed that 20 tonnes (44,000 lb) may be the maximum weight possible for land mammals, and

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vertebrae of the neck were wider than in most modern rhinoceroses, with space for strong ligaments and muscles that would be needed to hold up the large head. The rest of the vertebrae were also very wide, and had large

2005:(gap). This feature is found in mammals where the incisors and cheek teeth have different specialisations. The upper molars, except for the third upper molar that was V-shaped, had a pi-shaped (π) pattern and a reduced 1794:

fossils, the skeleton of the animal has been reconstructed in several different ways since its discovery. In 1923, Matthew supervised an artist to draw a reconstruction of the skeleton based on the even less complete

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was a browser with a diet consisting of relatively soft leaves and shrubs. Later rhinoceroses were grazers, with high-crowned teeth because their diets contained grit that quickly wore down their teeth. Studies of

1956:, mastoid-paroccipital processes that were relatively thin and placed back on the skull, a lambdoid crest, which extended less back, and an occipital condyle with a horizontal orientation, which it shared with 1901:, which are otherwise found in horned and tusked animals that need strong muscles to push and fight. It also had a deep pit for the attachment of nuchal ligaments, which hold up the skull automatically. The 2293:

have been found in early to late Oligocene (34–23 million years ago) formations across Eurasia, in modern-day China, Mongolia, India, Pakistan, Kazakhstan, Georgia, Turkey, Romania, Bulgaria, and the

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might have been taller than any proboscidean. Its shoulder height was estimated as 5.25 m (17.2 ft) at the shoulders by Granger and Gregory, but 4.8 m (15.7 ft) by the palaeontologist

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period, 23 to 34 million years ago. The genus is distinguished from other indricotheres by its large size, nasal incision that would have supported a muscular snout, and its down-turned

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may have migrated while continuously foraging from tall trees, which smaller mammals could not reach. Osborn suggested that its mode of foraging would have been similar to that of the high-browsing

1779:. The ears of elephants enlarge the body's surface area and are filled with blood vessels, making the dissipation of excess heat easier. According to Prothero, this would have been true for 1678:

in the Oligocene. These researchers did not find Hyracodontidae to form a natural group, and found Paraceratheriidae to be closer to Rhinocerotidae, unlike previous studies.

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Antoine, P. O.; Ibrahim Shah, S. M.; Cheema, I. U.; Crochet, J. Y.; Franceschi, D. D.; Marivaux, L.; Métais, G. G.; Welcomme, J. L. (2004). "New remains of the baluchithere

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from the late Oligocene of Kazakhstan may be too incomplete for its position to be resolved in relation to the other species; the same applies to proposed species such as

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Wang, Y.; Deng, T. (2005). "A 25 m.y. Isotopic record of paleodiet and environmental change from fossil mammals and paleosols from the NE margin of the Tibetan Plateau".

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based on a palate and other fragments from Dera Bugti, thought to belong to a giant member of that genus. These fossils are now thought to have belonged to an aberrant

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Martin, C.; Bentaleb, I.; Antoine, P. -O. (2011). "Pakistan mammal tooth stable isotopes show paleoclimatic and paleoenvironmental changes since the early Oligocene".

4675: 3198:"New data on Amynodontidae (Mammalia, Perissodactyla) from Eastern Europe: Phylogenetic and palaeobiogeographic implications around the Eocene-Oligocene transition" 2504:

proboscideans from Africa in the late Oligocene (between 28 and 23 million years ago) may have considerably changed the habitats they entered, like

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are around 1.3 metres (4.3 ft) long, 33 to 38 centimetres (13 to 15 in) at the back of the skull, and 61 centimetres (24 in) wide across by the

1384:-shaped (π) pattern on the crown, and each lower molar has paired L-shapes. Various skull features are also used for identification of fossil rhinoceroses. 4774: 2174:

was close to this limit. The reasons mammals cannot reach the much larger size of sauropod dinosaurs are unknown. The reason may be ecological instead of

3505: 3479: 3401: 1411:(natural) grouping. Radinsky's scheme is the prevalent hypothesis today. The hyracodont family contains long-legged members adapted to running, such as 2561: 653:, have been named, but no complete specimens exist, making comparison and classification difficult. Most modern scientists consider these genera to be 4764: 850:. His rationale for this reclassification was the species' distinctly down-turned lower tusks. In 1913, Forster-Cooper named a new genus and species, 787:(in modern-day Pakistan) in 1846 by a soldier named Vickary, but these fragments were unidentifiable at the time. The first fossils now recognised as 3073: 4662: 4306: 2845: 2798: 1074:

and Forster-Cooper himself had expressed similar doubts few years earlier. Although it had already been declared a junior synonym, the genus name

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continue to be discovered across Eurasia, but the political situation in Pakistan had become too unstable for further excavations to occur there.

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dispersed westward to Kazakhstan during the early Oligocene from the ancestral area of Mongolia, where the most primitive member of the genus,

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in Balochistan in 1907–1908. His material consisted of an upper jaw, lower teeth, and the back of a jaw. The fossils were collected in the

2245:; it would extract relatively little nutrition from its food and would have to eat large volumes to survive. Like other large herbivores, 1417:, and were distinguished by incisor characteristics. Indricotheres are distinguished from other hyracodonts by their larger size and the 1089:

and Jay C. Sobus published a revision of indricothere taxa, which was subsequently followed by western scientists. They concluded that

803:, where Pilgrim had previously been exploring. In 1908, he used the fossils as basis for a new species of the extinct rhinoceros genus 4779: 2474: 2420:

was the only species of the genus represented in the Oligocene of western Pakistan, while the genus was highly diversified across the

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in 1966. Previously, they had been regarded as a subfamily within Rhinocerotidea, or even a full family, Indricotheriidae. In a 1999

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epoch (34–23 million years ago). The first fossils were discovered in what is now Pakistan, and remains have been found across

986:, was named by the Chinese palaeontologist Yong-Xiang Li and colleagues based on jaw elements from the Hanjiajing Formation in the 2490: 1968:, thick mastoid-paroccipital processes, a lambdoid crest that extended back, and occipital condyles with a vertical orientation. 4744: 3054:"New fossils of paraceratheres (Perissodactyla, Mammalia) from the Early Oligocene of the Lanzhou Basin, Gansu Province, China" 1922:

shows it was only 8 percent of the skull length, while the brain of the Indian rhinoceros is 17.7 percent of its skull length.

1367:. The diversity within the rhinocerotoid group was much larger in prehistoric times; they ranged from dog-sized to the size of 3696:"Earliest known unequivocal rhinocerotoid sheds new light on the origin of Giant Rhinos and phylogeny of early rhinocerotoids" 725:

that are now known to be contemporaneous were of different ages. Many genera were named on the basis of subtle differences in

4224: 2461:, and through lowlands in the area, some of which were possibly under 2,000 m (6,600 ft) in elevation at the time. 1262:

In contrast to the revision by Lucas and Sobus, a 2003 paper by Chinese palaeontologist Jie Ye and colleagues suggested that

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trees, while Siberia and Kazakhstan also had walnut trees. Dera Bugti in Pakistan had dry, temperate to subtropical forest.

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in 1922. Also in 1923, Borissiak created the subfamily Indricotheriinae to include the various related forms known by then.

821:; it included several unrelated species of hornless rhinoceros, many of which have since been moved to other genera. Fossil 157: 2826:, a new genus of perissodactyles from the Upper Oligocene deposits of the Bugti hills of Baluchistan. —Preliminary notice" 914: 141: 641:

of the genus and the species within has a long and complicated history. Other genera of Oligocene indricotheres, such as

2067:-like openings (hollow parts of the bone) in their pre-sacral vertebrae, which probably helped to lighten the skeleton. 3196:

Tissier, J.; Becker, D.; Codrea, V.; Costeur, L.; Fărcaș, C.; Solomon, A.; Venczel, M.; Maridet, O.; Smith, T. (2018).

