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A mastodon ( 'breast' + 'tooth') is any
proboscidea The Proboscidea (; , ) are a taxonomic order of afrotherian mammals containing one living family ( Elephantidae) and several extinct families. First described by J. Illiger in 1811, it encompasses the elephants and their close relatives. Fr ...
n belonging to the extinct genus ''Mammut'' (family Mammutidae). Mastodons inhabited North and Central America during the late
Miocene The Miocene ( ) is the first 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 means "less recen ...
or late
Pliocene The Pliocene ( ; also Pleiocene) is the epoch in the geologic time scale that extends from 5.333 million to 2.58Pleistocene The Pleistocene ( , often referred to as the ''Ice age'') is the geological Epoch (geology), epoch that lasted from about 2,580,000 to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was fina ...
10,000 to 11,000 years ago. They lived in herds and were predominantly forest-dwelling animals. They generally had a
browsing Browsing is a kind of orienting strategy. It is supposed to identify something of relevance for the browsing organism. When used about human beings it is a metaphor taken from the animal kingdom. It is used, for example, about people browsing o ...
diet, distinct from that of the contemporary
Columbian mammoth The Columbian mammoth (''Mammuthus columbi'') is an extinct species of mammoth that inhabited the Americas as far north as the Northern United States and as far south as Costa Rica during the Pleistocene epoch. It was one of the last in a line ...
, which tended towards
grazing In agriculture, grazing is a method of animal husbandry whereby domestic livestock are allowed outdoors to roam around and consume wild vegetations in order to convert the otherwise indigestible (by human gut) cellulose within grass and ot ...
. ''M. americanum'', the American mastodon, and ''M. pacificus'', the Pacific mastodon, are the youngest and best-known species of the genus. Mastodons disappeared from North America as part of a
mass extinction An extinction event (also known as a mass extinction or biotic crisis) is a widespread and rapid decrease in the biodiversity on Earth. Such an event is identified by a sharp change in the diversity and abundance of multicellular organisms. I ...
of most of the
Pleistocene megafauna Pleistocene megafauna is the set of large animals that lived on Earth during the Pleistocene epoch. Pleistocene megafauna became extinct during the Quaternary extinction event resulting in substantial changes to ecosystems globally. The role of ...
, widely believed to have been caused by a combination of climate changes at the end of the Pleistocene and
overexploitation Overexploitation, also called overharvesting, refers to harvesting a renewable resource to the point of diminishing returns. Continued overexploitation can lead to the destruction of the resource, as it will be unable to replenish. The term ap ...
by Paleo-Indians.


History

A Dutch tenant farmer found the first recorded remnant of ''Mammut'', a tooth some in weight, in the village of
Claverack, New York Claverack is a town in Columbia County, New York, United States. The population was 6,021 at the 2010 census. The town name is a corruption for the Dutch word “Klaverakker” for "Clover Fields" or "Clover Reach". In 1705, the first discovery ...
, in 1705. The mystery animal became known as the "incognitum". In 1739 French soldiers at present-day
Big Bone Lick State Park Big Bone Lick State Park is located at 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 lick deposited around t ...
, Kentucky, found the first bones to be collected and studied scientifically. They carried them to the
Mississippi River The Mississippi River is the List of longest rivers of the United States (by main stem), second-longest river and chief river of the second-largest Drainage system (geomorphology), drainage system in North America, second only to the Hudson B ...
, from where they were transported to the
National Museum of Natural History The National Museum of Natural History is a natural history museum administered by the Smithsonian Institution, located on the National Mall in Washington, D.C., United States. It has free admission and is open 364 days a year. In 2021, with 7. ...
in
Paris Paris () is the capital and most populous city of France, with an estimated population of 2,165,423 residents in 2019 in an area of more than 105 km² (41 sq mi), making it the 30th most densely populated city in the world in 2020. Si ...
. Similar teeth were found in South Carolina, and some of the
African slaves Slavery has historically been widespread in Africa. Systems of servitude and slavery were common in parts of Africa in ancient times, as they were in much of the rest of the ancient world. When the trans-Saharan slave trade, Indian Ocean ...
there supposedly recognized them as being similar to the teeth of African elephants. There soon followed discoveries of complete bones and tusks in Ohio. People started referring to the "incognitum" as a "mammoth", like the ones that were being dug out in Siberia – in 1796 the French anatomist
Georges Cuvier Jean Léopold Nicolas Frédéric, Baron Cuvier (; 23 August 1769 – 13 May 1832), known as Georges Cuvier, was a French naturalist and zoologist, sometimes referred to as the "founding father of paleontology". Cuvier was a major figure in na ...
proposed the radical idea that mammoths were not simply elephant bones that had been somehow transported north, but a species which no longer existed.
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 ...
assigned the scientific name ''Mammut'' to the American "incognitum" remains in 1799, under the assumption that they belonged to mammoths. Other anatomists noted that the teeth of mammoths and elephants differed from those of the "incognitum", which possessed rows of large conical cusps, indicating that they were dealing with a distinct species. In 1817 Cuvier named the "incognitum" ''Mastodon''. Cuvier assigned the name ''mastodon'' (or ''mastodont'') – meaning "breast tooth" (
Ancient Greek Ancient Greek includes the forms of the Greek language used in ancient Greece and the ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Dark Ages (), the Archaic pe ...
: μαστός "breast" and ὀδούς, "tooth"), – for the nipple-like projections on the crowns of the molars.


