Elasmotherium
Encyclopedia
Elasmotherium is an extinct
genus
of giant rhinoceros
endemic to Eurasia
during the Late Pliocene through the Pleistocene
, documented from 2.6 mya to as late as 50,000 years ago, possibly later, in the Late Pleistocene
, an approximate span of slightly less than 2.6 million years. Three species are recognised. The best known, E. sibiricum was the size of a mammoth and is thought to have borne a large, thick horn on its forehead which was used for defense, attracting mates, driving away competitors, sweeping snow from the grass in winter and digging for water and plant roots. Like all rhinoceroses, elasmotheres were herbivorous. Unlike any others, its high-crowned molars were ever-growing. Its legs were longer than those of other rhinos and were designed for galloping, giving it a horse-like gait. The Russian paleontologists of the 19th century who discovered and named the initial fossils were influenced by ancient legends of a huge unicorn roaming the steppes of Siberia. To date no evidence either contradicts or confirms the possibility that Elasmotherium survived into legendary times.
, the Dirécteur Perpétuel of the Natural History Museum, Moscow University, at a presentation before the Societé Impériale des Naturalistes in 1808. The next year in the Mémoires of the society he reported what he had said in the Programme d'invitation:
Then he noted on his derivation of the name:
All he had before him was one lower jaw donated to the museum by Yekaterina Romanovna Vorontsova-Dashkova
, which he named Elasmotherium sibiricum, lamenting that it was the sole species of which he knew. The molars, the only teeth in the jaw, had formed in layers like tree rings, except the "rings," or lamellae, were highly corrugated. The edges in the grinding surface were elaborately sinuous to better break down the grasses on which the animal fed.
and was rather larger than the contemporary Woolly Rhinoceros
.
E. sibiricum had a measured shoulder height of approximately 2 metres (6.6 ft). To it, however, must be added the height of a massive hump anchored on the fin-like transverse processes extending from the top of the cervical vertebrae, a maximum of 53 centimetres (1.7 ft). The total height then was in excess of 253 centimetres (8.3 ft). The measured length of sibiricum (from a nearly complete skeleton found at Gaevskaya) is 4.5 metres (14.8 ft). Extrapolation
from the greater size of caucasicum molar
s obtains a length of 5 metres (16.4 ft)—5.2 metres (17.1 ft) for caucasicum.
The science of estimation is more advanced than it was when the 19th century estimates and reconstructions were made. A 1986 study of mammalian communities by Serge Legendre
correlated body weight with the M1 area, which, in the code of dental formulae, means the area of the first lower molar
. A correlation typically is expressed by a curve
in a graph showing two (in this case) quantitative variables on a modified Cartesian coordinate system. The data is graphed as a scatter of points to which a smooth curve
is then fitted in regression analysis
. The curve fitted by Legendre for body weight of perissodactyls in this case is the function
where x is the weight, y is the M1 area (also called the ml area, the length of the molar grinding surface times its width), and the two numbers are constants applying only to perissodactyls. According to Legendre's formula, E. sibiricum had a mass of over 4000 kilogram; E. caucasicum, 5000 kilogram. These weights place Elasmotherium in the "really huge" category of all Rhinocerotidae and therefore the animal was "strongly brachyopodial;" that is, they required feet of large contact area to prevent sinking into the soil. The feet were unguligrade, the front larger than the rear: purely tridactyl on Digits II-IV in the rear, but with an extra vestigial digit, I, in the front.
s can be divided into two general groups on the basis of nutrition, which grade into each other morphologically: "foregut fermenters
" and "hindgut fermenters
." The border region is correlated to bulk: up to 600 kilograms (1,322.8 lb) – 1200 kilograms (2,645.5 lb) are the former; over it, the latter. In foregut fermentation the animal must "browse" to select the most nutritious plants and then ruminate
to make up for the shorter digestive tract. The hind-gut fermenters are "bulk-feeders:" they ingest large quantities of low-nutrient food, which they process for a longer time in a much longer intestine. The main food in that category is grass, indicating that Elasmotherium, like the elephants, was probably a grassland "grazer" moving over long distances to take advantage of the growth phases of grass in different regions. The standard is not without exception, as Indricotherium, the largest land mammal ever, with a weight of 15–20 tons, subsisted by browsing the treetops.
There are also paleontological indications of the grazing. In general, the normal position of the head can be determined by the angle between a vertical plane coinciding with the occiput
of the cranium, which is always vertical, and a plane through the base of the cranium. A right or acute angle would indicate a head held high for browsing leaves at various heights. Elasmotherium had the most obtuse angle of the Rhinocerotids. It could only reach the lowest levels and therefore must have grazed habitually. This morphological feature favors the identification of the one-horned beast depicted in Rouffignac
Cave (shown in this article) as Elasmotherium and lends some validity to the bison-like restorations based on it.
s and three molar
s (originally taken for five molars), no incisor
s, no canine
s. Where some of the browsers kept the incisors in the form of tusks, E, had instead a spoon-like symphysis
, or tip, of the lower mandible
and a rostrum
, or beak, of the upper, which served as a bony basis for a soft-tissue labial
grasping and tearing mechanism. Grass, a very tough, fibrous material, contains phytolith
s, microscopic granules mainly of silica, which act as sandpaper on the molars of grazers. Their response in geologic time is to evolve cheek teeth with large crowns (hypsodonty). There appears to be a correlation between grazing and hypsodonty:
Vladimir Onufryevich Kovalevsky first proposed a connection between hypsodonty and grazing for horses in 1873. Since then the concept has been expanded to all mammalian grazers at any time and has further been elaborated into hypsodonty or proto-hypsodonty and hypselodonty or euhypsodonty. The euhypsodonts, of which, surprisingly for its bulk, E. was one, have ever-growing high-crowned teeth. Most other examples are to be found among diminutive mammals such as Rodentia, which already casts doubt on the correlation, as they do not generally graze grass.
Teeth form from the top down through the deposition of enamel on a cement core by formative soft tissue in the jaw. The enamel of hypsodont Perissodactyla is highly rugose rather than sharp. In brachydont species, such as humans, when the crown is complete, the roots are deposited and finally the completed tooth erupts. Hypselodonty is a condition of tooth eruption and continued crown formation before a delayed root formation. In its most developed variety, the roots never form. Only rare fossils of E. show any sign of a root, and that on a premolar. No molars have roots, or, in the terminology of some, the roots are "open."
of the Mesozoic
, a group of mammals so primitive that he describes their cheek teeth as "molariform," as they are neither clearly molars nor premolars. Molariform hypsodonty "cannot be correlated with a grass diet, since grasses were not present." Instead he suggests for his example, Sudamerica ameghinioi, that it lived a "semiaquatic and perhaps a burrowing way of life." Modern species provide many examples from beavers to hippopotamuses.
Attempts have been made to link the wear on E. teeth to grazing. In 1938, H.E. Wood, a Rhinocerotid tooth specialist, pointed out that interproximal wear, or loss of tooth surface between teeth, due to abrasion during mastication, of E. is similar to that of the White Rhinoceros
, the only remaining Rhinocerotid grazer, which has hypsodont teeth. Data such as this led to an intuitive concept of some sort of correlation between grass-eating and hypsodonty, but it has been difficult to isolate mathematically.
A 2007 study by Mendoza and Palmqvist compared the habitats and diets of 134 species of living ungulate
s, for which this data is known, both Artiodactyla and Perissodactyla, against "thirty-two craniodental measurements" to discover what correlations exist for the group studied and to test hypotheses concerning hypsodonty and mode of life. Habitats considered were open (savanna, deserts), mixed (wooded savanna, brush) and closed (riverine and forest). Diets considered were grazing, mixed grazing and browsing, browsing, omnivory and special niches, such as treetop browsing. Measurements included the Hypsodonty Index (HI) and Muzzle Width (MZW). The results showed that, except for the "high-level" browsers, hypsodonty is correlated to "open and mixed habitats." The HI was not precise enough to discriminate between open and mixed. However, high MZW is correlated to grazing in the open category, although some forest species also have wide muzzles. Grazers therefore are distinguished by a combination of high-crowned cheek teeth and wide muzzles, both of which are possessed by E. Life in the open is implied.
By distal they mean "furthest outward;" that is, the extremities. Cursorial animals are unequivocally "runners" although the authors did not examine what sort of gait scientifically should be proposed as "running." They selected caucasicum for study because of the availability of a few dozen limb bone fragments from Nihewan, China. These made possible a selective comparison with the fewer bones remaining of other fossil rhinocerotids. In comparison with them, the long legs of E. are the most derived; that is, the others did not have the same cursorial capabilities. The authors approach but do not solve the problem of how to reconcile the weight with the supposed mobility. They say elsewhere in the article that the legs of caucasicum are to be distinguished from those of other fossil Rhinocerotids at Nihewan by their "enormous size."
The White Rhinoceros
at an estimated weight of 2.5 ton has been photographed galloping at a speed of about 30 kph. In a gallop
, all feet are off the ground ("ballistic phase") twice a cycle, a feat that elephants, at 2.5-11 ton, cannot perform. They can walk up to 20 kph; however, their straight, relatively inflexible legs are those of striders, not the bent and spring-like legs of gallopers, which utilize haunches, ankle mobility and knee flexion to spring off the ground on alternative legs of a pair. E. legs are sufficiently like those of the White Rhino to hypothesize a similar gait even though E. weighed 4.5-5 ton.
, or horn
s and hooves, made of keratin
, the same substance of which hair
is made. These keratin structures appear to have formed in the Mesozoic
, a remnant in humans being the nail
s. A keratin horn is to be distinguished from a bone horn and a tusk
. Bone forms the base of most horns but in some cases the horn is entirely of bone. A tusk is a modified canine or incisor tooth. Rhinocerotidae have had tusks, but not E. Two open questions are whether they were horned or hornless, hairy or hairless. Most Rhinocerotidae have and have had horns, but there are some instances of hornlessness, and most are or were hairy, such as the Wooly Rhinoceros, but no instances of hair or horn have yet been found for E. Only circumstantial evidence of them exists.
The main evidence suggestive of a horn on E. is a frontal protuberance, which struck the attention of the late 19th century paleontologists and was immediately interpreted as the bony basis for a horn by most investigators from that time forward. A skull of E. sibiricum from the Volga region (cast shown in this article's lead picture) described by Alexander Brandt in the Russian journal, Niwa, and reported in Nature in 1878 offers the following description of the protuberance: hemispherical, 5 inches (12.7 cm) deep, furrowed surface, circumference of 3 foot (0.9144 m). The furrows are interpreted as the seats of blood vessels for the tissues that generated the horn:
Brandt was already familiar with the legend of a unicorn among the Tatars of Siberia with a horn so large it required a sledge for transport, and made the connection in interpreting the bump as the base of a horn. He also interpreted the rostrum of the upper mandible as the basis of a nasal horn, a hypothesis now rejected in favor of the cropping labia. In any case the non-circular base indicates a section through the horn would not have been circular. This possibility is supported by another fossil with a non-circular partially healed puncture wound in the base, chiefly interpreted as the result of dueling other males with the horn.