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Clauss, M.; Frey, R.; Kiefer, B.; Lechner-Doll, M.; Loehlein, W.; Polster, C.; Rössner, G. E.; Streich, W. J. (2003).

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in Kazakhstan; it was the most complete indricothere skeleton known, but it lacked the skull. It is mounted in the

2367:), all adapted to arid environments. Trees were rare, and concentrated near groundwater. The parts of China where 2001:

otherwise found behind the incisors were lost. The incisors were separated from the row of cheek teeth by a large

1093:, as the oldest name, was the only valid indricothere genus from the Oligocene, and contained four valid species, 4408:

and other vertebrates from Oligocene and middle Miocene deposits of the Kağızman-Tuzluca Basin, Eastern Turkey".

971:, including the legs of a specimen standing in an upright position, indicating that it had died while trapped in 1322: 834:

In 1910, more partial fossils were discovered in Dera Bugti during an expedition by the British palaeontologist

3667:, a Middle to Late Eocene hyracodontid (Perissodactyla: Rhinocerotoidea) from Asia and Western North America". 2342:

appears to have varied across its range, based on the types of geological formations it has been found in. The

3866: 3520: 3460: 3386: 1826: 1166: 858:, based on larger fossils from the same excavations (some of which he had earlier suggested to belong to male 3428:"A giant rhinocerotoid (Mammalia, Perissodactyla) from the Late Oligocene of north-central Anatolia (Turkey)" 3354: 1783:, as indicated by the robust bones around the ear openings. The palaeontologists Pierre-Olivier Antoine and 2070:

The limbs were large and robust to support the animal's large weight, and were in some ways similar to and

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below follows the 1989 analysis of Indricotheriinae by Lucas and Sobus, and shows the closest relatives of

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were made through various colonial links to Asia. The first known indricothere fossils were collected from

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elephants and rhinoceroses are largely hairless. Prothero has proposed that, contrary to most depictions,

1071: 940: 887: 2237:, which are mainly leaves. Like its perissodactyl relatives, the horses, tapirs, and other rhinoceroses, 1914:

has a domed forehead, whereas others have flat foreheads, possibly because of sexual dimorphism. A brain

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Meng, J.; McKenna, M. C. (1998). "Faunal turnovers of Palaeogene mammals from the Mongolian Plateau".

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with much room for muscles, tendons, ligaments, and nerves, to support the head, neck, and spine. The

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in 1997. The neck was estimated at 2 to 2.5 m (6.6 to 8.2 ft) long by the palaeontologists

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skull AMNH 18650 (left), and front views of a cast of same, showing the incisors (center and right).

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belongs, was first classified as part of the family Hyracodontidae by the American palaeontologist

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Borissiak, A. A. (1924). "Über die Unterfamilie Indricotheriinae Boriss. = Baluchitheriinae Osb".

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rhinoceroses are comparatively short-limbed. The last vertebra of the lower back was fused to the

1421:

structure of their snouts, incisors and canines. The earliest known indricothere is the dog-sized

4503: 4484: 4441: 4298: 4000: 3961: 3829: 3676: 3578: 2941: 2600: 2409: 1066:, because these specimens were collected at the same locality and were possibly part of the same 1051: 702: 638: 355: 152: 2178:, and perhaps related to reproduction strategies. Movement, sound, and other behaviours seen in 1403:, the American palaeontologist Luke Holbrook found indricotheres to be outside the hyracodontid 4706: 4502:

Leopold, E. B.; Liu, G.; Clay-Poole, S. (1992), "Low-biomass vegetation in the Oligocene?", in

2381: 2006: 4688: 4680: 4573: 4511: 4433: 4290: 4166: 3924: 3733: 3619: 3368: 3358: 3237: 3143: 2592: 2505: 2494: 2421: 2242: 1902: 1890:. A distinguishing feature was that the nasal incision was retracted to the P2-P3 premolars. 1879: 1741: 1138: 1047: 879: 818: 718: 690: 555: 551: 222: 4716: 4693: 4476: 4425: 4386: 4382: 4351: 4282: 4205: 4156: 4148: 4116: 4086: 3992: 3951: 3914: 3904: 3821: 3787: 3723: 3715: 3609: 3570: 3439: 3312: 3273: 3227: 3217: 3178: 3133: 3125: 3065: 2931: 2880: 2837: 2790: 2750: 2584: 2299: 2226: 2158: 2137: 1875: 1858: 1776: 1751: 1509: 1121: 1086: 867: 627: 2017: 144:; this is the most completely known skeleton, but the skull is a cast of a specimen at the 3561:

Radinsky, L. B. (1966). "The families of the Rhinocerotoidea (Mammalia, Perissodactyla)".

2425: 1737: 1524: 1423: 1342: 1224: 547: 4533:

Putshkov, P. V. (2001). ""Proboscidean agent" of some Tertiary megafaunal extinctions".

4472: 4421: 4347: 4278: 4201: 3711: 3308: 3213: 2779:, a new genus of Rhinocerotidae from the Bugti Hills of Baluchistan.—Preliminary notice" 2746: 4404:

Sen, S.; Antoine, P. O.; Varol, B.; Ayyildiz, T.; Sözeri, K. (2011). "Giant rhinoceros

4161: 4136: 3919: 3892: 3791: 3728: 3695: 3506:"Indricotheres (Perissodactyla, Mammalia) from Oligocene in Linxia Basin, Gansu, China" 3260:

Granger, W.; Gregory, W. K. (1936). "Further notes on the gigantic extinct rhinoceros,

3232: 3197: 3138: 3109: 2449: 2038: 1898: 1724: 1482: 1459:. It had also lost the second and third lower incisors, lower canines, and lower first 1364: 1360: 1339: 1059: 987: 906: 654: 209: 3856:"Dinosaur models: The good, the bad, and using them to estimate the mass of dinosaurs" 4733: 4488: 4302: 4152: 4004: 3444: 3427: 3069: 2604: 2517: 2153: 2047: 1965: 1574: 1439: 1356: 1126: 991: 930: 780: 726: 310: 86: 4445: 3965: 2703:(1910). "Notices of new mammalian genera and species from the Tertiaries of India". 2277: 1817: 1643:

Lucas and colleagues had reached similar conclusions in a previous 1981 analysis of

917:. In 1916, based on these remains, Aleksei Alekseeivich Borissiak erected the genus 3833: 2500:

Putshkov and Andrzej H. Kulczicki instead suggested in 1995 and 2001 that invading

2316: 2312: 2269:, rather than to modern rhinoceroses, whose heads are carried close to the ground. 2175: 2079:

had short upper limb bones and long hand and foot bones—except for the disc-shaped

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is complex due to the fragmentary nature of the known fossils and because Western,

585: 580: 263: 251: 1694:(olive green) compared with that of humans, other large mammals, and the dinosaur 4355: 3346: 3222: 2731:: a genus of aberrant rhinoceroses from the Lower Miocene deposits of Dera Bugti" 878:

were so fragmentary that Foster-Cooper was only able to identify it as a kind of

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only in the anatomy of the rear portion of the jaw, as well as its larger size.

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Gromova published a more complete skeletal reconstruction in 1959, based on the

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remained popular in various media because of the publicity surrounding Osborn's

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tooth characteristics—features that vary within populations of other rhinoceros

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skeleton from the Aral Formation, but this also lacked several neck vertebrae.

1747: 1125:). They considered most other names to be junior synonyms of those taxa, or as 733:—and are therefore not accepted by most scientists for distinguishing species. 4549: 4429: 4286: 4077:

Granger, W.; Gregory, W. K. (1935). "A revised restoration of the skeleton of

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Fortelius, M.; Kappelman, J. (1993). "The largest land mammal ever imagined".

3182: 2884: 2841: 2794: 2588: 2570:: Imperial and International Networks in Early-Twentieth Century Paleontology" 2458: 2376: 2360: 2352: 2064: 1871: 1715:

in 1959, but the former estimate is now considered exaggerated. The weight of

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from the late Oligocene of Kazakhstan, Mongolia, and northern China included

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Deng, T.; Lu, X.; Wang, S.; Flynn, L. J.; Sun, D.; He, W.; Chen, S. (2021).

