Woolly rhinoceros

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Woolly rhinoceros
Skeleton of a woolly rhinoceros

Skeleton of a woolly rhinoceros

Temporal occurrence
Middle to Upper Pleistocene
550,000 to 12,000 years
Locations
Systematics
Subclass : Higher mammals (Eutheria)
Superordinate : Laurasiatheria
Order : Unpaired ungulate (Perissodactyla)
Family : Rhinoceros (Rhinocerotidae)
Genre : Woolly rhinos ( Coelodonta )
Type : Woolly rhinoceros
Scientific name
Coelodonta antiquitatis
( Blumenbach , 1799)

The extinct woolly rhinoceros ( Coelodonta antiquitatis ), also called woolly rhinoceros or fur rhinoceros , was a species of rhinoceros that was widespread in the Ice Age cold steppes between Western Europe and East Asia during the Middle and Young Pleistocene . It belongs to the group of Dicerorhinina, Eurasian rhinos with two horns, of which only the Sumatran rhinoceros ( Dicerorhinus sumatrensis ) still exists today.

Along with the woolly mammoth, the woolly rhinoceros was one of the character animals of the Ice Age landscapes. With its clearly low-hanging head and high-crowned molars, the rhinoceros species was very well adapted to the hard grass of the steppes. Due to numerous fossil finds , but also mummified carcasses from the permafrost soil of Siberia, the further way of life is very well known. In addition, woolly rhinos were depicted by early groups of hunters and gatherers in cave paintings , as engravings on stone and bones, and as small statuettes .

The rhinoceros species died out at the end of the Pleistocene around 12,000 years ago during the Quaternary wave of extinction, along with numerous other larger animal species.

distribution and habitat

Live reconstruction of a woolly rhinoceros in the Natural History Museum Vienna

Dissemination in space and time

Distribution area of ​​the woolly rhinoceros, above: entire distribution area in Eurasia during the Vistula glacial period , including: distribution in Europe (center) and Northeast Asia (below) with a representation of important sites (black dots)

As a cold-adapted animal, the woolly rhinoceros mainly preferred a boreal climate and inhabited the steppes of northern Eurasia . The core area was thus largely in the northern part of Central Asia (probably in Transbaikalia ), in southern Siberia and in Mongolia . However, it is difficult to narrow down due to the lack of reliably dated finds. During the cold periods of the Pleistocene, the woolly rhinoceros often expanded its habitat from these core areas far to the south, south-west and south-east. One of the earliest records of the rhino species comes from northeast China from the Zhoukoudian site and is more than 500,000 years old; however, the species only appeared there for a short time and possibly in the course of an early expansion of the habitat. A first, much more extensive expansion phase occurred during the Elster Glaciation (around 400,000 to 320,000 years ago), during which the woolly rhinoceros first appeared in Central Europe . So it reached Central Germany, among others, and is proven in the Harz area. When this icing phase ended, however, it returned to its original realm.

A further expansion phase took place during the Saale Glaciation (300,000 to 126,000 years ago). At that time the woolly rhinoceros reached France and the British Isles and reached its southernmost borders in the northern part of the Iberian Peninsula and in Romania . It has also been found for the first time in the Don and Volga rivers . But even at the end of this cold phase, the populations left their newly acquired habitats.

With the end of the Eem warm period and the beginning of the Vistula glacial period (115,000 to 11,700 years ago), the woolly rhinoceros again expanded its refuge from Inner Asia far south and in this phase reached its largest distribution area, which was also the largest contiguous territory. that was ever colonized by an extinct or still living rhinoceros species. In western Eurasia it penetrated to the Iberian, Apennines and the Balkan Peninsula or the Peloponnese and reached its extreme southern limit at about 37 to 40 ° north latitude. Its occurrence in the Caucasus and Central Asia ( Kazakhstan , Kyrgyzstan ) is also documented. In eastern Eurasia, finds from northeast China and the Korean Peninsula are known; in China, the species even penetrated to the 33rd north latitude, while the most easterly found points are on Kamchatka . At the same time there was also a northern expansion, in the course of which the woolly rhinoceros advanced to Yakutia as far as Kolyma and Wrangel Island .

Thus, the distribution area covers almost the entire northern Eurasia during its maximum extent; the populated area is given as approximately 23.7 million square kilometers. The regions that were ice-covered at the time ( Fennos Scandinavia , Northeast Europe, Northwest Asia) are excluded . It is noteworthy, however, that the woolly rhinoceros did not appear - or only very sporadically - in the extreme northeast of Asia and thus did not reach the Bering Bridge , which would have been necessary to penetrate to America . It is not entirely clear why the woolly rhinoceros did not succeed in this, even though other animal species such as the woolly mammoth , the bison and ultimately humans took this hurdle at the same time. Some researchers speculate that the species nevertheless crossed the Bering Bridge in small numbers and that there is only a lack of evidence in Alaska . In their opinion, it certainly did not cross the American ice sheets, which blocked the further way south. It has long been assumed that the woolly rhinoceros only penetrated north-east Asian regions in climatically favorable phases, which were obviously characterized by greater moisture penetration of the cold-age landscapes of Beringia . The formation of stagnant water on the permafrost soil then probably counteracted further eastward expansion, creating a migration barrier for the animal, which had adapted to the dry climate. More recent studies on the ecological conditions of the Bering Bridge during the last glacial period support this view, whereby, in addition to a more humid climate, a poorer food supply was possibly the reason why the woolly rhinoceros avoided crossing the Bering Bridge.

habitat

The woolly rhinoceros lived mainly in lowland areas , but also occurred on plateaus and plateaus. It seems to have avoided higher mountain regions due to the inaccessibility. Its main habitats were the wide lowlands and river valleys. The species only visited higher altitudes in favorable climatic phases. Furthermore, it was tied to largely open landscapes with a dry to arid climate .

The mammoth steppe of Eurasia was an important and classic habitat for the woolly rhinoceros . Characteristic for this steppe form, which no longer exists in this form today, were extensive, almost treeless grass and bush landscapes, which, due to the high levels of solar radiation, had vegetation rich in species and nutrients . This mammoth steppe was inhabited by the Mammuthus Coelodonta Faunenkomplex , which reached its greatest extent in the Young Pleistocene. In addition to the woolly rhinoceros, the woolly mammoth, the giant deer , the reindeer , the saiga antelope and the bison were other important accompanying elements of this fauna.

Due to its large distribution area, the woolly rhinoceros often coexisted with other rhino species. In the outgoing warm periods, when the woolly rhinoceros slowly advanced into the European areas, it partially shared its habitat with the steppe rhinoceros ( Stephanorhinus hemitoechus ), in Eastern Europe and Western Asia it occurred together with Elasmotherium . In East Asian Middle Pleistocene North Tsingling-Faunenkomplex early representative of the woolly rhinoceros were with the rhinoceros ( Stephanorhinus kirchbergensis socialized).

Climatic adaptation

Live reconstruction (illustration)

The woolly rhinoceros is generally considered to be the classic animal of the cold-age steppes of Eurasia, alongside the woolly mammoth. The ideas about this animal species are mainly shaped by the finds in the Siberian permafrost and the storage in mainly cold-age sediments of the last cold period. Their presence is thus also used as a marker for the climatic reconstruction of a site.

