Plains rhinoceros

from Wikipedia, the free encyclopedia
Plains rhinoceros
skull

skull

Temporal occurrence
Middle to Young Pleistocene
500,000 to 10,000 years
Locations
  • Southern and Central Europe
  • Middle East
Systematics
Higher mammals (Eutheria)
Laurasiatheria
Unpaired ungulate (Perissodactyla)
Rhinoceros (Rhinocerotidae)
Stephanorhinus
Plains rhinoceros
Scientific name
Stephanorhinus hemitoechus
( Falconer , 1859, 1868)

The extinct steppe rhinoceros ( Stephanorhinus hemitoechus ) was a Pleistocene rhinoceros species of Eurasia and North Africa . It was first described scientifically in 1868 by the Scottish paleontologist Hugh Falconer (1808-1865) under the name Rhinoceros hemitoechus . In contrast to the closely related forest rhinoceros ( Stephanorhinus kirchbergensis ), the steppe rhinoceros lived in more open landscapes and for the most part lived on hard grass. Like the forest rhinoceros, it did not become extinct until the late Pleistocene , but according to recent regional studies it survived until the beginning of the Holocene .

distribution and habitat

The steppe rhinoceros lived in large parts of Europe and Asia in the Middle and Young Pleistocene , from the Iberian Peninsula to far into Central Asia . It is also known from sites in the Levant ( Lebanon , Syria , Israel ) and North Africa ( Morocco , Libya ). In North Africa, however, the steppe rhinoceros did not appear until the Young Pleistocene, after it probably immigrated from the north via the Middle East .

Like its close relative, the forest rhinoceros, the steppe rhinoceros was also a form adapted to sub- Mediterranean to warm-temperate climates . Its ancestral habitats were southern Europe and the west and central Asian steppes , bush or forest areas. It is unclear whether, as its name suggests, it was really a typical steppe animal or rather an inhabitant of open forest habitats and park landscapes. Above all, the fact that steppe rhinos are relatively common in fossil records in Europe, but were rarely found in Asia, where steppe areas were much more widespread in the Pleistocene, speaks against a strong adaptation to steppes. However, this animal species seems to have been significantly more resistant to cooler and dry climates than the forest rhinoceros, as it has also been detected in late interglacial or cooler phases of the warm periods north of the Alps and then sometimes occurs together with the woolly rhinoceros ( Coelodonta antiquitatis ). During the warm periods of the Central and Young Pleistocene, such as the Holstein warm period (340,000 to 325,000 years ago) and the Eem warm period (126,000 to 115,000 years ago), it also spread far into northern Alpine Europe ( Central and Eastern Europe) ), with sites in the Netherlands (Zwarte Water), England (Illford, Barrington) and Wales (Minchin Cave ) marking its northernmost occurrence. In the cold times, however, it withdrew from the areas north of the Alps to the south and was completely replaced by the woolly rhinoceros in the areas that became free.

Because of its widespread distribution in warm-time Europe, the steppe rhinoceros coexisted with the forest rhinoceros. In areas where both animal species coexisted, the steppe rhinoceros took a back seat as its relative was obviously better able to assert itself. Exceptions are regions that the wood rhinoceros did not reach, such as England during the Eem warm period or the Iberian Peninsula in general. In these areas, the steppe rhinoceros probably inhabited less open landscapes and, due to a lack of competition, may also have penetrated forest areas .

Sites where the steppe rhinoceros appear are quite widespread in Europe, with around a dozen sites each, it was most frequently registered in Germany and Spain . Well known are the eemzeitlichen Observations of Neumark-Nord 1 from the former open pit Muecheln in Geiseltal ( Saxony-indication ), where a largely complete skeleton was found in addition to two skulls and 1988 days construction, which with some flint artifacts was connected to what a human manipulation of the carcass speaks. Also in the travertine of Ehringsdorf ( Thuringia ), the 200,000 year old site of the Ehringsdorf primitive man , numerous remains of the steppe rhinoceros have been handed down, including an almost completely preserved skull. In the fauna of the archaeological site of Bilzingsleben in northern Thuringia , which has been scientifically researched since 1969 , in which Homo erectus or Homo heidelbergensis has also been detected and which is around 350,000 years old, the steppe rhinoceros (together with the wood rhinoceros) occurs relatively frequently. The bones and mostly remains of teeth, however, are severely dismembered and rarely complete, which prompted the excavators of that time, with reference to other references, to interpret them as remains of the hunting prey of early humans. Other important sites in Germany are Steinheim an der Murr ( Baden-Württemberg ), known for the Steinheim prehistoric people , and Mosbach ( Wiesbaden , Hesse ). On the latter, one of the earliest records of the steppe rhinoceros in Europe, it is only represented by a few remains, since otherwise the Hundsheim rhinoceros dominates here, which will soon die out.

