Narrow-nosed rhinoceros

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Narrow-nosed rhinoceros
Temporal range: Middle Pleistocene–Late Pleistocene
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Dicerorhinus hemitoechus.JPG
Skull of Stephanorhinus hemitoechus
Hemitoechus2011.jpg
Stephanorhinus hemitoechus life restoration
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Perissodactyla
Family: Rhinocerotidae
Genus: Stephanorhinus
Species:
S. hemitoechus
Binomial name
Stephanorhinus hemitoechus
Falconer, 1859
Stephanorhinus kirchbergensis range map.svg
Range of Stephanorhinus hemitoechus (blue) and Stephanorhinus kirchbergensis (red), with overlapping range in purple
Synonyms
  • Dicerorhinus hemitoechusFalconer, 1859
  • Rhinoceros hemitoechus Falconer, 1859
  • Rhinoceros binagadensisDzhafarov, 1955
  • Rhinoceros subinermis Pomel, 1895

The narrow-nosed rhinoceros (Stephanorhinus hemitoechus), 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 [1] during the Pleistocene. It first appeared in Europe around 500,000 years ago during the Middle Pleistocene and survived there until at least 34,000 years Before Present. It was native to temperate and Mediterranean environments, where it fed on low growing plants and to a lesser extent woody plants. Evidence has been found that it was exploited for food by archaic humans, including Neanderthals.

Contents

Taxonomy

The species was originally described by Hugh Falconer in 1859 based on remains found in cave deposits in Glamorganshire, south Wales in Great Britain, dating to the Eemian/Last interglacial, around 130-115,000 years ago. The species like other Stephanorhinus species was previously assigned historically to the genus Dicerorhinus , which includes the Sumatran rhinoceros, [2] which genetic evidence indicates is the closest living relative of Stephanorhinus. [3]

Description

Skulls of the narrow-nosed rhinocers (middle) compared to S. kirchbergensis, (top) and the woolly rhinoceros (bottom), showing the difference in head angle Skulls of pleistocene rhinos.png
Skulls of the narrow-nosed rhinocers (middle) compared to S. kirchbergensis, (top) and the woolly rhinoceros (bottom), showing the difference in head angle

The narrow-nosed rhinoceros was a large animal, reaching a body mass of approximately 1,500 kilograms (3,300 lb), making it around the same size/slightly larger than S. hundsheimensis, but smaller than S. kirchbergensis (Merck's rhinoceros) and the woolly rhinoceros (Coelodonta antiquitatis). [4] The size of the species was somewhat variable depending on local conditions. [5] [6] The skull is low slung, with the cranium being downwardly directed. [7] The nasal notch (narial incison) is deep, extending back as far as the first upper molar (M1). [8] The teeth are relatively high crowned (hypsodont) compared to other Stephanorhinus species, with the third molars being relatively enlarged. [7] The teeth are relatively shifted forwards in the jaw. [8] Like other species of Stephanorhinus, S. hemitoechus is thought to have borne two horns, [9] though the attachment regions for the horns are "weakly developed" similar to S. etruscus but unlike S. hundsheimensis. [8] The lower jaw is relatively slender in comparison to S. kirchbergensis, and tapers, becoming slender and narrow towards the front (the mandibular symphysis). [8] Compared to the earlier species S. jeanvireti, S. etruscus, and S. hundsheimensis the limb joints are relatively shallow, and the limb bones relatively broad and short. [7]

Distribution

In comparison to the widespread Merck's rhinoceros (Stephanorhinus kirchbergensis), the narrow-nosed rhinoceros had a less geographically extensive distribution, including much of Europe, [10] [11] as well as West Asia [12] and North Africa. [1] In Europe the species is known from abundant remains in the Iberian Peninsula in the west (where S. kirchbergensis appears to have been rare or absent), [13] eastwards to Italy, [5] Bulgaria [14] and Greece. [15] [16] Its range extended northwards into northern France, [17] Britain (as far north as North Yorkshire [18] ), Germany [2] and Slovakia [19] during warm interglacial periods. [20]