Taxonomy

''Mastodon'' as a genus name is obsolete; the valid name is ''Mammut'', as that name preceded Cuvier's description, making ''Mastodon'' a
junior synonym The Botanical and Zoological Codes of nomenclature treat the concept of synonymy differently. * In botanical nomenclature, a synonym is a scientific name that applies to a taxon that (now) goes by a different scientific name. For example, Linn ...
. The change was met with resistance, and authors sometimes applied ''"Mastodon"'' as an informal name; consequently it became the common term for members of the genus. "Mastodon" is also used informally to describe other non-elephant proboscideans, such as
gomphothere Gomphotheres are any members of the diverse, extinct taxonomic family Gomphotheriidae. Gomphotheres were elephant-like proboscideans, but do not belong to the family Elephantidae. They were widespread across Afro-Eurasia and North America dur ...
s. Species include: * ''M. americanum'', the American mastodon, is one of the best known and among the last species of ''Mammut''. Its earliest occurrences date from the early-middle Pliocene (early Blancan stage). It was formerly regarded (see below) as having a continent-wide distribution, especially during the Pleistocene epoch, known from fossil sites ranging from present-day Alaska, Ontario and New England in the north, to Florida, southern California, and as far south as Honduras.
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 ...
also suggests they ranged as far northeast as
Greenland Greenland ( kl, Kalaallit Nunaat, ; da, Grønland, ) is an island country in North America that is part of the Kingdom of Denmark. It is located between the Arctic and Atlantic oceans, east of the Canadian Arctic Archipelago. Greenland ...
during the warmer conditions of the
early Pleistocene The Early Pleistocene is an unofficial sub-epoch in the international geologic timescale in chronostratigraphy, being the earliest division of the Pleistocene Epoch within the ongoing Quaternary Period. It is currently estimated to span the time ...
. The American mastodon has been widely thought to have resembled a
woolly mammoth The woolly mammoth (''Mammuthus primigenius'') is an extinct species of mammoth that lived during the Pleistocene until its extinction in the Holocene epoch. It was one of the last in a line of mammoth species, beginning with '' Mammuthus s ...
in appearance. However, consideration of the long tail (usually present in animals living in warm climates), size, body mass and environment implies the animal was not similarly hairy, and there is scant preserved evidence of body hair (what little has been recovered suggests a
semiaquatic In biology, semiaquatic can refer to various types of animals that spend part of their time in water, or plants that naturally grow partially submerged in water. Examples are given below. Semiaquatic animals Semi aquatic animals include: * Ve ...
lifestyle). It had
tusk Tusks are elongated, continuously growing front teeth that protrude well beyond the mouth of certain mammal species. They are most commonly canine teeth, as with pigs and walruses, or, in the case of elephants, elongated incisors. Tusks share ...
s that sometimes exceeded in length; they curved upwards, but less dramatically than those of the woolly mammoth. Its main habitat was cold
spruce A spruce is a tree of the genus ''Picea'' (), a genus of about 35 species of coniferous evergreen trees in the family Pinaceae, found in the northern temperate and boreal (taiga) regions of the Earth. ''Picea'' is the sole genus in the subfam ...
woodlands, and it is believed to have browsed in herds. It became extinct at the end of the Pleistocene approximately 11,000 years ago. * ''M. matthewi'' — found in the Snake Creek Formation of Nebraska, dating to the late Hemphillian. Some authors consider it practically indistinguishable from ''M. americanum''. There is one report of it in China. * ''M. pacificus'' — based on a 2019 analysis, Pleistocene specimens from California and southern Idaho have been transferred from ''M. americanum'' to this new species. It differs from the eastern population in having narrower molars, six as opposed to five sacral vertebrae, a thicker femur, and a consistent absence of mandibular tusks. * ''M. raki'' — Its remains were found in the Palomas Formation, near
Truth or Consequences, New Mexico Truth or Consequences (often abbreviated as T or C) is a city in New Mexico, and the county seat of Sierra County. In 2020, the population was 6,052. It has frequently been noted on lists of unusual place names for having chosen to rename itse ...
, dating from the early-middle Pliocene, between 4.5 and 3.6 Ma. It coexisted with '' Equus simplicidens'' and '' Gigantocamelus'' and differs from ''M. americanum'' in having a relatively longer and narrower third molar, similar to the description of the defunct genus ''Pliomastodon'', which supports its arrangement as an early species of ''Mammut''. However, like ''M. matthewi'', some authors do not consider it sufficiently distinct from ''M. americanum'' to warrant its own species. * ''M. cosoensis'' — found in the Coso Formation of California, dating to the
Late Pliocene Late may refer to: * LATE, an acronym which could stand for: ** Limbic-predominant age-related TDP-43 encephalopathy, a proposed form of dementia ** Local-authority trading enterprise, a New Zealand business law ** Local average treatment effe ...
, originally a species of '' Pliomastodon'', it was later assigned to ''Mammut''. Since a tentative 1977 report of ''M. matthewi'' in China, there have been no reports of currently recognized ''Mammut'' species outside of North America according to Paleobiology database (which does not recognize '' M. borsoni''). However, the status of ''Mammut'' or '' Zygolophodon borsoni'' in the literature appears equivocal.


