Mastodon
Mastodon | |
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Mounted M. americanum skeleton ("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
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Other species | |
Species pending reassessment
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Synonyms | |
Genus synonymy
Synonyms of M. americanum
Synonyms of M. matthewi Synonyms of M. vexillarius
Synonyms of M. raki
Synonyms of M. nevadanum
Synonyms of M. cosoensis
Synonyms of "M." borsoni Synonyms of "M." obliquelophus
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A mastodon (mastós 'breast' + odoús 'tooth') is a member of the genus Mammut (German for "mammoth"), which strictly defined, was endemic to North America and lived from the late Miocene to the early Holocene. Mastodons belong to the order Proboscidea, the same order as elephants and mammoths (which belong to the family Elephantidae). Mammut is the type genus of the extinct family Mammutidae, which diverged from the ancestors of modern elephants at least 25 million years ago, during the Oligocene.
M. americanum, known as an "American mastodon" or simply "mastodon," had a long and complex paleontological history spanning all the way back to 1705 when the first fossils were uncovered from Claverack, New York in the American colonies. Naturalists struggled to explain the affinities of the proboscidean because of its uniquely shaped molars, 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 the American Revolution. American historians of the 21st century have argued that findings and displays of its fossils had helped to bolster American nationalism and 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 by Charles Willson Peale in 1804.
Taxonomically, M. americanum was first recognized as a distinct species by
As a member of the Mammutidae, it is defined by zygodont molars which have remained evolutionarily conservative (little changed) throughout the evolution of the family. In comparison to its likely ancestor
Mastodons disappeared along with many other North American animals, predominantly megafauna, as part of the Late Pleistocene extinctions, the causes typically being attributed to Clovis culture hunting, severe climatic phases like the Younger Dryas, or some combination of the two. Of note is that unlike most other extinct North American species, the American mastodon has a recorded last occurrence in the earliest Holocene, making it amongst the last terminal Pleistocene North American faunas to have gone extinct. Today, the American mastodon is one of the most well-known fossil species in both academic research and the public perception, the result of its inclusions in American popular culture.
Research history
Earliest finds
In a letter dating to 1813, Edward Hyde, 3rd Earl of Clarendon (known also as Lord Cornbury) from New York reported to the Royal Society learned society of Great Britain that in 1705, a large-sized tooth was found near the side of the Hudson River by a Dutch country-fellow and was sold to New York General Assembly member Van Bruggen for a gill of rum, and Bruggen eventually gave it to Cornbury. He then stated that he sent Johannis Abeel, a recorder of Albany, New York to dig near the original site of the tooth to find more bones.[1][2]
Abeel reported in a later that he went to the town of Claverack, New York where the original bones were found. American historian Paul Semonin said that the accounts written by Cornbury and Abeel match up with that written by in the July 30, 1705 entry in The Boston News-Letter.[3] The account reported skeletal evidence of an antediluvian (or biblical) "giant" uncovered from Claverack. The femur and one of the teeth both dissolved before they could be further observed, however.[4][1]
Big Bone Lick
In 1739, a French military expedition under the command of Charles III Le Moyne (known also as "Longueil") explored the locality of "Big Bone Lick" (located in what is now the US state of Kentucky) and gathered fossil bones and teeth there.[5] The French naturalist Louis Jean-Marie Daubenton examined the fossil collection brought by Longueuil and compared it with specimens of extant elephants and Siberian mammoths in 1762. Daubenton said that the bones were discovered by Native Americans (probably Abenaki hunter-warriors). He came to the conclusion that the femur and tusk belonged to an elephant while the molars (or cheek teeth) came from a separate giant hippopotamus.[6][7][8]
In Shawnee tradition, the proboscideans roamed in herds and were hunted by giants, who both eventually died out. The accounts told by the Shawnee individuals in 1762 are the oldest known documented interpretations of the "Ohio" fossils, although the traditions may have had been told for generations.[9][10]
In 1767, Peter Collinson credited Irish trader George Croghan for having sent him and Benjamin Franklin fossil evidence of the mysterious proboscideans, using them for his studies. He concluded that the peculiar grinders (the molars) were built for herbivorous diets of branches of trees and shrubs as well as other vegetation, a view later followed by Franklin.[11][12]
