Elasmotherium

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Elasmotherium
Skeletal reconstruction of Elasmotherium caucasicum

Skeletal reconstruction of Elasmotherium caucasicum

Temporal occurrence
Upper Pliocene to Young Pleistocene
3.6 million years to 50,000 years
Locations
Systematics
Higher mammals (Eutheria)
Laurasiatheria
Unpaired ungulate (Perissodactyla)
Rhinocerotoidea
Rhinoceros (Rhinocerotidae)
Elasmotherium
Scientific name
Elasmotherium
Fischer , 1808

Elasmotherium is a genus of rhinoceros thatoccurredin the Pliocene and Pleistocene in northern Eurasia and belongs to the Elasmotheriini group, a form of rhinoceros with usually only one horn. It survived at least until the beginning of the New Pleistocene and probably died out during the last glacial period . Their area of ​​distribution included the steppe areas and river bank landscapes of Eastern Europe as well as Central and East Asia . Elasmotherium is one of the largest known representatives of the rhinos. The animals were of massive build and adapted to the open regions. A prominent feature can be found in the dome-like bonybulgeon the frontal bone of the skull, on which a single horn sat. Due to the lack of fossil conservation, it is unclear how massive this horn was. In addition, the animals had the most reduced number of teeth of all rhinos. The teeth were extremely high crowned andcharacterizedby conspicuously folded enamel on the chewing surface. Throughout the teeth structure can Elasmotherium thus be regarded as extremely specialized, mainly for the phylogenetically applies youngest members of the genus. Little is known about the way of life. The animals probably lived in open landscapes, which were interspersed with water-rich areas. There they probably lived on marsh plants. Overall, the individual species of Elasmotherium rarely appear in the fossil record compared to other, simultaneously existing rhino forms such as the woolly rhinoceros . The genus was first described in 1808 using a lower jaw. Their affiliation with the rhinos was only recognized later. The generic name is made up of the Greek words έλασμα (elasma) for "plate" and θηρίον (thēríon) for "animal" and refers to the special structure of the teeth. There are four species recognized today.

features

Elasmotherium is one of the least studied fossil rhino genera. All in all, they were very robust and large rhinos. The limbs were relatively long in relation to the body. The forelegs ended in three toes each ( metacarpus II - IV), as with the recent rhinos , with the middle one being the largest. A fourth toe (Metacarpus V) was only rudimentary . The hind legs also had three toes each. The rest of the physique was relatively strong: the spinous processes on the thoracic vertebrae, where the massive muscles used to stabilize the head, reached lengths of over 50 cm.

Skull of Elasmotherium sibiricum in the Museum für Naturkunde Berlin

The skull was also very massive and large with a sturdy lower jaw. He had a wide and elongated occiput with a strongly developed bone crest. The occiput, which is clearly stretched and thus angled when viewed from the side, caused the animal to hold its head down at an angle, similar to what is now the case with the white rhinoceros ( Ceratotherium simum ). The nasal bone , on the other hand, was built slender, making the skull clearly wedge-shaped when viewed from above. A long and narrow symphysis on the lower jaw corresponded to this . Furthermore, the nasal septum was completely ossified in the adult animal. This feature does not occur in the rhinoceros species living today and is otherwise only known in fossil form in the genus Coelodonta , here especially in the woolly rhinoceros ( Coelodonta antiquitatis ), and partly from some representatives of the genus Stephanorhinus . Since Elasmotherium is not directly related to each other with the two genera, this characteristic must have developed independently of each other ( convergent evolution ).

A dome-like or dome-like elevation on the frontal bone was striking, protruding about 15 cm above the skull and having a diameter of 25 to 35 cm. This elevation represented the base of the horn, inside it consisted of the enlarged frontal sinuses, which were in connection with the nasal cavity, possibly it also served as an expansion space for the olfactory organs to make it easier to find the food . Nothing is known about the appearance of the keratin horn due to the lack of soft tissue tradition, but the dimensions of the horn base suggest a massive and large horn. The Horn location on the forehead differs Elasmotherium significantly from the recent one-horned rhinos as the Indian rhinoceros ( Rhinoceros unicornis ), who wear their horn on the nose.

More recently it has been possible to demonstrate a sexual dimorphism based on the skull, at least for the late representatives of Elasmotherium . Thus, cows were generally characterized by a slimmer skull structure with a less pronounced occipital ridge. Furthermore, the dome-like elevation of the skull was less prominent.

Even the bite of Elasmotherium showed characteristic properties because it was greatly reduced and only two Vormahlzähnen and three molars was ever arch. Accordingly, the dental formula for the permanent dentition : . The presence of the alveoli of the incisors , albeit in a rudimentary state, suggests that incisors in the deciduous teeth of the young animals were still developed. The lack of incisors in the permanent dentition was compensated for by flat plate-like bone formations on the upper and lower jaw, which may have been lip-like formations and are interpreted as an organ for plucking grass, similar to the case with the recent African rhinos. Noteworthy are the molars , which had a distinctly rectangular shape with distinctively folded enamel , with the fold lines in the younger representatives also showing a sinusoidal course. The strong folding led to the name of the genus. In addition, the molars were extremely crowned and rootless ( euhypsodont ). The extreme high crown is the strongest known specialization in the rhinoceros family and the order of the odd-toed ungulate and is only surpassed among the mammals by some representatives of the rodents .