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.

From the steppe rhinoceros, as from the forest rhinoceros, only bones and tooth remnants are known, in some cases more or less complete surviving skeletons are available. On the basis of this, however, its appearance can be reconstructed more precisely with the aid of anatomical examinations and comparisons. The body was strongly built, but its limbs were much shorter and wider than the forest rhinoceros and had weak joints, which is an adaptation to life in open landscapes. In terms of body size was slightly smaller than also be fossil relative, the rhinoceros, which as the largest member of the genus Stephanorhinus valid and comparable to today's Indian rhinoceros is. In general, the steppe rhinoceros is described as quite slender or graceful for an animal of its size. The reconstructed live weight is given as around 1.4 tons to 3.4 tons.

The skull of the steppe rhinoceros, which was up to 76 cm long, was greatly elongated at the occiput , so that it carried the head diagonally downwards. Thereby it differs very markedly from the other species of the Stephanorhinus line, which kept the head much more erect, and is more similar to the woolly rhinoceros or today's white rhinoceros ( Ceratotherium simum ). The species had two horns , which were on the nasal bone and the middle skull. In the fossils found, the horns, made of keratin but not preserved, are marked by cauliflower-like overgrown attachment surfaces. The shape of these roughened surfaces shows that the anterior horn in particular must have been very large, which is also indicated by the nasal septum , which is more ossified than the forest rhinoceros .

Lower jaw fragment of the steppe rhinoceros

The lower jaw measured up to 56 cm in length and was solidly built, with the lower jaw body being up to 11 cm high. The bite was characterized as with all representatives of the Stephanorhinus -line by the absence of front teeth out, furthermore by the reduced number of Vorbackenzähne (three arch ) and the actual molars (also three). The resulting tooth formula reads: Both the second premolar and the third molar were particularly delicate in this species. An abnormal peculiarity that has been observed in isolated cases in the plains rhinoceros is the formation of a fourth molar, which is actually typical for marsupials . The teeth were generally quite high-crowned and had a thick layer of cement  - a feature that was most pronounced in this species of animal within the Stephanorhinus line - and thus, like the posture of the skull, speak for a mainly grass-eating diet. However, since the teeth are not always ground evenly horizontally, which can be observed in all grass-eating mammals due to the hard silica contained in the grass , but also occasionally trough-like depressions occur, it is assumed that the steppe rhinoceros also have softer food such as leaves , flowers or fruits has consumed.

The postcranial skeleton is far less known. The spine comprised at least 7 cervical, 18 thoracic and 4 lumbar vertebrae, the exact number of sacral and caudal vertebrae is not known. The humerus could be up to 46 cm long, the ulna 51 cm. The femur reached 50 cm in length and was relatively robust. Presumably the hands and feet, as in today's modern rhinos but also in fossil species of Stephanorhinus , were constructed with three rays, with the central ray (Metapodium III) being particularly pronounced. A largely complete foot from the Geiseltal had a metatarsal bone 18 cm long .

Little is known about the further appearance of this species, as there are no surviving soft tissues. No information can be given about the size and shape of the ears or the appearance of the skin . The assumption is speculative that this species had a fur , but the Sumatran rhinoceros as its closest living relative is hairy. Whether this also applies to the Pleistocene steppe rhinoceros cannot be said without further ado. Its occurrence in cooler climatic sections of the warm periods, which are also associated with cold winters, especially in northern Alpine Europe, could make this likely.

Systematics and tribal history

The genus Stephanorhinus next to the steppe rhino includes the simultaneously occurring with it rhinoceros ( Stephanorhinus kirchbergensis ) as well as the phylogenetically older types Etruscan rhinoceros ( Stephanorhinus etruskus ) which Hundsheimer Rhino ( Stephanorhinus hundsheimensis ) and others. Together with the genus Dicerorhinus and Coelodonta , Stephanorhinus belongs to the Dicerorhinina, a group of two-horned rhinos whose only representative still alive today is the Sumatran rhinoceros ( Dicerorhinus sumatrensis ), which is endangered in its population .

Little is known about the origin of the species. It is believed that it developed from a late form of the Etruscan rhinoceros in Asia and later immigrated to Europe. It first appeared in Europe between 450,000 and 500,000 years ago in the Middle Pleistocene. The oldest finds come from Wiesbaden (Hesse, Mosbach 3 ) and the Arago Cave ( France ). Older sites associated with the steppe rhinoceros, such as Süßenborn (Thuringia), are most likely confused with the Hundsheim rhinoceros or the Etruscan rhinoceros, both of which are phylogenetically older forms.