In 1768, Scottish anatomist William Hunter recorded that he and his brother John Hunter observed that the teeth were not like those of modern elephants. He determined that the "grinders" from Ohio were of a carnivorous animal but believed that the tusks belonged to the same animal. After examining fossils from Franklin and Lord Shelburne, Hunter was convinced that the "pseudo-elephant", or "animal incognitum" (shortened as "incognitum"), was an animal species separate from elephants that might have also been the same as the proboscideans found in Siberia. He concluded his article with the opinion that although regrettable to philosophers, humanity should be thankful to heaven that the animal, if truly carnivorous, was extinct.[13]
Early American observations
In 1785, Reverend Robert Annan wrote an account recalling an event in which workers discovered bones in his farm near the Hudson River in New York in fall of 1780. The workers found four molars in addition to another that was broken and thrown away. They also uncovered bones, including vertebrae that broke shortly after. 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.[14]
American statesman
In 1799, laborers recovered a thighbone while digging a marl pit at John Masten's farm in Newburgh, New York, and subsequent excavations were observed by a crowd of over a hundred people.[18] American painter and exhibitionist Charles Willson Peale visited the locality in 1801, where he first sketched the fossils then purchased excavation privileges and full ownership of the fossils from Masten and borrowed a loan from the American Philosophical Society (APS) in Philadelphia, 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 his museum in Philadelphia in 1804, and its exhibit was open first to invited members of the American Philosophical Society on December 24 then to the general public on December 25 for an exhibit admission fee in addition to the general admission fee.[19]
The special exhibition attracted thousands of visitors, and the skeleton became a US national symbol.[20] Charles Peale's son Rembrandt Peale took the skeleton to Europe used to promote the fossil proboscidean and have it used as support for Jefferson's final rebuttals against Buffon's arguments for supposed inferiority of American faunas. Author Keith Stewart Thomson argued that the promotion of the "mastodon" skeleton made it a symbol of the strength of American nationalism and that "mammoth" as a term became associated with gigantism. Decades later, the museum bankrupted, and the first skeleton's specimens were sold to some German spectators in around 1848, who eventually sold it to Hessisches Landesmuseum Darmstadt in Germany where it is now displayed. The second skeleton's specimens landed eventually at the American Museum of Natural History.[21]
Other skeletons of Mammut americanum were excavated within the United States in the first half of the 19th century. One of them was collected by American showman
In 1845, another skeleton was excavated from Newburgh by laborers hired by Nathaniel Brewster initially to remove lacustrine deposits to fertilize the neighboring fields. They were observed by a large amount of spectators and uncovered relatively complete fossil evidence of M. americanum.[25][26] The skeleton was exhibited in New York City and other New England towns then was acquired by John Collins Warren for study.[27][28] After Warren's death in 1856, the skeleton was sent to Warren's family but was traded to Harvard Medical School for John Warren's skeleton. The "Warren mastodon", under the request of American paleontologist Henry Fairfield Osborn, was purchased by the American financier J. P. Morgan for $30,000 in 1906 and donated to the American Museum of Natural History where it is exhibited today.[29][26]
Early taxonomic history
In the 1790s, the "American incognitum" was subject to research by multiple taxonomists. Scottish writer Robert Kerr erected the species name Elephas americanus in 1792 based on fossil tusks and "grinders" from the Big Bone Lick locality. He stated that the tusks were similar to elephants while the molars were completely different because they were covered with enamel and had a double row of high conical cusp processes. Kerr was unsure about the taxonomic affinities of the molars and referenced that Thomas Pennant supposed that they belong to an unknown species within the genus Elephas, giving the common name "American elephant."[30]
German naturalist
French naturalist Georges Cuvier also described known fossil proboscidean species back in 1796, although his account was later published in 1799. He considered that the remains uncovered from Siberia were true "mammoths" that had similar dentitions to extant elephants but had some morphological differences. He mentioned the fossil remains that were brought back by Longueil from Ohio back in 1739 and several researchers from previous decades who noted the unusual molars and thought that they belonged to different animals like hippopotamuses. He followed recognition in the previously established species "Elephas americanus" and argued that the species was different from elephants and mammoths and cannot be found amongst living animals due to extinction from catastrophism.[32][33]
The proboscidean species was subject to several other species names given by other taxonomists within the earliest 18th century as well as the genus name Harpagmotherium by the Russian naturalist Gotthelf Fischer von Waldheim in 1808.[24]
Cuvier's taxonomy
