Forest rhinoceros

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Forest rhinoceros
skull

skull

Temporal occurrence
Middle Pleistocene to Young Pleistocene
600,000 to 100,000 years
Locations
  • South, Central and Eastern Europe
  • West, Central, East Asia
Systematics
Higher mammals (Eutheria)
Laurasiatheria
Unpaired ungulate (Perissodactyla)
Rhinoceros (Rhinocerotidae)
Stephanorhinus
Forest rhinoceros
Scientific name
Stephanorhinus kirchbergensis
( Hunter , 1839)

The rhinoceros ( Stephanorhinus kirchbergensis ), after its discoverer, Carl Heinrich Merck also Merck'sches rhino or Merck rhinoceros called, was a Pleistocene rhinoceros in northern Eurasia . It lived at the same time as the steppe rhinoceros ( Stephanorhinus hemitoechus ), but, in contrast to this, inhabited more wooded biotopes and mainly fed on soft vegetable food. Both rhino species became extinct in the course of the Young Pleistocene.

distribution and habitat

Distribution area of Stephanorhinus kirchbergensis in northern Eurasia . It shows the maximum extent of the forest rhinoceros during the warm periods. The black points represent important sites
Lower jaw and pelvic bones of a forest rhinoceros, found in Flurlingen near Schaffhausen

The wood rhinoceros was distributed over large parts of Eurasia from Western Europe to Southern Siberia , Central Asia and China in the Middle to Young Pleistocene . Its easternmost occurrence is known from the Korean Peninsula . As a heat-loving animal, it required at least a sub-Mediterranean climate and therefore had its core area mainly in the Mediterranean region, in the Middle East and Central Asia. It is noteworthy, however, that this species was not found on the Iberian Peninsula . In the Pleistocene warm periods , in which the temperatures in the climatic optimum were sometimes up to 3 ° higher than today, the forest rhinoceros spread far into the northern Eurasian areas. In northern Alpine Europe it reached its northernmost point with England (Illford, Grays and Clacton-on-Sea ) during the Holstein warm period (340,000 to 325,000 years ago) . In the Eem warm period (126,000 to 115,000 years ago), however, it was unable to cross the English Channel again and was restricted to mainland Europe. Also in its eastern distribution area, the wood rhinoceros expanded its habitat from its core areas (probably Central Asia and northern China) far north to Siberia during the warm periods. Remnants of the skull and bones of this rhinoceros species were found here in the alluvial lands of the Don and Volga in the Russian steppe , as well as in the Kuznetsk Basin and near Irkutsk on Lake Baikal . The northern limit of distribution is usually at 53 to 54 degrees north latitude. Two teeth were recovered from the Wiljui river basin in the Sakha Republic in Yakutia ; the site is located near the 64th parallel north. Such a high northern occurrence has so far rarely been documented, but a skull was discovered on the Chondon River in the Jana Indigirka lowlands in Yakutia at the 70th parallel north.

The wood rhinoceros was a typical warm-time fauna element. Its presence at sites is used as a measure of the change in vegetation belts in the course of cold and warm periods. In western Eurasia it was part of the so-called Palaeoloxodon antiquus fauna complex , which includes not only the eponymous European forest elephant but also the aurochs , red deer and fallow deer , roe deer and hippopotamus . The steppe rhinoceros , with which it co-existed, was also a typical representative of this faunal community. Unlike its cousin that appeared at the same time, the forest rhinoceros only appeared during the climatic optimum. When it cooled, it withdrew very quickly to the south into warmer climes. Since no remains of the forest rhinoceros have been found in Portugal or Spain so far, it is assumed that this region was not part of his core refuge. Rather, it seems to have migrated back from northern Alpine Europe to Asia or Southeast Europe every time .

