Paraceratherium

Paraceratherium Temporal range:
Skeleton of Paraceratherium transouralicum
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Order:	Perissodactyla
Family:	†Hyracodontidae
Subfamily:	†Indricotheriinae Borissiak, 1923
Genus:	†Paraceratherium Forster Cooper, 1911
Species
P. bugtiense P. orgosense P. prohorovi P. transouralicum P. zhajremensis
Synonyms
Baluchitherium Forster Cooper, 1913 Indricotherium Borissiak, 1916 Thaumastotherium Forster Cooper, 1913 Aralotherium Borissiak, 1939 Dzungariotherium Xu and Wang, 1973

Paraceratherium, also commonly known as Indricotherium or Baluchitherium (see taxonomic discussion below), is an extinct genus of gigantic hornless rhinoceros-like mammals of the family Hyracodontidae, endemic to Eurasia and Asia during the Eocene to Oligocene 37.2—23.030 Mya, existing for approximately 14.17 million years. It was first discovered in the Baluchistan province of Pakistan, hence the name, by Sir Clive Forster Cooper.

Background

Restoration

Paraceratherium is the largest land mammal known, larger than the largest species of mammoths (Mammuthus sungari, which may have approached it in size and weight). It is also known as the "giraffe rhinoceros". Adult Paraceratherium are estimated to have been 5.5 metres (18 ft) tall at the shoulder, 9 metres (30 ft) in length from nose to rump, a maximum raised head height of about 8 metres (26 ft), and a skull length of 1.5 metres (4.9 ft). Weight estimates vary greatly, but most realistic and reliable weight estimates are about 20 tonnes. This puts it in the weight range of some medium-sized sauropod dinosaurs.

It was a herbivore that stripped leaves from trees with its down-pointing, tusk-like upper teeth that occluded forward-pointing lower teeth. It had a long, low, hornless skull and vaulted frontal and nasal bones. Its front teeth were reduced to a single pair of incisors in either jaw, but they were conical and so large that they looked like small tusks. The upper incisors pointed straight downwards, while the lower ones jutted outwards. The upper lip was evidently extremely mobile. The neck was very long, the trunk robust, and the limbs long and thick, column-like.

Its type of dentition, its mobile upper lip and its long legs and neck indicate that it was a browser that lived on the leaves and twigs of trees and large shrubs.

Taxonomy

While more distinct at the species level, there is uncertainty and disagreement with regard to the genus level of taxonomy.

Paraceratherium was first described by Forster Cooper in 1911. The genus Baluchitherium was first described by Forster Cooper in 1913. The genus Indricotherium was first described by Borissiak in 1915.

Baluchitherium is now widely regarded as a synonym of (i.e. the same as) either Paraceratherium or Indricotherium.

File:Paraceratherium size.jpg

P. bugtiense compared to a man

However, there has been disagreement over whether Indricotherium is a distinct genus from Paraceratherium. Lucas and Sobus in their 1989 review of the subfamily Indricotheriinae (see reference below), argue for synonymy, and consider that the differences between the two are of species level at most, and may even be the result of sexual dimorphism in a single species, with the larger more robust Indricotherium with larger incisors being probably the male, and the more gracile Paraceratherium the female. Others, however, have expressed doubts about this (concerning the interpretation of the shape of the skull). Even if these two do turn out to be distinct genera, they would still be similar in size and appearance.

If they are considered the same genus, then Indricotherium would become a junior synonym of Paraceratherium, because, according to the priority principle of scientific classification, the first publication, and hence the oldest valid name, takes priority and the name Paraceratherium predates the other.

Here Lucas and Sobus are followed. They consider Indricotherium, Baluchitherium, Thaumastotherium Forster Cooper, 1913a, Aralotherium Borissiak, 1939, and Dzungariotherium Xu and Wang, 1973 all as junior synonyms of Paraceratherium.

Paraceratherium transouralicum skull

Lucas and Sobus recognise four valid species of Paraceratherium. One more (P. zhajremensis) has been tentatively added. The Paraceratherium species are:

Skull of Paraceratherium bugtiense.

Paraceratherium bugtiense (Pilgrim, 1908) from the Oligocene of Pakistan is the type species of Paraceratherium. Baluchitherium osborni Forster Cooper, 1913a is a junior synonym. It was first found in the Chitarwata Formation of the Bugti Hills, Balochistan, after which it was originally named. New specimens of P. bugtiense were unearthed in the last decade by a French-Pakistani team (Antoine et al., 2004; Métais et al., 2009).