The ungulate
s typically combine keratin and bone in various structures. If horns are keratinous, they have a bone core. Rhinocerotids horns, however, are uniquely derived. Hieronymus, an expert in Rhinoceros dermatology
, says:
He defines rhinocerotid horns as:
This tissue is "strikingly convergent" with other "cornified epidermis" in horses, cetaceans, artiodactyls and birds. The horn is not attached to the bone of the boss but grows from the surface of a dense dermal tissue. The top layer keratinizes itself to form tubules about 1-2 millimeter high, the cells of which then die. The next layer forms below it. As the layers age the horn loses diameter by degradation of the keratin due to ultraviolet light, desiccation and mechanical wear from contact with objects and agonistic behavior. However, melanin
and calcium deposits in the center harden the keratin there, causing differential wear and shaping of the horn.
The dermis generating the horn is anchored to the boss by interpenetration between rugosities – various irregularities of bone, which it creates by deposition. This tissue is a specialization of dermal armor, which, whenever it attaches to bone, deposits the rugosities to strengthen the attachment. The author states that rugosity is "a bony signature of dermal armor." Cranial rugisoty is an indication, but not a sure sign, of a horn. If, on the other hand, an annular (ring-shaped) pattern is visible in the rugosity, it is due to "stress concentration at the edges of horns" and is "the signature for epidermal horns."
Hieronymus found annular rugosities in all living and some fossil Rhinocerotidae. The rings had previously been noted on additional fossils. To date E. has not been examined for rings under lighting designed to show them up; however, based on the observations of other paleontologists, the author says "squamosal
rugosity" is the "most pronounced cranial rugosity in the elasmotherine lineage." This fact suggests an especially firm attachment was required, which, combined with the extraordinarily large hump of muscle for managing the head, suggest a large and heavy horn. In the early 19th century the state of the fossils had not yet revealed the presence of a horn. By 1910 and for all the time since then the paleontologists have not ventured a mathematical estimate but have preferred to refer to the horn as immense, enormous, great or huge. As the size and shape of the horn depended on the concentration of melanin and calcium, and no known indicator of those remains, any further estimate of horn morphology is purely speculative.
In 1968 Björn Kurtén's The Age of Dinosaurs pronounced, without a clue of the reasoning followed, that E. "carried a single frontal horn two meters in length." This figure is often repeated in non-technical books and articles, as it is the only one available. The number is speculative, but coincidentally it happens to be the shoulder height of E. sibiricum. If in fact E. swept snow with the horn, or excavated holes in the terrain with it, the length can hardly have been less to reach the ground.
alone has 30 sites of E. sibiricum. Dozens of crania have been reconstructed and given archaeological identifiers. The division into species is based mainly on the fine distinctions of the teeth and jaws and the shape of the skull. The finds can be dated only by context.
In 2002 P. O. Antoine performed a cladistic analysis using 282 "cranial, dental and postcranial characters" of 28 "terminal taxa" of "elasmotheriines from China and Mongolia" and four outgroups. He found that the elasmotheres were a monophyletic group. He says:
Antoine was following the now discontinued practice of considering the Miocene ancestral species as elasmotheres. They were not, however, distinguished by the features of an elasmothere. The original ancestors were "minute brachydont animals." Elasmotherium was a "mammoth-sized hypsodont." In his view,
V.A. Terjaev in 1948 had a different view of the cause of the hypsodont
y, maintaining that it was caused by the heavier grains in soil on plants pulled from moister environments, and that consequently Elasmotherians lived in "riparian biotopes." Noskova points out that Elasmotheriums diet was comparable to that of the concurrent Archidiskon, which ranged over both steppe and riverland.
genus
of Central Asia
, Sinotherium
. Elasmotherium is thought to be the most derived genus of elasmothere, with E. caucasicum in turn being more derived than E. sibiricum. The two Chinese fossils, formerly considered distinct species, E. inexpectatum and E. peii, defined by Chow in 1958, have been sunk into E. caucasicum. They were found in northern China
from the Early Pleistocene
Nihewan Faunal Assemblage
(from the same valley as nearby Nihewan in Shanxi
Province) and were extinct at approximately 1.6 Ma.
E. caucasicum, defined by Borissjak in 1914, flourished in the Black Sea
region, as a member of the Early Pleistocene
Tamanian Faunal Unit (1.1–0.8 Mya, Taman Peninsula
). However, an elasmotherian species turned up in the preceding Khaprovian or Khaprov Faunal Complex, which was at first taken to be caucasicum, and then on the basis of the dentition was redefined as a new species, E. chaprovicum (Shvyreva, 2004), named after the Khaprov Faunal Complex. The Khaprov is in the MIddle Villafranchian, MN17, which spans the Piacenzian of the Late Pliocene and the Gelasian of the Early Pleistocene
of Northern Caucasus, Moldova
and Asia and has been dated to 2.6–2.2 Mya.
E. sibiricum, described by Johann Fischer von Waldheim
in 1808 and chronologically the latest species of the sequence, coming from E. caucasicum in the Middle Pleistocene, ranged from southwestern Russia to western Siberia and southward into Ukraine
and Moldova
. It appears in the Middle Pleistocene
Khazar Faunal Complex of the Sea of Azov
region, which has "no exact stratigraphic situation."
The end of the elasmotheres is questionable; new evidence continues to turn up. The latest is from two caves in southern Siberia
, a sibericum tooth in Smelovskaya Cave 11 and remains from Batpak, both associated with Middle Pleistocene
relict species, including large herbivores and predators. They must have been dragged into the caves by some predator, perhaps even modern man. The elasmothere tooth and one of a Cave Hyena
from Smelovskaya were carbon dated to slightly greater than 50,000 BP in the Late Pleistocene
. A recent study of relict Late Pleistocene
remains in the Beringia region (Alaska, Eastern Siberia) identified a pattern of "imbedded micrometeorites" consistent with an extinction event similar to the Tunguska event
of 1908, only carbon-dated to 37,000 BP. A Siberian Elasmotherium skull in a museum was found to have this pattern.
, both being facets of the evolution of mammalian herbivores during the Cenozoic
. The overall pattern is based on the evolutionary development of grass
and grassland
, and further the changing distribution of C3
and C4
types of plant metabolism, so-called because they generate a molecule containing 3 or 4 Carbon
atoms by different metabolic pathways (sequences of chemical reactions).
and subsequently the Hyracodontidae
and Amynodontidae
show no sign of dermal armor or horns. The oldest known genus of Rhinocerotids, or true Rhonoceroses, is Teletaceras from the Middle Eocene
of North America and the Late Eocene of Asia. Due to the timing of bridges, it does not appear in South America. It features the first known Rhinocerotid tusks, a derived feature, which are still extant in three species and are "the primary offensive weapons" of those species. They appear to have been sexually dimorphic features convergent with the development of tusks in pigs, hippopotamoses and elephant seals.
The first dermal armor is indicated by cranial rugosity on Trigonias
osborni, Late Eocene
, 42-32 Mya. It appears more developed in Subhyracodon
, early Oligocene
, 33 Mya, both of North America. There appears to have been a correlation between the tusks and the armor later (4 Mya later) defending against them.
Having begun to diversify (increase in species and genera) in America the Rhinocerotids spread rapidly over Beringia when that bridge appeared in the Early Oligocene
to Eurasia, where subsequently they became highly diverse. The family reached a maximum floruit in the Miocene
. At that time they entered Africa, but never were very diverse there. In the Pliocene
they declined, disappearing from America. The Pleistocene
brought extinction from Eurasia but African and Southeast Asian species survived.
Elasmotherium appeared in the Late Pliocene
, apparently deriving from the Miocene—Pliocene Sinotherium
, sometimes informally aggregated with it. Sinotherium in turn appeared to be related to Iranotherium
; that is, Elasmotherium was believed to have been an iranotheriine. A 1995 cladistic analysis by Cerdeño asserted that Elasmotherium was most closely related to Coelodonta, the "Wooly Rhinoceros," whose soft-tissue corpus has been found in a few locations. By implication, Sinotherium was not an iranothere. However, Antoine's 2002 analysis reaffirmed that the elasmotheriine line was iranotheriine. Certainty concerning the exact line of descent remains elusive; fossils and techniques of the future will perhaps resolve it.
The grass of grasslands belongs predominantly to the Poaceae
family, also called the "true grasses." Species can be divided into "warm season grasses," which utilize C4, and "cool season grasses," which utilize C3. The evolution of grasses preceded the evolution of grasslands, which were created by the spreading of grasses from the forest onto the plains in the Miocene
. These initial grasslands were C3, which happen to be more nutritious than C4 species.
The evolution of Rhinocerotidae and Equidae in the Miocene grasslands proceeded along parallel lines from similar causes. Diminutive, cursorial, browsing, brachydont species expanded their ranges from the Oligocene
forests into the nutritious C3 Miocene grasslands and there over the millions of years of the Miocene evolved diverse and flourishing populations on the ample food supply, gradually increasing in size. Some of these species exploited the grazing niche. Hispanotherium
, Chilotherium
and Teleoceras
"show a clear tendency to hypsodonty, and a grazing feeding adaptation has usually been attributed to them."
By the late Miocene, C4 metabolism was well established on the plains. In place of the prehistoric C3 grasslands (whuch are no longer extant) was the savanna
, today's subtropical mixture of grasses and sedge
s. The deciding factors in this turn of evolution were a documented increase in temperature and aridity over the period. Faced with scant nutrition from browsing, the smaller Rhinocerotidae and Equidae declined in diversity, the gate in that direction, so to speak, now being closed.
The evolutionary response in the Pliocene
was in the direction of bulk-feeding. Massive grazers, such as Elasmotherium, maximally hypsodont, moved across vast ranges consuming C4 grass in bulk. While the horses and other more nearly modern grazers could rely on gallop
ing for mobility in any circumstance, there is a question as to what degree the brachypodial giants can be said to be "cursorial." Elephants, for example, must keep one foot on the ground, where true gallopers are entirely off it for brief intervals. The horns and impenetrable body armor insured that the giants would not be required to gallop away for any predator. At least until the arrival of man, they were, as the legends relate, lords of the savanna.
or mysterious beast with one horn, as opposed to the delicate imaginary unicorns of Europe based on narwhal
horns, are scattered across the former range of Elasmotherium from China to Eastern Europe. They have been noted there since the first known literature of the Middle Ages
.
and Mongolic languages
and lore. The writers of works in those languages have not known how to depict these strange beasts. A common theme is that they have a single horn, are dark and are huge.