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species in 2021, including some that had previously been declared synonyms,

972: 681:, so most reconstructions of the genus are based on it. Differences between 623: 611: 563: 169: 111: 55: 48: 4437: 4294: 4188:

from the Late/latest Oligocene of the Bugti hills, Balochistan, Pakistan".

4170: 3928: 3825: 3737: 3623: 3614: 3598:"The phylogeny and classification of tapiromorph perissodactyls (Mammalia)" 3597: 3317: 3292: 3241: 3147: 2936: 2911: 2755: 2726: 2596: 4641: 3956: 3943: 3167:, sp. n., from the Dera Bugti deposits of Baluchistan.—preliminary notice" 1046:

specimen that lacked the M3 molar. In 1936, the American palaeontologists

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Most terrestrial predators in their habitat were no bigger than a modern

2002: 1949: 1915: 1894: 1460: 1331: 995: 967:. Various indricothere remains were found in formations of the Mongolian 960: 910: 619: 189: 101: 96: 81: 76: 66: 3996: 4667: 3680: 3582: 2295: 2262: 822: 770: 603: 571: 567: 116: 91: 3719: 3387:"A new genus of giant rhinoceros from oligocene of Dzungaria, Sinkang" 3297:

Philosophical Transactions of the Royal Society B: Biological Sciences

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Philosophical Transactions of the Royal Society B: Biological Sciences

2122:

due to its size. According to Prothero, the best living analogues for

2118:

suggested in 1988 that overheating may have been a serious problem in

975:, as well as a very complete skull. These remains became the basis of 4535:

Terra Degli Elefanti Congresso Internazionale: The World of Elephants

4120: 4090: 3909: 3663:

Lucas, S. G.; Schoch, R. M.; Manning, E. (1981). "The systematics of

3052:

Yong-Xiang, L.; Yun-Xiang, Z.; Ji, L.; Zhi-Chao, L.; Kun, X. (2017).

2965:

Bulletin de la Société des Naturalistes de Moscou, Section Géologique

2945: 2379:, and the most common plant fossils are leaves of the desert-adapted 2332: 2052: 2037:

has yet been found and the tail is completely unknown. The atlas and

1428: 1191:

from the late Oligocene of Pakistan included junior synonyms such as

599: 559: 199: 179: 4586: 3574: 3277: 3162: 622:

and extant rhinoceroses. Because of its size, it would have had few

3855: 2448:

in Linxia during the late Oligocene, and it is possible that these

2385:. Trees in Mongolia and China included birch, elm, oaks, and other 2165:

may have been lengthy and individuals may have had long lifespans.

4480: 2468: 2391: 2276: 2266: 2195: 2095: 1984:

Unlike those of most primitive rhinocerotoids, the front teeth of

1924: 1887: 1746: 1685: 1549: 1433: 1404: 1321: 1018: 939: 618:

may have been similar to that of modern large mammals such as the

543: 3893:"Why sauropods had long necks; and why giraffes have short necks" 2140:(a grazer) exploit different niches in the same areas of Africa. 2136:; it is used to explain how the black rhinoceros (a browser) and 4654: 2302:

development of the Alpine-Himalayan mountain belt. The range of

2144: 1345:, which includes modern rhinoceroses, can be traced back to the 730: 4590: 4137:"Biology of the sauropod dinosaurs: The evolution of gigantism" 3426:

Antoine, P. O.; Karadenizli, L.; Saraç, G. E.; Sen, S. (2008).

1349:—about 50 million years ago—with early precursors such as 1215:

from the middle and late Oligocene of northwest China included

1137:

was domed, while others were flat at the top was attributed to

3981:"There were giants upon the earth in those days (Book Review)" 1964:

had robust maxillae and premaxillae, upturned zygomata, domed

1437:

is known from the middle Eocene; by the late Eocene the genus

614:(trunk). The legs were long and pillar-like. The lifestyle of 2436:, lived, and descendants may have continued to South Asia as 1670:

40 million years ago, with the resulting stock evolving into

2416:

based on their phylogenetic analys in 2021. They found that

1893:

The back of the skull was low and narrow, without the large

1723:, with the largest complete skeleton known belonging to the 1002:, based on a complete skull with an associated mandible and 825:

that Pilgrim had previously assigned to the unrelated genus

4510:, Princeton: Princeton University Press, pp. 399–420, 1231:

may be distinct enough to warrant its original genus name

4225:"All-time giants: the largest animals and their problems" 2924:

Philosophical Transactions of the Royal Society of London

3649: 3647: 3645: 3643: 3641: 3639: 3637: 3635: 3633: 3255: 3253: 3251: 2984:

Zentralblatt für Mineralogie, Geologie und Paläontologie

763:, which was the basis for its separation from the genus 3807:"Shoulder height, body mass and shape of proboscideans" 2672: 2670: 2668: 2666: 2664: 2662: 2660: 2658: 2221:

teeth confirm the creatures had a diet of soft leaves;

1278:. They also recognised the validity of species such as 562:

that has ever existed and lived from the early to late

3547: 3545: 3543: 3541: 2656: 2654: 2652: 2650: 2648: 2646: 2644: 2642: 2640: 2638: 2520:(between 23 and 16 million years ago). 1944:

are mainly discernible through skull characteristics.

661:, and it is thought to contain the following species; 4570:

Rhinoceros Giants: The Palaeobiology of Indricotheres

3944:"Estimating body mass from the astragalus in mammals" 2686: 2684: 2682: 2628: 2626: 2624: 2622: 2620: 2618: 2616: 2614: 2541: 2539: 2537: 2535: 2533: 882:, but he mentioned the possibility of confusion with 846:, meaning "near the hornless beast", in reference to 27:

Extinct genus of hornless rhinocerotoids from Eurasia

3340: 3338: 3336: 3334: 3332: 3330: 3328: 2768: 2766: 1981:

prominent hind end, and a smaller upper incisor I1.

