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A mastodon, from
Ancient Greek Ancient Greek (, ; ) includes the forms of the Greek language used in ancient Greece and the classical antiquity, ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Greek ...
μαστός (''mastós''), meaning "breast", and ὀδούς (''odoús'') "tooth", is a member of the genus ''Mammut'' (German for 'mammoth'), which was endemic to North America and lived from the late
Miocene The Miocene ( ) is the first epoch (geology), geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and mea ...
to the early
Holocene The Holocene () is the current geologic time scale, geological epoch, beginning approximately 11,700 years ago. It follows the Last Glacial Period, which concluded with the Holocene glacial retreat. The Holocene and the preceding Pleistocene to ...
. Mastodons belong to the order
Proboscidea Proboscidea (; , ) is 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. Three l ...
, the same order as
elephant Elephants are the largest living land animals. Three living species are currently recognised: the African bush elephant ('' Loxodonta africana''), the African forest elephant (''L. cyclotis''), and the Asian elephant ('' Elephas maximus ...
s and
mammoth A mammoth is any species of the extinct elephantid genus ''Mammuthus.'' They lived from the late Miocene epoch (from around 6.2 million years ago) into the Holocene until about 4,000 years ago, with mammoth species at various times inhabi ...
s (which belong to the family
Elephantidae Elephantidae is a family (biology), family of large, herbivorous proboscidean mammals which includes the living Elephant, elephants (belonging to the genera ''Elephas'' and ''Loxodonta''), as well as a number of extinct genera like ''Mammuthus'' ...
). ''Mammut'' is the
type genus In biological taxonomy, the type genus (''genus typica'') is the genus which defines a biological family and the root of the family name. Zoological nomenclature According to the International Code of Zoological Nomenclature, "The name-bearin ...
of the extinct family
Mammutidae Mammutidae is an extinct family of proboscideans belonging to Elephantimorpha. It is best known for the mastodons (genus ''Mammut''), which inhabited North America from the Late Miocene (around 8 million years ago) until their extinction at the ...
, which diverged from the ancestors of modern elephants at least 27–25 million years ago, during the
Oligocene The Oligocene ( ) is a geologic epoch (geology), epoch of the Paleogene Geologic time scale, Period that extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that defin ...
. Like other members of Mammutidae, the molar teeth of mastodons have zygodont morphology (where parallel pairs of cusps are merged into sharp ridges), which strongly differ from those of elephantids. In comparison to its likely ancestor ''
Zygolophodon ''Zygolophodon'' is an extinct genus of mammutid proboscidean that lived during the Miocene in Africa, Eurasia, and North America. Description As with other mammutids, the molars have a zygodont morphology. The fused front region of the lower ...
'', ''Mammut'' is characterized by particularly long and upward curving upper tusks, reduced or absent tusks on the lower jaw, as well as the shortening of the
mandibular symphysis In human anatomy, the facial skeleton of the skull the external surface of the mandible is marked in the median line by a faint ridge, indicating the mandibular symphysis (Latin: ''symphysis menti'') or line of junction where the two lateral ha ...
(the frontmost part of the lower jaw), the latter two traits also having evolved in parallel separately in elephantids. Mastodons had an overall stockier skeletal build, a lower-domed skull, and a longer tail compared to elephantids. Fully grown male ''M. americanum'' are thought to have been have been to at shoulder height and from to in body mass on average. The size estimates suggest that American mastodon males were on average heavier than any living elephant species; they were typically larger than
Asian elephant The Asian elephant (''Elephas maximus''), also known as the Asiatic elephant, is the only living ''Elephas'' species. It is the largest living land animal in Asia and the second largest living Elephantidae, elephantid in the world. It is char ...
s and
African forest elephant The African forest elephant (''Loxodonta cyclotis'') is one of the two living species of African elephant, along with the African bush elephant. It is native to humid tropical forests in West Africa and the Congo Basin. It is the smallest of the ...
s of both sexes but shorter than male
African bush elephant The African bush elephant (''Loxodonta africana''), also known as the African savanna elephant, is a species of elephant native to sub-Saharan Africa. It is one of three extant elephant species and, along with the African forest elephant, one ...
s. ''M. americanum'', known as an "American mastodon" or simply "mastodon," had a long and complex paleontological history spanning all the way back to 1705 when the first fossils were uncovered from
Claverack, New York Claverack is a town in Columbia County, New York, United States. The population was 6,058 at the 2020 census.US Census Bureau, 2020 Census Results, Claverack town, Columbia County, New York QuickFacts https://www.census.gov/quickfacts/claverackt ...
, in the American colonies. Because of the uniquely shaped molars with no modern analogues in terms of large animals, the species caught wide attention of European researchers and influential Americans before and after the
American Revolution The American Revolution (1765–1783) was a colonial rebellion and war of independence in which the Thirteen Colonies broke from British America, British rule to form the United States of America. The revolution culminated in the American ...
to the point of, according to American historians Paul Semonin and
Keith Stewart Thomson Keith Stewart Thomson (born 1938; B.SC. Birmingham, AM, Ph.D. Harvard) was from 2003 to 2012 a senior research fellow of the American Philosophical Society and is, starting in 2012, the Executive Officer of the American Philosophical Society and i ...
, bolstering
American nationalism American nationalism is a form of civic, ethnic, cultural or economic influences * * * * * * * found in the United States. Essentially, it indicates the aspects that characterize and distinguish the United States as an autonomous political com ...
and contributing to a greater understanding of extinctions. Taxonomically, it was first recognized as a distinct species by Robert Kerr in 1792 then classified to its own genus ''Mammut'' by
Johann Friedrich Blumenbach Johann Friedrich Blumenbach (11 May 1752 – 22 January 1840) was a German physician, naturalist, physiologist and anthropologist. He is considered to be a main founder of zoology and anthropology as comparative, scientific disciplines. He has be ...
in 1799, thus making it amongst the first fossil mammal genera to be erected with undisputed taxonomic authority. The genus served as a
wastebasket taxon Wastebasket taxon (also called a wastebin taxon, dustbin taxon or catch-all taxon) is a term used by some taxonomists to refer to a taxon that has the purpose of classifying organisms that do not fit anywhere else. They are typically defined by e ...
for proboscidean species with superficially similar molar teeth morphologies but today includes 7 definite species, 1 of questionable affinities, and 4 other species from Eurasia that are pending reassessments to other genera. Mastodons are considered to have had a predominantly
browsing Browsing is a kind of orienting strategy. It is supposed to identify something of relevance for the browsing organism. In context of humans, it is a metaphor taken from the animal kingdom. It is used, for example, about people browsing open sh ...
-based diet on leaves, fruits, and woody parts of plants. This allowed mastodons to niche partition with other members of Proboscidea in North America, like
gomphothere Gomphotheres are an extinct group of proboscideans related to modern elephants. First appearing in Africa during the Oligocene, they dispersed into Eurasia and North America during the Miocene and arrived in South America during the Pleistocene a ...
s and the Columbian mammoth, who had shifted to mixed feeding or
grazing In agriculture, grazing is a method of animal husbandry whereby domestic livestock are allowed outdoors to free range (roam around) and consume wild vegetations in order to feed conversion ratio, convert the otherwise indigestible (by human diges ...
by the late
Neogene The Neogene ( ,) is a geologic period and system that spans 20.45 million years from the end of the Paleogene Period million years ago ( Mya) to the beginning of the present Quaternary Period million years ago. It is the second period of th ...
-
Quaternary The Quaternary ( ) is the current and most recent of the three periods of the Cenozoic Era in the geologic time scale of the International Commission on Stratigraphy (ICS), as well as the current and most recent of the twelve periods of the ...
. It is thought that mastodon behaviors were not much different from elephants and mammoths, with females and juveniles living in herds and adult males living largely solitary lives plus entering phases of aggression similar to the
musth Musth or must (from Persian, ) is a periodic condition in bull (male) elephants characterized by aggressive behavior in animals, aggressive behavior and accompanied by a large rise in reproductive hormones. It has been known in Asian elephan ...
exhibited by modern elephants. ''Mammut'' achieved maximum species diversity in the
Pliocene The Pliocene ( ; also Pleiocene) is the epoch (geology), epoch in the geologic time scale that extends from 5.33 to 2.58Late Pleistocene The Late Pleistocene is an unofficial Age (geology), age in the international geologic timescale in chronostratigraphy, also known as the Upper Pleistocene from a Stratigraphy, stratigraphic perspective. It is intended to be the fourth division ...
. Mastodons for at least a few thousand years prior to their extinction coexisted with
Paleoindian Paleo-Indians were the first peoples who entered and subsequently inhabited the Americas towards the end of the Late Pleistocene period. The prefix ''paleo-'' comes from . The term ''Paleo-Indians'' applies specifically to the lithic period in ...
s, who were the first humans to have inhabited North America. Evidence has been found that Paleoindians (including those of the
Clovis culture The Clovis culture is an archaeological culture from the Paleoindian period of North America, spanning around 13,050 to 12,750 years Before Present (BP). The type site is Blackwater Draw locality No. 1 near Clovis, New Mexico, where stone too ...
) hunted mastodons based on the finding of mastodon remains with cut marks and/or with lithic artifacts. Mastodons disappeared along with many other North American animals, including most of its largest animals (
megafauna In zoology, megafauna (from Ancient Greek, Greek μέγας ''megas'' "large" and Neo-Latin ''fauna'' "animal life") are large animals. The precise definition of the term varies widely, though a common threshold is approximately , this lower en ...
), as part of the end-Pleistocene extinction event around the end of the Late Pleistocene-early Holocene, the causes typically being attributed to human hunting, severe climatic phases like the
Younger Dryas The Younger Dryas (YD, Greenland Stadial GS-1) was a period in Earth's geologic history that occurred circa 12,900 to 11,700 years Before Present (BP). It is primarily known for the sudden or "abrupt" cooling in the Northern Hemisphere, when the ...
, or some combination of the two. The American mastodon had its last recorded occurrence in the earliest Holocene around 11,000 years ago, which is considerably later than other North American megafauna species. Today, the American mastodon is one of the most well-known fossil species in both academic research and public perception, the result of its inclusion in American popular culture.


Taxonomy


Research history


Earliest finds

In a letter dating to 1713,
Edward Hyde, 3rd Earl of Clarendon Edward Hyde, 3rd Earl of Clarendon (28 November 1661 – 31 March 1723), styled Viscount Cornbury between 1674 and 1709, was an English Army officer, politician and colonial administrator. He was propelled into the forefront of English polit ...
(known also as Lord Cornbury) from
New York New York most commonly refers to: * New York (state), a state in the northeastern United States * New York City, the most populous city in the United States, located in the state of New York New York may also refer to: Places United Kingdom * ...
reported to the
Royal Society The Royal Society, formally The Royal Society of London for Improving Natural Knowledge, is a learned society and the United Kingdom's national academy of sciences. The society fulfils a number of roles: promoting science and its benefits, re ...
learned society A learned society ( ; also scholarly, intellectual, or academic society) is an organization that exists to promote an academic discipline, profession, or a group of related disciplines such as the arts and sciences. Membership may be open to al ...
of
Great Britain Great Britain is an island in the North Atlantic Ocean off the north-west coast of continental Europe, consisting of the countries England, Scotland, and Wales. With an area of , it is the largest of the British Isles, the List of European ...
that in 1705, a large-sized tooth was found near the side of the
Hudson River The Hudson River, historically the North River, is a river that flows from north to south largely through eastern New York (state), New York state. It originates in the Adirondack Mountains at Henderson Lake (New York), Henderson Lake in the ...
by a
Dutch Dutch or Nederlands commonly refers to: * Something of, from, or related to the Netherlands ** Dutch people as an ethnic group () ** Dutch nationality law, history and regulations of Dutch citizenship () ** Dutch language () * In specific terms, i ...
country-fellow and was sold to
New York General Assembly The General Assembly of New York, commonly known internationally as the New York General Assembly, and domestically simply as General Assembly, was the Parliamentary sovereignty, supreme Legislature, legislative body of the Province of New York d ...
member Van Bruggen for a
gill A gill () is a respiration organ, respiratory organ that many aquatic ecosystem, aquatic organisms use to extract dissolved oxygen from water and to excrete carbon dioxide. The gills of some species, such as hermit crabs, have adapted to allow r ...
of rum, and Bruggen eventually gave it to Cornbury. He then stated that he sent Johannis Abeel, a recorder of Albany, New York to dig near the original site of the tooth to find more bones. Abeel reported later that he went to the town of Claverack, New York where the original bones were found. American historian Paul Semonin said that the accounts written by Cornbury and Abeel match up with that written by in the July 30, 1705 entry in ''
The Boston News-Letter ''The Boston News-Letter'', first published on April 24, 1704, is regarded as the first continuously published newspaper in the Province of Massachusetts Bay, colony of Massachusetts. It was heavily subsidized by the British government, with a l ...
''. The account reported skeletal evidence of an
antediluvian The antediluvian (alternatively pre-diluvian or pre-flood) period is the time period chronicled in the Bible between the fall of man and the Genesis flood narrative in biblical cosmology. The term was coined by Thomas Browne (1605–1682). The n ...
(or biblical) "
giant In folklore, giants (from Ancient Greek: ''wiktionary:gigas, gigas'', cognate wiktionary:giga-, giga-) are beings of humanoid appearance, but are at times prodigious in size and strength or bear an otherwise notable appearance. The word ''gia ...
" uncovered from Claverack. The
femur The femur (; : femurs or femora ), or thigh bone is the only long bone, bone in the thigh — the region of the lower limb between the hip and the knee. In many quadrupeds, four-legged animals the femur is the upper bone of the hindleg. The Femo ...
and one of the teeth both dissolved before they could be further observed, however.


