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''Iranotherium'' ("Beast of
Iran Iran, officially the Islamic Republic of Iran, and also called Persia, is a country located in Western Asia. It is bordered by Iraq and Turkey to the west, by Azerbaijan and Armenia to the northwest, by the Caspian Sea and Turkmeni ...
") was a genus of large
elasmotheriine A rhinoceros (; ; ), commonly abbreviated to rhino, is a member of any of the five extant species (or numerous extinct species) of odd-toed ungulates in the family Rhinocerotidae. (It can also refer to a member of any of the extinct species o ...
rhinoceros A rhinoceros (; ; ), commonly abbreviated to rhino, is a member of any of the five extant species (or numerous extinct species) of odd-toed ungulates in the family Rhinocerotidae. (It can also refer to a member of any of the extinct specie ...
, comparable in
size Size in general is the magnitude or dimensions of a thing. More specifically, ''geometrical size'' (or ''spatial size'') can refer to linear dimensions ( length, width, height, diameter, perimeter), area, or volume. Size can also be me ...
to a modern
white rhino The white rhinoceros, white rhino or square-lipped rhinoceros (''Ceratotherium simum'') is the largest extant species of rhinoceros. It has a wide mouth used for grazing and is the most social of all rhino species. The white rhinoceros consists ...
. It is known from the
Late Miocene The Late Miocene (also known as Upper Miocene) is a sub-epoch of the Miocene Epoch made up of two stages. The Tortonian and Messinian stages comprise the Late Miocene sub-epoch, which lasted from 11.63 Ma (million years ago) to 5.333 Ma. The ...
( Tortonian) of
Maragha Maragheh ( fa, مراغه, Marāgheh or ''Marāgha''; az, ماراغا ) is a city and capital of Maragheh County, East Azerbaijan Province, Iran. Maragheh is on the bank of the river Sufi Chay. The population consists mostly of Iranian Azerba ...
,
Iran Iran, officially the Islamic Republic of Iran, and also called Persia, is a country located in Western Asia. It is bordered by Iraq and Turkey to the west, by Azerbaijan and Armenia to the northwest, by the Caspian Sea and Turkmeni ...
and the middle part of the Liushu formation of
northwestern China Northwest China () is a statistical region of China which includes the autonomous regions of Xinjiang and Ningxia and the provinces of Shaanxi, Gansu and Qinghai. It has an area of 3,107,900 km2. The region is characterized by a (semi-)arid con ...
. It was a precursor to the related ''
Sinotherium ''Sinotherium'' ("Chinese Beast") is a genus of single- horned elasmotheriine rhinoceros that lived from the late Miocene (Tortonian - Messinian) to Early Pliocene. It was ancestral to ''Elasmotherium,'' demonstrating a very important evolutiona ...
'' and may have been ultimately outcompeted by its descendant. This species is most well known for showing unique
sexual dimorphism Sexual dimorphism is the condition where the sexes of the same animal and/or plant species exhibit different morphological characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most an ...
among rhinos.


Description

''Iranotherium'' is only known from
cranial Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position prov ...
material, and rare
postcrania Postcrania (postcranium, adjective: postcranial) in zoology and vertebrate paleontology is all or part of the skeleton apart from the skull. Frequently, fossil remains, e.g. of dinosaurs or other extinct tetrapods, consist of partial or isolated sk ...
l elements.


Skull and Horn

The
skull The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, th ...
is long and dorsally concave, with a length of 775 mm. The parietal crests are broad, the
nasal bone The nasal bones are two small oblong bones, varying in size and form in different individuals; they are placed side by side at the middle and upper part of the face and by their junction, form the bridge of the upper one third of the nose. Ea ...
is long and wide, with a shallow U-shaped nasal notch, the
orbit In celestial mechanics, an orbit is the curved trajectory of an object such as the trajectory of a planet around a star, or of a natural satellite around a planet, or of an artificial satellite around an object or position in space such as ...
is prominent and has a high position close to the
skull The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, th ...
roof, the first half of the
zygomatic arch In anatomy, the zygomatic arch, or cheek bone, is a part of the skull formed by the zygomatic process of the temporal bone (a bone extending forward from the side of the skull, over the opening of the ear) and the temporal process of the zygo ...
is narrow. The
skull The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, th ...
s also show distinct
sexual dimorphism Sexual dimorphism is the condition where the sexes of the same animal and/or plant species exhibit different morphological characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most an ...
: The base from which the horn grows is large and strong in the
male Male (symbol: ♂) is the sex of an organism that produces the gamete (sex cell) known as sperm, which fuses with the larger female gamete, or ovum, in the process of fertilization. A male organism cannot reproduce sexually without access to ...
, while it is small and weak in the female, and there is strong and rough rugosities present on the back half part of the
zygomatic arch In anatomy, the zygomatic arch, or cheek bone, is a part of the skull formed by the zygomatic process of the temporal bone (a bone extending forward from the side of the skull, over the opening of the ear) and the temporal process of the zygo ...
of the
male Male (symbol: ♂) is the sex of an organism that produces the gamete (sex cell) known as sperm, which fuses with the larger female gamete, or ovum, in the process of fertilization. A male organism cannot reproduce sexually without access to ...
. The back half of the zygoma is also uplifted and higher than the skull roof.


