page_title: Mammoth
section_title: Evolution
hierarchical_section_title: Mammoth / Evolution
caption_attribution_description: English: A Woolly mammoth (left) and an American mastodon (right) facing each other, showing the physical differences between the two animals.
context_page_description: 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 and, in northern species, a covering of long hair. They lived from the Pliocene epoch into the Holocene at about 4,000 years ago, and various species existed in Africa, Europe, Asia, and North America. They were members of the family Elephantidae, which also contains the two genera of modern elephants and their ancestors.
  The oldest representative of Mammuthus, the South African mammoth, appeared around 5 million years ago during the early Pliocene in what is now southern and eastern Africa. Descendant species of these mammoths moved north and continued to propagate into numerous subsequent species, eventually covering most of Eurasia before extending into the Americas at least 600,000 years ago.
context_section_description: The earliest known proboscideans, the clade that contains the elephants, existed about 55 million years ago around the Tethys Sea area. The closest relatives of the Proboscidea are the sirenians and the hyraxes. The family Elephantidae is known to have existed six million years ago in Africa, and includes the living elephants and the mammoths. Among many now extinct clades, the mastodon is only a distant relative of the mammoths, and part of the separate Mammutidae family, which diverged 25 million years before the mammoths evolved.
  The following cladogram shows the placement of the genus Mammuthus among other proboscideans, based on hyoid characteristics:
  Since many remains of each species of mammoth are known from several localities, it is possible to reconstruct the evolutionary history of the genus through morphological studies. Mammoth species can be identified from the number of enamel ridges on their molars; the primitive species had few ridges, and the amount increased gradually as new species evolved and replaced the former ones. At the same time, the crowns of the teeth became longer, and the skulls become higher from top to bottom and shorter from the back to the front over time to accommodate this.
   
  The first known members of the genus Mammuthus are the African species Mammuthus subplanifrons from the Pliocene and Mammuthus africanavus from the Pleistocene. The former is thought to be the ancestor of later forms. Mammoths entered Europe around 3 million years ago; the earliest known type has been named M. rumanus, which spread across Europe and China. Only its molars are known, which show it had 8–10 enamel ridges. A population evolved 12–14 ridges and split off from and replaced the earlier type, becoming M. meridionalis. In turn, this species was replaced by the steppe mammoth, M. trogontherii, with 18–20 ridges, which evolved in East Asia ca. 1.8 million years ago. Steppe mammoth populations replaced M. meridionalis in Europe between 1 and 0.7 million years ago.  The Columbian mammoth, M. columbi, evolved from a population of M. trogontherii that had entered North America over 1 million years ago. Mammoths derived from M. trogontherii evolved molars with 26 ridges between 800,000 and 400,000 years ago in Siberia, becoming the woolly mammoth, M. primigenius. The woolly mammoth would replace the steppe mammoth in Europe during the late Middle Pleistocene around 200,000 years ago. A 2011 genetic study showed that two examined specimens of the Columbian mammoth were grouped within a subclade of woolly mammoths. This suggests that the two populations interbred and produced fertile offspring. It also suggested that a North American form known as "M. jeffersonii" may be a hybrid between the two species.
  By the late Pleistocene, mammoths in continental Eurasia had undergone a major transformation, including a shortening and heightening of the cranium and mandible, increase in molar hypsodonty index, increase in plate number, and thinning of dental enamel. Due to this change in physical appearance, it became customary to group European mammoths separately into distinguishable clusters:
  Early Pleistocene – Mammuthus meridionalis
  Middle Pleistocene – Mammuthus trogontherii
  Late Pleistocene – Mammuthus primigenius
  There is speculation as to what caused this variation within the three chronospecies. Variations in environment, climate change, and migration surely played roles in the evolutionary process of the mammoths. Take M. primigenius for example: Woolly mammoths lived in opened grassland biomes. The cool steppe-tundra of the Northern Hemisphere was the ideal place for mammoths to thrive because of the resources it supplied. With occasional warmings during the ice age, climate would change the landscape, and resources available to the mammoths altered accordingly.
caption: Comparison of a woolly mammoth (left) and an American mastodon (right).
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