334: 317: 300: 287: 258: 4599: 759:1911 illustrations of the down-turned lower jaw of 717:" have also contributed to the problem. Inaccurate 30:"Baluchitherium" redirects here. For the song, see 3266:Bulletin of the American Museum of Natural History 2424:, northwestern China, and Kazakhstan north to the 1997:. The incisors may have been larger in males. The 4336:Palaeogeography, Palaeoclimatology, Palaeoecology 3110:"An Oligocene giant rhino provides insights into 2400:species during the early (yellow) and late (red) 2055:, a feature found in advanced rhinoceroses. Like 1431:of western North America and Asia. The cow-sized 1235:, though its exact position requires evaluation. 578:means "near the hornless beast", in reference to 4329: 4327: 2508:do today. This would have made food scarcer for 2298:. Their distribution may be correlated with the 598:is unknown because of the incompleteness of the 4081:, gigantic fossil rhinoceros of Central Asia". 1070:variable species. The American palaeontologist 831:were later shown to belong to the new species. 4508:Eocene-Oligocene Climatic and Biotic Evolution 4111:, a giant hornless rhinoceros from Mongolia". 4102: 4100: 3756:Trudy Paleontology Institut Akademii Nauk SSSR 3754:Gromova, V. L. (1959). "Gigantskie nosorogi". 3749: 3747: 3694:Wang, H.; Bai, B.; Meng, J.; Wang, Y. (2016). 3047: 3045: 2720: 2718: 1775:had large elephant-like ears that it used for 959:led a well-documented expedition to China and 929:, until 1923, but the Russian palaeontologist 838:. Based on these remains, Foster-Cooper moved 4052:blogs.scientificamerican.com/tetrapod-zoology 4046:Sea Monsters on Medieval and Renaissance Maps 1787:have expressed scepticism towards this idea. 1035:In 1922 Forster-Cooper named the new species 721:previously led scientists to believe various 8: 3773: 3771: 3769: 3353:, New York, New York & Oxford, England: 3349:, in Prothero, D. R.; Schoch, R. M. (eds.), 2208:The simple, low-crowned teeth indicate that 866:later that year because the former name was 606:and a nasal incision that suggests it had a 4528: 4526: 1355:. Rhinocerotoidea contains three families; 4587: 3504:Qui, Z.-X.; Wang, B.-Y.; Deng, T. (2004). 3465:in the northern Junggar Basin of Xinjiang" 2963:n. sp. provenant du district de Tourgay". 1031:), in rhinoceros-like and slender versions 693:, which would make them the same species. 127: 38: 4160: 3955: 3918: 3908: 3780:Zoological Journal of the Linnean Society 3727: 3613: 3443: 3432:Zoological Journal of the Linnean Society 3316: 3293:"The extinct rhinoceroses of Baluchistan" 3231: 3221: 3137: 3103: 3101: 3099: 3097: 3095: 3093: 3091: 2935: 2754: 2705:Records of the Geological Survey of India 2440:, dispersing through the Tibetan region. 2408:Deng and colleagues speculated about the 1387:The subfamily Indricotheriinae, to which 1149:represents the male of the same species. 948:skull (AMNH 18650), formerly assigned to 791:were discovered by the British geologist 2555: 2553: 2551: 2404:, according to Deng and colleagues, 2021 2249:would have had a large digestive tract. 2016: 1948:had features such as relatively slender 1731:). Despite its roughly equivalent mass, 1223:. In 2013, the American palaeontologist 1062:(an invalid name for the same taxon) of 998:and colleague described the new species 990:of China; the name refers to the nearby 894:was named, suggested it may have been a 2529: 2444:existed in Xinjiang and Kazakhstan and 1790:Due to the fragmentary nature of known 1443:of Asia had almost reached the size of 1085:In 1989, the American palaeontologists 994:. In 2021, the Chinese palaeontologist 921:named for a mythological monster, the " 677:. The most completely-known species is 414: 370: 4239:from the original on 27 September 2016 3171:Annals and Magazine of Natural History 2873:Annals and Magazine of Natural History 2830:Annals and Magazine of Natural History 2783:Annals and Magazine of Natural History 2457:freely along the eastern coast of the 1175:Skull (left) and vertebrae (right) of 905:expedition later found fossils in the 4572:. Indiana: Indiana University Press. 4011:from the original on 12 November 2014 2801:from the original on 20 November 2015 1375:forms adapted for running and squat, 925:". He did not assign a species name, 7: 3891:Taylor, M. P.; Wedel, M. J. (2013). 3843:from the original on 24 August 2016. 3459:Ye, Y.; J., Meng; Yu, W. W. (2003). 3407:from the original on 22 October 2014 3076:from the original on 12 October 2017 1719:was similar to that of some extinct 1145:fossils represent the female, while 870:, as it had already been used for a 4371:Earth and Planetary Science Letters 4058:from the original on 6 October 2014 3345:Lucas, S. G.; Sobus, J. C. (1989), 3264:, from the Oligocene of Mongolia". 2891:from the original on 12 August 2018 2848:from the original on 7 October 2015 1660:evolved from a clade consisting of 1565: 1540: 1515: 1488: 1477: 1451:itself lived in Eurasia during the 1407:, and wrote that they may not be a 4775:Taxa named by Clive Forster-Cooper 3863:Dinofest International Proceedings 3792:10.1111/j.1096-3642.1993.tb02560.x 3347:"The systematics of indricotheres" 2475:Pakistan Museum of Natural History 2285:, by Elizabeth Rungius Fulda, 1923 2225:studies have yet to be conducted. 1184:According to Lucas and Sobus, the 965:American Museum of Natural History 435:Forster-Cooper, 1913 (preoccupied) 146:American Museum of Natural History 25: 3872:from the original on 4 March 2016 3485:from the original on 3 March 2016 3030:from the original on 9 March 2016 2577:Journal of the History of Biology 1935:Musée d'Histoire Naturelle, Paris 1874:upper lip similar to that of the 1023:1923 skeletal reconstructions of 4765:Rupelian genus first appearances 4552: 4312:from the original on 8 June 2019 4153:10.1111/j.1469-185X.2010.00137.x 4032:"Tet Zoo Bookshelf: van Grouw's 3653:Prothero, 2013. pp. 107–121 3445:10.1111/j.1096-3642.2007.00366.x 3070:10.19615/j.cnki.1000-3118.170922 1897:crests at the top and along the 1834: 1825: 1816: 1330:with other rhinos, according to 1165: 1156: 944:Preparator Otto Falkenbach with 749: 740: 156: 4190:Journal of Asian Earth Sciences 3396:. 