Stephanorhinus kirchbergensis

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Stephanorhinus kirchbergensis
Temporal range: Middle Pleistocene–Late Pleistocene
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Dicerorhinus kirchbergensis.JPG
Mostly complete skull from Germany
Stephanorhinus kirchbergensis skull.svg
Diagram of skull from Russia
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Perissodactyla
Family: Rhinocerotidae
Genus:† Stephanorhinus
Species:
†S. kirchbergensis
Binomial name
†Stephanorhinus kirchbergensis
(Jäger, 1839)
Synonyms
List
  • Coelodonta merckii
    (Kaup, 1841)
  • Dicerorhinus merckii
    (Kaup, 1841)
  • Dc. m. var. brachycephala
    Schroeder, 1903
  • Dc. kirchbergensis
    (Jäger, 1839)
  • Dc. k. var. brachycephala
    Schroeder, 1903
  • Dihoplus kirchbergensis
    (Jäger, 1839)
  • R. incisivus
    Merck, 1784
  • R. kirchbergense
    Jäger, 1839
  • R. leptorhinus
    Cuvier, 1836
  • R. megarhinus
    de Christol, 1834
  • R. merckii (or mercki, merki)
    Kaup, 1841
  • R. m. var. brachycephala
    Schroeder, 1903
  • R. (Tichorhinus) merckii
    (Kaup, 1841)
  • Stephanorhinus yunchuchenensis(Chow, 1963)

Stephanorhinus kirchbergensis, also known as Merck's rhinoceros or the forest rhinoceros, is an extinct species of rhinoceros belonging to the genus Stephanorhinus from the Middle to Late Pleistocene of Eurasia. Its range spanned from western Europe to eastern Asia. Among the last members of the genus, it co-existed alongside Stephanorhinus hemitoechus (the narrow-nosed or steppe rhinoceros) in the western part of its range.

Contents

Etymology and taxonomy

The first part of the genus name is derived from that of King Stephen I of Hungary, and the second part from 'rhinos' (ρινος, meaning "nose"), as with Dicerorhinus. The species name was given by Georg Friedrich von Jäger in 1839 for Kirchberg an der Jagst in Baden-Württemberg, Germany where the type specimens had been found. [1] It is often known in English (and equivalents in other languages) as Merck's rhinoceros after Carl Heinrich Merck, who gave the initial name to the species in 1784 as Rhinoceros incisivus, that is now considered a nomen oblitum , and who after a widely used junior synonym of the species, Rhinoceros merckii (historically several alternate spellings) was named by Johann Jakob Kaup in 1841. [2]

Description

Comparison of the head angle of Merck's rhinoceros (top) with that of Stephanorhinus hemitoechus and the woolly rhinoceros (middle and bottom, respectively) Skulls of pleistocene rhinos.png
Comparison of the head angle of Merck's rhinoceros (top) with that of Stephanorhinus hemitoechus and the woolly rhinoceros (middle and bottom, respectively)

Merck's rhinoceros was a large rhinoceros, with a particularly large specimen from Poland reaching an estimated height at the withers of 1.82 metres (6.0 ft). [3] The bones of the skeleton are robust and massive. The skull of Merck's rhinoceros is elongated, with the septum nasalis ossified only towards its anterior (front) end. The mandibular symphysis is relatively long and the mandible has a horizontal high, thick branch. [4]

Dental anatomy

The enamel of the teeth is very thick, and often bright coloured and smooth, with very thin or absent coronal cement. The buccal (cheek-facing) sides of the teeth often have sub-vertical bluish lines. Tooth dimensions are highly variable in comparison to other Stephanorhinus species. The upper teeth, especially the molars, are much higher towards the buccal side than to the lingual (towards the tongue) side. The ectolophs of the first and second upper molars have shallower folds, especially the fold between the paracone and mesostyle, than those of S. hemitoechus, resulting in a less pronounced undulation. In comparison to other species of Stephanorhinus, the premolars of S. kirchbergensis are mesially (towards the front of the tooth) broad and relatively lingually short. The upper premolar ectoloph folds are shallow, and have narrow anterior valleys. The ectoloph curves strongly mesially and often distally (towards the hind portion of the tooth) towards the inside of the tooth. In both upper molars and premolars, the metalophs and the protolophs are distinctly bulbous. The lower premolars and molars are similar and hard to distinguish. [4]

