Mastodon

This article is about the genus. For the social networking platform, seeMastodon (social network).For the band, seeMastodon (band).For other uses, seeMastodon (disambiguation).

"Mammut" redirects here. For other uses, seeMammut (disambiguation).

Amastodon(mastós'breast' +odoús'tooth') is a member of the genusMammut(German for "mammoth" ), which, strictly defined, was endemic to North America and lived from the lateMioceneto the earlyHolocene.Mastodons belong to the orderProboscidea,the same order aselephantsandmammoths(which belong to the familyElephantidae).Mammutis the type genus of the extinct familyMammutidae,which diverged from the ancestors of modern elephants at least 27-25 million years ago, during theOligocene.

Mastodon Temporal range: LateMiocene– earlyHolocene(Possible earliest record of up to ~10 Ma)
MountedM. americanumskeleton ( "Warren mastodon" ),American Museum of Natural History
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Order:	Proboscidea
Family:	†Mammutidae
Genus:	†Mammut Blumenbach,1799
Type species
†Elephas americanus (= †Mammut americanum) Kerr,1792
Other species
†M. matthewiOsborn,1921 †M. vexillariusMatthew,1930 †M. rakiFrick,1933 †M. nevadanumStock,1936 †M. cosoensisSchultz, 1937 †M? furlongiShotwell & Russell, 1963 †M. pacificumDooley et al., 2019 Species pending reassessment †M. borsoniHays,1834 †M. obliquelophusMucha, 1980 †M. lufugenseZhang, 1982 †M. zhupengensisZhang et. al., 1991
Synonyms
Genus synonymy HarpagmotheriumFischer von Waldheim,1808 MastotheriumFischer von Waldheim, 1814 MastodonCuvier,1817 TetracaulodonGodman,1830 MissouriumKoch,1840 LeviathanKoch, 1841 PliomastodonOsborn, 1926 Synonyms ofM. americanum Elephas americanusKerr, 1792 Mammut ohioticumBlumenbach, 1799 Elephas macrocephalusCamper, 1802 Harpagmotherium canadenseFischer de Waldheim, 1808 Elephas mastodontusBarton,1810 Mastotherium megalodonFischer de Waldheim, 1814 Tapirus mastodontoidesHarlan,1825 Tetracaulodon mastodontoideumGodman, 1830 Mastodon ohioticumEichwald,1832 Mastodon cuvieriHays, 1834 Mastodon jeffersoniHays, 1834 Tetracaulodon collinsiiHays, 1834 Tetracaulodon godmaniHays, 1834 Tetracaulodon tapyroidesHays, 1834 Elephas ohioticusde Blainville,1839–1864 Missourium kochiiKoch, 1840 Leviathan missouriiKoch, 1840 Tetracaulodon osagiiKoch, 1841 Tetracaulodon kochiiKoch, 1841 Tetracaulodon bucklandiiGrant, 1842 Missourium theristocaulodonKoch, 1843 Mastodon rugatumKoch, 1845 Elephas rupertianusRichardson,1854 Trilophodon ohioticusFalconer,1868 Mammut progeniumHay,1914 Mastodon americanus plicatusOsborn, 1926 Mammut oregonenseHay, 1926 Mastodon moodieiBarbour,1931 Mastodon americanus alaskensisFrick, 1933 Mastodon acutidensOsborn, 1936 Synonyms ofM. matthewi Mastodon matthewiOsborn, 1921 Pliomastodon sellardsiSimpson,1930 Pliomastodon adamsiHibbard,1944 Synonyms ofM. vexillarius Pliomastodon vexillariusMatthew, 1930 Synonyms ofM. raki Mastodon rakiFrick, 1933 Synonyms ofM. nevadanum Pliomastodon nevadanusStock, 1936 Synonyms ofM. cosoensis Pliomastodon cosoensisSchultz, 1937 Synonyms of "M."borsoni Mastodon vellavusAymard,1847 Mastodon vialletiAymard, 1847 Mastodon buffonisPomel,1848 Mastodon affinisPomel, 1859 Zygolophodon borsoniOsborn, 1926 Mastodon pavlowiOsborn, 1936 Mammut shansienseChow & Chang, 1961 Synonyms of "M."obliquelophus M. praetypicum?Schlesinger, 1917

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 fromClaverack,New Yorkin the American colonies. Naturalists struggled to explain the affinities of the proboscidean because of its uniquely shapedmolars,which have no modern analogues in terms of large mammals. The American mastodon caught the attention of not only European researchers but also influential Americans before and after theAmerican Revolution.American historians of the 21st century, such as Paul Semonin andKeith Stewart Thomson,have argued that findings and displays of its fossils had helped to bolsterAmerican nationalismand contributed to a greater understanding of extinctions. It is known by many skeletons which are now typically on display in American museums, a trend first started byCharles Willson Pealein 1804.

Taxonomically,M. americanumwas first recognized as a distinct species byRobert Kerrin 1792 then classified to its own genusMammutbyJohann Friedrich Blumenbachin 1799, thus making it amongst the first fossil mammal genera to be erected with undisputed taxonomic authority. However, the original genus name was overshadowed by "Mastodon"in the 19th century, which was established formally by the FrenchnaturalistGeorges Cuvierin 1817. The genus became problematic as it served as awastebasket taxonfor over 20 fossil proboscidean species whose dentitions more closely resembled that ofM. americanumthan those of elephantids ordeinotheresup to the early 20th century. After major revisions of proboscidean taxonomy, the genusMammuttoday is defined as the valid genus name and includes 7 definite species, 1 of questionable affinities, and 4 other species from Eurasia that are pending reassessments to other genera.