11 (in Chinese and English). 3351:The Evolution of Perissodactyls 2727:"On the skull and dentition of 2676:Prothero, 2013. pp. 87–106 2493:, vegetational change, and low 2311:included other rhinocerotoids, 1054:proposed that Forster-Cooper's 955:In 1922, the American explorer 886:. The American palaeontologist 3551:Prothero, 2013. pp. 53–66 3012:"The extinct giant rhinoceros 2690:Prothero, 2013. pp. 35–52 2632:Prothero, 2013. pp. 67–86 2545:Prothero, 2013. pp. 17–34 1647:, wherein they still retained 1326:Phylogenetic relationships of 773:and part of a mandible (right) 512:Pristinotherium brevicervicale 1: 4770:Fossil taxa described in 1911 3948:Acta Palaeontologica Polonica 3814:Acta Palaeontologica Polonica 3000:Prothero, 2013. pp. 1–16 2961:Indricotherium transouralicum 1744:and Mathew J. Wedel in 2013. 915:Moscow Paleontological Museum 472:Indricotherium transouralicum 142:Moscow Paleontological Museum 4755:Prehistoric placental genera 4750:Aquitanian genus extinctions 4356:10.1016/j.palaeo.2011.07.010 3519:(3): 177–192. Archived from 3223:10.1371/journal.pone.0193774 3016:of Western and Central Asia" 2869:"Correction of generic name" 1270:were valid genera, and that 862:), but he renamed the genus 457:Paraceratherium zhajremensis 4785:Oligocene mammals of Europe 4760:Extinct animals of Pakistan 3291:Forster-Cooper, C. (1934). 3161:Forster-Cooper, C. (1922). 2910:Forster-Cooper, C. (1923). 2867:Forster-Cooper, C. (1913). 2820:Forster-Cooper, C. (1913). 2773:Forster-Cooper, C. (1911). 1363:("true rhinoceroses"), and 1255:is now a junior synonym of 979:, named by Osborn in 1923. 903:Russian Academy of Sciences 4801: 4391:10.1016/j.epsl.2005.05.006 4210:10.1016/j.jaes.2003.09.005 4030:Naish, D. (29 June 2013). 3515:(in Chinese and English). 3474:(in Chinese and English). 3130:10.1038/s42003-021-02170-6 2396:Map showing localities of 2188:are entirely conjectural. 1708:largest known land mammals 1371:. There were long-legged, 1314:as synonyms of the genus. 525:Benaratherium callistratum 29: 4780:Oligocene mammals of Asia 4506:; Berggren, W.A. (eds.), 4430:10.1007/s00114-011-0786-z 4287:10.1007/s00442-003-1254-z 4223:Alexander, R. M. (1998). 4186:Paraceratherium bugtiense 4113:American Museum Novitates 4083:American Museum Novitates 3183:10.1080/00222932208632717 2885:10.1080/00222931308693431 2842:10.1080/00222931308693412 2795:10.1080/00222931108693085 2777:Paraceratherium bugtiense 2729:Paraceratherium bugtiense 2589:10.1007/s10739-014-9395-y 2473:Life sized model outside 2428:. They hypothesised that 2147:and were not a threat to 1952:and premaxillae, shallow 1588: 1570: 1563: 1545: 1538: 1520: 1513: 1493: 1486: 1427:from the middle and late 1064:Paraceratherium bugtiense 1044:Paraceratherium bugtiense 584:, the genus in which the 361: 354: 284: 279: 257: 250: 153:Scientific classification 151: 135: 126: 41: 4269:(Submitted manuscript). 2918:Indricotherium turgaicum 2824:Thaumastotherium osborni 2579:(Submitted manuscript). 2273:Distribution and habitat 1251:. Though the genus name 982:In 2017, a new species, 550:belonging to the family 488:Indricotherium asiaticum 433:Thaumastotherium osborni 4383:2005E&PSL.236..322W 4109:Baluchitherium grangeri 4107:Osborn, H. F. (1923). " 3979:Antoine, P. O. (2014). 3805:Larramendi, A. (2016). 3669:Journal of Paleontology 3385:Zhan-Xiang, Q. (1973). 3355:Oxford University Press 2200:Skull and lower jaw of 2157:. As in elephants, the 1674:in the late Eocene and 1247:, as well as the genus 977:Baluchitherium grangeri 874:insect. The fossils of 591:was originally placed. 504:Indricotherium grangeri 480:Baluchitherium grangeri 4745:Oligocene rhinoceroses 3942:Tsubamoto, T. (2012). 3826:10.4202/app.00136.2014 3615:10.1006/clad.1999.0107 3513:Vertebrata PalAsiatica 3472:Vertebrata PalAsiatica 3394:Vertebrata PalAsiatica 3318:10.1098/rstb.1934.0013 3165:Metamynodon bugtiensis 3118:Communications Biology 3058:Vertebrata PalAsiatica 3010:Osborn, H. F. (1923). 2937:10.1098/rstb.1924.0002 2914:Baluchitherium osborni 2756:10.1098/rstb.1924.0009 2725:Cooper, C. F. (1924). 2477: 2405: 2289:Remains assignable to 2286: 2205: 2182:documentaries such as 2111: 2026: 1937: 1853:The largest skulls of 1758: 1729:Mammuthus trogontherii 1700: 1335: 1141:; it is possible that 1072:William Diller Matthew 1056:Baluchitherium osborni 1032: 952: 888:Henry Fairfield Osborn 570:between China and the 449:Metamynodon bugtiensis 443:(Forster-Cooper, 1913) 441:Baluchitherium osborni 425:Aceratherium bugtiense 4702:Paleobiology Database 4568:Prothero, D. (2013). 3957:10.4202/app.2011.0067 3596:Holbrook, L. (1999). 2959:Pavlova, M. (1922). " 2472: 2395: 2280: 2199: 2134:competitive exclusion 2099: 2020: 1928: 1750: 1689: 1325: 1022: 943: 933:had already named it 779:Early discoveries of 723:geological formations 715:lumping and splitting 32:Baluchitherium (song) 3854:Paul, G. S. (1997). 3563:Journal of Mammalogy 3357:, pp. 358–378, 3303:(494–508): 569–616. 2741:(391–401): 369–394. 2487:inadaptive evolution 2344:Hsanda Gol Formation 2013:Postcranial skeleton 1334:and colleagues, 2021 1015:Species and synonyms 854:("wonderful beast") 836:Clive Forster-Cooper 797:Chitarwata Formation 496:Indricotherium minus 451:Forster-Cooper, 1922 376:Forster-Cooper, 1913 4560:Paleontology portal 4473:1998Natur.394..364M 4422:2011NW.....98..407S 4410:Naturwissenschaften 4348:2011PPP...311...19M 4279:2003Oecol.136...14C 4202:2004JAESc..24...71A 3997:10.18563/pv.38.1.e4 3712:2016NatSR...639607W 3309:1934RSPTB.223..569F 3214:2018PLoSO..1393774T 2747:1924RSPTB.212..369F 2560:Manias, C. (2014). 2185:Walking With Beasts 2116:Robert M. Alexander 2029:No complete set of 1393:Leonard B. Radinsky 957:Roy Chapman Andrews 793:Guy Ellcock Pilgrim 554:. It is one of the 4141:Biological Reviews 4038:Eternity of Eagles 3700:Scientific Reports 3526:on 22 October 2014 2930:(391–401): 35–66. 