Origin

The earliest definitive records are from Choukoutien Locality 13, in Fangshan District near Beijing at around the Early-Middle Pleistocene transition. [5] Stephanorhinus yunchuchenensis from Shanxi, China, likely represents a junior synonym of S. kirchbergensis, its precise age is uncertain, but it has been suggested to date to the late Early Pleistocene. [6] S. kirchbergensis appears in Europe during the early Middle Pleistocene between 0.7 and 0.6 million years ago, existing alongside the already present S. hundsheimensis. [7] Mitochondrial and nuclear genomes obtained from a permafrost specimen [8] [9] and a dental proteome [10] suggest that it is more closely related to the woolly rhinoceros than the Sumatran rhinoceros. A 2023 morphological study suggested its closest relative was the narrow-nosed rhinoceros (S. hemitoechus). [11]

Relationships among Late Pleistocene and modern rhinoceros genera, based on nuclear DNA, after Liu et al, 2021: [9]

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, after (Pandolfi, 2023) Note: This excludes living African rhinoceros species. [11]

Hyrachyus eximius

Tapirus terrestris (South American tapir)

Rhinocerotidae

Trigonias osborni

Ronzotherium filholi

Aceratheriini
Teleoceratina
Rhinocerotina

Distribution

Range of Merck's rhinoceros, excluding arctic localities Distribution of S. kirchbergensis2.png
Range of Merck's rhinoceros, excluding arctic localities

Its range spans from Europe to East Asia, but appears to be absent from the Iberian Peninsula. [12] [13] It was predominantly present in Europe during interglacial periods where it formed part of the Palaeoloxodon antiquus assemblage, where it occurred alongside the straight-tusked elephant (Palaeoloxodon antquus) the narrow-nosed rhinoceros (Stephanorhinus hemitoechus), and the hippopotamus (Hippopotamus amphibius). [14] It is presumed to have had a preference for closed forest and woodland habitats, as opposed the to open grassland habitats favoured by S. hemitoechus. [14] Its range extended into the Arctic Circle, with a 70–48,000 year old skull known from arctic Yakutia in the Chondon River valley [8] and a late Middle Pleistocene aged lower jaw from the Yana River valley. [15] Teeth are known from caves in Primorsky Krai , suggested to date between 50,000 and 25,000 years ago based on dates of other bones found in the deposit, which are the easternmost known records, [16] along with a record from the Middle Pleistocene of western Japan. [17] A tooth of S. cf. kirchbergensis of an unknown age is known from the Lut Desert in eastern Iran. [18] It is fairly common throughout the Pleistocene in North China, [19] but is a rarer component of South Chinese assemblages, [20] being known from around 30 localities in the region. [5] Antoine (2012) states that D.choukoutienensis, D. lantianensis, and D. yunchuchenensis are local names for the taxon, without elaboration. [19] Its range was strongly controlled by glacial cycles, with the species experiencing repeated cycles of expansion and contraction as the ice sheets advanced, this accounts for the relative rarity of its remains in comparison to the woolly rhinoceros. [12] During the Last Glacial Period, the species range contracted. The timing of its extinction in Europe is uncertain, thought it postdates the end of the Last Interglacial around 115,000 years ago. [21] Radiocarbon dated remains from the Altai date to around 40,000 years ago. [22] The youngest reliable records in China are from the Rhino Cave in Hubei, which is early Late Pleistocene in age. [20] Though less definitive remains are known from near Harbin in Heilongjiang, which are thought to be 20 kya in age. [5] Records from Migong Cave just south of the Yangtze River in the Three Gorges area are suggested to date to MIS 2 (29,000-14,000 years ago). [23]

