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Mastodons and related early elephants

The American Mastodon, Mammut americanum

Photo: Sergiodlarosa, GNU Free Documentation License, Version 1.2 or later

The lineage shown in the diagram at left was determined from sequencing the mitochondrial genome of the mastodon Mammut americanum, a recently extinct relative of the living elephants that diverged about 26 million years ago. The sequence was obtained from a tooth dated to 50 000–130 000 years ago, increasing the specimen age for which such palaeogenomic analyses have been done by almost a complete glacial cycle.

Using this sequence, together with mitochondrial genome sequences from two African elephants, two Asian elephants, and two woolly mammoths (all of which have been previously sequenced), the authors showed that mammoths are more closely related to Asian than to African elephants. They also used a calibration point lying outside the Elephantidae radiation (elephants and mammoths), which enabled us to estimate accurately the time of divergence of African elephants from Asian elephants and mammoths (about 7.6 million years ago) and the time of divergence between mammoths and Asian elephants (about 6.7 million years ago).

These dates are strikingly similar to the divergence time for humans, chimpanzees, and gorillas, and raise the possibility that the speciation of mammoth and elephants and of humans and African great apes had a common cause. Despite the similarity in divergence times, the substitution rate within primates is more than twice as high as in proboscideans.

Photo and text adapted from: Rohland et al (2007)

(to see the full size version click a few times on the image, depending on your browser)

Mastodon expansion

Distribution of Proboscideans

The distribution history of proboscideans is characterised by two major waves of dispersal: that of the mastodons (ancient elephants) in the Miocene epoch, approximately 25 million years ago, and that of the true elephants in the Pleistocene epoch, beginning approximately 2 million years ago.

Originating in North Africa, proboscideans spread throughout Africa and Eurasia during the Miocene. These migrations were significantly influenced by their ability to adapt to the prevailing climatic conditions. While the warmth-loving deinotheres remained confined to Africa and Eurasia, some groups of mastodons reached North America via the Bering Strait, which was then a landlocked strait located in a cooler climate. Thus, at the Miocene/Pliocene boundary and during the Pliocene epoch, gomphotheres, platybelodonts, and zygolophodonts migrated to North America.

Among the mastodons, however, there were also groups that did not overcome the climatic barrier between Eurasia and North America and remained in their tropical to subtropical habitat: chorolophodonts, stegodons, and stegotetrabelodonts.

That warm climates also represented barriers to dispersal is demonstrated by the example of the North American mastodon populations. Adapted to colder climates in the Pleistocene (evidenced by the discovery of fur), they did not cross the Central American land bridge, which lies in a warm climate. The stegomastodons, on the other hand, were widespread, with Cuvieronius being widespread as far south as South America during the Pleistocene.

Climate adaptations were certainly not the only factors influencing the spread of proboscideans. Otherwise, it would be incomprehensible that proboscideans, such as the Anancus group, which was distributed as far north as Eurasia, never reached North America despite the existence of the Bering Strait.

The second wave of dispersal comprised the elephants themselves. They originated in East Africa during the Pliocene and spread throughout Africa and Eurasia during the Pleistocene. The cold-adapted mammoths even advanced as far as North America during the last Ice Age.

Thus, proboscideans reached their greatest extent during the Pleistocene. All continents were populated except for Australia and Antarctica, which by that time were already impossibly far removed from the other continents. The present-day habitats in South and Southeast Asia (Indian elephant) and in sub-Saharan Africa (African elephant) represent the last remaining refuges of proboscideans.

The world map shows the two waves of proboscidean dispersal (excluding deinotheres):

● Dispersal of mastodons (ancient elephants) beginning approximately 25 million years ago, indicated by green and dark brown arrows.

● Spread of the actual elephants beginning about 2 million years ago with light reddish-brown arrows, which are spatially separated by bridges to show the time interval of this dispersal wave.

Photo and text: Senckenberg Naturmuseum, Frankfurt, Germany.

