Stephanorhinus

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Stephanorhinus
Temporal range: Late Pliocene to Late Pleistocene 3.4–0.04  Ma
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Stephanorhinus etruscus skeleton 23 (cropped).jpg
Stephanorhinus etruscus skeleton
Abhandlungen der Geologischen Bundesanstalt (1902) (16144915223).jpg
Stephanorhinus hundsheimensis skeleton
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
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 North Africa that lived during the Late 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. The last two species of Stephanorhinus – Merck's rhinoceros (S. kirchbergensis) and the narrow-nosed rhinoceros (S. hemitoechus) – went extinct during the last glacial period.

Contents

Etymology

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

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. [2] The genus was named by Miklós Kretzoi in 1942. [3] Genomes obtained from Stephanorhinus kirchbergensis suggests that Stephanorhinus is more closely related to Dicerorhinus (which contains the living Sumatran rhinoceros) and Coelodonta (which contains the woolly rhinoceros), than it is to other living rhinoceroses, and is more closely related to Coelodonta than to Dicerorhinus, with the date of divergence between Coelodonta and Stephanorhinus estimated at around 5.5 million years ago, with the estimated split between their last common ancestor and Dicerorhinus estimated at around 9.4 million years ago. [4] The genus is also closely related to the fossil rhinoceros genera Dihoplus and Pliorhinus , known from the Late Miocene and Pliocene of Eurasia, which may be ancestral to Stephanorhinus. [5] Although a study based on dental proteomes suggested that the genus was paraphyletic with respect to Coelodonta, [6] a 2023 morphological study recovered Stephanorhinus as monophyletic. [5]

Placement of Stephanorhinus kirchbergensis among recent and subfossil rhinoceros species based on nuclear genomes (Liu, 2021): [4]

Elasmotheriinae

† Elasmotherium sibiricum

Rhinocerotinae

Black rhinoceros (Diceros bicornis)

White Rhinoceros (Ceratotherium simum)

Indian rhinoceros (Rhinoceros unicornis)

Javan rhinoceros (Rhinoceros sondaicus)

Sumatran rhinoceros (Dicerorhinus sumatrensis)

† Woolly rhinoceros (Coelodonta antiquitatis)

† Merck's rhinoceros (Stephanorhinus kirchbergensis)

Bayesian morphological phylogeny (Pandolfi, 2023) Note: This excludes living African rhinoceros species. [5]

Description

Species of Stephanorhinus were large-sized rhinoceroses, with body masses estimated between 1,500–3,000 kilograms (3,300–6,600 lb). [7] Stephanorhinus species have proportionally long (dolichocephalic) skulls. They had two horns, a frontal and a nasal horn. The nasal septum was partially ossified (turned to bone), which connected the nasal bones with the premaxillary bones. The incisors were either lost completely or very heavily reduced in size. [2]

Species and evolution

The oldest known species of the genus are from the Pliocene of Europe. [8] The species "S." pikermiensis and "S." megarhinus that were formerly considered to belong to Stephanorhinus are currently considered to belong to Dihoplus and Pliorhinus , respectively. [9] [10] [11] “Stephanorhinus” miguelcrusafonti from the Early Pliocene of Western Europe has also been assigned to Pliorhinus in recent studies. [10]

The position of Stephanorhinus? africanus from the Middle Pliocene of Tunisia and Chad is uncertain. [12] Some authors have suggested that Stephanorhinus likely originated from members of the genus Pliorhinus. [5]

Stephanorhinus jeanvireti, also known as S. elatus [13] 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, [14] France, Italy, [15] Slovakia [16] and Greece, [17] and the Early Pleistocene of Romania, [18] with its temporal span being around 3.4 to 2 million years ago (Ma). [5]

Stephanorhinus etruscus first appears in the latest Pliocene in the Iberian Peninsula, around 3.3 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. [19] Stephanorhinus etruscus is thought to have had a browsing based diet. [20]

Remains of Stephanorhinus not assigned to species have been reported from the Dmanisi site in the Caucasus (Georgia), dating to around 1.8 Ma. The remains appear to belong to two morphotypes, which may represent distinct species. [21] Some of these remains may be closely related to the later species Stephanorhinus hundsheimensis. [22]

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. [23] 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. [24] Stephanorhinus hundsheimensis is typically suggested to have gone extinct at around 0.5 Ma, [5] though a 2023 study suggested that the species may have persisted as recently as the latest Middle Pleistocene-earliest Late Pleistocene around 130,000 years ago, based on fossils found in Spain. [25]