Diet

Restoration of two Merck's rhinoceros in open wooded landscape with oak trees during the Eemian interglacial in Europe Stephanorhinus kirchbergensis Eemian landscape.jpg
Restoration of two Merck's rhinoceros in open wooded landscape with oak trees during the Eemian interglacial in Europe

Merck's rhinoceros has been interpreted as a browser or a mixed feeder, consuming both browse such as branches and leaves of trees and shrubs, as well as low-lying vegetation. Its diet appears to have varied according to local conditions. [24] [25] [7] Analysis of plant material embedded within teeth from the Neumark-Nord locality in Germany found remains of Populus (poplar or aspen) Quercus (oak), Crataegus (hawthorn), Pyracantha , Urtica (nettles) and Nymphaea (water lilies) as well as indeterminate remains of Betulaceae, Rosaceae, and Poaceae (grass). [26] Preserved plant remains found with the teeth on the arctic Chondon skull included twigs of Salix (willow), Betula (birch) and abundant Larix (larch) alongside fragments of Ericaceae (heather); sedges were notably absent. [8] A specimen from Eemian aged deposits in Gorzów Wielkopolski in Poland had twigs of Corylus (hazel), Carpinus (hornbeam), and Viscum (mistletoe), alongside fruit scales of birch, with hazel and birch dominating amongst the pollen. [24] The pollen from a specimen found at Spinadesco in Italy was dominated (~50%) by trees, particularly Alnus (alder) and Fagus (beech), with Hippophae rhamnoides (sea buckthorn), dominating amongst the shrubs, with around 30% of the total contribution being from a variety of herbaceous plants. [27]

Human exploitation

Cut marks are known on bones of S. kirchbergensis from the Guado San Nicola site in central Italy, which dates to the late Middle Pleistocene, around 400-345,000 years ago. [28] Remains of S. kirchbergensis with cut marks have also been reported from the Medzhibozh locality in western Ukraine, dating to MIS 11, around 425-375,000 years ago. [29] At the Taubach travertine site in Thuringia, Germany, which dates to the Eemian (approximately 130,000-115,000 years ago) abundant remains of Merck's rhinoceros with cut marks are known. The vast majority of remains were of young subadults, alongside a much smaller number of adults. It has been suggested that the rhinoceroses were killed and butchered on site by Neanderthals. [30]

Related Research Articles

<i>Elasmotherium</i> Genus of extinct rhinoceroses

Elasmotherium is an extinct genus of large rhinoceros endemic to Eurasia 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, simply known as woolly rhino, 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 species range 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.

<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>Dicerorhinus</i> Genus of mammals

Dicerorhinus is a genus of the family Rhinocerotidae, consisting of a single extant species, the two-horned Sumatran rhinoceros, and several extinct species. The genus likely originated in the Mid to Late Pliocene of Northern Indochina and South China. Many species previously placed in this genus probably belong elsewhere.

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

Rhinoceros is a genus comprising one-horned rhinoceroses. This scientific name was proposed by Swedish taxonomist Carl Linnaeus in 1758. The genus contains two species, the Indian rhinoceros and the Javan rhinoceros. Although both members are threatened, the Javan rhinoceros is one of the most endangered large mammals in the world with only 60 individuals surviving in Java (Indonesia). The word 'rhinoceros' is of Greek origin meaning "nose-horn".

<span class="mw-page-title-main">Gomphothere</span> Extinct family of proboscidean mammals

Gomphotheres are an extinct group of proboscideans related to modern elephants. They were widespread across Afro-Eurasia and North America during the Miocene and Pliocene epochs and dispersed into South America during the Pleistocene as part of the Great American Interchange. Gomphotheres are a paraphyletic group that is ancestral to Elephantidae, which contains modern elephants, as well as Stegodontidae. While most famous forms such as Gomphotherium had long lower jaws with tusks, which is the ancestral condition for the group, some later members developed shortened (brevirostrine) lower jaws with either vestigial or no lower tusks, looking very similar to modern elephants, an example of parallel evolution, which outlasted the long-jawed gomphotheres. By the end of the Early Pleistocene, gomphotheres became extinct in Afro-Eurasia, with the last two genera, Cuvieronius ranging from southern North America to western South America, and Notiomastodon having a wide range over most of South America until the end of the Pleistocene around 12,000 years ago, when they became extinct following the arrival of humans.