Way of life

Live reconstruction of Elasmotherium

Elasmotherium is one of the largest known mammals of the Pliocene and Pleistocene in Eurasia . Due to its massive body, it should therefore be adapted to productive, nutrient-rich landscapes, for example to long grassy steppes or to the edge of the river. The high-crowned teeth with the conspicuously folded tooth enamel indicate highly abrasive food material. Such teeth are often interpreted as an adaptation to grass forage. Since grasses contain fewer nutrients, an animal has to eat more accordingly, which accelerates the abrasion of the teeth. This is additionally reinforced by a high silica content, which makes the high tooth crowns necessary for food exclusively containing grass. However, the narrow snout of Elasmotherium speaks against full specialization in grass food . This was well suited to looking for food plants in damp subsoil. The view is also supported by the fact that at least the late representatives of Elasmotherium were found in large parts of their range, for example in water-rich southern Siberia, in alluvial forests on river banks or on lake banks. As a result, it is likely that the animals fed on marsh plants, the lower sections of which were enriched with soil and thus had a high mineral content , which in turn caused abrasion of the molars. In the case of root-rich food, which may have been torn from the ground along with the earth, this effect was even more pronounced . However, some early forms show a wear pattern on the teeth that suggests a mixed vegetable diet of hard and soft components. Overall, the reconstructed low position of the head indicates a predominant search for food on the ground. This view is supported by isotope analyzes , which resulted in extremely high nitrogen and carbon values for Elasmotherium . The values ​​are outside the variation of other rhinos such as the woolly rhinoceros and are comparable to those of some local populations of today's saigas . In this way, among other things, an increased consumption of foxtail plants and / or roots can be assumed. However, contrary to some other postulated opinions, Elasmotherium was not adapted to a semi-aquatic way of life. Body proportions in general and long legs in particular speak against such a way of life. These mostly represent adaptations to open grasslands. Apparently, the animals moved on the migrations from their feeding grounds in the floodplain or bank areas through adjacent or intermediate steppe landscapes. The habitat of the rhinoceros genus can thus be assumed to be an open landscape with numerous enclosed water biotopes.

Little is known about the further way of life. The position of the horn on the forehead suggests that it was not used or only to a limited extent in dominance or territorial fights, in contrast to today's rhinos. Since the (larger) horn is on the nose of today's rhinos, it can be used much more effectively, as more force is transferred to the horn via the neck muscles when the head is moved. It is therefore assumed that fights between individual representatives of Elasmotherium were highly ritualized. In addition, due to its location, the horn probably had no function in foraging, as is sometimes the case with today's rhinos. Elasmotherium probably did not have natural enemies due to its size. However, the skull of Atyrau (formerly Gurjew) shows a round skull injury in the forehead area, which however healed again during the life of the animal. It is also assumed that the animals may have had fur due to their relatively northern distribution .

Systematics

Internal systematics of the Elasmotheriini according to Sanisidro et al. 2012 and Deng 2008
  Rhinocerotinae  

 Menoceratini


   
  Elasmotheriini  

 Bugtirhinus


   

 Kenyatherium


   

 Caementodon


   

 Hispanotherium


   

 Procoelodonta


   

 Huaqingtherium


   

 Iranotherium


   

 Ningxiatherium


   

 Parelasmotherium


   

 Sinotherium


   

 Elasmotherium










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 Rhinocerotini




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Relationship of rhinos based on genetic data from Margaryan et al. 2020
  Rhinocerotinae  
  Elasmotheriini 

 Elasmotherium


  Rhinocerotini 
  Dicerorhinina 


 Stephanorhinus


   

 Coelodonta



   

 Dicerorhinus



   
  Dicerotina 

 Ceratotherium


   

 Diceros



  Rhinocerotina 

 Rhinoceros





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Elasmotherium is a genus of the rhinoceros family . Within the family it belongs to the subfamily Rhinocerotinae and to the tribe of the Elasmotheriini. The tribe includes rhinos with a generally large to very large body and extremely high tooth crowns. With reference to the latter characteristic, the Elasmotherien represent, at least in their late development phase, the most specialized rhinoceros representatives. They also form the sister group of the Rhinocerotini, which in turn includes the rhinoceros species living today. Both rhinoceros lines can show a long independent development history, as their separation according to molecular genetic data had already taken place in the Paleogene . Depending on the study, the determined period ranges from the Middle Eocene , about 47 million years ago, to the Lower Oligocene , 31 million years ago. The closest relative of Elasmotherium is Sinotherium , both forms are combined by some scientists to form the sub-tribus of Elasmotheriina. Their characteristic feature can be found in the formation of horns on the forehead. Opposite the Elasmotheriina are the Iranotheriina, which are older Elasmotheriina with a nasal horn. In addition, the latter generally have less high-crowned molars and less folded enamel. These sub-tribus are assigned to the eponymous Iranotherium, among others, Ningxiatherium , Parelasmotherium or Hispanotherium . The distinction between Elasmotheriina and Iranotheriina was first made by the Hungarian paleontologist Miklós Kretzoi (1907-2005) in 1943.

Four species from the genus Elasmotherium are currently recognized. These include the largely oberpliozäne Elasmotherium chaprovicum , the two forms occur in the Early Pleistocene Elasmotherium caucasicum or Elasmotherium PEII and to Upper Pleistocene living from Central sibiricum Elasmotherium . Some authors reduce the number of species by referring to metric features on the skeletal material to only two species, mostly Elasmotherium caucasicum and Elasmotherium sibiricum , whereby the other two species are regarded as synonymous with the former, but this is often rejected.

species

Elasmotherium chaprovicum

Distribution of the four species of Elasmotherium in northern Eurasia. The markings represent important sites.

The oldest representative of this rhinoceros genus is Elasmotherium chaprovicum , which was widespread in the Black Sea region in the Upper Pliocene 3.6 to 2.6 million years ago . Only a few bones and teeth of this type are known to date. It belongs to the Chapri fauna complex and is associated with early representatives of the southern elephant , the Etruscan rhinoceros ( Stephanorhinus etruscus ) and Paracamelus . Type sources are Chapri and Liwenzowka (both Ukraine ). At first it was counted to the species Elasmotherium caucasicum , but differences to this species were quickly recognized. These mainly affect the dental apparatus; The size of the teeth is comparable to that of Elasmotherium caucasicum and significantly larger than that of Elasmotherium sibiricum , but they differ from the former in the structure and structure of the tooth enamel. Furthermore, the limbs are significantly shorter than those of the Caucasian variant and reach the dimensions of the Siberian representative, but are significantly more massive than this. However, as there is too little find material so far, it is currently difficult to differentiate. First described was Elasmotherium chaprovicum of Anna Konstantinovna Schwyrjowa 2004 in the light of teeth from Liwenzowka.