In 1806, Cuvier wrote multiple extended research articles on fossil proboscideans of Eurasia and the Americas. He stated that the bones that Buffon previously described from North America were not of elephants but another animal that he referred to as the "mastodonte," or the "animal of Ohio."[34] He reinforced the idea that the extinct "mastodon" was an animal close in relationship to elephants that differed by jaws with large tubercles. He suggested that "mammoth" and "carnivorous elephant" be discontinued as names for the species and that it receive a new genus name instead. Cuvier said that for "mastodonte," he derived the name's etymology (compound μαστός (mastós, "breast") + ὀδούς (odoús, "tooth") from Ancient Greek to mean "nipple tooth," since he thought that it expressed the characteristic form of the teeth.[35]
In 1817, the French naturalist officially established the genus name Mastodon, reaffirming that it is extinct and has left no living descendants. He established that it had an overall body form similar to elephants but had molars more similar to hippopotamuses and pigs that did not serve to grind meat. The first species he erected within Mastodon was Mastodon giganteum, giving it the informal name "great mastodon" and writing that that it is designated to the Ohio proboscidean with abundant fossil evidence, equal size but greater proportions to modern elephants, and diamond-shaped points of the molars. The naturalist also created the second species name Mastodon angustidens and gave it the informal name "narrow-toothed mastodon," diagnosing it as having narrower molars, smaller sizes compared to M. giganteum, and range distributions in Europe and South America.[36] Cuvier also erected several other species of Mastodon originating from other continents in 1824.[37] Despite Cuvier's genus name being younger than multiple other genus names, Mastodon became the most commonly used genus name for the 19th century.[38][24]
Taxonomic problems
"Mastodon" was riddled with major taxonomic problems since species now determined as belonging to other proboscidean genera were classified to Mastodon on the basis of similar dentitions to that of "Mastodon giganteum" (= Mammut americanum), effectively making it a wastebasket taxon.[36][37][39] Various fossil proboscidean species were classified into Mastodon in the 19th century before eventually being reclassified into distinct genera.[24] 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.[40][41][42] Also, many species names erected based on M. americanum remains were erected. As a result, M. americanum has many synonymous names. The issue of synonymous species names were especially apparent in the first half of the 19th century.[24]
Today, the genera that include species formerly classified into Mastodon include Gomphotherium (G. angustidens, G. pyrenaicum, G. productum, G. libycum, G. subtapiroideum, G. steinheimense),[43][44][45] Zygolophodon (Z. turicensis, Z. proavus),[46][47] Cuvieronius (C. hyodon),[48] Stegodon (S. elephantoides),[49] Stegolophodon (S. latidens, S. cautleyi),[50] Anancus (A. avernensis, A. sivalensis, A. perimensis),[51] Tetralophodon (T. longirostris),[52] Choerolophodon (C. pentelici),[53] Stegomastodon (S. mirificus),[54] Rhynchotherium ("R." euhypodon),[41] Stenobelodon (S. floridanus),[55] and Notiomastodon (N. platensis).[40]
In 1830, American naturalist John Davidson Godman created the genus Tetracaulodon plus its species T. Mastodontoideum based on what he determined to be differences between it and Mastodon based on the skull and dentition.[56] Both Richard Harlan and William Cooper pointed out that except for the tusks, all other characteristics of the specimens were consistent with M. giganteum. They therefore argued that there was no reason to assume that the tusks were not just individual variations, a view followed also by George William Featherstonhaugh. Isaac Hays comparatively defended Godman's taxon, which led to a bitter debate regarding the validity of the genus amongst American naturalists.[57]
The validities of both Tetracaulodon and Missourium were rejected by Owen in 1842, although he retained the former name informally.[58] By 1869, American paleontologist Joseph Leidy determined that Mastodon americanus is the senior species synonym and listed M. giganteum as a junior synonym. He also listed Mammut, Harpagmotherium, Mastotherium, Missourium, and Leviathan as synonyms of Mastodon. He also noted that M. americanum as a species was highly variable in morphology.[59][60]
In 1902, American paleontologist Oliver Perry Hay listed Mammut as the prioritized genus name given its status as the oldest genus name, making Mastodon, Tetracaulodon, and Missourium classified as junior synonyms. He also established M. americanum as the type species.[38] The genus name Mastodon was subsequently abandoned by many American paleontologists in favor of Mammut within the early 20th century.[61][62][63][24] In 1942, American paleontologist George Gaylord Simpson said that for his study, he prioritized the historic plus taxonomically correct name Mammut over Mastodon.[64] He continued prioritizing Mammut in 1945, stating that people were generally aware of its taxonomic priorities over Mastodon and that people had refused to use it. He stated that he did not want to either but reluctantly set aside his personal preferences to follow taxonomic rules.[65]
Additional species
In 1921, Osborn created the species name Mastodon matthewi based on distinct molars from the Snake Creek Formation of western Nebraska, naming it in honor of William Diller Matthew. He also erected another species M. merriami from the Thousand Creek Formation in Nevada, which was eventually synonymized with Zygolophodon proavus.[66][47] Osborn in 1926 followed up for Mastodon matthewi by establishing the genus Pliomastodon for the species based on cranial differences from "Miomastodon" (= Zygolophodon).[67]