As a member of the Mammutidae, it is defined by its molars which have a zygodont morphology (where the cusps of the teeth are merged into ridges) which have remained evolutionarily conservative (little changed) throughout the evolution of the family. In comparison to its likely ancestorZygolophodon,Mammutis evolutionarilyderivedwith a reduction to loss of the lower tusks, shortening of themandibular symphysis(the frontmost part of the lower jaw), and particularly long and upward curving upper tusks. As a result of its conservative evolutionary history,Mammutconsistently occupied specializations tobrowsingon leaves, fruits, and woody plants. This allowed mastodons toniche partitionwith other members of Proboscidea in North America, likegomphotheresand theColumbian mammoth,who had shifted to mixed feeding or grazing by the lateNeogene-Quaternary.Mammutas a result was a highly successful proboscidean that had achieved maximum diversity in thePliocenethen is known from abundant fossil evidence in theLate Pleistocene.It is thought to have had behaviors not much different from elephants and mammoths, living in herds, displaying strong degrees ofsexual dimorphism,and entering phases of aggression undermusthin the case of males.

Archeological evidence reveals that mastodons for at least a few thousand years coexisted withPaleoindians,who were the first humans to have dispersed to North America and at least sometimes hunted them based onkill sitesfound in both pre-Clovis andCloviscultural phases.

Mastodons disappeared along with many other North American animals, including most of its largest animals (megafauna), as part of theLate Pleistocene extinctionsaround the end of the Late Pleistocene-early Holocene, the causes typically being attributed to Clovis culture hunting, severe climatic phases like theYounger Dryas,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.

Research history

Main article:Research history ofMammut

Earliest finds

Mammut americanummolartooth,Rotunda Museum

In a letter dating to 1713,Edward Hyde, 3rd Earl of Clarendon(known also as Lord Cornbury) fromNew Yorkreported to theRoyal Societylearned societyofGreat Britainthat in 1705, a large-sized tooth was found near the side of theHudson Riverby aDutchcountry-fellow and was sold toNew York General Assemblymember Van Bruggen for agillof rum, and Bruggen eventually gave it to Cornbury. He then stated that he sent Johannis Abeel, a recorder ofAlbany,New York to dig near the original site of the tooth to find more bones.

Abeel reported in a later that he went to the town ofClaverack,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 inThe Boston News-Letter.The account reported skeletal evidence of anantediluvian(or biblical) "giant"uncovered from Claverack. Thefemurand one of the teeth both dissolved before they could be further observed, however.

Big Bone Lick

Engravings of the femurs of an unspecified extant elephant species (top),M. americanum(middle), and a "Siberian" mammoth (bottom), 1764

In 1739, a French military expedition under the command ofCharles III Le Moyne(known also as "Longueil" ) explored the locality of "Big Bone Lick"(located in what is now the US state ofKentucky) and gathered fossil bones and teeth there.The French naturalistLouis Jean-Marie Daubentonexamined the fossil collection brought by Longueuil and compared it with specimens of extantelephantsand Siberianmammothsin 1762. Daubenton said that the bones were discovered by Native Americans (probablyAbenakihunter-warriors). He came to the conclusion that the femur and tusk belonged to an elephant while themolars(or cheek teeth) came from a separate gianthippopotamus.

InShawneetradition, 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"fossils, although the traditions may have had been told for generations.

In 1767,Peter Collinsoncredited Irish traderGeorge Croghanfor having sent him andBenjamin Franklinfossil 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 anatomistWilliam Hunterrecorded that he and his brotherJohn Hunterobserved 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

The 1806–1808 paintingThe Exhumation of the MastodonbyCharles Willson Peale

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 statesmanThomas Jeffersonstated his thoughts onNotes on the State of Virginia(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 Americansocial degeneracybyGeorges-Louis Leclerc, Comte de Buffon,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 inspireAmerican nationalismand counter against the theory of American degeneracy.

Coloredlithographof the "Missourium"(=Mammut) skeleton, ca. 1845

In 1799, laborers recovered a thighbone while digging amarlpit at John Masten's farm inNewburgh,New York, and subsequent excavations were observed by a crowd of over a hundred people.American painter and exhibitionistCharles Willson Pealevisited 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 theAmerican Philosophical Society(APS) inPhiladelphia,Pennsylvania.In addition to the first skeleton, the second was excavated using a mill-like device to drain a 12 ft (3.7 m) deep marl pit. Peale assembled a complete skeleton in hisPhiladelphia Museumin 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 sonRembrandt Pealetook 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 toHessisches Landesmuseum Darmstadtin Germany where it is now displayed. The second skeleton's specimens landed eventually at theAmerican Museum of Natural History.

Excavation of a specimen in agolf courseinHeath, Ohio,1989 (left) and a replica of the "Burning Tree mastodon"complete skeleton (right)

Other skeletons ofMammut americanumwere excavated within the United States in the first half of the 19th century. One of them was collected by American showmanAlbert C. Kochin what is today theMastodon State Historic SiteatMissouriin 1839. He hypothesized in 1840 that the proboscidean, which he classified asMissourium,was much larger than an elephant, had horizontal tusks plus trunks, and occupied aquatic habitats.He acquired additional fossils from a spring on thePomme de Terre Riverto assemble a mounted skeleton of the "Missouri Leviathan"and briefly exhibited it atSt. Louis.After exhibiting the skeleton throughout Europe, he sold the skeleton to theBritish Museum of Natural History.Richard Owenthen 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 removelacustrine depositsto fertilize the neighboring fields. They were observed by a large amount of spectators and uncovered relatively complete fossil evidence ofM. americanum.The skeleton was exhibited inNew York Cityand other New England towns then was acquired byJohn Collins Warrenfor study.After Warren's death in 1856, the skeleton was sent to Warren's family but was traded toHarvard Medical Schoolfor John Warren's skeleton. The "Warren mastodon", under the request of American paleontologistHenry Fairfield Osborn,was purchased by the American financierJ. P. Morganfor $30,000 in 1906 and donated to the American Museum of Natural History where it is exhibited today.

Early taxonomic history

Mammutskeleton previously displayed by Charles Peale at his museum, now on display atHessisches Landesmuseum Darmstadt

In the 1790s, the "American incognitum"was subject to research by multiple taxonomists. Scottish writerRobert Kerrerected the species nameElephas americanusin 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 conicalcuspprocesses. Kerr was unsure about the taxonomic affinities of the molars and referenced thatThomas Pennantsupposed that they belong to an unknown species within the genusElephas,giving the common name "American elephant."