Stephanorhinus migrated from its origin in western Eurasia into eastern Eurasia during the Early Pleistocene, [5] with remains of Stephanorhinus including those of S. etruscus being known from the Early Pleistocene of Kazakhstan and Tajikistan in Central Asia, with the earliest remains of the genus in China dating to around 1.6 Ma. [22] Stephanorhinus yunchuchenensis is known from a single specimen in probably late Early Pleistocene aged deposits in Yushe, Shaanxi, China, while Stephanorhinus lantianensis is also known from a single specimen from late Early Pleistocene (1.15 Ma) deposits in Lantian, also in Shaanxi. [26] These may be synonymous with other named Stephanorhinus species, with a 2022 study suggesting that they were likely synonyms of S. kirchbergensis and S. etruscus respectively. [22]

The first definitive record of Stephanorhinus kirchbergensis (Merck's rhinoceros) is in China at Zhoukoudian (Choukoutien; near Beijing), around the Early–Mid-Pleistocene transition at 0.8 Ma. [26]

Approximate time averaged range of Stephanorhinus kirchbergensis (red) and Stephanorhinus hemitoechus (blue), with overlapping range in purple. Stephanorhinus kirchbergensis range map.svg
Approximate time averaged range of Stephanorhinus kirchbergensis (red) and Stephanorhinus hemitoechus (blue), with overlapping range in purple.

S. kirchbergensis appeared in Europe between 0.7-6 Ma with S. hemitoechus (the narrow-nosed rhinoceros) first appearing in Europe around 0.6-0.5 Ma. S. kirchbergensis and S. hemitoechus are typically interpreted mixed feeders tending towards browsing and grazing, 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. [27]

S. kirchbergensis was broadly distributed over northern Eurasia from Western Europe to East Asia, while S. hemitoechus was generally confined to the western Palearctic, including Europe, West Asia, and North Africa. [28] [29] [12]

In Europe, the timing of the extinction of S. kirchbergensis is uncertain, though it is sometime after 115,000 years ago. [30] The latest records of S. hemitoechus in Europe are known from the Iberian Peninsula, where they survived until at least 34,000 years ago, [31] with the species possibly surviving as late as 15,500 years ago in the Levant. [32] [33] In the Altai region, S. kirchbergensis survived until at least 40,000 years ago. [34] In South China, the species may have survived into Marine Isotope Stage 2 (~29-14,000 years ago). [35]

Relationship with humans

Remains of Stephanorhinus species have been found in sites across Europe with break or cut marks indicating that they were butchered by archaic humans. [36] [37] [38] [39] [40] [41] The earliest such site is Vallonnet Cave in France dating to around 1.2 to 1.1 million years ago, where remains of S. hundsheimensis have been reported with cut marks. [38] Another early site is Boxgrove in England, dating to around 500,000 years ago, where an indeterminate species of Stephanorhinus was found with cut marks thought to have been created by Homo heidelbergensis . [42] The youngest sites are known from the late Middle Paleolithic (around 100-40,000 years ago), which were created by Neanderthals. [39] At some sites hunting is suggested to be the more likely than scavenging based on mortality profiles. [38]

An astragalus attributed to Stephanorhinus was discovered in the Southwest Quarter of Mycenae, along with a collection of other artifacts. Its placement there was dated to the thirteenth century B.C.E. [43]

Related Research Articles

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Elasmotherium is an extinct genus of large rhinoceros endemic to Eastern Europe and Central Asia with isolated finds from East Asia during Late Miocene through to the Late Pleistocene, with the youngest reliable dates around 39,000 years ago. It was the last surviving member of Elasmotheriinae, a distinctive group of rhinoceroses separate from the group that contains living rhinoceros (Rhinocerotinae).

<span class="mw-page-title-main">Woolly rhinoceros</span> Extinct species of rhinoceros of northern Eurasia

The woolly rhinoceros is an extinct species of rhinoceros that inhabited northern Eurasia during the Pleistocene epoch. The woolly rhinoceros was a member of the Pleistocene megafauna. The woolly rhinoceros was covered with long, thick hair that allowed it to survive in the extremely cold, harsh mammoth steppe. It had a massive hump reaching from its shoulder and fed mainly on herbaceous plants that grew in the steppe. Mummified carcasses preserved in permafrost and many bone remains of woolly rhinoceroses have been found. Images of woolly rhinoceroses are found among cave paintings in Europe and Asia. The range of the woolly rhinoceros contracted towards Siberia beginning around 17,000 years ago, with the youngest known records being around 14,000 years old in northeast Siberia, coinciding with the Bølling–Allerød warming, which likely disrupted its habitat, with environmental DNA records possibly extending the range of the species around 9,800 years ago. Its closest living relative is the Sumatran rhinoceros.