Elasmotherium caucasicum

Elasmotherium caucasicum was the largest member of the genus and reached a length of 5 to 5.2 m. The reconstructed weight of 5 t is comparable to today's fully grown Asian elephant , making the rhinoceros one of the largest of the Pleistocene. So far, the species is only known about individual skeletal elements, including teeth and long bones - complete skeletons have not yet been found. In general, the physique was similar to that of Elasmotherium sibiricum . The limbs, however, were more powerfully built and much longer, and they also showed some ancient features compared to their Siberian relatives. Compared to the arm and leg bones of other rhinos, those of Elasmotherium caucasicum stand out due to their enormous size. Those of the white rhinoceros are around 50% smaller and 30% narrower. The teeth were characterized by a slightly smaller crown height and their general size, while the enamel was clearly folded, but the fold lines did not yet have such a distinctive sinus-like shape as was the case with the later Elasmotherium sibiricum . The last premolar tooth, which was clearly massive, is remarkable, which suggests that this species was not yet as well adapted to life in open terrain as the later Elasmotherium sibiricum .

The rhinoceros species occurred in Central Asia and the Black Sea region and has been documented in the younger Old Pleistocene; it occurred here 1.1 to 0.8 million years ago. A relatively dense detection area is on the Taman Peninsula between the Black Sea and the Sea of ​​Azov . Here alone there are at least 230 fossil finds of Elasmotherium caucasicum , the classic Sinaya Balka site alone contained around eleven individuals of various ages. A skeletal reconstruction made up of different individuals has a shoulder height of 2.4 and a length of 4.2 m. The Caucasian form formed part of the Taman fauna complex , which continues to include the southern elephant and the Taman bison . Other finds come from Salcia ( Moldova ) , among others . The rhinoceros species is also documented in Kazakhstan , with some teeth from the village of Lebjazhje on the Irtysh .

The species was described in 1914 by Alexei Alexejewitsch Borissiak . The description was based on several large teeth of an individual from Sinaya Balka on the Taman Peninsula on the Sea of ​​Azov, morphologically different from Elasmotherium sibiricum .

Elasmotherium peii

The species Elasmotherium peii was first known from some sites in the Nihewan Basin in the north of the Chinese provinces of Hebei and Shanxi and from Huáng Hé in the south of Shanxi, later remnants from Eastern Europe were added. It was a smaller representative, which is mainly occupied by teeth and some limb bones. It first appeared at the end of the Upper Pliocene in East Asia, where it appeared among other things in the Nihewan Basin with early representatives of the woolly rhinoceros ( Coelodonta nihowanensis ) within the Nihewan Faunenkomplex . However, it disappeared again in the early Old Pleistocene 1.6 million years ago. The species was described in 1958 by Chow Minchen with reference to an upper left row of teeth with the preserved teeth from the second milk premolar through the third permanent premolar to the third permanent molar. In the same article, Chow established another independent species, Elasmotherium inexpectatum , based on a single upper molar. But today it is placed with Elasmotherium peii . Some of the finds had already been collected by Teilhard de Chardin in northern China in the 1930s and 1940s .

At the beginning of the 21st century, some scientists revoked the species status of Elasmotherium peii , citing analyzes on the teeth and on the long bones, which were not previously documented in detail, and saw the form as identical to Elasmotherium caucasicum . More recent, very well-preserved finds, such as teeth and foot bones from Shanshenmiaozui and Heitugou in the Nihewan Basin, among others, made it possible to separate both representatives at the species level. Accordingly has Elasmotherium PEII less hochkronige teeth, a thicker enamel pattern and clearly formed roots.

Elasmotherium sibiricum

The best known and so far most frequently found species is Elasmotherium sibiricum . The species reached a length of 4 to 4.5 m (without tail), with a height at the withers of 2 m. The skull alone measured 95 cm. With a calculated body weight of 4 t, it was slightly smaller than the older Elasmotherium caucasicum . Other differences to this species are that Elasmotherium sibiricum had longer legs in relation to the body. The row of teeth was also much shorter, with the last premolar looking relatively delicate. The teeth had strongly folded enamel with clearly curved lines of folds and were clearly higher crowned. In relation to Elasmotherium caucasicum , however, the teeth were noticeably smaller.

The distribution area of Elasmotherium sibiricum reached from the Black Sea to eastern Kazakhstan. The southernmost limit of distribution was around 44 degrees latitude in present-day Uzbekistan , while the species advanced to the north as far as Siberia at the mouth of the Kama near the city of Irbit at 57 degrees latitude. A total of over 60 sites are known to date, half of which are in Kazakhstan. Usually only single bones and teeth were found - to this day there is no complete skeleton of this species, so that individual skeletal elements are still unknown. The most completely preserved skeleton to date comes from Selenokumsk ( Russia ) in the northern foothills of the Caucasus , where it was found in 1966 in a brickworks pit on the right bank of the Kuma . The skeleton, the remains of which were scattered over an area of ​​10 m², comprises a complete skull, parts of the spine, ribs and fore and hind limbs. In 1968 two more individuals of this rhinoceros appeared only 200 m away. As early as 1964 , a relatively complete skeleton was found near Stavropol , also in southern Russia, which was later exhibited in the local museum as a skeleton assembly. One of the most extensive find complexes of Elasmotherium sibiricum was found in 1938 on the right bank of the Great Karaman , a tributary of the Volga . It consists of several individuals, including a partially complete skeleton. In relative proximity to this is the site of Irgis 1 on the banks of the Great Irgis , also a tributary of the Volga. The remains of at least seven individuals of all ages were found here. The geological and paleontological data suggest that the animals died suddenly if the carcasses were quickly covered. Analyzes of the signs of wear on the teeth indicate that the animals switched from the general grass diet to a soft vegetable diet in their last phase of life. The scientists explain the unusual change in diet with a harsh Dsud combined with heavy snowfall. The resulting inaccessible grass food forced the animals to resort to leaves and branches, which led to malnutrition and ultimately to their death. There are also several skull finds: One skull comes from the left bank terrace of the Sakmara near Chelyabinsk (Russia), four more were found near Lutschka near the mouth of the Volga (also Russia) and two are from Kazakh sites near Atyrau and Nur-Sultan ( until 2019 Astana) .