German naturalistJohann Friedrich Blumenbachalso 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 speciesElephas primigenius?(now known asMammuthus 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 genusMammutand erected the speciesMammut ohioticumbased 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 genusMammutin 1799 makes it the second or third genus to be recognized with taxonomic authority given thatMegalonyxhad been named the same year.

French naturalistGeorges Cuvieralso 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 fromcatastrophism.

The proboscidean species was subject to several other species names given by other taxonomists within the earliest 18th century as well as the genus nameHarpagmotheriumby the Russian naturalistGotthelf Fischer von Waldheimin 1808.

Cuvier's taxonomy

Sketch of the skeleton ofMammut,labeled as "Mastodonte"

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 (compoundμαστός(mastós,"breast" ) +ὀδούς(odoús,"tooth" ) fromAncient Greekto mean "nipple tooth," since he thought that it expressed the characteristic form of the teeth.

In 1817, the French naturalist officially established the genus nameMastodon,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 withinMastodonwasMastodon 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 nameMastodon angustidensand gave it the informal name "narrow-toothed mastodon," diagnosing it as having narrower molars, smaller sizes compared toM. giganteum,and range distributions in Europe and South America.Cuvier also erected several other species ofMastodonoriginating from other continents in 1824.Despite Cuvier's genus name being younger than multiple other genus names,Mastodonbecame the most commonly used genus name for the 19th century.

Taxonomic problems

M. americanumskeleton,Natural History Museum, London.The skeleton was initially assembled byAlbert C. Kochas "Missourium"or"Leviathan",both now synonymous withMammut.

"Mastodon"was riddled with major taxonomic problems since species now determined as belonging to other proboscidean genera were classified toMastodonon the basis of similar dentitions to that of "Mastodon giganteum"(=Mammut americanum), effectively making it awastebasket taxon.Various fossil proboscidean species were classified intoMastodonin 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 onM. americanumremains were erected. As a result,M. americanumhas 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 intoMastodonincludeGomphotherium(G. angustidens,G. pyrenaicum,G. productum,G. libycum,G. subtapiroideum,G. steinheimense),Zygolophodon(Z. turicensis,Z. proavus),Cuvieronius(C. hyodon),Stegodon(S. elephantoides),Stegolophodon(S. latidens,S. cautleyi),Anancus(A. avernensis,A. sivalensis,A. perimensis),Tetralophodon(T. longirostris),Choerolophodon(C. pentelici),Stegomastodon(S. mirificus),Rhynchotherium( "R."euhypodon),Stenobelodon(S. floridanus),andNotiomastodon(N. platensis).

In 1830, American naturalistJohn Davidson Godmancreated the genusTetracaulodonplus its speciesT. Mastodontoideumbased on what he determined to be differences between it andMastodonbased on the skull and dentition.BothRichard HarlanandWilliam Cooperpointed out that except for the tusks, all other characteristics of the specimens were consistent withM. giganteum.They therefore argued that there was no reason to assume that the tusks were not just individual variations, a view followed also byGeorge 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 bothTetracaulodonandMissouriumwere rejected by Owen in 1842, although he retained the former name informally.By 1869, American paleontologistJoseph Leidydetermined thatMastodon americanusis the senior species synonym and listedM. giganteumas a junior synonym. He also listedMammut,Harpagmotherium,Mastotherium,Missourium,andLeviathanas synonyms ofMastodon.He also noted thatM. americanumas a species was highly variable in morphology.

In 1902, American paleontologistOliver Perry HaylistedMammutas the prioritized genus name given its status as the oldest genus name, makingMastodon,Tetracaulodon,andMissouriumclassified as junior synonyms. He also establishedM. americanumas the type species.The genus nameMastodonwas subsequently abandoned by many American paleontologists in favor ofMammutwithin the early 20th century.In 1942, American paleontologistGeorge Gaylord Simpsonsaid that for his study, he prioritized the historic plus taxonomically correct nameMammutoverMastodon.He continued prioritizingMammutin 1945, stating that people were generally aware of its taxonomic priorities overMastodonand 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

Sketch of the reconstructed skull of "Pliomastodon vexillarius"(=Mammut vexillarius), 1930

In 1921, Osborn created the species nameMastodon matthewibased on distinct molars from theSnake Creek Formationof westernNebraska,naming it in honor ofWilliam Diller Matthew.He also erected another speciesM. merriamifrom theThousand Creek FormationinNevada,which was eventually synonymized withZygolophodon proavus.Osborn in 1926 followed up forMastodon matthewiby establishing the genusPliomastodonfor the species based on cranial differences from "Miomastodon"(=Zygolophodon).

In 1930, Matthew erected a second species forPliomastodonnamedP. vexillariusbased on fossil material from the locality ofElephant HillinCalifornia,determining that it differs fromMammutby differences in the skull and that the etymology of the species name was made in honor of paleontological contributions by theStandard Oil Companyof California.

In 1933,Childs Fricknamed the speciesMastodon rakifrom the locality ofTruth or Consequences,New Mexicobased on differences on theheeland M₃tooth fromM. americanus,otherwise having proportions similar to it.In 1936,Chester Stockpublished the species namePliomastodon nevadanusbased on fossils from the Thousand Creek Beds of northwestern Nevada.In 1937, John R. Schultz created the species namePliomastodon? cosoensis,naming it after theCoso MountainsinInyo County,California where skull fossils were recovered.

In 1963, J. Arnold Shotwell and Donald E. Russell created another speciesMammut (Pliomastodon) furlongi,assigning it to fossils collected from theJuntura Formationof Oregon. The species name was created in honor ofEustace L. Furlong,who made early fossil collections from the western side of the Juntura Basin.

The genusPliomastodonwas synonymized withMammutwhileMiomastodonwas synonymized withZygolophodonbyJeheskel ShoshaniandPascal Tassyin a 1996 appendix,a view that was followed by other authors in later years.