For example, a book of divination in the Old Turkic language
uses kälän käyik, "tawny beast," (the kilin is altered to kälän by vowel harmony
). The Han I Araha Manju Gisun I Buleku Bithe, "Mirror of the Manchu Language," dating from the 18th century, defines the kilin as
The Chinese had other unicorns, such as the Xiezhi, but the latter was less like a real animal and more a symbolic figure. It had the ability to discriminate right from wrong and therefore a representaton of it was placed in the courtrooms of the Han Dynasty
and, in the 3rd and 4th centuries, on the caps of judges. It looks like a seated or standing lion with a horn rising from the top of the head. In contrast to this mythical beast, the k'i-lin is presented as an animal actually caught by hunting parties of the past. The Shiji
("Historical Records") of Sima Qian
in the 2nd century mentions the capture of a "deer-like" animal with one horn in 122 BC. The Chunqin, an encyclopedia of the 3rd century BC, states that a unicorn was captured in 481 BC, but without any description. Chinese representations of unicorns vary quite a lot, but an engraving on a bronze vessel of the Warring States Period
, shows an animal very like the cave paintngs supposed to be of Elasmotherium: head down for grazing, horn protruding horizontally from the forehead, neck and shoulders humped.
In the northeast of the range, in 1866 Vasily Radlov
reported a legend among the Yakuts
of Siberia
of a "huge black bull" killed by spear, who had "a single horn" so large it required a sledge for transportation. In 1878 A. F. Brandt suggested the beast was an Elasmotherium.
, a collection of ballads has survived, combined into a spiritual theme deriving ultimately from Zoroastrianism
, but with Christian overtones, called the Stikh o Golubinioi knige, or in another version the Golubiniai kniga, "The Book of the Dove." The "dove," or golub of the Christian work is probably an alteration of glub, "depth," so that the reconstructed name, *Glubinnaia kniga, is on the same theme as the Zend Avesta and similar works in Iran
, the struggle between cosmic good and evil.
The force of pravda, or "rightness," is symbolized by a beast called mainly the Indrik
, but also Indra, Beloiandrikh, Kondryk, Edinrog and Edinor. The name is a calque
: Greek mono-keros ("single-horn") > ino-rog (in, "one"; rogom, "horn") > edinrog > Indrik. As a unicorn struggling with a lion representing krivda, "lies," the Indrik has the body of a goat or a horse in representations.
An Indrik composite from all the variants of the ballads depicts the Indrik as dwelling in the Holy Mountain, which it consumes for sustenance. It is the mother or the father of all animals. It wanders unseen on the plain by day. When other beasts encounter it they must do obeisance. The Indrik saves the world from draught by digging springs of pure water. By night it wanders in the subterranean world, forging a path with its horn. There it purifies all the waters and releases them from any blockages.
The righteous but fantastic unicorn appears also in the Bundahišn
or Zand Āgāhīh, a Middle Persian
Zoroastrian composition concerning the "deep and miraculous utterances" about the creation, the archetype of the *Gubinnaia kniga, "Book of the Deep." Its unicorn is an ass of 3 feet, 6 eyes, 9 mouths, 2 ears, 1 horn, a dark blue head and a white body, which is a symbolic rather than a real entity. It stands in the sea of Fraxwkand (wherever that is) and purifies the waters by urinating in them so that life may continue. The Russian concept of a unicorn was apparently based on a different experience.
In fact the Arabo-Persian word for unicorn in the region was karkadann
, which extended also to India
, and was loaned into the Turkmen language
as kergeden and the Ottoman Turkish language
as gergeden, among others. The word had an equivocal meaning: unicorn and rhinoceros. This rhinoceros has capabilities and habits not possessed by the Indian rhinoceros
: it indulges in mortal combat with elephants, which it kills by goring underneath with its horn. The symbol of evil is thus transferred to the elephant in that milieu; however, the two conduct their combats over grasslands never populated by either in historical times. The root of the word is Sanskrit
Khadga-, Khadga dhenu-, "rhinoceros." In Christianity the single horn becomes a symbol of monotheism
.
Science writer and cryptozoologist Willy Ley
theorised that these legends have simply been passed down since a hunting association with the beast in prehistoric times, and that the animals did not actually survive into recorded history. One argument for survival is the testimony by the medieval traveller, Ibn Fadlan. His account states:
Extinction
In biology and ecology, extinction is the end of an organism or of a group of organisms , normally a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and recover may have been lost before this point...
genus
Genus
In biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...
of giant rhinoceros
Rhinoceros
Rhinoceros , also known as rhino, is a group of five extant species of odd-toed ungulates in the family Rhinocerotidae. Two of these species are native to Africa and three to southern Asia....
endemic to Eurasia
Eurasia
Eurasia is a continent or supercontinent comprising the traditional continents of Europe and Asia ; covering about 52,990,000 km2 or about 10.6% of the Earth's surface located primarily in the eastern and northern hemispheres...
during the Late Pliocene through the Pleistocene
Pleistocene
The Pleistocene is the epoch from 2,588,000 to 11,700 years BP that spans the world's recent period of repeated glaciations. The name pleistocene is derived from the Greek and ....
, documented from 2.6 mya to as late as 50,000 years ago, possibly later, in the Late Pleistocene
Late Pleistocene
The Late Pleistocene is a stage of the Pleistocene Epoch. The beginning of the stage is defined by the base of the Eemian interglacial phase before the final glacial episode of the Pleistocene 126,000 ± 5,000 years ago. The end of the stage is defined exactly at 10,000 Carbon-14 years BP...
, an approximate span of slightly less than 2.6 million years. Three species are recognised. The best known, E. sibiricum was the size of a mammoth and is thought to have borne a large, thick horn on its forehead which was used for defense, attracting mates, driving away competitors, sweeping snow from the grass in winter and digging for water and plant roots. Like all rhinoceroses, elasmotheres were herbivorous. Unlike any others, its high-crowned molars were ever-growing. Its legs were longer than those of other rhinos and were designed for galloping, giving it a horse-like gait. The Russian paleontologists of the 19th century who discovered and named the initial fossils were influenced by ancient legends of a huge unicorn roaming the steppes of Siberia. To date no evidence either contradicts or confirms the possibility that Elasmotherium survived into legendary times.
Discovery
The fossil received its name from Johann Fischer von WaldheimJohann Fischer von Waldheim
Johann Gotthelf Fischer von Waldheim was a German anatomist, entomologist and paleontologist....
, the Dirécteur Perpétuel of the Natural History Museum, Moscow University, at a presentation before the Societé Impériale des Naturalistes in 1808. The next year in the Mémoires of the society he reported what he had said in the Programme d'invitation:
"L'elasmotherium est un animal à tête elongée sans dents incisives et sans canines, de chaque coté 5 molaires à lames contournées."
Then he noted on his derivation of the name:
"Du mot grec ἐλασμος, lamelle, pour designer la forme lamelleuse des dents molaires."
All he had before him was one lower jaw donated to the museum by Yekaterina Romanovna Vorontsova-Dashkova
Yekaterina Romanovna Vorontsova-Dashkova
Princess Yekaterina Romanovna Vorontsova-Dashkova was the closest female friend of Empress Catherine the Great and a major figure of the Russian Enlightenment...
, which he named Elasmotherium sibiricum, lamenting that it was the sole species of which he knew. The molars, the only teeth in the jaw, had formed in layers like tree rings, except the "rings," or lamellae, were highly corrugated. The edges in the grinding surface were elaborately sinuous to better break down the grasses on which the animal fed.
Description
Various theories of Elasmothere morphology, nutrition and habits have been the cause of wide variation in reconstruction. Some show the beast trotting like a horse with a horn; others hunched over with head to the ground, like a bison, and still others immersed in swamps like a hippopotamus. The use of the horn and whether or not there was one, and how large, have been popular topics. The statistical correlations of modern paleontology have taken much of the speculation out of the subject, although some details remain undetermined.Morphology
The most reconstructed species is perhaps E. sibiricum by generations of scientists working at the Paleontological Museum in Moscow and elsewhere in Russia. The majority of the fossils fall or have fallen within their national jurisdiction. The dimensions and morphology of the various reconstructions vary considerably. They are for the most part estimating the gross details from the minutiae. However, they all agree on the general order of magnitude, that sibiricum was comparable to a MammothMammoth
A mammoth is any species of the extinct genus Mammuthus. These proboscideans are members of Elephantidae, the family of elephants and mammoths, and close relatives of modern elephants. They were often equipped with long curved tusks and, in northern species, a covering of long hair...
and was rather larger than the contemporary Woolly Rhinoceros
Woolly Rhinoceros
The woolly rhinoceros is an extinct species of rhinoceros that was common throughout Europe and Asia during the Pleistocene epoch and survived the last glacial period. The genus name Coelodonta means "cavity tooth"...
.
E. sibiricum had a measured shoulder height of approximately 2 metres (6.6 ft). To it, however, must be added the height of a massive hump anchored on the fin-like transverse processes extending from the top of the cervical vertebrae, a maximum of 53 centimetres (1.7 ft). The total height then was in excess of 253 centimetres (8.3 ft). The measured length of sibiricum (from a nearly complete skeleton found at Gaevskaya) is 4.5 metres (14.8 ft). Extrapolation
Extrapolation
In mathematics, extrapolation is the process of constructing new data points. It is similar to the process of interpolation, which constructs new points between known points, but the results of extrapolations are often less meaningful, and are subject to greater uncertainty. It may also mean...
from the greater size of caucasicum molar
Molar (tooth)
Molars are the rearmost and most complicated kind of tooth in most mammals. In many mammals they grind food; hence the Latin name mola, "millstone"....
s obtains a length of 5 metres (16.4 ft)—5.2 metres (17.1 ft) for caucasicum.
The science of estimation is more advanced than it was when the 19th century estimates and reconstructions were made. A 1986 study of mammalian communities by Serge Legendre
Serge Legendre
Serge Legendre is a research scientist in the field of paleobiology with the Institute of Paleoenvironment & Paleobiosphere, University of Lyonand Editor-in-Chief of Geobios, a scientific journal published bi-monthly.-Publications:...
correlated body weight with the M1 area, which, in the code of dental formulae, means the area of the first lower molar
Molar (tooth)
Molars are the rearmost and most complicated kind of tooth in most mammals. In many mammals they grind food; hence the Latin name mola, "millstone"....
. A correlation typically is expressed by a curve
Graph of a function
In mathematics, the graph of a function f is the collection of all ordered pairs . In particular, if x is a real number, graph means the graphical representation of this collection, in the form of a curve on a Cartesian plane, together with Cartesian axes, etc. Graphing on a Cartesian plane is...
in a graph showing two (in this case) quantitative variables on a modified Cartesian coordinate system. The data is graphed as a scatter of points to which a smooth curve
Curve
In mathematics, a curve is, generally speaking, an object similar to a line but which is not required to be straight...
is then fitted in regression analysis
Regression analysis
In statistics, regression analysis includes many techniques for modeling and analyzing several variables, when the focus is on the relationship between a dependent variable and one or more independent variables...
. The curve fitted by Legendre for body weight of perissodactyls in this case is the function
where x is the weight, y is the M1 area (also called the ml area, the length of the molar grinding surface times its width), and the two numbers are constants applying only to perissodactyls. According to Legendre's formula, E. sibiricum had a mass of over 4000 kilogram; E. caucasicum, 5000 kilogram. These weights place Elasmotherium in the "really huge" category of all Rhinocerotidae and therefore the animal was "strongly brachyopodial;" that is, they required feet of large contact area to prevent sinking into the soil. The feet were unguligrade, the front larger than the rear: purely tridactyl on Digits II-IV in the rear, but with an extra vestigial digit, I, in the front.
