Biology:Steppe mammoth

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Short description: Extinct species of mammal

Steppe mammoth
Temporal range: Early-Middle Pleistocene 1.7–0.2 Ma
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Steppe mammoth.jpg
Skeleton
Scientific classification edit
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Proboscidea
Family: Elephantidae
Genus: †Mammuthus
Species:
†M. trogontherii
Binomial name
†Mammuthus trogontherii
(Pohlig, 1885) [1]
Synonyms
  • Elephas trogontherii Pohlig, 1885
  • Euelephas protomammonteus Matsumoto, 1924
  • Mammuthus protomammonteus (Matsumoto, 1924)
  • Mammuthus sungari Zhou, 1959
  • Mammuthus trogontherii chosaricus Dubrovo, 1966
  • Mammuthus armeniacus Falconer, 1857

Mammuthus trogontherii, sometimes called the steppe mammoth, is an extinct species of mammoth that ranged over most of northern Eurasia during the Early and Middle Pleistocene, approximately 1.7 million-200,000 years ago. One of the largest mammoth species, it evolved in East Asia during the Early Pleistocene, around 1.8 million years ago, before migrating into North America around 1.5 million years ago, and into Europe during the Early/Middle Pleistocene transition, around 1 to 0.7 million years ago. It was the ancestor of the woolly mammoth and Columbian mammoth of the later Pleistocene.

Taxonomy

There was historically confusion about the correct scientific name for the steppe mammoth, either Mammuthus armeniacus (Falconer 1857) or Mammuthus trogontherii (Pohlig 1885). Falconer used material from Asian sources while Pohlig worked with fossil remains from Europe and both names appear in scientific publications, adding to the confusion. A first taxonomical overhaul was done by Maglio (1973) who decided that both names were synonyms, armeniacus being the older, hence the preferred name.[2] However, in Shoshani & Tassy (1996) it was decided that the description of Pohlig prevailed, and consequently the correct name for the steppe mammoth is M. trogontherii.[3] The status of Mammuthus armeniacus as a synonym of Mammuthus trogontherii has been supported by most recent authors.[4] The type specimens of the species are molars from the Süssenborn (also spelled Süßenborn) locality in Germany, dating to the early Middle Pleistocene, (MIS 16, approximately 676-621,000 years ago).[5]

Several Japanese mammoth varieties from the early Pleistocene have been named, but all are now thought to be synonyms of M. trogontherii.[6]

Description

Skeletal diagram of the 3.89 metre tall Zhalainuoer III specimen, including side on view (centre), top-down (above) and from the front minus the head (left)
Size comparison of the fragmentary "Mosbach mammoth" estimated to be 4.5 metres tall.

Mammuthus trogontherii was one of the largest mammoth species, with males on average being about 4 m (13.1 ft) tall at the shoulders and about 11 tonnes (10.8 long tons; 12.1 short tons) in weight and females on average being about 3.7 m (12.1 ft) tall at the shoulders and about 9.5 tonnes (9.3 long tons; 10.5 short tons) in weight, considerably exceeding the size of modern elephants.[7][8] A largely complete specimen (Zhalainuoer III) from Inner Mongolia, China, was estimated to have had a shoulder height of around 3.69 m (12.1 ft) measured at the top of the scapula, which represents a flesh shoulder height of 3.89 m (12.8 ft), with a body mass estimated via volumetric analysis at 10.4 tonnes (10.2 long tons; 11.5 short tons).[9][7] A larger bull, (Azov I), estimated to be 3.96 m (13.0 ft) tall at the shoulder (previously erroneously estimated as 4.5 m (15 ft) due to incorrect mounting) was estimated to weigh 11.5 tonnes (11.3 long tons; 12.7 short tons) via volumetric analysis. Another individual represented by a single giant humerus 1.46 m (4.8 ft) long and an associated pelvis found in Mosbach Sande, Germany, is estimated to have had a shoulder height of 4.5 metres (14.8 ft) and a weight of 14.3 tonnes (14.1 long tons; 15.8 short tons) via regression analysis.[7]

Jaw with molar teeth

The skull was high-domed and short, and bore twisted tusks. The lower jaw was short and deep.[9] The number of lamellae on the third molars is around 18–22, noticeably lower than the number typically present in M. primigenius (though some European specimens of M. primigenius have counts which overlap with those of M. trogontherii).[5] The body has around 19 thoracic vertebrae and 5 or 6 sacral vertebrae, with the first few thoracic vertebrae having long neural spines.[9] The tusks were proportionally large, among the largest known among proboscideans, with one large tusk from the Kostolac Basin in Serbia measuring 4.2 metres (14 ft) in length, with an estimated mass of 213 kilograms (470 lb).[10]

Sequenced genomes suggests that Early Pleistocene M. trogontherii specimens from Siberia, around 1 million years old, had already developed many of the genetic changes thought to be responsible for traits that were adaptations for living in cold environments characteristic of woolly mammoths.[11] Due to the cold climates it inhabited and short tail, Mammuthus trogontherii is suggested to have borne a coat of fur, which was probably somewhat thinner than that of the woolly mammoth.[9]

