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''Iranotherium'' ("Beast of
Iran Iran, officially the Islamic Republic of Iran (IRI) and also known as Persia, is a country in West Asia. It borders Iraq to the west, Turkey, Azerbaijan, and Armenia to the northwest, the Caspian Sea to the north, Turkmenistan to the nort ...
") is an extinct genus of large elasmotheriine
rhinocerotid A rhinoceros ( ; ; ; : rhinoceros or rhinoceroses), commonly abbreviated to rhino, is a member of any of the five extant taxon, extant species (or numerous extinct species) of odd-toed ungulates (perissodactyls) in the family (biology), famil ...
s, comparable in
size Size in general is the Magnitude (mathematics), magnitude or dimensions of a thing. More specifically, ''geometrical size'' (or ''spatial size'') can refer to three geometrical measures: length, area, or volume. Length can be generalized ...
to a modern
white rhino The white rhinoceros, also known as the white rhino or square-lipped rhinoceros (''Ceratotherium simum''), is the largest extant species of rhinoceros and the most Sociality, social of all rhino species, characterized by its wide mouth adapted f ...
. It is known from the
Late Miocene The Late Miocene (also known as Upper Miocene) is a sub-epoch of the Miocene epoch (geology), Epoch made up of two faunal stage, stages. The Tortonian and Messinian stages comprise the Late Miocene sub-epoch, which lasted from 11.63 Ma (million ye ...
(
Tortonian The Tortonian is in the geologic time scale an age or stage of the late Miocene that spans the time between 11.608 ± 0.005 Ma and 7.246 ± 0.005 Ma (million years ago). It follows the Serravallian and is followed by the Messinian. The Tort ...
) of Maragha,
Iran Iran, officially the Islamic Republic of Iran (IRI) and also known as Persia, is a country in West Asia. It borders Iraq to the west, Turkey, Azerbaijan, and Armenia to the northwest, the Caspian Sea to the north, Turkmenistan to the nort ...
and the middle part of the Liushu formation of northwestern China. It was a precursor to the related '' Sinotherium'' 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 sexes of the same species exhibit different Morphology (biology), morphological characteristics, including characteristics not directly involved in reproduction. The condition occurs in most dioecy, di ...
among rhinos.


Description

''Iranotherium'' is only known from
cranial Standard anatomical terms of location are used to describe unambiguously the anatomy of humans and other animals. The terms, typically derived from Latin or Greek language, Greek roots, describe something in its standard anatomical position. Thi ...
material, and rare
postcrania The postcranium ("behind the cranium"; plural: postcrania) or postcranial skeleton in zoology and vertebrate paleontology is the skeleton apart from the skull. The postcranium encompasses the axial skeleton, which includes the entirety of the verte ...
l elements.


Skull and Horn

The
skull The skull, or cranium, is typically a bony enclosure around the brain of a vertebrate. In some fish, and amphibians, the skull is of cartilage. The skull is at the head end of the vertebrate. In the human, the skull comprises two prominent ...
is long and
dorsally Standard anatomical terms of location are used to describe unambiguously the anatomy of humans and other animals. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position provi ...
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. Eac ...
is long and wide, with a shallow U-shaped nasal notch, the
orbit In celestial mechanics, an orbit (also known as orbital revolution) 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 ...
is prominent and has a high position close to the
skull The skull, or cranium, is typically a bony enclosure around the brain of a vertebrate. In some fish, and amphibians, the skull is of cartilage. The skull is at the head end of the vertebrate. In the human, the skull comprises two prominent ...
roof, the first half of the
zygomatic arch In anatomy, the zygomatic arch (colloquially known as the cheek bone), is a part of the skull formed by the zygomatic process of temporal bone, zygomatic process of the temporal bone (a bone extending forward from the side of the skull, over the ...
is narrow. The
skull The skull, or cranium, is typically a bony enclosure around the brain of a vertebrate. In some fish, and amphibians, the skull is of cartilage. The skull is at the head end of the vertebrate. In the human, the skull comprises two prominent ...
s also show distinct
sexual dimorphism Sexual dimorphism is the condition where sexes of the same species exhibit different Morphology (biology), morphological characteristics, including characteristics not directly involved in reproduction. The condition occurs in most dioecy, di ...
: The base from which the horn grows is large and strong in the
male Male (Planet symbols, symbol: ♂) is the sex of an organism that produces the gamete (sex cell) known as sperm, which fuses with the larger female gamete, or Egg cell, ovum, in the process of fertilisation. A male organism cannot sexual repro ...
, 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 (colloquially known as the cheek bone), is a part of the skull formed by the zygomatic process of temporal bone, zygomatic process of the temporal bone (a bone extending forward from the side of the skull, over the ...
of the
male Male (Planet symbols, symbol: ♂) is the sex of an organism that produces the gamete (sex cell) known as sperm, which fuses with the larger female gamete, or Egg cell, ovum, in the process of fertilisation. A male organism cannot sexual repro ...
. The back half of the
zygoma The term zygoma generally refers to the zygomatic bone, a bone of the human skull that is commonly referred to as the cheekbone or malar bone, but it may also refer to: * The zygomatic arch, a structure in the human skull formed primarily by parts ...
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 mi ...
), 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 Tooth (human), teeth, or bicuspids, are transitional teeth located between the Canine tooth, canine and Molar (tooth), molar teeth. In humans, there are two premolars per dental terminology#Quadrant, quadrant in ...
s are significantly shortened, however, the
cheek teeth Cheek teeth or postcanines 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 ...
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, tear ...
, it was recognized that this
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
was similar to the
rhino A rhinoceros ( ; ; ; : rhinoceros or rhinoceroses), commonly abbreviated to rhino, is a member of any of the five extant taxon, extant species (or numerous extinct species) of odd-toed ungulates (perissodactyls) in the family (biology), famil ...
'' Elasmotherium'' 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 ...
of the latter (This was before '' Sinotherium'' 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 (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...
was designated for the genus. Specimen MNHN 1905–10, and atlas from Maragha, Iran was chosen as the
Lectotype In biology, a type is a particular specimen (or in some cases a group of specimens) of an organism to which the scientific name of that organism is formally associated. In other words, a type is an example that serves to anchor or centralizes ...
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 dentition 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 (: faunae or faunas) is all of the animal life present in a particular region or time. The corresponding terms for plants and fungi are ''flora'' and '' funga'', respectively. Flora, fauna, funga and other forms of life are collectively ...
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 feliformia, feliform carnivoran mammals belonging to the Family (biology), family Hyaenidae (). With just four extant species (each in its own genus), it is the fifth-smallest family in the orde ...
('' Ictitherium sp., Hyaenictitherium wongii, H. hyaenoides, Dinocrocuta gigantea''), two
cats The cat (''Felis catus''), also referred to as the domestic cat or house cat, is a small domesticated carnivorous mammal. It is the only domesticated species of the family Felidae. Advances in archaeology and genetics have shown that the ...
('' Amphimachairodus hezhengensis'', '' Felis sp.''), one
proboscidea Proboscidea (; , ) is 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. Three l ...
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 Monotypic prehistoric placental genera Fossils of Iran {{paleo-oddtoedungulate-stub