Gomphotherium arrived in Western Europe approximately 19 to 18 million years ago during the Early Miocene.This major migration event is famously known as the 'Gomphotherium Event' or 'Proboscidean Event'. It marked a radical ecological shift in Europe when lowering sea levels created the 'Gomphotherium land bridge', allowing early proboscideans and other large mammals to disperse outward from Africa and quickly spread across Eurasian habitats.For more context on the arrival and evolution of these ancient relatives of modern elephants, read about the Miocene: The Elephants Reach Europe overview or explore the Gomphotheriidae fossil record.

The southern mammoth (Archidiskodon meridionalis, now often classified as Mammuthus meridionalis) arrived in Western Europe around 2.6 to 2.2 million years ago during the Early Pleistocene.The spread of this species into Europe was a massive, continent-wide faunal turnover event. Fossil evidence indicates it crossed over from Africa via the Middle East, eventually establishing itself in Mediterranean and Western European locations, such as the Upper Valdarno region of Tuscany, Italy and the southern coast of England.

Photo: Senckenberg Naturmuseum, Frankfurt, Germany.
Text: Various sources, including Markov (2012) and Lister et al. (2004)

Zygolophodon, a primitive mastodon, arrived in Western Europe around 18 to 17.5 million years ago. They migrated from Africa into Eurasia during the early Miocene epoch in what palaeontologists call the 'Proboscidean Datum Event', often crossing via the Gomphotherium Land Bridge.

Anancus (Anancus arvernensis) arrived in Western and broader Europe approximately 7.2 million years ago, migrating from Asia during the late Miocene epoch. These ancient, long-tusked relatives of modern elephants subsequently flourished across the continent and survived in the region until the Early Pleistocene.

Photo: Senckenberg Naturmuseum, Frankfurt, Germany.
Text: Various sources, including Wikipedia

The Mammuthus genus arrived in Europe in distinct waves. Early species (like the southern mammoth) arrived roughly 3 million years ago. However, the iconic woolly mammoth (Mammuthus primigenius) evolved in Siberia and migrated into northern Europe significantly later, first appearing there around 200 000 years ago.

The timeline of mammoth presence in northern Europe unfolded in several distinct waves:

3 Million Years BP: The earliest ancestors of the mammoth lineage (Mammuthus rumanus) arrived in Europe from Africa.

200 000 BP: Fully evolved woolly mammoths migrated westward from northeastern Asia and made their way across northern Europe.

Gomphotherium arrived in North America (via the Bering Land Bridge) approximately 16 to 15 million years ago. Following its arrival, it quickly migrated south, reaching southern Canada and sweeping across the rest of the continent by the mid-Miocene epoch (about 16 to 14 million years ago)

Mammoths first arrived in North America via the Bering Land Bridge between 1.5 million and 100 000 years ago, depending on the species.

While Woolly mammoths generally remained in the north, Columbian mammoths migrated deep into the continent, ranging as far south as Nicaragua and Honduras in Central America.

The mammoth genus (Mammuthus) crossed the Bering Land Bridge in multiple waves, leading to two distinct migration profiles in North America:

1. Columbian Mammoths (Mammuthus columbi)

Arrival: The earliest ancestors crossed over from Asia into Alaska roughly 1.5 million years ago.

Southern Limit: They migrated throughout the United States and deep into Mexico. Their southernmost fossil remains have been discovered as far south as Nicaragua and Honduras in Central America. They never crossed into South America.

2. Woolly Mammoths (Mammuthus primigenius)

Arrival: A later, cold-adapted wave of Woolly mammoths crossed the land bridge about 100 000 to 200 000 years ago.

Southern Limit: Unlike their Columbian cousins, Woolly mammoths were uniquely adapted to the harsh tundra-steppe. They primarily remained in Alaska, Canada, and the northern United States.

Ancestral mastodons crossed the Bering Land Bridge into North America multiple times over millions of years.

As glaciers expanded, they moved south and eventually roamed all the way to Central Mexico and Honduras.