Elasmotherium sibiricum first appeared in the Middle Pleistocene, it probably emerged from Elasmotherium caucasicum . Especially in Central Asia , it was part of the Koshkurgan fauna complex , which includes the southern elephant, the Schoetensack bison , the forest rhinoceros and the Mosbach horse . In Eastern Europe it belongs to the late Middle Pleistocene Chazar fauna complex of the Volga- Don area and occurs here together with the steppe mammoth and the forest rhinoceros. The most recent find in Eastern Europe comes from the Volga island of Khoroshevsky, which was flooded with the completion of the Saratov reservoir , and dates to the Mikulino warm period (synonym for the Eem warm period ; 128,000 to 115,000 years ago). The maxillary molar found there also has some peculiarities that are considered pathological changes due to the lack of similar comparative findings . However, Elasmotherium sibiricum survived longer in its easternmost distribution areas . Finds from the Smelovskaya II cave in southwestern Siberia came to light, they are associated with the woolly mammoth and the woolly rhinoceros and, according to radiometric data , are more than 50,000 years old. Other late Pleistocene bone remains are documented from Batpak and Voronovka, also in southwestern Siberia. The age of finds near the place Kozhamzhar in the Pavlodar region in Kazakhstan , including not only a skull of Elasmotherium sibiricum but also the remains of the steppe and woolly mammoth, was dated to 27,500 to 29,000 years and could thus make the date of extinction significantly compared to the previous ones move accepted data. However, there are methodological problems with the measurement, so that the values ​​must initially be considered uncertain. In comparison, 19 analyzed bone remains from western Asia and eastern Europe yielded radiocarbon dates between 50,000 and 36,520 years. According to this, Elasmotherium sibiricum would have disappeared well before the maximum of the last glacial period .

This lower jaw, described in 1808, forms the holotype specimen of Elasmotherium .

The first scientific description of the genus Elasmotherium with the species Elasmotherium sibiricum was provided by the German zoologist Gotthelf Fischer von Waldheim (1771-1853). It appeared in 1808 as a short essay, which was published again the following year in a larger frame with the same illustrations. The basis of the description was formed by a lower jaw from the Natural History Museum in Moscow , which belonged to the fossil collection of Princess Ekaterina Daschkowa . Daschkowa had donated the collection to Moscow University three years earlier with the express wish to make it accessible to the public. The lower jaw is considered a holotype , while a much more complete, 97 cm long skull, discovered as early as 1877, is considered a lectotype by Lutschka. An Elasmotherium tooth, which was also presented in the first description, was lost in 1812. In the course of the history of research there have been various names for the species Elasmotherium sibiricum :

  • Elasmotherium fischerii Desmarest , 1820
  • Elasmotherium keyserlingii Fischer von Waldheim , 1842
  • Stereocerus galli Duvernoy , 1855
  • Enigmatherium stavropolitanum Pavlova , 1916.

Elasmotherium fischerii is a name used by Anselme Gaëtan Desmarest in 1820, which is based on Fischer von Waldheim's description. It was used several times in the 19th century, but is invalid due to the later name. The form Elasmotherium keyserlingii refers to Alexander Graf Keyserling and is based on a tooth from the Caspian Sea, which Keyserling received from a Kyrgyz prince in 1841. Fischer von Waldheim then established the name in honor of Keyserling in the following year, but Johann Friedrich von Brandt reunited it with Elasmotherium sibiricum in 1864 . The description of Enigmatherium stavropolitanum is based on a single tooth from the northern Caucasus region, which, however, does not show any characteristic differences to Elasmotherium sibiricum , which is why this name is no longer used. The Stereocerus galli species is problematic , it goes back to a rear skull fragment from the natural history collection of the doctor Franz Joseph Gall (1758–1828), the origin of which was given as “Rhine Valley”. This led to the assumption that Elasmotherium sibiricum would have spread as far as Central or Western Europe. However, the actual origin of these early research finds is unclear, as is the case with similarly old Elasmotherium fossils from Hungary or Italy.

Tribal history

Origin and development

The genus Elasmotherium first appeared in the late Pliocene of Asia and goes back to Sinotherium of the Upper Miocene and Lower Pliocene. Both genera form a clade , which - together with its sister clade Parelasmotherium and Ningxiatherium and the further outlying Iranotherium - belongs to the tribe of the Elasmotheriini , which is mostly assigned to the subfamily Rhinocerotinae, but also to an independent subfamily, the Elasmotheriinae, by some scientists. All of the rhinos mentioned are characterized by a large body, a tendency towards high crowning of the teeth with strongly folded enamel and a lack of front teeth. These features were not yet developed in the earliest representatives of the Elasmotheriini in the Middle Miocene and only crystallized in the late Miocene and early Pliocene. These changes, in particular the increasing body size and tooth morphology, go hand in hand with a greater spread of open landscapes due to the increasing aridization of the climate and the associated change in diet to grasses. A clear parallel can be seen in the development of the closely related genera Stephanorhinus with the steppe and forest rhinoceros ( Stephanorhinus hemitoechus and Stephanorhinus kirchbergensis ) and Coelodonta with the woolly rhinoceros ( Coelodonta antiquitatis ) during the Pleistocene, but these belong to the group of rhinocerotini and thus not in the immediate family of Elasmotherium . Here, too, as a result of the desertification of the landscape, there was a change in diet and thus an increasing crowning of the molars, which reached its peak in Elasmotherium and Coelodonta ( but was even more developed in Elasmotherium ). It is possible that this strong specialization was also responsible for the comparatively early extinction of Elasmotherium , as the isotope analyzes available so far indicate a less flexible diet. Accordingly, the animals were probably only moderately capable of reacting to the greater climatic fluctuations of the last glacial period combined with a changing supply of food crops. The extinction process was exacerbated by the low population density and the long succession of generations that is common in rhinos.