In Central Asia, the forest rhinoceros also belonged to the Koshkurgan fauna complex , named after a Middle Pleistocene archaeological travertine site in the south of Kazakhstan , which also includes finds from Shoktas, also Kazakhstan, and Lachuti ( Tajikistan ). Here it was with the Südelefanten , the Mosbacher horse , the Schoetensack - Bison and the Elasmotherium socialized. In southern Siberia, for example In the Kuznetsk Basin, for example, the wood rhinoceros was part of the Tatar fauna complex , which is also Middle Pleistocene , which in turn includes the red deer, the giant deer but also the mammoth . In East Asia, this animal species - which is locally referred to by the scientific name Stephanorhinus (Dicerorhinus) choukoutienensis - is a companion of the Equus Euctenoceros / Megaloceros Faunenkomplexes (also North Tsingling Faunenkomplex ), which is mainly distributed in northern China and in of Mongolia had. In addition to the eponymous horse and the giant deer, this complex also includes Palaeoloxodon namadicus and some northern elements, such as the marmot or the musk ox . Occasionally there are also rhinos of the genus Coeledonta , which, however, do not belong to the typical woolly rhinoceros and are therefore not regarded as cold climatic indicators like this one. In mainly in southern China and Indochina -spread Stegodon-Ailuropoda Faunenkomplex (also South Tsingling-Faunenkomplex called), named after the elephantine Stegodon and the giant panda , the rhinoceros had not previously been detected and should here according to previous opinion by the also now extinct Chinese rhinoceros ( Rhinoceros sinensis ) have been represented. More recent investigations, including from the rhinoceros cave near Shennongjia ( Hubei Province ), a very fossil-rich site, show that the wood rhinoceros occasionally appear in this fauna complex and obviously sought ecological niches that corresponded to its northern refuges. It advanced south in eastern China to around 30 degrees of latitude.

Fossils of this type are found relatively rarely compared to other large Pleistocene mammals. The reasons for this could either be the poor fossilization conditions or the species was actually relatively rare. In general, it is found more frequently in western Eurasia than in eastern Eurasia. In Germany, the wood rhinoceros is known from more than a dozen sites. An almost complete skeleton was recovered from the Pleistocene cover layers of the lignite mine of Neumark-Nord in the Geiseltal ( Saxony-Anhalt ) and belongs to the Eem warm period. In addition, the forest rhinoceros became known in the warm-time deposits of the important, up to 600,000 year old fossil site Mosbacher Sande in the city of Wiesbaden ( Hesse ) and from the 300,000 year old early human discovery site Steinheim an der Murr ( Baden-Württemberg ). Other important sites here are the 350,000-year-old Homo erectus site in Bilzingsleben ( Thuringia ), the site where the Schöninger Speere ( Lower Saxony ), which was roughly the same age, and the Ehringsdorf Neanderthal station, which was dated to around 220,000, possibly only 120,000 years ago (also in Thuringia).

Physique and diet

Schematic representation of the head posture of the three rhinoceros species occurring in the Middle and Young Pleistocene of Europe . Above: forest rhinoceros ( Stephanorhinus kirchbergensis ), middle: steppe rhinoceros ( Stephanorhinus hemitoechus ), below: woolly rhinoceros ( Coelodonta antiquitatis ). Without lower jaw , each shown in left side view
Lower jaw fragment from the forest rhinoceros

As there are no cave drawings or frozen remains of the forest rhinoceros in contrast to the Middle and Young Pleistocene woolly rhinoceros ( Coelodonta antiquitatis ), much less is known about its appearance than that of its northern relative. Anatomical research shows that it had a strong build and very long and narrow front and rear legs, the joints of which were pronounced. The last two features in particular show that the forest rhinoceros moved in areas that had a rather closed vegetation cover. Within the Stephanorhinus line, the forest rhinoceros was their largest representative. Reconstructed body weights vary between 1.6 t and a maximum of 2.9 t and are thus within the range of variation of the Indian rhinoceros .

The head is between 70 and 80 cm long and looks relatively small due to the enormous body size, which in the past has often led to discussions about the actual body size. A striking feature of the skull is the shortened and therefore more rectangular occiput, which caused the forest rhinoceros to keep its head clearly upright. This makes it similar to the earlier Stephanorhinus species, but differs markedly from its relative, the steppe rhinoceros, which held its head much further down. In this characteristic, the forest rhinoceros also resembles today's black rhinoceros ( Diceros bicornis ) or its closest living relative, the Sumatran rhinoceros ( Dicerorhinus sumatrensis ). Like the other representatives of Stephanorhinus , it had two horns on the nose and on the middle skull, the attachment points of which on the bone surfaces are roughened like cauliflower. The front horn was larger because of these roughened surfaces and rose steeply to the top. Presumably, however, it was not as big as in the steppe rhinoceros, which, in addition to the smaller size and shape of the attachment surfaces, is also indicated by the nasal septum , which, in contrast to its close relative, is only ossified in the front third. Another feature is the extremely solidly built lower jaw , which in terms of robustness far exceeds that of its relatives and was up to 60 cm long.

