Mammuthus trogontherii, commonly 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 to 200,000 years ago. The evolution of the steppe mammoth marked the initial adaptation of the mammoth lineage towards cold environments, with the species probably being covered in a layer of fur. One of the largest mammoth species, it evolved in East Asia during the Early Pleistocene, around 1.7 million years ago, before migrating into North America around 1.3 million years ago, and into Europe during the Early/Middle Pleistocene transition, around 1 to 0.7 million years ago (replacing the earlier mammoth species Mammuthus meridionalis). It was the ancestor of the woolly mammoth, Columbian mammoth and dwarf Sardinian mammoth of the later Pleistocene. In Europe, its range overlapped with that of the temperate adapted straight-tusked elephant (Palaeoloxodon antiquus), with steppe mammoths and straight-tusked elephants generally alternately occupying northern Europe during glacial periods and interglacials, respectively, though at rare intervals they lived alongside each other at some locations.
Taxonomy There was historically confusion about the correct scientific name for the steppe mammoth, either Mammuthus armeniacus , named by Hugh Falconer in 1857 or Mammuthus trogontherii, named by Hans Pohlig in 1885. Falconer described M. armeniacus based on molar teeth collected from near Erzurum in eastern Turkey, of uncertain age, while Pohlig described M. trogontherii from fossil remains found in Europe. 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. The status of Mammuthus armeniacus as a synonym of Mammuthus trogontherii has been supported by most recent authors. The type specimens of the species are molars from the Süssenborn (also spelled Süßenborn) locality in Germany, dating to the early Middle Pleistocene, (Marine Isotope Stage/MIS 16, approximately 676-621,000 years ago). Several Early Pleistocene Japanese mammoth species and subspecies (including Mammuthus protomammonteus, Mammuthus paramammonteus shigensis, Mammuthus meridionalis shigensis and Mammuthus meridionalis proximus) are now thought to be synonyms of M. trogontherii. The species M. sungari named by Zhou 1959 from specimens found in Zalainuoer, Inner Mongolia, China, that was formerly widely used for mammoths in China is now also recognised as a synonym for M. trogontherii. Analysis of ancient DNA showing deep genetic divergences between early steppe-like mammoths in Siberia, dating around one million years ago, has led to questions about what material should be attributed to the species. In a 2024 review, Adrian Lister and Love Dalén argued that the species should be retained for now in a broad morphospecies sense for mammoth remains found across Eurasia spanning both the Early and Middle Pleistocene.
Description
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 (24,000 lb) in weight and females on average being about 3.7 m (12.1 ft) tall at the shoulders and about 9.5 tonnes (21,000 lb) in weight, considerably exceeding the size of modern elephants. 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 (23,000 lb). 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 (25,000 lb) 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 (32,000 lb) via regression analysis. Steppe mammoths from the late Middle Pleistocene of Europe were considerably smaller than these "typical" M. trogontherii specimens, with the smallest M. trogontherii population being from Stanton Harcourt, Oxfordshire, England, dating to MIS 7 (around 200,000 years ago), among the last records of the species in Europe, which have an estimated shoulder height of only 2.1–2.9 m (6.9–9.5 ft).
The skull was high-domed and short, and bore twisted tusks. The lower jaw was short and deep. The number of lamellae/plates on the third molars is around 18–22, significantly higher than the number in earlier mammoth species, but noticeably lower than the number typically present in woolly mammoths (M. primigenius), though some European specimens of M. primigenius have counts which overlap with those of M. trogontherii. Compared to M. primigenius, the teeth of calves of M. trogontherii were proportionally larger. The body has around 19 thoracic vertebrae and 5 or 6 sacral vertebrae, with the first few thoracic vertebrae having long neural spines. 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). 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. 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.
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