In 2019, Alton C. Dooley Jr. et al. establishedMammut pacificusbased on fossils collected from theDiamond Valley LakeinHemet,California. They also stated thatM. oregonenseis anomen dubiumand that further analysis needs to be done to confirm whether or notM. furlongibelongs toZygolophodoninstead.

In 2023, Wighart von Koenigswald et al. reviewed the North American species ofZygolophodonandMammut.They synonymizedP. adamsiandP. sellardsiwithMammut matthewiand emendedM. nevadanusandM. pacificustoM. nevadanumandM. pacificum,respectively. They also said that they were uncertain of the taxonomic status ofM. furlongi,specifically whether or not it was a variant ofsexual dimorphismofZ. proavus.Some authors have consideredM. nevadanumto be synonymous withM. matthewiwhile others had retained validity of the species name.

Several mammutid species outside of North America are classified toMammut(or "Pliomastodon"), namelyM. borsoni,M. obliquelophus,M. zhupengensis,andM. lufugense(possibly synonymous withM. obliquelophus).Recent research such as that of von Koenigswald et al. in 2023 warned that the genusMammutshould be carefully used for non-North American species.

Classification and evolution

Portrait ofJohann Friedrich Blumenbach,who erected the genusMammutin 1799

Mammutis thetype genusof theMammutidae,the sole family of theelephantimorphcladeMammutida(the other elephantimorph clade isElephantida). The Mammutidae is characterized by molars with zygodont-form crests, which have remained morphologically conservative throughout the evolutionary history of the family.Mammutis 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 theDeinotheriidaedue to major differences in dentition and emergence of adult teeth.The Mammutidae is identified as amonophyleticclade, 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 theGomphotheriidaeisparaphyletic(or ancestral to more derived descendant groups in the cladistic sense) in relation to the derivedelephantoidfamiliesStegodontidaeandElephantidae(elephants, mammoths, and relatives).

Although the separation of the Mammutida and Elephantida is strongly supported based on morphological differences, their origins within the latePaleogeneremain uncertain. One hypothesis asserts that the Elephantimorpha is monophyletic if the primitiveElephantiformesgenusPhiomiawas truly ancestral to both the Elephantida and Mammutida. An alternate hypothesis suggests that the Elephantimorpha is diphyletic becausePhiomiais ancestral to gomphotheres whilePalaeomastodonis ancestral to mammutids.The earliest undisputed mammutid genusLosodokodonis recorded inKenya,Africa and firmly establishes the earliest presence of mammutids in the lateOligocene(~27-24 Ma). The Mammutidae, like other Paleogene proboscideans, was therefore an endemic radiation within the continent akin to other endemic mammals likearsinoitheres,hyracoids,andcatarrhineprimatesplus non-endemics such asanthracotheresandhyaenodonts.

In the earlyNeogenephase of evolution,Eozygodonmade an appearance in the earliestMiocene(~23-20 Ma) of Africa afterLosodokodon.Eozygodonwas subsequently succeeded byZygolophodonby 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.

Mammutas currently definedsensu lato(in a broad sense) is most likelypolyphyletic(comprising several unrelated groups). This is because the inclusion of Eurasian mammutid species intoMammutimplies that they share a common origin with North AmericanMammut,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,the last Eurasian mammutid, became extinct during the earliestPleistocene,around 2.5-2 million years ago.

Skeletons of an adult and calfM. americanum,George C. Page Museum

The oldest evidence of mammutids in North America is of a fragmentary molar ofZygolophodonsp. fromMassacre Lake,Nevada, dating to 16.5-16.4 Ma (during theHemingfordianstage of theNorth American land mammal ages(NALMA)). The only definitively defined species ofZygolophodonfrom North America isZ. proavus,which occurs in theBarstovianandClarendonianstages.M? furlongifrom 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 ofMammut,then its earliest temporal range is recorded at about 10 Ma. The earliest undisputed appearance ofMammutis ofM. nevadanumfrom Thousand Creek Beds, dating back to the earlyHemphillian,or 8.0-7.1 Ma. Historically, North American paleontologists considered that North AmericanZygolophodonevolved intoMammutin an endemic fashion while European workers generally thought thatMammutwas a Eurasian immigrant that replaced North AmericanZygolophodonduring the Miocene orPliocene.Current evidence supports an endemic origin of North AmericanMammutfromZygolophodonwithout later migration because of the gradual appearance ofMammutmorphologies and a lack of solid evidence thatMammutsensu stricto(in a strict sense) ever dispersed outside of North America.

M. matthewiis recorded from the late Hemphillian to earlyBlancanstages. Mammutid specimens of the Hemphillian and Blancan had typically previously been assigned toM. matthewi,but this is seemingly the result of overreliance on stratigraphic positions to define taxa.M. vexillarius,M. raki,andM. cosoensisare definitively recorded from the Blancan, andM. rakispecifically is thought to not be synonymous withM. pacificum.M. americanum(known popularly as an "American mastodon" or simply "mastodon" ) is also stratigraphically recorded first from the early Blancan of theRingold Formation,Washington.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 theIrvingtonianto theRancholabrean(from around 1.6 million to 11,000 years ago), onlyM. americanumand the newly appearingM. pacificumare 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 fromendocasts,includingM. americanum:

Proboscidea	Phosphatherium esculliei Numidotherium koholense Moeritherium lyonsi Deinotheriidae Elephantiformes Palaeomastodon beadnelli Elephantimorpha Mammutida Mammut americanum Zygolophodon borsoni Choerolophodon pentelici Gomphotherium augustidens Cuvieronius andium Stegomastodon humboldti Elephantoidea Stegodon insignis Elephantidae Mammuthus meridionalis Mammuthus primigenius Mammuthus columbi Elephas maximus Loxodonta africana Palaeoloxodon Palaeoloxodon antiquus Palaeoloxodon falconeri	"plesielephantiforms" "mastodonts"

Description

Skull

ArticulatedM. americanumskull at thePorter County Museum(left) and an unarticulatedcraniumplus tusks ofM. pacificum(right)

Mammutis diagnosed and differentiated in terms of the skull fromZygolophodonas having a shortened bottom skull base (basicranium) and a high-domedcranium.It is also diagnosed as having an "elephantoid"mandiblewith a shortenedmandibular symphysis(or "brevirostrine" ) and a protruding angular process in the mandible. The diagnosis accounts for both trueMammutspecies andMammutspecies 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 fromZygolophodontoMammut.However, mandibular remains with characteristics ofMammutare not known from any anywhere within the Hemphillian, thus making the transition poorly understood.It differs fromSinomammutby the shortened mandibular symphysis, althoughMammutsometimes retained lower tusks unlike the other genus.

