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The steppe mammoth (''Mammuthus trogontherii'', sometimes ''Mammuthus armeniacus'') is an extinct species of
Elephantidae Elephantidae is a family of large, herbivorous proboscidean mammals collectively called elephants and mammoths. These are terrestrial large mammals with a snout modified into a trunk and teeth modified into tusks. Most genera and species in the ...
that ranged over most of northern
Eurasia Eurasia (, ) is the largest continental area on Earth, comprising all of Europe and Asia. Primarily in the Northern and Eastern Hemispheres, it spans from the British Isles and the Iberian Peninsula in the west to the Japanese archipelag ...
during the late Early and Middle
Pleistocene The Pleistocene ( , often referred to as the ''Ice age'') is the geological Epoch (geology), epoch that lasted from about 2,580,000 to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was fina ...
, approximately 1.8 million-200,000 years ago. It evolved in
Siberia Siberia ( ; rus, Сибирь, r=Sibir', p=sʲɪˈbʲirʲ, a=Ru-Сибирь.ogg) is an extensive geographical region, constituting all of North Asia, from the Ural Mountains in the west to the Pacific Ocean in the east. It has been a part ...
during the Early Pleistocene from ''
Mammuthus meridionalis ''Mammuthus meridionalis'', or the southern mammoth, is an extinct species of mammoth native to Europe and Central Asia from the Gelasian stage of the Early Pleistocene, living from 2.5–0.8 mya. Taxonomy The taxonomy of extinct ele ...
''. It was the first stage in the evolution of the
steppe In physical geography, a steppe () is an ecoregion characterized by grassland plains without trees apart from those near rivers and lakes. Steppe biomes may include: * the montane grasslands and shrublands biome * the temperate gras ...
and
tundra In physical geography, tundra () is a type of biome where tree growth is hindered by frigid temperatures and short growing seasons. The term ''tundra'' comes through Russian (') from the Kildin Sámi word (') meaning "uplands", "treeless mou ...
elephant Elephants are the largest existing land animals. Three living species are currently recognised: the African bush elephant, the African forest elephant, and the Asian elephant. They are the only surviving members of the family Elephantida ...
s and the ancestor of the
woolly mammoth The woolly mammoth (''Mammuthus primigenius'') is an extinct species of mammoth that lived during the Pleistocene until its extinction in the Holocene epoch. It was one of the last in a line of mammoth species, beginning with '' Mammuthus s ...
and
Columbian mammoth The Columbian mammoth (''Mammuthus columbi'') is an extinct species of mammoth that inhabited the Americas as far north as the Northern United States and as far south as Costa Rica during the Pleistocene epoch. It was one of the last in a line ...
of the later Pleistocene. Populations of steppe mammoth may have persisted in
northern China Northern China () and Southern China () are two approximate regions within China. The exact boundary between these two regions is not precisely defined and only serve to depict where there appears to be regional differences between the climate ...
and Mongolia as late as 33,000 years ago.


Taxonomy

There is 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. 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''. 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''.


Genetics

DNA has been sequenced from two Siberian mammoth
molars The molars or molar teeth are large, flat teeth at the back of the mouth. They are more developed in mammals. They are used primarily to grind food during chewing. The name ''molar'' derives from Latin, ''molaris dens'', meaning "millstone to ...
(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 woolly mammoth (''Mammuthus primigenius'') is an extinct species of mammoth that lived during the Pleistocene until its extinction in the Holocene epoch. It was one of the last in a line of mammoth species, beginning with '' Mammuthus s ...
, 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 The Columbian mammoth (''Mammuthus columbi'') is an extinct species of mammoth that inhabited the Americas as far north as the Northern United States and as far south as Costa Rica during the Pleistocene epoch. It was one of the last in a line ...
.


Description

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 in old bulls; females on the other hand had thinner and slightly curved tusks. With several individuals reaching tall at the shoulders, it is smaller than the largest
proboscidea The Proboscidea (; , ) are a taxonomic order of afrotherian mammals containing one living family ( Elephantidae) and several extinct families. First described by J. Illiger in 1811, it encompasses the elephants and their close relatives. Fr ...
ns ever to have lived (''
Palaeoloxodon namadicus ''Palaeoloxodon namadicus'' or the Asian straight-tusked elephant, is an extinct species of prehistoric elephant known from the early Middle to Late Pleistocene of the Indian subcontinent, and possibly also elsewhere in Asia. Some authorities r ...
'' reached 22 tonnes and shoulder heights of ), but was larger than other
mammoth A mammoth is any species of the extinct elephantid genus ''Mammuthus'', one of the many genera that make up the order of trunked mammals called proboscideans. The various species of mammoth were commonly equipped with long, curved tusks an ...
s. A skeleton mounted on the Azov Museum reaches at the shoulder, though this figure might be overestimated because the vertebrae have been placed between the tips of the shoulder blades. Another individual represented by a single humerus long found in Mosbach Sande,
Germany Germany,, officially the Federal Republic of Germany, is a country in Central Europe. It is the second most populous country in Europe after Russia, and the most populous member state of the European Union. Germany is situated betwee ...
, is estimated to have an in-the-flesh shoulder height of , weighed between and might be the largest mammoth found yet. Another estimate gives a shoulder height of and a weight of for the species.