Big Bone Lick

In 1739, a French military expedition under the command of Charles III Le Moyne (known also as "Longueil") explored the locality of "
Big Bone Lick Big Bone Lick State Park is located at Big Bone, Kentucky, Big Bone in Boone County, Kentucky. The name of the park comes from the Pleistocene megafauna fossils found there. Mammoths are believed to have been drawn to this location by a salt lic ...
" (located in what is now the US state of
Kentucky Kentucky (, ), officially the Commonwealth of Kentucky, is a landlocked U.S. state, state in the Southeastern United States, Southeastern region of the United States. It borders Illinois, Indiana, and Ohio to the north, West Virginia to the ...
) and gathered fossil bones and teeth there. The French naturalist
Louis Jean-Marie Daubenton Louis Jean-Marie Daubenton (; 29 May 1716 – 1 January 1800) was a French natural history, naturalist and contributor to the ''Encyclopédie, Encyclopédie ou Dictionnaire raisonné des sciences, des arts et des métiers''. Biography Daubent ...
examined the fossil collection brought by Longueuil and compared it with specimens of extant
elephant Elephants are the largest living land animals. Three living species are currently recognised: the African bush elephant ('' Loxodonta africana''), the African forest elephant (''L. cyclotis''), and the Asian elephant ('' Elephas maximus ...
s and Siberian
mammoth A mammoth is any species of the extinct elephantid genus ''Mammuthus.'' They lived from the late Miocene epoch (from around 6.2 million years ago) into the Holocene until about 4,000 years ago, with mammoth species at various times inhabi ...
s in 1762. Daubenton said that the bones were discovered by Native Americans (probably
Abenaki The Abenaki ( Abenaki: ''Wαpánahki'') are Indigenous people of the Northeastern Woodlands of Canada and the United States. They are an Algonquian-speaking people and part of the Wabanaki Confederacy. The Eastern Abenaki language was pred ...
hunter-warriors). He came to the conclusion that the femur and tusk belonged to an elephant while the
molars The molars or molar teeth are large, flat tooth, teeth at the back of the mouth. They are more developed in mammal, mammals. They are used primarily to comminution, grind food during mastication, chewing. The name ''molar'' derives from Latin, '' ...
(or cheek teeth) came from a separate giant
hippopotamus The hippopotamus (''Hippopotamus amphibius;'' ; : hippopotamuses), often shortened to hippo (: hippos), further qualified as the common hippopotamus, Nile hippopotamus and river hippopotamus, is a large semiaquatic mammal native to sub-Sahar ...
. In
Shawnee The Shawnee ( ) are a Native American people of the Northeastern Woodlands. Their language, Shawnee, is an Algonquian language. Their precontact homeland was likely centered in southern Ohio. In the 17th century, they dispersed through Ohi ...
tradition, the proboscideans roamed in herds and were hunted by giants, who both eventually died out. The accounts told by the Shawnee individuals in 1762 are the oldest known documented interpretations of the "
Ohio Ohio ( ) is a U.S. state, state in the Midwestern United States, Midwestern region of the United States. It borders Lake Erie to the north, Pennsylvania to the east, West Virginia to the southeast, Kentucky to the southwest, Indiana to the ...
" fossils, although the traditions may have had been told for generations. In 1767, Peter Collinson credited Irish trader
George Croghan George Croghan (c. 1718 – August 31, 1782) was an Kingdom of Ireland, Irish-born fur trader in the Ohio Country of North America (current United States) who became a key early figure in the region. In 1746 he was appointed to the Onondaga Cou ...
for having sent him and
Benjamin Franklin Benjamin Franklin (April 17, 1790) was an American polymath: a writer, scientist, inventor, statesman, diplomat, printer, publisher and Political philosophy, political philosopher.#britannica, Encyclopædia Britannica, Wood, 2021 Among the m ...
fossil evidence of the mysterious proboscideans, using them for his studies. He concluded that the peculiar grinders (the molars) were built for herbivorous diets of branches of trees and shrubs as well as other vegetation, a view later followed by Franklin. In 1768, Scottish anatomist William Hunter recorded that he and his brother
John Hunter John Hunter may refer to: Politics *John Hunter (British politician) (1724–1802), British Member of Parliament for Leominster * John Hunter (Canadian politician) (1909–1993), Canadian Liberal MP for Parkdale, 1949–1957 *Sir John Hunter ( ...
observed that the teeth were not like those of modern elephants. He determined that the "grinders" from Ohio were of a carnivorous animal but believed that the tusks belonged to the same animal. After examining fossils from Franklin and Lord Shelburne, Hunter was convinced that the "''pseudo-elephant''", or "''animal incognitum''" (shortened as "''incognitum''"), was an animal species separate from elephants that might have also been the same as the proboscideans found in Siberia. He concluded his article with the opinion that although regrettable to philosophers, humanity should be thankful to heaven that the animal, if truly carnivorous, was extinct.


Early American observations

In 1785, Reverend Robert Annan wrote an account recalling an event in which workers discovered bones in his farm near the Hudson River in New York in fall of 1780. The workers found four molars in addition to another that was broken and thrown away. They also uncovered bones, including vertebrae that broke shortly thereafter. Annan expressed his confusion at what the animal could be but speculated based on its "grinders" that it was carnivorous in diet. He speculated also that it was probably extinct due to some catastrophe within the globe. American statesman
Thomas Jefferson Thomas Jefferson (, 1743July 4, 1826) was an American Founding Fathers of the United States, Founding Father and the third president of the United States from 1801 to 1809. He was the primary author of the United States Declaration of Indepe ...
stated his thoughts on ''
Notes on the State of Virginia ''Notes on the State of Virginia'' (1785) is a book written by the American statesman, philosopher, and planter Thomas Jefferson. He completed the first version in 1781 and updated and enlarged the book in 1782 and 1783. It originated in Jeffers ...
'' (published by 1785) that the fossil proboscideans may have been carnivorous, still exist in the northern parts of North America, and are related to mammoths whose remains were found in Siberia. Jefferson referenced the theory of American
social degeneracy Social degeneration was a widely influential concept at the interface of the social and biological sciences in the 18th and 19th centuries. During the 18th century, scientific thinkers including Georges-Louis Leclerc, Comte de Buffon, Johann Fri ...
by
Georges-Louis Leclerc, Comte de Buffon Georges-Louis Leclerc, Comte de Buffon (; 7 September 1707 – 16 April 1788) was a French Natural history, naturalist, mathematician, and cosmology, cosmologist. He held the position of ''intendant'' (director) at the ''Jardin du Roi'', now ca ...
, countering it by using extant and extinct animal measurements, including those of "mammoths," as proof that North America faunas were not "degenerative" in size. Semonin pointed out that social degeneracy was an offensive concept to Anglo-American naturalists and that the American proboscidean fossils were used as political tools to inspire
American nationalism American nationalism is a form of civic, ethnic, cultural or economic influences * * * * * * * found in the United States. Essentially, it indicates the aspects that characterize and distinguish the United States as an autonomous political com ...
and counter against the theory of American degeneracy. In 1799, laborers recovered a thighbone while digging a
marl Marl is an earthy material rich in carbonate minerals, Clay minerals, clays, and silt. When Lithification, hardened into rock, this becomes marlstone. It is formed in marine or freshwater environments, often through the activities of algae. M ...
pit at John Masten's farm in Newburgh, New York, and subsequent excavations were observed by a crowd of over a hundred people. American painter and exhibitionist
Charles Willson Peale Charles Willson Peale (April 15, 1741 – February 22, 1827) was an American painter, military officer, scientist, and naturalist. In 1775, inspired by the American Revolution, Peale moved from his native Maryland to Philadelphia, where he set ...
visited the locality in 1801, where he first sketched the fossils then purchased excavation privileges and full ownership of the fossils from Masten and borrowed a loan from the
American Philosophical Society The American Philosophical Society (APS) is an American scholarly organization and learned society founded in 1743 in Philadelphia that promotes knowledge in the humanities and natural sciences through research, professional meetings, publicat ...
(APS) in
Philadelphia Philadelphia ( ), colloquially referred to as Philly, is the List of municipalities in Pennsylvania, most populous city in the U.S. state of Pennsylvania and the List of United States cities by population, sixth-most populous city in the Unit ...
,
Pennsylvania Pennsylvania, officially the Commonwealth of Pennsylvania, is a U.S. state, state spanning the Mid-Atlantic (United States), Mid-Atlantic, Northeastern United States, Northeastern, Appalachian, and Great Lakes region, Great Lakes regions o ...
. In addition to the first skeleton, the second was excavated using a mill-like device to drain a deep marl pit. Peale assembled a complete skeleton in his Philadelphia Museum in 1804, and its exhibit was open first to invited members of the American Philosophical Society on December 24 then to the general public on December 25 for an exhibit admission fee in addition to the general admission fee. The special exhibition attracted thousands of visitors, and the skeleton became a US national symbol. Charles Peale's son
Rembrandt Peale Rembrandt Peale (February 22, 1778 – October 3, 1860) was an American artist and museum keeper. A prolific portrait painter, he was especially acclaimed for his likenesses of presidents George Washington and Thomas Jefferson. Peale's style wa ...
took the skeleton to Europe used to promote the fossil proboscidean and have it used as support for Jefferson's final rebuttals against Buffon's arguments for supposed inferiority of American faunas. Author Keith Stewart Thomson argued that the promotion of the "mastodon" skeleton made it a symbol of the strength of American nationalism and that "mammoth" as a term became associated with gigantism. Decades later, the museum bankrupted, and the first skeleton's specimens were sold to some German spectators in around 1848, who eventually sold it to
Hessisches Landesmuseum Darmstadt Hessisches Landesmuseum Darmstadt (HLMD) is a large multidisciplinary museum in Darmstadt, Germany. The museum exhibits Rembrandt, Beuys, a primeval horse and a mastodon under the slogan "The whole world under one roof". As one of the oldest publ ...
in Germany where it is now displayed. The second skeleton's specimens landed eventually at the
American Museum of Natural History The American Museum of Natural History (AMNH) is a natural history museum on the Upper West Side of Manhattan in New York City. Located in Theodore Roosevelt Park, across the street from Central Park, the museum complex comprises 21 interconn ...
. Other skeletons of ''Mammut americanum'' were excavated within the United States in the first half of the 19th century. One of them was collected by American showman Albert C. Koch in what is today the Mastodon State Historic Site at
Missouri Missouri (''see #Etymology and pronunciation, pronunciation'') is a U.S. state, state in the Midwestern United States, Midwestern region of the United States. Ranking List of U.S. states and territories by area, 21st in land area, it border ...
in 1839. He hypothesized in 1840 that the proboscidean, which he classified as ''Missourium'', was much larger than an elephant, had horizontal tusks plus trunks, and occupied aquatic habitats. He acquired additional fossils from a spring on the Pomme de Terre River to assemble a mounted skeleton of the "''Missouri Leviathan''" and briefly exhibited it at After exhibiting the skeleton throughout Europe, he sold the skeleton to the
British Museum of Natural History The Natural History Museum in London is a museum that exhibits a vast range of specimens from various segments of natural history. It is one of three major museums on Exhibition Road in South Kensington, the others being the Science Museum (Lo ...
.
Richard Owen Sir Richard Owen (20 July 1804 – 18 December 1892) was an English biologist, comparative anatomy, comparative anatomist and paleontology, palaeontologist. Owen is generally considered to have been an outstanding naturalist with a remarkabl ...
then properly reassembled the skeleton, and it today is on display there. In 1845, another skeleton was excavated from Newburgh by laborers hired by Nathaniel Brewster initially to remove
lacustrine deposits Lacustrine deposits are sedimentary rock formations which formed in the bottom of ancient lakes. A common characteristic of lacustrine deposits is that a river or stream channel has carried sediment into the basin. Lacustrine deposits form in all ...
to fertilize the neighboring fields. They were observed by a large number of spectators and uncovered relatively complete fossil evidence of ''M. americanum''. The skeleton was exhibited in
New York City New York, often called New York City (NYC), is the most populous city in the United States, located at the southern tip of New York State on one of the world's largest natural harbors. The city comprises five boroughs, each coextensive w ...
and other New England towns then was acquired by
John Collins Warren John Warren may refer to: Entertainment * John F. Warren (1909–2000), American cinematographer * John Warren (actor) (1916–1977), British screenwriter and actor * John Warren (musician) (born 1938), Canadian baritone saxophonist and compos ...
for study. After Warren's death in 1856, the skeleton was sent to Warren's family but was traded to
Harvard Medical School Harvard Medical School (HMS) is the medical school of Harvard University and is located in the Longwood Medical and Academic Area, Longwood Medical Area in Boston, Massachusetts. Founded in 1782, HMS is the third oldest medical school in the Un ...
for John Warren's skeleton. The "Warren mastodon", under the request of American paleontologist
Henry Fairfield Osborn Henry Fairfield Osborn, Sr. (August 8, 1857 – November 6, 1935) was an American paleontologist, geologist and eugenics advocate. He was professor of anatomy at Columbia University, president of the American Museum of Natural History for 25 y ...
, was purchased by the American financier
J. P. Morgan John Pierpont Morgan Sr. (April 17, 1837 – March 31, 1913) was an American financier and investment banker who dominated corporate finance on Wall Street throughout the Gilded Age and Progressive Era. As the head of the banking firm that ...
for $30,000 in 1906 and donated to the American Museum of Natural History where it is exhibited today.


Early taxonomic history

In the 1790s, the "''American incognitum''" was subject to research by multiple taxonomists. Scottish writer Robert Kerr erected the species name ''Elephas americanus'' in 1792 based on fossil tusks and "grinders" from the Big Bone Lick locality. He stated that the tusks were similar to elephants while the molars were completely different because they were covered with enamel and had a double row of high conical
cusp A cusp is the most pointed end of a curve. It often refers to cusp (anatomy), a pointed structure on a tooth. Cusp or CUSP may also refer to: Mathematics * Cusp (singularity), a singular point of a curve * Cusp catastrophe, a branch of bifu ...
processes. Kerr was unsure about the taxonomic affinities of the molars and referenced that
Thomas Pennant Thomas Pennant (16 December 1798) was a Welsh natural history, naturalist, traveller, writer and antiquarian. He was born and lived his whole life at his family estate, Downing Hall, near Whitford, Flintshire, in Wales. As a naturalist he had ...
supposed that they belong to an unknown species within the genus ''
Elephas ''Elephas'' is a genus of elephants and one of two surviving genera in the Family (biology), family Elephantidae, comprising one extant species, the Asian elephant (''E. maximus''). Several extinct species have been identified as belonging to t ...
'', giving the common name "American elephant". German naturalist
Johann Friedrich Blumenbach Johann Friedrich Blumenbach (11 May 1752 – 22 January 1840) was a German physician, naturalist, physiologist and anthropologist. He is considered to be a main founder of zoology and anthropology as comparative, scientific disciplines. He has be ...
also followed up with more taxonomic descriptions of fossil proboscideans in 1799. The first fossil species, recovered from Germany, was described as belonging to the newly erected species ''Elephas primigenius''? (now known as '' Mammuthus primigenius''). The second was what he considered to be an unknown "colossal land monster of the prehistoric world," considering it to be the "mammoth." He created the genus ''Mammut'' and erected the species ''Mammut ohioticum'' based on fossil bones dug up from Ohio in North America. He said that the species was distinguished from other animals of the prehistoric world based on the unusual shapes of the large molars. The genus name "Mammut" refers to the German translation for "mammoth." The naming of the genus ''Mammut'' in 1799 makes it the second or third genus to be recognized with taxonomic authority given that ''
Megalonyx ''Megalonyx'' (Greek, "great-claw") is an extinct genus of ground sloths of the family Megalonychidae, native to North America. It evolved during the Pliocene Epoch and became extinct at the end of the Late Pleistocene, living from ~5 million ...
'' had been named the same year. French naturalist
Georges Cuvier Jean Léopold Nicolas Frédéric, baron Cuvier (23 August 1769 – 13 May 1832), known as Georges Cuvier (; ), was a French natural history, naturalist and zoology, zoologist, sometimes referred to as the "founding father of paleontology". Cuv ...
also described known fossil proboscidean species back in 1796, although his account was later published in 1799. He considered that the remains uncovered from Siberia were true "mammoths" that had similar dentitions to extant elephants but had some morphological differences. He mentioned the fossil remains that were brought back by Longueil from Ohio back in 1739 and several researchers from previous decades who noted the unusual molars and thought that they belonged to different animals like hippopotamuses. He followed recognition in the previously established species "''Elephas americanus''" and argued that the species was different from elephants and mammoths and cannot be found amongst living animals due to extinction from
catastrophism In geology, catastrophism is the theory that the Earth has largely been shaped by sudden, short-lived, violent events, possibly worldwide in scope. This contrasts with uniformitarianism (sometimes called gradualism), according to which slow inc ...
. The proboscidean species was subject to several other species names given by other taxonomists within the earliest 18th century as well as the genus name ''Harpagmotherium'' by the Russian naturalist
Gotthelf Fischer von Waldheim Gotthelf Fischer von Waldheim (; 13 October 1771 – 18 October 1853) was a Saxon anatomist, entomologist and paleontologist. Fischer was born as Gotthilf Fischer in Waldheim, Saxony, the son of a linen weaver. He studied medicine at Leipz ...
in 1808.