As with all Stephanorhinus TYPES missing teeth in the incisors , still occurred each arch three Vorbackenzähne and three molars on the dental formula thus read: . Of all the species of the Stephanorhinus line, the forest rhinoceros has the largest teeth in relation to head size, with the dimensions of the teeth increasing from front to back. The third molar particularly stands out here, as it was extremely voluminous. This characteristic indicates that this animal species mainly consumed large amounts of plant-based food, which, however , offered little nutrients overall . The teeth are relatively high crowned , with the crown height decreasing from the premolars to the molars, but do not reach the values ​​of the steppe rhinoceros. However, they have less dental cement , which is characteristic of animal species that mainly eat soft foods such as leaves , flowers , berries or fruits . This is also indicated by the tooth surfaces, which are always trough-shaped in the middle, which is caused by abrasion when chewing the food. Investigations on food residues found on the teeth of the forest rhinoceros show that this species of animal, at least in Central Europe, mainly fed on plant species such as birch , rose , poplar , oak , hawthorn and firethorn, but also on water lilies and sometimes grass . Using plant remains from the teeth of the skull of the Chondon River in Yakutia, the animal mainly fed on leaves and twigs . Among them were the remains of larches , blueberries , birches and star mosses . Mosses made up more than 20% of the remains found, but no evidence of sour grasses was found. The proven food spectrum differs significantly from that of the woolly rhinoceros in the same region and is more similar in its general composition to that of the black rhinoceros .

Remnants of the trunk skeleton and the limbs are rarely passed down and far less known than from the related Pleistocene rhinos of Eurasia. The spine comprised at least 7 cervical, 18 thoracic and 4 lumbar vertebrae, but the exact number of sacral and caudal vertebrae is unclear. The radius was up to 46 cm long. The thigh bone was very massive at 56 cm, the shin reached 46 cm in length. Most likely, the forest rhinoceros, like today's modern rhinos and the other fossil species of Stephanorhinus, had three-pointed hands and feet with a particularly powerful central ray (Metapodium III). The third metacarpal bone was 25 cm long , the third metatarsal bone 21 cm long.

Since there are no surviving soft tissues, nothing is known about the future appearance of the forest rhinoceros. It is also unclear whether this rhinoceros had a fur; there is still a lack of information about the texture of the skin , the possible formation of skin folds or the size and shape of the ears . In addition, there is little evidence of the way of life of this animal species. The skeleton structure and the way of eating make you think of a forest dweller. It is unclear whether the forest rhinoceros actually lived in dense forests or rather in the transition area from the forest edges to the open landscapes.

Systematics and tribal history

The forest rhinoceros was one of the last representatives of the genus Stephanorhinus , as its contemporary the steppe rhinoceros ( Stephanorhinus hemitoechus ) also survived until the late Pleistocene . The related Sumatran rhinoceros ( Dicerorhinus sumatrensis ) has survived to this day in the forests of Southeast Asia , although human influences have decimated it to a few hundred animals. The genus Coelodonta , which includes the woolly rhinoceros , is also closely related . All three genera belong to the tribe Dicerorhinina, which represent a rather primitive group within today's modern rhinos. Common features are the two horns and the partially or completely fused nasal septum, Dicerorhinus differs from the other genera by the developed anterior teeth. Above all, the ossification of the nasal septum is unique and does not occur in the other recent rhino representatives.