M. americanumis diagnosed as having a long plus low skull and a shortened mandible.Thefrontal bone(or forehead) gives off a flattened appearance compared to extant elephants.The skull ofM. americanumhas manyplesiomorphies(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 backsideinfraorbital foramen.At least some of these features are thought to have been acquired fromPhiomia.The nasal aperture ofM. americanumis oval, whereas that of the skull of "M."cf.obliquelophusis more trapezoidal.M. americanumis also more derived based on the lack of a strong proximal constriction of the incisive fossa of theincisive foramen.M. americanumalso has a high and narroworbitwith a somewhat rectangular outline, but it is less rectangular than that ofEozygodon.The North American mammutid retains a primitive trait in the form of the orbit containing alacrimal bonewith a hole known as the lacrimal foramen. Unlike elephantidans, it has another primitive trait of a short and high-positionedtemporal fossa,a trait shared withEozygodon.

Endocast anatomy

Illustration of theendocastofM. americanumwithout any visibleolfactory bulbs,1906

M. americanumis 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 ofMammut,recording that it was elephantlike in both size and shape.According to Shoshani et al. in 2006, the endocast ofM. americanumfeatures theolfactory bulbsprotruding in front of thefrontal lobe.They also drew several proboscidean brains to scale, in which the brain ofM. americanumwas much larger than that ofMoeritheriumlyonsibut smaller than that of theAsian elephant(Elephas maximus).

Julien Benoit et al. in 2022 explained that while the front tips of the olfactory bulbs of "M."borsoniare partially visible in the brain's back (or dorsal) area, its visibility inM. americanumis 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 asM. americanum,had a 30% lowerencephalization quotient(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 ofMoeritheriumandPalaeomastodon.The endocast volume and brain size of the brainM. americanumare larger than those ofStegodonbut smaller than those of derived elephantids. It has an EQ that is higher than those of Paleogene proboscideans and "M."borsonibut lower than those of elephantids (extant and extinct) and stegodonts.

The type species is also known from endocasts ofear petrosals.According to Eric G. Ekdale, the ear petrosals ofMammutcannot automatically be distinguished fromMammuthusalone. Thesubarcuate fossais absent from the cerebellar surface of the inner ear. The ear petrosals ofMammutare relatively incomplete, leaving several traits to be unable to be observed.

Dentition

M. americanumlower jaw and molars,Phillips Park (Aurora, Illinois)

Front view of the "Warren mastodon" (M. americanum). Note the presence of a single vestigial mandibular tusk.

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 (bunodontmolars that evolutionarily convert to being thin and platelike) and the Deinotheriidae (tapir-likelophodontto 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 ofparallel evolution.The Mammutidae was not the only proboscidean family to have acquired zygodont crested molars, as Neogene species of the gomphothereSinomastodondisplay moderate to weak zygodont crests. Pleistocene species ofSinomastodondo not display zygodont crests, however.

The dentition ofMammutis 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 acingulum.The upper tusks (or upperincisors) ofMammutdiffer from those ofZygolophodonby the generally larger sizes, tendency to either straighten or curve up, and the typical lack of any enamel band, althoughM. vexillariusretains a very narrow strip of enamel in the upper tusks. The lower (or mandibular) tusks tend to be reduced in comparison.M. nevadanumrepresents 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 fromM. americanumandM. pacificumby 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 ofconvergent evolution,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 ofMammut.PleistoceneM. americanumcomparatively often lacks mandibular tusks, andM. pacificumis always devoid of them.The presence of lower tusks inM. rakiseparates it as a species fromM. pacificum.M. pacificumdiffers fromM. americanumin part by the narrower molars. Both species have broader molars compared to the "narrow-toothed"M. nevadanum,M. raki,andM. cosoensis.

Like its relative "M."borsoni,M. americanumhad very large tusks, with some records suggesting lengths of 3 m (9.8 ft) and diameters exceeding 200 mm (7.9 in) were not unusual.In the skull of the earlier-appearingM. matthewi,itsdental alveolusof the right tusk from the locality ofHermiston,Oregon suggests a tusk diameter of approximately 200 mm (7.9 in).Similar to modern elephants,M. americanumalso has degrees of sexual dimorphism indicated by the sizes of the upper tusks. Adult males have tusks 1.15 to 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 ofM. pacificumare thought to have been smaller in length and circumstance than that ofM. americanumand may have similarly exhibited degrees of sexual dimorphism.

Postcranial skeleton

"Cohoes Mastodon" skeleton,New York State Museum

As a result of proboscidean diagnoses focusing mostly on dentition, the postcranial anatomies of fossil proboscideans likeMammutare underrepresented in academic literature. Jennifer A. Hodgson et al. compared the anatomies ofMammutandMammuthus,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. americanumis typically depicted as stocky based on postcranial evidence.

Thevertebral column(also known as the backbone or spine) ofMammutis documented as having a highest point located in the shoulder's front likeMammuthus,but the spines gradually decrease in length then increase slightly in the rear area. The number of ribs and vertebrae ofMammutis not well-documented in paleontological literature and may vary by individual.Mammutusually has 20thoracic vertebraewhereasMammuthususually has 19, but both have documented individuals with 18 of them. The reduction of thoracic vertebrae inMammuthusis considered a derived trait also present in modern elephants. The "Watkins Glen mastodon," for example, has 7cervical vertebrae,20 thoracic vertebrae, 3lumbar vertebrae,and 5sacral vertebrae.They believed thatMammutcould have had as many as 20ribsand that the back ribs were shorter and broader than that ofMammuthus.The tail ofMammutmay have been made up of as many as up to 27caudal vertebrae,suggesting that it had a long tail compared to gomphotheres and elephantids.