Cuvier's taxonomy

In 1806, Cuvier wrote multiple extended research articles on fossil proboscideans of Eurasia and the Americas. He stated that the bones that Buffon previously described from North America were not of elephants but another animal that he referred to as the "''mastodonte''," or the "''animal of Ohio''." He reinforced the idea that the extinct "mastodon" was an animal close in relationship to elephants that differed by jaws with large tubercles. He suggested that "mammoth" and "carnivorous elephant" be discontinued as names for the species and that it receive a new genus name instead. Cuvier said that for "''mastodonte''," he derived the name's etymology (compound (, "breast") + (, "tooth")) from
Ancient Greek Ancient Greek (, ; ) includes the forms of the Greek language used in ancient Greece and the classical antiquity, ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Greek ...
to mean "nipple tooth," since he thought that it expressed the characteristic form of the teeth. In 1817, the French naturalist officially established the genus name ''Mastodon'', reaffirming that it is extinct and has left no living descendants. He established that it had an overall body form similar to elephants but had molars more similar to hippopotamuses and pigs that did not serve to grind meat. The first species he erected within ''Mastodon'' was ''Mastodon giganteum'', giving it the informal name "great mastodon" and writing that that it is designated to the Ohio proboscidean with abundant fossil evidence, equal size but greater proportions to modern elephants, and diamond-shaped points of the molars. The naturalist also created the second species name ''Mastodon angustidens'' and gave it the informal name "narrow-toothed mastodon," diagnosing it as having narrower molars, smaller sizes compared to ''M. giganteum'', and range distributions in Europe and South America. Cuvier also erected several other species of ''Mastodon'' originating from other continents in 1824. Despite Cuvier's genus name being younger than multiple other genus names, ''Mastodon'' became the most commonly used genus name for the 19th century.


Taxonomic problems

"''Mastodon''" was riddled with major taxonomic problems since species now determined as belonging to other proboscidean genera were classified to ''Mastodon'' on the basis of similar dentitions to that of "''Mastodon giganteum''" (= ''Mammut americanum''), effectively making it a
wastebasket taxon Wastebasket taxon (also called a wastebin taxon, dustbin taxon or catch-all taxon) is a term used by some taxonomists to refer to a taxon that has the purpose of classifying organisms that do not fit anywhere else. They are typically defined by e ...
. Various fossil proboscidean species were classified into ''Mastodon'' in the 19th century before eventually being reclassified into distinct genera. In addition to still-valid species names, several synonymous or dubious species names ultimately belonging to different genera were erected within the Americas as well throughout the 19th century. Also, many species names erected based on ''M. americanum'' remains were erected. As a result, ''M. americanum'' has many synonymous names. The issue of synonymous species names were especially apparent in the first half of the 19th century. Today, the genera that include species formerly classified into ''Mastodon'' include ''
Gomphotherium ''Gomphotherium'' (; "nail beast" for its double set of straight tusks) is an extinct genus of gomphothere proboscidean from the Neogene of Eurasia, Africa and North America. It is the most diverse genus of gompothere, with over a dozen valid sp ...
'' (''G. angustidens'', ''G. pyrenaicum'', ''G. productum'', ''G. libycum'', ''G. subtapiroideum'', ''G. steinheimense''), ''
Zygolophodon ''Zygolophodon'' is an extinct genus of mammutid proboscidean that lived during the Miocene in Africa, Eurasia, and North America. Description As with other mammutids, the molars have a zygodont morphology. The fused front region of the lower ...
'' (''Z. turicensis'', ''Z. proavus''), ''
Cuvieronius ''Cuvieronius'' is an extinct New World genus of gomphothere which ranged from southern North America to western South America during the Pleistocene epoch. Reaching a shoulder height of and a body mass of , it was on average shorter but compara ...
'' (''C. hyodon''), ''
Stegodon ''Stegodon'' (from the Ancient Greek στέγω (''stégō''), meaning "to cover", and ὀδούς (''odoús''), meaning "tooth", named for the distinctive ridges on the animal's molars) is an extinct genus of proboscidean, related to elephants ...
'' (''S. elephantoides''), ''
Stegolophodon ''Stegolophodon'' is an extinct genus of Stegodontidae, stegodontid proboscideans. It lived during the Miocene epoch in Asia. The earliest fossils are known from the Early Miocene, with one of the oldest fossils being from Japan, estimated to be ...
'' (''S. latidens'', ''S. cautleyi''), ''
Anancus ''Anancus'' is an extinct genus of "tetralophodont gomphothere" native to Afro-Eurasia, that lived from the Tortonian stage of the late Miocene until its extinction during the Early Pleistocene, roughly from 8.5–2 million years ago. Taxonomy ...
'' (''A. avernensis'', ''A. sivalensis'', ''A. perimensis''), ''
Tetralophodon ''Tetralophodon'' ("four-ridged tooth") is an extinct genus of "tetralophodont gomphothere" belonging to the superfamily Elephantoidea, known from the Miocene of Afro-Eurasia. Taxonomy and evolution The genus ''Tetralophodon'' (meaning "four-rid ...
'' (''T. longirostris''), ''
Choerolophodon ''Choerolophodon'' is an extinct genus of proboscidean that lived during the Miocene of Eurasia and Africa. Fossils of ''Choerolophodon'' have been found in Africa, Southeast Europe, Turkey, Iraq, Iran, the Indian subcontinent, and China.Chunxia ...
'' (''C. pentelici''), ''
Stegomastodon ''Stegomastodon'' ('roof breast tooth') is an extinct genus of gomphotheres. It ranged throughout North America from the Pliocene (early Blancan ~4 Ma), to the Early Pleistocene (early Irvingtonian, ~1.2 Ma). The former South American species h ...
'' (''S. mirificus''), ''
Rhynchotherium ''Rhynchotherium'' is an extinct genus of proboscidea endemic to North America and Central America during the Miocene through Pliocene from 13.650 to 3.6 Annum, Ma, living for approximately . This gomphothere had two tusks and may have evolved f ...
'' ("''R.''" ''euhypodon''), ''
Stenobelodon ''Stenobelodon'' is an extinct genus of amebelodont proboscidean from the Miocene of North America. The only known species was formerly considered to belong to the genus ''Amebelodon ''Amebelodon'' is a genus of extinct proboscidean belongin ...
'' (''S. floridanus''), and ''
Notiomastodon ''Notiomastodon'' is an extinct genus of gomphothere proboscidean (related to modern elephants), endemic to South America from the Pleistocene to the early Holocene. ''Notiomastodon'' specimens reached a size similar to that of the modern Asian ...
'' (''N. platensis''). In 1830, American naturalist John Davidson Godman created the genus ''Tetracaulodon'' plus its species ''T. Mastodontoideum'' based on what he determined to be differences between it and ''Mastodon'' based on the skull and dentition. Both
Richard Harlan Richard Harlan (September 19, 1796 – September 30, 1843) was an American paleontologist, anatomist, and physician. He was the first American to devote significant time and attention to vertebrate paleontology and was one of the most importan ...
and
William Cooper William Cooper may refer to: Business *William Cooper (accountant) (1826–1871), founder of Cooper Brothers * William Cooper (businessman) (1761–1840), Canadian businessman * William Cooper (co-operator) (1822–1868), English co-operator * Wil ...
pointed out that except for the tusks, all other characteristics of the specimens were consistent with ''M. giganteum''. They therefore argued that there was no reason to assume that the tusks were not just individual variations, a view followed also by George William Featherstonhaugh. Isaac Hays comparatively defended Godman's taxon, which led to a bitter debate regarding the validity of the genus amongst American naturalists. The validities of both ''Tetracaulodon'' and ''Missourium'' were rejected by Owen in 1842, although he retained the former name informally. By 1869, American paleontologist
Joseph Leidy Joseph Mellick Leidy (September 9, 1823 – April 30, 1891) was an American paleontologist, parasitologist and anatomist. Leidy was professor of anatomy at the University of Pennsylvania, later becoming a professor of natural history at Swarth ...
determined that ''Mastodon americanus'' is the senior species synonym and listed ''M. giganteum'' as a junior synonym. He also listed ''Mammut'', ''Harpagmotherium'', ''Mastotherium'', ''Missourium'', and ''Leviathan'' as synonyms of ''Mastodon''. He also noted that ''M. americanum'' as a species was highly variable in morphology. In 1902, American paleontologist
Oliver Perry Hay Oliver Perry Hay (May 22, 1846 – November 2, 1930) was an American herpetologist, ichthyologist, and paleontologist. Hay was born in Jefferson County, Indiana, to Robert and Margaret Hay. In 1870, Hay graduated with a bachelor of arts from ...
listed ''Mammut'' as the prioritized genus name given its status as the oldest genus name, making ''Mastodon'', ''Tetracaulodon'', and ''Missourium'' classified as junior synonyms. He also established ''M. americanum'' as the type species. The genus name ''Mastodon'' was subsequently abandoned by many American paleontologists in favor of ''Mammut'' within the early 20th century. In 1942, American paleontologist
George Gaylord Simpson George Gaylord Simpson (June 16, 1902 – October 6, 1984) was an American paleontologist. Simpson was perhaps the most influential paleontologist of the twentieth century, and a major participant in the modern synthesis, contributing '' Tempo ...
said that for his study, he prioritized the historic plus taxonomically correct name ''Mammut'' over ''Mastodon''. He continued prioritizing ''Mammut'' in 1945, stating that people were generally aware of its taxonomic priorities over ''Mastodon'' and that people had refused to use it. He stated that he did not want to either but reluctantly set aside his personal preferences to follow taxonomic rules.


Additional species

In 1921, Osborn created the species name ''Mastodon matthewi'' based on distinct molars from the Snake Creek Formation of western
Nebraska Nebraska ( ) is a landlocked U.S. state, state in the Midwestern United States, Midwestern region of the United States. It borders South Dakota to the north; Iowa to the east and Missouri to the southeast, both across the Missouri River; Ka ...
, naming it in honor of
William Diller Matthew William Diller Matthew FRS (February 19, 1871 – September 24, 1930) was a vertebrate paleontologist who worked primarily on mammal fossils, although he also published a few early papers on mineralogy, petrological geology, one on botany, one on ...
. He also erected another species ''M. merriami'' from the Thousand Creek Formation in
Nevada Nevada ( ; ) is a landlocked state in the Western United States. It borders Oregon to the northwest, Idaho to the northeast, California to the west, Arizona to the southeast, and Utah to the east. Nevada is the seventh-most extensive, th ...
, which was eventually synonymized with ''Zygolophodon proavus''. Osborn in 1926 followed up for ''Mastodon matthewi'' by establishing the genus ''Pliomastodon'' for the species based on cranial differences from "''Miomastodon''" (= ''Zygolophodon''). In 1930, Matthew erected a second species for ''Pliomastodon'' named ''P. vexillarius'' based on fossil material from the locality of Elephant Hill in
California California () is a U.S. state, state in the Western United States that lies on the West Coast of the United States, Pacific Coast. It borders Oregon to the north, Nevada and Arizona to the east, and shares Mexico–United States border, an ...
, determining that it differs from ''Mammut'' by differences in the skull and that the etymology of the species name was made in honor of paleontological contributions by the
Standard Oil Company Standard Oil Company was a corporate trust in the petroleum industry that existed from 1882 to 1911. The origins of the trust lay in the operations of the Standard Oil Company (Ohio), which had been founded in 1870 by John D. Rockefeller. The ...
of California. In 1933,
Childs Frick Childs Frick (March 12, 1883 - May 8, 1965) was an American vertebrate paleontologist. He was a trustee of the American Museum of Natural History and a major benefactor of its Department of Paleontology, which in 1916 began a long partnership wi ...
named the species ''Mastodon raki'' from the locality of
Truth or Consequences ''Truth or Consequences'' is an American game show originally hosted on NBC radio by Ralph Edwards (1940–57) and later on television by Edwards (1950–54), Jack Bailey (1954–56), Bob Barker (1956–75), Steve Dunne (1957–58), Bob Hi ...
,
New Mexico New Mexico is a state in the Southwestern United States, Southwestern region of the United States. It is one of the Mountain States of the southern Rocky Mountains, sharing the Four Corners region with Utah, Colorado, and Arizona. It also ...
based on differences on the
heel The heel is the prominence at the posterior end of the foot. It is based on the projection of one bone, the calcaneus or heel bone, behind the articulation of the bones of the lower leg. Structure To distribute the compressive forces exerted ...
and M3 tooth from ''M. americanus'', otherwise having proportions similar to it. In 1936, Chester Stock published the species name ''Pliomastodon nevadanus'' based on fossils from the Thousand Creek Beds of northwestern Nevada. In 1937, John R. Schultz created the species name ''Pliomastodon? cosoensis'', naming it after the Coso Mountains in
Inyo County Inyo County () is a county in the eastern central part of the U.S. state of California, located between the Sierra Nevada and the state of Nevada. In the 2020 census, the population was 19,016. The county seat is Independence. Inyo County is ...
, California where skull fossils were recovered. In 1963, J. Arnold Shotwell and Donald E. Russell designated another species ''Mammut (Pliomastodon) furlongi'', assigning it to fossils collected from the Juntura Formation of Oregon. The species name was created in honor of Eustace L. Furlong, who made early fossil collections from the western side of the Juntura Basin. The genus ''Pliomastodon'' was synonymized with ''Mammut'' while ''Miomastodon'' was synonymized with ''Zygolophodon'' by
Jeheskel Shoshani Jeheskel "Hezy" Shoshani (; January 23, 1943 – May 21, 2008) was an evolutionary biologist who studied elephants and their relatives for over 35 years. Life and work Early life and career Shoshani was born in what is now Tel Aviv, Israel ...
and Pascal Tassy in a 1996 appendix, a view that was followed by other authors in later years. In 2019, Alton C. Dooley Jr. et al. established ''Mammut pacificus'' based on fossils collected from the
Diamond Valley Lake Diamond Valley Lake is a man-made off-stream reservoir located near Hemet, California, United States. It is one of the largest reservoirs in Southern California and one of the newest. It has a capacity of . The lake nearly doubled the area's s ...
in
Hemet Hemet is a city in the San Jacinto Valley in Riverside County, California, United States. It covers a total area of , about half of the valley, which it shares with the neighboring city of San Jacinto. The population was 89,833 at the 2020 ...
, California. They also stated that ''M. oregonense'' is a ''
nomen dubium In binomial nomenclature, a ''nomen dubium'' (Latin for "doubtful name", plural ''nomina dubia'') is a scientific name that is of unknown or doubtful application. Zoology In case of a ''nomen dubium,'' it may be impossible to determine whether a ...
'' and that further analysis needs to be done to confirm whether or not ''M. furlongi'' belongs to ''Zygolophodon'' instead. In 2023, Wighart von Koenigswald et al. reviewed the North American species of ''Zygolophodon'' and ''Mammut''. They synonymized ''P. adamsi'' and ''P. sellardsi'' with ''Mammut matthewi'' and emended ''M. nevadanus'' and ''M. pacificus'' to ''M. nevadanum'' and ''M. pacificum'', respectively. They also said that they were uncertain of the taxonomic status of ''M. furlongi'', specifically whether or not it was a variant of
sexual dimorphism Sexual dimorphism is the condition where sexes of the same species exhibit different Morphology (biology), morphological characteristics, including characteristics not directly involved in reproduction. The condition occurs in most dioecy, di ...
of ''Z. proavus''. Some authors have considered ''M. nevadanum'' to be synonymous with ''M. matthewi'' while others had retained validity of the species name. Several mammutid species outside of North America are classified to ''Mammut'' (or "''Pliomastodon''"), namely ''M. borsoni'', ''M. obliquelophus'', ''M. zhupengensis'', and ''M. lufugense'' (possibly synonymous with ''M. obliquelophus''). Recent research such as that of von Koenigswald et al. in 2023 warned that the genus ''Mammut'' should be carefully used for non-North American species.