The development of the horn of Elasmotherium

Elasmotherium represents the final link in the development of Elasmotheria and is characterized by a single horn rising up on the forehead, which also stands on a large dome-like elevation. For a long time it was unclear how this horn and its bony substructure had evolved . Most of the ancestral predecessors of Elasmotherium had a single horn on the nose and a flat or straight forehead line. Only in the case of Sinotherium was the formation of the horn not known due to the fact that the skull was only fragmented, but based on the anatomical features of the skull, the rhinoceros representative was one of Elasmotherium's closest relatives . Due to the discovery of a partial skull from the Linxia Basin in 2012, the skull morphology of all more modern Elasmotheria is now largely known, which means that the horn development within this rhinoceros group can be better understood. It began with comparatively small representatives with an equally small horn on the tip of the nose, as can be seen in Hispanotherium from the transition from the lower to the middle Miocene 18 million years ago. With the help of the fossils, the slow increase in body size of the animals, but also of the horns through the expansion of the roughened attachment points, can be understood. In the course of the evolution of the Elasmotheria in the history of the phylogenetic tree, the anterior horn was shifted slightly backwards, so that it was no longer located directly on the tip of the nose, but a little further back on the nasal bone: This is known above all from the late Miocene form Parelasmotherium , which was born before 10th centuries Lived up to around 5 million years ago. In Sinotherium the horn base was at the transition from the nasal to the frontal bone; an additional, much smaller horn base lay directly behind and was easily separated. Both showed a clearly dome-like structure, as is also known from Elasmotherium from the Plio and Pleistocene . With this, the base of the horn rose directly on the forehead. During the development of Elasmotherium , these two dome-like elevations must have merged. On the base of the horn of Elasmotherium a slight bone suture can be seen, the origin of which has not yet been explained, but could have its origin in the merging of the two separate elevations in Sinotherium .

Parallel to the backward displacement of the horn, the elasmotheria also caused a deformation of the nasal area. Elasmotherium has a completely ossified nasal septum , just like Sinotherium ; this is largely unknown in older Elasmotheria. Only Ningxiatherium from the late Miocene has an ossified septum in the front third. It is unknown whether these adhesions are related to the relocation of the horn. To stabilize the forehead and the rear nasal area through the load of the massive horn, Sinotherium had numerous transverse bony ribs on the palate area. Ossified nasal septa in rhinos otherwise only occur in the Coelodonta - Stephanorhinus line, which are particularly evident in the woolly rhinoceros and the steppe rhinoceros.

Research history

The first living reconstruction of Elasmotherium by Alexander Fjodorowitsch Brandt from 1878

When Fischer von Waldheim introduced the genus Elasmotherium in 1808 , only one lower jaw was available to him. He therefore did not yet associate the new form with an extinct rhinoceros, but compared it with other mammal species that no longer exist today, which in his opinion had no relatives among the recent representatives. These included the American mastodon (a primeval trunk animal), Megatherium and Megalonyx (two ground-living sloths) or Palaeotherium (a related horse). In the subsequent period the relationship of Elasmotherium remained unclear without additional finds. Richard Owen therefore led the genus in his important work Odontography , which was published from 1840 to 1845, only within the ungulates . However, as early as 1840 , Johann Jakob Kaup assumed , including the skull fragment from the collection of Franz Joseph Gall, that Elasmotherium is a representative of the rhinos. ( Georges Louis Duvernoy used the same skull fragment to set up Stereocerus galli , which he also assigned to the rhinos, but the shape was not generally recognized.) Johann Friedrich von Brandt followed this up almost a quarter of a century later with a similar argument, but only first the skull found near Lutschka in 1877 enabled Brandt to clearly relate Elasmotherium to the rhinos. In 1878 he submitted a comprehensive description of the skull, in which he not only emphasized the forehead dome as a special education, but also expressed speculations about the appearance and size of the animal. In the same year his son, the Russian scientist Alexander Fjodorowitsch Brandt , published a first pictorial reconstruction based on the previous finds of Elasmotherium , which mainly contained a massive frontal horn.

In the following years Elasmotherium was often interpreted in an external form similar to the woolly rhinoceros, but with only one front horn, due to the lack of comprehensive skeletal material. Konstantin K. Fljorow reconstructed the animals in the 1950s as steppe dwellers who churned up the ground with their snouts in search of food. The head was dominated by a horn that sat like a cap on the dome-shaped bulge of the skull. Fljorow also interpreted the forehead dome as an extended olfactory organ that was helpful in the search for food. Around the same time, VA Teryaev saw Elasmotherium as a hippopotamus -like animal that inhabited swamps. In its living reconstruction it no longer resembled a rhinoceros, the dome-like bulge on the forehead was hornless, and the forefeet consisted of four toes, comparable to tapirs , in order to move better in the swampy underground. Teryaev justified his interpretation with the frequent finding of fossil finds of Elasmotherium in former river landscapes. He also saw the bulge as too delicate to exercise a load-bearing function. More recently, Teryaev's views have mostly been revised from an anatomical point of view.

