Woolly Mammoth (Mammuthus primigenius)

jefferson and mammoth

Symbol of the Pleistocene Epoch, the woolly mammoth (Mammuthus primigenius) —along with the American mastodon— played a crucial role in the emergence of the science of paleontology and the realization of an extensive, prehistoric past.

The remains of mammoths were long familiar to Siberian aboriginals who often encountered remains along the banks of rivers and sold their tusks for the ivory trade. They believed them to be a giant underground mole-like animal that died upon emerging from their burrows (1). Scattered accounts of these remains reached Western Europeans during the 17th century. They variously attributed these remains to human giants, elephants or the biblical behemoth and introduced transliterations of the local names for these animals: mammut, mammount, and ultimately mammoth. However, detailed descriptions or actual specimens weren't available until the first half of the 18th century.

In 1728, the British anatomist Hans Sloane (2) reported on fossil teeth and tusks he examined from Siberia and elsewhere. He conclusively demonstrated that they came from elephants rather than giants or the behemoth. But Sloane had to explain the presence of tropical animals so far north. He concluded that their demise was the result of a drastic climatic change and that their corpses were buried during the biblical flood. Others concluded that the mammoths were killed during the flood and transported by its waters to their final resting place.

mammoth skull
Mammoth skull, based on
Messerschmidt's drawing
(after Cuvier, 1812)

In 1738, Philosophical Magazine published a correspondence by the German anatomist Johann Breyne that included his analysis of fossils and illustrations provided by Daniel Messerschmidt (3). Breyne corroborated Sloane's conclusion that the Siberian mammoth was an elephant and concluded that it was a victim of the flood. More significantly, the article featured Messerschmidt's detailed drawings of the animal's skull, molar, tusk and femur.

The identity of the Siberian mammoth was substantially revised in 1796 by George Cuvier. In the first paper he published Cuvier demonstrated—with the assistance of Messerschmidt's drawings—that the Siberian mammoth was distinct from living elephants. It was a different species. It was also extinct (4). The Siberian mammoth was formally named Elephas primigenius by Johann Friedrich Blumenbach in 1799. Much later, authorities realized that the mammoth was sufficiently distinct to warrant its own genus. It's now known as Mammuthus primigenius.

In the same year that Blumenbach named the mammoth, a hunter discovered one entombed in permafrost in a bank of the Lena River, Siberia. Over the next few years thawing freed enough of the carcass from the ice that the hunter was able to remove the tusks, which he sold to a merchant in the city of Yakust. In 1806 Mikhail (Michael) Adams, a Scottish botanist attached to the Russian Academy of Science, received word of the mammoth while traveling through Yakust. When his party reached it, they found a badly decomposed carcass. However, it still had considerable patches of skin and hair (5) and most of the skeleton was intact. The find was transported to St. Petersburg, where it was mounted at the Zoological Institute. It became known as the Adams mammoth.

Adams mammoth
The Adam's Mammoth (after Tilesius, 1815)

Other, albiet less spectacular, discoveries of the mammoth fossils were made at about the same time in North America. In 1743, Mark Catesby reported on several teeth found in Carolina that black slaves identified as elephant molars. In 1786, David Rittenhouse reported to the American Philosophical Society on a massive molar he collected from Tioga, Pennsylvania. And in 1807, several mammoth molars were collected from Big Bone Lick during the Clark-Jefferson expedition (6). Both William Clark, who excavated the fossils, and Caspar Wistar, who examined them at the White House, realized they belonged to the Siberian "elephant" (mammoth).

Now commonly known as the woolly mammoth, Mammuthus primigenius, is the best known of fossil vertebrates. Significant finds of carcasses, skeletons and other fossils have been reported for over a hundred sites in Eurasia and North America. Frozen and mummified carcasses have enabled the analyses of tissues, DNA, hair, diet and parasites, while extensive skeletal remains have enabled extensive pathology and growth studies. We even have cave paintings of these animals.