Diet
HerbivoreHerbivore
Herbivores are organisms that are anatomically and physiologically adapted to eat plant-based foods. Herbivory is a form of consumption in which an organism principally eats autotrophs such as plants, algae and photosynthesizing bacteria. More generally, organisms that feed on autotrophs in...
s can be divided into two general groups on the basis of nutrition, which grade into each other morphologically: "foregut fermenters
Foregut fermentation
Foregut fermentation is a form of digestion that occurs in the foregut of some animals. It has evolved independently in several groups of mammals, and also in the hoatzin bird. All ruminants use foregut fermentation, whereas only some rodents and marsupials do. It has also evolved in colobine...
" and "hindgut fermenters
Hindgut fermentation
Hindgut fermentation is a digestive process seen in monogastric herbivores, animals with a simple, single-chambered stomach. Cellulose is digested with the aid of symbiotic bacteria. The microbial fermentation occurs in the digestive organs that follow the small intestine, namely the large...
." The border region is correlated to bulk: up to 600 kilograms (1,322.8 lb) – 1200 kilograms (2,645.5 lb) are the former; over it, the latter. In foregut fermentation the animal must "browse" to select the most nutritious plants and then ruminate
Ruminant
A ruminant is a mammal of the order Artiodactyla that digests plant-based food by initially softening it within the animal's first compartment of the stomach, principally through bacterial actions, then regurgitating the semi-digested mass, now known as cud, and chewing it again...
to make up for the shorter digestive tract. The hind-gut fermenters are "bulk-feeders:" they ingest large quantities of low-nutrient food, which they process for a longer time in a much longer intestine. The main food in that category is grass, indicating that Elasmotherium, like the elephants, was probably a grassland "grazer" moving over long distances to take advantage of the growth phases of grass in different regions. The standard is not without exception, as Indricotherium, the largest land mammal ever, with a weight of 15–20 tons, subsisted by browsing the treetops.
There are also paleontological indications of the grazing. In general, the normal position of the head can be determined by the angle between a vertical plane coinciding with the occiput
Occiput
The occiput is the anatomical term for the posterior portion of the head, in insects the posterior part of those head capsule.-Clinical significance:Trauma to the occiput can cause a basilar skull fracture....
of the cranium, which is always vertical, and a plane through the base of the cranium. A right or acute angle would indicate a head held high for browsing leaves at various heights. Elasmotherium had the most obtuse angle of the Rhinocerotids. It could only reach the lowest levels and therefore must have grazed habitually. This morphological feature favors the identification of the one-horned beast depicted in Rouffignac
Rouffignac
Rouffignac is a commune in the Charente-Maritime department in southwestern France.-Population:-References:*...
Cave (shown in this article) as Elasmotherium and lends some validity to the bison-like restorations based on it.
Dentition
A third type of evidence that Elasmotherium was a grazer is that of tooth wear and morphology. Like all Rhinocerotidae, E. had cheek teeth evolved for herbivory: two premolarPremolar
The premolar teeth or bicuspids are transitional teeth located between the canine and molar teeth. In humans, there are two premolars per quadrant, making eight premolars total in the mouth. They have at least two cusps. Premolars can be considered as a 'transitional tooth' during chewing, or...
s and three molar
Molar (tooth)
Molars are the rearmost and most complicated kind of tooth in most mammals. In many mammals they grind food; hence the Latin name mola, "millstone"....
s (originally taken for five molars), no incisor
Incisor
Incisors are the first kind of tooth in heterodont mammals. They are located in the premaxilla above and mandible below.-Function:...
s, no canine
Canine tooth
In mammalian oral anatomy, the canine teeth, also called cuspids, dogteeth, fangs, or eye teeth, are relatively long, pointed teeth...
s. Where some of the browsers kept the incisors in the form of tusks, E, had instead a spoon-like symphysis
Symphysis
A symphysis is a fibrocartilaginous fusion between two bones. It is a type of cartilaginous joint, specifically a secondary cartilaginous joint.1.A symphysis is an amphiarthrosis, a slightly movable joint.2.A growing together of parts or structures...
, or tip, of the lower mandible
Mandible
The mandible pronunciation or inferior maxillary bone forms the lower jaw and holds the lower teeth in place...
and a rostrum
Rostrum (anatomy)
The term rostrum is used for a number of unrelated structures in different groups of animals:*In crustaceans, the rostrum is the forward extension of the carapace in front of the eyes....
, or beak, of the upper, which served as a bony basis for a soft-tissue labial
Lip
Lips are a visible body part at the mouth of humans and many animals. Lips are soft, movable, and serve as the opening for food intake and in the articulation of sound and speech...
grasping and tearing mechanism. Grass, a very tough, fibrous material, contains phytolith
Phytolith
Some plants can take up silica in the soil, whereupon it is deposited within different intracellular and extracellular structures of the plant. After these plants decay, silica is redeposited in the soil in the form of phytoliths , which are rigid, microscopic structures of varying sizes and shapes...
s, microscopic granules mainly of silica, which act as sandpaper on the molars of grazers. Their response in geologic time is to evolve cheek teeth with large crowns (hypsodonty). There appears to be a correlation between grazing and hypsodonty:
"As a general rule, extant herbivores with low-crowned teeth are predominantly browsers and species with high-crowned teeth are predominantly grazers."
Vladimir Onufryevich Kovalevsky first proposed a connection between hypsodonty and grazing for horses in 1873. Since then the concept has been expanded to all mammalian grazers at any time and has further been elaborated into hypsodonty or proto-hypsodonty and hypselodonty or euhypsodonty. The euhypsodonts, of which, surprisingly for its bulk, E. was one, have ever-growing high-crowned teeth. Most other examples are to be found among diminutive mammals such as Rodentia, which already casts doubt on the correlation, as they do not generally graze grass.
Teeth form from the top down through the deposition of enamel on a cement core by formative soft tissue in the jaw. The enamel of hypsodont Perissodactyla is highly rugose rather than sharp. In brachydont species, such as humans, when the crown is complete, the roots are deposited and finally the completed tooth erupts. Hypselodonty is a condition of tooth eruption and continued crown formation before a delayed root formation. In its most developed variety, the roots never form. Only rare fossils of E. show any sign of a root, and that on a premolar. No molars have roots, or, in the terminology of some, the roots are "open."
Habitats
If all grazers are hypsodont, not all hypsodonts are grazers. The supposed correlation between grass-eating and hypsodonty proved difficult to support in a number of instances. Koenigswald, for example, pointed out that hypsodonty had occurred among the GondwanatheriaGondwanatheria
Gondwanatheria is an extinct group of mammals that lived during the Upper Cretaceous through the Eocene in the Southern Hemisphere, including Antarctica...
of the Mesozoic
Mesozoic
The Mesozoic era is an interval of geological time from about 250 million years ago to about 65 million years ago. It is often referred to as the age of reptiles because reptiles, namely dinosaurs, were the dominant terrestrial and marine vertebrates of the time...
, a group of mammals so primitive that he describes their cheek teeth as "molariform," as they are neither clearly molars nor premolars. Molariform hypsodonty "cannot be correlated with a grass diet, since grasses were not present." Instead he suggests for his example, Sudamerica ameghinioi, that it lived a "semiaquatic and perhaps a burrowing way of life." Modern species provide many examples from beavers to hippopotamuses.
Attempts have been made to link the wear on E. teeth to grazing. In 1938, H.E. Wood, a Rhinocerotid tooth specialist, pointed out that interproximal wear, or loss of tooth surface between teeth, due to abrasion during mastication, of E. is similar to that of the White Rhinoceros
White Rhinoceros
The White Rhinoceros or Square-lipped rhinoceros is one of the five species of rhinoceros that still exist. It has a wide mouth used for grazing and is the most social of all rhino species...
, the only remaining Rhinocerotid grazer, which has hypsodont teeth. Data such as this led to an intuitive concept of some sort of correlation between grass-eating and hypsodonty, but it has been difficult to isolate mathematically.
A 2007 study by Mendoza and Palmqvist compared the habitats and diets of 134 species of living ungulate
Ungulate
Ungulates are several groups of mammals, most of which use the tips of their toes, usually hoofed, to sustain their whole body weight while moving. They make up several orders of mammals, of which six to eight survive...
s, for which this data is known, both Artiodactyla and Perissodactyla, against "thirty-two craniodental measurements" to discover what correlations exist for the group studied and to test hypotheses concerning hypsodonty and mode of life. Habitats considered were open (savanna, deserts), mixed (wooded savanna, brush) and closed (riverine and forest). Diets considered were grazing, mixed grazing and browsing, browsing, omnivory and special niches, such as treetop browsing. Measurements included the Hypsodonty Index (HI) and Muzzle Width (MZW). The results showed that, except for the "high-level" browsers, hypsodonty is correlated to "open and mixed habitats." The HI was not precise enough to discriminate between open and mixed. However, high MZW is correlated to grazing in the open category, although some forest species also have wide muzzles. Grazers therefore are distinguished by a combination of high-crowned cheek teeth and wide muzzles, both of which are possessed by E. Life in the open is implied.
Cursorial distal limbs
The preponderance of evidence is that E., as far as is known now, was a grassland grazer. It is in this context that Deng and Zheng, experts in the few surviving leg bones, conclude, concerning the morphology of the legs:"Combined with hypsodont cheek teeth with much cement and strong enamel plication, the slender distal limb bones of E. caucasicum indicate that it is cursorial and dwells in an open steppe as a typical grazer."
By distal they mean "furthest outward;" that is, the extremities. Cursorial animals are unequivocally "runners" although the authors did not examine what sort of gait scientifically should be proposed as "running." They selected caucasicum for study because of the availability of a few dozen limb bone fragments from Nihewan, China. These made possible a selective comparison with the fewer bones remaining of other fossil rhinocerotids. In comparison with them, the long legs of E. are the most derived; that is, the others did not have the same cursorial capabilities. The authors approach but do not solve the problem of how to reconcile the weight with the supposed mobility. They say elsewhere in the article that the legs of caucasicum are to be distinguished from those of other fossil Rhinocerotids at Nihewan by their "enormous size."
The White Rhinoceros
White Rhinoceros
The White Rhinoceros or Square-lipped rhinoceros is one of the five species of rhinoceros that still exist. It has a wide mouth used for grazing and is the most social of all rhino species...
at an estimated weight of 2.5 ton has been photographed galloping at a speed of about 30 kph. In a gallop
Gait
Gait is the pattern of movement of the limbs of animals, including humans, during locomotion over a solid substrate. Most animals use a variety of gaits, selecting gait based on speed, terrain, the need to maneuver, and energetic efficiency...