As with the steppe rhinoceros, little is known about the origin. Presumably it descends from Stephanorhinus magarhinus and together with him forms a sister clade of the line Stephanorhinus etruscus - Stephanorhinus hundsheimensis - Stephanorhinus hemitoechus . It is still unclear exactly when the species originated. Here, too, an origin in Asia is assumed. It first appeared in Europe around 600,000 years ago, including in the Mosbacher Sanden (Wiesbaden, Hesse; more precisely Mosbach 2 ). and Ponte Molle ( Italy ). Older sources such as B. Soleilhac ( France ) or Tegelen ( Netherlands ), are likely to be confused with other representatives of the Stephanorhinus line. In East Asia, the forest rhinoceros appeared with certainty about 570,000 to 580,000 years ago at the famous early human discovery site Zhoukoudian (China, locality 1 ). Although it was also described from the Gongwangling site , a site more than 700,000 years old, the remains there are too sparse. In Central Asia, the forest rhinoceros can only be detected 500,000 to 550,000 years ago based on the sites of Lachuti (Tajikistan) and Koshkurgan (Kazakhstan).

The earliest forest rhinos had a very small third molar tooth that only increased in size over time. It then reached its greatest dimensions in the late Middle and early New Pleistocene. These changes in the dentition are often taken as biostratigraphic indicators. In contrast to the steppe rhinoceros, however, it did not increase in size; in contrast, numerous populations, especially in southern Europe, developed smaller forms.

In the cooler phases of the end of the Eem warm period, the forest rhinoceros withdrew from the northern refuges to the south, as with previous climatic changes. Evidence of the species from this late period comes from the Kůlna cave in the Moravian Karst (Czech Republic, layer 9b ), from the Veternica cave ( layer j ) and the Vindija cave ( layer K ) (both Croatia ) and from Grimaldi (Italy ). However, shortly after the beginning of the last glaciation period ( Vistula Glaciation ) it must have died out, as it can no longer be detected in the further course of the Young Pleistocene. Recent finds include the skull of the Chondon River in Yakutia, which dates between 70,000 and 48,000 years ago. The same fate, albeit later, also befell the European forest elephant and the steppe rhinoceros. ( see Quaternary extinction wave ).

Research history

Georg Friedrich von Jäger

The forest rhinoceros was first scientifically described as a fossil species in 1839 by the Stuttgart doctor and palaeontologist Georg Friedrich von Jäger (1785–1866). The scientific name Rhinoceros kirchbergensis introduced by Jäger was based on dental finds from Kirchberg an der Jagst in Württemberg . The later used genus name Dicerorhinus was replaced in 1942 by the name Stephanorhinus introduced by the Hungarian paleontologist Miklós Kretzoi (1907-2005) . He justified this with numerous existing anatomical differences to the Sumatran rhinoceros, which is still alive today, as the last representative of the genus Dicerorhinus .

In the course of time, the wood rhinoceros came under various scientific names:

  • Rhinoceros incisivus Merck 1784
  • Rhinoceros megarhinus de Christol 1834
  • Rhinoceros leptorhinus Cuvier 1836
  • Rhinoceros kirchbergensis hunter 1839
  • Rhinoceros Merckii ( merckii, mercki, merki, Mercki ) Kaup 1841
  • Dicerorhinus mercki Kaup 1841
  • Rhinoceros leptorhinus Owen 1850
  • Rhinoceros (Tichorhinus) Merckii Brandt 1877
  • Rhinoceros Mercki (Merckii) var.Brachycephala Schroeder 1903
  • Coelodonta merckii Abel 1919
  • Dicerorhinus kirchbergensis Hooijer 1947
  • Dicerorhinus mercki (kirchbergensis) (hunter) var.Brachycephalus Schroeder
  • vel Dicerorhinus merckii Mayer 1971

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 .

Web links

Commons : Forest rhinoceros ( Stephanorhinus )  - Collection of images, videos and audio files