M. americanumskeleton,Nova Scotia Museum of Natural History

Thescapula(or shoulder blade) ofMammuthas a straight vertebral border, contrasting with a more concave vertebral border ofMammuthus.Hodgson et al. disagreed with the claim by Olsen in 1972 that the neck of the scapula is more constricted inMammuthus primigeniusthanMammut americanum,since neither of the twoM. americanumscapulae observed by the researchers have any high constriction there. Thepelvisallows for identification of the sex of the species, as maleMammutindividuals have a smallerpelvic outletand wideriliumthan female individuals.

Mammuthas shorter and more robust limb bones compared to those of derived elephantids, probably the result of it retaining primitive anatomical traits. Both thehumerusandradiusof the mammutid genus are robust for instance. Theulnahas a slightly more developedolecranon processand a deepertrochlear notch.Thefemuris somewhat thick, short, and appears to have more expandedcondyles.Possibly, sexual dimorphism could be a factor behind the size of the femur itself. Thetibiadoes not appear much different in bothMammutandMammuthus,whereas thefibulamay 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. pacificumdiffers fromM. americanumby 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

Restoration of a mastodon with fur. The hypothesis thatMammuthad thick coats of fur has been questioned.

The American mastodon (M. americanum) has typically been depicted as having shaggy and brown-colored fur in reconstructions, especially in over a century ofpaleoart.Despite this, there is little direct evidence supporting the idea thatMammutwas 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 ofWisconsinnear the city ofMilwaukee.These have only been described briefly in the original literature and have never been figured beyond one hair from ascanning electron microscope(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 whetherMammutneeded thick coats for body warmth for their upper ranges at theArcticandSubarcticand mentioned that it would not have needed them in subtropical climates like inFlorida.

Davis et al. referenced that becauseColumbian 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 ofM. americanumhaving 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 justgreen algaefilaments. He concluded that the long tail and large body mass both contradict the hypothesis thatM. americanumwas covered with thick coats of fur, considering it to be probably exaggerated.

Size and weight

M. americanummale ( "Beusching mastodon," left) and female ( "Owosso mastodon," right),University of Michigan Museum of Natural History

According to Larramendi, the mammutids of the genusMammutwere amongst the largest proboscideans to have ever existed. This was especially the case with "M."borsoni,which had a body mass of 14–16 t (14–16 long tons; 15–18 short tons) and probably exceptionally large body sizes.M. americanumin comparison to its relative was much smaller, but it was still large in its own right compared to extant elephants. It was not inherently taller than extant elephants, but it was much more robust in body build than them. The Warren mastodon produces a body mass of nearby 7.8 t (7.7 long tons; 8.6 short tons) and had a shoulder height measuring 289 cm (114 in). If having the same shoulder heights as extant elephants,M. americanumcould have been up to 80% heavier than them. Larger than average individuals may have possibly had a shoulder height of 325 cm (128 in) and weighed up to 11 t (11 long tons; 12 short tons). At mean calculations,M. americanumranged at 275 cm (108 in) to 305 cm (120 in) at shoulder height and 6.8 t (6.7 long tons; 7.5 short tons) to 9.2 t (9.1 long tons; 10.1 short tons) in body mass. The average measurements ofM. americanumare 2.9 m (9 ft 6 in) in shoulder height and 8 t (7.9 long tons; 8.8 short tons) in body mass. These average estimates are larger than those of the Asian elephant.M. americanumhad a smaller average shoulder height and a larger body mass compared to theAfrican bush elephant(Loxodonta africana).

Skeletal diagram of the "Warren mastodon" specimen, an adult bull ofM. americanumcompared to a human

The size of the "Overmyer Mastodon," an individual skeleton recovered from the farm of Robert Overmyer northwest ofRochester,Indianain 1976, was estimated by Neal Woodman and Jon W. Branstrator in 2008. They estimated based on the length of the humerus (829 mm (32.6 in)) that the shoulder height of the individual was 230.2 cm (90.6 in), 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 ofMammutsp. from theGray Fossil SiteinTennessee,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 thanM. americanumand even several species ofMammuthus.The specimens are still being prepared for further studies.

Paleobiology

Diet

Restoration of an American mastodon without fur byHeinrich Harder

The zygodont molar morphologies of mammutids suggest that they consistently occupied adaptations tofolivorousdiets throughout their evolutionary history. This means that mammutids such asMammut,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 ofM. americanumindicate that the species consumedspruce needles,pine cones,grass, and occasionallygourdsplusvine leaves.Of note is that whereas mammutids of Eurasia went extinct by the early Pleistocene in association with more seasonal climates,Mammutsurvived in North America and became abundant, although the reason for the latter faunal trend does not have any offered explanation.The browsing specialization ofMammutis supported further by thecoprolites(or fossil dung) ofM. 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, consumingC₃as opposed toC₄plants, and occupying closed forests versus more open habitats. This dietary inflexibility may have prevented them from invading South America during theGreat American Interchange,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 denseconiferous forestsmade up of spruces (Picea) and pines (Pinus) within most of eastern North America. In Florida, it consumed twigs of the genusTaxodiumas well as other woody plants and fruits. Based oncarbon isotopic analysesof mastodons in Florida, they had low δ13C values which indicate C₃browsing specialization.The dietary preferences of North AmericanMammutare thought to have mirrored those of the olderZygolophodon,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" ofMichiganand the "Burning Tree mastodon"of Ohio, mastodons may have also consumed swamp grasses (GlyceriaandZizania) as well as semiaquatic and aquatic plants such as sedge marshes (Carex) that surrounded lakes. They may have additionally ingested other aquatic plants and aquatic invertebrates while consuming more than 100 L (22 imp gal; 26 US gal) 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 thatM. americanumwas 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,Mammutoccupied an ecological niche that allowed it to activelyniche partition(or occupy similar but niche ecological spaces) with other proboscideans of North America in the Neogene-Quaternary.In the Blancan,M. rakishowed few morphological changes. In stark contrast, the contemporary gomphothereStegomastodonshowed progressive developments in response to increasingly arid and extensive grasslands from the Blancan up to the early Irvingtonian, with molar complexities resembling those ofMammuthus.The morphology ofStegomastodonsuggests 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 C₃resource usages, although whether this represents browsing or grazing in the case of mammoths remains unclear.