In North Africa, where the species was previously known as Rhinoceros subinermis [15] , remains are known from Cyrenacia in northeast Libya [1] as well as the Maghreb in Morocco [1] and northern Algeria. [21] In West Asia, the range of the species extends from the Levant, including Israel, [22] Palestine, Lebanon, [16] Jordan [23] and Syria [24] in the west, to western Iran, [25] [26] and Azerbaijan in the east, [27] where some remains were previously referred to as the species Rhinoceros binagadensis. [15]

Ecology

A Middle Pleistocene landscape in Spain, including the narrow-nosed rhinoceros (far right) as well the straight-tusked elephant (background centre-left), the extinct fallow deer Dama celiae (foreground) wild horse (left), bison, (background centre) and aurochs (background right) Middle Pleistocene landscape in Manzanares valley.png
A Middle Pleistocene landscape in Spain, including the narrow-nosed rhinoceros (far right) as well the straight-tusked elephant (background centre-left), the extinct fallow deer Dama celiae (foreground) wild horse (left), bison, (background centre) and aurochs (background right)

The morphology of the skull suggests that the species was adapted towards a grazing diet. [2] Tooth wear analysis suggests that the narrow-nosed rhinoceros had a variable diet tending towards grazing or mixed feeding, and clearly distinct from the more browsing focused diet inferred for S. kirchbergensis on average. [28] [29] [30] [31] Although the species has been referred to as the "steppe rhinoceros" and presumed to have had a preference for open habitats, the species was ecologically plastic, and occurred in both open grasslands and forested environments. [32] In Europe the species was found in temperate as well as cool climates, and seems to have been intolerant of cold climate conditions like those typically inhabited by the woolly rhinoceros. [19] Its dietary flexibility likely helped it tolerate cool glacial periods. [31] In the Middle East the species inhabited warm, open xeric habitats. [22]

The narrow-nosed rhinoceros probably behaved similarly to living rhinoceroses. A 1993 study suggested that the species may have been sedentary (having a home range and not migratory) and territorial, and probably engaged in ritualized confrontations between males, which may have sometimes broken out into full-on fighting. [9] A rib of a narrow-nosed rhinoceros from Neumark Nord in Germany has been found with a healed fracture, which may have been the result of such a fight. [33]

Across its range, the narrow-nosed rhinoceros lived alongside other megafauna species, including both animals living today, like red deer, fallow deer, roe deer, wild boar, wolves, brown bears, leopards, wild horse and bison, and those that are extinct, like European wild ass, aurochs, cave hyena, Irish elk, cave lions and straight-tusked elephants. [34] S. hemitoechus juveniles represented likely prey items for cave lions and cave hyenas at least on occasion. [31]

Relationship with humans

Finds at a number of sites suggest that the narrow-nosed rhinoceros was exploited for food by archaic humans. Specimens of S. hemitoechus from the Middle Pleistocene (Marine Isotope Stage 12, 478,000-424,000 years ago) Arago Cave (Caune de l'Arago) site in Southern France shows extensive evidence of butchery (presumably by Tautavel Man, which is found at the same site). The ratios of skeletal elements implies that only the parts of the body with the most meat were carried to the site. The profile of ages of rhino bones in the cave resembles natural mortality curves, suggesting that there was not selective hunting, and the fact that marks of other carnivores are rare implies that the carcasses were acquired by hunting or active scavenging. [35] At the Shishan Marsh site in the Azraq Oasis in northeast Jordan, dating to around 250,000 years ago, stone tools at the site have been found to have protein residue from the butchery of rhinoceroses. As S. hemitoechus is the only rhinoceros species known from the site, it is probable that it was the species butchered. [36]