The woolly mammoth had about the same height (9-11 ft. 2.8-3.4 m) and weight (4-6 tons) as the Indian elephants, but its overall shape was conspicuously different. The hind legs were much shorter than the forelegs, resulting in a pronounced slope to the back. The skull was narrower from front to back than those of modern elephants and there was a large dome on the top. As indicated by the name, woolly mammoths had a dense coat of hair. The outer layer consisted of long, coarse guard hairs, while a dense layer of fine wool lay underneath.

The tusks were substantially larger than those of modern elephants, sometimes exceeding 13.5 ft. (4.2 m) in length. Whereas elephant tusks are generally straight, those of mammoths are strongly curved and spirally twisted. These tusks swept down and then in, and in some old individuals actually overlapped.

mammoth jaw
Lower jaw of the woolly mammoth
(after Cuvier, 1799)

The molars of the woolly mammoth, like those of other mammoths and elephants were large, specialized structures with a flattened grinding surface. Low ridges of dense enamel run across the surface of the teeth, making them ideal for processing grasses. As with mastodons, a mammoth will have a series of six cheek teeth (premolars and molars) on each side of the jaw (a total of 24 teeth for both upper and lower jaws.) Younger individuals will have three teeth on each side while most adults have two and old individuals have one. As the teeth were worn down, they were replaced by teeth growing in from the back. These teeth were larger and had more ridges than earlier teeth.

The earliest records of woolly mammoth occur in Eurasia about 250,000 years ago. These mammoths were transitional forms. The more advanced woolly mammoths appeared sometime later, and by 100,000 years ago it extended from the British Isles through Siberia and into North America; it moved across the Bering Land Bridge into North America at the beginning of the Rancholabraean Land Mammal Age.

The woolly mammoth was a dominant inhabitant of the "mammoth steppe", a immense dry and cold grassland that extended from Western Europe through most of Siberian and into Beringia (eastern Siberia, Alaska and Yukon). A secondary mammoth steppe —separated from Beringia by the massive Wisconsinan Ice Sheet— was located in the north-central United States and southern Canada. Grasses and sagebrush (Artemesia) dominated the steppe vegetation. The grinding teeth of the woolly mammoth were well suited to feed on steppe grasses. Indeed, stomach contents from well-preserved Siberian specimens indicated a preponderance of grasses and low-lying herbs, although woodier growth may form an important part of its winter diet.

The fossil evidence isn't conclusive, but woolly mammoths probably lived (like modern elephants) in sex-segregated herds. The short and intense growing season of the mammoth steppe suggest that they would have had highly constricted mating and birthing periods. This way, the calves would be born when food was most available. The high food demands of these massive animals suggest they migrated in search of food. The large and strongly curved trunks, which are considerably larger in males, were well suited for male-male contests (7).

The woolly mammoth (Mammuthus primigenius) was one of several species of mammoths (Mammuthus). Unlike the American mastodon, mammoths evolved rapidly into several different and more specialized forms. In fact, the taxonomy of the genus is still a matter of some disagreement, in part because of the prevalence of transitional forms, which some authorities recognize as separate species. A general trend from the more primitive to the more derived forms is indicated by progressive increases in the number of enamel ridges on the molars and the progressive reduction in the width of individual ridges.

The earliest mammoths have been recorded about 4 million years ago from several localities in Africa. Between 3-3.5 million years ago mammoths expanded into Europe. This first non-African species, the southern mammoth (Mammuthus meridionalis) extended throughout much of Eurasia and entered North America in the early Pleistocene. It was about 14 ft (4 m) at the shoulder, lived in woodlands and fed mainly on tree and shrub browse. This widely distributed species is the direct ancestor of both the Columbian mammoth (Mammuthus columbi) of North and Central America, and the steppe mammoth (Mammuthus trogontherii or M. armeniacus) of western Eurasia. The steppe mammoth itself is regarded as an intermediate between the Southern mammoth and the woolly mammoth, while the Columbian mammoth is regarded as an intermediate of the Jefferson mammoth (Mammuthus jeffersonii). The steppe, Columbian and Jefferson mammoths were larger than the woolly mammoth, but two dwarf forms are recognized. Mammuthus exilis is the name for dwarf descendants of Columbian mammoths isolated on the Channel Islands off California. It was about 4-6 ft (1.2-1.8 m) at the shoulders. A dwarf form of the woolly mammoth lived on Wrangle Island, which lies off the coast of northern Russia.