Ancestors of the mastodons crossed from Asia over the Bering Land Bridge in multiple waves spanning between 15 million and 2 million years ago. The American mastodon evolved in North America about 3.5 million years ago.

Mastodons preferred forests and wetlands to browse on leaves and twigs, so they only occupied the far north during brief, warm interglacial periods.

As ice sheets grew, mastodons migrated south and their geographic range eventually extended through the contiguous United States, deep into Central Mexico, and even down to the tropics of Honduras.

They suffered complete extinction alongside other megafauna around 10 000 years ago.

Stegomastodon (an extinct type ofGomphothere) evolved in North America from earlier ancestors (like Gomphotherium) roughly 4 million years ago. In terms of its geographic spread, it lumbered across the continent and eventually reached as far south as Central America, with notable fossil finds in places like Mexico (e.g., Jalisco and Michoacán).

The Stegomastodon lineage diverged and became distinct in the Americas around 5.49 million years ago, firmly establishing itself across the United States and Mexico by the early Blancan epoch (about 4 million years ago).

The confirmed genus Stegomastodon got as far down as Central America, with fossils recovered from central Mexico (e.g., Lake Chapala, Jalisco). While older palaeontology literature suggested they crossed the Isthmus of Panama into South America, modern consensus reclassifies those southern animals as distinct, related genera like Notiomastodon.

Extinction: The North American Stegomastodon died out in the early Pleistocene (around 1.2 million years ago), largely pushed to extinction by competition from migrating, more efficient grazers like the true mammoth.

Photo: Senckenberg Naturmuseum, Frankfurt, Germany.
Text: Various sources.

Notiomastodon was an extinct genus of gomphothere (a distant relative of modern elephants) that evolved in South America after crossing the Isthmus of Panama. It evolved into a continent-spanning, lowland-dwelling herbivore that dispersed from Colombia down to Chile, surviving until the end of the Pleistocene.

As a group, Gomphotheres originated in North America and migrated to South America during the late Pliocene, following the formation of the Isthmus of Panama.

The Andean Split: Notiomastodon shared the continent with only one other Gomphothere, Cuvieronius.

While Cuvieronius stayed in the high, cold Andean highlands, Notiomastodon evolved along an eastern, lowland route to become an ecological generalist.

Similar in size to an Asian elephant (up to 2.5 metres tall at the shoulder), Notiomastodon had a stocky build, a domed head, and upper tusks that varied greatly in size and curvature depending on the individual. It had a highly flexible mixed diet, allowing it to adapt to local conditions ranging from scrublands to rainforests.

Notiomastodon ranged more widely across South America than any other proboscidean. Its fossils have been discovered from Colombia in the northwest, across the Brazilian savannas, down into the Pampean region of Argentina, and as far south as the Zona Sur of Chile.Notiomastodon

It primarily inhabited lowlands, seasonally dry open forests, savannahs, and coastal plains. For example, fossils found in the Valle del Cauca of Colombia and in central Chile highlight how these giants used river valleys and ancient lake basins (like Lake Tagua Tagua) as migratory and feeding corridors.

Isotopic and microscopic dental wear analyses indicate Notiomastodon was a generalist mixed feeder, surviving on a diet of C3 shrubs, C4 grasses, and fleshy fruits. In forested ecosystems, it served as an essential seed disperser for large fruit-bearing plants, acting as a prehistoric keystone species.

Becoming extinct around 11 000 to 10 000 years ago, Notiomastodon vanished during the Quaternary megafauna extinction event. Archaeological finds, including a juvenile specimen from Brazil discovered with a stone artefact embedded directly into its skull, suggest that early human hunting played a notable role in their disappearance.

Photo: © cisiopurple
Text: Various sources, including Wikipedia

Cuvieronius is an extinct New World genus of gomphotheres which ranged from southern North America to northwestern South America during the Pleistocene epoch. Reaching a shoulder height of 2.3 metres and a body mass of 3.5 tonnes, it was comparable in size to an Asian elephant.