Elasmotherium and man

It is unclear whether the giant rhinoceros used to be prey for human hunter-gatherer groups, such as Homo erectus or the Neanderthal , but it seems to have rarely occurred at archaeological sites. The classic Sinaya Balka site on the Taman Peninsula yielded not only the type specimen of Elasmotherium caucasicum and other individuals but also stone artifacts from the Old Paleolithic . From some travertine sites in the south of Kazakhstan, such as Koshkurgan and Shoktas, there are also ancient Plaeolithic stone artifacts together with remains of Elasmotherium sibiricum . The finds are somewhat younger than those of Sinaya Balka and date to the early Middle Pleistocene .

Alleged representation of Elasmotherium in the Rouffignac cave

The interpretation of some characteristic rhinoceros portraits of the Upper Paleolithic cave art, such as from the Rouffignac cave as a representation of Elasmotherium, is rather problematic . Usually these assumptions relate to individual rhino drawings with only one horn. Provided that the art depictions of the Upper Palaeolithic are most likely not used to reproduce realistic objects, all rhinoceros depictions can be assigned to the woolly rhinoceros . In addition, there is currently no clear evidence that Elasmotherium was so widespread both spatially and temporally.

In some cases, Elasmotherium is also associated with the legends about the supposed unicorn . A pioneer for this was Alexander Fyodorowitsch Brandt, who reported in his popular article from 1878 about stories of some East Siberian peoples about the killing of huge black, one-horned bulls. The first written evidence of the mythical creature can be found in ancient Greece. Even before that, unicorn-like beings were depicted on seals in the cities of Mohenjo-Daro and Harappa of the Indus culture , which are a little over 4000 years old. It is unclear, however, whether they actually represent unicorns or only suggest this through their strict side representation. Elasmotherium as a prototype of the unicorn can only be justified if there had been a complete oral tradition from the time of its existence to more recent periods.

literature

  • Анна К. Швырева: Исқопаемые носороги зласмотерии. Stavropol, 1995, pp. 1-104
  • Anna K. Schvyreva: On the importance of the representatives of the genus Elasmotherium (Rhinocerotidae, Mammalia) in the biochronology of the Pleistocene of Eastern Europe. Quaternary International 379, 2015, pp. 128-134
  • Vladimir Zhegallo, Nikolay Kalandatze, Andrey Shapovalov, Zoya Bessudnova, Natalia Noskova and Ekaterina Tesakova: On the fossil rhinoceros Elasmotherium (Including the collections of the Russian Academy of sciences). Cranium 22 (1), 2005, pp. 17-40

