Stephanorhinus

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Stephanorhinus
Temporal range: Pliocene to Late Pleistocene 5–0.04 Ma
Stephanorhinus etruscus skeleton 23.jpg
Stephanorhinus etruscus skeleton
Abhandlungen der Geologischen Bundesanstalt (1902) (16144915223).jpg
Stephanorhinus hundsheimensis skeleton
Scientific classification e
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Perissodactyla
Family: Rhinocerotidae
Tribe: Dicerorhinini
Genus: Stephanorhinus
Kretzoi, 1942
Type species
Rhinoceros etruscus
Falconer, 1868
Species
  • S. etruscus (Falconer, 1868) Etruscan rhinoceros
  • S. hemitoechus (Falconer, 1859) Narrow-nosed rhinoceros
  • S. hundsheimensis (Toula, 1902) Hundsheim rhinoceros
  • S. jeanvireti (Falconer, 1859)
  • S. kirchbergensis (Jäger, 1839) Merck's rhinoceros
  • S. lantianensis (Hu and Qi, 1978)
  • S. yunchuchenensis (Chow, 1963)

Stephanorhinus is an extinct genus of two-horned rhinoceros native to Eurasia and possibly North Africa that lived during the Pliocene to Late Pleistocene. Species of Stephanorhinus were the predominant and often only species of rhinoceros in much of temperate Eurasia, especially Europe, for most of the Pleistocene. Two species of StephanorhinusMerck's rhinoceros (S. kirchbergensis) and the narrow-nosed rhinoceros (S. hemitoechus) – persisted into the last glacial period.

Etymology[edit]

The first part of the name, Stephano-, honours Stephen I, the first king of Hungary.[1] (The genus name was coined by Kretzoi, a Hungarian.) The second part is from rhinos (Greek for "nose"), a typical suffix of rhinoceros genus names.

Taxonomy[edit]

The taxonomic history of Stephanorhinus is long and convoluted, as many species are known by numerous synonyms and different genera – typically Rhinoceros and Dicerorhinus – for the 19th and most of the early 20th century. The genus was named by Miklós Kretzoi in 1942.[2] It is thought that Stephanorhinus is more closely related to the Sumatran rhinoceros and woolly rhinoceros than other rhino species. A complete mitochondrial genome of S. kirchbergensis obtained from a 70,000–48,000-year-old skull preserved in permafrost in arctic Yakutia showed that it was more closely related to the woolly rhinoceros than the Sumatran rhinoceros, with the tree species forming a clade to the exclusion of other living rhinoceros species.[3] In 2019 a study of dental proteomes proposed that Stephanorhinius was paraphyletic as currently defined, with the proteome sequence obtained from the enamel of a 1.77 million year old Stephanorhinus tooth from Dmanisi belonging to an indeterminate species found outside the clade containing the woolly rhinoceros and S. kirchbergensis, suggesting that the genus Coelodonta was derived from an early diverging lineage within Stephanorhinus.[4] A 2021 study based on nuclear genomes including those of S. kirchbergensis found the same result as the mitochondrial genome study, with strong support, with the estimated split between the woolly rhinoceros and S. kirchbergensis occurring around 5.5 million years ago.[5]

Species and evolution[edit]

The oldest known species of the genus are from the Pliocene of Europe, the species S. pikermiensis and S. megarhinus that were formerly considered to belong to Stephanorhinus are currently considered to belong to Dihoplus,[6] while the positions of “Stephanorhinusmiguelcrusafonti from the Early Pliocene of Western Europe and Stephanorhinus? africanus from the Middle Pliocene of Tunisia and Chad are uncertain.[7]

Skull of Stephanorhinus jeanvireti

Stephanorhinus jeanvireti, also known as S. elatus[8] is known from the Late Pliocene and Early Pleistocene of Europe. Its remains are relatively rare in comparison to other Stephanorhinus species. Specimens are known from the Late Pliocene of Germany,[9] France, Italy,[10] Slovakia[11] and Greece,[12] and the Early Pleistocene of Romania.[13]

Skull of Stephanorhinus etruscus

Stephanorhinus etruscus first appears in the latest Pliocene in the Iberian Peninsula, around 3.3 million years ago (Ma) at Las Higueruelas in Spain and before 3 Ma at Piedrabuena, and during the latest Pliocene at Villafranca d’Asti and Castelnuovo di Berardenga in Italy and is abundant during most of the Villafranchian period in Europe, and is the sole rhinoceros species in Europe between 2.5 and around 1.3 Ma. A specimen is known from the Early Pleistocene (1.6-1.2 Ma) Ubeidiya locality in Israel. During the late Early Pleistocene, it is largely replaced by S. hundsheimensis. The last known records of the species are from the latest Early Pleistocene of the Iberian peninsula, around 0.9-0.8 Ma.[14]