Classification and evolution

''Mammut'' is the
type genus In biological taxonomy, the type genus (''genus typica'') is the genus which defines a biological family and the root of the family name. Zoological nomenclature According to the International Code of Zoological Nomenclature, "The name-bearin ...
of the
Mammutidae Mammutidae is an extinct family of proboscideans belonging to Elephantimorpha. It is best known for the mastodons (genus ''Mammut''), which inhabited North America from the Late Miocene (around 8 million years ago) until their extinction at the ...
, the sole family of the elephantimorph clade Mammutida (the other elephantimorph clade is
Elephantida Elephantimorpha is a clade of proboscideans that contains the Mammutidae (mastodons), as well as Elephantida ( amebelodonts, choerolophodonts, gomphotheres, stegodontids and elephantids). All members of Elephantimorpha have the horizontal toot ...
). The Mammutidae is characterized by molars with zygodont-form crests, which have remained morphologically conservative throughout the evolutionary history of the family. ''Mammut'' is considered to be a derived genus of the family because of strong zygodont development. As a family of the Elephantimorpha clade, it is only distantly related to the
Deinotheriidae Deinotheriidae ("terrible beasts") is a family of prehistoric elephant-like proboscideans that lived during the Cenozoic era, first appearing in Africa during the Oligocene then spreading across Europe and the lower latitudes of Asia during the ...
due to major differences in dentition and emergence of adult teeth. The Mammutidae is identified as a
monophyletic In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent co ...
clade, meaning that it did not leave any derived descendant groups in its evolutionary history. The monophyly of the Mammutidae makes it differ from the Elephantida, where the
Gomphotheriidae Gomphotheres are an extinct group of proboscideans related to modern elephants. First appearing in Africa during the Oligocene, they dispersed into Eurasia and North America during the Miocene and arrived in South America during the Pleistocene ...
is
paraphyletic Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...
(or ancestral to more derived descendant groups in the cladistic sense) in relation to the derived elephantoid families
Stegodontidae Stegodontidae is an extinct family of proboscideans from Africa and Asia (with a single occurrence in Europe) from the Early Miocene (at least 17.3 million years ago) to the Late Pleistocene. It contains two genera, the earlier ''Stegolophodon' ...
and
Elephantidae Elephantidae is a family (biology), family of large, herbivorous proboscidean mammals which includes the living Elephant, elephants (belonging to the genera ''Elephas'' and ''Loxodonta''), as well as a number of extinct genera like ''Mammuthus'' ...
(elephants, mammoths, and relatives). Although the separation of the Mammutida and Elephantida is strongly supported based on morphological differences, their origins within the late
Paleogene The Paleogene Period ( ; also spelled Palaeogene or Palæogene) is a geologic period and system that spans 43 million years from the end of the Cretaceous Period Ma (million years ago) to the beginning of the Neogene Period Ma. It is the fir ...
remain uncertain. One hypothesis asserts that the Elephantimorpha is monophyletic if the primitive
Elephantiformes Elephantiformes is a suborder within the order Proboscidea. Members of this group are primitively characterised by the possession of upper tusks, an elongated mandibular symphysis (the frontmost part of the lower jaw) and lower tusks, and the ret ...
genus ''
Phiomia ''Phiomia'' (after the Ancient Greek ''phiom'' "lake", an ancient name for the Fayum), is an extinct genus of basal elephantiform proboscidean that lived in what is now Northern Africa during the Late Eocene to Early Oligocene some 37–30 milli ...
'' was truly ancestral to both the Elephantida and Mammutida. An alternate hypothesis suggests that the Elephantimorpha is diphyletic because ''Phiomia'' is ancestral to gomphotheres while ''
Palaeomastodon ''Palaeomastodon'' ("ancient mastodon") is an extinct genus of basal proboscideans from the Oligocene of North Africa. The first specimen discovered was recovered from strata belonging to the Fayum fossil deposits of Egypt. It was described and ...
'' is ancestral to mammutids. The earliest undisputed mammutid genus ''
Losodokodon ''Losodokodon'' is an extinct genus of large herbivorous mammals belonging to the family Mammutidae. It was first described in 2009 by David Tab Rasmussen and Mercedes Gutiérrez from fossils found in the Erageleit Formation of northwestern Keny ...
'' is recorded in
Kenya Kenya, officially the Republic of Kenya, is a country located in East Africa. With an estimated population of more than 52.4 million as of mid-2024, Kenya is the 27th-most-populous country in the world and the 7th most populous in Africa. ...
, Africa and firmly establishes the earliest presence of mammutids in the late
Oligocene The Oligocene ( ) is a geologic epoch (geology), epoch of the Paleogene Geologic time scale, Period that extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that defin ...
(~27-24 Ma). The Mammutidae, like other Paleogene proboscideans, was therefore an endemic radiation within the continent akin to other endemic mammals like arsinoitheres, hyracoids, and
catarrhine The parvorder Catarrhini (known commonly as catarrhine monkeys, Old World anthropoids, or Old World monkeys) consists of the Cercopithecoidea and apes (Hominoidea). In 1812, Geoffroy grouped those two groups together and established the name Ca ...
primates Primates is an order of mammals, which is further divided into the strepsirrhines, which include lemurs, galagos, and lorisids; and the haplorhines, which include tarsiers and simians ( monkeys and apes). Primates arose 74–63  ...
plus non-endemics such as
anthracothere Anthracotheriidae is a paraphyletic family of extinct, hippopotamus-like artiodactyl ungulates related to hippopotamuses and whales. The oldest genus, '' Elomeryx'', first appeared during the middle Eocene in Asia. They thrived in Africa and Eura ...
s and hyaenodonts. In the early
Neogene The Neogene ( ,) is a geologic period and system that spans 20.45 million years from the end of the Paleogene Period million years ago ( Mya) to the beginning of the present Quaternary Period million years ago. It is the second period of th ...
phase of evolution, ''
Eozygodon ''Eozygodon'' is an extinct genus of proboscidean in the family Mammutidae. It is a monotypic genus that contains the single species ''E. morotoensis'', named in 1983. It is known from the Early Miocene of Africa (Kenya, Uganda, Namibia) and we ...
'' made an appearance in the earliest
Miocene The Miocene ( ) is the first epoch (geology), geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and mea ...
(~23-20 Ma) of Africa after ''Losodokodon''. ''Eozygodon'' was subsequently succeeded by ''Zygolophodon'' by the early Miocene, and the latter dispersed into Eurasia by around 19-18 million years ago, and into North America by the middle Miocene. The dispersal of mammutids between Africa and Eurasia may have occurred multiple times. The Mammutidae eventually went extinct in Africa prior to the late Miocene. ''Mammut'' as currently defined ''sensu lato'' (in a broad sense) is most likely
polyphyletic A polyphyletic group is an assemblage that includes organisms with mixed evolutionary origin but does not include their most recent common ancestor. The term is often applied to groups that share similar features known as Homoplasy, homoplasies ...
(comprising several unrelated groups). This is because the inclusion of Eurasian mammutid species into ''Mammut'' implies that they share a common origin with North American ''Mammut'', but this relationship has been doubted. As a result, these Eurasian species may belong to either other existing mammutid genera or entirely new genera. ''
"Mammut" borsoni ''"Mammut" borsoni'' (sometimes called Borson's mastodon) is an extinct species of mammutid proboscidean known from the Late Miocene to Early Pleistocene of Eurasia, spanning from western Europe to China. It is the last known mammutid in Eurasi ...
,'' the last Eurasian mammutid, became extinct during the earliest
Pleistocene The Pleistocene ( ; referred to colloquially as the ''ice age, Ice Age'') is the geological epoch (geology), epoch that lasted from to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was fin ...
, around 2.5-2 million years ago. The oldest evidence of mammutids in North America is of a fragmentary molar of ''Zygolophodon'' sp. from Massacre Lake, Nevada, dating to 16.5-16.4 Ma (during the
Hemingfordian The Hemingfordian on the geologic timescale is the North American faunal stage according to the North American Land Mammal Ages chronology (NALMA), typically set from 20,600,000 to 16,300,000 years BP. It is usually considered to overlap the lates ...
stage of the
North American land mammal age The North American land mammal ages (NALMA) establishes a geologic timescale for North American fauna beginning during the Late Cretaceous and continuing through to the present. These periods are referred to as ages or intervals (or stages when ref ...
s (NALMA)). The only definitively defined species of ''Zygolophodon'' from North America is ''Z. proavus'', which occurs in the
Barstovian The Barstovian North American Stage on the geologic timescale is the North American faunal stage according to the North American Land Mammal Ages chronology (NALMA), typically set from 16,300,000 to 13,600,000 years BP, a period of . It is usua ...
and
Clarendonian The Clarendonian North American Stage on the geologic timescale is the North American faunal stage according to the North American Land Mammal Ages chronology (NALMA), typically set from 13,600,000 to 9,000,000 years BP, a period of . It is us ...
stages. ''M? furlongi'' from the Black Butte in Oregon also dates back to the Clarendonian stage, but the affinities of the species remains unclear. If it truly is a species of ''Mammut'', then its earliest temporal range is recorded at about 10 Ma. The earliest undisputed appearance of ''Mammut'' is of ''M. nevadanum'' from Thousand Creek Beds, dating back to the early
Hemphillian The Hemphillian North American Stage on the geologic timescale is a North American faunal stage according to the North American Land Mammal Ages chronology (NALMA), typically set from 10,300,000 to 4,900,000 years BP. It is usually considered t ...
, or 8.0-7.1 Ma. Historically, North American paleontologists considered that North American ''Zygolophodon'' evolved into ''Mammut'' in an endemic fashion while European workers generally thought that ''Mammut'' was a Eurasian immigrant that replaced North American ''Zygolophodon'' during the Miocene or
Pliocene The Pliocene ( ; also Pleiocene) is the epoch (geology), epoch in the geologic time scale that extends from 5.33 to 2.58 ''M. matthewi'' is recorded from the late Hemphillian to early
Blancan The Blancan North American Stage on the geologic timescale is the North American faunal stage according to the North American Land Mammal Ages chronology (NALMA), typically set from 4,750,000 to 1,806,000 years BP, a period of . ''M. americanum'' (known popularly as an "American mastodon" or simply "mastodon") is also stratigraphically recorded first from the early Blancan of the
Ringold Formation The Ringold Formation is a Formation (geology), geologic formation in Eastern Washington (state), Washington, United States. The formation consists of sediment laid down by the Columbia River following the flood basalt eruptions of the Columbia R ...
,
Washington Washington most commonly refers to: * George Washington (1732–1799), the first president of the United States * Washington (state), a state in the Pacific Northwest of the United States * Washington, D.C., the capital of the United States ** A ...
. The age of the formation where the mammutid specimen was found dates to about 3.75 Ma. It is also known from multiple other Blancan sites such as Fish Springs Flat in Nevada. From the
Irvingtonian The Irvingtonian North American Land Mammal Age on the geologic timescale is the North American faunal stage according to the North American Land Mammal Ages chronology (NALMA), spanning from 1.8 million – 250,000 years Before Present, BP.
to the
Rancholabrean The Rancholabrean North American Land Mammal Age on the geologic timescale is a North American faunal stage in the North American Land Mammal Ages chronology (NALMA),Sanders, A.E., R.E. Weems, and L.B. Albright III (2009) Formalization of the mid- ...
(from around 1.6 million to 11,000 years ago), only ''M. americanum'' and the newly appearing ''M. pacificum'' are recorded, the former having an exceptional level of diversity based on abundant skeletal evidences from the late Pleistocene that is unusual for the typical mammutid fossil record. The following cladogram defines the phylogeny of certain proboscideans, a majority known from
endocast An endocast is the internal cast of a hollow object, often referring to the cranial vault in the study of brain development in humans and other organisms. Endocasts can be artificially made for examining the properties of a hollow, inaccessible ...
s, including ''M. americanum'':


Description


Skull

''Mammut'' is diagnosed and differentiated in terms of the skull from ''Zygolophodon'' as having a shortened bottom skull base (basicranium) and a high-domed
cranium The skull, or cranium, is typically a bony enclosure around the brain of a vertebrate. In some fish, and amphibians, the skull is of cartilage. The skull is at the head end of the vertebrate. In the human, the skull comprises two prominent ...
. It is also diagnosed as having an "elephantoid"
mandible In jawed vertebrates, the mandible (from the Latin ''mandibula'', 'for chewing'), lower jaw, or jawbone is a bone that makes up the lowerand typically more mobilecomponent of the mouth (the upper jaw being known as the maxilla). The jawbone i ...
with a shortened
mandibular symphysis In human anatomy, the facial skeleton of the skull the external surface of the mandible is marked in the median line by a faint ridge, indicating the mandibular symphysis (Latin: ''symphysis menti'') or line of junction where the two lateral ha ...
(or "brevirostrine") and a protruding angular process in the mandible. The diagnosis accounts for both true ''Mammut'' species and ''Mammut'' species pending reassessments. The shortening of the symphysis is one of the major evolutionary trends observed in Neogene mammutids, making it critical in understanding the evolutionary transition from ''Zygolophodon'' to ''Mammut''. However, mandibular remains with characteristics of ''Mammut'' are not known from any anywhere within the Hemphillian, thus making the transition poorly understood. It differs from ''
Sinomammut ''Sinomammut'' (meaning "Chinese ''Mammut''") is a mammutid proboscidean from the Miocene of China. Only one species, ''S. tobieni'', is known, named in 2016. Discovery and naming It was known from the holotype, specimen GIOTC 0984-9-178, whi ...
'' by the shortened mandibular symphysis, although ''Mammut'' sometimes retained lower tusks unlike the other genus. The shortened mandibular symphysis in ''Mammut'' and the similarity of its skull with modern elephants would have allowed for an elephant-like prehensile trunk perhaps long enough to reach the ground. ''M. americanum'' is diagnosed as having a long plus low skull and a shortened mandible. The
frontal bone In the human skull, the frontal bone or sincipital bone is an unpaired bone which consists of two portions.'' Gray's Anatomy'' (1918) These are the vertically oriented squamous part, and the horizontally oriented orbital part, making up the bo ...
(or forehead) gives off a flattened appearance compared to extant elephants. The skull of ''M. americanum'' has many
plesiomorphies In phylogenetics, a plesiomorphy ("near form") and symplesiomorphy are synonyms for an ancestral character shared by all members of a clade, which does not distinguish the clade from other clades. Plesiomorphy, symplesiomorphy, apomorphy, an ...
(or ancestral traits) that can be observed, namely the low and flat brain case, a slightly vertical basicranium, a narrow nasal aperture inlet of the nose with no step-like perinasal fossa, and a backside
infraorbital foramen In human anatomy, the infraorbital foramen is one of two small holes in the skull's upper jawbone ( maxillary bone), located below the eye socket and to the left and right of the nose. Both holes are used for blood vessels and nerves. In anatomic ...
. At least some of these features are thought to have been acquired from ''Phiomia''. The nasal aperture of ''M. americanum'' is oval, whereas that of the skull of "''M.''" cf. ''obliquelophus'' is more trapezoidal. ''M. americanum'' is also more derived based on the lack of a strong proximal constriction of the incisive fossa of the
incisive foramen In the human mouth, the incisive foramen (also known as: "''anterior palatine foramen''", or "''nasopalatine foramen''") is the opening of the incisive canals on the hard palate immediately behind the incisor teeth. It gives passage to blood ves ...
. ''M. americanum'' also has a high and narrow
orbit In celestial mechanics, an orbit (also known as orbital revolution) is the curved trajectory of an object such as the trajectory of a planet around a star, or of a natural satellite around a planet, or of an artificial satellite around an ...
with a somewhat rectangular outline, but it is less rectangular than that of ''Eozygodon''. The North American mammutid retains a primitive trait in the form of the orbit containing a
lacrimal bone The lacrimal bones are two small and fragile bones of the facial skeleton; they are roughly the size of the little fingernail and situated at the front part of the medial wall of the orbit. They each have two surfaces and four borders. Several bon ...
with a hole known as the lacrimal foramen. Unlike elephantidans, it has another primitive trait of a short and high-positioned
temporal fossa The temporal fossa is a fossa (shallow depression) on the side of the skull bounded by the temporal lines above, and the zygomatic arch below. Its floor is formed by the outer surfaces of four bones of the skull. The fossa is filled by the te ...
, a trait shared with ''Eozygodon''.