Social behaviors

American mastodon ( "Perry mastodon" ) skeleton with silhouette in back including the trunk,Wheaton College (Illinois)

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 nearFort Wayne,Indiana in 1998. The mastodon individual lived during the later part of theBølling–Allerød warmingperiod 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 frommusth,although it may have been season-specific compared to living elephants given climatic conditions in North America. He likely engaged inintraspecific competitionlate 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 4 cm (1.6 in) to 5 cm (2.0 in) 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

North American map of the distributions ofM. americanum(blue) andM. pacificum(red) fossil localities of theIrvingtonian-Rancholabrean

The range of most species ofMammutis 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. americanumfossil sites range in time from theBlancantoRancholabreanfaunal 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. americanumwas most common in the eastern United States but rarer in the western US in comparison.M. pacificumis known across California and present as far north as southernIdaho,but it was apparently absent from both theSonoran DesertandMojave Desertregions. The elevated-controlled distributions of coniferous forests within theRocky Mountainregion may have limited populations ofMammutcompared to the other Plio-Pleistocene proboscideans.The easternmost range of the species was in what is nowMontanain the Irvingtonian but may have been extirpated from the area as a result ofIllinoian glaciation.An isolated record ofM. americanumis known fromHonduras,where the genus is not recorded to have extended beyond.

M. matthewiis known by a wide distribution range, its westernmost range being in California from theHorned Toad Formationin the late Hemphillian.It has also apparently been identified from the latest Hemphillian based on skull material from thePascagoula FormationinTunica Hills,Louisiana.This suggests thatMammutalready 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) inBrewster,Florida in the latest Hemphillian whileMammutsp. is recorded from the Gray Fossil Site in Tennessee.

The American mastodon was only present in the far north of North America duringinterglacialperiods, with mitochondrial genome analysis suggesting that separate populations repeatedly colonised the region before becomingextirpatedduring glacial periods.A 2022 study of ancientenvironmental DNAfrom theKap Kobenhavn Formationof northernGreenland,dating theEarly Pleistocene,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 theHolocene,with aboreal foresthosting a species assemblage with no modern analogue. These are among the oldest DNA fragments ever sequenced.

Late Neogene-Quaternary North America

Teleocerasfossigerskeleton,Natural History Museum of Los Angeles County.Mammutcoexisted withrhinocerotidsup to the Pliocene.

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), C₄grasslands 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.

Megalonyxjeffersoniiskeleton,Natural History Museum of Utah.Megalonyxmostly likely descended fromPliometanastesand was present in North America since the late Hemphillian.

The earliest undisputed record ofMammutsensu strictowas ofM. nevadanumin the Thousand Creek Formation in Nevada.Coexistent with the mammutid species were a large variety of other mammals, namely those of theArtiodactyla(antilocaprids,camelids,tayassuids),Carnivora(canids,felids,mustelids,ursids),Eulipotyphla(talpids),Lagomorpha(leporids),Perissodactyla(equids,rhinocerotids), andRodentia(aplodontiids,castorids,geomyids,heteromyids,cricetids,mylagaulids,andsciurids).The latest Hemphillian of Florida based on the Palmetto Fauna of the Bone Valley Formation records the coexistence ofM. matthewiwith similar types of faunas, namelyPilosa(megalonychids), Eulipotyphla (talpids), Lagomorpha (leporids), Carnivora (borophaginecanids,caninecanids, ursids,procyonids,mustelids includinglutrines,felinefelids,machairodontinefelids), Proboscidea (gomphotheres), Perissodactyla (tapirs, rhinocerotids,hipparionineequids), and Artiodactyla (tayassuids,protoceratids,camelids, "pseudoceratines,"cervids,antilocaprids).North America in the late Neogene is understood to have undergone a long-term decline in large mammal diversity (i.e. theDromomerycidae,"Blastomerycinae,"Rhinocerotidae) as a result of C₄grassland expansion, cooler climates, and increased seasonality.

The Blancan fossil record suggests a maximum known diversity of four species ofMammut(M. americanum,M. vexillarius,M. raki,andM. cosoensis).However, the Blancan record ofMammutis relatively rare.M. rakifrom thePalomas Formationof Truth or Consequences in New Mexico is recorded with a few other mammalian faunas, namely the megalonychidground slothMegalonyx,the pocket gopherGeomys,the cricetidSigmodon,the equinEquus,the hipparionineNannippus,and the camelidCamelops.A late Blancan locality known as the Fish Springs Flat Fauna in Nevada reveals that fossils ofM. americanumwere found with those of the leporidHypolagus,lutrineSatherium,equidEquus,camelidGigantocamelus,gopherThomomys,and the ground squirrelSpermophilus.

In the Irvingtonian, onlyM. americanumis recorded to have crossed past the Blancan whileM. pacificumreplaced the other Blancan species.By this time,Mammutwould have coexisted with the elephantidMammuthusand the gomphotheresCuvieroniusandStegomastodon,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.ThePort Kennedy Bone Caveof Pennsylvania is of Irvingtonian age (Middle Pleistocene) and reveals that during this time,M. americanumwas present with the megalonychidMegalonyx wheatleyi,thetremarctinebearArctodus pristinus,thejaguar(Panthera onca), the felidMiracinonyx inexpectatus,and the machairodontineSmilodon gracilis.The Big Bone Lick locality in Kentucky, which dates to the latest Pleistocene (Rancholabrean), indicates the coexistence of the American mastodon with the extantreindeer(Rangifer tarandus) along with various other extinctmegafaunalike ancientbison(Bison antiquus), thecaprinebovidBootherium bombifrons,mylodontidground slothParamylodon harlani,megalonychidMegalonyx jeffersoni,true deerCervalces scotti,equidEquus complicatus,and the Columbian mammoth.