Individual evidence

  1. ↑ a b c d e Deng Tao: A new elasmothere (Perissodactyla, Rhinocerotidae) from the late Miocene of the Linxia Basin in Gansu, China. Geobios 41, 2008, pp. 719-728
  2. ↑ a b c d e Deng Tao and Zheng Ming: Limb bones of Elasmotherium (Rhinocerotidae, Perissodactyla) from Nihewan (Hebei, China). Vertebrata Palasiatica 43, 2005, pp. 110-121
  3. ↑ a b c d e f g h i j k l m n o p q Vladimir Zhegallo, Nikolay Kalandatze, Andrey Shapovalov, Zoya Bessudnova, Natalia Noskova and Ekaterina Tesakova: On the fossil rhinoceros Elasmotherium (Including the collections of the Russian Academy of sciences). Cranium 22 (1), 2005, pp. 17-40
  4. ↑ a b E. I. Belyaeva: About the hyoideum, sternum and metacarpale V bones of Elasmotherium sibiricum Fischer (Rhinocerotidae). Journal of the Palaeontological Society of India 20, 1977, pp. 10-15
  5. ↑ a b c d Paul Mazza and A. Azzaroli: Ethological inferences on Pleistocene rhinoceroses of Europe; Inferenze sull'etologia di rinoccrontipleistocenici d'Europa. Rendiconti Lincei 4, (2) 1993, pp. 127-137
  6. ↑ a b Jan van der Made and René Grube: The rhinoceroses from Neumark-Nord and their nutrition. In: Harald Meller (Ed.): Elefantenreich - Eine Fossilwelt in Europa. Halle / Saale, 2010, pp. 382–394
  7. ↑ Esperanza Cerdeño: Cladistic analysis of the family Rhinocerotidae (Perissodactyla). American Museum Novitates 3143, 1995, pp. 1-25
  8. ↑ a b c d e f g Baldyrgan S. Kožamkulova: Elasmotherium sibiricum and its distribution area on the territory of the USSR. Quaternary Palaeontology 4, 1981, pp. 85-91
  9. ↑ a b c d Ekaterina Petrova: New data about the skull of the Elasmotherium sibiricum. In: Abstract volume: Quaternary stratigraphy and paleontology of the Southern Russia: connections between Europe, Africa and Asia. 2010 annual meeting INQUA-SEQS, Rostov-on-Don, Russia, June 21–26, 2010. Rostov-on-Don, 2010, pp. 121–122
  10. ^ Mikael Fortelius: Ecological aspects of dental functional morphology in the Plio-Pleistocene rhinoceroses of Europe. In: B. Kurtén (Ed.): Teeth: Form, function and evolution. New York, 1982, pp. 163-181
  11. ^ A b Donald R. Prothero, Claude Guérin and Earl Manning: The history of Rhinocerotoidea. In Donald R. Prothero and RM Schoch (eds.): The evolution of the Perissodactyls. New-York, 1989, pp. 321-340
  12. ↑ a b c V. S. Baigusheva, GI Timonina and VV Titov: Some biological features of Elasmotherium caucasicum (Mammalia, Rhinocerotidae) from the Late Early Pleistocene of Eastern Europe. In: Femke Holwerda, Anneke Madern, Dennis Voeten, Anneke van Heteren, Hanneke Meijer and Natasja den Ouden (eds.): XIV Annual Meeting of the European Association of Vertebrate Palaeontologists 6-10 July 2016, Haarlem, The Netherlands Program and Abstract Book . Haarlem, 2016, p. 43
  13. ↑ a b c d Pavel Kosintsev, Kieren J. Mitchell, Thibaut Devièse, Johannes van der Plicht, Margot Kuitems, Ekaterina Petrova, Alexei Tikhonov, Thomas Higham, Daniel Comeskey, Chris Turney, Alan Cooper, Thijs van Kolfschoten, Anthony J. Stuart and Adrian M. Lister: Evolution and extinction of the giant rhinoceros Elasmotherium sibiricum sheds light on late Quaternary megafaunal extinctions. Nature Ecology & Evolution 3, 2019, pp. 31-38, doi: 10.1038 / s41559-018-0722-0
  14. ↑ a b В. А. Теряев: Был ли Elasmotherium трехпалым? Ежегодник 8, 1930, pp. 77–82
  15. ↑ a b В. А. Теряев: Геологическое положение горболобого носорога (зласмотерия). Советская Геология 34, 1948, pp. 81–89
  16. ↑ a b c d e f g h N. G. Noskova: Elasmotherians - evolution, distribution and ecology. In: G. Cavarretta et al. (Eds.): The World of Elephants - International Congress. Consiglio Nazionale delle Ricerche (Rome) 2001, pp. 126–128
  17. ↑ Oscar Sanisidro, María Teresa Alberdi and Jorge Morales: The First Complete Skull of Hispanotherium matritense (Prado, 1864) (Perissodactyla, Rhinocerotidae) from the Middle Miocene of the Iberian Peninsula. Journal of Vertebrate Paleontology, 32 (2), 2012, pp. 446-455
  18. ^ A b Ashot Margaryan, Mikkel-Holger S. Sinding, Shanlin Liu, Filipe Garrett Vieira, Yvonne L. Chan, Senthilvel KSS Nathan, Yoshan Moodley, Michael W. Bruford and M. Thomas P. Gilbert: Recent mitochondrial lineage extinction in the critically endangered Javan rhinoceros. Zoological Journal of the Linnean Society 190 (1), 2020, pp. 372-383, doi: 10.1093 / zoolinnean / zlaa004
  19. ↑ Miklós Kretzoi: Gobitherium ng (Mamm., Rhinoc.). Földtany Közlony 73, 1943, pp. 268–271 ( [1] )
  20. ↑ a b Anna K. Schvyreva: On the importance of the Representatives of the genus Elasmotherium (Rhinocerotidae, Mammalia) in the biochronology of the Pleistocene of Eastern Europe. Quaternary International 379, 2015, pp. 128-134
  21. ↑ a b c В. В. Титов: Крупные млекопитающие позднего плиоцена Зеверо-Восточного Приазовья. Rostov-on-Don, 2008
  22. ↑ a b c Vera Baigusheva and Vadim Titov: Pleistocene large mammals associations of the Sea of ​​Azov and adjacents regions. In: Abstract volume: Quaternary stratigraphy and paleontology of the Southern Russia: connections between Europe, Africa and Asia. 2010 annual meeting INQUA-SEQS, Rostov-on-Don, Russia, June 21-26, 2010. Rostov-on-Don, 2010, pp. 24-27
  23. ↑ Vera S. Bajgusheva and Vadim V. Titov: Results of the Khapry Faunal Unit revision. In: L. Maul and RD Kahlke (Eds.): Late Neogene and Quaternary biodiversity and evolution: Regional developments and interregional correlations. Conference Volume. 18th International Senckenberg Conference. VI International Palaeontological Colloquium in Weimar. Conference in Weimar, 25. – 30. April, 2004. Terra Nostra, Writings of the Alfred Wegener Foundation (Berlin), 2004
  24. ↑ Vitaliy Logvynenko: The development of Late Pliocene to early Middle Pleistocene large mammal fauna of Ukraine. In: L. Maul and RD Kahlke (Eds.): Late Neogene and Quaternary biodiversity and evolution: Regional developments and interregional correlations. Conference Volume. 18th International Senckenberg Conference. VI International Palaeontological Colloquium in Weimar. Conference in Weimar, 25. – 30. April, 2004. Terra Nostra, Writings of the Alfred Wegener Foundation (Berlin), 2004
