Stephanorhinus

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Stephanorhinus
Skull of the steppe rhinoceros (Stephanorhinus hemitoechus)

Skull of the steppe rhinoceros ( Stephanorhinus hemitoechus )

Temporal occurrence
Upper Miocene to Upper Pleistocene
8.2 million years to 10,000 years
Locations

northern Eurasia

Systematics
Higher mammals (Eutheria)
Laurasiatheria
Unpaired ungulate (Perissodactyla)
Rhinocerotoidea
Rhinoceros (Rhinocerotidae)
Stephanorhinus
Scientific name
Stephanorhinus
Kretzoi , 1942

Stephanorhinus is a now extinct genus from the rhinoceros family. It was distributedmainly in northern Eurasia from the Upper Miocene to the end of the Pleistocene 8 million to around 10,000 years ago. Is related Stephanorhinus with Dicerorhinus whose only living representatives today in Southeast Asia living and highly endangered Sumatran rhinoceros ( Dicerorhinus sumatrensis is). Both genera belong to the Dicerorhinina, rhinos with two horns that were predominantly found in Eurasia. The genus also appeared occasionally in the northernmost part of Africa .

features

The representatives of the genus Stephanorhinus were medium-sized to very large rhinos that occupied various ecological niches and therefore had a variable appearance. In general, they were quite strong, reminiscent of today's rhinos. Forest dwellers were somewhat more graceful with long, slender limbs, while inhabitants of the forest edge areas or open landscapes had a stronger physique and rather short and wide limbs. The size of the individual species varied considerably and, in the case of small representatives such as the Hundsheimer rhinoceros ( Stephanorhinus hundsheimensis ) and the Etruscan rhinoceros ( Stephanorhinus etruscus ), are 700 to 900 kg, while the largest forms, such as the steppe rhinoceros ( Stephanorhinus hemitoechus ) and the forest rhinoceros ( Stephanorhinus) kirchbergensis ), up to 3 t live weight. However, some species went through a marked increase in size during their development. In the postcranial skeleton structure, Stephanorhinus resembled today's rhinos. The most important feature are the three-pronged hands and feet, which the rhinoceros genus represents to the modern Rhinocerotini.

The skull varied in length from 56 to 81 cm, depending on the size of the species, and was very large and massive with protruding cheekbones . The occipital bone was like pulled in most rhinos back, but had mostly a right angle on the Hinterhauptswulst to the joint surfaces on which exaggerated the cervical spine. This resulted in a high to horizontal head position. The only exception here is the steppe rhinoceros, which had an acute angle and thus had a low or deep head position. The nasal bone was clearly rounded and had cauliflower-like roughened surfaces as a point of attachment for the anterior horn. A similar surface structure on the frontal bone , which was mostly less clear, marked the rear second horn. A common feature of all Stephanorhinus species was the partially ossified nasal septum , which in the majority of species grew together in the front third, in the steppe rhinoceros up to two thirds. Only the genus Coelodonta with the woolly rhinoceros ( Coelodonta antiquitatis ), whose nasal septum was completely ossified, possessed a similar characteristic within the modern rhinos . In contrast to this, the nasal bone of Stephanorhinus did not sit directly on the intermaxillary bone .

The dentition of Stephanorhinus is significantly reduced, so that the anterior teeth are usually completely missing. The dental formula of adult animals was: . The premolars were largely similar to the molars , so they were clearly molarized. Since the majority of the early species had low tooth crowns ( brachyodont ), later these were slightly raised. Only the steppe rhinoceros developed very high tooth crowns ( hypsodont ) with a high proportion of tooth cement.

distribution

Stephanorhinus was particularly widespread in northern Eurasia and was partially adapted to cooler climates than Dicerorhinus . It occurred from the Iberian Peninsula in the west across all of Europe , the Russian steppe landscapes to East Asia in the east. However, individual species remained restricted to western or eastern Eurasia, only the forest rhinoceros had a Pan-Eurasian distribution, while the steppe rhinoceros has been detected as far as Lake Baikal . The southernmost limit in the western distribution area is North Africa , where the genus did not appear until the Young Pleistocene . In East Asia representatives of Stephanorhinus penetrated up to the 30th northern parallel, more southern areas were then inhabited by the genera Rhinoceros and Dicerorhinus . A skull find from Yakutia suggests that the genus also partially penetrated into the arctic regions of Asia, where the woolly rhinoceros is more likely to be detected. Remains of Staphanorhinus have also been reported from some sites in Japan , but no precise species assignment has been made so far.