, all feet are off the ground ("ballistic phase") twice a cycle, a feat that elephants, at 2.5-11 ton, cannot perform. They can walk up to 20 kph; however, their straight, relatively inflexible legs are those of striders, not the bent and spring-like legs of gallopers, which utilize haunches, ankle mobility and knee flexion to spring off the ground on alternative legs of a pair. E. legs are sufficiently like those of the White Rhino to hypothesize a similar gait even though E. weighed 4.5-5 ton.
Ceratomorphology
The Ceratomorpha are so-called because their families, such as the Rhinocerotidae, of which E. is undisputably one, are characterized by the presence of hoovesHoof
A hoof , plural hooves or hoofs , is the tip of a toe of an ungulate mammal, strengthened by a thick horny covering. The hoof consists of a hard or rubbery sole, and a hard wall formed by a thick nail rolled around the tip of the toe. The weight of the animal is normally borne by both the sole...
, or horn
Horn (anatomy)
A horn is a pointed projection of the skin on the head of various animals, consisting of a covering of horn surrounding a core of living bone. True horns are found mainly among the ruminant artiodactyls, in the families Antilocapridae and Bovidae...
s and hooves, made of keratin
Keratin
Keratin refers to a family of fibrous structural proteins. Keratin is the key of structural material making up the outer layer of human skin. It is also the key structural component of hair and nails...
, the same substance of which hair
Hair
Hair is a filamentous biomaterial, that grows from follicles found in the dermis. Found exclusively in mammals, hair is one of the defining characteristics of the mammalian class....
is made. These keratin structures appear to have formed in the Mesozoic
Mesozoic
The Mesozoic era is an interval of geological time from about 250 million years ago to about 65 million years ago. It is often referred to as the age of reptiles because reptiles, namely dinosaurs, were the dominant terrestrial and marine vertebrates of the time...
, a remnant in humans being the nail
Nail (anatomy)
A nail is a horn-like envelope covering the dorsal aspect of the terminal phalanges of fingers and toes in humans, most non-human primates, and a few other mammals. Nails are similar to claws, which are found on numerous other animals....
s. A keratin horn is to be distinguished from a bone horn and a tusk
Tusk
Tusks are elongated, continuously growing front teeth, usually but not always in pairs, that protrude well beyond the mouth of certain mammal species. They are most commonly canines, as with warthogs, wild boar, and walruses, or, in the case of elephants and narwhals, elongated incisors...
. Bone forms the base of most horns but in some cases the horn is entirely of bone. A tusk is a modified canine or incisor tooth. Rhinocerotidae have had tusks, but not E. Two open questions are whether they were horned or hornless, hairy or hairless. Most Rhinocerotidae have and have had horns, but there are some instances of hornlessness, and most are or were hairy, such as the Wooly Rhinoceros, but no instances of hair or horn have yet been found for E. Only circumstantial evidence of them exists.
The main evidence suggestive of a horn on E. is a frontal protuberance, which struck the attention of the late 19th century paleontologists and was immediately interpreted as the bony basis for a horn by most investigators from that time forward. A skull of E. sibiricum from the Volga region (cast shown in this article's lead picture) described by Alexander Brandt in the Russian journal, Niwa, and reported in Nature in 1878 offers the following description of the protuberance: hemispherical, 5 inches (12.7 cm) deep, furrowed surface, circumference of 3 foot (0.9144 m). The furrows are interpreted as the seats of blood vessels for the tissues that generated the horn:
"The whole analogy with the rhinoceros points with the greatest certainty to the previous existence of a horn, which, to judge from the size of the blood vessels once encircling the base, must have possessed enormous dimensions."
Brandt was already familiar with the legend of a unicorn among the Tatars of Siberia with a horn so large it required a sledge for transport, and made the connection in interpreting the bump as the base of a horn. He also interpreted the rostrum of the upper mandible as the basis of a nasal horn, a hypothesis now rejected in favor of the cropping labia. In any case the non-circular base indicates a section through the horn would not have been circular. This possibility is supported by another fossil with a non-circular partially healed puncture wound in the base, chiefly interpreted as the result of dueling other males with the horn.
The ungulate
Ungulate
Ungulates are several groups of mammals, most of which use the tips of their toes, usually hoofed, to sustain their whole body weight while moving. They make up several orders of mammals, of which six to eight survive...
s typically combine keratin and bone in various structures. If horns are keratinous, they have a bone core. Rhinocerotids horns, however, are uniquely derived. Hieronymus, an expert in Rhinoceros dermatology
Dermatology
Dermatology is the branch of medicine dealing with the skin and its diseases, a unique specialty with both medical and surgical aspects. A dermatologist takes care of diseases, in the widest sense, and some cosmetic problems of the skin, scalp, hair, and nails....
, says:
"...extant rhinocerotids are unique in possessing a massive entirely keratinous horn that approximates the functions of keratin-and-bone horns such as those of bovidBovidA bovid is any of almost 140 species of cloven-hoofed ruminant mammal at least the males of which bear characteristic unbranching horns covered in a permanent sheath of keratin....
artiodactyls ...."
He defines rhinocerotid horns as:
"...cylindrical blocks of constantly growing cornified papillary epidermisEpidermis (zoology)The Epidermis is an epithelium that covers the body of an eumetazoan . Eumetazoa have a cavity lined with a similar epithelium, the gastrodermis, which forms a boundary with the epidermis at the mouth.Sponges have no epithelium, and therefore no epidermis or gastrodermis...
."
This tissue is "strikingly convergent" with other "cornified epidermis" in horses, cetaceans, artiodactyls and birds. The horn is not attached to the bone of the boss but grows from the surface of a dense dermal tissue. The top layer keratinizes itself to form tubules about 1-2 millimeter high, the cells of which then die. The next layer forms below it. As the layers age the horn loses diameter by degradation of the keratin due to ultraviolet light, desiccation and mechanical wear from contact with objects and agonistic behavior. However, melanin
Melanin
Melanin is a pigment that is ubiquitous in nature, being found in most organisms . In animals melanin pigments are derivatives of the amino acid tyrosine. The most common form of biological melanin is eumelanin, a brown-black polymer of dihydroxyindole carboxylic acids, and their reduced forms...
and calcium deposits in the center harden the keratin there, causing differential wear and shaping of the horn.
The dermis generating the horn is anchored to the boss by interpenetration between rugosities – various irregularities of bone, which it creates by deposition. This tissue is a specialization of dermal armor, which, whenever it attaches to bone, deposits the rugosities to strengthen the attachment. The author states that rugosity is "a bony signature of dermal armor." Cranial rugisoty is an indication, but not a sure sign, of a horn. If, on the other hand, an annular (ring-shaped) pattern is visible in the rugosity, it is due to "stress concentration at the edges of horns" and is "the signature for epidermal horns."
Hieronymus found annular rugosities in all living and some fossil Rhinocerotidae. The rings had previously been noted on additional fossils. To date E. has not been examined for rings under lighting designed to show them up; however, based on the observations of other paleontologists, the author says "squamosal
Squamosal
The squamosal is a bone of the head of higher vertebrates. It is the principal component of the cheek region in the skull, lying below the temporal series and otic notch and bounded anteriorly by postorbital. Posteriorly, the squamosal articulates with the posterior elements of the palatal complex,...
rugosity" is the "most pronounced cranial rugosity in the elasmotherine lineage." This fact suggests an especially firm attachment was required, which, combined with the extraordinarily large hump of muscle for managing the head, suggest a large and heavy horn. In the early 19th century the state of the fossils had not yet revealed the presence of a horn. By 1910 and for all the time since then the paleontologists have not ventured a mathematical estimate but have preferred to refer to the horn as immense, enormous, great or huge. As the size and shape of the horn depended on the concentration of melanin and calcium, and no known indicator of those remains, any further estimate of horn morphology is purely speculative.
In 1968 Björn Kurtén's The Age of Dinosaurs pronounced, without a clue of the reasoning followed, that E. "carried a single frontal horn two meters in length." This figure is often repeated in non-technical books and articles, as it is the only one available. The number is speculative, but coincidentally it happens to be the shoulder height of E. sibiricum. If in fact E. swept snow with the horn, or excavated holes in the terrain with it, the length can hardly have been less to reach the ground.
Taxonomy
The genus is known from hundreds of find sites, mainly of cranial fragments and teeth, but in some cases nearly complete skeletons of post-cranial bones, scattered over Eurasia from Eastern Europe to China. For example, KazakhstanKazakhstan
Kazakhstan , officially the Republic of Kazakhstan, is a transcontinental country in Central Asia and Eastern Europe. Ranked as the ninth largest country in the world, it is also the world's largest landlocked country; its territory of is greater than Western Europe...
alone has 30 sites of E. sibiricum. Dozens of crania have been reconstructed and given archaeological identifiers. The division into species is based mainly on the fine distinctions of the teeth and jaws and the shape of the skull. The finds can be dated only by context.
Classification within the rhinoceros family
The classification of all the species and genera of the Rhinoceros family below the level of Rhinoceratidae has changed more rapidly in the first decade of the 21st century due to first the continued discovery of fossils representing new species and genera from China and Mongolia and second the number of points of morphology to be compared in computerized phylogenetic analysis. Two branches are generally recognized, that leading to the smaller Rhinoceroses and the Pliocene/Pleistocene branch of the larger elasmotheres. Both began from small species in the steppes of the Far East, but those in turn are comparable to North American species. Whether the branch point is considered to be at the subfamily level (Elasmotheriinae versus Rhinocerotinae) or at the tribe level (Elasmotheriini versus Rhinocerotini, the currently accepted version) or whether the elasmotheres existed at the subtribe level (Elasmotheriina, a discontinued taxon) is a matter of the status quo in publications, fossils and software programs.In 2002 P. O. Antoine performed a cladistic analysis using 282 "cranial, dental and postcranial characters" of 28 "terminal taxa" of "elasmotheriines from China and Mongolia" and four outgroups. He found that the elasmotheres were a monophyletic group. He says:
"The main characters of derived elasmotheres (huge size, frontal horn, ossified nasal septum, enamel folding, hypsodonty, loss of anterior dentition, lengthening of the molar series ...) are absent in Middle Miocene Elasmotheriina from China and Mongolia. Most of these features appear somewhat later, during the Late Miocene or the Pliocene, in Parelasmotherium, Sinotherium and then Elasmotherium.
Antoine was following the now discontinued practice of considering the Miocene ancestral species as elasmotheres. They were not, however, distinguished by the features of an elasmothere. The original ancestors were "minute brachydont animals." Elasmotherium was a "mammoth-sized hypsodont." In his view,
"Such cranio-dental evolution demonstrates unequivocally the increasing proportion of grass-eating in the elasmotherine diet throughout the Middle Miocene."
V.A. Terjaev in 1948 had a different view of the cause of the hypsodont
Hypsodont
Hypsodont dentition is characterized by high-crowned teeth and enamel which extends past the gum line. This provides extra material for wear and tear. Some examples of animals with hypsodont dentition are cows, horses and deer; all animals that feed on gritty, fibrous material. The opposite...
y, maintaining that it was caused by the heavier grains in soil on plants pulled from moister environments, and that consequently Elasmotherians lived in "riparian biotopes." Noskova points out that Elasmotheriums diet was comparable to that of the concurrent Archidiskon, which ranged over both steppe and riverland.