  25. ↑ a b c Esperanza Cerdeño: Diversity and evolutionary trends of the family Rhinocerotidae (Perissodactyla). Palaeo 141, 1998, pp. 13-34
  26. ↑ a b c d Piruza Tleuberdina, Gulzhan Nazymbetova: Distribution of Elasmotherium in Kazakhstan. In: Abstract volume: Quaternary stratigraphy and paleontology of the Southern Russia: connections between Europe, Africa and Asia. 2010 annual meeting INQUA-SEQS, Rostov-on-Don, Russia, June 21–26, 2010. Rostov-on-Don, 2010, pp. 171–173
  27. ↑ А. Борисякъ: О зубномъ аппарате Elasmotherium caucasicum n. Sp .. Исвестя Императорскоӣ Академиӣ Наук 6 (9), 1914, pp. 555-584
  28. ↑ a b Chow Minchen: New elasmotherine rhinoceroses from Shansi. Vertebrata Palasiatica 2 (2/3), 1958, pp. 132-142
  29. ↑ a b Tong Haowen, Wang Fagang, Zheng Min and Chen Xi: New Fossils of Stephanorhinus kirchbergensis and Elasmotherium peii from the Nihewan Basin. Vertebrata Palasiatica 33 (3), 2014, pp. 369-388
  30. ↑ a b c Анна К. Швырева: Исқопаемые носороги зласмотерии. Stavropol, 1995, pp. 1-104
  31. ↑ Florent Rivals, Natalya E. Prilepskaya, Ruslan I. Belyaev and Evgeny M. Pervushov: Dramatic change in the diet of a late Pleistocene Elasmotherium population during its last days of life: Implications for its catastrophic mortality in the Saratov region of Russia. Palaeogeography, Palaeoclimatology, Palaeoecology 556, 2020, p. 109898, doi: 10.1016 / j.palaeo.2020.109898
  32. ↑ a b Anatoly P. Derevianko, Valery T. Petrin, Zhaken. K. Taimagambetov and Marcel Otte: Early Palaeolithic assemblages in travertine, Southern Kazakhstan (a variant of an adaptation model). Anthropologie 36, 1998, pp. 137-164
  33. ↑ a b Diana Pushkina: The Pleistocene easternmost distribution in Eurasia of the species associated with the Eemian palaeoloxodon antiquus assemblage. Mammal Review 37 (3), 2007, pp. 224-245
  34. ↑ a b А. А. Хромов: Зуб эласмотерия с нетипичной из среднеплейстоценовых отложений Саратовского поволжья. In: В. М. Подбина, С. А. Родыгин, Н. И. Савина and Г. М. Татьянин: Эволюция жизни на Земле. Материалы II Международного симпозиума 12-15 ноября 2001 г. (Evolution of life on the earth. Proceedings of the II International symposium November 12-15, 2001) Tomsk, 2001, pp. 534-535
  35. ^ Pavel Kosintsev: Relict mammal species of the Middle Pleistocene in Late Pleistocene Fauna of Western Siberia south. In: Abstract volume: Quaternary stratigraphy and paleontology of the Southern Russia: connections between Europe, Africa and Asia. 2010 annual meeting INQUA-SEQS, Rostov-on-Don, Russia, June 21–26, 2010. Rostov-on-Don, 2010, pp. 78–79
  36. ^ PA Kosintsev: Elasmotherium (Elasmotherium sibiricum Fischer, 1808): new data on the period of existence and geographic range. In: AV Borodin, EA Markova and TV Strukova (eds.): The Quaternary of the Urals: global trends and Pan-European Quaternary records. International conference INQUA-SEQS 2014 Ekaterinburg, Russia, September 10–16, 2014. Ekaterinenburg, 2014, pp. 67–68
  37. ↑ Andrei Valerievich Shpansky, Valentina Nurmagambetovna Aliyassova and Svetlana Ilyina Anatolievna: The Quaternary Mammals from Kozhamzhar Locality (Pavlodar region, Kazakhstan). American Journal of Applied Sciences 13 (2), 2016, pp. 189–199 ( [2] )
  38. ^ Paula Jo Reimer and Svetlana Vladimirovna Svyatko: Comment on Shpansky et al. 2016, 'The Quaternary Mammals from Kozhamzhar Locality (Pavlodar Region, Kazakhstan). American Journal of Applied Sciences 13 (4), 2016, pp. 477-478
  39. ^ A b Gotthelf Fischer von Waldheim: Sur l'Elasmotherium et le Trogontherium deux animaux fossiles et insonnus de la Russie. Mémoires de la Société des naturalistes de Moscou 2, 1809, pp. 251–268 ( [3] )
  40. ^ A b c Gotthelf Fischer von Waldheim: Addendum to the history of the Elasmotherium. Bulletin de la Société impériale des naturalistes de Moscou 15, 1842, pp. 457–461 ( [4] )
  41. ↑ Anselme Gaëtan Desmarest: Mammalogie ou description des espèces des Mammifères. Paris, 1820, pp. 1–555 (p. 546) ( [5] )
  42. ^ A b c Johann Friedrich von Brandt: Observationes de elasmotherii reiquiis. Mémoires de l'Académie impériale des sciences de St.-Pétersbourg 7, 8 (4), 1864, pp. 1–34 ( [6] )
  43. ↑ Alexander Graf Keyserling: Observation of an Elasmotherium. Bulletin de la Société impériale des naturalistes de Moscou 15, 1842, pp. 454–457 ( [7] )
  44. ^ A b Johann Jakob Kaup: About the Elasmotherium. New yearbook for mineralogy, geognosy, geology and petrafacts customer, 1840, pp. 453–456 ( [8] )
  45. ^ Mikael Fortelius and Kurt Heissig: The phylogenetic relationships of the Elasmotheriini (Rhinocerotidae, Mamm.). Communications from the Bavarian State Collection for Paleontology and Historical Geology 29, 1989, pp. 227–233
  46. ↑ Kurt Heissig and Oldřich Fejfar: The fossil rhinos (Mammalia, Rhinocerotidae) from the Lower Miocene of Tuchorice in northwestern Bohemia. Sborník Národního Muzea v Praze. Acta Musei Nationalis Pragae (series B, Natural History) 63 (1), 2007, pp. 19-64
  47. ↑ a b Pierre-Oliver Antoine: Middle Miocene elasmotheriine Rhinocerotidae from China and Mongolia: taxonomic revision and phylogenetic relationships. The Norwegian Academy of Science and Letters - Zoologica Scripta 32, 2003, pp. 95-118
  48. ↑ a b Tao Deng, ShiQi Wang and SuKuan Hou: A bizarre tandem-horned elasmothere rhino from the Late Miocene of northwestern China and origin of the true elasmothere. Chinese Science Bulletin 2012, pp. 1-7
  49. ↑ Richard Owen: Odontography; or, A treatise on the comparative anatomy of the teeth; their physiological relations, mode of development, and microscopic structure, in the vertebrate animals. Hippolyte Bailliere, London, 1840–1845, pp. 1–655 (p. 587) ( [9] )
  50. ^ Georges Louis Duvernoy: Nouvelles études sur les rhinocéros fossiles. Archives du Muséum d'Histoire Naturelle 7, 1855, pp. 1–482 (pp. 125–130) ( [10] )
  51. ^ Johann Friedrich von Brandt: On the history of the gradually gained knowledge of the Elasmotherium. Mémoires de l'Académie impériale des sciences de St.-Pétersbourg 7, 26 (6), 1878, pp. 1–31 ( [11] )
  52. ^ WT Schaurte: Representation of an Elasmotherium in the rock painting of Rouffignac. Natur und Museum 94 (9), 1964, pp. 354–356

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