<i>Panthera spelaea</i> Extinct species of lion

Panthera spelaea, also known as the cave lion or steppe lion, is an extinct Panthera species that most likely evolved in Europe after the third Cromerian interglacial stage, less than 600,000 years ago. Genetic analysis of ancient DNA has revealed that while closely related, it was a distinct species genetically isolated from the modern lion occurring in Africa and Asia, with the genetic divergence between the two species variously estimated between 1.9 million and 600,000 years ago. It is closely related and probably ancestral to the American lion. The species ranged from Western Europe to eastern Beringia in North America, and was a prominent member of the mammoth steppe fauna. It became extinct about 13,000 years ago.

<i>Coelodonta</i> Extinct genus of rhinoceros

Coelodonta is an extinct genus of rhinoceros that lived in Eurasia between 3.7 million years to 14,000 years ago, in the Pliocene and the Pleistocene epochs. It is best known from the type species, the woolly rhinoceros, which ranged throughout northern Eurasia during the Pleistocene. The earliest known species, Coelodonta thibetana, lived in Tibet during the Pliocene, with the genus spreading to the rest of Eurasia during the Pleistocene.

The Chibanian, widely known as the Middle Pleistocene, is an age in the international geologic timescale or a stage in chronostratigraphy, being a division of the Pleistocene Epoch within the ongoing Quaternary Period. The Chibanian name was officially ratified in January 2020. It is currently estimated to span the time between 0.770 Ma and 0.126 Ma, also expressed as 770–126 ka. It includes the transition in palaeoanthropology from the Lower to the Middle Paleolithic over 300 ka.

<span class="mw-page-title-main">Straight-tusked elephant</span> Extinct species of elephant

The straight-tusked elephant is an extinct species of elephant that inhabited Europe and Western Asia during the Middle and Late Pleistocene. It was larger than any living elephant, with adult males suggested to reach 3.81–4.2 metres (12.5–13.8 ft) in shoulder height, and 11.3–15 tonnes in weight. Like modern elephants, the straight-tusked elephant lived in herds, flourishing during interglacial periods, when its range would extend as far north as Great Britain. Skeletons found in association with stone tools and wooden spears suggest they were scavenged and hunted by early humans, including Neanderthals. It is the ancestral species of most dwarf elephants that inhabited islands in the Mediterranean.

<i>Anancus</i> Genus of proboscideans

Anancus is an extinct genus of "tetralophodont gomphothere" native to Afro-Eurasia, that lived from the Tortonian stage of the late Miocene until its extinction during the Early Pleistocene, roughly from 8.5–2 million years ago.

<i>Cuvieronius</i> Extinct genus of mammals

Cuvieronius is an extinct New World genus of gomphothere which ranged from southern North America to western South America during the Pleistocene epoch. Among the last gomphotheres, it became extinct at the end of the Pleistocene, approximately 12,000 years ago, following the arrival of humans to the Americas.

<i>Equus namadicus</i> Extinct species of mammal

Equus namadicus is a prehistoric equid, known from remains dating to the Middle and Late Pleistocene from across the Indian subcontinent, with its last dated records being approximately 29-14,000 years ago. It is considered a "stenonine horse", meaning that it is probably more closely related to zebras and asses than true horses. It is relatively large in size. It is very similar to the earlier Equus sivalensis, also from the Indian subcontinent, from which it only differs in size and in subtle aspects of dental anatomy, and it has sometimes been suggested to be a synonym of it.

<i>Hippopotamus antiquus</i> Extinct species of mammal

Hippopotamus antiquus is an extinct species of Hippopotamus that ranged across Europe during the Early and Middle Pleistocene.

<i>Stephanorhinus</i> Extinct genus of rhinoceros

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 and the narrow-nosed rhinoceros – went extinct during the last glacial period.