Dentition

''Iranotherium'' teeth are hypsodont, (they were covered and filled by
cement A cement is a binder, a chemical substance used for construction that sets, hardens, and adheres to other materials to bind them together. Cement is seldom used on its own, but rather to bind sand and gravel (aggregate) together. Cement mixe ...
), meaning that ''Iranotherium'' was a grazer that fed on grasses. Its upper
incisor Incisors (from Latin ''incidere'', "to cut") are the front teeth present in most mammals. They are located in the premaxilla above and on the mandible below. Humans have a total of eight (two on each side, top and bottom). Opossums have 18, wher ...
s are completely absent and the
premolar The premolars, also called premolar teeth, or bicuspids, are transitional teeth located between the canine and molar teeth. In humans, there are two premolars per quadrant in the permanent set of teeth, making eight premolars total in the mouth ...
s are significantly shortened, however, the
cheek teeth Cheek teeth or post-canines comprise the molar and premolar teeth in mammals. Cheek teeth are multicuspidate (having many folds or tubercles). Mammals have multicuspidate molars (three in placentals, four in marsupials, in each jaw quadrant) and p ...
are longer in the male than in the female.


Discovery and History


Original Discovery

When ''Iranotherium'' was originally discovered from
teeth A tooth ( : teeth) is a hard, calcified structure found in the jaws (or mouths) of many vertebrates and used to break down food. Some animals, particularly carnivores and omnivores, also use teeth to help with capturing or wounding prey, ...
, it was recognized that this
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriat ...
was similar to the rhino ''
Elasmotherium ''Elasmotherium'' is an extinct genus of large rhinoceros endemic to Eurasia during Late Miocene through the Pleistocene, existing at least as late as 39,000 years ago in the Late Pleistocene. A more recent date of 26,000 BP is considered ...
'' and was the
ancestor An ancestor, also known as a forefather, fore-elder or a forebear, is a parent or ( recursively) the parent of an antecedent (i.e., a grandparent, great-grandparent, great-great-grandparent and so forth). ''Ancestor'' is "any person from w ...
of the latter (This was before ''
Sinotherium ''Sinotherium'' ("Chinese Beast") is a genus of single- horned elasmotheriine rhinoceros that lived from the late Miocene (Tortonian - Messinian) to Early Pliocene. It was ancestral to ''Elasmotherium,'' demonstrating a very important evolutiona ...
'' was discovered), however in 1908, when it was published as a new species, it was strangely referred to the genus ''Rhinoceros''. In 1924, the material of ''R. morgani'' from Maragha was further described. Ringström suggested that this species did not belong in ''Rhinoceros'', referring it to his new genus ''Iranotherium'' as ''I. morgani''. ''Iranotherium'' as a genus was originally described in 1908 by Mecquenem where no
holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of seve ...
was designated for the genus. Specimen MNHN 1905–10, and atlas from Maragha, Iran was chosen as the Lectotype for this genus.


Later Discoveries

In 2005, three specimens of ''Iranotherium'' were described by Tao Deng. Two of these specimens were complete skulls that showed evidence of unique sexual dimorphism. These specimens are: HMV 0979, a complete young male skull, HMV 1098, a complete adult female skull and finally HMV 1099, known only from a mandible belonging to an adult that is missing the ascending ramus. All specimens were found from the lower part of the upper Miocene Liushu Formation in the Linxia Basin (Gansu, China). HMV 0979 and HMV 1099 were discovered at Houshan village, and HMV 1098 at Shanzhuang village. The studied skulls and mandible from the Linxia Basin, especially HMV 0979 and HMV 1099, are found to be generally similar to the skull and mandible of Iranotherium morgani from Maragha, with minor differences, like the length ratio of lower premolars to molars. This difference is considered to be due to individual or sexual variations and is not sufficient for establishing a new species. Hence, all three specimens described have been assigned to the genus Iranotherium by Deng. Based on the two skulls from the study, the Maragha specimen is identified as a male due to the presence of hemispherical rugosities on its zygoma.


Paleobiology


Feeding

Iranotherium was a grazer since its denition is that of a hypsodont. In life it would've had wide and flat lips like those of the extant White Rhino. The absolute body size of an herbivore is very important in determining the fiber/protein ratio that an animal will be able to tolerate in its diet, as larger animals require proportionately less protein and will be able to tolerate a larger proportion of cellulose. Therefore, the giant body size of I. morgani implies that this species grazed high-fiber grasses. I. morgani obviously inhabited an open grassland.