Evolution

''Mammut'' is a genus of the extinct family Mammutidae, closely related to the
proboscidean The Proboscidea (; , ) are a taxonomic order of afrotherian mammals containing one living family (Elephantidae) and several extinct families. First described by J. Illiger in 1811, it encompasses the elephants and their close relatives. From ...
family
Elephantidae Elephantidae is a family of large, herbivorous proboscidean mammals collectively called elephants and mammoths. These are terrestrial large mammals with a snout modified into a trunk and teeth modified into tusks. Most genera and species in the ...
(mammoths and elephants), from which it originally diverged approximately twenty-seven million years ago. The following
cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to ...
shows the placement of the American mastodon among other proboscideans, based on
hyoid The hyoid bone (lingual bone or tongue-bone) () is a horseshoe-shaped bone situated in the anterior midline of the neck between the chin and the thyroid cartilage. At rest, it lies between the base of the mandible and the third cervical vertebra. ...
characteristics: Over the years, several fossils from localities in North America, Africa and Asia have been attributed to ''Mammut'', but only the North American remains have been named and described, one of them being ''M. furlongi'', named from remains found in the Juntura Formation of Oregon, dating from the late Miocene. However, it is no longer considered valid, leaving only five valid species. A complete
mitochondrial DNA Mitochondrial DNA (mtDNA or mDNA) is the DNA located in mitochondria, cellular organelles within eukaryotic cells that convert chemical energy from food into a form that cells can use, such as adenosine triphosphate (ATP). Mitochondrial D ...
(mtDNA) sequence has been obtained from the tooth of an ''M. americanum'' skeleton found in
permafrost Permafrost is ground that continuously remains below 0 °C (32 °F) for two or more years, located on land or under the ocean. Most common in the Northern Hemisphere, around 15% of the Northern Hemisphere or 11% of the global surface ...
in northern Alaska. The remains are thought to be 50,000 to 130,000 years old. This sequence has been used as an outgroup to refine divergence dates in the evolution of the Elephantidae. The rate of mtDNA sequence change in proboscideans was found to be significantly lower than in primates. A 2020 analysis of mtDNA from American mastodon remains collected in eastern
Beringia Beringia is defined today as the land and maritime area bounded on the west by the Lena River in Russia; on the east by the Mackenzie River in Canada; on the north by 72 degrees north latitude in the Chukchi Sea; and on the south by the tip ...
indicated they belonged to two genetically divergent
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English ter ...
s. The clades were dated to different
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 ...
s, suggesting a repeating pattern of
colonization Colonization, or colonisation, constitutes large-scale population movements wherein migrants maintain strong links with their, or their ancestors', former country – by such links, gain advantage over other inhabitants of the territory. When ...
during an interglacial followed by extirpation during the subsequent glacial advance. The Beringian clades had less genetic diversity than populations present south of the ice sheets, suggesting they were founded by relatively small migrating populations.