Species
Elasmotherium is believed to have descended from the Late PliocenePliocene
The Pliocene Epoch is the period in the geologic timescale that extends from 5.332 million to 2.588 million years before present. It is the second and youngest epoch of the Neogene Period in the Cenozoic Era. The Pliocene follows the Miocene Epoch and is followed by the Pleistocene Epoch...
genus
Genus
In biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...
of Central Asia
Central Asia
Central Asia is a core region of the Asian continent from the Caspian Sea in the west, China in the east, Afghanistan in the south, and Russia in the north...
, Sinotherium
Sinotherium
Sinotherium was a genus of single-horned rhinoceri of the late Miocene and Pliocene. It was ancestral to Elasmotherium, and its fossils have been found in western China....
. Elasmotherium is thought to be the most derived genus of elasmothere, with E. caucasicum in turn being more derived than E. sibiricum. The two Chinese fossils, formerly considered distinct species, E. inexpectatum and E. peii, defined by Chow in 1958, have been sunk into E. caucasicum. They were found in northern China
China
Chinese civilization may refer to:* China for more general discussion of the country.* Chinese culture* Greater China, the transnational community of ethnic Chinese.* History of China* Sinosphere, the area historically affected by Chinese culture...
from the Early Pleistocene
Early Pleistocene
Calabrian is a subdivision of the Pleistocene Epoch of the Geologic time scale. ~1.8 Ma.—781,000 years ago ± 5,000 years, a period of ~.The end of the stage is defined by the last magnetic pole reversal and plunge in to an ice age and global drying possibly colder and drier than the late Miocene ...
Nihewan Faunal Assemblage
Faunal assemblage
Faunal Assemblage is the archaeological or paleontological term for a group of associated animal fossils found together in a given stratum.The principle of faunal succession is used in biostratigraphy to determine each biostratigraphic unit, or biozone...
(from the same valley as nearby Nihewan in Shanxi
Shanxi
' is a province in Northern China. Its one-character abbreviation is "晋" , after the state of Jin that existed here during the Spring and Autumn Period....
Province) and were extinct at approximately 1.6 Ma.
E. caucasicum, defined by Borissjak in 1914, flourished in the Black Sea
Black Sea
The Black Sea is bounded by Europe, Anatolia and the Caucasus and is ultimately connected to the Atlantic Ocean via the Mediterranean and the Aegean seas and various straits. The Bosphorus strait connects it to the Sea of Marmara, and the strait of the Dardanelles connects that sea to the Aegean...
region, as a member of the Early Pleistocene
Early Pleistocene
Calabrian is a subdivision of the Pleistocene Epoch of the Geologic time scale. ~1.8 Ma.—781,000 years ago ± 5,000 years, a period of ~.The end of the stage is defined by the last magnetic pole reversal and plunge in to an ice age and global drying possibly colder and drier than the late Miocene ...
Tamanian Faunal Unit (1.1–0.8 Mya, Taman Peninsula
Taman peninsula
The Taman Peninsula is a peninsula in the present-day Krasnodar Krai of Russia. It is bounded on the north by the Sea of Azov, on the west by the Strait of Kerch and on the south by the Black Sea. The peninsula has evolved over the past two millennia from a chain of islands into the peninsula it is...
). However, an elasmotherian species turned up in the preceding Khaprovian or Khaprov Faunal Complex, which was at first taken to be caucasicum, and then on the basis of the dentition was redefined as a new species, E. chaprovicum (Shvyreva, 2004), named after the Khaprov Faunal Complex. The Khaprov is in the MIddle Villafranchian, MN17, which spans the Piacenzian of the Late Pliocene and the Gelasian of the Early Pleistocene
Early Pleistocene
Calabrian is a subdivision of the Pleistocene Epoch of the Geologic time scale. ~1.8 Ma.—781,000 years ago ± 5,000 years, a period of ~.The end of the stage is defined by the last magnetic pole reversal and plunge in to an ice age and global drying possibly colder and drier than the late Miocene ...
of Northern Caucasus, Moldova
Moldova
Moldova , officially the Republic of Moldova is a landlocked state in Eastern Europe, located between Romania to the West and Ukraine to the North, East and South. It declared itself an independent state with the same boundaries as the preceding Moldavian Soviet Socialist Republic in 1991, as part...
and Asia and has been dated to 2.6–2.2 Mya.
E. sibiricum, described by Johann Fischer von Waldheim
Johann Fischer von Waldheim
Johann Gotthelf Fischer von Waldheim was a German anatomist, entomologist and paleontologist....
in 1808 and chronologically the latest species of the sequence, coming from E. caucasicum in the Middle Pleistocene, ranged from southwestern Russia to western Siberia and southward into Ukraine
Ukraine
Ukraine is a country in Eastern Europe. It has an area of 603,628 km², making it the second largest contiguous country on the European continent, after Russia...
and Moldova
Moldova
Moldova , officially the Republic of Moldova is a landlocked state in Eastern Europe, located between Romania to the West and Ukraine to the North, East and South. It declared itself an independent state with the same boundaries as the preceding Moldavian Soviet Socialist Republic in 1991, as part...
. It appears in the Middle Pleistocene
Middle Pleistocene
The Middle Pleistocene, more specifically referred to as the Ionian stage, is a period of geologic time from ca. 781 to 126 thousand years ago....
Khazar Faunal Complex of the Sea of Azov
Sea of Azov
The Sea of Azov , known in Classical Antiquity as Lake Maeotis, is a sea on the south of Eastern Europe. It is linked by the narrow Strait of Kerch to the Black Sea to the south and is bounded on the north by Ukraine mainland, on the east by Russia, and on the west by the Ukraine's Crimean...
region, which has "no exact stratigraphic situation."
The end of the elasmotheres is questionable; new evidence continues to turn up. The latest is from two caves in southern Siberia
Siberia
Siberia is an extensive region constituting almost all of Northern Asia. Comprising the central and eastern portion of the Russian Federation, it was part of the Soviet Union from its beginning, as its predecessor states, the Tsardom of Russia and the Russian Empire, conquered it during the 16th...
, a sibericum tooth in Smelovskaya Cave 11 and remains from Batpak, both associated with Middle Pleistocene
Middle Pleistocene
The Middle Pleistocene, more specifically referred to as the Ionian stage, is a period of geologic time from ca. 781 to 126 thousand years ago....
relict species, including large herbivores and predators. They must have been dragged into the caves by some predator, perhaps even modern man. The elasmothere tooth and one of a Cave Hyena
Cave Hyena
The Cave Hyena is an extinct subspecies of spotted hyena native to Eurasia, ranging from Northern China to Spain and into the British Isles...
from Smelovskaya were carbon dated to slightly greater than 50,000 BP in the Late Pleistocene
Late Pleistocene
The Late Pleistocene is a stage of the Pleistocene Epoch. The beginning of the stage is defined by the base of the Eemian interglacial phase before the final glacial episode of the Pleistocene 126,000 ± 5,000 years ago. The end of the stage is defined exactly at 10,000 Carbon-14 years BP...
. A recent study of relict Late Pleistocene
Late Pleistocene
The Late Pleistocene is a stage of the Pleistocene Epoch. The beginning of the stage is defined by the base of the Eemian interglacial phase before the final glacial episode of the Pleistocene 126,000 ± 5,000 years ago. The end of the stage is defined exactly at 10,000 Carbon-14 years BP...
remains in the Beringia region (Alaska, Eastern Siberia) identified a pattern of "imbedded micrometeorites" consistent with an extinction event similar to the Tunguska event
Tunguska event
The Tunguska event, or Tunguska blast or Tunguska explosion, was an enormously powerful explosion that occurred near the Podkamennaya Tunguska River in what is now Krasnoyarsk Krai, Russia, at about 7:14 a.m...
of 1908, only carbon-dated to 37,000 BP. A Siberian Elasmotherium skull in a museum was found to have this pattern.
Evolution
The evolution of Rhinocerotidae follows a very similar pattern to the evolution of the EquidaeEquidae
Equidae is the taxonomic family of horses and related animals, including the extant horses, donkeys, and zebras, and many other species known only from fossils. All extant species are in the genus Equus...
, both being facets of the evolution of mammalian herbivores during the Cenozoic
Cenozoic
The Cenozoic era is the current and most recent of the three Phanerozoic geological eras and covers the period from 65.5 mya to the present. The era began in the wake of the Cretaceous–Tertiary extinction event at the end of the Cretaceous that saw the demise of the last non-avian dinosaurs and...
. The overall pattern is based on the evolutionary development of grass
Grass
Grasses, or more technically graminoids, are monocotyledonous, usually herbaceous plants with narrow leaves growing from the base. They include the "true grasses", of the Poaceae family, as well as the sedges and the rushes . The true grasses include cereals, bamboo and the grasses of lawns ...
and grassland
Grassland
Grasslands are areas where the vegetation is dominated by grasses and other herbaceous plants . However, sedge and rush families can also be found. Grasslands occur naturally on all continents except Antarctica...
, and further the changing distribution of C3
C3 carbon fixation
carbon fixation is a metabolic pathway for carbon fixation in photosynthesis. This process converts carbon dioxide and ribulose bisphosphate into 3-phosphoglycerate through the following reaction:...
and C4
C4 carbon fixation
C4 carbon fixation is one of three biochemical mechanisms, along with and CAM photosynthesis, used in carbon fixation. It is named for the 4-carbon molecule present in the first product of carbon fixation in these plants, in contrast to the 3-carbon molecule products in plants. fixation is an...
types of plant metabolism, so-called because they generate a molecule containing 3 or 4 Carbon
Carbon
Carbon is the chemical element with symbol C and atomic number 6. As a member of group 14 on the periodic table, it is nonmetallic and tetravalent—making four electrons available to form covalent chemical bonds...
atoms by different metabolic pathways (sequences of chemical reactions).
Evolution of Rhinocerotidae
The Rhinocerotoids of the early EoceneEocene
The Eocene Epoch, lasting from about 56 to 34 million years ago , is a major division of the geologic timescale and the second epoch of the Paleogene Period in the Cenozoic Era. The Eocene spans the time from the end of the Palaeocene Epoch to the beginning of the Oligocene Epoch. The start of the...
and subsequently the Hyracodontidae
Hyracodontidae
Hyracodontidae is an extinct family of rhinoceroses endemic to North America, Europe, and Asia during the Eocene through early Miocene living from 55.8—20 mya, existing for approximately .They are typified as having long limbs and having no horns...
and Amynodontidae
Amynodontidae
The Amynodonts were a group of hippo-like perissodactyls, related to true rhinoceri, that were descended from the Hyracodontidae. They ranged from North America, Europe and Asia during the Late Eocene to Miocene living from 46.2 Ma—7 Ma years ago and existed for approximately .The last species died...
show no sign of dermal armor or horns. The oldest known genus of Rhinocerotids, or true Rhonoceroses, is Teletaceras from the Middle Eocene
Eocene
The Eocene Epoch, lasting from about 56 to 34 million years ago , is a major division of the geologic timescale and the second epoch of the Paleogene Period in the Cenozoic Era. The Eocene spans the time from the end of the Palaeocene Epoch to the beginning of the Oligocene Epoch. The start of the...
of North America and the Late Eocene of Asia. Due to the timing of bridges, it does not appear in South America. It features the first known Rhinocerotid tusks, a derived feature, which are still extant in three species and are "the primary offensive weapons" of those species. They appear to have been sexually dimorphic features convergent with the development of tusks in pigs, hippopotamoses and elephant seals.