Sexual variation

In 2005, two skulls of Iranotherium were described by Tao Deng. One skull was from an adult female Iranotherium and the other from a young adult male. These skulls, when compared showed significant, sexually dimorphic differences. The male had rugosities present on each zygomatic arch, whereas the females lacked them entirely. This unique feature is virtually unknown among other rhinocerotids. Male Iranotherium are hypothesized to bear it for supporting larger masseteric and temporalis musculature. Additional sexually dimorphic features include the zygomatic arch of the male being thicker than that of the female. The anterior part of the nasals are wider and rougher in the male, but narrow and smooth in the female; this difference is related to the huge horn boss in the male and the smaller horn boss in the female. Because the nasal horn boss of the male is a large dome, its nasal groove is distinctly deeper than that of the female. The nasal horn boss of the male is much rougher and likely to have supported a larger horn than that of the female. The skull roof of the male is less curved than that of the female, and the frontal region of the male is more convex than that of the female. Therefore, the overall structure of the whole skull of the male is more robust than that of the female. These features, especially the huge nasal horn, could be used for defense or competition for mates. The zygomatic rugosities of the male I. morgani could have supported a horn-like callused structure for combat or for display. Because these hypertrophies are different from the horn boss rugosities associated with the true horns in rhinos, they are probably indicative of the presence of larger masseteric and temporal muscles and the overall robustness of the skull.


Distribution and Paleoecology

''Iranotherium'' is so far only known from Iran and China. It lived in a forested environment in Iran whereas it occupied a more open habitat in China. In Iran, ''Iranotherium'' is known from the middle local biozones of the Maragha fauna, from an age of between 9 and 7.6 Ma. Over there, it coexisted with ''Hipparion'' ''prostylum'', '' Mesopithecus pentelici'' and '' Indarctos maraghanus'' In China, ''Iranotherium'' is known from the middle part of the Liushu formation of the Linxia Basin. ''I. morgani'' appeared in Linxia earlier than in Maragha. ''I. morgani'' is likely to have first appeared in northwestern China and then dispersed westward to central Asia. The
fauna Fauna is all of the animal life present in a particular region or time. The corresponding term for plants is '' flora'', and for fungi, it is '' funga''. Flora, fauna, funga and other forms of life are collectively referred to as '' biota''. ...
that ''Iranotherium'' was a part of is known as the Dashengou fauna. It was contemporary with one glire ( ''Pararhizomys hipparionum''), four mustelids ('' Promephitis sp.'', '' P. hootoni, Melodon majori, Sinictis sp.''), four
hyaenids Hyenas, or hyaenas (from Ancient Greek , ), are feliform carnivoran mammals of the family Hyaenidae . With only four extant species (each in its own genus), it is the fifth-smallest family in the Carnivora and one of the smallest in the clas ...
('' Ictitherium sp., Hyaenictitherium wongii, H. hyaenoides,
Dinocrocuta gigantea ''Dinocrocuta'' is an extinct genus of hyena-like feliform carnivores. It lived in Asia, and Africa, during the Miocene epoch. It had very strong jaws that were able to crush bones. It is estimated that their weight was around and their height ...
''), two
cats The cat (''Felis catus'') is a domestic species of small carnivorous mammal. It is the only domesticated species in the family Felidae and is commonly referred to as the domestic cat or house cat to distinguish it from the wild members o ...
( ''Amphimachairodus giganteus'', '' Felis sp.''), one
proboscidea The Proboscidea (; , ) are a taxonomic order of afrotherian mammals containing one living family ( Elephantidae) and several extinct families. First described by J. Illiger in 1811, it encompasses the elephants and their close relatives. Fr ...
n ( Tetralophodon exoletus), four perissodactyls ('' Hipparion chiai, H. weihoensis, Acerorhinus hezhengensis, Chilotherium wimani''), seven artiodactyls ('' Chleuastochoerus stehlini, Dicrocerus sp., Samotherium sp., Honanotherium schlosseri, Gazella sp., Hezhengia bohlini, and Miotragocerus sp.'') Based on pollen analysis, the vegetation of the Liushu Formation belonged to an arid steppe.


References


Bibliography

*Deng, T. (2005). New discovery of ''Iranotherium morgani'' (Perissodactyla, Rhinocerotidae) from the late Miocene of the Linxia Basin in Gansu, China, and its sexual dimorphism. Journal of vertebrate Paleontology, 25(2), 442–450. * Miocene rhinoceroses Miocene mammals of Asia Fossil taxa described in 1924 Prehistoric monotypic mammal genera Fossils of Iran {{paleo-oddtoedungulate-stub