The origin of the woolly rhinoceros lies in the arid steppes of eastern Central and western East Asia. Adaptations to this open landscape form show the wide, short limbs, but also the characteristic diet with the resulting high-crowned teeth and the low head posture. These traits were already developed to varying degrees in his direct ancestors. However, some paleontologists argue that the woolly rhinoceros was not fully adapted to the glacial conditions. An indication of this is the not so high northern distribution compared to the woolly mammoth, as well as the fact that it did not make it to America. Rather, in the opinion of those palaeontologists, this animal species was tied to dry climatic conditions, as its frequent occurrence in Transbaikalia shows. A very continental climate prevailed there during the cold periods , which resulted in extensive steppes and partly semi-deserts .

Dry climatic conditions also meant that representatives of the woolly rhinoceros occasionally appeared in warm climatic sections in the early Middle Pleistocene in East Asia, for example at the important early human site of Zhoukoudian near Beijing (sometimes referred to as the subspecies Coelodonta antiquitatis yenshanensis ). These East Asian representatives of the woolly rhinoceros shared their habitat with Palaeoloxodon namadicus , an Asian old elephant species , the forest rhinoceros and sometimes also with the orangutan ( Pongo ). In Europe, too, the woolly rhinoceros can occasionally be detected in warm-period deposits of the Middle and Young Pleistocene. It was documented in the more than 200,000 year old deposits of La Fage (France), where it lived in a forest steppe. Further examples are finds of the woolly rhinoceros from the upper travertines of Ehringsdorf (Thuringia) and from the Geiseltal . At the latter site, which is assigned to the Eem warm period, a strongly continental climate can be reconstructed on the basis of macroscopic plant remains, which finds its current ecological equivalents in the Alföld or in the southern Russian steppe regions. In Crayford, England , the woolly rhinoceros was identified for the late Eem warm period. The species obviously not only tolerated boreal climates, but also managed to cope with warmer climates, but their occurrence in the Central and Young Pleistocene warm periods was probably linked to cooler or very dry phases.

During the Glaciation of the Vistula, the woolly rhinoceros not only reached the far north of Siberia due to its huge range, it also colonized landscape areas that reached far to the south. The woolly rhinoceros was widespread over large parts of southern Europe in the second half of the Glacial Vistula, so that its area partially overlapped with the retreats of some warm-period fauna elements. The woolly rhinoceros, for example, has been passed down in sandy deposits near Ingarano (southern Italy; layer b ) together with the European forest elephant and the hippopotamus .

Finds

Locations in general

Mounted woolen rhinoceros skeleton, found during construction work in the Rhine-Herne Canal and exhibited in the Ruhr Museum in Essen

Finds of the woolly rhinoceros are among the most common fossil finds of Pleistocene animal species in Eurasia . The mostly isolated bones and teeth come mainly from gravel, sand and clay pits, which indicate former rivers or lake shores. In addition, some finds are also known from caves . In addition, the remains of woolly rhinos are repeatedly found submarine from shelf seas such as the North and Baltic Seas , which were dry at times during the cold periods. In Poland alone , more than 50 sites are known, while the species has been registered in at least 20 sites in Spain. It is also known from Italy in a little more than half a dozen places. In eastern Eurasia, the woolly rhinoceros is the most common of all known fossil rhinoceros species in China with more than 70 sites, which include more than 60 taxa from the Eocene to the Pleistocene.

The woolly rhinoceros is also represented in Germany with a high density of finds, which includes more than 30 known sites in Westphalia alone . However, articulated skeletal remains or even complete skeletons are extremely rare. One of the earliest skeletons found comes from Pohlitz ( Thuringia ) and is now exhibited in the Museum of Natural History in Gera (also Thuringia). Another was found in a gravel and sand pit near Petershagen ( North Rhine-Westphalia ) and comprises 66 skeletal elements including a skull; it is now in the Bielefeld Natural History Museum . A third, formerly articulated skeleton from the Bavarian State Collection for Paleontology and Geology in Munich was destroyed during the Second World War. Numerous other mounted skeletons, for example in the Geological-Paleontological Museum of the Westphalian Wilhelms University in Münster , in the Museum of Prehistory and Local History, Eiszeithalle Quadrat Bottrop , or in the Ruhr Museum in Essen , on the other hand, mostly represent reconstructions from several skeletal elements in the respective region of found animals.

Finds of mummified carcasses

Sites of mummified carcasses of the woolly rhinoceros, star: carcasses in permafrost , star in a circle: carcasses in oil shale
A mummified carcass of a woolly rhinoceros was found in Starunia (now Ukraine ), 1929

As with the woolly mammoth, there are also preserved carcasses of the woolly rhinoceros in today's permafrost area, which provide information about the soft tissue anatomy. With around half a dozen scientifically registered finds, all of which come from Yakutia and date to the last glacial period, these are much rarer than those of the mammoth. One of the first known frozen carcasses was discovered in 1771 on the Viljui River . In 1858 an almost complete skeleton with partially preserved skin remains came to light in sandy deposits on the Wiljui. Only a few years later, in 1877, a Siberian trader discovered a rhinoceros mummy on the Chalbui River, a tributary of the Bytantai , but only a few parts of which he was able to recover, including the head with well-preserved skin; the rest of the skeleton was washed away in the following spring flood.

One of the most significant and informative finds is that of Churaptscha between the rivers Lena and Amga from 1972. While only individual skull and rib fragments were initially found during civil engineering work, further investigations in the same year at a depth of around 2.5 to 3.6 m in the frozen ground revealed an almost complete skeleton with preserved soft tissues on the right hind limbs. A short time later, in 1976, during an excursion on the left bank of the Aldan , students found a skull, a horn, individual vertebrae and other rhinoceros bones. Follow-up examinations revealed a partially preserved skeleton, including the skull with both horns and teeth, the lower jaw and vertebrae.

In 2007 an ice mummy was found on the lower reaches of the Kolyma. This includes a skeleton from which the skull and the left body part were preserved, with remains of skin with hair on the limbs. The finds from the river bed of the Great Chukotschja from 2008, which include an almost complete skeleton with a skull and both horns, are even more recent . In the same year, partially mummified rhino remains were discovered by a local resident near the Amga and professionally recovered in a two-year excavation in 2009 and 2010. This revealed pelvic bones, fore and hind limbs along with finger and toe bones, caudal vertebrae and ribs.

In September 2014, the ice mummy of a young animal was found on an unnamed tributary of the Semjuljach River in Abyjski ulus, Yakut . The parts that had already thawed out of the permafrost were no longer preserved due to animal damage. The head with fur and horn as well as parts of the soft tissue could be recovered from the permafrost. The two finders gave their find to the Academy of Sciences of Yakutia. According to their first assessment, the remains are well preserved. The age of the young animal is estimated based on the head-trunk length of about 172 cm at 12 to 18 months. Due to the state of preservation, scientists hope to obtain useful DNA.