Cuvieronius inhabited subtropical and tropical latitudes in environments ranging from grasslands to tropical rainforest. Among the last gomphotheres along with the South American Notiomastodon, it became extinct as part of the end Pleistocene extinction event, approximately 12 - 11 000 years ago, along with most other large mammals in the Americas. The extinctions followed the arrival of humans to the Americas, and evidence has been found for human hunting of Cuvieronius, which may have been a factor in its extinction.

Photo: Senckenberg Naturmuseum, Frankfurt, Germany.
Text: Wikipedia

This is a superb reconstruction of Phiomia by Blue Rhino Studio, who do hyper-realistic reconstructions of extinct or living animals for major museums such as the Field Museum.

Phiomia is one of the Elephantiformes, as per the first table above.

Proboscidea → Elephantiformes → Phiomia

Photo: http://www.rhinocentral.com/html/prehistoricsculpt/prehistorics_1.html

Phiomia serridens

Phiomia is an extinct genus of basal proboscid that lived in what is now Northern Africa during the Late Eocene to Early Oligocene some 36-35 million years ago. "Phiomia serridens" means "saw-toothed animal of Faiyum". Faiyum is in Egypt, 130 km southwest of Cairo.

Phiomia was around 2.5 metres (8.2 ft) high, and vaguely resembled a modern elephant, although, based on the shape of its nasal bones, it had only a very short trunk. It had short tusks on the upper jaw and also short shovel-like tusks on the lower jaw that were most likely used for gathering food. These were similar to those of the Miocene Platybelodon, Archaeobelodon, and Amebelodon, but considerably smaller. The tusks in the upper jaw may have been used in defence, or scraping bark off trees.

Photo: LadyofHats
Source: Facsimile, Muséum national d'Histoire naturelle, Paris
Permission: This photograph has been released into the public domain.
Text: Wikipedia

The text here reads:

Coming originally from Africa, the mastodons invaded Europe and Asia 18 million years ago. They made their way to North America via the Bering Bridge ten million years ago and finally reached South America two million years ago.

The development of the tusks and of the trunk reflects the evolution of the mastodons during the Miocene.

Photo: Don Hitchcock 2008

Source: Display at the Vienna Natural History Museum

One version of what Phiomia looked like, with a small trunk, and not showing tusks.

Photo: Don Hitchcock 2008

Source: Display at the Vienna Natural History Museum

Phiomia wintoni

Lower jaw of a prehistoric elephant - from Faiyum, Egypt, located 130 km southwest of Cairo.

Note the shovel-like appearance of the tusks, possibly used to dig up roots and tubers.

Photo: Don Hitchcock 2008

Source: Apparently original, Display at the Vienna Natural History Museum

Phiomia wintoni

Upper jaw of the same species of mastodon, also from Faiyum, Egypt.

Photo: Don Hitchcock 2008

Source: Apparently original, Display at the Vienna Natural History Museum

Gomphotherium

Proboscidea → Elephantiformes → Elephantoidea → Elephantida → Gomphotheriidae → Gomphotherium

Photo: Zdeněk Burian

Zdeněk Michael František Burian (February 11, 1905, Kopřivnice, Moravia - July 1, 1981 Prague) was a Czech painter and book illustrator whose work played a central role in the development of palaeontological reconstructions during a remarkable career spanning five decades.

Originally recognised only in his native Czechoslovakia, Burian's fame later spread to an international audience, and a number of artists later attempted to emulate his style. He is regarded by many as the most influential palaeo-artist of the modern era.

Text: Wikipedia

Gomphotherium angustidens

Teeth of the upper jaw of a mastodon, from Dornbach, Austria

Photo: Don Hitchcock 2008

Source: Apparently original, Display at the Vienna Natural History Museum

Tetralophodon

Proboscidea → Elephantiformes → Elephantoidea → Elephantida → Tetralophodon

Photo: Zdeněk Burian

Tetralophodon longirostris

Photo: Don Hitchcock 2008

Source: Facsimile, Display at the Vienna Natural History Museum

Tetralophodon longirostris

Tetralophodon ("four-ridged tooth") is an extinct gomphothere genus (family Gomphotheriidae). Like typical gomphotheres, Tetralophodon had four tusks and a trunk, and was a browser.