At the late Middle Pleistocene Gran Dolina site in Spain, a handful of S. cf. hemitoechus bones display cut marks. [37] At Biache-Saint-Vaast in northeast France, dating to MIS 7, around 240,000 years ago, [38] remains of at least 33 individuals of S. hemitoechus were found in association with human artifacts, with a significant proportion displaying cut marks. The mortality profile, which is heavily skewed towards juveniles, with no old adults, may suggest selective hunting of juveniles by Neanderthals. [35] At the collapsed cave of Payre in southeast France, dating to the late Middle Pleistocene, numerous remains of rhinoceroses, primarily S. kirchbergensis along with a smaller amount of S. hemitoechus have been found, which display marks indicative of butchery and are suggested to have been accumulated at the site by Neanderthals. The abundance of teeth found at the site (though other skull material is largely absent) suggests that the Neanderthals may have been using them as tools. Mortality profiles found that young and old individuals were the most abundant at the site. [39] The late Middle Pleistocene sites of Great Yeldham and Grays Thurrock in southern Britain (both probably dating to around 300,000 years ago) where remains of S. hemitoechus have been found have also been suggested as butchery sites. [40]

A skull from Cueva Des-Cubierta in central Spain, dating to the early-mid Late Pleistocene (MIS 4-early MIS 3, ~71-43,000 years ago), exhibits fracturing and cut marks consistent with butchery by Neanderthals. The missing pieces of the skull were not found in the cave, suggesting that it had been butchered off-site. It has been proposed that the skull was kept as a hunting trophy along with the skulls of aurochs and bison. [41] Several other sites in Spain demonstrate the exploitation of S. hemitoechus by Neanderthals during the early-mid Late Pleistocene, including Navalmaíllo Rock Shelter (which represents a single individual brought to a hunting camp and butchered) [42] and Abric Romani (minimum of two individuals). [43] Exploitation of narrow-nosed rhinoceros by Neanderthals was relatively infrequent compared to other types of prey. [42]

Evolution and extinction

The earliest remains of the species in Europe date to the early-mid Middle Pleistocene, around 500,000 years ago, [44] with one of the oldest if not the oldest record in Europe being from Campagna Romana near Rome in Central Italy, dating to Marine Isotope Stage 13. [45] It is suggested to have evolved outside of Western Europe before later migrating into the region. The species may have evolved from the earlier Stephanorhinus etruscus , [27] though other authors suggest it may have evolved from S. hundsheimensis. [45] In North Africa, remains of the narrow-nosed rhinoceros are known dating between 109-53,000 years ago. [1] In Europe, the narrow-nosed rhinoceros survived latest in the southern parts of its range. The last records in Italy date to around 41,000 years ago, [31] while remains dating to 40,000 years ago are known from Bacho Kiro cave in Bulgaria. [14] In the Iberian Peninsula, the latest directly dated records of the species date to approximately 34,000 years ago (Portalón del Tejadilla and Arbreda in Spain) [46] with indirectly dated remains found at Gruta da Figueira Brava in Portugal possibly dating to somewhat later. [31] In the Levant, the species may have survived as recently as 15,500 years ago based on remains found in Hayonim Cave, Israel. [47] [22] Its extinction in southern Europe was suggested in a 2017 study to be due at least in part to climatic change causing habitat fragmentation resulting in population fragmentation, with small populations more likely to become extinct for a variety of reasons, "including loss of genetic variability, inbreeding, genetic drift, demographic fluctuations and environmental variations or natural catastrophes". The study found little evidence of systematic hunting of narrow-nosed rhinoceroses by humans at the time of its extinction. [31]