Outside Yakutia and today's permafrost area, there are two complete female woolly rhinos and the remains of two other individuals from Starunia (now Ukraine ), which were found in a bitumen or oil shale pit and are also attributable to the last phase of icing. The first carcass with a full body and skull and two intact horns came to light in 1907. Another mummy was found in 1929, which was also complete - but this time without horns - and was discovered together with the other two, only partially preserved rhinoceros remains. Both carcasses showed very good soft tissue preservation, but without traditional fur cover. The second complete carcass find has been exhibited as a dermoplastic in the Natural History Museum in Krakow since the 1930s.

features

Skeleton of a woolly rhinoceros without a horn

Habitus

Information about the physique, diet and lifestyle of the woolly rhinoceros is much more extensive than for any other Pleistocene rhinoceros species. This is due to the fact that, in addition to isolated bones and teeth, occasionally complete skeletons have also come down to us; In addition, there are ice mummies preserved in the permafrost of Siberia, as well as carcasses preserved in brine and oil shale, and cave paintings and drawings carved into bones and stones. The woolly rhinoceros had a head-trunk length of 340 to 360 cm - in addition there was a 50 cm long tail - reached a height at the withers of 150 to 170 cm and had a body weight between 1.5 and 2.9 t. Furthermore, it was characterized by a robust physique and strong, broad limbs with clearly defined muscle attachment points, which is seen as an adaptation to its way of life in open landscapes. In addition, the woolly rhinoceros had a high and strongly pronounced neck hump, similar to the white rhinoceros ( Ceratotherium simum ), which gave it a characteristic body shape. As the frozen carcasses also show, it had a thick fur.

Skull and dentition features

Skull with horns

The skull measured between 70 and 90 cm and was very elongated. Characteristic was the pronounced backward pulling out back of the head , whereby the woolly rhinoceros carried its head clearly downwards. In this characteristic it was similar to today's white rhinoceros and its fossil relative, the steppe rhinoceros , and Elasmotherium had a similar head posture. Strong muscles attached to the greatly elongated occiput, which contributed to the shaping of the already mentioned neck hump and were necessary to hold the massive skull.

Like all Dicerorhinina, the woolly rhinoceros had two horns, which were located on the nose and the middle of the skull. The attachment points on the bone are roughened in the manner of pearl or cauliflower, the extensive expression of these surface roughening shows that the front horn was considerably larger than the rear. The front horn must have been particularly large compared to other rhinoceros, as the extent of these surfaces reached the highest values ​​within the Dicerorhinina group and also exceeded that of the steppe rhinoceros. A special feature is the completely overgrown nasal septum in adult individuals , which may have served to stabilize the large horn. This also occurs in older forms of Coelodonta , and to a less pronounced extent in the sister group Stephanorhinus ; Such a bony formation does not occur in the recent rhinoceros species. Unique to the woolly rhinoceros is the fusion of the nasal bone , which is rounded forwards, with the intermaxillary bone , which is not the case with older Coelodonta species - as is the case with today's rhino representatives.

The lower jaw was very massive and up to 60 cm long, while the lower jaw bone was up to 10 cm high. The dentition resembled that of its fossil relatives from the Stephanorhinus line, that is, the incisors were missing - these are usually only found in the deciduous dentition and are rudimentary there - while the premolar and molar teeth were each present three times per arch. Thus, the woolly rhinoceros following dental formula had: . The molars were very high crowned ; in this they surpassed those of the steppe rhinoceros and had a lot of root cement . It is particularly noteworthy that the second premolar was particularly graceful.

horns

Fossil horn of the woolly rhinoceros from Yakutia , for the first time in 1849 by Johann Friedrich von Brandt published

The horns have also been handed down in a frozen state, although the larger front horn (nasal horn) is more often preserved. This was clearly curved in shape and reached lengths of up to 90 cm, with the front, convex (outwardly curved) edge over the arch being up to 123 cm long. The weight is around 11 kg. The rear horn (frontal horn) has also been preserved from time to time. This often had a triangular to oval cross-section, was sometimes slightly curved and reached lengths of up to 40 cm. The weight of this horn is around 4.5 kg.

It is noteworthy that both horns are not round in cross-section as in today's rhinoceros species and as the overgrown attachment points suggested, but rather appear laterally compressed or sometimes flat as a board. It has not yet been clarified whether this shape corresponds to the original one or whether it resulted from subsequent shrinkage processes. The leading edge of the horn extends to the tip of the nose. Interestingly enough, it often shows strong abrasion marks. These were probably caused by the animal pushing away the snow cover with its front horn in order to get to the plants below, or simply wearing the horn off while eating and the associated ground contact.

Both horns are built up by horizontal bands, the width of which often varies. So far, a maximum of 35 such bands have been counted on a single horn. Early assumptions linked these ligaments to periods of horn growth that were interrupted when food was scarce. Such horn bands have also been found in recent rhinos, but far less often. Recent studies show that the bands - similar to the annual rings of trees or the concentric rings of elephant tusks  - are annual growth rates, with each band consisting of two parts: a darker section, which represents the summer months, and a lighter area representing the winter months.

Originally, it was thought that the horns, which consist of thick keratin threads (so-called filaments) running lengthways to the horn axis, were not used for rival fights due to their great length and flattened cross-section, as lateral shear forces would have made it easy to kink. On the basis of recent studies on the horn structure of the woolly rhinoceros, however, it has been determined that the inside of the horn has a heterogeneous structure, with the core of the horn being made significantly harder than the outer areas due to longer and more densely grown filaments.

Features of the trunk skeleton

Forelegs of the woolly rhinoceros with the characteristic three-pronged hands

The body skeleton is well known due to the numerous finds and especially the partially complete carcasses. The spine consisted of 7 cervical, 18 thoracic, 4 lumbar and 4 sacral vertebrae. The cervical vertebrae in particular were short and very high. The foremost cervical vertebrae were massive, the spinous process could be up to 36 cm long here. The humerus was massive and wide and at 43 cm in length only slightly smaller than that of today's white rhinoceros. The joint head rose only a little from the shaft. The ulna reached an even greater length at 46 cm. It exceeded the radius by one and a half times. The largest long bone, however, was the thigh bone with 52 cm, the shaft of which was extremely straight and had an additional trochanter in the middle for strong leg muscles. The shin , which was 36 cm long, was partially fused with the fibula . However, since both bones are quite often found separately, it is known that the adhesion must have taken place quite late in the individual development of the animals. The limbs each ended in three toes, as is typical of today's rhinos. The central ray (Metapodium III) was particularly massive, 16 cm long on the hand ( metacarpal bone ) and slightly smaller on the foot ( metatarsal bone ). The shafts had a straight shape, while the rays lying on the inside and outside (each Metapodium II and IV) were shorter and slightly curved.

Soft tissue morphology

Above all, the well-preserved finds from Starunia yielded numerous insights into the design of soft tissue ; further clues were added by the permafrost mummies. The skin in general was very wrinkled, but not segmented as in some of today's species, but interspersed with the hair follicles of the fur. It reached a thickness of 5 to 15 mm, it was particularly thin on the skull and extremely thick in the trunk area. The head was characteristic, especially the muzzle, which had a wide mouth with a mouth opening of up to 35 cm, similar to that of the recent white rhinoceros, as well as string-like lips , which replaced the incisors and were suitable for plucking grass. The mouth differed significantly from that of the other recent rhinos with their pointed upper lip. This could be proven both in the findings of Starunia and that of 1877 on the Chalbui River. As with the white rhinoceros, there were numerous wrinkles around the eyes. The ears were very far back on the head, were long and narrow - up to 24 cm long and 8 cm wide - and very pointed at the end. The prominent neck hump attached to the back of the head was not only created by the strong muscles that contributed to the stability of the skull, but also served as a fat store, which could also be examined particularly clearly in the Starunia finds. Comparable to today's African rhinos, the skin on the body shows hardly any wrinkles. The short tail, on the other hand, was very broad at its base, which is a parallel to the woolly mammoth as an adaptation to colder climates. The fur on the Yakut ice mummies can be observed particularly well. This consisted of long, stubbly outer hairs that were particularly well developed on the neck and shoulders. These covered a thick, insulating undercoat . The fur on the limbs, in contrast, was much shorter. Among the fur remnants found, rust-brown hair dominated. The calf from the Semjuljach River, rescued in 2014, had light-ash-gray to yellowish-blonde hair, which leads to the assumption that young animals may have a lighter coat than old animals. In addition, the fur on calves was significantly shorter.