Stephanorhinus yunchuchenensis is known from a single specimen in Early Pleistocene deposits in Yushe, Shaanxi, while Stephanorhinus lantianensis is also known from a single specimen from late Early Pleistocene (1.15 Ma) deposits in Lantian, also in Shaanxi.[15] These are stated to be synonyms of Stephanorhinus kirchbergensis by some sources.[16]

The first record of Stephanorhinus kirchbergensis is in China at Zhoukoudian (Choukoutien; near Beijing), around the Early–Mid-Pleistocene transition at 0.8 Ma.[15]

Skull of Stephanorhinus hundsheimensis

Stephanorhinus hundsheimensis first definitively appears in the fossil record in Europe and Anatolia at around 1.2 Ma, with possible records in Iberia around 1.6 Ma and 1.4-1.3 Ma. The earliest confirmed appearance in Italy around 1 Ma.[17] The diet of S. hundsheimensis was flexible and ungeneralised, with two different early Middle Pleistocene populations under different climatic regimes (having tooth wear analyses suggesting contrasting browsing and grazing habits).[18] The more specialised S. kirchbergensis and S. hemitoechus, appear in Europe between 0.7-6 Ma and 0.6-0.5 Ma respectively, and replace S. hundsheimensis. S. kirchbergensis and S. hemitoechus are typically interpreted as a browsing form and grazing form, respectively. The evolution of more specialized diets is possibly due to the change to the 100 Kyr cycle after the Mid-Pleistocene Transition, which resulted in environmental stability allowing the development of more specialized forms.[19]

Skulls from top to bottom. S. kirchbergensis, S. hemitoechus and the woolly rhinoceros, showing the difference in head angle

From the late Middle Pleistocene onwards, the large Merck's rhinoceros (Stephanorhinus kirchbergensis) and the narrow-nosed rhinoceros (Stephanorhinus hemitoechus) were the only species of Stephanorhinus. S. kirchbergensis was broadly distributed over northern Eurasia, while S. hemitoechus was generally confined to the western Palearctic.[20] S. kirchbergensis is relatively rare in fossil record and known from few Italian, French, German, British, and East-European localities, mostly of the middle Pleistocene. In Asia it is known from Siberia, Central Asia (e.g. southeastern Kazakhstan), central Korea, and China. It may have also occurred in Israel and Lebanon, but here it is unclear if it was really S. kirchbergensis or a similar species.[21] In eastern China, S. kirchbergensis was present throughout the Pleistocene between 30°N and 40°N.[16] It is stated by some sources that S. hemitoechus occurs in North Africa.[7]

In eastern Europe, S. kirchbergensis disappeared during the earliest Late Pleistocene. Almost complete remains (more than 100 bones - 85%) was discovered in Eemian lake sediments in the Gorzów Plain, near Gorzów Wielkopolski (NW Poland).[22] In the forests of the Caucasus, it may have survived even until the early Weichselian. Apart from Europe, it is also known from Syria, Israel, the Caucasus and from one Late Pleistocene locality close to Lake Baikal. The latest fossils of the narrow-nosed rhinoceros are known from the Balkans, where it survived until the early late Weichselian.[20] The last records of S. hemitoechus in Italy date to around 41 kya.[23] A late record of S. hemitoechus is known from 40,000 years ago in Bacho Kiro cave in Bulgaria.[24] Remains of S. kirchbergensis in the Russian Far East and South China are suggested to date to marine isotope stage 3 (~60-27,000 years ago) and 2 (~29-14,000 years ago), respectively.[25][26]

References[edit]