Paleobiology

The predominant species, especially in the late Miocene , in the Pliocene and in the early Pleistocene, were adapted to closed forests due to their more delicate physique with slender limbs, later forms developed that also preferred more open landscapes. The steppe rhinoceros from the late Middle Pleistocene was the only species that also colonized open steppe areas and accordingly had stronger and short legs. The mostly low-crowned teeth and the high head posture advocate a diet based on soft plant-based food ( browsing ), which can also be demonstrated by the traces of abrasion on the chewing surfaces of the teeth with trough-like depressions. The Hundsheim rhinoceros preferred a mixed diet. In the forest rhinoceros, in turn, food residues are passed down in the teeth that come from birch , rose , poplar , oak , hawthorn , firethorn and water lily . The steppe rhinoceros, on the other hand, has very high tooth crowns and a large proportion of dental cement . Characteristic horizontal sanding patterns on the chewing surfaces speak for a specialized, hard and siliceous grass food ( grazing ) that had to be picked up from the ground. In evolution, this led to the lengthening of the occiput and a permanently low head posture, as is still the case today with the white rhinoceros ( Ceratotherium simum ).

Stephanorhinus typically had two horns, of which the anterior (nasal or nasal horn) was attached to the nasal bone and the posterior (frontal or frontal lobe) to the frontal bone, as can be seen on the roughened bone surfaces, the keratin horn itself but not fossilized. Such so-called tandem horns are still found today in the Sumatran rhinoceros and the more distantly related African rhinos. The extent of the roughened surfaces suggests the size of the former horn; in general, the nasal horns were larger than the frontal horns. Early Stephanorhinus representatives tended to have small horns, while later ones had larger ones. Originally, the overgrown nasal septum was associated with the increase in the size of the horns, today's African rhinos sometimes have very long horns, but do not show any ossification of the nasal septum. In addition to the Stephanorhinus - Coelodonta line, the fossil record is the more primitive Elasmotherium , which does not belong to the modern rhinos, and which also had such ossification. However, this had no nasal horn at all, only a forehead horn possibly up to two meters long.

Systematics

Closer relationship of Stephanorhinus according to Welker et al. 2017
 Rhinocerotini  
  Dicerotina  

 Ceratotherium (white rhinoceros)


   

 Diceros (black rhinoceros)



   
  Dicerorhinina  

 Coelodonta (woolly rhinoceros and others (†))


   

 Dicerorhinus (Sumatran rhinoceros)


   

 Stephanorhinus (†)


Template: Klade / Maintenance / 3

  Rhinocerotina  

 Rhinoceros (armored and Java rhinoceros)




Template: Klade / Maintenance / Style

In today's system of rhinos is Stephanorhinus to subtribe Dicerorhinina provided, which in turn to the tribes of Rhinocerotini is provided, to which all extant rhinos. Closely related species are Coelodonta with the woolly rhinoceros , as well as the phylogenetically older forms Dihoplus and Lartetotherium . The next representative still alive today is the Sumatran rhinoceros ( Dicerorhinus sumatrensis ), from whose generic name the name of the sub-tribus is derived. The close relationship of Stephanorhinus with both Coelodonta and Dicerorhinus could be confirmed with the help of protein sequence studies on Middle Pleistocene finds from Schöningen and Neumark-Nord in the Geiseltal . Molecular genetic analyzes on a Young Pleistocene skull find from Yakutia revealed a sister group relationship between Stephanorhinus and Coelodonta . Another study of protein sequences on an approximately 1.8 million year old fossil find from Dmanissi in Georgia showed that Stephanorhinus and Coleodonta probably form a common group. Both genera are descended from a common ancestor. Representatives of the Stephanorhinus line have been known at least since the end of the Miocene in western Eurasia , while Coelodonta was formed in eastern Eurasia in the Pliocene . As a consequence of this, Stephanorhinus is most likely to be understood as paraphyletic and would have to be resolved in Coelodonta . The synonymity of the two genera was previously considered based on the same anatomical features, but a direct combination has so far been rejected.

Closer relationship of Stephanorhinus after 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

All anatomical, genetic and biochemical investigations carried out to date identify the Sumatran rhinoceros as the closest living relative of Stephanorhinus and Coelodonta . Its direct phylogenetic ancestor is still unknown today. Due to the preservation of the anterior dentition, however, it is a rather primitive member in terms of phylogenetic history. Dicerorhinus split off from the Asian rhinoceros line 26 million years ago, and Coelodonta separated from the Dicerorhinus line 21 million years ago .