Discovery

Fossilized teeth are recovered, but
skeletal A skeleton is the structural frame that supports the body of an animal. There are several types of skeletons, including the exoskeleton, which is the stable outer shell of an organism, the endoskeleton, which forms the support structure inside ...
parts are rare. The most complete skeleton of a steppe mammoth yet found was discovered in 1996 in
Kikinda Kikinda ( sr-Cyrl, Кикинда, ; hu, Nagykikinda) is a city and the administrative center of the North Banat District in Serbia . The city urban area has 38,069 inhabitants, while the city administrative area has 59,453 inhabitants. The ...
,
Serbia Serbia (, ; Serbian: , , ), officially the Republic of Serbia ( Serbian: , , ), is a landlocked country in Southeastern and Central Europe, situated at the crossroads of the Pannonian Basin and the Balkans. It shares land borders with Hu ...
. It was mounted, and put on display in 2005. The specimen is a female, which was about high, in length and with long tusks. 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 Dick "Sir Mammoth" Mol (born June 26, 1955) is a Dutch paleontologist - a specialist in the field of mammoths for almost three decades. He is a research associate of several museums. Mol's primary focus is on mammals of the Quaternary period, incl ...
and Frédéric Lacombat in the
Musée Crozatier The Musée Crozatier is a museum in Le Puy-en-Velay in the French Auvergne. Inaugurated in 1868, its collection comprises art and archaeological artifacts from Velay and the Haute-Loire region. The museum has undergone a major renovation from 2 ...
in
Le Puy-en-Velay Le Puy-en-Velay (, literally ''Le Puy in Velay''; oc, Lo Puèi de Velai ) is the prefecture of the Haute-Loire department in the Auvergne-Rhône-Alpes region of south-central France. Located near the river Loire, the city is famous for its c ...
. In 1959 Zhou, M. Z described what he called a new species of mammoth, ''M. sungari'',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. that gained recent notoriety as the largest proboscidean due to a tall and 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 The Botanical and Zoological Codes of nomenclature treat the concept of synonymy differently. * In botanical nomenclature, a synonym is a scientific name that applies to a taxon that (now) goes by a different scientific name. For example, Linn ...
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. There are 8 skeletons in Russia museums, including 3 complete ones. Recently one was discovered in
Okhansk rayon Okhansky District (russian: Оханский райо́н) is an administrative district ( raion) of Perm Krai, Russia; one of the thirty-three in the krai.Law #416-67 Municipally, it is incorporated as Okhansky Municipal District.Law #1878-407 ...
, Perm kray.


Evolution

''M. trogontherii'' is derived from ''Mammuthus meridionalis'', the oldest records of the species are known from China, at around 1.8-2 ma in age from the Nihewan Formation near Majuangou,
Hebei Hebei or , (; alternately Hopeh) is a northern province of China. Hebei is China's sixth most populous province, with over 75 million people. Shijiazhuang is the capital city. The province is 96% Han Chinese, 3% Manchu, 0.8% Hui, and ...
. Steppe mammoths arrived in North America across
Beringia Beringia is defined today as the land and maritime area bounded on the west by the Lena River in Russia; on the east by the Mackenzie River in Canada; on the north by 72 degrees north latitude in the Chukchi Sea; and on the south by the tip ...
around 1.5 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). Steppe mammoths replaced ''Mammuthus meridionalis'' between 1-0.7 million years ago in Europe, in a complex
diachronous In geology, a diachronism ( Greek ''dia'', "through" + ''chronos'', "time" + ''-ism''), or diachronous deposit, is a sedimentary rock formation in which the material, although of a similar nature, varies in age with the place where it was deposit ...
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'' morphology by 400,000 years ago. ''M. primigenius'' replaced ''M. trogontherii'' in Europe around 200 kya. 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 ka BP and Inner Mongolia to c. 33.7 ka BP.


See also

*''
Elephas recki ''Palaeoloxodon recki'' is an extinct species of elephant native to Africa during the Pliocene and Pleistocene. At up to 14 feet (4.27 metres) in shoulder height, it was one of the largest elephant species to have ever lived. It is believed that ...
'' *'' Mammuthus columbi'' *''
Palaeoloxodon ''Palaeoloxodon'' is an extinct genus of elephant. The genus originated in Africa during the Pliocene era, and expanded into Eurasia during the Pleistocene era. The genus contains some of the largest known species of elephants, over four metres t ...
'' *''
Paraceratherium ''Paraceratherium'' is an extinct genus of hornless rhinoceros. It is one of the largest terrestrial mammals that has existed and lived from the early to late Oligocene epoch (34–23 million years ago). The first fossils were discovered ...
'' *
West Runton Mammoth The West Runton Mammoth is a fossilized skeleton of a steppe mammoth (''Mammuthus trogontherii'') found in the cliffs of West Runton in the county of Norfolk, England in 1990. The find is the largest nearly complete mammoth skeleton known, and is ...


References


Further reading

* * * * (English and French)


External links


The Kikinda mammoth
{{Taxonbar, from=Q113284 Pleistocene mammals of Europe Mammoths Pleistocene proboscideans Fossils of Serbia