Individual evidence

  1. ^ A b Johann Jakob Kaup: Files of the primeval world or osteology of primeval mammals and amphibians. Darmstadt, publisher's publisher, 1841
  2. ↑ a b c d e f g h Emmanuel ME Billia: Revision of the fossil material attributed to Stephanorhinus kirchbergensis (Jäger 1839) (Mammalia, Rhinocerotidae) preserved in the museum collections of the Russian Federation. Quaternary International, Volume 179, Issue 1, March 2008, pp. 25-37
  3. ↑ 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 et al. (Ed.): Neumark-Nord: An interglacial ecosystem of the Middle Palaeolithic people. Publications of the State Museum for Prehistory 62. Halle / Saale 2010, pp. 433–527
  4. ↑ a b c Mikael Fortelius, Paul Mazza and Benedetto Sala: Stephanorhinus (Mammalia: Rhinocerotidae) of the Western European Pleistocene, with a revision of S. etruscus (Falconer, 1868). Palaeontographia Italica, 80, 1993, pp. 63-155
  5. ↑ Emmanuel ME Billia: First records of Stephanorhinus kirchbergensis (Jäger, 1839) (Mammalia, Rhinocerotidae) from the Kuznetsk Basin (Kemerovo region, Kuzbass area, South-East of Western Siberia). Bollettino della Società Paleontologica Italiana, 46 (2-3), 2007, pp. 95-100
  6. ^ Emmanuel ME Billia: The skull of Stephanorhinus kirchbergensis (Jaeger 1839) (Mammalia, Rhinocerotidae) from the Irkutsk region (Southwest Eastern Siberia). Quaternary International, Volume 179, Issue 1, March 2008, pp. 20-24
  7. ^ IV Kirillova, OF Chernova, VV Kukarskikh, FK Shidlovskiy and OG Zanina: The First Finding of a Rhinoceros of the Genus Stephanorhinus in Arctic Asia. Doklady Biological Sciences 471, 2016, pp. 300-303
  8. ↑ a b c Irina V. Kirillova, Olga F. Chernova, Jan van der Made and Vladimir V. Kukarskih: Discovery of the skull of Stephanorhinus kirchbergensis (Jäger, 1839) above the Arctic Circle. Quaternary Research, 2017 doi: 10.1017 / qua.2017.53
  9. ↑ Donald R. Prothero, Robert M. Schoch: Horns, tusks, and flippers . The evolution of hoofed mammals. Johns Hopkins Univ Pr, Baltimore 2003, ISBN 0-8018-7135-2 , pp. 272 .
  10. ^ Wighart von Koenigswald: Lebendige Eiszeit. Climate and fauna in transition. Stuttgart, 2002, pp. 153-155.
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  12. ↑ a b c Frederic Lacombat: Phylogeny of the genus Stephanorhinus in the Plio-Pleistocene of Europe. Hallesches Jahrbuch für Geoswissenschaften 23, 2007, pp. 63–65
  13. ^ A b Hans-Dietrich Kahlke: On the chronological position of the Choukoutien culture. Alt-Thüringen 6, 1963, pp. 22-41
  14. ^ Jean S. Aigner: Archaeological remains in Pleistocene China. AVA research 1, Bonn, 1981
  15. ↑ a b Tong HaoWen and WU XianZhu: Stephanorhinus kirchbergensis (Rhinocerotidae, Mammalia) from the Rhino Cave in Shennongjia, Hubei. Chinese Science Bulletin 55 (4), 2010, pp. 1157-1168
  16. ↑ Pierre Olivier Antoine: Pleistocene and holocene rhinocerotids (Mammalia, Perissodactyla) from the Indochinese Peninsula. In: Comptes Rendus Palevol. 2011, pp. 1-10.
  17. ↑ a b c d e f g H.K. Loose: Pleistocene Rhinocerotidae of Western Europe with reference to the recent two-horned species of Africa and SE Asia. Proefschrift [dissertation], Leiden, Scripta Geologica 33, 1975, pp. 1-59
  18. ^ Emmanuel ME Billia: Occurences of Stephanorhinus kirchbergensis (JÄGER, 1839) in Eurasia - An account. Acta Palaeontologica Romaniae 7, 2011, pp. 17-40.
  19. ↑ Karl-Dietrich Adam: The primeval man of Steinheim an de Murr and his environment. A picture of life from a quarter of a million years ago. Yearbook of the Roman-Germanic Central Museum 35, 1988, pp. 1–23