Genetic traits

In the genome of the woolly rhinoceros there are various indications of adaptations to cold climatic conditions. This includes individual TRP channels such as TRPA1, which activates receptors in the skin that react to ambient cold. In many species, the proteins expelled in the process are normally suppressed by KCNK17, a variant of the KCNK channels , which leads to sensitivity to cold in these species. In the woolly rhinoceros, however, KCNK17 has become functionless due to a mutation , so its inactivity probably contributed to the animals' adaptation to the cold. Similar mechanisms are also known from the woolly mammoth .

Paleobiology

Diet

Numerous anatomical features suggest that the woolly rhinoceros feed mainly grazing. These features include the sloping position of the head, the high crown of the teeth with their high cement content and the delicate formation of the second premolar. Furthermore, an often horizontal abrasion of the teeth was observed, which is caused by the silica contained in grasses . In addition to these skeletal anatomical findings, the preservation of the snout area in some mummies provides further information.

Furthermore, plant residues from the teeth of individual woolly rhinos and fossil stomach contents from the Churaptscha rhinoceros from 1972 could be obtained and analyzed, which confirm the assumption of a largely grass-eating diet and also show that this animal species did not generally disdain bushes and branches. So among grasses and daisy family such as Artemisia to their food spectrum, as well as conifers , willows and alders . Recent isotope studies using fossil horns continued to show a possible seasonal diet. The different growth zones on the horns suggested different food offers. In summer, the woolly rhinoceros feed mainly grazing, while in winter it partly ate bushes and branches.

Way of life

The woolly rhinoceros reached an age of 40 to 45 years and does not differ in this from the more recent rhinos. The individual development was just as similar to that of today's relatives, which was mainly demonstrated by studies on changing teeth based on comparisons with the white and black rhinoceros . Accordingly, ten age levels were proposed for the structure of the ontogeny. Sexual maturity probably occurred at the age of five. Adult woolly rhinos certainly had no natural enemies or were rarely attacked by large predators such as the cave lion . Calves and young animals, however, were exposed to such a danger as is still the case with modern rhinos. Some juvenile skeletons have marks in the throat and neck area that can be traced back to large cat bites . In addition, there were other dangers that the woolly rhinoceros faced: poorly accessible terrain, steep or slippery slopes and the like could lead to serious falls or even death of individual individuals.

It is not known whether the woolly rhinoceros was a loner, most of today's rhinos are. However, the only rhinoceros that specialize in grass food today, the white rhinoceros, lives in small, matriarchal groups. Bulls are predominantly loners and live territorial, defending their territory against food competitors, both in intra- (concerning their own species) and interspecific (concerning other animal species) competition. Some woolen rhinoceros skulls show injuries indicative of fights with other rhinos. For example, perforations occur in the skull, often in the parietal bone , but also in the orbit or the maxilla . Parts of these injuries later led to osteomyelitis or arthrosis . Broken and overgrown lower jaws have also been handed down, as well as broken and partly healed ribs, which also suggest such fights. As a rule, injuries from ritual or territorial fighting were not fatal, and most woolly rhinos apparently regenerated. The relative frequency of such combat injuries compared to the recent rhinos is attributed to the rapidly changing climatic conditions during the last glacial period, whereby the woolly rhinoceros was exposed to an increased stress situation in competition with other large and medium-sized herbivores of the mammoth steppe . Apart from such injuries, osteomas have also been observed in the woolly rhinoceros , as well as tooth anomalies in the form of excess premolars and molars or the occurrence of genetically caused congenital enamel hypoplasia .

Woolly rhinoceros and human

The woolly rhinoceros as a supplier of food and raw materials

Direct references to the hunting of the woolly rhinoceros by early humans, comparable with the wild horse with the Schöninger spears and with the European forest elephant with the lance of Lehringen , are not yet known. The number of sites where humans and woolly rhinoceros appear together is relatively high, but a matching deposit is not always guaranteed. Caves, for example, in which the woolly rhinoceros is often found, such as the Bockstein cave ( Baden-Württemberg ) or the Kůlna cave ( Czech Republic ), but which do not reflect the natural habitat, used not only humans but also other, often large predators , such as hyenas , who also come into question as those who bring such residues. In addition, at many sites there are mostly only single teeth or bone fragments from the woolly rhinoceros, which can rarely be assigned to more than one individual. Traces of manipulation on the bones by humans are a sure sign of the use of the woolly rhinoceros by early human hunter-gatherer groups, whether the raw materials were captured by active hunting or by collecting at the animals' dying places.

Early common occurrences of human remains and woolly rhinoceros remains in Europe are Markkleeberg ( Saxony ), or Romain-la-Roche (France) from the Saale Glaciation . Such early discovery sites are relatively rare and often represent natural and thus accidental associations. Only with the Vistula glacial period did the number of common sites increase rapidly. But at only a few of these sites from the early phase of the last glaciation cycle, which are generally assigned to the late Middle Paleolithic and whose bearer was the Neanderthal man , clearly artificially broken rhinoceros bones with striking traces of cut appear, including in the Gudenus Cave ( Austria ) or at the Königsaue open-air station ( Saxony-Anhalt ). This breaking of bones is in part related to the acquisition of food in order to get to the high-energy bone marrow . On the other hand, early humans also smashed animal bones and reworked them in order to later use them as tools or weapons, which goes hand in hand with the increasing use of organic raw materials such as bones, antlers or ivory for the manufacture of devices. For example, in Zwoleń ( Poland ), the battered pelvis of a woolly rhinoceros was redesigned into a device.

In the subsequent Upper Paleolithic , connected with the first appearance of anatomically modern humans ( Homo sapiens ) in Europe, only a few sites are known for the use of the woolly rhinoceros as a source of raw materials, although this period is characterized by numerous innovations in the field of tool manufacture or techniques of hunting weapons, and organic raw material was used to an increasing extent and much more frequently than before. Rhino remains are relatively common in the early part of the early Paleolithic, the Aurignacian . In the Vogelherd cave (Baden-Württemberg), for example, at least 124 remains of bones from at least twelve woolly rhinoceros individuals have survived, although it is also discussed here whether these bone finds go back to active hunting or whether they represent remnants of the utilization of raw materials.

After the retreat of the inland glaciers , which reached their maximum extent 20,000 to 16,000 years ago, and the increasing repopulation of northern Alpine Europe by carriers of the Magdalenian in the late Upper Paleolithic, the woolly rhinoceros plays only a minor role in subsistence . This is due to the now relative rarity and the rather rapid later disappearance of this species in Europe. Remnants of the woolly rhinoceros are therefore only found in a few cases, such as in Gönnersdorf ( Rhineland-Palatinate ).

The woolly rhinoceros in Upper Paleolithic art

Part of the “Panneau of Horses” from the Chauvet Cave (replica). Approx. 31,000 BP, possibly Aurignacia. In addition to the horses, three rhinos, two apparently facing each other - but probably without any reference to each other - and three aurochs are shown.