Description

Modern reconstructions based on partial and skeletal remains reveal that mastodons were very similar in appearance to elephants and, to a lesser degree,
mammoth A mammoth is any species of the extinct elephantid genus ''Mammuthus'', one of the many genera that make up the order of trunked mammals called proboscideans. The various species of mammoth were commonly equipped with long, curved tusks an ...
s, though not closely related to either one. Compared to mammoths and extant elephants, mastodons had a longer and wider body but were not as tall due to their shorter legs. Their limbs were more heavily muscled and had considerably thicker limb bones, making mastodons much more robust in comparison. As in modern elephants, the females were smaller than the males. The average shoulder height for male ''M. americanum'' was about with an average body mass of ; large males were up to in height and in weight. They had a low and long skull with long curved tusks, with those of the males being more massive and more strongly curved. Mastodons had cusp-shaped teeth, very different from mammoth and elephant teeth (which have a series of enamel plates), well-suited for chewing leaves and branches of trees and shrubs. Mastodons are typically depicted with a thick woolly mammoth-like coat of hair, but there is no preserved evidence for this.


Paleobiology


Social behavior

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.


Diet

Mastodons have been characterized as predominantly browsing animals. Of New World proboscids, they appear to have been the most consistent in browsing rather than
grazing In agriculture, grazing is a method of animal husbandry whereby domestic livestock are allowed outdoors to roam around and consume wild vegetations in order to convert the otherwise indigestible (by human gut) cellulose within grass and ot ...
, consuming C3 as opposed to C4 plants, and in 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 lan ...
, due to the need to cross areas of grassland to do so. Most accounts of gut contents have identified coniferous twigs as the dominant element in their diet. Other accounts (e.g., the Burning Tree mastodon) have reported no coniferous content and suggest selective feeding on low, herbaceous vegetation, implying a mixed browsing and grazing diet, with evidence provided by studies of isotopic bone chemistry indicating a seasonal preference for browsing. Study of mastodon teeth microwear patterns indicates that mastodons could adjust their diet according to the ecosystem, with regionally specific feeding patterns corresponding to boreal forest versus cypress swamps, while a population at a given location was sometimes able to maintain its dietary niche through changes in climate and browse species availability.