During its early appearance, the steppe rhinoceros was still relatively graceful and resembled the early representatives of the genus Stephanorhinus in terms of variation . Sites with such shapes include Bilzingsleben (Thuringia), Steinheim an der Murr (Baden-Württemberg) and Orgnac ( France ). It then increased in robustness, especially in the late Middle Pleistocene and New Pleistocene. Such large steppe rhinos have been found in Ehringsdorf (Thuringia), Geiseltal (Saxony-Anhalt) and Crayford (England). For this reason, two subspecies were introduced as early as the 1960s, Stephanorhinus hemitoechus falconeri comprising the older and Stephanorhinus hemitoechus aretinus the younger.

After the Eem warm period, the steppe rhinoceros, like many heat-adapted animals, retreated south to the Mediterranean region. Finds from this late period of this animal species, which then only occurred in isolated populations, come from the karst caves of Melpigano in the southern Italian municipality of Cursi and from the Pontine plain near Rome and are between 100,000 and 70,000 years old. It is not possible to say exactly when the plains rhinoceros finally became extinct. Originally it was assumed that it disappeared shortly after the beginning or in the first half of the Vistula Glaciation (115,000 to 11,700 years ago). More recent finds, including from La Ventana near Madrid (Spain), however, suggest that the steppe rhinoceros could have survived until the end of the Late Paleolithic ( Epipalaeolithic ) or even the beginning of the Holocene . ( see also main article: Quaternary extinction wave )

Research history

Hugh Falconer

The first description as Rhinoceros hemitoechus was made by Hugh Falconer in 1868. Its processing was based on finds from the Minchin Cave on the Gower Peninsula in Glamorgan (Wales). But already in 1859 Falconer had presented rhinoceros fossils from "Caves in Glamorganshire" (Wales) with the same name, which is also mentioned in several publications of the time. The finds from the "Caves in Glamorganshire" therefore represent the syntype and those from the Minchin Cave the lectotype of this species.

Even before Falconer, in 1846, the English naturalist Richard Owen (1804-1892) had introduced the name Rhinoceros leptorhinus for the steppe rhinoceros , but this is no longer valid today. For his description, he had used finds from Clacton-on-Sea (England), which, however, no longer belong to the type specimens. The name is also a homonym of the same name used by the French naturalist Georges Cuvier (1769-1832) as early as 1822.

The genus name Rhinoceros originally used was later replaced by Dicerorhinus . Due to numerous morphological differences to the Sumatran rhinoceros as the only recent representative, the Hungarian paleontologist Miklós Kretzoi (1907-2005) introduced the genus name Stephanorhinus , which is valid today . This genus includes most of the fossil Plio- and Pleistocene rhinos of northern Eurasia and is monophyletically related to the genus Coelodonta , which also includes the woolly rhinoceros, which also occurs in the Middle and Young Pleistocene.

In terms of research history, there has often been a great deal of confusion regarding the steppe rhinoceros and its related forest rhinoceros. In the course of time, the steppe rhinoceros came under various scientific names, some of which overlap with those of the forest rhinoceros:

  • Rhinoceros leptorhinus Cuvier, 1822
  • Rhinoceros megarhinus De Christol, 1834
  • Dicerorhinus mercki Kaup , 1841
  • Rhinoceros Mercki Kaup, 1841
  • Rhinoceros leptorhinus Owen, 1846
  • Rhinoceros lunellensis Gervais, 1848-1852
  • Rhinoceros hemitoechus Falconer, 1868
  • Rhinoceros subinermis Pomel, 1895
  • Rhinoceros hemitoechus aretinus Azzaroli, 1962
  • Rhinoceros hemitoechus falconeri Azzaroli, 1962.

literature

  • Wighart von Koenigswald: Living Ice Age. Climate and fauna in transition. Theiss-Verlag, Stuttgart 2002, ISBN 3-8062-1734-3 .