It is from the late Miocene-Pliocene (3-2 million years ago) and was about 8 feet (2.5 metres) high and one ton in weight.

It had four tusks and large, four-cusped teeth.

The "tetra" in Tetralophodon refers to this prehistoric elephant's unusually large, four-cusped cheek teeth, but it could apply equally well to Tetralophodon's four tusks, which mark it as a "gomphothere" proboscid (and thus a close relative of the better-known Gomphotherium).

Like Gomphotherium, Tetralophodon enjoyed an unusually wide distribution during the late Miocene and early Pliocene epochs; fossils of various species have been found as far afield as North and South America, Africa and Eurasia.

Photo: Don Hitchcock 2008

Text: Adapted from Wikipedia and http://dinosaurs.about.com/

Source: Apparently original, Display at the Vienna Natural History Museum

Tetralophodon longirostris

This genus of animals stood about ten feet tall and was a very widespread and successful proboscidean. Tetralophodon lived through the Miocene and Pliocene epoches. These animals have been found in North America, Europe, Asia, and Africa.

The member species are T. longirostris, T. fricki, T. atticus, and T. punjabensis.

Photo: Don Hitchcock 2008

Text: Adapted from Wikipedia

Source: Apparently original, Display at the Vienna Natural History Museum

Tetralophodon longirostris

Lower jaw from Stellenhof, Austria.

Photo: Don Hitchcock 2008

Source: Apparently original, Display at the Vienna Natural History Museum

Tetralophodon longirostris

Upper jaw teeth from Wien - St. Marx, Austria.

Photo: Don Hitchcock 2008

Source: Apparently original, Display at the Vienna Natural History Museum

Tetralophodon longirostris

Upper jaw teeth (maxillaries) of a juvenile from Wien - St. Marx, Austria.

Photo: Don Hitchcock 2008

Source: Apparently original, Display at the Vienna Natural History Museum

Mammut americanum

A superb reconstruction based on the available fossils.

Proboscidea → Elephantiformes → Elephantoidea → Mammutidae → Mammut

Photo: Sergiodlarosa
Permission: GNU Free Documentation License, Version 1.2 or later

Mammut americanum

Upper jaw of a mastodon from St. Louis USA

The American mastodon, Mammut americanum, is an extinct North American proboscidean that lived from about 3.7 million years ago until about 10 000 BC. It was the last surviving member of the mastodon family.

Photo: Don Hitchcock 2008
Source: Apparently original, Display at the Vienna Natural History Museum
Text: Adapted from Wikipedia

Mammut americanum

Third Molar of a mastodon from St. Louis USA

Photo: Don Hitchcock 2008
Source: Apparently original, display at the Vienna Natural History Museum

American mastodon, Mammut americanum

This Ice Age relative of the elephant roamed North America until 13 000 BP.

This specimen is from Missouri, USA, circa 30 000 BP - 40 000 BP.

These magnificent beasts co-existed with humans until they were eventually driven to extinction by a combination of hunting, habitat loss and climate change.

This specimen was originally toured by an American entrepreneur as an 'aquatic monster', with added vertebrae and horizontal tusks.

Acquired by the Museum in 1844, Sir Richard Owen, the first Director of the NHMUK, repositioned the mastodon into its current, accurate position, revealing its relationship to living elephants.

It is known from fossils found ranging from present-day Alaska and New England in the north, to Florida, southern California, and as far south as Honduras and El Salvador. Its main habitat was cold spruce woodlands, and it is believed to have browsed in herds.