Despite the relatively rare evidence of the use of the woolly rhinoceros by hunter-gatherer groups, this imposing animal must have made an impression on people of the time, especially from the Upper Paleolithic, as the representations of this animal in the emerging cave and mobile art show. The reproduction of the woolly rhinoceros takes place in - often left - side representation and not only traces the characteristic back line with the high neck hump, as it is also proven by the mummified carcass, but also reflects the low position of the head. In addition, there are also the representations of the ears and especially the long horns; in some cases the fur is also indicated. The partly very realistic drawings and engravings give a deep insight into the appearance of this Pleistocene animal species in addition to the biological investigations.

The earliest images include those from the Chauvet cave (France), which, with an age of more than 31,000 years, probably belong to the Aurignacia. There are 47 known representations on several panels, around a quarter of all animal drawings in the cave. They are alternately drawn in red or black, but also shown as an engraving. Particularly noteworthy are the drawings of two rhinos facing each other - a very rare scene in Upper Palaeolithic art - in the panneau of horses or those of four animals following each other in the panneau of positive hands . Repeating features of the representations are ears indicated by two curved lines or a black band dividing the body. The frequent depiction of long horns should be emphasized.

Other noteworthy representations are those from the Rouffignac cave , which apparently show the animals grazing, one by Font-de-Gaume with a noticeably higher head posture, which suggests a more attentive position, or one made of black pigments, very realistic in the shaft of the dead man in the cave of Lascaux (all France). Woolly rhinos are also depicted in other caves in the area of Franco-Cantabrian cave art , such as La Colombière, Les Rebières and Les Combarelles (all France) and El Castillo (Spain). Despite the numerous well-known cave drawings of the woolly rhinoceros, with only around 1% of all animal drawings in caves, it is relatively rare. Outside the caves of Frankokantabria, the drawings of woolly rhinos from the Kapova cave (Urals, Russia) made with red pigments are known, which can be assigned to the late Upper Paleolithic.

In addition to cave paintings, the woolly rhinoceros was also depicted several times in the mobile art of the - often late - Upper Palaeolithic. Particularly revealing representations come from the Gönnersdorf outdoor station (Rhineland-Palatinate) and are engraved on slate . At least ten images of this animal species are known from here, but in terms of numbers they lag far behind those of the horse and mammoth. Other rhinoceros depictions come from the Teufelsbrücke near Saalfeld and from the Kniegrotte near Döbritz (both Thuringia); the former was also carved into a piece of slate, the latter is on a piece of reindeer antler. A very rare woolen rhinoceros statuette made of clay was found in Dolní Věstonice in Moravia .

Tribal history

Schematic representation of the change in head posture of the woolly rhinoceros during its evolution, top: Coelodonta nihowanensis , middle: Coelodonta tologoijensis , below Coelodonta antiquitatis

The genus Coelodonta developed in East Asia . The oldest known representative, Coelodonta thibetana , lived in the middle Pliocene 3.7 million years ago in the highlands of Tibet . Coelodonta nihowanensis, the next younger member, comes from Mongolia and Northern China and is around 2.16 to 2.55 million years old; important sites here are Longdan and Shitougu (both province of Gansu , China). Compared to the later woolly rhinoceros, both species were still relatively small rhinos with a slim physique. Their occurrence goes hand in hand with an increasingly dry climate in eastern Asia, which at least in the case of Coelodonta nihowanensis is indicated by the simultaneous formation of aeolian loess deposits in the region. The most recent finds of Coelodonta nihowanensis to date are around 1 million years old. From this, Coelodonta tologoijensis developed , which is first detected in deposits around 750,000 years old in Tologoj on the Selenga River ( Buryatia , Russia ). Compared to the later woolly rhinoceros, this one is also much more delicate and shows slimmer and longer limbs. In addition, the teeth are less high-crowned and the occiput is not as elongated. Early Middle Pleistocene finds from the Kuznetsk Basin indicate a first westward movement of the genus Coelodonta , a first northern expansion up to the Lena occurred a little more than 600,000 years ago. The first evidence of Coelodonta antiquitatis in eastern Asia has been handed down more than 500,000 years ago .

There is a considerable gap in time and space between the first appearance of the early Coelodonta species in East and Central Asia and the earliest appearance of the woolly rhinoceros in western Eurasia in the Middle Pleistocene around 400,000 years ago. The earliest European finds are from the Elster Cold Age and come from Brașov (Romania) and Central Germany, mainly from the region around the Harz Mountains. The finds are mainly assigned to the classic woolly rhinoceros Coelodonta antiquitatis , but the material from this period is sparse and limited to individual tooth remains and bone fragments; only an almost complete skull is available from Bad Frankenhausen ( Thuringia ), which still has a relatively short occiput. Only from the subsequent Saale Glaciation have significantly more fossils come down to us. The finds from La Fage and Romain-la-Roche (both France) should be emphasized here. The latter site contained around 225 rhinoceros remains from nine individuals that belong to the end of the Saale Cold Age.

Differences between the older Late Middle Pleistocene and the later Young Pleistocene representatives of the European woolly rhinoceros had been identified early on. These mainly concern the degree of robustness of both the body and the limbs and the length of the occiput, all of which increased over time. This gave Claude Guérin an opportunity in 1980 to subdivide it into two subspecies, with Coelodonta antiquitatis praecursor representing the more delicate Middle Pleistocene and Coelodonta antiquitatis antiquitatis the more robust form. More recent research would like to assign the older subspecies to a later form of Coelodonta tologoijensis and only view the Young Pleistocene finds from the Vistula glacial period as the classic woolly rhinoceros Coelodonta antiquitatis . But this interpretation is controversial. A similarly motivated division into two subspecies as in Western Europe was carried out in Russia at the end of the 1960s on the basis of Yakut finds and the older Coelodonta antiquitatis jacuticus was separated from the younger Coelodonta antiquitatis humilis ; However, this classification is rarely used due to problems in taxonomic naming.

In terms of tribal history , the wool rhinoceros can be derived from the earliest Coelodonta finds in East Asia. It developed from a relatively small and slender rhino species to a large, robust one. Associated with this are skeletal morphological changes to the skull, such as the gradual lengthening of the occiput and the associated lowering of the head, combined with a thinning of the skull. By relocating the eyes to the back of the skull, the woolly rhinoceros could look forward when the head was lowered. Furthermore, there was an increase in the crown height of the teeth with a simultaneous increase in the proportion of cement and a stronger development of the nose area, which probably led to an increase in the size of the horn. With regard to the musculoskeletal system, a development from long, narrow limbs in relation to the body to short, wide limbs with strong muscles can be seen, which can be seen as a progressive adaptation to life in open landscapes. The evolutionary changes from Coelodonta nihowanensis to Coelodonta tologoijensis to Coelodonta antiquitatis , which culminated in the typical development of the woolly rhinoceros in the Glaciation of the Vistula, are regarded as gradual.

The woolly rhinoceros died out at the end of the last glacial period as part of the Quaternary extinction wave . The last representatives of this species can be found in the British Isles around 35,000 years ago. It probably disappeared from continental Europe around 14,000 years ago during the Elder Dryas . The most recent dates include those from Gönnersdorf , which date from 16,640 to 15,800 years ago; the finds from Vaumarcus ( Switzerland ) are only slightly older . In the Urals and Western Siberia, the woolly rhinoceros seems to have survived into the Younger Dryas period , as dates from Lugovskoy (about 30 km west of Khanty-Mansiysk ) are between 13,200 and 11,990 years ago. For eastern Siberia, the most recent data for the woolly rhinoceros range from 18,480 to 15,340 years ago. More recent finds have been reported from East Asia - for example, the date is 7,400 years ago. These dates are rejected on the one hand due to problems in the dating methodology, on the other hand, in contrast to Western Eurasian finds, no fossil has been directly dated so far.