2478: 2410:palaeobiogeography 2406: 2287: 2241:would have been a 2206: 2112: 2057:sauropod dinosaurs 2027: 1938: 1905:was very wide and 1759: 1701: 1690:Estimated size of 1336: 1274:did not belong in 1211:. By this scheme, 1052:William K. Gregory 1033: 953: 594:The exact size of 465:P. transouralicum: 4740:Paraceratheriidae 4727: 4726: 4689:Open Tree of Life 4593:Taxon identifiers 4579:978-0-253-00819-0 4517:978-0-691-02542-1 4467:(6691): 364–367. 3720:10.1038/srep39607 3364:978-0-19-506039-3 2506:African elephants 2495:reproduction rate 2422:Mongolian Plateau 2283:P. transouralicum 2281:Foraging herd of 2243:hindgut fermenter 2202:P. transouralicum 2102:P. transouralicum 2100:Restoration of a 2023:P. transouralicum 1962:P. transouralicum 1931:P. transouralicum 1920:P. transouralicum 1912:P. transouralicum 1903:occipital condyle 1880:Indian rhinoceros 1845:P. transouralicum 1802:P. transouralicum 1797:P. transouralicum 1764:P. transouralicum 1756:P. transouralicum 1742:Michael P. Taylor 1692:P. transouralicum 1640: 1639: 1631: 1630: 1622: 1621: 1613: 1612: 1604: 1603: 1201:P. transouralicum 1147:P. transouralicum 1139:sexual dimorphism 1131:P. transouralicum 1099:P. transouralicum 1029:P. transouralicum 963:sponsored by the 946:P. transouralicum 935:I. transouralicum 880:odd-toed ungulate 842:to the new genus 819:wastebasket taxon 719:geological dating 691:sexual dimorphism 687:P. transouralicum 679:P. transouralicum 667:P. transouralicum 552:Paraceratheriidae 535: 534: 528: 515: 507: 499: 491: 483: 475: 460: 452: 444: 436: 428: 411: 402: 393: 385: 377: 367: 348: 331: 314: 305:P. transouralicum 297: 246: 223:Paraceratheriidae 138:P. transouralicum 16:(Redirected from 4792: 4720: 4719: 4710: 4709: 4697: 4696: 4684: 4683: 4671: 4670: 4658: 4657: 4645: 4644: 4635: 4634: 4633: 4620: 4619: 4618: 4588: 4583: 4562: 4557: 4556: 4555: 4539: 4538: 4530: 4521: 4520: 4499: 4493: 4492: 4456: 4450: 4449: 4401: 4395: 4394: 4377:(1–2): 322–338. 4366: 4360: 4359: 4331: 4322: 4321: 4319: 4317: 4311: 4264: 4255: 4249: 4248: 4246: 4244: 4235:(6): 1231–1245. 4220: 4214: 4213: 4181: 4175: 4174: 4164: 4131: 4125: 4124: 4104: 4095: 4094: 4074: 4068: 4067: 4065: 4063: 4034:Unfeathered Bird 4027: 4021: 4020: 4018: 4016: 3985:Palaeovertebrata 3976: 3970: 3969: 3959: 3939: 3933: 3932: 3922: 3912: 3910:10.7717/peerj.36 3888: 3882: 3881: 3879: 3877: 3871: 3860: 3851: 3845: 3844: 3842: 3811: 3802: 3796: 3795: 3775: 3764: 3763: 3751: 3742: 3741: 3731: 3691: 3685: 3684: 3660: 3654: 3651: 3628: 3627: 3617: 3593: 3587: 3586: 3558: 3552: 3549: 3536: 3535: 3533: 3531: 3525: 3510: 3501: 3495: 3494: 3492: 3490: 3484: 3469: 3456: 3450: 3449: 3447: 3423: 3417: 3416: 3414: 3412: 3406: 3391: 3382: 3376: 3375: 3342: 3323: 3322: 3320: 3288: 3282: 3281: 3257: 3246: 3245: 3235: 3225: 3193: 3187: 3186: 3158: 3152: 3151: 3141: 3105: 3086: 3085: 3083: 3081: 3049: 3040: 3039: 3037: 3035: 3007: 3001: 2998: 2992: 2991: 2979: 2973: 2972: 2956: 2950: 2949: 2939: 2907: 2901: 2900: 2898: 2896: 2864: 2858: 2857: 2855: 2853: 2817: 2811: 2810: 2808: 2806: 2770: 2761: 2760: 2758: 2722: 2713: 2712: 2697: 2691: 2688: 2677: 2674: 2633: 2630: 2609: 2608: 2574: 2557: 2546: 2543: 2300:palaeogeographic 2227:Isotope analysis 2192:Diet and feeding 2159:gestation period 2138:white rhinoceros 1993:and the related 1958:Dzungariotherium 1929:Upper molars of 1876:black rhinoceros 1859:zygomatic arches 1838: 1829: 1820: 1777:thermoregulation 1752:Life restoration 1566: 1541: 1516: 1510:Indricotheriinae 1489: 1478: 1306:, while keeping 1284:P. tienshanensis 1268:Dzungariotherium 1245:P. tienshanensis 1233:Dzungariotherium 1169: 1160: 1122:Dzungariotherium 1087:Spencer G. Lucas 852:Thaumastotherium 753: 744: 628:gestation period 523: 510: 502: 494: 486: 478: 470: 459:Bayshashov, 1988 455: 447: 439: 431: 423: 410:Species synonymy 409: 400: 391: 383: 375: 365: 342: 336: 325: 319: 308: 302: 295: 289: 260: 241: 234: 221: 161: 160: 131: 121: 58: 47:Temporal range: 39: 21: 4800: 4799: 4795: 4794: 4793: 4791: 4790: 4789: 4730: 4729: 4728: 4723: 4715: 4713: 4705: 4700: 4692: 4687: 4679: 4674: 4666: 4661: 4653: 4648: 4640: 4638: 4631:Paraceratherium 4629: 4628: 4623: 4614: 4613: 4608: 4601:Paraceratherium 4595: 4580: 4567: 4558: 4553: 4551: 4548: 4543: 4542: 4532: 4531: 4524: 4518: 4504:Prothero, D. R. 4501: 4500: 4496: 4458: 4457: 4453: 4406:Paraceratherium 4403: 4402: 4398: 4368: 4367: 4363: 4333: 4332: 4325: 4315: 4313: 4309: 4262: 4257: 4256: 4252: 4242: 4240: 4222: 4221: 4217: 4183: 4182: 4178: 4133: 4132: 4128: 4106: 4105: 4098: 4076: 4075: 4071: 4061: 4059: 4029: 4028: 4024: 4014: 4012: 3978: 3977: 3973: 3941: 3940: 3936: 3890: 3889: 3885: 3875: 3873: 3869: 3858: 3853: 3852: 3848: 3840: 3809: 3804: 3803: 3799: 3777: 3776: 3767: 3753: 3752: 3745: 3693: 3692: 3688: 3665:Forstercooperia 3662: 3661: 3657: 3652: 3631: 3595: 3594: 3590: 3575:10.2307/1377893 3560: 3559: 3555: 3550: 3539: 3529: 3527: 3523: 3508: 3503: 3502: 3498: 3488: 3486: 3482: 3467: 3463:Paraceratherium 3458: 3457: 3453: 3425: 3424: 3420: 3410: 3408: 3404: 3389: 3384: 3383: 3379: 3365: 3344: 3343: 3326: 3290: 3289: 3285: 3259: 3258: 3249: 3208:(4): e0193774. 3195: 3194: 3190: 3177:(53): 617–620. 3160: 3159: 3155: 3112:Paraceratherium 3107: 3106: 3089: 3079: 3077: 3051: 3050: 3043: 3033: 3031: 3020:Natural History 3009: 3008: 3004: 2999: 2995: 2981: 2980: 2976: 2958: 2957: 2953: 2909: 2908: 2904: 2894: 2892: 2866: 2865: 2861: 2851: 2849: 2836:(70): 376–381. 2819: 2818: 2814: 2804: 2802: 2789:(48): 711–716. 