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References

  1. 1 2 3 4 5 Faith, J. Tyler (2014). "Late Pleistocene and Holocene mammal extinctions on continental Africa". Earth-Science Reviews. 128: 105–121. Bibcode:2014ESRv..128..105F. doi:10.1016/j.earscirev.2013.10.009.
  2. 1 2 3 van der Made, J. (2010). "The rhinos from the middle Pleistocene of Neumark-Nord (Saxony-Anhalt)" (PDF). Veröffentlichungen des Landesamt für Denkmalpflege und Archäologie. 62: 433–527.
  3. 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 (16 September 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 . PMID   34433011. S2CID   237273079.
  4. Saarinen, J; Eronen, J; Fortelius, M; Seppä, H; Lister, A (2016). "Patterns of diet and body mass of large ungulates from the Pleistocene of Western Europe, and their relation to vegetation". Palaeontologia Electronica. doi:10.26879/443. ISSN   1094-8074.
  5. 1 2 Pandolfi, Luca; Petronio, Carmelo (2011). "The small-sized rhinoceroses from the Late Pleistocene of Apulia (southern Italy)". Rivista Italiana di Paleontologia e Stratigrafia. 117 (3): 509–520. doi:10.13130/2039-4942/5989.
  6. Pandolfi, Luca; Tagliacozzo, Antonio (March 2015). "Stephanorhinus hemitoechus (Mammalia, Rhinocerotidae) from the Late Pleistocene of Valle Radice (Sora, Central Italy) and re-evaluation of the morphometric variability of the species in Europe". Geobios. 48 (2): 169–191. Bibcode:2015Geobi..48..169P. doi:10.1016/j.geobios.2015.02.002.
  7. 1 2 3 Giaourtsakis, Ioannis X. (2022), Vlachos, Evangelos (ed.), "The Fossil Record of Rhinocerotids (Mammalia: Perissodactyla: Rhinocerotidae) in Greece", Fossil Vertebrates of Greece Vol. 2, Cham: Springer International Publishing, pp. 409–500, doi:10.1007/978-3-030-68442-6_14, ISBN   978-3-030-68441-9 , retrieved 2024-06-13
  8. 1 2 3 4 Fortelius, M.; Mazza, P.; Sala, B. Stephanorhinus (Mammalia-Rhinocerotidae) of the Western European Pleistocene, with a revision of S. etruscus (Falconer, 1868). Palaeontogr. Ital.1993, 80, 63–155.
  9. 1 2 P. Mazza "Ethological inferences on Pleistocene rhinoceroses of Europe" Rend. Lincei Cl. Sci. Fis. Mat. Nat. Accad. Naz., 9 (1993), pp. 127-137
  10. Pandolfi, Luca; Tagliacozzo, Antonio (1 March 2015). "Stephanorhinus hemitoechus (Mammalia, Rhinocerotidae) from the Late Pleistocene of Valle Radice (Sora, Central Italy) and re-evaluation of the morphometric variability of the species in Europe". Geobios . 48 (2): 169–191. Bibcode:2015Geobi..48..169P. doi:10.1016/j.geobios.2015.02.002. ISSN   0016-6995 . Retrieved 13 January 2024 via Elsevier Science Direct.
  11. 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
  12. Pokines, James T.; Lister, Adrian M.; Ames, Christopher J. H.; Nowell, April; Cordova, Carlos E. (March 2019). "Faunal remains from recent excavations at Shishan Marsh 1 (SM1), a Late Lower Paleolithic open-air site in the Azraq Basin, Jordan". Quaternary Research . 91 (2): 768–791. Bibcode:2019QuRes..91..768P. doi:10.1017/qua.2018.113. ISSN   0033-5894. S2CID   134216492.