It is controversial in science whether climate changes, hunting by the Young and Late Paleolithic humans or both were responsible for the extinction of the woolly rhinoceros. Genetic studies from 2020 on more than a dozen Siberian woolly rhinos show that the population there has experienced a continuous decline since the Eem warm period , around 29,500 years ago, i.e. shortly before the icing peak of the last cold period, but experienced a significant increase and over remained relatively stable for the next 13,000 years. In this period before the species immediately disappeared, there was no decrease in genetic diversity or a stronger incest caused by a potential decline in populations, in contrast to the wool-haired mammoths on Wrangel Island shortly before their extinction. This also means that there is no evidence of greater pressure on the population, which would, for example, indicate more intensive hunting on the part of humans. The authors of the study explain the eventual extinction of the species with the rapid climatic changes at the end of the last glacial period, which began around 16,000 years ago. In the Bølling- and Allerød-Interstadial , these led to an increasing expansion of forest habitats along with the decline of open grasslands as the primary habitat of the woolly rhinoceros. Combined with the more frequent snowfall under more humid climatic conditions , this could have severely impaired the nutritional basis of the woolly rhinoceros and thus have been one of the main factors for the extinction of the species.

Systematics

Relationship of the woolly rhinoceros to other representatives of the Dicerorhinina. In addition to the representatives of the genus Coelodonta, the Sumatran rhinoceros and the Western Eurasian representatives of the Stephanorhinus line are shown.

The woolly rhinoceros is a kind from the genus Coelodonta within the family of rhinos (Rhinocerotidae) currently five recent includes representatives in Africa and Asia. Due to the endemic distribution of the woolly rhinoceros in Eurasia and individual anatomical similarities, a closer relationship with the Asian than with the African rhinos was suspected early on. Molecular genetic studies of the mitochondrial DNA of woolly rhinos from the Scladina grotto in Belgium showed that this species is most closely related to the Sumatran rhinoceros , which was also confirmed by subsequent studies. According to this, the extinct form can be placed in the sub-tribus of the Dicerorhinina, a group of two-horned rhinos. According to the genetic data, both lines separated in the late Oligocene or early Miocene around 21 to 26 million years ago, although some studies also assume a much more recent date. Already more than 29 million years ago, the common ancestor of Dicerorhinina and Rhinocerotina to which the Indian split rhinoceros ( Rhinoceros unicornis ) and Javan rhinoceros ( Rhinoceros sondaicus ) include, from the African from rhinos.

Closer relationship of Coelodonta according to Cappellini et al. 2019
 Rhinocerotini  

  Dicerorhinina  



 Coelodonta (woolly rhinoceros and others (†))


   

 Stephanorhinus kirchbergensis (†)



   

 Stephanorhinus sp. (†)



   

 Dicerorhinus (Sumatran rhinoceros)



  Dicerotina  

 Ceratotherium (white rhinoceros)


   

 Diceros (black rhinoceros)




  Rhinocerotina  

 Rhinoceros (armored and Java rhinoceros)



Template: Klade / Maintenance / Style

In addition to the genera Coelodonta and Dicerorhinus , Stephanorhinus (occasionally also called Brandtorhinus ) belongs to the Dicerorhinina, the two last representatives of which, the forest rhinoceros ( Stephanorhinus kirchbergensis ) and the steppe rhinoceros ( Stephanorhinus hemitoechus ), also in the Middle and Young Pleistocene, but in different biocene northern Eurasia lived. From an anatomical point of view , Coelodonta and Stephanorhinus are more closely related to each other than to other genera and form a clade ; A Coelodonta-Stephanorhinus group is often used . A main argument for this is the ossification of the nasal septum, which only occurs in these two genera. These ossifications begin relatively early in the evolution of the Stephanorhinus forms and progress from the forest rhinoceros to the steppe rhinoceros until they finally reach their peak in the woolly rhinoceros. Another common feature of Coelodonta and Stephanorhinus is the lack of incisors in the permanent dentition, while they are rudimentary in the deciduous dentition. Dicerorhinus, on the other hand, has permanent incisors, albeit in fewer numbers. Therefore, some researchers advocate merging the genera Stephanorhinus and Coelodonta . However, it is often pointed out that both have been described as separate genera. Molecular genetic analyzes and protein sequence studies confirm the close relationship of the three genera. A biochemical study of dental material from the 1.8 million year old site of Dmanissi published in 2019 comes to the conclusion that Stephanorhinus is polyphyletic with regard to Coelodonta , which would advocate the union of both genera. Both Coelodonta and Stephanorhinus can therefore be traced back to a common ancestor. The latter genus has been documented at least since the end of the Miocene in western Eurasia, the former developed in the Pliocene in eastern Eurasia.

Research history

Johann Friedrich Blumenbach
The Handbook of Natural History by Johann Friedrich Blumenbach from 1799 (6th edition) with the first description of the woolly rhinoceros

One of the earliest scientific mentions of a rhino species that once lived in northern Eurasia was made in 1769 when the German naturalist Peter Simon Pallas (1741–1811) wrote a report on his expeditions to Siberia and found a skull and two horns of such an animal from the permafrost there mentioned. The name Rhinoceros lenenesis , which was introduced in 1772, also came from him , whereby the investigations leading to the naming were based on several skeletal finds from Siberia including the ice mummy from the Wiljui River discovered in 1771. This scientific name is largely no longer in use today; some recent Russian researchers have occasionally used it for Yakut finds, but this is not generally accepted. Only a few years later, in 1782, Johann Heinrich Merck (1741–1791) determined fossil rhino finds from Germany for the first time, including from the Odenwald . The first scientific description that is valid today , then under the name Rhinoceros antiquitatis , was made in 1799 by the German zoologist and anthropologist Johann Friedrich Blumenbach (1752-1840). The syntype mainly includes finds from the area around Göttingen in Lower Saxony. These include an incomplete upper arm from Scharzfeld and several bones (including an occiput, a second cervical vertebra and several long bones) as well as teeth (two premolars) from the gypsum karst landscape of Hainholz near Düna (Osterode am Harz) , all of which were discovered in the middle of the 18th century . In addition, the syntype also includes a fragmented skull from the former Ufa governorate (Russia), which the Baron Georg Thomas von Asch ( Russian. Егор Фёдорович Аш , Jegor Fjodorowitsch Asch; 1729–1807) had given to the University of Göttingen around 1790 and who was only rediscovered in 2005.

The name Rhinoceros tichorhinus comes from the German paleontologist Johann Gotthelf Fischer von Waldheim (1771-1853) , the species name referring to the characteristic, completely ossified nasal septum. Tichorhinus was later occasionally used as a generic name, but is also not valid. The recognized generic name Coelodonta, however, was established in 1831 by the German geologist Heinrich Georg Bronn (1800–1862). The name Coelodonta antiquitatis , which is valid today , first appeared during the 1930s. The generic name Coelodonta comes from the Greek words κοιλία ( koilía 'cave') and ὀδούς ( odoús 'tooth') and refers to the characteristic indentation in the middle of the molars, while antiquus is of Latin origin and means "old".