Photo: Don Hitchcock 2018
Catalog: NHMUK PV OR 15913
Source and text: Original, display at The Natural History Museum, Cromwell Road, London SW7 5BD
Additional text: Adapted from Wikipedia

Anancus arvernensis

First described from the middle Pliocene of Puy-de-Dome, Perrier (France) as Mastodon arvernensis (or Mastodon from Auvergne) by Croizet et al. (1828) from a fragment of a lower jaw and two milk premolars.

Proboscidea → Elephantiformes → Elephantoidea → Elephantida → Anuncus

Photo: Zdeněk Burian

Fossil remains of various members of the genus Anancus are known from the Late Miocene - Middle Pliocene of India and China, and from the Pliocene, early Pliocene and early Pleistocene of North, Central, Eastern and Southern Africa, and form the Middle to Late Pliocene, from a number of countries south of the former Soviet Union (Moldova, Ukraine, North Caucasus and Transcaucasia, Kazakhstan).

(The text above translated from http://age-of-mammals.ucoz.ru/index/anancus/0-242)

Anancus arvernensis

Molar of a mastodon, locality unknown.

Proboscidea → Elephantiformes → Elephantoidea → Elephantida → Anancus

Photo: Don Hitchcock 2008
Source: Apparently original, display at the Vienna Natural History Museum

Anancus arvernensis, known as 'Pippo' discovered in 1825 near Montecarlo in Tuscany (Italy)

Standing around 3 m (9.8 ft) tall and weighing about 5 tons, it closely resembled a modern elephant. It had two tusks, whereas most other gomphotheres had four. Aside from its somewhat shorter legs, Anancus was also different from modern elephants in that its tusks were much longer, up to 4 m (13 ft) in length. The molars were not composed of lamellae like those of true elephants, but had cusps, like tapir and pig molars. Anancus appears to have lived in forests, eating from trees and shrubs and digging out tubers and roots in the forest floor, and it died out when these forests gave way to grasslands.

Photo: http://age-of-mammals.ucoz.ru/index/anancus/0-242
Source: Apparently original, display at an unknown museum.
Text: http://carnivoraforum.com/

Zygolophodon turicensis

Lower jaw of a mastodon, found at Wolkersdorf. It is closely related to the Mammut genus.

Proboscidea → Elephantiformes → Elephantoidea → Mammutidae → Zygolophodon

Photo: Don Hitchcock 2008
Source: Apparently original, display at the Vienna Natural History Museum

Zygolophodon turicensis

Wolkersdorf im Weinviertel is a town in the district of Mistelbach in the Austrian state of Lower Austria. It is situated in the Weinviertel, about 15 km north of Vienna, on the main road from Vienna to Brno.

Photo: Don Hitchcock 2008
Source: Apparently original, display at the Vienna Natural History Museum

The Manis Mastodon - A newly analysed mastodon rib bone shows that Native Americans were using bone-pointed weapons to take down big game nearly a thousand years earlier than thought, according to a new study. Images of the rib show a broken bone projectile point still stuck where a hunter drove it in 13 800 years ago. The rib was found near the town of Sequim on Washington’s Olympic Peninsula, in the late 1970s. Radiocarbon analysis, DNA samples, genetics work, and other modern techniques recently revealed its true age.

References

1. Croizet l'Abbé, Jobert A. , 1828: Recherches sur les Ossemens Fossiles du Departement du Puy-de-Dome, Chez les principaux libraires 1828, Paris
2. Lister, A., van Essen, H., 2004: The earliest mammoths in Europe, Terra Nostra, 2004, 4. pp 152-154.
3. Markov, G.,, 2012: Mammuthus rumanus, early mammoths, and migration out of Africa: Some interrelated problems, Quaternary International, Volumes 276–277, 25 October 2012, Pages 23-26
4. Rohland N., Malaspinas A., Pollack J., Slatkin M., Matheus P., Hofreiter M., 2007: Proboscidean Mitogenomics: Chronology and Mode of Elephant Evolution Using Mastodon as Outgroup, PLoS Biol 5(8): e207. doi:10.1371/journal.pbio.0050207

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