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  21. ↑ Hartmut Thieme (ed.): The Schöninger Speere. Humans and hunting 400,000 years ago. Exhibition catalog. Stuttgart, 2007
  22. ^ Walter Steiner: The travertine of Ehringsdorf and its fossils. Lutherstadt Wittenberg 1981
  23. ↑ Mikael Fortelius (coordinator). Neogene of the Old World Database of Fossil Mammals (NOW). University of Helsinki, 2003 http://www.helsinki.fi/science/now/
  24. ^ Andreas Wagner: Reconstruction of body masses of Pleistocene Rhinocerotidae in the Königswald collection. Frankfurt am Main, 2007
  25. ↑ a b 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, pp. 382–394
  26. ↑ René Grube: Vegetable food remains of the fossil elephants and rhinos from the interglacial of Neumark-Nord. In: Jan Michal Burdukiewicz, Lutz Fiedler, Wolf-Dieter Heinrich, Antje Justus and Enrico Brühl (eds.): Knowledge hunters . Festschrift for Dietrich Mania. Publications of the State Museum for Prehistory in Halle 57. Halle / Saale 2003, pp. 221–236
  27. ^ FE Zeuner: New reconstructions of the whooly rhinoceros and Merck's rhinoceros. Proceedings of the Linnean Society of London 156 (3), 1945, pp. 183-195
  28. ↑ a b c Claude Guérin: Les Rhinocerotidae (Mammalia, Perissodactyla) du Miocène terminal au Pléistocène supérieur d'Europe occidentale comparés aux espèces actuelles: tendances évolutives et relations phylogénétiques. Géobios 15, 1982, pp. 599-605
  29. ^ A b Emmanuel ME Billia and Carmelo Petronio: Selected records of Stephanorhinus kirchbergensis (Jäger, 1839) (Mammalia, Rhinocerotidae) in Italy. Bollettino della Società Paleontologica Italiana, 48, 2009, pp. 1-12.
  30. ↑ Xinzhi Wu and Frank E. Poirier: Human evolution in China. New York, Oxford 1995
  31. ↑ Jan van der Made: Biostratigraphy - "Large mammals". In: Harald Meller (Hrsg.): Elefantenreich - Eine Fossilwelt in Europa. Halle / Saale, pp. 83–92
  32. ↑ Frederic Lacombat: Pleistocene rhinoceroses in Mediterranean Europe and Massif Central (France). Courier des Forschungs-Institut Senckenberg 256, 2006, pp. 57–69
  33. ^ Rudolf Musil: The environment of the Middle Palaeolithic sites in Central and Eastern Europe. In: Jan Michał Burdukiewicz and Andrzej Wiśniewski (eds.): Middle Palaeolithic human activity and palaeoecology: New discoveries and ideas. Studia Archeologiczne 41, Wrocław 2010, pp. 121–180
  34. ↑ Georg Friedrich Jäger: About the fossil mammals that have been found in Württemberg in various formations, along with geognostic remarks about these formations. C. Erhard Verlag, Stuttgart, 1835–39
  35. ↑ Miklós Kretzoi: Comments on the system of post-Miocene rhinoceros genera. Földtany Közlöny 72, 1942, pp. 309-323
  36. ↑ Christol, J. de: Recherches sur les grandes espèces de rhinocéros fossiles. Montpellier, J. Martel, 1834, 170 pages, 31 ills.
  37. ^ Owen, R .: A History of British fossil mammals and birds. London, Joh Van Voorst: IXLVII
  38. ↑ Brandt, JF, attempt at a monograph of the Tichorhinus rhinoceroses with comments on Rhinoceros leptorhinus Cuv. u. s. w. Mém. Ac. Imp. Sc. St. Pétersb., Sér. VII, XXIV, 4, 1877
  39. ↑ H. Schroeder: The vertebrate fauna of the Mosbacher Sand, I genus Rhinoceros. Dep. Kings Preuss. Geol. Landesanst., NF 18, 1903, p. 1143.
  40. ↑ Hooijer, D. Α .: Notes on some fossil mammals of the Netherlands. Arch. Mus. Teyler, 1947, Ser. 3, 10, 1
  41. ^ Mayer, G .: Contributions to the history of the Baden state collections for natural history in Karlsruhe, III. The skull of Dicerorhinus mercki (kirchbergensis) (Jaeger) var. Brachycephalus Schroeder von Daxlanden and its history. Contribution natural Research South Dtl., 30/2, 1971, pp. 157-163.