During research on the woolly rhinoceros different scientific names were used for this species:

  • Rhinoceros lenensis Pallas 1773
  • Rhinoceros antiquitatis Blumenbach 1799
  • Rhinoceros tichorhinus Fischer 1814
  • Rhinoceros Pallasi Desmarest 1822
  • Coelodonta bojei Bronn 1831
  • Rhinoceros tichorhinus Cuvier 1834–1836
  • Rhinoceros tichorhinus Brandt 1849
  • Rhinoceros tichorhinus pediment 1851
  • Rhinoceros antiquitatis Falconer 1868
  • Rhinoceros jourdani Lortet and Chantre 1876
  • Rhinoceros (Tichorhinus) antiquitatis Brandt 1877
  • Rhinoceros Mercki Schrenck 1880
  • Rhinoceros antiquitatis tichorhinus Niezabitowski 1914
  • Tichorhinus antiquitatis (= tichorhinus) Wüst 1922
  • Rhinoceros antiquitatis Hilzheimer 1924
  • Tichorhinus antiquitatis Zeuner 1934

Coelodonta tologoijensis was first described in 1966 by E. I. Beliajewa based on finds from Tologoj Mountain on the Selenga in the Transbaikalia region (Russia). The name Coelodonta nihowanensis, on the other hand, was introduced by the German paleontologist Hans-Dietrich Kahlke in 1967 on finds from the Nihewan Basin ( Hebei Province , China), without this name being recognized; the same applies to the homonymous name given by Chow B. S. 1978. However, it was later pointed out that the first description of this species was carried out in compliance with the nomenclature rules. At the moment this species name is considered to be the noun nudum . The first description of Coelodonta thibetana appeared in 2011, citing an almost complete skull from Tibet.

Cultural history

Finds of large ice age mammals often stimulated the imagination of people in earlier times, such as the use of so-called dragon bones , ground into powder, viewed as a medical panacea in East Asia, or how the interpretation of the skulls of pygmy elephants on the eastern Mediterranean islands as remains of cyclops in Greek mythology shows . Rhinoceros remains in the Siberian permafrost have long been known to the indigenous peoples of Yakutia. Until the 19th century they used the fossil horn of the woolly rhinoceros to improve the elasticity of their hunting bows . In addition, they interpreted the horns as the claws of a huge bird due to the flat cross-section and the curved shape . The zoologist G. Fischer von Waldheim linked these Siberian legends with the ancient stories about the griffin , who, according to Herodotus, lived as an eagle-lion hybrid in the north between the Scythians and the mythical Hyperboreans . According to Fischer, the origin of this legend was to be found in the Urals. As early as 1823, these reports had prompted the German natural scientist Gotthilf Heinrich von Schubert (1780–1860) to introduce the name Gryphus antiquitatis for this supposed “fossil griffin”.

In Europe, too, fossils of the woolly rhinoceros were found very early, for example a rhinoceros skull came to light in 1335 in the Zollfeld near Klagenfurt . Around 250 years later, this was used by the sculptor Ulrich Vogelsang as a model for the dragon's head, which he was to design at the Lindwurmbrunnen in Klagenfurt.

literature

  • Paul S. Martin, Richard G. Klein (Eds.): Quaternary Extinctions. A Prehistoric Revolution. The University of Arizona Press, Tucson AZ 1984, ISBN 0-8165-1100-4 .
  • Arno Hermann Müller: Textbook of paleozoology. Volume 3: Vertebrates. Part 3: Mammalia. 2nd, revised and expanded edition. Fischer, Jena 1989, ISBN 3-334-00223-3 .
  • Wighart von Koenigswald: Living Ice Age. Climate and fauna in transition. Theiss, Stuttgart 2002, ISBN 3-8062-1734-3 .