Distribution and habitat

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 the North American faunal stage according to the North American Land Mammal Ages chronology (NALMA), typically set from less than 240,000 years to 11,000 years BP, a pe ...
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, with an isolated record from Honduras, probably reflecting the results of the maximum expansion achieved by the American mastodon during the Late Pleistocene. A few isolated reports tell of mastodons being found along the east coast up to the New England region, with high concentrations in the Mid-Atlantic region. There is strong evidence indicating that the members of ''Mammut'' were forest dwelling proboscideans, predominating in woodlands and forests, and browsed on trees and shrubs. They apparently did not disperse southward to South America, it being speculated that this was because of a dietary specialization on a particular type of vegetation. A 2022 study of ancient (
Early Pleistocene The Early Pleistocene is an unofficial sub-epoch in the international geologic timescale in chronostratigraphy, being the earliest division of the Pleistocene Epoch within the ongoing Quaternary Period. It is currently estimated to span the time ...
, 2 million years ago)
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 The Kap Kobenhavn Formation is a geologic formation in Greenland. It preserves fossils dating back to the Neogene period. Analysis of environmental DNA in samples from a 2006 expedition to the formation have shown the DNA is two million years o ...
of northern
Greenland Greenland ( kl, Kalaallit Nunaat, ; da, Grønland, ) is an island country in North America that is part of the Kingdom of Denmark. It is located between the Arctic and Atlantic oceans, east of the Canadian Arctic Archipelago. Greenland ...
identified preserved DNA fragments of mastodons, assigned to ''M. americanum''. This suggests that mastodons 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 geological epoch. It began approximately 11,650 cal years Before Present (), after the Last Glacial Period, which concluded with the Holocene glacial retreat. The Holocene and the preceding Pleistocene togeth ...
, with a
boreal forest Taiga (; rus, тайга́, p=tɐjˈɡa; relates to Mongolic and Turkic languages), generally referred to in North America as a boreal forest or snow forest, is a biome characterized by coniferous forests consisting mostly of pines, spruc ...
hosting a species assemblage with no modern analogue. These are among the oldest DNA fragments ever sequenced.


Extinction

Fossil evidence indicates that mastodons probably disappeared from North America about 10,500 years ago as part of a
mass extinction An extinction event (also known as a mass extinction or biotic crisis) is a widespread and rapid decrease in the biodiversity on Earth. Such an event is identified by a sharp change in the diversity and abundance of multicellular organisms. I ...
of most of the
Pleistocene megafauna Pleistocene megafauna is the set of large animals that lived on Earth during the Pleistocene epoch. Pleistocene megafauna became extinct during the Quaternary extinction event resulting in substantial changes to ecosystems globally. The role of ...
that is widely believed to have been a result of human hunting pressure. The latest Paleo-Indians entered the Americas and expanded to relatively large numbers 13,000 years ago, and their hunting may have caused a gradual attrition of the mastodon population. Analysis of tusks of mastodons from the American Great Lakes region over a span of several thousand years prior to their extinction in the area shows a trend of declining age at maturation; this is contrary to what one would expect if they were experiencing stresses from an unfavorable environment, but is consistent with a reduction in intraspecific competition that would result from a population being reduced by human hunting. On the other hand, environmental DNA sequencing indicates that disappearance of megafaunal DNA in North America correlates in time with major changes in plant DNA, suggesting a key role of climate change. Modeling based on the whole of the proboscid fossil record also suggests climate was the more important factor, though with human hunting imposing a "double jeopardy" on mastodons and their kin.


See also

*
Coats–Hines site The Coats–Hines–Litchy site (formerly Coats–Hines) is a paleontological site located in Williamson County, Tennessee, in the Southeastern United States. The site was formerly believed to be archaeological, and identified as one of only a very ...
* List of museums and colleges with mastodon fossils on display * Manis Mastodon site *
Snowmastodon site The Snowmastodon site, also known as the Ziegler Reservoir fossil site, is the location of an important Ice Age fossil excavation near Snowmass Village, Colorado. Fossils were first discovered on October 14, 2010, during the construction of a r ...


Notes


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'


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


* ttp://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 ''Scientific American'', informally abbreviated ''SciAm'' or sometimes ''SA'', is an American popular science magazine. Many famous scientists, including Albert Einstein and Nikola Tesla, have contributed articles to it. In print since 1845, it ...
'',
The Chicago Mastodon
, 18 September 1880, p. 175 {{Authority control Messinian first appearances Miocene proboscideans Pleistocene extinctions Pleistocene proboscideans Pliocene proboscideans Miocene mammals of North America Pliocene mammals of North America Pleistocene mammals of North America Symbols of Michigan Extinct animals of Canada Paleontology in Michigan Taxa named by Johann Friedrich Blumenbach Ringold Formation Miocene Fauna