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The steppe mammoth had a short skull compared with ''[[Mammuthus meridionalis|M. meridionalis]]'' as well as a smaller jaw. The males had spiral tusks with a recurved tip that could grow as long as {{convert|4.9|m|ft}} in old bulls; females on the other hand had thinner and slightly curved tusks.
The steppe mammoth had a short skull compared with ''[[Mammuthus meridionalis|M. meridionalis]]'' as well as a smaller jaw. The males had spiral tusks with a recurved tip that could grow as long as {{convert|4.9|m|ft}} in old bulls; females on the other hand had thinner and slightly curved tusks.


With individuals not uncommonly reaching {{convert|4|m|ft|1|abbr=on}} tall at the shoulders, ''M. trogontherii'' was considerably larger than modern elephants.<ref name="proboscideans" /> A skeleton mounted on the Azov Museum reaches {{convert|4.5|m|ft|1|abbr=on}} at the shoulder, though this figure might be overestimated because the vertebrae have been placed between the tips of the shoulder blades.<ref>{{citation |last1=Mol |first1=Dick |last2=van den Bergh |first2=Gert D. |last3=de Vos |first3=John |title=Fossil Proboscideans from The Netherlands, the North Sea and the Oosterschelde Estuary |url=http://www.hetnatuurhistorisch.nl/fileadmin/user_upload/documents-nmr/Publicaties/Deinsea/Deinsea_06/Deinsea_6_10_Mol__.pdf |volume=6 |work=Mammoths and the Mammoth Fauna: Studies of an Extinct Ecosystem |publisher=Deinsea |accessdate=15 August 2016 |date=May 17, 1999 |page=135 }}</ref> Another individual represented by a single giant humerus {{convert|1.46|m|ft|abbr=on}} long and an associated pelvis found in Mosbach Sande, [[Germany]], is estimated to have had a shoulder height of {{Convert|4.5|m|ft|1}} and a weight of {{Convert|14.3|t}}.<ref name="proboscideans">{{Cite journal | last1 = Larramendi | first1 = A. | year = 2016 | title = Shoulder height, body mass and shape of proboscideans | journal = Acta Palaeontologica Polonica | volume = 61 | doi = 10.4202/app.00136.2014 | url = https://www.app.pan.pl/archive/published/app61/app001362014.pdf | doi-access = free }}</ref> 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.<ref name=":0" />
With individuals not uncommonly reaching {{convert|4|m|ft|1|abbr=on}} tall at the shoulders, ''M. trogontherii'' was considerably larger than modern elephants.<ref name="proboscideans" /> A skeleton mounted on the Azov Museum reaches {{convert|4.5|m|ft|1|abbr=on}} at the shoulder, though this figure might be overestimated because the vertebrae have been placed between the tips of the shoulder blades.<ref>{{citation |last1=Mol |first1=Dick |last2=van den Bergh |first2=Gert D. |last3=de Vos |first3=John |title=Fossil Proboscideans from The Netherlands, the North Sea and the Oosterschelde Estuary |url=http://www.hetnatuurhistorisch.nl/fileadmin/user_upload/documents-nmr/Publicaties/Deinsea/Deinsea_06/Deinsea_6_10_Mol__.pdf |volume=6 |work=Mammoths and the Mammoth Fauna: Studies of an Extinct Ecosystem |publisher=Deinsea |accessdate=15 August 2016 |date=May 17, 1999 |page=135 }}</ref> A largely complete specimen (Zhalainuoer III) from Inner Mongolia, China, was estimated to have had a shoulder height of around {{convert|3.69|m|ft|abbr=on}} metres measured at the top of the [[scapula]], which represents a flesh shoulder height of {{convert|3.89|m|ft|abbr=on}}, with a body mass estimated via volumetric analysis at 10.5 tons.<ref name=":1">{{Cite journal |last=Larramendi |first=Asier |date=2015-06 |title=Skeleton of a Late Pleistocene steppe mammoth (Mammuthus trogontherii) from Zhalainuoer, Inner Mongolian Autonomous Region, China |url=http://link.springer.com/10.1007/s12542-014-0222-8 |journal=Paläontologische Zeitschrift |language=en |volume=89 |issue=2 |pages=229–250 |doi=10.1007/s12542-014-0222-8 |issn=0031-0220}}</ref> Another individual represented by a single giant humerus {{convert|1.46|m|ft|abbr=on}} long and an associated pelvis found in Mosbach Sande, [[Germany]], is estimated to have had a shoulder height of {{Convert|4.5|m|ft|1}} and a weight of {{Convert|14.3|t}} via [[regression analysis]].<ref name="proboscideans">{{Cite journal | last1 = Larramendi | first1 = A. | year = 2016 | title = Shoulder height, body mass and shape of proboscideans | journal = Acta Palaeontologica Polonica | volume = 61 | doi = 10.4202/app.00136.2014 | url = https://www.app.pan.pl/archive/published/app61/app001362014.pdf | doi-access = free }}</ref> 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.<ref name=":0" /> 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.<ref name=":1" />