2772: 2771: 2764: 2724: 2723: 2716: 2699: 2698: 2694: 2689: 2680: 2675: 2636: 2631: 2612: 2572: 2559: 2558: 2549: 2544: 2531: 2526: 2510:Paraceratherium 2482:Paraceratherium 2467: 2426:Tibetan Plateau 2414:Paraceratherium 2398:Paraceratherium 2369:Paraceratherium 2340:Paraceratherium 2338:The habitat of 2309:Paraceratherium 2304:Paraceratherium 2291:Paraceratherium 2275: 2259:Paraceratherium 2255:Paraceratherium 2247:Paraceratherium 2239:Paraceratherium 2233:fed chiefly on 2231:Paraceratherium 2219:Paraceratherium 2210:Paraceratherium 2194: 2172:Paraceratherium 2167:Paraceratherium 2163:Paraceratherium 2149:Paraceratherium 2129:Paraceratherium 2124:Paraceratherium 2120:Paraceratherium 2104:pair, with two 2094: 2077:Paraceratherium 2061:Paraceratherium 2035:Paraceratherium 2015: 1991:Paraceratherium 1986:Paraceratherium 1942:Paraceratherium 1940:The species of 1907:Paraceratherium 1868:Paraceratherium 1863:Paraceratherium 1855:Paraceratherium 1851: 1850: 1849: 1848: 1841: 1840: 1839: 1831: 1830: 1822: 1821: 1810: 1792:Paraceratherium 1781:Paraceratherium 1773:Paraceratherium 1738:Gregory S. Paul 1733:Paraceratherium 1717:Paraceratherium 1704:Paraceratherium 1684: 1676:Paraceratherium 1662:Forstercooperia 1649:Paraceratherium 1645:Forstercooperia 1641: 1632: 1623: 1614: 1605: 1593:Paraceratherium 1525:Forstercooperia 1472:Paraceratherium 1449:Paraceratherium 1445:Paraceratherium 1424:Forstercooperia 1389:Paraceratherium 1369:Paraceratherium 1343:Rhinocerotoidea 1328:Paraceratherium 1320: 1292:Paraceratherium 1276:Paraceratherium 1257:Paraceratherium 1227:suggested that 1225:Donald Prothero 1197:P. zhajremensis 1182: 1181: 1180: 1179: 1172: 1171: 1170: 1162: 1161: 1119:(originally in 1109:(originally in 1101:(originally in 1091:Paraceratherium 1068:morphologically 1017: 1009:Paraceratherium 884:Paraceratherium 844:Paraceratherium 789:Paraceratherium 777: 776: 775: 774: 769:(left), and an 756: 755: 754: 746: 745: 707:Paraceratherium 699: 659:Paraceratherium 655:junior synonyms 651:Pristinotherium 616:Paraceratherium 596:Paraceratherium 576:Paraceratherium 539:Paraceratherium 531: 498:Borissiak, 1923 490:Borissiak, 1923 413: 412: 405: 392:Birkjukov, 1953 389:Pristinotherium 384:Borissiak, 1916 369: 368: 296:(Pilgrim, 1908) 275: 269: 240: 237:Paraceratherium 232: 219: 155: 122: 120: 119: 114: 109: 104: 99: 94: 89: 84: 79: 74: 69: 64: 53: 52: 45: 43:Paraceratherium 35: 28: 23: 22: 15: 12: 11: 5: 4798: 4796: 4788: 4787: 4782: 4777: 4772: 4767: 4762: 4757: 4752: 4747: 4742: 4732: 4731: 4725: 4724: 4722: 4721: 4711: 4698: 4685: 4672: 4659: 4646: 4636: 4621: 4605: 4603: 4597: 4596: 4591: 4585: 4584: 4578: 4564: 4563: 4547: 4544: 4541: 4540: 4522: 4516: 4494: 4451: 4416:(5): 407–423. 4396: 4361: 4342:(1–2): 19–29. 4323: 4250: 4215: 4176: 4147:(1): 117–155. 4126: 4096: 4079:Baluchitherium 4069: 4044:, Van Duzer's 4022: 3971: 3934: 3883: 3846: 3797: 3765: 3758:(in Russian). 3743: 3686: 3675:(4): 826–841. 3655: 3629: 3608:(3): 331–350. 3588: 3569:(4): 631–639. 3553: 3537: 3496: 3461:"Discovery of 3451: 3438:(3): 581–592. 3418: 3377: 3363: 3324: 3283: 3262:Baluchitherium 3247: 3188: 3153: 3087: 3064:(4): 367–381. 3041: 3014:Baluchitherium 3002: 2993: 2974: 2951: 2920:, Borrissyak)" 2902: 2859: 2812: 2762: 2714: 2701:Pilgrim, G. E. 2692: 2678: 2634: 2610: 2568:Indricotherium 2564:Baluchitherium 2547: 2528: 2527: 2525: 2522: 2491:climate change 2466: 2463: 2450:sister species 2274: 2271: 2193: 2190: 2114:The zoologist 2093: 2090: 2014: 2011: 1972:differed from 1970:P. huangheense 1899:sagittal crest 1886:(trunk) as in 1843: 1842: 1833: 1832: 1824: 1823: 1815: 1814: 1813: 1812: 1811: 1809: 1806: 1725:steppe mammoth 1706:is one of the 1683: 1680: 1653:Indricotherium 1638: 1637: 1634: 1633: 1629: 1628: 1625: 1624: 1620: 1619: 1616: 1615: 1611: 1610: 1607: 1606: 1602: 1601: 1598: 1597: 1587: 1584: 1583: 1580: 1579: 1569: 1564: 1562: 1559: 1558: 1555: 1554: 1544: 1539: 1537: 1534: 1533: 1530: 1529: 1519: 1514: 1512: 1505: 1504: 1501: 1500: 1492: 1487: 1485: 1483:Hyracodontidae 1476: 1365:Hyracodontidae 1361:Rhinocerotidae 1319: 1316: 1312:Baluchitherium 1308:Indricotherium 1264:Indricotherium 1253:Indricotherium 1241:I. intermedium 1217:D. turfanensis 1174: 1173: 1164: 1163: 1155: 1154: 1153: 1152: 1151: 1135:Indricotherium 1103:Indricotherium 1076:Baluchitherium 1060:junior synonym 1048:Walter Granger 1016: 1013: 988:Gansu Province 984:P. huangheense 919:Indricotherium 907:Aral Formation 890:, after which 876:Baluchitherium 864:Baluchitherium 817:was by then a 758: 757: 748: 747: 739: 738: 737: 736: 735: 698: 695: 689:may be due to 671:P. huangheense 647:Indricotherium 643:Baluchitherium 548:rhinocerotoids 542:is an extinct 533: 532: 530: 529: 521: 519:Dubious names: 516: 514:Birjukov, 1953 508: 506:(Osborn, 1923) 500: 492: 484: 476: 468: 461: 453: 445: 437: 429: 421: 408: 407: 406: 404: 403: 394: 386: 381:Indricotherium 378: 373:Baluchitherium 366:Genus synonymy 364: 363: 362: 359: 358: 352: 351: 350: 349: 332: 322:P. huangheense 315: 298: 282: 281: 277: 276: 270: 255: 254: 248: 247: 243:Forster-Cooper 230: 226: 225: 217: 213: 212: 210:Perissodactyla 207: 203: 202: 197: 193: 192: 187: 183: 182: 177: 173: 172: 167: 163: 162: 149: 148: 133: 132: 124: 123: 115: 110: 105: 100: 95: 90: 85: 80: 75: 70: 65: 60: 59: 46: 26: 24: 18:Indricotherium 14: 13: 10: 9: 6: 4: 3: 2: 4797: 4786: 4783: 4781: 4778: 4776: 4773: 4771: 4768: 4766: 4763: 4761: 4758: 4756: 4753: 4751: 4748: 4746: 4743: 4741: 4738: 4737: 4735: 4718: 4712: 4708: 4703: 4699: 4695: 4690: 4686: 4682: 4677: 4673: 4669: 4664: 4660: 4656: 4651: 4647: 4643: 4637: 4632: 4626: 4622: 4617: 4611: 4607: 4606: 4604: 4602: 4598: 4594: 4589: 4581: 4575: 4571: 4566: 4565: 4561: 4550: 4545: 4536: 4529: 4527: 4523: 4519: 4513: 4509: 4505: 4498: 4495: 4490: 4486: 4482: 4481:10.1038/28603 4478: 4474: 4470: 4466: 4462: 4455: 4452: 4447: 4443: 4439: 4435: 4431: 4427: 4423: 4419: 4415: 4411: 4407: 4400: 4397: 4392: 4388: 4384: 4380: 4376: 4372: 4365: 4362: 4357: 4353: 4349: 4345: 4341: 4337: 4330: 4328: 4324: 4308: 4304: 4300: 4296: 4292: 4288: 4284: 4280: 4276: 4272: 4268: 4261: 4254: 4251: 4238: 4234: 4230: 4229:Palaeontology 4226: 4219: 4216: 4211: 4207: 4203: 4199: 4195: 4191: 4187: 4180: 4177: 4172: 4168: 4163: 4158: 4154: 4150: 4146: 4142: 4138: 4130: 4127: 4122: 4118: 4114: 4110: 4103: 4101: 4097: 4092: 4088: 4084: 4080: 4073: 4070: 4057: 4053: 4049: 4047: 4043: 4039: 4035: 4026: 4023: 4010: 4006: 4002: 3998: 3994: 3990: 3986: 3982: 3975: 3972: 3967: 3963: 3958: 3953: 3949: 3945: 3938: 3935: 3930: 3926: 3921: 3916: 3911: 3906: 3902: 3898: 3894: 3887: 3884: 3868: 3864: 3857: 3850: 3847: 3839: 3835: 3831: 3827: 3823: 3819: 3815: 3808: 3801: 3798: 3793: 3789: 3785: 3781: 3774: 3772: 3770: 3766: 3761: 3757: 3750: 3748: 3744: 3739: 3735: 