Individual evidence

  1. ↑ a b Нина В. Гарутт и Геннади Г. Боескоров: Шерстистые носороги: к истории рода . In: Geos . 2001, p. 157-167 .
  2. ↑ a b c d e f g h i j Ralf-Dietrich Kahlke: The origin, development and distribution history of the Upper Pleistocene Mammuthus-Coelodonta Faunenkomplex in Eurasia (large mammals) . In: Treatises of the Senckenbergische Naturforschenden Gesellschaft . tape 546 . Frankfurt am Main 1994.
  3. ↑ a b c Diego J. Álvarez-Lao and Nuria García: Geographical distribution of Pleistocene cold-adapted large mammal faunas in the Iberian Peninsula . In: Quaternary International . tape 233 , no. 2 , February 15, 2011, p. 159–170 , doi : 10.1016 / j.quaint.2010.04.017 .
  4. ^ 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, p. 321-340 .
  5. ↑ a b c d e Gennady Boeskorov: Woolly rhino (Coelodonta antiquitatis) distribution in Northeast Asia . In: Deinsea . tape 8 , 2001, p. 15-20 ( PDF ).
  6. ↑ a b c d e f g h i j Ralf-Dietrich Kahlke and Frédéric Lacombat: The earliest immigration of woolly rhinoceros (Coelodonta tologoijensis, Rhinocerotidae, Mammalia) into Europe and its adaptive evolution in Palaearctic cold stage mammal faunas . In: Quaternary Science Reviews . tape 27 , 2008, p. 1951–1961 , doi : 10.1016 / j.quascirev.2008.07.013 .
  7. ↑ a b c d e f H. Loose: Pleistocene Rhinocerotidae of W. Europe with reference to the recent two-horned species of Africa and SE Asia . In: Scripta Geologica . tape 33 , 1975, pp. 1-59 .
  8. ^ Scott A. Elias, Barnaby Crocker: The Bering Land Bridge: a moisture barrier to the dispersal of steppe – tundra biota? In: Quaternary Science Reviews . tape 27 , 2008, p. 2473-2483 , doi : 10.1016 / j.quascirev.2008.09.011 .
  9. ↑ a b c d Wighart von Koenigswald: Lebendige Eiszeit. Climate and fauna in transition . Stuttgart 2002, p. 54-61 .
  10. ↑ Diana Pushkina: The Pleistocene easternmost distribution in Eurasia of the species associated with the Eemian Palaeoloxodon antiquus assemblage . In: Mammal Review . tape 37 , no. 3 , 2007, p. 224-245 ( [1] ).
  11. ↑ Hans-Dietrich Kahlke: On the chronological position of the Choukoutien culture . In: Old Thuringia . tape 6 , 1963, pp. 22-41 .
  12. ↑ a b c d e f g h i j Jan van der Made: The rhinos from the Middle Pleistocene of Neumark-Nord (Saxony-Anhalt) . In: Dietrich Mania u. a. (Ed.): Neumark-Nord: An interglacial ecosystem of the Middle Paleolithic man. Publications of the State Museum for Prehistory . tape 62 . Halle / Saale 2010, p. 433-527 .
  13. ↑ a b Tong Haowen: Palaeoenvironmental significance of Coelodonta in different fossil assemblages . In: Acta Anthropologica Sinica . tape 23 , 2004, pp. 306-314 .
  14. ↑ a b c d e f Claude Guérin: Coelodonta antiquitatis praecursor (Rhinocerotidae) du Pléistocène moyen final de l'aven de Romain-la-Roche (Doubs, France) . In: Revue de Paléobiologie . tape 29 , 2010, p. 697-746 .
  15. ^ Walter Steiner: The travertine of Ehringsdorf and its fossils . Ed .: Dong Wei. tape 23 . Lutherstadt Wittenberg 1981.
  16. ^ Dietrich Mania and Dieter-Hans Mai: On the climatic character of the warm period of Neumark-Nord 1 . In: Harald Meller (Hrsg.): Elefantenreich - Eine Fossilwelt in Europa . Halle / Saale 2010, p. 174-185 .
  17. ^ Anthony J. Stuart: The extinction of woolly mammoth (Mammuthus primigenius) and straight-tusked elephant (Palaeoloxodon antiquus) in Europe. In: Quaternary International: Volume 126-128, 2005, pp 171-177.
  18. ↑ a b c Cajus G. Diedrich: A skeleton of an injured Coelodonta antiquitatis from the Late Pleistocene of north-western Germany . In: Cranium . tape 25 , 2008, p. 9-23 .
  19. ↑ a b c d Magdalena Borsuk-Białynicka: Studies on the Pleistocene Rhinoceros Coelodonta antiquitatis (Blumenbach) . In: Palaeontologia Polonica . tape 29 . Warsaw-Krakow 1973.
  20. ^ A b c Emmanuel ME Billia: A brief review of the "woolly rhino" Coelodonta antiquitatis (Blumenbach, 1799) (Mammalia, Rhinocerotidae) in Italy (Southern Europe) . In: VM Podobina (Ed.): Evolution of life on the earth: Proceedings of the IV international symposium November 10-12, 2010 . Tomsk 2010, p. 529-533 .
  21. ↑ Tong Haowen and Anne-Marie Moigne: Quaternary rhinoceros of China . In: Acta Anthropologica Sinica . tape 19 , 2000, pp. 257-263 ( PDF ).
  22. ↑ Tong Haowen: Rhinocerotids in China - systematics and material analysis . In: Geobios . tape 34 , no. 5 , 2001, p. 585-591 , doi : 10.1016 / S0016-6995 (01) 80071-1 .
  23. ^ A b Cajus G. Diedrich: Bone remnants brought in and gnawed by Coelodonta antiquitatis (Blumenbach 1807) from the Upper Pleistocene spotted hyena nest Perick caves in northern Sauerland and contribution to the taphonomy of woolly rhinoceros carcasses in northwest Germany . In: Communications of the Association of German Caves and Caves Karst explorers . tape 54 , no. 4 , 2008, p. 100-117 .
  24. ↑ a b c П. А. Лазарев, С. Е. Григорев и В. В. Плотников: Шерстистые носороги Якутии . In: VM Podobina (Ed.): Evolution of life on the earth: Proceedings of the IV international symposium, November 10-12, 2010 . Tomsk 2010, p. 555-558 .
  25. ↑ a b c Gennady G. Boeskorov, PA Lazarev, NT Bakulina, MV Shchelchkova, SP Davydov and NG Solomonov: Preliminary study of woolly rhinoceros from the lower reaches of the Kolyma river . In: Doklady Biological Sciences . tape 424 , 2009, p. 53-56 .
  26. ↑ Gennady G. Boeskorov, Peter A. Lazarev, Andrei V. Sher, Sergei P. Davydov, Nadezhda T. Bakulina, Marina V. Shchelchkova, Jonas Binladen, Eske Willerslev, Bernard Buigues and Alexey N. Tikhonov: Woolly rhino discovery in the lower Kolyma River . In: Quaternary Science Reviews . tape 30 , no. 17 , 2011, p. 2262–2272 , doi : 10.1016 / j.quascirev.2011.02.010 .
  27. ↑ a b c Irina V. Kirillova and Fedor K. Shidlovskiy: Estimation of individual age and season of death in woolly rhinoceros, Coelodonta antiquitatis (Blumenbach, 1799), from Sakha-Yakutia, Russia . In: Quaternary Science Reviews . tape 29 , no. 23 , 2010, p. 3106-3114 , doi : 10.1016 / j.quascirev.2010.06.036 .
  28. ↑ Prometheus: Baby woolly rhinoceros found - Siberian permafrost soil releases sensational discovery. March 1, 2015 ( [2] ).
  29. ↑ Anna Liesowska: Meet Sasha - the world's only baby woolly rhino; Siberian Times. March 1, 2015 ( [3] ).
  30. ↑ a b OF Chernova, AV Protopopov, TV Perfilova, IV Kirillova and GG Boeskorov: Hair Microstructure of the First Time Found Calf of Woolly Rhinoceros Coelodonta antiquitatis. In :: Doklady Biological Sciences. Volume 471, 2016, pp. 291-295.
  31. ↑ a b c Нина В. Гарүтт: К истории изучения шерстистого носорога Coelodonta antiquitatis (Blumenbach, 1799) . In: Geos . 2000, p. 22-33 .
  32. ↑ Maciej J. Kotarba, Marek Dzieniewicz, Włodzimierz J. Mościcki and Henryk Sechman: Unique Quaternary environment for discoveries of woolly rhinoceroses in Starunia, fore-Carpathian region, Ukraine: Geochemical and geoelectric studies . In: Geology . tape 36 , no. 7 , 2008, p. 567-570 .
  33. ^ Henryk Kubiak and Daniel M. Drygant: The Starunia collections in Lviv and Kraków Natural History Museums and history of paleontological studies . In: MJ Kotarba (Ed.): Geological Studies 2004–2005 at Starunia - The area of ​​discoveries of woolly rhinoceroses . Warszawa-Kraków 2005, p. 37-44 .
  34. ^ Mikael Fortelius (coordinator). Neogene of the Old World Database of Fossil Mammals (NOW) . University of Helsinki, 2003.
  35. ↑ Jan van der Made and René Grube: The rhinoceroses from Neumark-Nord and their nutrition . In: Harald Meller (Hrsg.): Elefantenreich - Eine Fossilwelt in Europa . Halle / Saale 2010, p. 382-394 .
  36. ↑ a b c Tao Deng, Xiaoming Wang, Mikael Fortelius, Qiang Li, Yang Wang, Zhijie J. Tseng, Gary T. Takeuchi, Joel E. Saylor, Laura K. Säilä and Guangpu Xie: Out of Tibet: Pliocene Woolly Rhino Suggests High-Plateau Origin of Ice Age Megaherbivores . In: Science . tape 333 , 2011, pp. 1285-1288 .
  37. ^ A b c Nina V. Garutt: Dental ontogeny of the wolly rhino Coelodonta antiquitatis (Blumenbach, 1799) . In: Cranium . tape 11 , 1994, pp. 37-48 .
  38. ↑ a b c d e f g Mikael Fortelius: The morphology and paleobiological significance of the horns of Coelodonta antiquitatis (Mammalia: Rhinocerotidae) . In: Journal of Vertebrate Paleontology . tape 3 , no. 2 , 1983, p. 125-135 .
  39. ↑ a b c Nina Garutt: New information about the horn of the fur rhinoceros Coelodonta antiquitatis . In: Deinseas . tape 4 , 1998, pp. 25-39 .
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  82. ↑ Е. И. Беляева: Семейство Rhinocerotidae Owen 1845 . In: А. В. Вангенгейм, Е. И. Беляева, Е. Гарутт, Е. Л. Дмитриева и В. С. Зажигин (Ed.): Млекопитающие Еоплейстоцена западного Забайкалья. Труды Геологического Института Академии Наук СССР . tape 152 . Moscow 1966, p. 92-143 .
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This article was added to the list of excellent articles on December 17th, 2012 in this version .