In 1930, Matthew erected a second species for Pliomastodon named P. vexillarius based on fossil material from the locality of
In 1933,
In 1963, J. Arnold Shotwell and Donald E. Russell created another species Mammut (Pliomastodon) furlongi, assigning it to fossils collected from the Juntura Formation of Oregon. The species name was created in honor of Eustace L. Furlong, who made early fossil collections from the western side of the Juntura Basin.[72]
The genus Pliomastodon was synonymized with Mammut while Miomastodon was synonymized with Zygolophodon by Jeheskel Shoshani and Pascal Tassy in a 1996 appendix,[73] a view that was followed by other authors in later years.[74][75][47]
In 2019, Alton C. Dooley Jr. et al. established Mammut pacificus based on fossils collected from the Diamond Valley Lake in Hemet, California. They also stated that M. oregonense is a nomen dubium and that further analysis needs to be done to confirm whether or not M. furlongi belongs to Zygolophodon instead.[75]
In 2023, Wighart von Koenigswald et al. reviewed the North American species of Zygolophodon and Mammut. They synonymized P. adamsi and P. sellardsi with Mammut matthewi and emended M. nevadanus and M. pacificus to M. nevadanum and M. pacificum, respectively. They also said that they were uncertain of the taxonomic status of M. furlongi, specifically whether or not it was a variant of sexual dimorphism of Z. proavus. Some authors have considered M. nevadanum to be synonymous with M. matthewi while others had retained validity of the species name.[47][75]
Several mammutid species outside of North America are classified to Mammut (or "Pliomastodon"), namely M. borsoni, M. obliquelophus, M. zhupengensis, and M. lufugense (possibly synonymous with M. obliquelophus).[76][77][78][79] Recent research such as that of von Koenigswald et al. in 2023 warned that the genus Mammut should be carefully used for non-North American species.[47]
Classification and evolution
Mammut is the
Although the separation of the Mammutida and Elephantida is strongly supported based on morphological differences, their origins within the late
In the early Neogene phase of evolution, Eozygodon made an appearance in the earliest Miocene (~23-20 Ma) of Africa after Losodokodon. Eozygodon was subsequently succeeded by Zygolophodon by the early Miocene, and the latter dispersed into Eurasia by the late MN3 or MN4 of the Mammal Neogene zones (early Miocene) 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 from Africa prior to the late Miocene.[85][86]
Mammut as currently defined sensu lato (in a loose sense) is most likely
The oldest evidence of mammutids in North America is of a fragmentary molar of Zygolophodon sp. from Massacre Lake, Nevada, dating to 16.5-16.4 Ma (during the Hemingfordian stage of the North American land mammal ages (NALMA)). The only definitively defined species of Zygolophodon from North America is Z. proavus, which occurs in the Barstovian and Clarendonian stages. M? furlongi from the Black Butte in Oregon also dates back to the Clarendonian stage, but the affinities of the species remains unclear. If it truly is a species of Mammut, then its earliest temporal range is recorded at about 10 Ma. The earliest undisputed appearance of Mammut is of M. nevadanum from Thousand Creek Beds, dating back to the early Hemphillian, or 8.0-7.1 Ma. Historically, North American paleontologists considered that North American Zygolophodon evolved into Mammut in an endemic fashion while European workers generally thought that Mammut was a Eurasian immigrant that replaced North American Zygolophodon during the Miocene or Pliocene. Current evidence supports an endemic radiation of North American mammutids because of the gradual appearance of Mammut morphologies and a lack of solid evidence that Mammut sensu stricto (in a strict sense) ever dispersed outside of North America.[47]
M. matthewi is recorded from the late Hemphillian to early Blancan stages. Mammutid specimens of the Hemphillian and Blancan had typically previously been assigned to M. matthewi, but this is seemingly the result of overreliance on stratigraphic positions to define taxa. M. vexillarius, M. raki, and M. cosoensis are definitively recorded from the Blancan, and M. raki specifically is thought to not be synonymous with M. pacificum.[47] M. americanum (known popularly as an "American mastodon" or simply "mastodon") is also stratigraphically recorded first from the early Blancan of the Ringold 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.[47][87][88] From the Irvingtonian to the Rancholabrean, only M. americanum and the newly appearing M. pacificum are recorded, the former having an exceptional level of diversity based on abundant skeletal evidences from the late Pleistocene that is unusual for the typical mammutid fossil record.[75][47]
The following cladogram defines the phylogeny of certain proboscideans, a majority known from
Proboscidea |
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"plesielephantiforms" "mastodonts" | ||||||||||||||||||||||||||||||||||||||||||
Description
Skull
Mammut is diagnosed and differentiated in terms of the skull from Zygolophodon as having a shortened bottom skull base (basicranium) and a high-domed