== Ecology ==
== Ecology ==

Revision as of 07:32, 27 June 2023

Not to be confused with Mammoth steppe.

Steppe mammoth
Temporal range: Early-Mid Pleistocene 1.8–0.2 Ma
↓
 Possible Late Pleistocene record
Skeleton
Skeletal diagram of the 3.89 metre tall Zhalainuoer III specimen
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Proboscidea
Family: Elephantidae
Genus: †Mammuthus
Species:
†M. trogontherii
Binomial name
†Mammuthus trogontherii
Synonyms
  • Mammuthus sungari Zhou, M.Z, 1959
  • Mammuthus protomammonteus
  • Mammuthus trogontherii chosaricus

The steppe mammoth (Mammuthus trogontherii, sometimes Mammuthus armeniacus) is an extinct species of mammoth that ranged over most of northern Eurasia during the Early and Middle Pleistocene, approximately 1.8 million-200,000 years ago. 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. Populations of steppe mammoth may have persisted in northern China and Mongolia as late as 33,000 years ago.

Taxonomy

Molar of Mammuthus trogontherii at the National Museum (Prague)

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] It is unclear whether both forms are indeed identical and authors tend to use the name M. trogontherii for European material and M. armeniacus for Asian remains.

Several Japanese mammoth varieties from the early Pleistocene have been named, but all are now thought to be synonyms of M. trogontherii.[4]

Genetics

DNA has been sequenced from two Siberian mammoth molars (which judging by morphology belong to the steppe mammoth); at over a million years old, this is the oldest ancient DNA ever recovered. The two genomes recovered belong to separate and distinct lineages: one is ancestral to the woolly mammoth, the other is a previously-unknown lineage. The latter lineage hybridized with woolly mammoths at least 420,000 years ago, giving rise to the Columbian mammoth.[5]

Description

Size comparison of the fragmentary "Mosbach mammoth" estimated to be 4.5 metres tall.

The steppe mammoth had a short skull compared with M. meridionalis as well as a smaller jaw. The males had spiral tusks with a recurved tip that could grow as long as 4.9 metres (16 ft) in old bulls; females on the other hand had thinner and slightly curved tusks.

With individuals not uncommonly reaching 4 m (13.1 ft) tall at the shoulders, M. trogontherii was considerably larger than modern elephants.[6] A skeleton mounted on the Azov Museum reaches 4.5 m (14.8 ft) at the shoulder, though this figure might be overestimated because the vertebrae have been placed between the tips of the shoulder blades.[7] 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) metres 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 10.5 tons.[8] 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.[6] 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.[5] 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.[8]

Ecology

Steppe mammoths are thought to have been grazers to grazing-predominant mixed feeders, distinct from Mammuthus meridionalis, whos diet was highly variable but on average included more browse.[9]