Paraceratherium transouralicum (Pavlova, 1922). Also known as Indricotherium transouralicum, this is the best known and most widespread species, known from the middle and late Oligocene of Kazakhstan, Mongolia, and Nei Monggol in northern China. Lucas and Sobus list the following species as synonyms: Baluchitherium grangeri Osborn, 1923, Indricotherium asiaticum Borissiak, 1923, Indricotherium minus Borissiak, 1923.

Paraceratherium orgosensis (Chiu, 1973) is the largest species, the teeth being at least a quarter again as big as P. transouralicum (see Lucas and Sobus p. 363/fig.19.2). It is known from the middle and late Oligocene of Xinjiang, northwest China. The three synonyms are Dzungariotherium orgosensis Chiu, 1973 and (each of the following named after a separate skull) Dzungariotherium turfanensis Xu & Wang, 1978 and Paraceratherium lipidus Xu & Wang, 1978. While there is some variation in details of the proportions of the skull (perhaps due to sexual dimorphism), all occur in a close geographical region and have distinct first and second upper molar crochets.

Paraceratherium prohorovi (Borissiak, 1939) from the late Oligocene or early Miocene of eastern Kazakhstan.

Paraceratherium zhajremensis (Osborn, 1923) from the Middle and late Oligocene of India.

Other remains referable to Paraceratherium or related taxa were found in Southeastern Europe and Asia Minor, notably in Turkey (Antoine et al., 2008).

Paraceratherium means "near horn animal" in old Greek, indicating the fact that is was hornless, but related to the rhinoceroses.

Indricotherium is named after a mythical Russian beast called the Indrik-Beast, considered the most powerful creature and the father of the animals.

Baluchitherium means "beast of Baluchistan", as it was first discovered in Baluchistan, in eastern Iran.

In popular culture

File:Indricothere CAS.jpg

Reconstruction at the California Academy of Sciences

Paraceratherium was the main focus in the third episode of the popular BBC documentary Walking with Beasts. In the programme the narrator always calls the creature an Indricothere, not a specific genus. Although in the accompanying book's information box about the animal, it states that the Indricothere is called Paraceratherium.

The creature is featured in Ratha's Courage, the fifth book of the Clan Ground series by Clare Bell.

The creature's name is also given to the title of an instrumental track from Van Halen's album, Balance.

References

• Antoine P.-O., Shah, S.M.I., Cheema, I.U., Crochet, J.-Y., de Franceschi, D., Marivaux, L., Métais, G., Welcomme, J.-L. (2004). New remains of the baluchithere Paraceratherium bugtiense (Pilgrim, 1910) from the Late/latest Oligocene of the Bugti Hills, Balochistan, Pakistan. Journal of Asian Earth Sciences 24: 71–77.
• Antoine, P.-O., Karadenizli, L., Saraç, G., Sen, S. (2008). A giant rhinocerotoid (Mammalia, Perissodactyla) from the Late Oligocene of north-central Anatolia (Turkey). Zoological Journal of the Linnean Society, London, 152: 581-592.
• Métais, G., Antoine, P.-O., Baqri, S.R.H., Marivaux, L., Welcomme, J.-L. (2009). Lithofacies, depositional environments, regional biostratigraphy, and age of the Chitarwata Formation in the Bugti Hills, Balochistan, Pakistan. Journal of Asian Earth Sciences, 34, 154-167.
• Lucas, S. G. & Sobus, J. C. (1989), The Systematics of Indricotheres. 358-378 in Prothero, D. R. & Schoch, R. M., (eds.) 1989: The Evolution of Perissodactyls, Oxford University Press, New York, New York & Oxford, England, ix-537 - argues that Indricotherium should be included under Paraceratherium
• "Indricotherium." Encyclopædia Britannica. 2009. Encyclopædia Britannica Online. 06 Apr. 2009 .
• Taggart, S. & Starr, E. (2009). Biology The Unit and Diversity of Life. Twelfth Edition. Cengage, Belmont, CA.

External links

• Indricotherium
• Discovery Channel - Answers from Dr. Mikael Fortelius
• Hyracodontidae: Indricotheriinae and Synonyms of Paraceratherium transouralicum - Mikko's Phylogeny Archive - follows Lucas and Sorbus
• sci.bio.paleontology newsgroup 20 July 1999
• "Indricotherium" Encyclopædia Britannica