M. americanum is diagnosed as having a long plus low skull and a shortened mandible.
Endocast anatomy
M. americanum is known by several brain endocasts stored in American museums, although they are seldom subjected to studies. In 1973, neuroscientist Harry J. Jerison studied an endocast of Mammut, recording that it was elephantlike in both size and shape.[97] According to Shoshani et al. in 2006, the endocast of M. americanum features the olfactory bulbs protruding in front of the frontal lobe. They also drew several proboscidean brains to scale, in which the brain of M. americanum was much larger than that of Moeritherium lyonsi but smaller than that of the Asian elephant (Elephas maximus).[98]
Julien Benoit et al. in 2022 explained that while the front tips of the olfactory bulbs of "M." borsoni are partially visible in the brain's back (or dorsal) area, its visibility in M. americanum is debated. Some authors had argued that the olfactory bulbs are visible in the brain's back area while some other authors did not portray them as being visible. The researchers confirmed based on one specimen that the olfactory bulbs are only partially visible in the brain's back area. They also observed that "M." borsoni, despite weighing twice as much as M. americanum, had a 30% lower encephalization quotient (EQ) compared to the other mammutid species, supporting the idea that the evolution of proboscidean encephalization is tied with phylogeny.[89] The Mammutida, as the most basal clade of the Elephantimorpha, has an EQ twice that of Moeritherium and Palaeomastodon. The endocast volume and brain size of the brain M. americanum are larger than those of Stegodon but smaller than those of derived elephantids. It has an EQ that is higher than those of Paleogene proboscideans and "M." borsoni but lower than those of elephantids (extant and extinct) and stegodonts.[99]
The type species is also known from endocasts of ear petrosals.[89] According to Eric G. Ekdale, the ear petrosals of Mammut cannot automatically be distinguished from Mammuthus alone. The subarcuate fossa is absent from the cerebellar surface of the inner ear. The ear petrosals of Mammut are relatively incomplete, leaving several traits to be unable to be observed.[100][101]
Dentition
The family Mammutidae is defined by zygolophodont molars with compressed and sharp transverse ridges plus lack of accessory conules (smaller cusps). The intermediate molars, or the first two molars, are consistently trilophodont, or three-cusped. The dental morphologies of the clade Mammutida contrast strongly with most members of both the Elephantida (
The dentition of Mammut is diagnosed as being strongly zygodont and having no conules. The lophs extend to the long axis of the molars. The first two molars in the dental row have no more than three lophs while the third molars have four lophs plus a cingulum. The upper tusks (or upper incisors) of Mammut differ from those of Zygolophodon by the generally larger sizes, tendency to either straighten or curve up, and the typical lack of any enamel band, although M. vexillarius retains a very narrow strip of enamel in the upper tusks. The lower (or mandibular) tusks tend to be reduced in comparison. M. nevadanum represents the earliest case of a North American mammutid species without any enamel band, although the possibility of it being worn off by wear cannot automatically be eliminated.[90][47] It differs from M. americanum and M. pacificum by the nearly straight but downward-facing upper tusk, whereas males of the latter two species have large and upward-facing upper tusks while females had upward or straight but frontward-directed upper tusks.[75] The reduction to loss of the lower tusks plus reduction of the mandibular symphysis of the derived Mammutidae and Elephantida is an instance of convergent evolution, correlating potentially with the need to reduce heat loss due to the decrease of global temperature and humidity during the late Miocene and Pliocene.[89] Despite the reductions of the lower tusks, they were still present in Neogene species of Mammut. Pleistocene M. americanum comparatively often lacks mandibular tusks, and M. pacificum is always devoid of them.[47] The presence of lower tusks in M. raki separates it as a species from M. pacificum. M. pacificum differs from M. americanum in part by the narrower molars. Both species have broader molars compared to the "narrow-toothed" M. nevadanum, M. raki, and M. cosoensis.[75]
Like its relative "M." borsoni, M. americanum had 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.