Numerous species were assigned to the genus Stephanorhinus . the following are recognized today:

The name Stephanorhinus was introduced in 1942 by the Hungarian paleontologist Miklós Kretzoi . The name is made up of the first name “ Stefan ” and the Greek word ῥίς ( rhīs “nose”; genitive rhinos ). "Stephan" is a tribute to the first king of Hungary and today's national saint of the country Stephan I ( István ). However, at that time Kretzoi did not include the steppe rhinoceros ( Stephanorhinus hemitoechus ) due to the differently built skull, but referred it to that of Genus Procerorhinus created independently for him . Initially, the name Stephanorhinus hardly caught on. The French researcher Claude Guérin did not recognize the name, instead he created his own in 1980 with Brandtorhinus , in which he included all European Plio- and Pleistocene Dicerorhinina forms (with the exception of Coelodonta ); however, Guérin took Brandtorhinus only as a subgenus to Dicerorhinus . In 1993 Mikael Fortelius published an extensive revision of the Western Eurasian Stephanorhinus representatives and thus helped the name to gain recognition. Today the name Stephanorhinus is widely accepted. In 2012 the East Eurasian species were revised.

Tribal history

The genus Stephanorhinus first appeared in the Upper Miocene around 8 million years ago, the oldest species is Stephanorhinus pikermiensis . Earlier finds come mainly from Greece , from Pikermi and the island of Samos . Especially in the Pliocene and Pleistocene , numerous species emerged. Basically, two lines of development can be traced in western Eurasia: One ran via Stephanorhinus megarhinus to Stephanorhinus kirchbergensis . The other included Stephanorhinus etruscus , from which Stephanorhinus jeanvireti , Stephanorhinus hundsheimensis and finally Stephanorhinus hemitoechus descended, but the process has not yet been clarified in detail. Especially in the Middle and Young Pleistocene, with their strongly fluctuating climatic phases, two highly specialized species developed, on the one hand the forest rhinoceros and on the other hand the steppe rhinoceros. While most species had already disappeared in the early Middle Pleistocene, the wood rhinoceros survived until the beginning of the Late Pistocene, while the steppe rhinoceros may not become extinct until the beginning of the Holocene . A very late record comes here from La Ventana in Spain .