3730: 3725: 3721: 3717: 3713: 3709: 3705: 3701: 3697: 3690: 3687: 3682: 3678: 3674: 3670: 3666: 3659: 3656: 3650: 3648: 3646: 3644: 3642: 3640: 3638: 3636: 3634: 3630: 3625: 3621: 3616: 3611: 3607: 3603: 3599: 3592: 3589: 3584: 3580: 3576: 3572: 3568: 3564: 3557: 3554: 3548: 3546: 3544: 3542: 3538: 3522: 3518: 3514: 3507: 3500: 3497: 3481: 3477: 3473: 3466: 3464: 3455: 3452: 3446: 3441: 3437: 3433: 3429: 3422: 3419: 3403: 3399: 3395: 3388: 3381: 3378: 3374: 3370: 3366: 3360: 3356: 3352: 3348: 3341: 3339: 3337: 3335: 3333: 3331: 3329: 3325: 3319: 3314: 3310: 3306: 3302: 3298: 3294: 3287: 3284: 3279: 3275: 3271: 3267: 3263: 3256: 3254: 3252: 3248: 3243: 3239: 3234: 3229: 3224: 3219: 3215: 3211: 3207: 3203: 3199: 3192: 3189: 3184: 3180: 3176: 3172: 3168: 3166: 3157: 3154: 3149: 3145: 3140: 3135: 3131: 3127: 3123: 3119: 3115: 3113: 3104: 3102: 3100: 3098: 3096: 3094: 3092: 3088: 3075: 3071: 3067: 3063: 3059: 3055: 3048: 3046: 3042: 3029: 3025: 3021: 3017: 3015: 3006: 3003: 2997: 2994: 2989: 2986:(in German). 2985: 2978: 2975: 2970: 2967:(in French). 2966: 2962: 2955: 2952: 2947: 2943: 2938: 2933: 2929: 2925: 2921: 2919: 2915: 2906: 2903: 2890: 2886: 2882: 2878: 2874: 2870: 2863: 2860: 2847: 2843: 2839: 2835: 2831: 2827: 2825: 2816: 2813: 2800: 2796: 2792: 2788: 2784: 2780: 2778: 2769: 2767: 2763: 2757: 2752: 2748: 2744: 2740: 2736: 2732: 2730: 2721: 2719: 2715: 2710: 2706: 2702: 2696: 2693: 2687: 2685: 2683: 2679: 2673: 2671: 2669: 2667: 2665: 2663: 2661: 2659: 2657: 2655: 2653: 2651: 2649: 2647: 2645: 2643: 2641: 2639: 2635: 2629: 2627: 2625: 2623: 2621: 2619: 2617: 2615: 2611: 2606: 2602: 2598: 2594: 2590: 2586: 2583:(2): 237–78. 2582: 2578: 2571: 2569: 2565: 2556: 2554: 2552: 2548: 2542: 2540: 2538: 2536: 2534: 2530: 2523: 2521: 2519: 2518:early Miocene 2515: 2511: 2507: 2503: 2498: 2496: 2492: 2488: 2483: 2476: 2471: 2464: 2462: 2460: 2455: 2451: 2447: 2446:P. linxiaense 2443: 2439: 2435: 2431: 2427: 2423: 2419: 2415: 2411: 2403: 2399: 2394: 2390: 2388: 2384: 2383: 2378: 2375:and abundant 2374: 2370: 2366: 2362: 2358: 2354: 2350: 2345: 2341: 2336: 2334: 2330: 2326: 2322: 2318: 2314: 2310: 2305: 2301: 2297: 2292: 2284: 2279: 2272: 2270: 2268: 2264: 2260: 2256: 2250: 2248: 2244: 2240: 2236: 2232: 2228: 2224: 2220: 2216: 2211: 2203: 2198: 2191: 2189: 2187: 2186: 2181: 2177: 2176:biomechanical 2173: 2168: 2164: 2160: 2156: 2155: 2154:Astorgosuchus 2150: 2146: 2141: 2139: 2135: 2130: 2125: 2121: 2117: 2109: 2108: 2103: 2098: 2092:Palaeobiology 2091: 2089: 2086: 2082: 2078: 2073: 2068: 2066: 2062: 2058: 2054: 2049: 2048:neural spines 2045: 2040: 2036: 2032: 2024: 2021:Hind foot of 2019: 2012: 2010: 2008: 2004: 2000: 1996: 1995:Urtinotherium 1992: 1987: 1982: 1979: 1978:P. linxiaense 1975: 1971: 1967: 1966:frontal bones 1963: 1959: 1955: 1951: 1947: 1943: 1936: 1932: 1927: 1923: 1921: 1917: 1913: 1908: 1904: 1900: 1896: 1891: 1889: 1885: 1882:, or a short 1881: 1877: 1873: 1869: 1864: 1860: 1856: 1846: 1837: 1828: 1819: 1807: 1805: 1803: 1798: 1793: 1788: 1786: 1782: 1778: 1774: 1768: 1765: 1757: 1753: 1749: 1745: 1743: 1739: 1734: 1730: 1726: 1722: 1721:proboscideans 1718: 1714: 1709: 1705: 1699: 1698: 1693: 1688: 1681: 1679: 1677: 1673: 1672:Urtinotherium 1669: 1668: 1663: 1659: 1654: 1650: 1646: 1636: 1635: 1627: 1626: 1618: 1617: 1609: 1608: 1600: 1599: 1596: 1595: 1594: 1586: 1585: 1582: 1581: 1578: 1577: 1576: 1575:Urtinotherium 1568: 1567: 1561: 1560: 1557: 1556: 1553: 1552: 1551: 1543: 1542: 1536: 1535: 1532: 1531: 1528: 1527: 1526: 1518: 1517: 1511: 1507: 1506: 1503: 1502: 1499: 1498: 1497:Triplopodinae 1491: 1490: 1484: 1480: 1479: 1475: 1473: 1469: 1464: 1462: 1458: 1454: 1450: 1446: 1442: 1441: 1440:Urtinotherium 1436: 1435: 1430: 1426: 1425: 1420: 1416: 1415: 1410: 1406: 1402: 1398: 1394: 1390: 1385: 1383: 1378: 1374: 1370: 1366: 1362: 1358: 1357:Amynodontidae 1354: 1353: 1348: 1344: 1341: 1333: 1329: 1324: 1317: 1315: 1313: 1309: 1305: 1301: 1297: 1293: 1289: 1285: 1281: 1277: 1273: 1269: 1265: 1260: 1258: 1254: 1250: 1249:Benaratherium 1246: 1242: 1238: 1234: 1230: 1229:P. orgosensis 1226: 1222: 1218: 1214: 1213:P. orgosensis 1210: 1206: 1202: 1198: 1194: 1190: 1187: 1178: 1177:P. linxiaense 1168: 1159: 1150: 1148: 1144: 1140: 1136: 1132: 1128: 1127:dubious names 1124: 1123: 1118: 1117:P. orgosensis 1114: 1113: 1108: 1104: 1100: 1096: 1092: 1088: 1083: 1081: 1077: 1073: 1069: 1065: 1061: 1058:was likely a 1057: 1053: 1049: 1045: 1041: 1039: 1030: 1026: 1021: 1014: 1012: 1010: 1005: 1001: 1000:P. linxiaense 997: 993: 992:Huanghe River 989: 985: 980: 978: 974: 970: 966: 962: 958: 951: 947: 942: 938: 936: 932: 931:Maria Pavlova 928: 924: 920: 916: 912: 908: 904: 899: 897: 893: 889: 885: 881: 877: 873: 869: 865: 861: 857: 853: 849: 845: 841: 837: 832: 830: 829: 824: 820: 816: 812: 808: 807: 802: 798: 794: 790: 786: 782: 781:indricotheres 772: 768: 767: 762: 752: 743: 734: 732: 728: 724: 720: 716: 712: 708: 704: 696: 694: 692: 688: 684: 680: 676: 675:P. linxiaense 672: 668: 664: 660: 656: 652: 648: 644: 640: 635: 633: 629: 625: 621: 617: 613: 610:upper lip or 609: 605: 601: 597: 592: 590: 587: 583: 582: 577: 573: 569: 565: 561: 557: 553: 549: 545: 541: 540: 527:Gabunia, 1955 526: 522: 520: 517: 513: 509: 505: 501: 497: 493: 489: 485: 481: 477: 474:Pavlova, 1922 473: 469: 467: 466: 462: 458: 454: 450: 446: 442: 438: 434: 430: 427:Pilgrim, 1908 426: 422: 420: 419: 418:P. bugtiense: 415: 401:Gabunia, 1955 399: 398:Benaratherium 395: 390: 387: 382: 379: 374: 371: 360: 357: 353: 346: 341: 340: 339:P. linxiaense 333: 329: 324: 323: 316: 312: 307: 306: 299: 294: 293: 286: 285: 283: 278: 273: 268: 267: 265: 256: 253: 249: 244: 239: 238: 231: 228: 227: 224: 218: 215: 214: 211: 208: 205: 204: 201: 198: 195: 194: 191: 188: 185: 184: 181: 178: 175: 174: 171: 168: 165: 164: 159: 154: 150: 147: 143: 139: 134: 130: 125: 118: 113: 108: 103: 98: 93: 88: 83: 78: 73: 68: 63: 57: 50: 44: 40: 37: 33: 19: 4600: 4569: 4546:Bibliography 4534: 4507: 4497: 4464: 4460: 4454: 4413: 4409: 4405: 4399: 4374: 4370: 4364: 4339: 4335: 4314:. 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