Discovery

Fossilized teeth are recovered, but skeletal parts are rare. The most complete skeleton of a steppe mammoth yet found was discovered in 1996 in Kikinda, Serbia. It was mounted, and put on display in 2005. The specimen is a female, which was about 3.7 metres (12.1 ft) high, 7 metres (23 ft) in length and with 2.7 metres (8.9 ft) long tusks.[10]

Another quite complete steppe mammoth was excavated in the cliffs of West Runton in Norfolk, UK; it preserves its jaws and teeth but is missing the upper part of its skull. A rare skull found in Auvergne, France, in 2008 will be examined by Dick Mol and Frédéric Lacombat in the Musée Crozatier in Le Puy-en-Velay.[11]

In 1959 Zhou, M. Z described what he called a new species of mammoth, M. sungari,[12] that gained recent notoriety as the largest proboscidean due to a 5.3 metres (17.4 ft) tall and 9.1 metres (29.9 ft) long composite skeletal mount based on two individuals found in 1980. However, Wei et al. (2010), who restudied the fossils referred to M sungari, considered this species to be a junior synonym of M. trogontherii. The authors state that some of the fossils are referrable to M. trogontherii, while the others can be referred to M. primigenius, according to morphological characters and measurements.[13]

There are 8 skeletons in Russia museums, including 3 complete ones.[14] Recently one was discovered in Okhansk rayon, Perm kray.

Evolution

M. trogontherii is suggested to have derived from Mammuthus meridionalis. The oldest records M. trogontherii are known from China, at around 1.8-2 million years ago, 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).[15] Steppe mammoths replaced Mammuthus meridionalis between 1-0.7 million years ago in Europe, in a complex diachronous mosaic pattern. European populations of M. trogontherii experienced a persistent size reduction towards the end of the Middle Pleistocene. A population of M. trogontherii in north east Siberia developed higher tooth plate count after 0.8 mya, reaching M. primigenius (woolly mammoth) morphology by 400,000 years ago.[16] 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 that likely reflects regional migration and introgression.[17] Relict populations of M. trogontherii may have persisted in Mongolia and North China well into the Last Glacial Period, with teeth of M. trogontherii like morphology in Shanxi being dated to 33,858–24,857 years BP and Inner Mongolia to c. 33,700 years Before Present.[18]