Postcranial skeleton
As a result of proboscidean diagnoses focusing mostly on dentition, the postcranial anatomies of fossil proboscideans like Mammut are underrepresented in academic literature. Jennifer A. Hodgson et al. compared the anatomies of Mammut and Mammuthus, mentioning that their postcranial anatomies were studied previously by Stanley John Olsen in 1972 and recognizing that the two genera were only distantly related to each other.[107][108] M. americanum is typically depicted as stocky based on postcranial evidence.[109]
The
The scapula (or shoulder blade) of Mammut has a straight vertebral border, contrasting with a more concave vertebral border of Mammuthus. Hodgson et al. disagreed with the claim by Olsen in 1972 that the neck of the scapula is more constricted in Mammuthus primigenius than Mammut americanum, since neither of the two M. americanum scapulae observed by the researchers have any high constriction there. The pelvis allows for identification of the sex of the species, as male Mammut individuals have a smaller pelvic outlet and wider ilium than female individuals.[107]
Mammut has shorter and more robust limb bones compared to those of derived elephantids, probably the result of it retaining primitive anatomical traits. Both the
External features
The American mastodon (M. americanum) has typically been depicted as having shaggy and brown-colored fur in reconstructions, especially in over a century of paleoart. Despite this, there is little direct evidence supporting the idea that Mammut was actually covered in hair. Supposedly, only one find of fur belonging to the mastodon is of a skull with two small hairy patches of skin from the state of Wisconsin near the city of Milwaukee. These have only been described briefly in the original literature and have never been figured beyond one hair from a scanning 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 whether Mammut needed thick coats for body warmth for their upper ranges at the Arctic and Subarctic and mentioned that it would not have needed them in subtropical climates like in Florida.[109][111][112]
Davis et al. referenced that because Columbian mammoths (Mammuthus columbi) were not thought to be hairy, it is unclear why mastodons would need thick coats in comparison. The former was typically depicted as hairless and the latter as hairy in paleoart, but the mastodon's preferences for closed or mixed habitats puts the speculations into question. They felt the need to portray the latter as hairy so that the average person could differentiate between the two species.[109]
The concept of M. americanum having thick coats of fur was also subjected to study by Asier Larramendi in 2015. He acknowledged that hair is important for thermoregulation in extant elephants but that there is a negative correlation between body size and hair density in mammals. Some mammals have broken this trend before, however, as woolly mammoths (Mammuthus primigenius) evolved to have thick coats of hair and a very short tail in response to cold climates. The idea that the American mastodon had hair is possible because of the seasonal climates, but there are few preserved soft tissues to support this idea, referencing the hairs found in Wisconsin. The supposed evidence of hair reported in the 19th century were actually just green algae filaments. He concluded that the long tail and large body mass both contradict the hypothesis that M. americanum was covered with thick coats of fur, considering it to be probably exaggerated.[110]
Size and weight
According to Larramendi, the mammutids of the genus Mammut were amongst the largest proboscideans to have ever existed. This was especially the case with "M." borsoni, which had a body mass of 14 t (14 long tons; 15 short tons) and probably exceptionally large body sizes. M. americanum in 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 8 t (7.9 long tons; 8.8 short tons) and had a shoulder height measuring 289 cm (114 in). If having the same shoulder heights as extant elephants, M. americanum could have been up to 80% heavier than them. Larger than average individuals may have possibly weighed up to 11 t (11 long tons; 12 short tons). At mean calculations, M. americanum ranged 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 of M. americanum are 2.89 m (9 ft 6 in) in shoulder height and 7.8 t (7.7 long tons; 8.6 short tons) in body mass. These average estimates are larger than those of the Asian elephant. M. americanum had a smaller average shoulder height and a larger body mass compared to the African bush elephant (Loxodonta africana).[110]
The size of the "Overmyer Mastodon," an individual skeleton recovered from the farm of Robert Overmyer northwest of Rochester, Indiana in 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.[113]
A relatively complete skeleton of Mammut sp. from the Gray Fossil Site in Tennessee, which was first uncovered in 2015, dates to the latest Hemphillian, and has an elongated mandibular symphysis and large mandibular tusks, is thought to have been several tonnes larger than M. americanum and even several species of Mammuthus. The specimens are still being prepared for further studies.[47][114]
Paleobiology
Diet
The zygodont molar morphologies of mammutids suggest that they consistently occupied adaptations to
The mastodon commonly browsed on woody plants (i.e. twigs) and fruits, occupying dense
As a result of the consistent browsing specializations of the genus, Mammut occupied an ecological niche that allowed it to actively
Social behaviors