The last of the large mammoths died out between 12,00-10,00 years ago; the Wrangle Island dwarfs survived until at least until 4700 years ago. The extinction of the larger forms has been attributed to both hunting by humans and climatic change. Proponents of climatic causes cite evidence of drastic changes in precipitation and temperature, which resulted in the severe contraction of the grasslands favored by mammoths. Proponents of overhunting cite numerous mass kill sites and the occurrence of spear points among the bones of mammoths (8).

Mammoths belong to the family Elephantidae a diverse family that includes three living species: the African elephant (Loxodonta africana), the forest African elephant (Loxodonta cyclotis) and the Indian elephant (Elephas maximas). Mammoths and elephants belong to the order Proboscidea, which also includes the American mastodon (Mammut americanum).


Woolly Mammoth by Benjamin Waterhouse Hawkins

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Web Pages:

  1. American Museum of Natural History's web page on Woolly Mammoth:
    www.amnh.org/science/biodiversity/extinction/Resources/
    Bestiary/Proboscidea.html#WoollyMammoth
  2. BBS's web article on the Jarvik Mammoth:
    news.bbc.co.uk/2/hi/science/nature/481571.stm
  3. Discovery Channel Canada's web page on Mammoths:
    exn.ca/mammoth/tour.cfm
  4. Explore North's web page on Mammoths:
    www.explorenorth.com/library/weekly/aa032400a.htm
  5. Illinois State Museum's web page on Mammoths:
    www.museum.state.il.us/exhibits/larson/mammuthus.html
  6. Mammoth Site Museum of Hot Springs, South Dakota:
    www.mammothsite.com/
  7. MegaFauna's web page on Mammoths and Mastodons:
    www.kokogiak.com/megafauna/woolly.asp
  8. UC Museum of Paleontology's web page on Mammoths:
    www.ucmp.berkeley.edu/mammal/mammoth/about_mammoths.html
  9. The Unmuseum's web page on Mastodons and Mammoths:
    unmuseum.mus.pa.us/mastodon.htm
  10. Yukon Beringia's web page on the Woolly Mammoth:
    www.beringia.com/02/02maina2.html

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Print Resources:

  1. Breyne, Johann Philip. 1741. "Observations, and a Description of Some Mammoth's Bones Dug Up in Siberia, Proving Them to Have Belonged to Elephants." Philosophical Transactions 40 (1737-1738): 124-139.
  2. Cohen, C. 2002. The Fate of the Mammoth: Fossils, Myths and History. Translated by William Rodarmor. Chicago: University of Chicago Press. Originally published as Le Destin du Mammouth. 1994. Editions du Seuil.
  3. Cuvier, Georges. 1796. "Mémoire sur les épèces d'elephans tant vivantes que fossils, lu à la séance publique de l'Institut National le 15 germinal, an IV." Magasin encyclopédique, 2e anée, 3: 440-445.
  4. Cuvier, Georges. 1799. "Mémoire sur les épèces d'elephans vivantes et fossils, lu le premier pluvose an 4 [January 21, 1796]". Mémoires de l'Institut National des Sciences et des Arts, sciences mathématiques et physiques (mémoires) 2: 1-32.
  5. Kurtén, B. and E. Anderson. 1980. Pleistocene Mammals of North America. New York: Columbia University Press.
  6. Lang, I.A. 2002. Ice Age Mammals of North America: A Guide to the Big, the Hairy, and the Bizarre. Missoula: Mounain Press.
  7. Lister, A. and P. Bahn. 1994. Mammoths. New York: Macmillian.
  8. Stone, R. 2001. Mammoth: The Resurrection of an Ice Age Giant. Cambridge: Perseus Publishing.
  9. Tilesius, G. 1815. "De sceleto mammonteo sivirico ad maris glacialis littora anno 1807, effoso cui praemissae elephantini generis specierum distinctiones." Sectio Primo. ... Conventui exhibuit die 10 Jan. 1810. Mémoires de l'Académie Impérial des Sciences de St. Petersbourg. 5:406-513.