  13. Sanisidro, Oscar; Cantalapiedra, Juan L. (2022-08-03). "The Rhinocerotidae fossil record in the Iberian Peninsula". Historical Biology. 34 (8): 1591–1610. Bibcode:2022HBio...34.1591S. doi:10.1080/08912963.2022.2051502. ISSN   0891-2963.
  14. 1 2 Stuart, A.J., Lister, A.M., 2007. Patterns of Late Quaternary megafaunal extinctions in Europe and northern Asia. In: Kahlke, R.-D., Maul, L.C., Mazza, P. (Eds.), Late Neogene and Quaternary Biodiversity and Evolution: Regional Developments and Interregional Correlations Vol. II, Proceedings of the 18th International Senckenberg Conference (VI International Palaeontological Colloquium in Weimar). Courier Forschungsinstitut Senckenberg 259, pp. 287-297.
  15. 1 2 3 Giaourtsakis, Ioannis X. (2022), Vlachos, Evangelos (ed.), "The Fossil Record of Rhinocerotids (Mammalia: Perissodactyla: Rhinocerotidae) in Greece", Fossil Vertebrates of Greece Vol. 2, Cham: Springer International Publishing, pp. 409–500, doi:10.1007/978-3-030-68442-6_14, ISBN   978-3-030-68441-9 , retrieved 2023-11-19
  16. 1 2 Tsoukala, E., and Guérin, C. (2016). The Rhinocerotidae and Suidae of the Middle Pleistocene from Petralona cave (Macedonia, Greece). Acta Zool. Bulgarica 68, 243–264.
  17. Auguste, Patrick (2009-12-01). "Évolution des peuplements mammaliens en Europe du Nord-Ouest durant le Pléistocène moyen et supérieur. Le cas de la France septentrionale". Quaternaire. 20 (4): 527–550. doi:10.4000/quaternaire.5361. ISSN   1142-2904.
  18. Boylan, Patrick J. (July 1981). "A New Revision of the Pleistocene Mammalian Fauna of Kirkdale Cave, Yorkshire". Proceedings of the Yorkshire Geological Society. 43 (3): 253–280. Bibcode:1981PYGS...43..253B. doi:10.1144/pygs.43.3.253. ISSN   0044-0604.
  19. 1 2 Stefaniak, Krzysztof; Kovalchuk, Oleksandr; Ratajczak-Skrzatek, Urszula; Kropczyk, Aleksandra; Mackiewicz, Paweł; Kłys, Grzegorz; Krajcarz, Magdalena; Krajcarz, Maciej T.; Nadachowski, Adam; Lipecki, Grzegorz; Karbowski, Karol; Ridush, Bogdan; Sabol, Martin; Płonka, Tomasz (November 2023). "Chronology and distribution of Central and Eastern European Pleistocene rhinoceroses (Perissodactyla, Rhinocerotidae) – A review". Quaternary International. 674–675: 87–108. Bibcode:2023QuInt.674...87S. doi:10.1016/j.quaint.2023.02.004. ISSN   1040-6182.
  20. Pushkina, Diana (July 2007). "The Pleistocene easternmost distribution in Eurasia of the species associated with the Eemian Palaeoloxodon antiquus assemblage". Mammal Review. 37 (3): 224–245. doi:10.1111/j.1365-2907.2007.00109.x. ISSN   0305-1838.
  21. Chelli-Cheheb, Razika; Merzoug, Souhila (2023-01-02). "The Aterian site of Phacochères (northern Algeria): a zooarchaeological perspective". Azania: Archaeological Research in Africa. 58 (1): 5–33. doi:10.1080/0067270X.2023.2187559. ISSN   0067-270X.
  22. 1 2 3 Marom, Nimrod; Lazagabaster, Ignacio A.; Shafir, Roee; Natalio, Filipe; Eisenmann, Vera; Horwitz, Liora Kolska (May 2022). "The Late Middle Pleistocene mammalian fauna of Oumm Qatafa Cave, Judean Desert: taxonomy, taphonomy and palaeoenvironment". Journal of Quaternary Science. 37 (4): 612–638. Bibcode:2022JQS....37..612M. doi:10.1002/jqs.3414. ISSN   0267-8179. PMC   9314136 . PMID   35915614.