Endocast anatomy

''M. americanum'' is known by several brain endocasts stored in American museums, although they are seldom subjected to studies. In 1973, neuroscientist Harry J. Jerison studied an endocast of ''Mammut'', recording that it was elephantlike in both size and shape. According to Shoshani et al. in 2006, the endocast of ''M. americanum'' features the
olfactory bulb The olfactory bulb (Latin: ''bulbus olfactorius'') is a neural structure of the vertebrate forebrain involved in olfaction, the sense of smell. It sends olfactory information to be further processed in the amygdala, the orbitofrontal cortex (OF ...
s protruding in front of the
frontal lobe The frontal lobe is the largest of the four major lobes of the brain in mammals, and is located at the front of each cerebral hemisphere (in front of the parietal lobe and the temporal lobe). It is parted from the parietal lobe by a Sulcus (neur ...
. They also drew several proboscidean brains to scale, in which the brain of ''M. americanum'' was much larger than that of ''
Moeritherium ''Moeritherium'' ("the beast from Lake Moeris") is an extinct genus of basal proboscideans from the Eocene of North and West Africa. The first specimen was discovered in strata from the Fayum fossil deposits of Egypt. It was named in 1901 by C ...
lyonsi'' but smaller than that of the
Asian elephant The Asian elephant (''Elephas maximus''), also known as the Asiatic elephant, is the only living ''Elephas'' species. It is the largest living land animal in Asia and the second largest living Elephantidae, elephantid in the world. It is char ...
(''Elephas maximus''). Julien Benoit et al. in 2022 explained that while the front tips of the olfactory bulbs of "''M.''" ''borsoni'' are partially visible in the brain's back (or dorsal) area, its visibility in ''M. americanum'' is debated. Some authors had argued that the olfactory bulbs are visible in the brain's back area while some other authors did not portray them as being visible. The researchers confirmed based on one specimen that the olfactory bulbs are only partially visible in the brain's back area. They also observed that "''M.''" ''borsoni'', despite weighing twice as much as ''M. americanum'', had a 30% lower
encephalization quotient Encephalization quotient (EQ), encephalization level (EL), or just encephalization is a relative brain size measure that is defined as the ratio between observed and predicted brain mass for an animal of a given size, based on nonlinear regre ...
(EQ) compared to the other mammutid species, supporting the idea that the evolution of proboscidean encephalization is tied with phylogeny. The Mammutida, as the most basal clade of the Elephantimorpha, has an EQ twice that of ''Moeritherium'' and ''Palaeomastodon''. The endocast volume and brain size of the brain ''M. americanum'' are larger than those of ''Stegodon'' but smaller than those of derived elephantids. It has an EQ that is higher than those of Paleogene proboscideans and "''M.''" ''borsoni'' but lower than those of elephantids (extant and extinct) and stegodonts. The type species is also known from endocasts of ear petrosals. According to Eric G. Ekdale, the ear petrosals of ''Mammut'' cannot automatically be distinguished from ''Mammuthus'' alone. The subarcuate fossa is absent from the cerebellar surface of the inner ear. The ear petrosals of ''Mammut'' are relatively incomplete, leaving several traits to be unable to be observed.


Dentition

The family Mammutidae is defined by zygolophodont molars with compressed and sharp transverse ridges plus lack of accessory conules (smaller cusps). The intermediate molars, or the first two molars, are consistently trilophodont, or three-cusped. The dental morphologies of the clade Mammutida contrast strongly with most members of both the Elephantida (
bunodont 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 tooth ...
molars that evolutionarily convert to being thin and platelike) and the Deinotheriidae (
tapir Tapirs ( ) are large, herbivorous mammals belonging to the family Tapiridae. They are similar in shape to a Suidae, pig, with a short, prehensile nose trunk (proboscis). Tapirs inhabit jungle and forest regions of South America, South and Centr ...
-like
lophodont 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 tooth ...
to bilophodont molars). The zygodont morphologies of the molars of mammutids were conservative, meaning that they hardly changed in the evolutionary history of the family. Mammutids also exhibited evidences of horizontal tooth displacement where milk teeth were gradually replaced by permanent molars, mirroring elephantidans in an instance of
parallel evolution Parallel evolution is the similar development of a trait in distinct species that are not closely related, but share a similar original trait in response to similar evolutionary pressure.Zhang, J. and Kumar, S. 1997Detection of convergent and pa ...
. The Mammutidae was not the only proboscidean family to have acquired zygodont crested molars, as Neogene species of the gomphothere ''
Sinomastodon ''Sinomastodon'' ("Chinese mastodont") is an extinct gomphothere genus (of order Proboscidea) known from the Late Miocene to Early Pleistocene of Asia, including China, Japan, Thailand, Myanmar, Indonesia and probably Kashmir. Extinct animals o ...
'' display moderate to weak zygodont crests. Pleistocene species of ''Sinomastodon'' do not display zygodont crests, however. The dentition of ''Mammut'' is diagnosed as being strongly zygodont and having no conules. The lophs extend to the long axis of the molars. The first two molars in the dental row have no more than three lophs while the third molars have four lophs plus a cingulum. The upper tusks (or upper
incisor Incisors (from Latin ''incidere'', "to cut") are the front teeth present in most mammals. They are located in the premaxilla above and on the mandible below. Humans have a total of eight (two on each side, top and bottom). Opossums have 18, wher ...
s) of ''Mammut'' differ from those of ''Zygolophodon'' by the generally larger sizes, tendency to either straighten or curve up, and the typical lack of any enamel band, although ''M. vexillarius'' retains a very narrow strip of enamel in the upper tusks. The lower (or mandibular) tusks tend to be reduced in comparison. ''M. nevadanum'' represents the earliest case of a North American mammutid species without any enamel band, although the possibility of it being worn off by wear cannot automatically be eliminated. It differs from ''M. americanum'' and ''M. pacificum'' by the nearly straight but downward-facing upper tusk, whereas males of the latter two species have large and upward-facing upper tusks while females had upward or straight but frontward-directed upper tusks. The reduction to loss of the lower tusks plus reduction of the mandibular symphysis of the derived Mammutidae and Elephantida is an instance of
convergent evolution Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last comm ...
, correlating potentially with the need to reduce heat loss due to the decrease of global temperature and humidity during the late Miocene and Pliocene. Despite the reductions of the lower tusks, they were still present in Neogene species of ''Mammut''. Pleistocene ''M. americanum'' comparatively often lacks mandibular tusks, and ''M. pacificum'' is always devoid of them. The presence of lower tusks in ''M. raki'' separates it as a species from ''M. pacificum''. ''M. pacificum'' differs from ''M. americanum'' in part by the narrower molars. Both species have broader molars compared to the "narrow-toothed" ''M. nevadanum'', ''M. raki'', and ''M. cosoensis''. Like its relative "''M.''" ''borsoni'', ''M. americanum'' had very large tusks, with some records suggesting lengths of and diameters exceeding were not unusual. In the skull of the earlier-appearing ''M. matthewi'', its
dental alveolus Dental alveoli (singular ''alveolus'') are sockets in the jaws in which the roots of teeth are held in the alveolar process with the periodontal ligament. The lay term for dental alveoli is tooth sockets. A joint that connects the roots of the t ...
of the right tusk from the locality of Hermiston, Oregon suggests a tusk diameter of approximately . Similar to modern elephants, ''M. americanum'' also has degrees of sexual dimorphism indicated by the sizes of the upper tusks. Adult males have tusks 1.15–1.25 times as large as those of adult females, also reflecting general body size differences between the two sexes. The sizes of the tusk also depend on the ages of the individuals, as older individuals have larger tusk circumferences than younger ones. Adult individuals of comparable ages have similar tusk sizes, but older individuals do not necessarily have larger tusk sizes. Tusk sizes may have depended on external factors like nutritional stress, geographic location, and reproductive status. The tusks of ''M. pacificum'' are thought to have been smaller in length and circumstance than that of ''M. americanum'' and may have similarly exhibited degrees of sexual dimorphism.


Postcranial skeleton

As a result of proboscidean diagnoses focusing mostly on dentition, the postcranial anatomies of fossil proboscideans like ''Mammut'' are underrepresented in academic literature. Jennifer A. Hodgson ''et al''. compared the anatomies of ''Mammut'' and ''Mammuthus'', mentioning that their postcranial anatomies were studied previously by Stanley John Olsen in 1972 and recognizing that the two genera were only distantly related to each other. ''M. americanum'' is typically depicted as stocky based on postcranial evidence. The
vertebral column The spinal column, also known as the vertebral column, spine or backbone, is the core part of the axial skeleton in vertebrates. The vertebral column is the defining and eponymous characteristic of the vertebrate. The spinal column is a segmente ...
(also known as the backbone or spine) of ''Mammut'' is documented as having a highest point located in the shoulder's front like ''Mammuthus'', but the spines gradually decrease in length then increase slightly in the rear area. The number of ribs and vertebrae of ''Mammut'' is not well-documented in paleontological literature and may vary by individual. ''Mammut'' usually has 20 
thoracic vertebrae In vertebrates, thoracic vertebrae compose the middle segment of the vertebral column, between the cervical vertebrae and the lumbar vertebrae. In humans, there are twelve thoracic vertebra (anatomy), vertebrae of intermediate size between the ce ...
whereas ''Mammuthus'' usually has 19, but both have documented individuals with 18 of them. The reduction of thoracic vertebrae in ''Mammuthus'' is considered a derived trait also present in modern elephants. The "Watkins Glen mastodon," for example, has 7 
cervical vertebrae In tetrapods, cervical vertebrae (: vertebra) are the vertebrae of the neck, immediately below the skull. Truncal vertebrae (divided into thoracic and lumbar vertebrae in mammals) lie caudal (toward the tail) of cervical vertebrae. In saurop ...
, 20 thoracic vertebrae, 3 
lumbar vertebrae The lumbar vertebrae are located between the thoracic vertebrae and pelvis. They form the lower part of the back in humans, and the tail end of the back in quadrupeds. In humans, there are five lumbar vertebrae. The term is used to describe t ...
, and 5 
sacral vertebrae The sacrum (: sacra or sacrums), in human anatomy, is a triangular bone at the base of the spine that forms by the fusing of the sacral vertebrae (S1S5) between ages 18 and 30. The sacrum situates at the upper, back part of the pelvic cavity, ...
. They believed that ''Mammut'' could have had as many as 20 
rib In vertebrate anatomy, ribs () are the long curved bones which form the rib cage, part of the axial skeleton. In most tetrapods, ribs surround the thoracic cavity, enabling the lungs to expand and thus facilitate breathing by expanding the ...
s and that the back ribs were shorter and broader than that of ''Mammuthus''. The tail of ''Mammut'' may have been made up of as many as up to 27 
caudal vertebrae Caudal vertebrae are the vertebrae of the tail in many vertebrates. In birds, the last few caudal vertebrae fuse into the pygostyle, and in apes, including humans, the caudal vertebrae are fused into the coccyx. In many reptiles, some of the caud ...
, suggesting that it had a long tail compared to gomphotheres and elephantids. The
scapula The scapula (: scapulae or scapulas), also known as the shoulder blade, is the bone that connects the humerus (upper arm bone) with the clavicle (collar bone). Like their connected bones, the scapulae are paired, with each scapula on either side ...
(or shoulder blade) of ''Mammut'' has a straight vertebral border, contrasting with a more concave vertebral border of ''Mammuthus''. Hodgson ''et al''. disagreed with the claim by Olsen in 1972 that the neck of the scapula is more constricted in ''Mammuthus primigenius'' than ''Mammut americanum'', since neither of the two ''M. americanum'' scapulae observed by the researchers have any high constriction there. The
pelvis The pelvis (: pelves or pelvises) is the lower part of an Anatomy, anatomical Trunk (anatomy), trunk, between the human abdomen, abdomen and the thighs (sometimes also called pelvic region), together with its embedded skeleton (sometimes also c ...
allows for identification of the sex of the species, as male ''Mammut'' individuals have a smaller
pelvic outlet The lower circumference of the lesser pelvis is very irregular; the space enclosed by it is named the inferior aperture or pelvic outlet. It is an important component of pelvimetry. Boundaries It has the following boundaries: * anteriorly: the pub ...
and wider ilium than female individuals. ''Mammut'' has shorter and more robust limb bones compared to those of derived elephantids, probably the result of it retaining primitive anatomical traits. Both the
humerus The humerus (; : humeri) is a long bone in the arm that runs from the shoulder to the elbow. It connects the scapula and the two bones of the lower arm, the radius (bone), radius and ulna, and consists of three sections. The humeral upper extrem ...
and
radius In classical geometry, a radius (: radii or radiuses) of a circle or sphere is any of the line segments from its Centre (geometry), center to its perimeter, and in more modern usage, it is also their length. The radius of a regular polygon is th ...
of the mammutid genus are robust for instance. The
ulna The ulna or ulnar bone (: ulnae or ulnas) is a long bone in the forearm stretching from the elbow to the wrist. It is on the same side of the forearm as the little finger, running parallel to the Radius (bone), radius, the forearm's other long ...
has a slightly more developed
olecranon process The olecranon (, ), is a large, thick, curved bony process on the proximal, posterior end of the ulna. It forms the protruding part of the elbow and is opposite to the cubital fossa or elbow pit (trochlear notch). The olecranon serves as a lever ...
and a deeper
trochlear notch The trochlear notch (), also known as semilunar notch and greater sigmoid cavity, is a large depression in the upper extremity of the ulna The ulna or ulnar bone (: ulnae or ulnas) is a long bone in the forearm stretching from the elbow to ...
. The
femur The femur (; : femurs or femora ), or thigh bone is the only long bone, bone in the thigh — the region of the lower limb between the hip and the knee. In many quadrupeds, four-legged animals the femur is the upper bone of the hindleg. The Femo ...
is somewhat thick, short, and appears to have more expanded
condyle A condyle (;Entry "condyle"
in
s. Possibly, sexual dimorphism could be a factor behind the size of the femur itself. The
tibia The tibia (; : tibiae or tibias), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two Leg bones, bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outsi ...
does not appear much different in both ''Mammut'' and ''Mammuthus'', whereas the
fibula The fibula (: fibulae or fibulas) or calf bone is a leg bone on the lateral side of the tibia, to which it is connected above and below. It is the smaller of the two bones and, in proportion to its length, the most slender of all the long bones. ...
may have only had subtle and complex differences within the two genera. The bones within both the front feet and back feet have their own subtle and complex differences by genus, but both have smaller and more narrow hind feet than fore feet so that the latter bears more weight of the proboscideans. In terms of postcranial anatomy, ''M. pacificum'' differs from ''M. americanum'' by the presence of six as opposed to five sacral vertebrae and the femur having a larger diameter of the middle shaft (or main cylindrical area).