<span class="mw-page-title-main">Rhinoceros</span> Family of mammals

A rhinoceros, commonly abbreviated to rhino, is a member of any of the five extant species of odd-toed ungulates in the family Rhinocerotidae; it can also refer to a member of any of the extinct species of the superfamily Rhinocerotoidea. Two of the extant species are native to Africa, and three to South and Southeast Asia.

<i>Diceros</i> Genus of Rhinocerotidae

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<i>Dicerorhinus</i> Genus of mammals

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<i>Rhinoceros</i> (genus) Genus of mammals

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<span class="mw-page-title-main">Narrow-nosed rhinoceros</span> Extinct species of rhinoceros

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References

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  3. ↑ 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
  4. 1 2 Liu, Shanlin; Westbury, Michael V.; Dussex, Nicolas; Mitchell, Kieren J.; Sinding, Mikkel-Holger S.; Heintzman, Peter D.; Duchêne, David A.; Kapp, Joshua D.; von Seth, Johanna; Heiniger, Holly; Sánchez-Barreiro, Fátima (August 2021). "Ancient and modern genomes unravel the evolutionary history of the rhinoceros family". Cell. 184 (19): 4874–4885.e16. doi: 10.1016/j.cell.2021.07.032 . hdl: 10230/48693 . ISSN   0092-8674. PMID   34433011. S2CID   237273079.
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  6. ↑ 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; Rakownikow Jersie-Christensen, Rosa; Mackie, Meaghan; Ginolhac, Aurélien; Ferring, Reid; Tappen, Martha (2019-09-11). "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.
  7. ↑ Jan van der Made und René Grube: The rhinoceroses from Neumark-Nord and their nutrition. In: Harald Meller (Hrsg.): Elefantenreich – Eine Fossilwelt in Europa. Halle/Saale 2010, S. 382–394
  8. ↑ Ballatore, Manuel; Breda, Marzia (March 2019). "Revision of the rhinoceros remains (Rhinocerotidae, Mammalia) from the late Pliocene of Etouaires (Auvergne, France) and the morphological distinction between the postcranial bones of Stephanorhinus elatus and S. etruscus". Comptes Rendus Palevol . 18 (2): 191–208. doi:10.1016/j.crpv.2018.06.003 . Retrieved 5 November 2024– via Elsevier Science Direct.
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  17. ↑ Guérin, Claude; Tsoukala, Evangelia (June 2013). "The Tapiridae, Rhinocerotidae and Suidae (Mammalia) of the Early Villafranchian site of Milia (Grevena, Macedonia, Greece)". Geodiversitas. 35 (2): 447–489. doi:10.5252/g2013n2a7. ISSN   1280-9659. S2CID   129164720.
  18. ↑ Pandolfi, Luca; Codrea, Vlad A.; Popescu, Aurelian (December 2019). "Stephanorhinus jeanvireti (Mammalia, Rhinocerotidae) from the early Pleistocene of Colțești (southwestern Romania)". Comptes Rendus Palevol. 18 (8): 1041–1056. Bibcode:2019CRPal..18.1041P. doi: 10.1016/j.crpv.2019.07.004 .
  19. ↑ 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. Bibcode:2017CRPal..16..762P. doi: 10.1016/j.crpv.2017.06.004 . hdl: 11336/57305 .
  20. ↑ Pandolfi, Luca; Bartolini-Lucenti, Saverio; Cirilli, Omar; Bukhsianidze, Maia; Lordkipanidze, David; Rook, Lorenzo (July 2021). "Paleoecology, biochronology, and paleobiogeography of Eurasian Rhinocerotidae during the Early Pleistocene: The contribution of the fossil material from Dmanisi (Georgia, Southern Caucasus)". Journal of Human Evolution. 156: 103013. Bibcode:2021JHumE.15603013P. doi:10.1016/j.jhevol.2021.103013. PMID   34030060. S2CID   235199139.
  21. ↑ Pandolfi, Luca; Bartolini-Lucenti, Saverio; Cirilli, Omar; Bukhsianidze, Maia; Lordkipanidze, David; Rook, Lorenzo (July 2021). "Paleoecology, biochronology, and paleobiogeography of Eurasian Rhinocerotidae during the Early Pleistocene: The contribution of the fossil material from Dmanisi (Georgia, Southern Caucasus)". Journal of Human Evolution. 156: 103013. Bibcode:2021JHumE.15603013P. doi:10.1016/j.jhevol.2021.103013. PMID   34030060.
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