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Notes:

  1. Woolly mammoths carcasses in Siberia are typically found either along high river banks or in yedomas (rounded hills of frozen silt). Both of these geologic features, which are largely composed of permafrost, erode rapidly with the summer thaw or following warm rains. The sight of a partially exposed mammoth carcass emerging from a river bank would lend some credence the stories of the mammut. [go back]
  2. Hans Sloan (1660-1753) was a physician and naturalist who played a major role in the advancement of British natural history. During an extended stay in Jamaica he observed and collected numerous plant and animal specimens. He later published a multi-volume Natural History of the island and served as president of the Royal Society. He amassed one of Europe's greatest natural history collection, that upon his death became the core of the British Museum's natural history collection. [go back]
  3. Daniel Messerschmidt (1785-1735) was a German naturalist sent by the Russian czar Peter the Great on an expedition to Siberia from 1720-1728. During this expedition Messerschmidt excavated a mammoth carcass from the banks of the Indigirka River. In addition to his drawings Messerschmidt supplied Johann Breyne with a molar, a tusk fragment, and testimony from a local Russian regarding the excavation.[go back]
  4. In 1796, Georges Cuvier presented his findings of living and extinct elephants. He convincingly demonstrated that, contrary to conventional taxonomy, the African and Indian elephants were in fact two species. He also convincingly demonstrated that the Siberian mammoth was different from the two modern species and that the "animal de l'Ohio" (American mastodon) belonged to a fourth species. He argued vigorously that both the mammoth and mastodon were extinct. (See Fossils and Extinction for more information.) [go back]
  5. The dense hair of the woolly mammoth demonstrated that it was suited for the frigid Siberian climate. It was no longer necessary to explain the presence of tropical elephants in the Far North. Georges Cuvier, who had earlier proclaimed the mammoth to be a lost (extinct) species, argued that its entombment in ice was evidence for a sudden catastrophe. (See Cuvier's Revolutions of the Globe for more information.) [go back]
  6. In 1807, Thomas Jefferson, then the president of the United States, enlisted William Clark to collect mastodon fossils from Big Bone Lick, Kentucky. Fossils from other species, including the mammoth were also collected during the expedition. (See Discovering the Mastodon: Part 8 - Fossils in the White House for more information.) [go back]
  7. Woolly mammoth males engaged in head-to-head contests in which the curving tusks could strike vulnerable parts of the shoulder and thorax. Presumably, this possibility for serious injury would discourage uneven contest and limit them to evenly matched opponents. In at least one occassion, however, a contest between to evenly matched male mammoths ended up in an unexpectly deadly stalemate. At one fossil site, the tusks of two male Columbia mammoths were interlocked and trapped the two constestants in a death embrace. [go back]
  8. The idea that human hunters caused the extinction of large mammals (megafauna) at the end of the Pleistocene first emerged among scientists early in the 19th century, but generally lost favor in the ensuing decades. The idea was rejuvenated in 1967 by Paul Martin. His "Blitzkreig" hypothesis proposed that the rapid die-off of North American megafauna following the retreat of the continental glaciers resulted from the entry of human hunters armed with lethal Clovis spear point technology. The theory is hotly debated by its many supporters and critics. See the National Humanities Center's web page on Paleoindian Pleistocene Overkill or Steven William's essay on Pleistocene Megafauna Extinctions for more information. [go back]

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