External features

The American mastodon (''M. americanum'') has typically been depicted as having shaggy and brown-colored fur in reconstructions, especially in over a century of
paleoart Paleoart (also spelled palaeoart, paleo-art, or paleo art) is any original artistic work that attempts to depict prehistoric life according to scientific evidence. Ansón, Fernández & Ramos (2015) pp. 28–34. Works of paleoart may be represen ...
. Despite this, there is little direct evidence supporting the idea that ''Mammut'' was actually covered in hair. Supposedly, only one find of fur belonging to the mastodon is of a skull with two small hairy patches of skin from the state of
Wisconsin Wisconsin ( ) is a U.S. state, state in the Great Lakes region, Great Lakes region of the Upper Midwest of the United States. It borders Minnesota to the west, Iowa to the southwest, Illinois to the south, Lake Michigan to the east, Michig ...
near the city of
Milwaukee Milwaukee is the List of cities in Wisconsin, most populous city in the U.S. state of Wisconsin. Located on the western shore of Lake Michigan, it is the List of United States cities by population, 31st-most populous city in the United States ...
. These have only been described briefly in the original literature and have never been figured beyond one hair from a
scanning electron microscope A scanning electron microscope (SEM) is a type of electron microscope that produces images of a sample by scanning the surface with a focused beam of electrons. The electrons interact with atoms in the sample, producing various signals that ...
(SEM). K.F. Hallin and D. Gabriel in 1981 speculated that mastodons were indeed hairy but were more suited for semiaquatic lifestyles than tolerance of colder climates. Matt Davis ''et al''. in 2022 were tentative in accepting the source as evidence for hairiness, as they questioned whether ''Mammut'' needed thick coats for body warmth for their upper ranges at the
Arctic The Arctic (; . ) is the polar regions of Earth, polar region of Earth that surrounds the North Pole, lying within the Arctic Circle. The Arctic region, from the IERS Reference Meridian travelling east, consists of parts of northern Norway ( ...
and
Subarctic The subarctic zone is a region in the Northern Hemisphere immediately south of the true Arctic, north of hemiboreal regions and covering much of Alaska, Canada, Iceland, the north of Fennoscandia, Northwestern Russia, Siberia, and the Cair ...
and mentioned that it would not have needed them in subtropical climates like in
Florida Florida ( ; ) is a U.S. state, state in the Southeastern United States, Southeastern region of the United States. It borders the Gulf of Mexico to the west, Alabama to the northwest, Georgia (U.S. state), Georgia to the north, the Atlantic ...
. Davis ''et al''. referenced that because Columbian mammoths (''Mammuthus columbi'') were not thought to be hairy, it is unclear why mastodons would need thick coats in comparison. The former was typically depicted as hairless and the latter as hairy in paleoart, but the mastodon's preferences for closed or mixed habitats puts the speculations into question. They felt the need to portray the latter as hairy so that the average person could differentiate between the two species. The concept of ''M. americanum'' having thick coats of fur was also subjected to study by Asier Larramendi in 2015. He acknowledged that hair is important for thermoregulation in extant elephants but that there is a negative correlation between body size and hair density in mammals. Some mammals have broken this trend before, however, as woolly mammoths (''Mammuthus primigenius'') evolved to have thick coats of hair and a very short tail in response to cold climates. The idea that the American mastodon had hair is possible because of the seasonal climates, but there are few preserved soft tissues to support this idea, referencing the hairs found in Wisconsin. The supposed evidence of hair reported in the 19th century were actually just
green algae The green algae (: green alga) are a group of chlorophyll-containing autotrophic eukaryotes consisting of the phylum Prasinodermophyta and its unnamed sister group that contains the Chlorophyta and Charophyta/ Streptophyta. The land plants ...
filaments. He concluded that the long tail and large body mass both contradict the hypothesis that ''M. americanum'' was covered with thick coats of fur, considering it to be probably exaggerated.


Size and weight

According to Larramendi, the mammutids of the genus ''Mammut'' were among the largest known proboscideans. This was especially the case with "''M.''" ''borsoni'', males of which are suggested to have had an average body mass of making it the largest known proboscidean alongside the extinct Indian elephant species ''
Palaeoloxodon namadicus ''Palaeoloxodon namadicus'' is an extinct species of prehistoric elephant known from the Chibanian, Middle Pleistocene to Late Pleistocene of the Indian subcontinent, and possibly also elsewhere in Asia. The species grew larger than any living el ...
,'' and one of the largest land mammals to have ever lived. ''M. americanum'' in comparison to "''M.''" ''borsoni'' was much smaller, but it was still large in its own right compared to extant elephants. The American mastodon did not grow taller than living elephants but it was much more robust in body build than them, in part due to its very broad pelvis. The Warren mastodon produces a body mass of nearby and had a shoulder height measuring . This robustness is so pronounced that ''M. americanum'' individuals could have been up to 80% heavier than an elephant with the same shoulder height. Larger than average individuals may have possibly had a shoulder height of and weighed up to . 90% of fully grown male ''M. americanum'' individuals are suggested to have had shoulder heights ranging from to and body masses ranging from to in body mass, with an average fully grown ''M. americanum'' male estimated at in shoulder height and in body mass. These estimates place males as larger on average in weight and shoulder height than those of both the living
Asian elephant The Asian elephant (''Elephas maximus''), also known as the Asiatic elephant, is the only living ''Elephas'' species. It is the largest living land animal in Asia and the second largest living Elephantidae, elephantid in the world. It is char ...
and
African forest elephant The African forest elephant (''Loxodonta cyclotis'') is one of the two living species of African elephant, along with the African bush elephant. It is native to humid tropical forests in West Africa and the Congo Basin. It is the smallest of the ...
, and heavier but somewhat shorter than average males of
African bush elephant The African bush elephant (''Loxodonta africana''), also known as the African savanna elephant, is a species of elephant native to sub-Saharan Africa. It is one of three extant elephant species and, along with the African forest elephant, one ...
s. The size of the "Overmyer Mastodon," an individual skeleton recovered from the farm of Robert Overmyer northwest of Rochester,
Indiana Indiana ( ) is a U.S. state, state in the Midwestern United States, Midwestern region of the United States. It borders Lake Michigan to the northwest, Michigan to the north and northeast, Ohio to the east, the Ohio River and Kentucky to the s ...
in 1976, was estimated by Neal Woodman and Jon W. Branstrator in 2008. They estimated based on the length of the humerus () that the shoulder height of the individual was , which they said was close to the average shoulder height of the species and comparable to a large female or small male. Similar to extant elephants, male American mastodon individuals tended to be larger than female individuals and tend to have larger and more strongly curved tusks, although the degree to which the body size is a factor in molar size is unclear. A relatively complete skeleton of ''Mammut'' sp. from the Gray Fossil Site in
Tennessee Tennessee (, ), officially the State of Tennessee, is a landlocked U.S. state, state in the Southeastern United States, Southeastern region of the United States. It borders Kentucky to the north, Virginia to the northeast, North Carolina t ...
, which was first uncovered in 2015, dates to the latest Hemphillian, and has an elongated mandibular symphysis and large mandibular tusks, is thought to have been several tonnes larger than ''M. americanum'' and even several species of ''Mammuthus''. The specimens are still being prepared for further studies.


Paleobiology


Diet

The zygodont molar morphologies of mammutids suggest that they consistently occupied adaptations to folivorous diets throughout their evolutionary history. This means that mammutids such as ''Mammut'', because they retained zygodont molars, were built to browse on higher vegetation and did not shift towards grazing specializations or consistent mixed feeding. The stomach contents of ''M. americanum'' indicate that the species consumed spruce needles,
pine cone A conifer cone, or in formal botanical usage a strobilus, : strobili, is a seed-bearing organ on gymnosperm plants, especially in conifers and cycads. They are usually woody and variously conic, cylindrical, ovoid, to globular, and have scal ...
s, grass, and occasionally
gourds Gourds include the fruits of some flowering plant species in the family Cucurbitaceae, particularly ''Cucurbita'' and ''Lagenaria''. The term refers to a number of species and subspecies, many with hard shells, and some without. Many gourds have ...
plus
vine leaves Grape leaves, the leaves of the grapevine plant, are used in the cuisines of a number of cultures. They may be obtained fresh, or preserved in jars or cans. The leaves are commonly rolled or stuffed with mixtures of meat and rice to produce d ...
. Of note is that whereas mammutids of Eurasia went extinct by the early Pleistocene in association with more seasonal climates, ''Mammut'' survived in North America and became abundant, although the reason for the latter faunal trend does not have any offered explanation. The browsing specialization of ''Mammut'' is supported further by the
coprolite A coprolite (also known as a coprolith) is fossilized feces. Coprolites are classified as trace fossils as opposed to body fossils, as they give evidence for the animal's behaviour (in this case, diet) rather than morphology. The name ...
s (or fossil dung) of ''M. americanum'', which are large-sized similar to extant elephants and predominantly consist of consumed woody contents but no grass. Of the Pleistocene New World proboscideans, the American mastodon appears to have been the most consistent in browsing rather than grazing, consuming C3 as opposed to C4 plants, and occupying closed forests versus more open habitats. This dietary inflexibility may have prevented them from invading South America during the
Great American Interchange The Great American Biotic Interchange (commonly abbreviated as GABI), also known as the Great American Interchange and the Great American Faunal Interchange, was an important late Cenozoic paleozoogeographic biotic interchange event in which land ...
, due to the need to cross areas of grassland to do so. The mastodon commonly browsed on woody plants (i.e. twigs) and fruits, occupying dense
coniferous forest Conifers () are a group of cone-bearing seed plants, a subset of gymnosperms. Scientifically, they make up the division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single extant class, Pinopsida. All e ...
s made up of spruces (''
Picea A spruce is a tree of the genus ''Picea'' ( ), a genus of about 40 species of coniferous evergreen trees in the family Pinaceae, found in the northern temperate and boreal (taiga) regions of the Northern hemisphere. ''Picea'' is the sole genus ...
'') and pines (''
Pinus A pine is any conifer tree or shrub in the genus ''Pinus'' () of the family Pinaceae. ''Pinus'' is the sole genus in the subfamily Pinoideae. ''World Flora Online'' accepts 134 species-rank taxa (119 species and 15 nothospecies) of pines as c ...
'') within most of eastern North America. In Florida, it consumed twigs of the genus ''
Taxodium ''Taxodium'' is a genus of one to three species (depending on taxonomic opinion) of extremely flood-tolerant conifers in the cypress family, Cupressaceae. The name is derived from the Latin word ''taxus'', meaning " yew", and the Greek word ' ...
'' as well as other woody plants and fruits. Based on carbon isotopic analyses of mastodons in Florida, they had low δ13C values which indicate C3 browsing specialization. The dietary preferences of North American ''Mammut'' are thought to have mirrored those of the older ''Zygolophodon'', which may have preferred living in closed forests and consuming conifers to avoid active competition with the bunodont gomphotheres and lophodont deinotheres in the Miocene of Europe. Most accounts of gut contents have identified coniferous twigs as the dominant element in their diet. In addition to twigs and leaves, as indicated by the "Heisler mastodon" of
Michigan Michigan ( ) is a peninsular U.S. state, state in the Great Lakes region, Great Lakes region of the Upper Midwest, Upper Midwestern United States. It shares water and land boundaries with Minnesota to the northwest, Wisconsin to the west, ...
and the "
Burning Tree mastodon The Burning Tree Mastodon site in Heath, Ohio, Heath, southern Licking County, Ohio, represents the location where the most complete skeleton of American mastodon was found. It is dated to about 11,500 BP. It is believed that there was human prese ...
" of Ohio, mastodons may have also consumed swamp grasses (''
Glyceria ''Glyceria'' is a widespread genus of Poaceae, grass family common across Eurasia, Australia, North Africa, and the Americas. ''Glyceria'' is known commonly as mannagrass in the United States, or, in the UK, sweet-grass. These are perennial Rhi ...
'' and ''
Zizania Wild rice, also called manoomin, mnomen, psíŋ, Canada rice, Indian rice, or water oats, is any of four species of Poaceae, grasses that form the genus ''Zizania'', and the grain that can be harvested from them. The grain was historically an ...
'') as well as semiaquatic and aquatic plants such as sedge marshes (''
Carex ''Carex'' is a vast genus of over 2,000 species of grass-like plants in the family (biology), family Cyperaceae, commonly known as sedges (or seg, in older books). Other members of the family Cyperaceae are also called sedges, however those of ge ...
'') that surrounded lakes. They may have additionally ingested other aquatic plants and aquatic invertebrates while consuming more than of water from lakes a day. The temporal shifts in molar and limb bone sizes in mastodon populations from Missouri and Florida as well as apparent differences in body size between western and eastern populations suggest that ''M. americanum'' was an adaptable species for local environmental shifts. Regardless, it depended heavily on forested environments similar to tapirs, so significant closed vegetation losses of any sort could have impacted them. As a result of the consistent browsing specializations of the genus, ''Mammut'' occupied an ecological niche that allowed it to actively niche partition (or occupy similar but niche ecological spaces) with other proboscideans of North America in the Neogene-
Quaternary The Quaternary ( ) is the current and most recent of the three periods of the Cenozoic Era in the geologic time scale of the International Commission on Stratigraphy (ICS), as well as the current and most recent of the twelve periods of the ...
. In the Blancan, ''M. raki'' showed few morphological changes. In stark contrast, the contemporary gomphothere ''Stegomastodon'' showed progressive developments in response to increasingly arid and extensive grasslands from the Blancan up to the early Irvingtonian, with molar complexities resembling those of ''Mammuthus''. The morphology of ''Stegomastodon'' suggests thus that it was grazing-specialized. A more well-known example of niche partitioning occurred between mastodons and mammoths within the later Pleistocene (Irvingtonian-Rancholabrean). Mammoths had a broader range of diets that allow them to occupy mixed feeding to specialized grazing habits whereas mastodons were specialized browsers that nonetheless still could have consumed a variety of plants. Mammoth diets varied by region whereas those of mastodons remain unclear still. Both at times overlapped in C3 resource usages, although whether this represents browsing or grazing in the case of mammoths remains unclear.


Social behaviors

American mastodons may have lived in herds, and it is possible that they were smaller than mammoth herds on average. Based on the characteristics of mastodon bone sites and strontium and oxygen isotopes from tusks, it can be inferred that, as in modern proboscideans, the mastodon social group consisted of adult females and young, living in bonded groups called mixed herds. The males abandoned the mixed herds once reaching sexual maturity and lived either alone or in male bond groupings. As in modern elephants, there probably was no seasonal synchrony of mating activity, with both males and females seeking out each other for mating when sexually active. Mastodons and other Pleistocene proboscideans may have used landscapes seasonally then migrated to suitable areas to mate or give birth. It is estimated that it may have taken 9 to 12 years for American mastodon females to become mature enough for reproduction, and they may have slowly reproduced single calves at a time. The social behaviors of male mastodon were inferred from one individual skeleton known as the "Buesching mastodon" (known informally as "Fred"), which was recovered from a peat farm near
Fort Wayne Fort Wayne is a city in Allen County, Indiana, United States, and its county seat. Located in northeastern Indiana, the city is west of the Ohio border and south of the Michigan border. The city's population was 263,886 at the 2020 United S ...
, Indiana in 1998. The mastodon individual lived during the later part of the Bølling–Allerød warming period when human populations were present. The Buesching mastodon's tusks grew for about 30 years, and he lived for 34 years total, an approximate lifespan comparable to other males. He may have had engaged in aggressive behavior from
musth Musth or must (from Persian, ) is a periodic condition in bull (male) elephants characterized by aggressive behavior in animals, aggressive behavior and accompanied by a large rise in reproductive hormones. It has been known in Asian elephan ...
, although it may have been season-specific compared to living elephants given climatic conditions in North America. He likely engaged in
intraspecific competition Intraspecific competition is an interaction in population ecology, whereby members of the same species compete for limited resources. This leads to a reduction in fitness for both individuals, but the more fit individual survives and is able to ...
late in his life with other males during the spring or early summer, and he had tusk fractures and may have been severely wounded from a to puncture to the right-sided temporal fossa. Multiple other males are recorded to have had severe wounds resulting from male-male musth fighting. The Buesching mastodon likely considered central Indiana his main home but went on seasonal migrations in his lifetime. He could have traveled hundreds of kilometers in the process and engaged with mates outside of the herd he was born from. Around his last moments, he probably wandered around in vagabondlike behaviors and spent little time in the area where his skeleton was found. His inferred behavior is quite similar to extant elephants.