The first dermal armor is indicated by cranial rugosity on Trigonias
Trigonias
Trigonias is an extinct genus of rhinoceros from the late Eocene some 35 million years ago of North America ....
osborni, Late Eocene
Eocene
The Eocene Epoch, lasting from about 56 to 34 million years ago , is a major division of the geologic timescale and the second epoch of the Paleogene Period in the Cenozoic Era. The Eocene spans the time from the end of the Palaeocene Epoch to the beginning of the Oligocene Epoch. The start of the...
, 42-32 Mya. It appears more developed in Subhyracodon
Subhyracodon
Subhyracodon is an extinct genus of cow-sized rhinoceroses. It was a medium sized herbivore on the plains of early Oligocene South Dakota 33 million years ago , smaller than only the Brontops and the chalicotheres. Subhyracodon had no horns, relying more on its speed to escape, but a species found...
, early Oligocene
Oligocene
The Oligocene is a geologic epoch of the Paleogene Period and extends from about 34 million to 23 million years before the present . As with other older geologic periods, the rock beds that define the period are well identified but the exact dates of the start and end of the period are slightly...
, 33 Mya, both of North America. There appears to have been a correlation between the tusks and the armor later (4 Mya later) defending against them.
Having begun to diversify (increase in species and genera) in America the Rhinocerotids spread rapidly over Beringia when that bridge appeared in the Early Oligocene
Oligocene
The Oligocene is a geologic epoch of the Paleogene Period and extends from about 34 million to 23 million years before the present . As with other older geologic periods, the rock beds that define the period are well identified but the exact dates of the start and end of the period are slightly...
to Eurasia, where subsequently they became highly diverse. The family reached a maximum floruit in the Miocene
Miocene
The Miocene is a geological epoch of the Neogene Period and extends from about . The Miocene was named by Sir Charles Lyell. Its name comes from the Greek words and and means "less recent" because it has 18% fewer modern sea invertebrates than the Pliocene. The Miocene follows the Oligocene...
. At that time they entered Africa, but never were very diverse there. In the Pliocene
Pliocene
The Pliocene Epoch is the period in the geologic timescale that extends from 5.332 million to 2.588 million years before present. It is the second and youngest epoch of the Neogene Period in the Cenozoic Era. The Pliocene follows the Miocene Epoch and is followed by the Pleistocene Epoch...
they declined, disappearing from America. The Pleistocene
Pleistocene
The Pleistocene is the epoch from 2,588,000 to 11,700 years BP that spans the world's recent period of repeated glaciations. The name pleistocene is derived from the Greek and ....
brought extinction from Eurasia but African and Southeast Asian species survived.
Elasmotherium appeared in the Late Pliocene
Pliocene
The Pliocene Epoch is the period in the geologic timescale that extends from 5.332 million to 2.588 million years before present. It is the second and youngest epoch of the Neogene Period in the Cenozoic Era. The Pliocene follows the Miocene Epoch and is followed by the Pleistocene Epoch...
, apparently deriving from the Miocene—Pliocene Sinotherium
Sinotherium
Sinotherium was a genus of single-horned rhinoceri of the late Miocene and Pliocene. It was ancestral to Elasmotherium, and its fossils have been found in western China....
, sometimes informally aggregated with it. Sinotherium in turn appeared to be related to Iranotherium
Iranotherium
Iranotherium was a large elasmothere rhinoceros, as big as a modern white rhino, found in Central Asia. It was a precursor of the related Sinotherium, and may have been ultimately outcompeted by its descendant....
; that is, Elasmotherium was believed to have been an iranotheriine. A 1995 cladistic analysis by Cerdeño asserted that Elasmotherium was most closely related to Coelodonta, the "Wooly Rhinoceros," whose soft-tissue corpus has been found in a few locations. By implication, Sinotherium was not an iranothere. However, Antoine's 2002 analysis reaffirmed that the elasmotheriine line was iranotheriine. Certainty concerning the exact line of descent remains elusive; fossils and techniques of the future will perhaps resolve it.
Evolution of grassland
C3 plant metabolism was chronologically first. It requires large amounts of water (which is lost through transpiration), moderate temperatures and moderate sunlight. About 95% of plants are C3. In places where the water supply is minimal, the temperature is too high, or the sunlight too bright, C3 plants cannot thrive, but C4 metabolism evolved at multiple locations convergently within the same families by a slight alteration of the metabolic process. It requires a fraction of the water, and thrives in warmer environments and bright sunlight. C4 plants spread where C3 plants must decline. About 5% of the Earth's plants are C4, far less than for grasses, for which the number is 46%.The grass of grasslands belongs predominantly to the Poaceae
Poaceae
The Poaceae is a large and nearly ubiquitous family of flowering plants. Members of this family are commonly called grasses, although the term "grass" is also applied to plants that are not in the Poaceae lineage, including the rushes and sedges...
family, also called the "true grasses." Species can be divided into "warm season grasses," which utilize C4, and "cool season grasses," which utilize C3. The evolution of grasses preceded the evolution of grasslands, which were created by the spreading of grasses from the forest onto the plains in the Miocene
Miocene
The Miocene is a geological epoch of the Neogene Period and extends from about . The Miocene was named by Sir Charles Lyell. Its name comes from the Greek words and and means "less recent" because it has 18% fewer modern sea invertebrates than the Pliocene. The Miocene follows the Oligocene...
. These initial grasslands were C3, which happen to be more nutritious than C4 species.
The evolution of Rhinocerotidae and Equidae in the Miocene grasslands proceeded along parallel lines from similar causes. Diminutive, cursorial, browsing, brachydont species expanded their ranges from the Oligocene
Oligocene
The Oligocene is a geologic epoch of the Paleogene Period and extends from about 34 million to 23 million years before the present . As with other older geologic periods, the rock beds that define the period are well identified but the exact dates of the start and end of the period are slightly...
forests into the nutritious C3 Miocene grasslands and there over the millions of years of the Miocene evolved diverse and flourishing populations on the ample food supply, gradually increasing in size. Some of these species exploited the grazing niche. Hispanotherium
Hispanotherium
Hispanotherium was a genus of rhinoceros of the tribe Elasmotheriini endemic to Europe and Asia during the Miocene living from 16—7.25 mya existing for approximately .-Taxonomy:...
, Chilotherium
Chilotherium
Chilotherium is an extinct genus of rhinoceros endemic to Eurasia and Asia during the Miocene through Pliocene living for 13.7—3.4 mya, existing for approximately . They are known for their large, protruding lower canine teeth.-Taxonomy:...
and Teleoceras
Teleoceras
Teleoceras is an extinct genus of grazing rhinoceros that lived in North America during the Miocene epoch, which ended about 5.3 million years ago, all the way to the early Pliocene epoch....
"show a clear tendency to hypsodonty, and a grazing feeding adaptation has usually been attributed to them."
By the late Miocene, C4 metabolism was well established on the plains. In place of the prehistoric C3 grasslands (whuch are no longer extant) was the savanna
Savanna
A savanna, or savannah, is a grassland ecosystem characterized by the trees being sufficiently small or widely spaced so that the canopy does not close. The open canopy allows sufficient light to reach the ground to support an unbroken herbaceous layer consisting primarily of C4 grasses.Some...
, today's subtropical mixture of grasses and sedge
Cyperaceae
Cyperaceae are a family of monocotyledonous graminoid flowering plants known as sedges, which superficially resemble grasses or rushes. The family is large, with some 5,500 species described in about 109 genera. These species are widely distributed, with the centers of diversity for the group...
s. The deciding factors in this turn of evolution were a documented increase in temperature and aridity over the period. Faced with scant nutrition from browsing, the smaller Rhinocerotidae and Equidae declined in diversity, the gate in that direction, so to speak, now being closed.
The evolutionary response in the Pliocene
Pliocene
The Pliocene Epoch is the period in the geologic timescale that extends from 5.332 million to 2.588 million years before present. It is the second and youngest epoch of the Neogene Period in the Cenozoic Era. The Pliocene follows the Miocene Epoch and is followed by the Pleistocene Epoch...
was in the direction of bulk-feeding. Massive grazers, such as Elasmotherium, maximally hypsodont, moved across vast ranges consuming C4 grass in bulk. While the horses and other more nearly modern grazers could rely on gallop
Gait
Gait is the pattern of movement of the limbs of animals, including humans, during locomotion over a solid substrate. Most animals use a variety of gaits, selecting gait based on speed, terrain, the need to maneuver, and energetic efficiency...
ing for mobility in any circumstance, there is a question as to what degree the brachypodial giants can be said to be "cursorial." Elephants, for example, must keep one foot on the ground, where true gallopers are entirely off it for brief intervals. The horns and impenetrable body armor insured that the giants would not be required to gallop away for any predator. At least until the arrival of man, they were, as the legends relate, lords of the savanna.
Possible historical witnesses
Legends and names referring to a special type of unicornUnicorn
The unicorn is a legendary animal from European folklore that resembles a white horse with a large, pointed, spiraling horn projecting from its forehead, and sometimes a goat's beard...
or mysterious beast with one horn, as opposed to the delicate imaginary unicorns of Europe based on narwhal
Narwhal
The narwhal, Monodon monoceros, is a medium-sized toothed whale that lives year-round in the Arctic. One of two living species of whale in the Monodontidae family, along with the beluga whale, the narwhal males are distinguished by a characteristic long, straight, helical tusk extending from their...
horns, are scattered across the former range of Elasmotherium from China to Eastern Europe. They have been noted there since the first known literature of the Middle Ages
Middle Ages
The Middle Ages is a periodization of European history from the 5th century to the 15th century. The Middle Ages follows the fall of the Western Roman Empire in 476 and precedes the Early Modern Era. It is the middle period of a three-period division of Western history: Classic, Medieval and Modern...
.
Eastern Eurasia
The Chinese k'i-lin, referring to some sort of beast, was loaned in various forms into the TurkicTurkic languages
The Turkic languages constitute a language family of at least thirty five languages, spoken by Turkic peoples across a vast area from Eastern Europe and the Mediterranean to Siberia and Western China, and are considered to be part of the proposed Altaic language family.Turkic languages are spoken...
and Mongolic languages
Mongolic languages
The Mongolic languages are a group of languages spoken in East-Central Asia, mostly in Mongolia and surrounding areas plus in Kalmykia. The best-known member of this language family, Mongolian, is the primary language of most of the residents of Mongolia and the Mongolian residents of Inner...
and lore. The writers of works in those languages have not known how to depict these strange beasts. A common theme is that they have a single horn, are dark and are huge.