Individual evidence

  1. ↑ a b Frederic Lacombat 2007: Phylogeny of the genus Stephanorhinus in the Plio-Pleistocene of Europe. Hallesches Jahrbuch für Geoswissenschaften 23, 63–65
  2. ^ Jean-Philip Brugal and Roman Croitor: Evolution, ecology and biochronology of herbivore associations in Europe during the last 3 million years. Quaternaire, 18 (2), 2007, pp. 129-152
  3. ↑ a b c d 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
  4. ↑ Frederic Lacombat: Rhinoceroses in Mediterranean Europe and Massif Central (France). Courier des Forschungs-Institut Senckenberg 256, 2006, pp. 57–69
  5. ^ M. Breda, SE Collinge, Simon A. Parfitt, and Adrian M. Lister: Metric analysis of ungulate mammals in the early Middle Pleistocene of Britain, in relation to taxonomy and biostratigraphy. I: Rhinocerotidae and Bovidae. Quaternary International 228, 2010, pp. 136-156
  6. ↑ Friedrich E. Zeuner: The relationships between skull shape and way of life in recent and fossil rhinos. Reports of the Natural Research Society in Freiburg 34, 1934, pp. 21–80
  7. ↑ 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
  8. ↑ a b c d e 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. ^ Emmanuel ME Billia: Occurrences of Stephanorhinus kirchbergensis (Jäger, 1839) (Mammalia, Rhinocerotidae) in Eurasia - An account. Acta Palaeontologica Romaniae 7, 2011, pp. 17-40
  10. ↑ Diana Pushkina: The Pleistocene easternmost distribution in Eurasia of the species associated with the Eemian palaeoloxodon antiquus assemblage. Mammal Review, 37 (3), 2007, pp. 224-245
  11. ↑ Denis Geraads: Rhinocerotidae. In: L. Werdelin and DJ Sanders (eds.): Cenozoic Mammals of Africa. Berkeley, 2010, pp. 669-683
  12. ↑ Pierre Olivier Antoine: Pleistocene and holocene rhinocerotids (Mammalia, Perissodactyla) from the Indochinese Peninsula. Comptes Rendus Palevol 2011, pp. 1-10
  13. ^ 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
  14. ↑ a b 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 88, 2017, pp. 537-550, doi: 10.1017 / qua.2017.53
  15. ↑ Naoto Handa, Naoki Kohno and Yuichiro Kudo: Reappraisal of a middle Pleistocene rhinocerotid (Mammalia, Perissodactyla) from the Matsugae Cave, Fukuoka Prefecture, southwestern Japan. Historical Biology, 2019, doi: 10.1080 / 08912963.2019.1604699
  16. ↑ Fredéric Lacombat and Thomas Mörs: The northernmost of the Late Pliocene rhinoceros Stephanorhinus jeanvireti (Mammalia, Perissodactyla). Neuer's Jahrbuch zur Geologie und Paläontologie, Abhandlungen 249 (2), 2008, pp. 157–165
  17. ↑ Ralf-Dietrich Kahlke and Thomas M. Kaiser: Generalism as a subsistence strategy: advantages and limitations of the highly flexible feeding traits of Pleistocene Stephanorhinus hundsheimensis (Rhinocerotidae, Mammalia). Quaternary Science Reviews 30, 2011, pp. 2250-2261 doi : 10.1016 / j.quascirev.2009.12.012 .
  18. ↑ HK 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
  19. ↑ a b Frido Welker, Geoff M. Smith, Jarod M. Hutson, Lutz Kindler, Alejandro Garcia-Moreno, Aritza Villaluenga, Elaine Turner and Sabine Gaudzinski-Windheuser: Middle Pleistocene protein sequences from the rhinoceros genus Stephanorhinus and the phylogeny of extant and extinct Middle / Late Pleistocene Rhinocerotidae. PeerJ 5, 2017, p. E3033, doi: 10.7717 / peerj.3033
  20. ↑ a b Enrico Cappellini, Frido Welker, Luca Pandolfi, Jazmín Ramos-Madrigal, Diana Samodova, Patrick L. Rüther, Anna K. Fotakis, David Lyon, J. Víctor Moreno-Mayar, Maia Bukhsianidze, Rosa Rakownikow Jersie-Christensen, Meaghan Mackie, Aurélien Ginolhac, Reid Ferring, Martha Tappen, Eleftheria Palkopoulou, Marc R. Dickinson, Thomas W. Stafford Jr., Yvonne L. Chan, Anders Götherström, Senthilvel KSS Nathan, Peter D. Heintzman, Joshua D. Kapp, Irina Kirillova , Yoshan Moodley, Jordi Agusti, Ralf-Dietrich Kahlke, Gocha Kiladze, Bienvenido Martínez-Navarro, Shanlin Liu, Marcela Sandoval Velasco, Mikkel-Holger S. Sinding, Christian D. Kelstrup, Morten E. Allentoft, Ludovic Orlando, Kirsty Penkman, Beth Shapiro, Lorenzo Rook, Love Dalén, M. Thomas P. Gilbert, Jesper V. Olsen, David Lordkipanidze and Eske Willerslev: Early Pleistocene enamel proteome from Dmanisi resolves Stephanorhinus phylogeny. Nature 574, 2019, pp. 103-107, doi: 10.1038 / s41586-019-1555-y
  21. ↑ a b Akira Fukuchi, Hideo Nakaya, Masanaru Takai and Shintaro Ogino: A preliminary report on the Pliocene rhinoceros from Udunga, Transbaikalia, Russia. Asian Paleoprimatology 5, 2009, pp. 61-98
  22. ↑ Christelle Tougard, Thomas Delefosse, Catherine Hänni and Claudine Montgelard: Phylogenetic Relationships of the Five Extant Rhinoceros Species (Rhinocerotidae, Perissodactyla) Based on Mitochondrial Cytochrome b and 12S rRNA Genes. Molecular Phylogenetics and Evolution 19, 2001, pp. 34-44
  23. ^ Ludovic Orlando, Jennifer A. Leonard, Aurélie Thenot, Vincent Laudet, Claude Guerin, and Catherine Hänni: Ancient DNA analysis reveals woolly rhino evolutionary relationships. Molecular Phylogenetics and Evolution 28, 2003, pp. 485-499
  24. ↑ Eske Willerslev, M Thomas P. Gilbert, Jonas Binladen, Simon YW Ho, Paula F. Campos, Aakrosh Ratan, Lynn P. Tomsho, Rute R. da Fonseca, Andrei Sher, Tatanya V. Kuznetsova, Malgosia Nowak-Kemp, Terri L. Roth, Webb Miller and Stephan C Schuster: Analysis of complete mitochondrial genomes from extinct and extant rhinoceroses reveals lack of phylogenetic resolution. BMC Evolutionary Biology 9, 2009, p. 95, doi: 10.1186 / 1471-2148-9-95
  25. ↑ a b Tong Haowen: Evolution of the non-Coelodonta dicerorhine lineage in China. Comptes Rendus Palevol 11 (8), 2012, pp. 1-8, doi: 10.1016 / j.crpv.2012.06.002 .
  26. ↑ Miklós Kretzoi: Comments on the system of the post-Miocene rhinoceros genera. Földtani Közlöni, Budapest 72 (4-12), 1942, pp. 309-318
  27. ↑ Claude Guérin: Les Rhinocerotidae (Mammalia, Perissodactyla) du miocene terminal au pleistocene superieur d'Europe occidentale compares aux especes actuelles: tendences evolutives et relations phylogenetiques. Geobios 15 (4), 1982, pp. 599-605
  28. ↑ Kurt Heissig: Family Rhinocerotidae. In: Gertrud E. Rössner and Kurt Heissig: The Miocene land mammals of Europe. Munich, 1999, pp. 175-188

Web links

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