Paleoecology


Distribution

The range of most species of ''Mammut'' is unknown as their occurrences are restricted to few localities, the exception being the American mastodon (''M. americanum''), which is one of the most widely distributed Pleistocene proboscideans in North America. ''M. americanum'' fossil sites range in time from the
Blancan The Blancan North American Stage on the geologic timescale is the North American faunal stage according to the North American Land Mammal Ages chronology (NALMA), typically set from 4,750,000 to 1,806,000 years BP, a period of .Rancholabrean The Rancholabrean North American Land Mammal Age on the geologic timescale is a North American faunal stage in the North American Land Mammal Ages chronology (NALMA),Sanders, A.E., R.E. Weems, and L.B. Albright III (2009) Formalization of the mid- ...
faunal stages and in locations from as far north as Alaska, as far east as Florida, and as far south as the state of Puebla in central Mexico. ''M. americanum'' was most common in the eastern United States but rarer in the western US in comparison. ''M. pacificum'' is known across California and present as far north as southern
Idaho Idaho ( ) is a landlocked U.S. state, state in the Pacific Northwest and Mountain states, Mountain West subregions of the Western United States. It borders Montana and Wyoming to the east, Nevada and Utah to the south, and Washington (state), ...
, but it was apparently absent from both the
Sonoran Desert The Sonoran Desert () is a hot desert and ecoregion in North America that covers the northwestern Mexican states of Sonora, Baja California, and Baja California Sur, as well as part of the Southwestern United States (in Arizona and California). It ...
and
Mojave Desert The Mojave Desert (; ; ) is a desert in the rain shadow of the southern Sierra Nevada mountains and Transverse Ranges in the Southwestern United States. Named for the Indigenous peoples of the Americas, indigenous Mohave people, it is located pr ...
regions. The elevated-controlled distributions of coniferous forests within the
Rocky Mountain The Rocky Mountains, also known as the Rockies, are a major mountain range and the largest mountain system in North America. The Rocky Mountains stretch in straight-line distance from the northernmost part of Western Canada, to New Mexico in ...
region may have limited populations of ''Mammut'' compared to the other Plio-Pleistocene proboscideans. The easternmost range of the species was in what is now
Montana Montana ( ) is a landlocked U.S. state, state in the Mountain states, Mountain West subregion of the Western United States. It is bordered by Idaho to the west, North Dakota to the east, South Dakota to the southeast, Wyoming to the south, an ...
in the Irvingtonian but may have been extirpated from the area as a result of Illinoian glaciation. An isolated record of ''M. americanum'' is known from
Honduras Honduras, officially the Republic of Honduras, is a country in Central America. It is bordered to the west by Guatemala, to the southwest by El Salvador, to the southeast by Nicaragua, to the south by the Pacific Ocean at the Gulf of Fonseca, ...
, where the genus is not recorded to have extended beyond. ''M. matthewi'' is known by a wide distribution range, its westernmost range being in California from the Horned Toad Formation in the late Hemphillian. It has also apparently been identified from the latest Hemphillian based on skull material from the Pascagoula Formation in Tunica Hills,
Louisiana Louisiana ( ; ; ) is a state in the Deep South and South Central regions of the United States. It borders Texas to the west, Arkansas to the north, and Mississippi to the east. Of the 50 U.S. states, it ranks 31st in area and 25 ...
. This suggests that ''Mammut'' already had an eastern range in the United States by the latest Miocene or earliest Pliocene. Similarly, the same species is recorded from the Palmetto Fauna locality ( Bone Valley Formation) in
Brewster Brewster may refer to: People *Brewster (surname) *Brewster Kahle (born 1960), American computer technologist * Brewster H. Shaw (born 1945), American astronaut * Grace Brewster Hopper (born 1906), American computer scientist, mathematician, and ...
, Florida in the latest Hemphillian while ''Mammut'' sp. is recorded from the Gray Fossil Site in Tennessee. The American mastodon was only present in the far north of North America during
interglacial An interglacial period (or alternatively interglacial, interglaciation) is a geological interval of warmer global average temperature lasting thousands of years that separates consecutive glacial periods within an ice age. The current Holocene i ...
periods, with mitochondrial genome analysis suggesting that separate populations repeatedly colonised the region before becoming
extirpated Local extinction, also extirpation, is the termination of a species (or other taxon) in a chosen geographic area of study, though it still exists elsewhere. Local extinctions are contrasted with global extinctions. Local extinctions mark a chan ...
during glacial periods. A 2022 study of ancient
environmental DNA Environmental DNA or eDNA is DNA that is collected from a variety of environmental samples such as soil, seawater, snow or air, rather than directly sampled from an individual organism. As various organisms interact with the environment, DNA ...
from the Kap Kobenhavn Formation of northern
Greenland Greenland is an autonomous territory in the Danish Realm, Kingdom of Denmark. It is by far the largest geographically of three constituent parts of the kingdom; the other two are metropolitan Denmark and the Faroe Islands. Citizens of Greenlan ...
, dating the
Early Pleistocene The Early Pleistocene is an unofficial epoch (geology), sub-epoch in the international geologic timescale in chronostratigraphy, representing the earliest division of the Pleistocene Epoch within the ongoing Quaternary Period. It is currently esti ...
, 2 million years ago, identified preserved DNA fragments of mastodons. This suggests that the mammutids ranged as far north as Greenland during optimal conditions. Around this time, northern Greenland was 11–19 °C warmer than the
Holocene The Holocene () is the current geologic time scale, geological epoch, beginning approximately 11,700 years ago. It follows the Last Glacial Period, which concluded with the Holocene glacial retreat. The Holocene and the preceding Pleistocene to ...
, with a
boreal forest Taiga or tayga ( ; , ), also known as boreal forest or snow forest, is a biome characterized by pinophyta, coniferous forests consisting mostly of pines, spruces, and larches. The taiga, or boreal forest, is the world's largest land biome. I ...
hosting a species assemblage with no modern analogue. These are among the oldest DNA fragments ever sequenced.


Late Neogene-Quaternary North America

The overall paleontological record of the Neogene of North America is relatively incomplete compared to other areas of the world. This is the result of a greater fossil record bias of western North America compared to eastern North America, meaning that the western half is better understood in terms of evolutionary and climatic trends while the eastern half is poorly understood. During the late Neogene (8-5 Ma), C4 grasslands spread throughout the North American continent and replaced woodland habitats. In eastern North America were relict woodlands in an increasingly drier climate followed by a large faunal turnover. There was a long-term decline of genus-level faunal diversity, with many large-sized herbivores going extinct. Many of the surviving herbivorous faunas were thus adapted for drier and more open habitats resulting from cooling and increase in seasonality. The earliest undisputed record of ''Mammut'' ''sensu stricto'' was of ''M. nevadanum'' in the Thousand Creek Formation in Nevada. Coexistent with the mammutid species were a large variety of other mammals, namely those of the
Artiodactyla Artiodactyls are placental mammals belonging to the order (biology), order Artiodactyla ( , ). Typically, they are ungulates which bear weight equally on two (an even number) of their five toes (the third and fourth, often in the form of a hoof ...
( antilocaprids,
camelid Camelids are members of the biological family (biology), family Camelidae, the only currently living family in the suborder Tylopoda. The seven extant taxon, extant members of this group are: dromedary, dromedary camels, Bactrian camels, wild Bac ...
s,
tayassuid Peccaries (also javelinas or skunk pigs) are pig-like ungulates of the family Tayassuidae (New World pigs). They are found throughout Central and South America, Trinidad in the Caribbean, and in the southwestern area of North America. Peccarie ...
s),
Carnivora Carnivora ( ) is an order of placental mammals specialized primarily in eating flesh, whose members are formally referred to as carnivorans. The order Carnivora is the sixth largest order of mammals, comprising at least 279 species. Carnivor ...
(
canid Canidae (; from Latin, ''canis'', "dog") is a family (biology), biological family of caniform carnivorans, constituting a clade. A member of this family is also called a canid (). The family includes three subfamily, subfamilies: the Caninae, a ...
s,
felid Felidae ( ) is the Family (biology), family of mammals in the Order (biology), order Carnivora colloquially referred to as cats. A member of this family is also called a felid ( ). The 41 extant taxon, extant Felidae species exhibit the gre ...
s,
mustelid The Mustelidae (; from Latin , weasel) are a diverse family of carnivoran mammals, including weasels, badgers, otters, polecats, martens, grisons, and wolverines. Otherwise known as mustelids (), they form the largest family in the suborde ...
s, ursids),
Eulipotyphla Eulipotyphla (, from '' eu-'' + '' Lipotyphla'', meaning truly lacking blind gut; sometimes called true insectivores) is an order of mammals comprising the Erinaceidae ( hedgehogs and gymnures); Solenodontidae (solenodons); Talpidae ( mole ...
(
talpid The family Talpidae () includes the true moles (as well as the shrew moles and desmans) who are small insectivorous mammals of the order Eulipotyphla. Talpids are all digging animals to various degrees: moles are completely subterranean animal ...
s),
Lagomorpha The lagomorphs () are the members of the taxonomic order Lagomorpha, of which there are two living families: the Leporidae (rabbits and hares) and the Ochotonidae ( pikas). There are 110 recent species of lagomorph, of which 109 species in t ...
(
leporid Leporidae () is the family of rabbits and hares, containing over 70 species of extant mammals in all. The family name comes from "Lepus", hare in Latin. Together with the pikas, the Leporidae constitute the mammalian order Lagomorpha. Leporidae ...
s),
Perissodactyla Perissodactyla (, ), or odd-toed ungulates, is an order of ungulates. The order includes about 17 living species divided into three families: Equidae (horses, asses, and zebras), Rhinocerotidae (rhinoceroses), and Tapiridae (tapirs). They t ...
(
equid Equidae (commonly known as the horse family) is the Taxonomy (biology), taxonomic Family (biology), family of Wild horse, horses and related animals, including Asinus, asses, zebra, zebras, and many extinct species known only from fossils. The fa ...
s,
rhinocerotid A rhinoceros ( ; ; ; : rhinoceros or rhinoceroses), commonly abbreviated to rhino, is a member of any of the five extant taxon, extant species (or numerous extinct species) of odd-toed ungulates (perissodactyls) in the family (biology), famil ...
s), and
Rodentia Rodents (from Latin , 'to gnaw') are mammals of the order Rodentia ( ), which are characterized by a single pair of continuously growing incisors in each of the upper and lower jaws. About 40% of all mammal species are rodents. They are na ...
( aplodontiids, castorids, geomyids, heteromyids, cricetids, mylagaulids, and sciurids). The latest Hemphillian of Florida based on the Palmetto Fauna of the Bone Valley Formation records the coexistence of ''M. matthewi'' with similar types of faunas, namely
Pilosa The Order (biology), order Pilosa is a clade of xenarthran placental mammals, native to the Americas. It includes anteaters and sloths (which include the extinct ground sloths). The name comes from the Latin word for "hairy". Origins and taxon ...
( megalonychids), Eulipotyphla (talpids), Lagomorpha (leporids), Carnivora ( borophagine canids,
canine Canine may refer to: Zoology and anatomy * Animals of the family Canidae, more specifically the subfamily Caninae, which includes dogs, wolves, foxes, jackals and coyotes ** ''Canis'', a genus that includes dogs, wolves, coyotes, and jackals ** Do ...
canids, ursids,
procyonid Procyonidae ( ) is a New World family (biology), family of the order (biology), order Carnivora. It includes the Procyon (genus), raccoons, ringtails, cacomistles, coatis, kinkajous, olingos, and olinguitos. Procyonids inhabit a wide range of e ...
s, mustelids including lutrines, feline felids,
machairodontine Machairodontinae (from Ancient Greek μάχαιρα ''machaira,'' a type of Ancient Greek sword and ὀδόντος ''odontos'' meaning tooth) is an extinct subfamily of carnivoran mammals of the cat family Felidae, representing the earliest d ...
felids), Proboscidea (gomphotheres), Perissodactyla (tapirs, rhinocerotids, hipparionine equids), and Artiodactyla (tayassuids, protoceratids, camelids, " pseudoceratines,"
cervid A deer (: deer) or true deer is a hoofed ruminant ungulate of the family Cervidae (informally the deer family). Cervidae is divided into subfamilies Cervinae (which includes, among others, muntjac, elk (wapiti), red deer, and fallow deer) a ...
s, antilocaprids). North America in the late Neogene is understood to have undergone a long-term decline in large mammal diversity (i.e. the
Dromomerycidae Dromomerycidae is an extinct family of Neogene ruminants belonging to the infraorder Pecora. Dromomerycids, known by two subfamilies, were endemic to North America from the later early Miocene up to the early Pliocene (~5 Ma), leaving no descenda ...
, " Blastomerycinae," Rhinocerotidae) as a result of C4 grassland expansion, cooler climates, and increased seasonality. The Blancan fossil record suggests a maximum known diversity of four species of ''Mammut'' (''M. americanum'', ''M. vexillarius'', ''M. raki'', and ''M. cosoensis''). However, the Blancan record of ''Mammut'' is relatively rare. ''M. raki'' from the Palomas Formation of Truth or Consequences in New Mexico is recorded with a few other mammalian faunas, namely the megalonychid
ground sloth Ground sloths are a diverse group of extinct sloths in the mammalian superorder Xenarthra. They varied widely in size with the largest, belonging to genera '' Lestodon'', ''Eremotherium'' and ''Megatherium'', being around the size of elephants. ...
''Megalonyx'', the pocket gopher ''
Geomys The genus ''Geomys'' contains 12 extant species of pocket gophersSearch results for "''Geomys''" on thASM Mammal Diversity Database often collectively referred to as the eastern pocket gophers. Like all pocket gophers, members of this genus are ...
'', the cricetid '' Sigmodon'', the equin '' Equus'', the hipparionine ''
Nannippus ''Nannippus'' is an extinct genus of three-toed horse endemic to North America during the Miocene through Pleistocene, about 13.3—1.8 million years ago (Mya), living around 11.5 million years. This ancient species of three-toed horse grew up to ...
'', and the camelid ''
Camelops ''Camelops'' is an extinct genus of camel that lived in North and Central America from the middle Pliocene (from around 4-3.2 million years ago) to the end of the Pleistocene (around 13-12,000 years ago). It is more closely related to living cam ...
''. A late Blancan locality known as the Fish Springs Flat Fauna in Nevada reveals that fossils of ''M. americanum'' were found with those of the leporid ''
Hypolagus ''Hypolagus'' is an extinct genus of lagomorph, first recorded in the Hemingfordian (early to middle Miocene) of North America. It entered Asia during the early Turolian and spread to Europe not much later, where it survived until the Middle Pl ...
'', lutrine ''
Satherium ''Satherium'' is an extinct genus of otters that lived in North America during the Pliocene and Pleistocene. Two species are known, '' Satherium piscinarium'' and ''Satherium ingens''. ''S. piscinarium'' was originally classified as a species of ...
'', equid ''Equus'', camelid '' Gigantocamelus'', gopher '' Thomomys'', and the ground squirrel ''
Spermophilus ''Spermophilus'' is a genus of ground squirrels in the Sciuridae, squirrel family. As traditionally defined the genus was very species-rich, ranging through Europe, Asia and North America, but this arrangement was found to be paraphyletic to ...
''. In the Irvingtonian, only ''M. americanum'' is recorded to have crossed past the Blancan while ''M. pacificum'' replaced the other Blancan species. By this time, ''Mammut'' would have coexisted with the elephantid ''Mammuthus'' and the gomphotheres ''Cuvieronius'' and ''Stegomastodon'', although the latter failed to survive past the early Irvingtonian. The Middle Pleistocene sites are scarce in North America compared to the Late Pleistocene sites, but from the Irvingtonian to the Rancholabrean, repeated glacial events occurred that led to repeated formations of major ice sheets in northern North America. The Port Kennedy Bone Cave of Pennsylvania is of Irvingtonian age (Middle Pleistocene) and reveals that during this time, ''M. americanum'' was present with the megalonychid ''Megalonyx wheatleyi'', the tremarctine bear '' Arctodus pristinus'', the
jaguar The jaguar (''Panthera onca'') is a large felidae, cat species and the only extant taxon, living member of the genus ''Panthera'' that is native to the Americas. With a body length of up to and a weight of up to , it is the biggest cat spe ...
(''Panthera onca''), the felid '' Miracinonyx inexpectatus'', and the machairodontine '' Smilodon gracilis''. The Big Bone Lick locality in Kentucky, which dates to the latest Pleistocene (Rancholabrean), indicates the coexistence of the American mastodon with the extant
reindeer The reindeer or caribou (''Rangifer tarandus'') is a species of deer with circumpolar distribution, native to Arctic, subarctic, tundra, taiga, boreal, and mountainous regions of Northern Europe, Siberia, and North America. It is the only re ...
(''Rangifer tarandus'') along with various other extinct
megafauna In zoology, megafauna (from Ancient Greek, Greek μέγας ''megas'' "large" and Neo-Latin ''fauna'' "animal life") are large animals. The precise definition of the term varies widely, though a common threshold is approximately , this lower en ...
like ancient bison (''Bison antiquus''), the Caprinae, caprine bovid ''Bootherium, Bootherium bombifrons'', mylodontid ground sloth ''Paramylodon, Paramylodon harlani'', megalonychid ''Megalonyx jeffersoni'', true deer ''Cervalces, Cervalces scotti'', equid ''Equus complicatus'', and the Columbian mammoth.