For example, a book of divination in the Old Turkic language
Old Turkic language
Old Turkic is the earliest attested form of Turkic, found in Göktürk and Uyghur inscriptions dating from about the 7th century to the 13th century....
uses kälän käyik, "tawny beast," (the kilin is altered to kälän by vowel harmony
Vowel harmony
Vowel harmony is a type of long-distance assimilatory phonological process involving vowels that occurs in some languages. In languages with vowel harmony, there are constraints on which vowels may be found near each other....
). The Han I Araha Manju Gisun I Buleku Bithe, "Mirror of the Manchu Language," dating from the 18th century, defines the kilin as
"...a quadruped with the body of a deer, the tail of a cow, the head of a sheep, the limbs of a horse, the hooves of a cow, and a big horn. His body is of five colors, his height a dozen feet. Of a gentle nature, he does not crush the worms or beak the plants."
The Chinese had other unicorns, such as the Xiezhi, but the latter was less like a real animal and more a symbolic figure. It had the ability to discriminate right from wrong and therefore a representaton of it was placed in the courtrooms of the Han Dynasty
Han Dynasty
The Han Dynasty was the second imperial dynasty of China, preceded by the Qin Dynasty and succeeded by the Three Kingdoms . It was founded by the rebel leader Liu Bang, known posthumously as Emperor Gaozu of Han. It was briefly interrupted by the Xin Dynasty of the former regent Wang Mang...
and, in the 3rd and 4th centuries, on the caps of judges. It looks like a seated or standing lion with a horn rising from the top of the head. In contrast to this mythical beast, the k'i-lin is presented as an animal actually caught by hunting parties of the past. The Shiji
Records of the Grand Historian
The Records of the Grand Historian, also known in English by the Chinese name Shiji , written from 109 BC to 91 BC, was the Magnum opus of Sima Qian, in which he recounted Chinese history from the time of the Yellow Emperor until his own time...
("Historical Records") of Sima Qian
Sima Qian
Sima Qian was a Prefect of the Grand Scribes of the Han Dynasty. He is regarded as the father of Chinese historiography for his highly praised work, Records of the Grand Historian , a "Jizhuanti"-style general history of China, covering more than two thousand years from the Yellow Emperor to...
in the 2nd century mentions the capture of a "deer-like" animal with one horn in 122 BC. The Chunqin, an encyclopedia of the 3rd century BC, states that a unicorn was captured in 481 BC, but without any description. Chinese representations of unicorns vary quite a lot, but an engraving on a bronze vessel of the Warring States Period
Warring States Period
The Warring States Period , also known as the Era of Warring States, or the Warring Kingdoms period, covers the Iron Age period from about 475 BC to the reunification of China under the Qin Dynasty in 221 BC...
, shows an animal very like the cave paintngs supposed to be of Elasmotherium: head down for grazing, horn protruding horizontally from the forehead, neck and shoulders humped.
In the northeast of the range, in 1866 Vasily Radlov
Vasily Radlov
Vasily Vasilievich Radlov or Friedrich Wilhelm Radloff was a German-born Russian founder of Turkology, a scientific study of Turkic peoples....
reported a legend among the Yakuts
Yakuts
Yakuts , are a Turkic people associated with the Sakha Republic.The Yakut or Sakha language belongs to the Northern branch of the Turkic family of languages....
of Siberia
Siberia
Siberia is an extensive region constituting almost all of Northern Asia. Comprising the central and eastern portion of the Russian Federation, it was part of the Soviet Union from its beginning, as its predecessor states, the Tsardom of Russia and the Russian Empire, conquered it during the 16th...
of a "huge black bull" killed by spear, who had "a single horn" so large it required a sledge for transportation. In 1878 A. F. Brandt suggested the beast was an Elasmotherium.
Western Eurasia
From medieval northern Russia, probably Veliky NovgorodVeliky Novgorod
Veliky Novgorod is one of Russia's most historic cities and the administrative center of Novgorod Oblast. It is situated on the M10 federal highway connecting Moscow and St. Petersburg. The city lies along the Volkhov River just below its outflow from Lake Ilmen...
, a collection of ballads has survived, combined into a spiritual theme deriving ultimately from Zoroastrianism
Zoroastrianism
Zoroastrianism is a religion and philosophy based on the teachings of prophet Zoroaster and was formerly among the world's largest religions. It was probably founded some time before the 6th century BCE in Greater Iran.In Zoroastrianism, the Creator Ahura Mazda is all good, and no evil...
, but with Christian overtones, called the Stikh o Golubinioi knige, or in another version the Golubiniai kniga, "The Book of the Dove." The "dove," or golub of the Christian work is probably an alteration of glub, "depth," so that the reconstructed name, *Glubinnaia kniga, is on the same theme as the Zend Avesta and similar works in Iran
Iran
Iran , officially the Islamic Republic of Iran , is a country in Southern and Western Asia. The name "Iran" has been in use natively since the Sassanian era and came into use internationally in 1935, before which the country was known to the Western world as Persia...
, the struggle between cosmic good and evil.
The force of pravda, or "rightness," is symbolized by a beast called mainly the Indrik
Indrik
In Russian folklore, the Indrik-Beast is a fabulous beast, the king of all animals, who lives on a mountain known as "The Holy Mountain" where no other foot may tread. When it stirs, the Earth trembles...
, but also Indra, Beloiandrikh, Kondryk, Edinrog and Edinor. The name is a calque
Calque
In linguistics, a calque or loan translation is a word or phrase borrowed from another language by literal, word-for-word or root-for-root translation.-Calque:...
: Greek mono-keros ("single-horn") > ino-rog (in, "one"; rogom, "horn") > edinrog > Indrik. As a unicorn struggling with a lion representing krivda, "lies," the Indrik has the body of a goat or a horse in representations.
An Indrik composite from all the variants of the ballads depicts the Indrik as dwelling in the Holy Mountain, which it consumes for sustenance. It is the mother or the father of all animals. It wanders unseen on the plain by day. When other beasts encounter it they must do obeisance. The Indrik saves the world from draught by digging springs of pure water. By night it wanders in the subterranean world, forging a path with its horn. There it purifies all the waters and releases them from any blockages.
The righteous but fantastic unicorn appears also in the Bundahišn
Bundahishn
Bundahishn, meaning "Primal Creation", is the name traditionally given to an encyclopædiaic collections of Zoroastrian cosmogony and cosmology written in Book Pahlavi. The original name of the work is not known....
or Zand Āgāhīh, a Middle Persian
Middle Persian
Middle Persian , indigenously known as "Pârsig" sometimes referred to as Pahlavi or Pehlevi, is the Middle Iranian language/ethnolect of Southwestern Iran that during Sassanid times became a prestige dialect and so came to be spoken in other regions as well. Middle Persian is classified as a...
Zoroastrian composition concerning the "deep and miraculous utterances" about the creation, the archetype of the *Gubinnaia kniga, "Book of the Deep." Its unicorn is an ass of 3 feet, 6 eyes, 9 mouths, 2 ears, 1 horn, a dark blue head and a white body, which is a symbolic rather than a real entity. It stands in the sea of Fraxwkand (wherever that is) and purifies the waters by urinating in them so that life may continue. The Russian concept of a unicorn was apparently based on a different experience.
In fact the Arabo-Persian word for unicorn in the region was karkadann
Karkadann
The Karkadann was a mythical creature said to live on the grassy plains of India, Persia, and North Africa. Referred to by Elmer Suhr as the "Persian version of the unicorn," its mention by Muslim writers may depend on earlier Greek texts and may have an origin in an account from the Mahabharata...
, which extended also to India
India
India , officially the Republic of India , is a country in South Asia. It is the seventh-largest country by geographical area, the second-most populous country with over 1.2 billion people, and the most populous democracy in the world...
, and was loaned into the Turkmen language
Turkmen language
Turkmen is the national language of Turkmenistan...
as kergeden and the Ottoman Turkish language
Ottoman Turkish language
The Ottoman Turkish language or Ottoman language is the variety of the Turkish language that was used for administrative and literary purposes in the Ottoman Empire. It borrows extensively from Arabic and Persian, and was written in a variant of the Perso-Arabic script...
as gergeden, among others. The word had an equivocal meaning: unicorn and rhinoceros. This rhinoceros has capabilities and habits not possessed by the Indian rhinoceros
Indian Rhinoceros
The Indian Rhinoceros is also called Greater One-horned Rhinoceros and Asian One-horned Rhinoceros and belongs to the Rhinocerotidae family...
: it indulges in mortal combat with elephants, which it kills by goring underneath with its horn. The symbol of evil is thus transferred to the elephant in that milieu; however, the two conduct their combats over grasslands never populated by either in historical times. The root of the word is Sanskrit
Sanskrit
Sanskrit , is a historical Indo-Aryan language and the primary liturgical language of Hinduism, Jainism and Buddhism.Buddhism: besides Pali, see Buddhist Hybrid Sanskrit Today, it is listed as one of the 22 scheduled languages of India and is an official language of the state of Uttarakhand...
Khadga-, Khadga dhenu-, "rhinoceros." In Christianity the single horn becomes a symbol of monotheism
Monotheism
Monotheism is the belief in the existence of one and only one god. Monotheism is characteristic of the Baha'i Faith, Christianity, Druzism, Hinduism, Islam, Judaism, Samaritanism, Sikhism and Zoroastrianism.While they profess the existence of only one deity, monotheistic religions may still...
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Science writer and cryptozoologist Willy Ley
Willy Ley
Willy Ley was a German-American science writer and space advocate who helped popularize rocketry and spaceflight in both Germany and the United States. The crater Ley on the far side of the Moon is named in his honor.-Life:...
theorised that these legends have simply been passed down since a hunting association with the beast in prehistoric times, and that the animals did not actually survive into recorded history. One argument for survival is the testimony by the medieval traveller, Ibn Fadlan. His account states:
See also
- AegyrcitheriumAegyrcitheriumAegyrcitherium was a genus of rhinoceros endemic to Europe during the Miocene living from 16.9—16 mya existing for approximately .-Taxonomy:...
- BugtirhinusBugtirhinusBugtirhinus was a genus of rhinoceros of the tribe Elasmotheriini endemic to Asia during the Miocene living from 20—16.9 mya existing for approximately .-Taxonomy:...
- HispanotheriumHispanotheriumHispanotherium was a genus of rhinoceros of the tribe Elasmotheriini endemic to Europe and Asia during the Miocene living from 16—7.25 mya existing for approximately .-Taxonomy:...
- SinotheriumSinotheriumSinotherium was a genus of single-horned rhinoceri of the late Miocene and Pliocene. It was ancestral to Elasmotherium, and its fossils have been found in western China....
- Tahash