  23. Pokines, James T.; Lister, Adrian M.; Ames, Christopher J. H.; Nowell, April; Cordova, Carlos E. (March 2019). "Faunal remains from recent excavations at Shishan Marsh 1 (SM1), a Late Lower Paleolithic open-air site in the Azraq Basin, Jordan". Quaternary Research. 91 (2): 768–791. Bibcode:2019QuRes..91..768P. doi:10.1017/qua.2018.113. ISSN   0033-5894.
  24. Stewart, Mathew; Louys, Julien; Price, Gilbert J.; Drake, Nick A.; Groucutt, Huw S.; Petraglia, Michael D. (May 2019). "Middle and Late Pleistocene mammal fossils of Arabia and surrounding regions: Implications for biogeography and hominin dispersals". Quaternary International. 515: 12–29. Bibcode:2019QuInt.515...12S. doi:10.1016/j.quaint.2017.11.052.
  25. Vahdati Nasab, Hamed; Berillon, Gilles; Hashemi, Seyyed Milad; Bahain, Jean-Jacques; Sévêque, Noémie; Jayez, Mozhgan; Bonilauri, Stéphanie; Jamet, Guillaume; Kharazian, Mohammad Akhavan; Nateghi, Asghar; Abdollahi, Alieh; Antoine, Pierre; Beheshti, Iraj; Boulbes, Nicolas; Chapon-Sao, Cécile (2024-05-23). "Qaleh Kurd Cave (Qazvin, Iran): Oldest Evidence of Middle Pleistocene Hominin Occupations and a Human Deciduous Tooth in the Iranian Central Plateau". Journal of Paleolithic Archaeology. 7 (1). doi:10.1007/s41982-024-00180-4. ISSN   2520-8217.
  26. Mashkour, M.; Monchot, H.; Trinkaus, E.; Reyss, J.-L.; Biglari, F.; Bailon, S.; Heydari, S.; Abdi, K. (November 2009). "Carnivores and their prey in the Wezmeh Cave (Kermanshah, Iran): a Late Pleistocene refuge in the Zagros". International Journal of Osteoarchaeology. 19 (6): 678–694. doi:10.1002/oa.997. ISSN   1047-482X.
  27. 1 2 Van der Made, Jan; Torres, Trinidad; Ortiz, Jose Eugenio; Moreno-Pérez, Laura; Fernández-Jalvo, Yolanda (2016), Fernández-Jalvo, Yolanda; King, Tania; Yepiskoposyan, Levon; Andrews, Peter (eds.), "The New Material of Large Mammals from Azokh and Comments on the Older Collections", Azokh Cave and the Transcaucasian Corridor, Cham: Springer International Publishing, pp. 117–162, doi:10.1007/978-3-319-24924-7_6, ISBN   978-3-319-24922-3 , retrieved 2024-06-27
  28. Fortelius, M.; Mazza, P.; Sala, B. (1993). "Stephanorhinus (Mammalia: Rhinocerotidae) of the Western European Pleistocene, with a revision of S. etruscus (Falconer, 1868)". Palaeontographia Italica, Pisa. 80: 63–155.
  29. Salari, L. (2019). "The Late Pleistocene faunal assemblage from Cava Muracci (Latium, Italy): Palaeoenvironmental implications for coastal central Italy during MIS 3". Comptes Rendus Palevol. 18 (1): 51–71. Bibcode:2019CRPal..18...51G. doi:10.1016/j.crpv.2018.04.006. S2CID   135071773.
  30. Rivals, Florent; Lister, Adrian M. (August 2016). "Dietary flexibility and niche partitioning of large herbivores through the Pleistocene of Britain". Quaternary Science Reviews. 146: 116–133. Bibcode:2016QSRv..146..116R. doi:10.1016/j.quascirev.2016.06.007.