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

The narrow-nosed rhinoceros, also known as the steppe rhinoceros is an extinct species of rhinoceros belonging to the genus Stephanorhinus that lived in western Eurasia, including Europe, as well as North Africa during the Pleistocene. It first appeared in Europe some 600,000 years ago during the Middle Pleistocene and survived there until at least 34,000 years Before Present.

<i>Bubalus murrensis</i> Extinct species of mammal

Bubalus murrensis, also known as European water buffalo, is an extinct buffalo species native to Europe during the Pleistocene epoch.

<i>Nesorhinus</i> Extinct genus of rhinoceros from the Pleistocene of the Philippines and Taiwan

Nesorhinus is an extinct genus of rhinoceros from the Pleistocene of Asia. It contains two species, Nesorhinus philippinensis from Luzon, Philippines and Nesorhinus hayasakai from Taiwan.

<i>Dihoplus</i> Extinct genus of rhinoceros

Dihoplus is an extinct genus of rhinoceros that lived in Eurasia from the Late Miocene to Pliocene.

<i>Pliorhinus</i> Extinct genus of rhinoceros

Pliorhinus is an extinct genus of rhinoceros known from the Late Miocene and Pliocene of Eurasia. The type species, Pliorhinus megarhinus, was previously assigned to Dihoplus.