Ecology

Steppe mammoths are thought to have been grazers to grazing-predominant mixed feeders, distinct from Mammuthus meridionalis, whose diet was highly variable but on average included more browse.[12] It was often associated with cold steppe environments, as its common name would suggest, but was not confined to them, as evidenced by the early Middle Pleistocene West Runton Mammoth specimen from Norfolk, England, which was associated with a temperate forested environment.[13]

Evolution

M. trogontherii is suggested to have derived from Mammuthus meridionalis. The oldest records M. trogontherii are known from China, around 1.7 million years old, from the Nihewan Formation near Majuangou, Hebei. Steppe mammoths arrived in North America across Beringia around 1.5-1.3 million years ago, giving rise to the Columbian mammoth (the ancestor was previously thought to be M. meridionalis but this was due to misinterpretation of tooth wear patterns).[14][11] Steppe mammoths replaced Mammuthus meridionalis between 1–0.7 million years ago in Europe, in a complex diachronous mosaic pattern, coincident with the arrival of the straight-tusked elephant (Palaeoloxodon antiquus) to Europe.[15] European populations of M. trogontherii experienced a persistent size reduction towards the end of the Middle Pleistocene, from around 400,000 years ago onwards.[15][11] Mammoths with M. primigenius (woolly mammoth)-type molar morphology had emerged in Northeast Siberia from M. trogontherii by around 600-500,000 years ago, reaching the typical morphology of M. primigenius around 400,000 years ago. Mammoths with M. primigenius type molar morphology displaced those of M. trogontherii type in Europe over the course of the late Middle Pleistocene, which was largely complete by 200,000 years ago (~MIS 7/6 boundary) in a protracted highly complex pattern including some molars with intermediate morphology between the two species that likely reflects gene flow from Siberian woolly mammoths into European M. trogontherii.[5] The replacement of European M. trogontherii by woolly mammoths is widely considered to mark the extinction of the species.[11]

Relationship with humans

At the Bełchatów coal mine in Poland, dating to the late Middle Pleistocene (in the interglacial period of either MIS 11 or MIS 9, around 425-300,000 years ago), remains of M. trogontherii have been found with cut marks, suggested to represent evidence of butchery by archaic humans.[16] Sites with evidence of both humans and M. trogontherii in Europe are rare, especially compared to the contemporaneous straight-tusked elephant, which is suggested to be the result of them primarily occupying different habitats.[17]