Relationship with humans

Clovisspearpoints,Cleveland Museum of Natural History

The exact timing ofhuman(Homo sapiens) arrival to temperate North America is unclear, but they likely arrived to North America ∼19,000–14,000calibratedyearsBefore Present.They are known within the archeological record asPaleoindiansand eventually gave rise to modern-day Native Americans.Of interest is that in theClovis culturephase, 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 definiteMammutkill sites compatible with Clovislithic technologyhave been recorded compared to 15 ofMammuthusand 1 ofCuvieronius.These two kill sites are thought to be fromKimmswick,Missouri and Pleasant Lake inWashtenaw 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 paleontologistDaniel C. Fisher,the "Heisler mastodon" site inCalhoun 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 aMoravianmissionary's testimony ofInuitretrieving 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.

Cast of a right rib of the "Manis mastodon" with an embedded object and healed wound,Sequim Museum & Arts.The wound has been hypothesized to be the result of pre-Clovis hunting from several sources.

In 2023,Michael R. Waterset al. suggested that theManis Mastodon siteinWashingtonstate 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 theCerutti Mastodon site,located inSan Diego Countyin California, is an archeological site involvingM. americanumthat dates to approximately 130,000 years ago. If true, they stated, the site would imply evidence of now-extinct species ofHomoin North America during theMarine 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 ofM. americanumwere 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 journalNaturedue to how highly prolific it is. Reporters from print presses and digital media published reactions of the article from various North American archeologists, withDonald K. Graysonstating that is was astonishingly bad,Jon M. Erlandsonarguing 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 ofHomo(i.e.H. erectus,Neanderthals,orDenisovans) 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.

Multiplepetroglyphssuggested 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

Summed probability distributions (SPDs) ofMammuthus,Mammut,Nothrotheriops,Equus,Smilodon,and humans in the latest Pleistocene of the United States

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.Mammutinitially occupied the region during theLast 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 theWisconsin 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 theLast Glacial Maximum.

The latest Pleistocene of North America records alarge extinction phasethat resulted in the disappearances of over 30 genera of mammals, the majority of which are considered "megafauna" (~45 kg (99 lb) or larger).Mammutwas 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 theYounger Dryascold 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 explained 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 thatMammuthuswas well-associated with archeological sites of North America. In comparison,Mammutand the peccaryPlatygonuswere far less frequently associated with human sites, potentially suggesting that Paleoindians hunted them less than mammoths. They stated that the current understanding ofMammutassociations with humans could shift if the supposed butchery sites were better understood while that ofPlatygonusis 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 calibratedradiocarbon dates.They determined based on the data that the declines ofMammuthus,Equus,andSmilodonwere correlated with Clovis culture hunting whileMammutand thenothrotheriidground slothNothrotheriopsdid 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 earlyHolocene.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

Political cartoon "Oblivion's Cave—Step Right In, Please" byWinsor McCay,1922

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 inNative 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 forslahal,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 theLewis and Clark Expeditionwould 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 byWalt Whitmanin a passage of the 1855 poem "Song of Myself."

Mastodon replica at theMastodon Ridgepark inStewiacke,Nova Scotia,Canada

The mastodon became the subject of a Michigan political campaign in 2000 whenWashtenaw Community Collegegeology instructor David P. Thomas Sr. aimed to make it the state fossil of Michigan. He, assisted by theSlauson Middle Schoolscience teacher Jeffrey Bradley, was sponsored by the state senatorThaddeus 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 theUniversity of Michigan Museum of Natural Historyto built a mastodon exhibit. In 2002, the mastodon became the state fossil, making it the fourteenthstate symbol.Similarly, the mastodon became the state fossil of Indiana as recently as 2022 due to House Bill 1013, authored by the representativeRandy Frye,passing unanimously.

In January 2024, Indiana senatorMike Braunand Michigan senatorGary Petersintroduced 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 underTitle 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 theMastodon Ridgepark in the Canadian town ofStewiacke,Nova Scotiais 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 ~1,400 kg (3,100 lb), is 3.5 m (11 ft) in shoulder height, and measures 7.5 m (25 ft) 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,4-8-0locomotivesof the late 19th century were originally named "Mastodons"before the name was eventually replaced with" 12-wheeler. "The name was a reference to the American mastodon. The4-10-0locomotive later became known also as "Mastodon."In the 1993-1995 showMighty Morphin Power Rangers,the Black RangerZack Taylorhad the mastodon ability and controlled the Mastodon Dinozord machine.The name "Mastodon" was also adopted by aheavy metalband when guitaristBill Kelliherwas asked by the guitarist-singerBrent Hindsasked him about the name of the "fossil elephant" after seeing his tattoo of aBanthaskull from theStar Warsfranchise, in which the members then agreed to it being theband's name."Mastodon" is also the name of a bloggingsocial networksite that also acquired its name from the extinct proboscidean species.

See also

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

External links

• The Rochester Museum of Science – Expedition Earth Glaciers & Giants
• Illinois State Museum – Mastodon
• Calvin College Mastodon Page
• American Museum of Natural History – Warren Mastodon
• BBC Science and Nature:Animals – American mastodon Mammut americanum
• BBC News – Greek mastodon find 'spectacular'
• Missouri State Parks and Historic Sites – Mastodon State Historic Site
• Saint Louis Front Page – Mastodon State Historic Site
• Story of the Randolph Mastodon (Earlham College)Archived2015-12-20 at theWayback Machine
• The Florida Museum of Natural History Virtual Exhibit – The Aucilla River Prehistory Project:When The First Floridians Met The Last Mastodons
• 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 ofmaleandfemalemastodon skeletons at the University of MichiganMammutidae digital fossil repository
• Scientific American,"The Chicago Mastodon",18 September 1880, p. 175