References

  1. ↑ Georg Friedrich Jäger: Über die fossilen Säugetiere welche in Württemberg in verschiedenen Formationen aufgefunden worden sind, nebst geognostischen Bemerkungen über diese Formationen. C. Erhard Verlag, Stuttgart, 1835–39
  2. ↑ Johann Jakob Kaup: Akten der Urwelt oder Osteologie der urweltlichen Säugethiere und Amphibien. Darmstadt, Verlag des Herausgebers, 1841
  3. ↑ Sobczyk, Artur; Borówka, Ryszard K.; Badura, Janusz; Stachowicz-Rybka, Renata; Tomkowiak, Julita; Hrynowiecka, Anna; Sławińska, Joanna; Tomczak, Michał; Pitura, Mateusz; Lamentowicz, Mariusz; Kołaczek, Piotr (May 2020). "Geology, stratigraphy and palaeoenvironmental evolution of the Stephanorhinus kirchbergensis -bearing Quaternary palaeolake(s) of Gorzów Wielkopolski (NW Poland, Central Europe)". Journal of Quaternary Science. 35 (4): 539–558. Bibcode:2020JQS....35..539S. doi:10.1002/jqs.3198. hdl: 10261/237944 . ISSN   0267-8179. S2CID   216264117.
  4. 1 2 Selected records of Stephanorhinus kirchbergensis (Jäger, 1839 (Mammalia, Rhinocerotidae) in Italy Emmanuel M.E. BILLIA & Carmelo PETRONIO Bollettino della Società Paleontologica Italiana, 48 (1), 2009, xx-xx. Modena
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  16. ↑ 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. PMID   32483707. S2CID   219156923.
  17. ↑ Handa, Naoto; Pandolfi, Luca (July 2016). "Reassessment of the Middle Pleistocene Japanese Rhinoceroses (Mammalia, Rhinocerotidae) and Paleobiogeographic Implications". Paleontological Research. 20 (3): 247–260. doi:10.2517/2015PR034. ISSN   1342-8144.
  18. ↑ Hashemi, N; Ashouri, A; Aliabadian, M; M. Gharaie, M; Sánchez Marco, A; Louys, J (2016). "First report of Quaternary mammals from the Qalehjough area, Lut Desert, Eastern Iran". Palaeontologia Electronica. doi: 10.26879/539 . ISSN   1094-8074.
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  20. 1 2 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. S2CID   67828905.
  21. ↑ Stefaniak, Krzysztof; Kotowski, Adam; Badura, Janusz; Sobczyk, Artur; Borówka, Ryszard K.; Stachowicz-Rybka, Renata; Moskal-del Hoyo, Magdalena; Hrynowiecka, Anna; Tomkowiak, Julita; Sławińska, Joanna; Przybylski, Bogusław; Ciszek, Dariusz; Alexandrowicz, Witold Paweł; Skoczylas-Śniaz, Sylwia; Ratajczak-Skrzatek, Urszula (2023-05-09). "A skeleton of peat-trapped forest rhinoceros Stephanorhinus kirchbergensis (Jäger, 1839) from Gorzów Wielkopolski, Northwestern Poland: a record of life and death of the Eemian large mammals". Neues Jahrbuch für Geologie und Paläontologie - Abhandlungen. 308 (1): 45–77. doi:10.1127/njgpa/2023/1129. ISSN   0077-7749.
  22. ↑ Кириллова, И. В.; Вершинина, А. О.; Зазовская, Э. П.; Занина, О. Г.; Катлер, С.; Косинцев, П. А.; Лаптева, Е. Г.; Чернова, О. Ф.; Шапиро, Б. (2021). "К вопросу о времени и среде обитания Stephanorhinus kirchbergensis Jäger 1839 (Mammalia, Rhinoceratidae) на Алтае и северо-востоке России" [To the question of time and environment of Stephanorhinus kirchbergensis Jäger 1839 (Mammalia, Rhinocerotidae) in Altai and North-East of Russia]. Зоологический Журнал (in Russian). 100 (5): 558–572. doi:10.31857/S0044513421050068. ISSN   0044-5134. S2CID   234970639.
  23. ↑ 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. Bibcode:2017QuInt.434...25P. doi:10.1016/j.quaint.2014.11.039.
  24. 1 2 Stefaniak, Krzysztof; Stachowicz-Rybka, Renata; Borówka, Ryszard K.; Hrynowiecka, Anna; Sobczyk, Artur; Moskal-del Hoyo, Magdalena; Kotowski, Adam; Nowakowski, Dariusz; Krajcarz, Maciej T.; Billia, Emmanuel M.E.; Persico, Davide (September 2020). "Browsers, grazers or mix-feeders? Study of the diet of extinct Pleistocene Eurasian forest rhinoceros Stephanorhinus kirchbergensis (Jäger, 1839) and woolly rhinoceros Coelodonta antiquitatis (Blumenbach, 1799)". Quaternary International. 605–606: 192–212. doi:10.1016/j.quaint.2020.08.039. hdl: 10261/255910 . S2CID   224984977.
  25. ↑ 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. doi:10.1016/j.jhevol.2021.103013. PMID   34030060.
  26. ↑ 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
  27. ↑ Burkanova, Elena M.; Billia, Emmanuel M.E.; Persico, Davide (July 2020). "Stephanorhinus kirchbergensis (Jäger, 1839) (Mammalia, Rhinocerotidae) from the Po valley (Lombardia, Northern Italy): possible diet/nutrition and living conditions". Quaternary International. 554: 164–169. Bibcode:2020QuInt.554..164B. doi:10.1016/j.quaint.2020.07.031. ISSN   1040-6182. S2CID   225542965.
  28. ↑ Berruti, Gabriele Luigi Francesco; Arzarello, Marta; Ceresa, Allison; Muttillo, Brunella; Peretto, Carlo (December 2020). "Use-Wear Analysis of the Lithic Industry of the Lower Palaeolithic Site of Guado San Nicola (Isernia, Central Italy)". Journal of Paleolithic Archaeology. 3 (4): 794–815. Bibcode:2020JPalA...3..794B. doi:10.1007/s41982-020-00056-3. ISSN   2520-8217.
  29. ↑ Stepanchuk, V.N.; Moigne, A.-M. (July 2016). "MIS 11-locality of Medzhibozh, Ukraine: Archaeological and paleozoological evidence". Quaternary International. 409: 241–254. Bibcode:2016QuInt.409..241S. doi:10.1016/j.quaint.2015.09.050.
  30. ↑ Bratlund, B. 1999. Taubach revisited. Jahrbuch des Römisch-Germanischen Zentralmuseums Mainz 46: 61-174.
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