American mastodons may have lived in herds, and it is possible that they were smaller than mammoth herds on average.[125] 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.[126][127] As in modern elephants,[128] there probably was no seasonal synchrony of mating activity, with both males and females seeking out each other for mating when sexually active.[127] 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.[126]
The social behaviors of male mastodon were inferred from one individual skeleton known as the "Buesching mastodon" (known informally as "Fred"),
Paleoecology
Distribution
The range of most species of Mammut is unknown as their occurrences are restricted to few localities, the exception being the American mastodon (M. americanum), which is one of the most widely distributed Pleistocene proboscideans in North America. M. americanum fossil sites range in time from the
M. matthewi is known by a wide distribution range, its westernmost range being in California from the Horned Toad Formation in the late Hemphillian.[47] It has also apparently been identified from the latest Hemphillian based on skull material from the Pascagoula Formation in Tunica Hills, Louisiana. This suggests that Mammut already had an eastern range in the United States by the latest Miocene or earliest Pliocene.[135] Similarly, the same species is recorded from the Palmetto Fauna locality (Bone Valley Formation) in Brewster, Florida in the latest Hemphillian while Mammut sp. is recorded from the Gray Fossil Site in Tennessee.[47]
The American mastodon was only present in the far north of North America during
Late Neogene-Quaternary North America
The overall paleontological record of the Neogene of North America is relatively incomplete compared to other areas of the world. This is the result of a greater fossil record bias of western North America compared to eastern North America, meaning that the western half is better understood in terms of evolutionary and climatic trends while the eastern half is poorly understood. During the late Neogene (8-5 Ma), C4 grasslands spread throughout the North American continent and replaced woodland habitats. In eastern North America were relict woodlands in an increasingly drier climate followed by a large faunal turnover.[139] 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.[140]
The earliest undisputed record of Mammut sensu stricto was of M. nevadanum in the Thousand Creek Formation in Nevada.
The Blancan fossil record suggests a maximum known diversity of four species of Mammut (M. americanum, M. vexillarius, M. raki, and M. cosoensis).
In the Irvingtonian, only M. americanum is recorded to have crossed past the Blancan while M. pacificum replaced the other Blancan species.
Relationship with humans
The exact timing of
As of present, 2 definite Mammut kill sites compatible with Clovis
According to the American paleontologist Daniel C. Fisher, the "Heisler mastodon" site in Calhoun County, Michigan, which recovered about 50% of the skeleton, was proof of meat caching in a pond by Paleoindians in the late Pleistocene. This hypothesis opposes the notion that proboscideans ended up unable to disentangle themselves in marsh wetlands, which he said there is no evidence of. His hypothesis was based on his experiment with partial carcasses of a horse that was preserved in a shallow lake then extracted as well as a Moravian missionary's testimony of Inuit retrieving caribou carcasses from lakes that they probably placed as storage in the cases of excess meat or future limited hunting successes. Fisher said that if his theory is true, then Paleoindian interactions with megafauna (hunting and scavenging) are far more complex than initially thought.[157][156]
In 2023, Michael R. Waters et al. suggested that the Manis Mastodon site in Washington state supported evidence of a mastodon hunt ~13,900 cal. years BP, some 900 years before Clovis culture. Their study was a continuation of a 2011 anatomical study that proposed that osseous (bone) pieces found in a right rib of a mastodon represented fragmented tips of a projectile point, but it had been repeatedly challenged by other authors. Based on anatomical reevaluations, they determined that the bone fragments were embedded in the Manis mastodon rib while it was alive, as evident by the visible healing around the wounded area. Waters and his colleagues stated that the bone pieces were from an external source, explainable by human-made projectile points. They rejected alternate explanations for why bone fragments ended up in the Manis mastodon rib. Based on this, they envisioned that the mastodon individual was wounded by pre-Clovis hunters and got away, giving it time to heal. Afterwards, it died either by natural causes and was scavenged by humans, or it was killed by them on another attack then butchered. This site proves the existence of pre-Clovis hunting technology that the earliest people brought with them when dispersing to North America and made localized adaptations of.[161][162]
In 2017, Steven R. Holen et al. published an article arguing that the
Multiple petroglyphs suggested to have depicted prehistoric proboscideans in North America like mastodons are known within the United States, but they are either fraudulent or depict entities other than mastodons. As a result, suggested rock art of mammoths and mastodons within North America are not sufficiently credible.[165]
Extinction