Relationship with humans

The exact timing of human (''Homo sapiens'') arrival to temperate North America is unclear, but they likely arrived in North America ∼19,000–14,000 radiocarbon calibration, calibrated years Before Present. They are known within the archeological record as
Paleoindian Paleo-Indians were the first peoples who entered and subsequently inhabited the Americas towards the end of the Late Pleistocene period. The prefix ''paleo-'' comes from . The term ''Paleo-Indians'' applies specifically to the lithic period in ...
s and eventually gave rise to modern-day Native Americans. Of interest is that in the
Clovis culture The Clovis culture is an archaeological culture from the Paleoindian period of North America, spanning around 13,050 to 12,750 years Before Present (BP). The type site is Blackwater Draw locality No. 1 near Clovis, New Mexico, where stone too ...
phase, there is evidence that Clovis hunters targeted contemporary proboscideans based on archeological "kill sites." Clovis projectile points and other artifacts have been found in association with both mammoths and mastodons. The former has more frequent evidence of having been hunted by Clovis hunters while mastodons have much fewer in comparison. Todd A. Surovell and Nicole M. Waguespack in 2008 hypothesized that Clovis hunters in North America hunted proboscideans more often than those in any other continent. They addressed that preservation biases of larger mammals in archeological sites may have caused higher representations of proboscidean kill sites but suggested that regardless, Clovis hunters were likely specialized in hunting large game. As of present, 2 definite ''Mammut'' kill sites compatible with Clovis lithic technology have been recorded compared to 15 of ''Mammuthus'' and 1 of ''Cuvieronius''. These two kill sites are thought to be from Kimmswick, Missouri and Pleasant Lake in Washtenaw County, Michigan. Whether various other sites can be confirmed as proboscidean butchery sites appear subjective, largely depending on the views of different authors. It is uncertain if Clovis people had hunting strategies of proboscideans similar to tribal Africans, but the Clovis points likely indicate usage as spears for thrusting or throwing at proboscideans (there are disagreements to whether they indicate multiple other usages, however). According to the American paleontologist Daniel Fisher (paleontologist), Daniel C. Fisher, the "Heisler mastodon" site in Calhoun County, Michigan, Calhoun County, Michigan, which recovered about 50% of the skeleton, was proof of meat caching in a pond by Paleoindians in the late Pleistocene. This hypothesis opposes the notion that proboscideans ended up unable to disentangle themselves in marsh wetlands, which he said there is no evidence of. His hypothesis was based on his experiment with partial carcasses of a horse that was preserved in a shallow lake then extracted as well as a Moravian Church, Moravian missionary's testimony of Inuit retrieving caribou carcasses from lakes that they probably placed as storage in the cases of excess meat or future limited hunting successes. Fisher said that if his theory is true, then Paleoindian interactions with megafauna (hunting and scavenging) are far more complex than initially thought. In 2023, Michael R. Waters et al. suggested that the Manis Mastodon site in
Washington Washington most commonly refers to: * George Washington (1732–1799), the first president of the United States * Washington (state), a state in the Pacific Northwest of the United States * Washington, D.C., the capital of the United States ** A ...
state supported evidence of a mastodon hunt ~13,900 cal. years BP, some 900 years before Clovis culture. Their study was a continuation of a 2011 anatomical study that proposed that osseous (bone) pieces found in a right rib of a mastodon represented fragmented tips of a projectile point, but it had been repeatedly challenged by other authors. Based on anatomical reevaluations, they determined that the bone fragments were embedded in the Manis mastodon rib while it was alive, as evident by the visible healing around the wounded area. Waters and his colleagues stated that the bone pieces were from an external source, explainable by human-made projectile points. They rejected alternate explanations for why bone fragments ended up in the Manis mastodon rib. Based on this, they envisioned that the mastodon individual was wounded by pre-Clovis hunters and got away, giving it time to heal. Afterwards, it died either by natural causes and was scavenged by humans, or it was killed by them on another attack then butchered. This site proves the existence of pre-Clovis hunting technology that the earliest people brought with them when dispersing to North America and made localized adaptations of. In 2017, Steven R. Holen et al. published an article arguing that the Cerutti Mastodon site, located in San Diego County in California, is an archeological site involving ''M. americanum'' that dates to approximately 130,000 years ago. If true, they stated, the site would imply evidence of now-extinct species of ''Homo'' in North America during the Marine Isotope Stage 5 (MIS 5e) temporal range of the early late Pleistocene. The proposal was highly controversial, as many archeologists were skeptical about the claim that the bones of ''M. americanum'' were broken by hominins, and alternate explanations have been offered. For instance, in the same year the article was published, Gary Haynes expressed concern of it being published in the journal ''Nature (journal), Nature'' due to how highly prolific it is. Reporters from print presses and digital media published reactions of the article from various North American archeologists, with Donald K. Grayson stating that it was astonishingly bad, Jon M. Erlandson arguing that the site was non-credible, and various other archeologists arguing that the claim is insufficiently supported. Haynes pointed out that the article's claim was "extraordinary" and must therefore be met with rigorous skepticism. He wrote that there were no traces of archeological structures typically built by archaic species of ''Homo'' (i.e. ''H. erectus'', Neanderthals, or Denisovans) in the Cerutti site. Additionally, he brought up the possibilities of the fossil bones being affected by sediment pressures or damage done by earth-moving construction equipments despite the original authors denying the latter possibility. Multiple petroglyphs suggested to have depicted prehistoric proboscideans in North America like mastodons are known within the United States, but they are either fraudulent or depict entities other than mastodons. As a result, suggested rock art of mammoths and mastodons within North America are not sufficiently credible.


Extinction

''Mammut'', or more specifically the American mastodon, experienced an initial decline in geographical range when it was extirpated from the northernmost ranges of North America ~75,000 years ago. ''Mammut'' initially occupied the region during the Last Interglacial (~125,000-75,000 years ago) back when suitable forested habitats were present there but was subsequently extirpated in correlation with environmental changes from the Wisconsin glaciation (MIS 4). The local extirpation, occurring long before human arrival, caused the mastodon range to be limited to areas south of North American ice sheets. The steppe-tundra faunas thrived there during the event whereas boreal forest-adapted faunas underwent declines. The trend of recolonization and extirpation appears to have had been a recurring trend in the Pleistocene correlated with repeated returns of forests and wetlands, but what is unclear is why faunas that were able to repeatedly recolonize northern North America during previous interglacial periods were unable to do so again after the Last Glacial Maximum. The latest Pleistocene of North America records a Late Pleistocene extinctions, large extinction phase that resulted in the disappearances of over 30 genera of mammals, the majority of which are considered "megafauna" (~ or larger). ''Mammut'' was one of the many genera recorded within North America whose extinction causes are currently unresolved. During the latest Pleistocene of North America, two major events occurred: the development of Clovis culture from 13,200 to 12,800 years ago and the onset of the
Younger Dryas The Younger Dryas (YD, Greenland Stadial GS-1) was a period in Earth's geologic history that occurred circa 12,900 to 11,700 years Before Present (BP). It is primarily known for the sudden or "abrupt" cooling in the Northern Hemisphere, when the ...
cold phase from 12,900 to 11,700 years ago. The extinctions of mammalian megafauna in North America are particularly high akin to those of South America and Australia rather than Eurasia and Africa. As a result, the extinctions that occurred in the latest Pleistocene of North America have been mainly attributed to human hunting, climate change, or some combination of the two (there are alternate but lesser-supported hypotheses). Many researchers have struggled to explain the North American extinctions, with both human hunting and climate change explanations alone being challenged. In recent years, research has shifted towards studying the extinctions of North American faunas by individual taxon and/or region rather as a homogenous group. The results vary in regions such as the northeast, with some authors suggesting that there was minimal evidence for Clovis hunting being the major factor behind proboscidean population drops and some others arguing that environmental shifts prior to human arrival were not detrimental enough to the proboscideans. Paul L. Koch and Anthony D. Barnosky in 2006 suggested that ''Mammuthus'' was well-associated with archeological sites of North America. In comparison, ''Mammut'' and the peccary ''Platygonus'' were far less frequently associated with human sites, potentially suggesting that Paleoindians hunted them less than mammoths. They stated that the current understanding of ''Mammut'' associations with humans could shift if the supposed butchery sites were better understood while that of ''Platygonus'' is stable and therefore unlikely to change. In 2018, Jack M. Broughton and Elic M. Weitzel calculated populated dynamics of some of the North American late Pleistocene megafauna based on summed probability distributions (SPDs) using calibrated radiocarbon dates. They determined based on the data that the declines of ''Mammuthus'', ''Equus'', and ''Smilodon'' were correlated with Clovis culture hunting while ''Mammut'' and the nothrotheriid ground sloth ''Nothrotheriops'' did not exhibit any significant population bust until after Clovis culture and during the Younger Dryas at ~12,650 years ago. They concluded that the declines of megafauna are of mixed causes and that the extinction processes and causes therefore vary by individual taxon and region. Of note is that there is a recorded latest survival of the American mastodon in the early
Holocene The Holocene () is the current geologic time scale, geological epoch, beginning approximately 11,700 years ago. It follows the Last Glacial Period, which concluded with the Holocene glacial retreat. The Holocene and the preceding Pleistocene to ...
. The Overmyer Mastodon individual, recovered from northern Indiana with 41-48% complete remains recovered, exhibits no evidence of weathering or gnawing by other animals. The individual dates from 11,795 to 11,345 years Before Present for a median of 11,576 calibrated years BP, therefore having a secure calibrated radiocarbon date dating to the early Holocene unlike most other extinct North American genera of the terminal Pleistocene. Neal Woodman and Nancy Beavan Athfield stressed that although the early Holocene survival of the species does not eliminate the possibilities that Clovis hunters and/or Younger Dryas impacted their populations in the long term, its survival meant that the genus was not immediately brought to extinction by either factor.


Cultural significance

Late Pleistocene proboscideans of the Americas such as the American mastodon could have been recognized in Native American oral histories, but they are unlikely to have referenced any specific species. Typically, they may have been depicted in Native American oral history as aggressive and antagonistic beasts. Mastodons may have played ancient roles in Indigenous peoples of the Pacific Northwest Coast, Native American cultures of the Pacific Northwest. In 1987, Carl E. Gustafson recovered fossil evidence of a late Pleistocene mastodon far away from where the species would typically roam, the radiocarbon dating confirming a date of about 13,800 years ago. The local tribal members identified the remains as being of game pieces for slahal, a gambling game for dispute settlements and entertainment. The bone sticks, carved from mastodon bones, are not easily interpretable archeologically, but tribal members saw the recovery of the items as evidence of the endurance of ancient cultural practices like slahal. The American mastodon had long been a stand-in within the United States for American nationalism since early American history, and Thomas Jefferson was famously known for having hoped that the Lewis and Clark Expedition would eventually yield evidence of living mastodons in the western frontier of the United States. It was a defining symbol of museums according to Brett Barney as evident by a mention of it by Walt Whitman in a passage of the 1855 poem "Song of Myself." The mastodon became the subject of a Michigan political campaign in 2000 when Washtenaw Community College geology instructor David P. Thomas Sr. aimed to make it the state fossil of Michigan. He, assisted by the Slauson Middle School (Ann Arbor, Michigan), Slauson Middle School science teacher Jeffrey Bradley, was sponsored by the state senator Thaddeus McCotter, arranged petition drives that collected thousands of signatures, and attended state hearings. Bradley's students participated in the "Mastodon for Michigan" campaign, which built a life-sized replica out of paper and raised $1,000 for the University of Michigan Museum of Natural History to build a mastodon exhibit. In 2002, the mastodon became the state fossil, making it the fourteenth Lists of United States state symbols, state symbol. Similarly, the mastodon became the state fossil of Indiana as recently as 2022 due to House Bill 1013, authored by the representative Randy Frye, passing unanimously. In January 2024, Indiana senator Mike Braun and Michigan senator Gary Peters introduced a bipartisan bill to make the mastodon the US national fossil is what is called the "National Fossil Act." Section 1 aims to define the bill's name, Section 2 would investigate the roles of the mastodon in American public life, and Section 3 would designate it as the national fossil under Title 36 of the United States Code. Peters justified that the mastodon represents a unique aspect of Michigan's history and American history, stating that he hoped that its establishment as the national fossil would preserve the histories and encourage new generations of scientists and other researchers to pursue their goals. Located in the Mastodon Ridge park in the Canadian town of Stewiacke, Nova Scotia is a large-sized replica of a mastodon based on a skeleton recovered from Nova Scotia. It was sculpted as a clay model, has a weight of ~, is in shoulder height, and measures long. The sculpture took about 8 weeks to be constructed and was sent to the Mastodon Ridge in January 1995. The name "mastodon" was adopted in different contexts within the United States. For instance, 4-8-0 locomotives of the late 19th century were originally named "''Mastodon (steam locomotive), Mastodons''" before the name was eventually replaced with "12-wheeler." The name was a reference to the American mastodon. The 4-10-0 locomotive later became known also as "''Mastodon''." In the 1993-1995 show ''Mighty Morphin Power Rangers'', the Black Ranger Zack Taylor had the mastodon ability and controlled the Mastodon Dinozord machine. The name "Mastodon" was also adopted by a heavy metal music, heavy metal band when guitarist Bill Kelliher was asked by the guitarist-singer Brent Hinds asked him about the name of the "fossil elephant" after seeing his tattoo of a Bantha skull from the Star Wars franchise, in which the members then agreed to it being the Mastodon (band), band's name. "Mastodon" is also the name of a blogging social network site that also acquired its name from the extinct proboscidean species. The mastodon is the mascot of the Massachusetts College of Art and Design.


See also

* Big Bone Lick State Park * Cerutti Mastodon site * Coats–Hines site * List of museums and colleges with mastodon fossils on display * Manis Mastodon site * Snowmastodon site


References


External links


The Rochester Museum of Science – Expedition Earth Glaciers & Giants





American Museum of Natural History – Warren Mastodon

BBC Science and Nature:Animals – American mastodon Mammut americanum

BBC News – Greek mastodon find 'spectacular'


* [http://www.slfp.com/Mastodon.htm Saint Louis Front Page – Mastodon State Historic Site]
Story of the Randolph Mastodon (Earlham College)


* [http://westerncentermuseum.org Western Center for Archaeology & Paleontology, home of the largest mastodon ever found in the Western United States]
Smithsonian Magazine Features Mammoths and Mastodons

360 View of a Mastodon Skull from Indiana State Museum
* 3-D Viewers o

an

mastodon skeletons at the University of Michiga
Mammutidae digital fossil repository
* ''Scientific American'',
The Chicago Mastodon
, 18 September 1880, p. 175 {{Authority control Mastodons, Extinct animals of Canada Fossil taxa described in 1799 Mammutidae Messinian first appearances Miocene mammals of North America Miocene proboscideans Paleontology in Michigan Pleistocene extinctions Pleistocene mammals of North America Pleistocene proboscideans Pliocene mammals of North America Pliocene proboscideans Ringold Formation Miocene Fauna Symbols of Indiana Symbols of Michigan