  1. ^ Tong, HaoWen; Wu, XianZhu (April 2010). "Stephanorhinus kirchbergensis (Rhinocerotidae, Mammalia) from the Rhino Cave in Shennongjia, Hubei". Chinese Science Bulletin. 55 (12): 1157–1168. Bibcode:2010ChSBu..55.1157T. doi:10.1007/s11434-010-0050-5. ISSN 1001-6538.
  2. ^ Miklós Kretzoi: Bemerkungen zur System der Nachmiozänen Nashorn-Gattungen (Comments on the system of the post Miocene rhinoceros genera) Földtani Közlöni, Budapest 72 (4-12), 1942, S. 309–318
  3. ^ Kirillova, Irina V.; Chernova, Olga F.; van der Made, Jan; Kukarskih, Vladimir V.; Shapiro, Beth; van der Plicht, Johannes; Shidlovskiy, Fedor K.; Heintzman, Peter D.; van Kolfschoten, Thijs; Zanina, Oksana G. (November 2017). "Discovery of the skull of Stephanorhinus kirchbergensis (Jäger, 1839) above the Arctic Circle". Quaternary Research. 88 (3): 537–550. Bibcode:2017QuRes..88..537K. doi:10.1017/qua.2017.53. ISSN 0033-5894.
  4. ^ Cappellini, Enrico; Welker, Frido; Pandolfi, Luca; Ramos-Madrigal, Jazmín; Samodova, Diana; Rüther, Patrick L.; Fotakis, Anna K.; Lyon, David; Moreno-Mayar, J. Víctor; Bukhsianidze, Maia; et al. (October 2019). "Early Pleistocene enamel proteome from Dmanisi resolves Stephanorhinus phylogeny". Nature. 574 (7776): 103–107. Bibcode:2019Natur.574..103C. doi:10.1038/s41586-019-1555-y. ISSN 0028-0836. PMC 6894936. PMID 31511700.
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  9. ^ Lacombat, Frédéric; Mörs, Thomas (2008-08-01). "The northernmost occurrence of the rare Late Pliocene rhinoceros Stephanorhinus jeanvireti (Mammalia, Perissodactyla)". Neues Jahrbuch für Geologie und Paläontologie - Abhandlungen. 249 (2): 157–165. doi:10.1127/0077-7749/2008/0249-0157.
  10. ^ Pandolfi, Luca (October 2013). "New and revised occurrences of Dihoplus megarhinus (Mammalia, Rhinocerotidae) in the Pliocene of Italy". Swiss Journal of Palaeontology. 132 (2): 239–255. doi:10.1007/s13358-013-0056-0. ISSN 1664-2376.
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  14. ^ Pandolfi, Luca; Cerdeño, Esperanza; Codrea, Vlad; Kotsakis, Tassos (September 2017). "Biogeography and chronology of the Eurasian extinct rhinoceros Stephanorhinus etruscus (Mammalia, Rhinocerotidae)". Comptes Rendus Palevol. 16 (7): 762–773. doi:10.1016/j.crpv.2017.06.004.
  15. ^ a b Tong, Hao-wen (November 2012). "Evolution of the non-Coelodonta dicerorhine lineage in China". Comptes Rendus Palevol. 11 (8): 555–562. doi:10.1016/j.crpv.2012.06.002.
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  17. ^ Pandolfi, Luca; Erten, Hüseyin (January 2017). "Stephanorhinus hundsheimensis (Mammalia, Rhinocerotidae) from the late early Pleistocene deposits of the Denizli Basin (Anatolia, Turkey)". Geobios. 50 (1): 65–73. doi:10.1016/j.geobios.2016.10.002.
  18. ^ Kahlke, Ralf-Dietrich; Kaiser, Thomas M. (August 2011). "Generalism as a subsistence strategy: advantages and limitations of the highly flexible feeding traits of Pleistocene Stephanorhinus hundsheimensis (Rhinocerotidae, Mammalia)". Quaternary Science Reviews. 30 (17–18): 2250–2261. Bibcode:2011QSRv...30.2250K. doi:10.1016/j.quascirev.2009.12.012.
  19. ^ van Asperen, Eline N.; Kahlke, Ralf-Dietrich (January 2015). "Dietary variation and overlap in Central and Northwest European Stephanorhinus kirchbergensis and S. hemitoechus (Rhinocerotidae, Mammalia) influenced by habitat diversity". Quaternary Science Reviews. 107: 47–61. doi:10.1016/j.quascirev.2014.10.001.
  20. ^ a b Diana Pushkina: The Pleistocene easternmost distribution in Eurasia of the species associated with the Eemian Palaeoloxodon antiquus assemblage. Mammal Review, 2007. Volume 37 Issue 3, Pages 224 - 245
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  22. ^ .Remains of rhinoceros (Stephanorhinus sp.) and fallow deer (Dama dama) discovered in Eemian lake sediments in the Gorzów Plain (NW Poland)
  23. ^ PANDOLFI, LUCA; BOSCATO, PAOLO; CREZZINI, JACOPO; GATTA, MAURIZIO; MORONI, ADRIANA; ROLFO, MARIO; TAGLIACOZZO, ANTONIO (2017-04-13). "LATE PLEISTOCENE LAST OCCURRENCES OF THE NARROW-NOSED RHINOCEROS STEPHANORHINUS HEMITOECHUS (MAMMALIA, PERISSODACTYLA) IN ITALY". Rivista Italiana di Paleontologia e Stratigrafia (Research In Paleontology and Stratigraphy). V. 123: N. 2 (2017). doi:10.13130/2039-4942/8300.
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  25. ^ Kosintsev, P. A.; Zykov, S. V.; Tiunov, M. P.; Shpansky, A. V.; Gasilin, V. V.; Gimranov, D. O.; Devjashin, M. M. (March 2020). "The First Find of Merck's Rhinoceros (Mammalia, Perissodactyla, Rhinocerotidae, Stephanorhinus kirchbergensis Jäger, 1839) Remains in the Russian Far East". Doklady Biological Sciences. 491 (1): 47–49. doi:10.1134/S0012496620010032. ISSN 0012-4966.
  26. ^ Pang, Libo; Chen, Shaokun; Huang, Wanbo; Wu, Yan; Wei, Guangbiao (April 2017). "Paleoenvironmental and chronological analysis of the mammalian fauna from Migong Cave in the Three Gorges Area, China". Quaternary International. 434: 25–31. doi:10.1016/j.quaint.2014.11.039.