Mammut, or more specifically the American mastodon, experienced an initial decline in geographical range when it was extirpated from the northernmost ranges of North America ~75,000 years ago. Mammut initially occupied the region during the Last Interglacial (~125,000-75,000 years ago) back when suitable forested habitats were present there but was subsequently extirpated in correlation with environmental changes from the Wisconsin glaciation (MIS 4). The local extirpation, occurring long before human arrival, caused the mastodon range to be limited to areas south of North American ice sheets. The steppe-tundra faunas thrived there during the event whereas boreal forest-adapted faunas underwent declines.[166][33] The trend of recolonization and extirpation appears to have had been a recurring trend in the Pleistocene correlated with repeated returns of forests and wetlands, but what is unclear is why faunas that were able to repeatedly recolonize northern North America during previous interglacial periods were unable to do so again after the Last Glacial Maximum.[136]
The latest Pleistocene of North America records a large extinction phase that resulted in the disappearances of over 30 genera of mammals, the majority of which are considered "megafauna" (~45 kg (99 lb) or larger). Mammut was one of the many genera recorded within North America whose extinction causes are currently unresolved.[167] During the latest Pleistocene of North America, two major events occurred: the development of Clovis culture from 13,200 to 12,800 years ago and the onset of the Younger Dryas cold phase from 12,900 to 11,700 years ago.[168] The extinctions of mammalian megafauna in North America are particularly high akin to those of South America and Australia rather than Eurasia and Africa.[169] 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.[170] 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.[171][172]
Paul L. Koch and Anthony D. Barnosky in 2006 suggested that Mammuthus was well-associated with archeological sites of North America. In comparison, Mammut and the peccary
Of note is that there is a recorded latest survival of the American mastodon in the early Holocene. 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.[174][168]
Cultural significance
Late Pleistocene proboscideans of the Americas such as the American mastodon could have been recognized in Native American oral histories, but they are unlikely to have referenced any specific species. Typically, they may have been depicted in Native American oral history as aggressive and antagonistic beasts.[175] Mastodons may have played ancient roles in Native American cultures of the Pacific Northwest. In 1987, Carl E. Gustafson recovered fossil evidence of a late Pleistocene mastodon far away from where the species would typically roam, the radiocarbon dating confirming a date of about 13,800 years ago. The local tribal members identified the remains as being of game pieces for slahal, a gambling game for dispute settlements and entertainment.[176] 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.[177]
The American mastodon had long been a stand-in within the United States for American nationalism since early American history,[21] and Thomas Jefferson was famously known for having hoped that the Lewis and Clark Expedition would eventually yield evidence of living mastodons in the western frontier of the United States.[178][179] It was a defining symbol of museums according to Brett Barney as evident by a mention of it by Walt Whitman in a passage of the 1855 poem "Song of Myself."[180]
The mastodon became the subject of a Michigan political campaign in 2000 when
In January 2024, Indiana senator Mike Braun and Michigan senator Gary Peters introduced a bipartisan bill to make the mastodon the US national fossil is what is called the "National Fossil Act." Section 1 aims to define the bill's name, Section 2 would investigate the roles of the mastodon in American public life, and Section 3 would designate it as the national fossil under Title 36 of the United States Code. Peters justified that the mastodon represents a unique aspect of Michigan's history and American history, stating that he hoped that its establishment as the national fossil would preserve the histories and encourage new generations of scientists and other researchers to pursue their goals.[184][185]
Located in the Mastodon Ridge park in the Canadian town of Stewiacke, Nova Scotia is a large-sized replica of a mastodon based on a skeleton recovered from Nova Scotia. It was sculpted as a clay model, has a weight of ~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.[186]
The name "mastodon" was adopted in different contexts within the United States. For instance, 4-8-0 locomotives of the late 19th century were originally named "Mastodons" before the name was eventually replaced with "12-wheeler." The name was a reference to the American mastodon. The 4-10-0 locomotive later became known also as "Mastodon."[187][188] In the 1993-1995 show Mighty Morphin Power Rangers, the Black Ranger Zack Taylor had the mastodon ability and controlled the Mastodon Dinozord machine.[189] The name "Mastodon" was also adopted by a heavy metal band when guitarist Bill Kelliher was asked by the guitarist-singer Brent Hinds asked him about the name of the "fossil elephant" after seeing his tattoo of a Bantha skull from the Star Wars franchise, in which the members then agreed to it being the band's name.[190] "Mastodon" is also the name of a blogging social network site that also acquired its name from the extinct proboscidean species.[191]
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
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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) Archived 2015-12-20 at the Wayback 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 of male and female mastodon skeletons at the University of Michigan Mammutidae digital fossil repository
- Scientific American, "The Chicago Mastodon", 18 September 1880, p. 175