  31. 1 2 3 4 5 6 Pandolfi, Luca; Boscato, Paolo; Crezzini, Jacopo; Gatta, Maurizio; Moroni, Adriana; Rolfo, Mario; Tagliacozzo, Antonio (July 2017). "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). 123: 177–192. doi:10.13130/2039-4942/8300.
  32. 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.
  33. J. van der Made, R. Grübe The rhinoceroses from Neumark-Nord and their nutrition (Die Nashörner von Neumark-Nord und ihre Ernährung) H. Meller (Ed.), Elefantenreich – Eine Fossilwelt in Europa. Landesamt. Denkmalpflege und Archäologie Sachsen-Anhalt und Landesmuseum Vorgeschichte (2010), pp. 383-394
  34. Davoli, Marco; Monsarrat, Sophie; Pedersen, Rasmus Østergaard; Scussolini, Paolo; Karger, Dirk Nikolaus; Normand, Signe; Svenning, Jens-Christian (January 2024). "Megafauna diversity and functional declines in Europe from the Last Interglacial to the present". Global Ecology and Biogeography. 33 (1): 34–47. Bibcode:2024GloEB..33...34D. doi:10.1111/geb.13778. hdl: 11573/1714498 . ISSN   1466-822X.
  35. 1 2 Chen, Xi; Moigne, Anne-Marie (November 2018). "Rhinoceros ( Stephanorhinus hemitoechus ) exploitation in Level F at the Caune de l'Arago (Tautavel, Pyrénéés-Orientales, France) during MIS 12". International Journal of Osteoarchaeology. 28 (6): 669–680. doi:10.1002/oa.2682. S2CID   80923883.
  36. Nowell, A.; Walker, C.; Cordova, C.E.; Ames, C.J.H.; Pokines, J.T.; Stueber, D.; DeWitt, R.; al-Souliman, A.S.A. (September 2016). "Middle Pleistocene subsistence in the Azraq Oasis, Jordan: Protein residue and other proxies". Journal of Archaeological Science. 73: 36–44. Bibcode:2016JArSc..73...36N. doi:10.1016/j.jas.2016.07.013.
  37. Blasco, Ruth; Rosell, Jordi; Fernández Peris, Josep; Arsuaga, Juan Luis; Bermúdez de Castro, José María; Carbonell, Eudald (June 2013). "Environmental availability, behavioural diversity and diet: a zooarchaeological approach from the TD10-1 sublevel of Gran Dolina (Sierra de Atapuerca, Burgos, Spain) and Bolomor Cave (Valencia, Spain)". Quaternary Science Reviews. 70: 124–144. Bibcode:2013QSRv...70..124B. doi:10.1016/j.quascirev.2013.03.008.
  38. Bahain, Jean-Jacques; Falguères, Christophe; Laurent, Michel; Dolo, Jean-Michel; Shao, Qingfeng; Auguste, Patrick; Tuffreau, Alain (October 2015). "ESR/U-series dating of faunal remains from the paleoanthropological site of Biache-Saint-Vaast (Pas-de-Calais, France)". Quaternary Geochronology. 30: 541–546. Bibcode:2015QuGeo..30..541B. doi:10.1016/j.quageo.2015.02.020.
  39. Daujeard, Camille; Daschek, Eva J.; Patou‑Mathis, Marylène; Moncel, Marie‑Hélène (2018-09-01). "Les néandertaliens de Payre (Ardèche, France) ont-ils chassé le rhinocéros?" [Did the Neanderthals of Payre (Ardèche, France) hunt rhinoceros?]. Quaternaire (in French). 29 (3): 217–231. doi:10.4000/quaternaire.10196. ISSN   1142-2904.
  40. Parfitt, Simon A. (2022-09-12). "A Middle Pleistocene Butchery Site at Great Yeldham, Essex, UK: Identifying Butchery Strategies and Implications for Mammalian Faunal History". Journal of Paleolithic Archaeology. 5 (1): 11. Bibcode:2022JPalA...5...11P. doi: 10.1007/s41982-022-00122-y . ISSN   2520-8217.