See also

References

  1. ↑ Todd, N. E. (January 2010). "New Phylogenetic Analysis of the Family Elephantidae Based on Cranial-Dental Morphology". The Anatomical Record (Wiley-Liss, Inc.) 293 (1): 74–90. doi:10.1002/ar.21010. PMID 19937636. 
  2. ↑ Maglio, V. J. (1973). "Origin and Evolution of the Elephantidae". Trans Am Philos Soc 63 (3): 1–149. doi:10.2307/1006229. 
  3. ↑ Shoshani, J.; Tassy, P., eds (1996). The Proboscidea : Evolution and Palaeoecology of Elephants and Their Relatives. Oxford: Oxford University Press. ISBN 0-19-854652-1. 
  4. ↑ Athanassiou, Athanassios (March 2012). "A skeleton of Mammuthus trogontherii (Proboscidea, Elephantidae) from NW Peloponnese, Greece" (in en). Quaternary International 255: 9–28. doi:10.1016/j.quaint.2011.03.030. https://linkinghub.elsevier.com/retrieve/pii/S1040618211001765. 
  5. ↑ 5.0 5.1 5.2 Lister, Adrian M. (October 2022). "Mammoth evolution in the late Middle Pleistocene: The Mammuthus trogontherii-primigenius transition in Europe" (in en). Quaternary Science Reviews 294: 107693. doi:10.1016/j.quascirev.2022.107693. Bibcode: 2022QSRv..29407693L. https://linkinghub.elsevier.com/retrieve/pii/S0277379122003249. 
  6. ↑ van de Greer, Alexandra; Lyras, George; de Vos, John; Dermitzakis, Michael (2011) (in da, en). Evolution of Island Mammals: Adaptation and Extinction of Placental Mammals on Islands. John Wiley & Sons. p. 240. ISBN 978-1444391282. https://books.google.com/books?id=JmSsNuwMAxgC&q=Mammuthus+paramammonteus. 
  7. ↑ 7.0 7.1 7.2 Larramendi, A. (2016). "Shoulder height, body mass and shape of proboscideans". Acta Palaeontologica Polonica 61. doi:10.4202/app.00136.2014. https://www.app.pan.pl/archive/published/app61/app001362014.pdf. 
  8. ↑ Larramendi, Asier; Palombo, Maria Rita; Marano, Federica (2017). "Reconstructing the life appearance of a Pleistocene giant: size, shape, sexual dimorphism and ontogeny of Palaeoloxodon antiquus (Proboscidea: Elephantidae) from Neumark-Nord 1 (Germany)". Bollettino della Società Paleontologica Italiana (3): 299–317. doi:10.4435/BSPI.2017.29. ISSN 0375-7633. https://www.paleoitalia.it/wp-content/uploads/2021/08/01_Larramendi_et_al_2017_BSPI_563.pdf. 
  9. ↑ 9.0 9.1 9.2 9.3 Larramendi, Asier (June 2015). "Skeleton of a Late Pleistocene steppe mammoth (Mammuthus trogontherii) from Zhalainuoer, Inner Mongolian Autonomous Region, China" (in en). Paläontologische Zeitschrift 89 (2): 229–250. doi:10.1007/s12542-014-0222-8. ISSN 0031-0220. http://link.springer.com/10.1007/s12542-014-0222-8. 
  10. ↑ Larramendi, Asier (2023-12-10). "Estimating tusk masses in proboscideans: a comprehensive analysis and predictive model" (in en). Historical Biology: 1–14. doi:10.1080/08912963.2023.2286272. ISSN 0891-2963. https://www.tandfonline.com/doi/full/10.1080/08912963.2023.2286272. 
  11. ↑ 11.0 11.1 11.2 11.3 van der Valk, Tom; Pečnerová, Patrícia; Díez-del-Molino, David; Bergström, Anders; Oppenheimer, Jonas; Hartmann, Stefanie; Xenikoudakis, Georgios; Thomas, Jessica A. et al. (17 February 2021). "Million-year-old DNA sheds light on the genomic history of mammoths" (in en). Nature 591 (7849): 265–269. doi:10.1038/s41586-021-03224-9. ISSN 1476-4687. PMID 33597750. Bibcode: 2021Natur.591..265V. 
  12. ↑ Rivals, Florent; Semprebon, Gina M.; Lister, Adrian M. (September 2019). "Feeding traits and dietary variation in Pleistocene proboscideans: A tooth microwear review" (in en). Quaternary Science Reviews 219: 145–153. doi:10.1016/j.quascirev.2019.06.027. Bibcode: 2019QSRv..219..145R. https://linkinghub.elsevier.com/retrieve/pii/S0277379119302641. 
  13. ↑ Lister, Adrian M.; Stuart, Anthony J. (December 2010). "The West Runton mammoth (Mammuthus trogontherii) and its evolutionary significance" (in en). Quaternary International 228 (1-2): 180–209. doi:10.1016/j.quaint.2010.07.032. https://linkinghub.elsevier.com/retrieve/pii/S1040618210003149. 
  14. ↑ Lister, A. M.; Sher, A. V. (2015-11-13). "Evolution and dispersal of mammoths across the Northern Hemisphere" (in en). Science 350 (6262): 805–809. doi:10.1126/science.aac5660. ISSN 0036-8075. PMID 26564853. Bibcode: 2015Sci...350..805L. 
  15. ↑ 15.0 15.1 Lister, Adrian M.; Sher, Andrei V.; van Essen, Hans; Wei, Guangbiao (January 2005). "The pattern and process of mammoth evolution in Eurasia". Quaternary International 126-128: 49–64. doi:10.1016/j.quaint.2004.04.014. ISSN 1040-6182. Bibcode: 2005QuInt.126...49L. 
  16. ↑ Haynes, Gary (March 2022). "Late Quaternary Proboscidean Sites in Africa and Eurasia with Possible or Probable Evidence for Hominin Involvement" (in en). Quaternary 5 (1): 18. doi:10.3390/quat5010018. ISSN 2571-550X. 
  17. ↑ Konidaris, George E.; Tourloukis, Vangelis (2021-04-14). "Proboscidea-Homo interactions in open-air localities during the Early and Middle Pleistocene of western Eurasia: a palaeontological and archaeolocigal perspective" (in en). Human-Elephant Interactions: From Past to Present. doi:10.15496/publikation-55599. https://publikationen.uni-tuebingen.de/xmlui/handle/10900/114224. 

Further reading

  • Benes, Josef (1979). Prehistoric Animals and Plants. London: Hamlyn. p. 271. ISBN 0-600-30341-1. 
  • Augusti, Jordi; Anton, Mauricio (2002). Mammoths, Sabertooths and Hominids 65 Million Years of Mammalian Evolution in Europe. Columbia University Press. ISBN 0-231-11640-3. 
  • Lister, Adrian; Bahn, Paul (1997) (in de). Mammuts : Riesen der Eiszeit. Sigmaringen: Thorbecke Verlag. ISBN 3-7995-9050-1. 
  • Mol, Dick; Lacombat, Frédéric (2010). Mammoths & Mastodons of Haute-Loire. Drukware. p. 271. ISBN 978-2-911794-97-1.  (English and French)

External links

Wikidata ☰ Q113284 entry



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