Web links

Commons : Stephanorhinus  - collection of images, videos and audio files

Individual evidence

  1. ↑ a b c d e f g H. Loose: Pleistocene Rhinocerotidae of W. Europe with reference to the recent two-horned species of Africa and SE Asia. Scripta Geologica 33, 1975, pp. 1-59
  2. ↑ Denise Geraads: Rhinocerotidae. In: L. Werdelin and DJ Sanders (eds.): Cenozoic Mammals of Africa. Berkeley, 2010, pp. 669-683
  3. ^ 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
  4. ↑ a b Wighart von Koenigswald: Lebendige Eiszeit. Climate and fauna in transition. Stuttgart, 2002, pp. 54-61
  5. ↑ Diana Pushkina: The Pleistocene easternmost distribution in Eurasia of the species associated with the Eemian Palaeoloxodon antiquus assemblage. Mammal Review 37, 2007, pp. 224-245
  6. ↑ a b 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. Quaternary Science Reviews 27, 2008, pp. 1951-1961
  7. ↑ Ralf-Dietrich Kahlke: The origins, development and distribution history of the upper-pleistozönen Mammuthus-Coelodonta fauna complex in Eurasia (large mammals). Treatises of the Senckenberg Natural Research Society 546 Frankfurt am Main, 1994
  8. ↑ a b c d e f 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
  9. ^ Dietrich Mania et al: Quaternary research in the Neumark-Nord opencast mine, Geiseltal (Saxony-Anhalt) and its results so far. In: Dietrich Mania et al. (Ed.): Neumark-Nord - An interglacial ecosystem of the Middle Palaeolithic people. Publications of the State Museum for Prehistory in Halle 62 Halle / Saale 2010, pp. 11–70
  10. ^ Walter Steiner: The travertine of Ehringsdorf and its fossils. Lutherstadt Wittenberg 1981
  11. ↑ Dietrich Mania: On the geology of the ancient Paleolithic find horizon of Bilzingsleben (Thuringia), taking into account the geological impact factor "man". Hercynia 37, 2004, pp. 143-184
  12. ↑ a b c Jan van der Made: A preliminary note on the rhinos from Bilzingsleben. Praehistoria Thuringica 4, 2000, pp. 41-64
  13. ↑ 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
  14. ↑ 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
  15. ↑ Frederic Lacombat: Phylogeny of the genus Stephanorhinus in the Pliocene-Pleistocene of Europe. Hallesches Jahrbuch für Geoswissenschaften 23, 2007, pp. 63–65
  16. ↑ 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 179, 2008, pp. 25-37
  17. ↑ Mikael Fortelius (coordinator). Neogene of the Old World Database of Fossil Mammals (NOW). University of Helsinki, 2003 http://www.helsinki.fi/science/now/
  18. ^ Andreas Wagner: Reconstruction of body masses of Pleistocene Rhinocerotidae in the Königswald collection. Frankfurt am Main, 2007
  19. ↑ Frederic Lacombat: Pleistocene rhinoceroses in Mediterranean Europe and Massif Central (France). Courier des Forschungs-Institut Senckenberg 256, 2006, pp. 57–69
  20. ↑ 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
  21. ↑ 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
  22. ↑ E. Cerdeño: Diversity and evolutionary trends of the family Rhinocerotidae Perissodactyla. Palaeo 141, 1997, pp. 13-34
  23. ↑ Ralf-Dietrich Kahlke, Nuria García, Dimitris S. Kostopoulos, Frédéric Lacombat, Adrian M. Lister, Paul PA Mazza, Nikolai Spassov and, Vadim V. Titov: Western Palaearctic palaeoenvironmental conditions during the Early and early Middle Pleistocene inferred from large mammal communities, and implications for hominin dispersal in Europe. Quaternary Science Reviews, 2010, pp. 1-28
  24. ↑ A. Azzaroli: Validità della specie Rhinoceros hemitoechus Falconer. Palaeontographia Italica 57, 1962, pp. 21-34
  25. ^ A b Carmelo Petronio and Luca Pandolfi: Stephanorhinus hemitoechus (FALCONER 1868) del Pleistocene Superiore dell'Area the Melpigano-Cursi E S. (Lecce, Italia). Geologica Romana 41, 2008, pp. 1-12
  26. ↑ Simone Farina: Late Pleistocene-Holocene mammals from “Canale delle Acque Alte (Canale Mussolini)” (Agro Pontino, Latium). Bollettino della Società Paleontologica Italiana, 50 (1), 2011, pp. 11-22
  27. ↑ 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
  28. ↑ A. Sánchez, S. Fraile, Jan van der Made, J. Morales, V. Quiralte, MJ Salesa, IM Sánchez, B. Sanchiz, D. Soria, J. Jiménez, LJ Barbadillo, C. Laplana, Z. Szyndlar : Primeros datos faunísticos del Neolítico madrileño: la cueva de la Ventana (Torrelaguna, Madrid). In: P. Arias Cabal, R. Ontañón Peredo, C. García-Moncó Piñeiro (eds.) III Congreso del Neolítico en la Península Ibérica. Monografias del Instituto Internacional de Investigaciones Prehistóricas de Cantabria, 1, 2005, pp. 155-165
  29. ^ Hugh Falconer: On the European Pliocene and Pleistocene species of the genus Rhinoceros. In: C. Murchison (Ed.): Palaeontological memories and notes of the late Hugh Falconer. Volume 2. London, 1868, pp. 309-403
  30. ↑ Miklós Kretzoi: Comments on the system of post-Miocene rhinoceros genera. Földtany Közlöny 72, 1942, pp. 309-323