  41. Baquedano, Enrique; Arsuaga, Juan L.; Pérez-González, Alfredo; Laplana, César; Márquez, Belén; Huguet, Rosa; Gómez-Soler, Sandra; Villaescusa, Lucía; Galindo-Pellicena, M. Ángeles; Rodríguez, Laura; García-González, Rebeca; Ortega, M.-Cruz; Martín-Perea, David M.; Ortega, Ana I.; Hernández-Vivanco, Lucía (March 2023). "A symbolic Neanderthal accumulation of large herbivore crania". Nature Human Behaviour. 7 (3): 342–352. doi:10.1038/s41562-022-01503-7. ISSN   2397-3374. PMC   10038806 . PMID   36702939.
  42. 1 2 Moclán, Abel; Huguet, Rosa; Márquez, Belén; Laplana, César; Galindo-Pellicena, María Ángeles; García, Nuria; Blain, Hugues-Alexandre; Álvarez-Lao, Diego J.; Arsuaga, Juan Luis; Pérez-González, Alfredo; Baquedano, Enrique (October 2021). "A neanderthal hunting camp in the central system of the Iberian Peninsula: A zooarchaeological and taphonomic analysis of the Navalmaíllo Rock Shelter (Pinilla del Valle, Spain)". Quaternary Science Reviews. 269: 107142. Bibcode:2021QSRv..26907142M. doi:10.1016/j.quascirev.2021.107142.
  43. Rosell, Jordi; Blasco, Ruth; Huguet, Rosa; Cáceres, Isabel; Saladié, Palmira; Rivals, Florent; Bennàsar, Maria; Bravo, Pilar; Campeny, Gerard (2012), Carbonell i Roura, Eudald (ed.), "Occupational Patterns and Subsistence Strategies in Level J of Abric Romaní", High Resolution Archaeology and Neanderthal Behavior, Dordrecht: Springer Netherlands, pp. 313–372, doi:10.1007/978-94-007-3922-2_8, ISBN   978-94-007-3921-5 , retrieved 2024-06-14
  44. Stefanelli, Dario; Mecozzi, Beniamino; Marino, Maria; Sardella, Raffaele; Breda, Marzia (2024-03-28). "The postcranial variability of Quaternary European rhinoceroses: the case study of Stephanorhinus hundsheimensis from the Middle Pleistocene site of Contrada Monticelli (Apulia, southern Italy)". Historical Biology: 1–20. doi:10.1080/08912963.2024.2328276. ISSN   0891-2963.
  45. 1 2 Pandolfi, L., Gaeta, M. & Petronio, C. The skull of Stephanorhinus hemitoechus (Mammalia, Rhinocerotidae) from the Middle Pleistocene of Campagna Romana (Rome, central Italy): Biochronological and paleobiogeographic implications. Bull. Geosci.88(1), 51–62 (2013).
  46. Sala, Nohemi; Pablos, Adrián; Rodríguez-Hidalgo, Antonio; Arriolabengoa, Martin; Alcaraz-Castaño, Manuel; Cubas, Miriam; Posth, Cosimo; Nägele, Kathrin; Pantoja-Pérez, Ana; Arlegi, Mikel; Rodríguez-Almagro, Manuel; Conde-Valverde, Mercedes; Cuenca-Bescós, Gloria; Arribas, Alfonso; Gómez-Olivencia, Asier (February 2021). "Cueva de los Torrejones revisited. New insights on the paleoecology of inland Iberia during the Late Pleistocene". Quaternary Science Reviews. 253: 106765. Bibcode:2021QSRv..25306765S. doi: 10.1016/j.quascirev.2020.106765 .
  47. Dembitzer, Jacob; Barkai, Ran; Ben-Dor, Miki; Meiri, Shai (January 2022). "Levantine overkill: 1.5 million years of hunting down the body size distribution". Quaternary Science Reviews. 276: 107316. Bibcode:2022QSRv..27607316D. doi:10.1016/j.quascirev.2021.107316.