See also

References

  1. ^ Todd, N. E. (January 2010). "New Phylogenetic Analysis of the Family Elephantidae Based on Cranial-Dental Morphology". The Anatomical Record. 293 (1). Wiley-Liss, Inc.: 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. JSTOR 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. ^ van de Greer, Alexandra; Lyras, George; de Vos, John; Dermitzakis, Michael (2011). Evolution of Island Mammals: Adaptation and Extinction of Placental Mammals on Islands (in Danish and English). John Wiley & Sons. p. 240. ISBN 978-1444391282.
  5. ^ a b van der Valk, Tom; Pečnerová, Patrícia; Díez-del-Molino, David; Bergström, Anders; Oppenheimer, Jonas; Hartmann, Stefanie; Xenikoudakis, Georgios; Thomas, Jessica A.; Dehasque, Marianne; Sağlıcan, Ekin; Fidan, Fatma Rabia (17 February 2021). "Million-year-old DNA sheds light on the genomic history of mammoths". Nature. 591 (7849): 265–269. Bibcode:2021Natur.591..265V. doi:10.1038/s41586-021-03224-9. ISSN 1476-4687. PMC 7116897. PMID 33597750.
  6. ^ a b Larramendi, A. (2016). "Shoulder height, body mass and shape of proboscideans" (PDF). Acta Palaeontologica Polonica. 61. doi:10.4202/app.00136.2014.
  7. ^ Mol, Dick; van den Bergh, Gert D.; de Vos, John (May 17, 1999), "Fossil Proboscideans from The Netherlands, the North Sea and the Oosterschelde Estuary" (PDF), Mammoths and the Mammoth Fauna: Studies of an Extinct Ecosystem, vol. 6, Deinsea, p. 135, retrieved 15 August 2016
  8. ^ a b Larramendi, Asier (2015-06). "Skeleton of a Late Pleistocene steppe mammoth (Mammuthus trogontherii) from Zhalainuoer, Inner Mongolian Autonomous Region, China". Paläontologische Zeitschrift. 89 (2): 229–250. doi:10.1007/s12542-014-0222-8. ISSN 0031-0220. {{cite journal}}: Check date values in: |date= (help)
  9. ^ Rivals, Florent; Semprebon, Gina M.; Lister, Adrian M. (September 2019). "Feeding traits and dietary variation in Pleistocene proboscideans: A tooth microwear review". Quaternary Science Reviews. 219: 145–153. Bibcode:2019QSRv..219..145R. doi:10.1016/j.quascirev.2019.06.027. S2CID 200073388.
  10. ^ Milivojevic, Milos (2011). "Excavation, reconstruction and conservation of steppe elephant from the clay pit of the building material factory "Toza Markovic" at Kikinda (Serbia)" (PDF). Bulletin of the Natural History Museum, 2011, 4: 51-64. Archived from the original (PDF) on 1 August 2019. Retrieved 23 June 2012.
  11. ^ Rincon, Paul (2 September 2008). "'Rare' mammoth skull discovered". BBC News. Retrieved 2 September 2008.
  12. ^ Zhou, M.Z., 1959. Proboscidea. In: Pleistocene mammalian fossils from the northeastern provinces: 22-34, pls. 6-15. Edited by Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, China.
  13. ^ GuangBiao Wei, SongMei Hu, KeFu Yu, YaMei Hou, Xin Li, ChangZhu Jin, Yuan Wang, JianXin Zhao, WenHua Wang; Hu; Yu; Hou; Li; Jin; Wang; Zhao; Wang (2010). "New materials of the steppe mammoth, Mammuthus trogontherii, with discussion on the origin and evolutionary patterns of mammoths". Science China Earth Sciences. 53 (7): 956–963. Bibcode:2010ScChD..53..956W. doi:10.1007/s11430-010-4001-4. S2CID 129568366.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  14. ^ "В Пермском крае нашли предка мамонта". 18 July 2015.
  15. ^ Lister, A. M.; Sher, A. V. (2015-11-13). "Evolution and dispersal of mammoths across the Northern Hemisphere". Science. 350 (6262): 805–809. Bibcode:2015Sci...350..805L. doi:10.1126/science.aac5660. ISSN 0036-8075. PMID 26564853. S2CID 206639522.
  16. ^ 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. Bibcode:2005QuInt.126...49L. doi:10.1016/j.quaint.2004.04.014. ISSN 1040-6182.
  17. ^ Lister, Adrian M. (October 2022). "Mammoth evolution in the late Middle Pleistocene: The Mammuthus trogontherii-primigenius transition in Europe". Quaternary Science Reviews. 294: 107693. Bibcode:2022QSRv..29407693L. doi:10.1016/j.quascirev.2022.107693. S2CID 252264887.
  18. ^ Wei, GuangBiao; Hu, SongMei; Yu, KeFu; Hou, YaMei; Li, Xin; Jin, ChangZhu; Wang, Yuan; Zhao, JianXin; Wang, WenHua (2010-06-09). "New materials of the steppe mammoth, Mammuthus trogontherii, with discussion on the origin and evolutionary patterns of mammoths". Science China Earth Sciences. 53 (7): 956–963. Bibcode:2010ScChD..53..956W. doi:10.1007/s11430-010-4001-4. ISSN 1674-7313. S2CID 129568366.

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). Mammuts : Riesen der Eiszeit (in German). 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

Retrieved from "https://en.wikipedia.org/w/index.php?title=Steppe_mammoth&oldid=1162144209"