View source for Woolly mammoth

{{Short description|Extinct species of mammoth}}
{{pp-semi-indef|small=yes}}
{{Featured article}}
{{Use dmy dates|date=December 2020}}
{{Use British English|date=April 2013}}
{{Speciesbox
| fossil_range = [[Middle Pleistocene]] – Late [[Holocene]] {{fossil range|0.40|0.0037}}
| image = Siegsdorfer Mammut.jpg
| image_upright = 1.1
| image_caption = Largest European specimen, a male at Südostbayerisches Naturkunde- und Mammut-Museum, [[Siegsdorf]]
| genus = Mammuthus
| species = primigenius
| range_map = Woolly Mammoth Late Pleistocene Dymaxion Biogeographic Distribution.png
| range_map_upright = 1.1
| range_map_caption = [[Dymaxion map]] showing the [[Late Pleistocene]] distribution of ''M. primigenius'' in blue (light blue [[Beringia|was land at the time]]), inferred from fossil finds
| authority = ([[Johann Friedrich Blumenbach|Blumenbach]], 1799)
| synonyms = {{collapsible list|bullets = true|title=List:
| ''Elephas primigenius'' Blumenbach, 1799
| ''Elephas mammonteus'' Cuvier, 1799
| ''Mammuthus boreus'' Brookes, 1828
| ''Mammonteus primigenius'' Osborn, 1924
| ''Elephas boreus'' Hay, 1924
}}
}}

The '''woolly mammoth''' ('''''Mammuthus primigenius''''') is an extinct species of [[mammoth]] that lived from the [[Middle Pleistocene]] until its extinction in the [[Holocene]] epoch. It was one of the last in a line of mammoth species, beginning with the African ''[[Mammuthus subplanifrons]]'' in the early [[Pliocene]].  The woolly mammoth began to diverge from the [[steppe mammoth]] about 800,000 years ago in [[Siberia]]. Its closest extant relative is the [[Asian elephant]]. The [[Columbian mammoth]] (''Mammuthus columbi'') lived alongside the woolly mammoth in North America, and [[DNA studies]] show that the two [[Hybrid (biology)|hybridised]] with each other. Mammoth remains were long known in Asia before they became known to Europeans. The origin of these remains was long debated and often explained as the remains of [[legendary creature]]s. The mammoth was identified as an extinct elephant species by [[Georges Cuvier]] in 1796.

The appearance and behaviour of the woolly mammoth are among the best studied of any [[Prehistory|prehistoric]] animal because of the discovery of frozen carcasses in Siberia and [[North America]], as well as skeletons, teeth, stomach contents, dung, and depiction from life in prehistoric [[cave painting]]s. It was roughly the same size as modern [[African elephant]]s. Males reached shoulder heights between {{cvt|2.67|and|3.49|m}} and weighed between {{cvt|3.9|and|8.2|MT}}. Females reached {{cvt|2.3|-|2.6|m}} in shoulder heights and weighed between {{cvt|2.8|-|4|MT}}. A newborn calf weighed about {{cvt|90|kg}}. The woolly mammoth was well adapted to the cold environments present during [[glacial period]]s, including the [[last ice age]]. It was covered in fur, with an outer covering of long guard hairs and a shorter undercoat. The colour of the [[coat (animal)|coat]] varied from dark to light. The ears and tail were short to minimise [[frostbite]] and heat loss. It had long, curved [[tusk]]s and four [[Molar (tooth)|molars]], which were replaced six times during the lifetime of an individual. Its behaviour was similar to that of modern elephants, and it used its tusks and [[Elephant#Trunk|trunk]] for manipulating objects, fighting, and foraging. The diet of the woolly mammoth was mainly grasses and [[Cyperaceae|sedges]]. Individuals could probably reach the age of 60. Its habitat was the [[mammoth steppe]], which stretched across northern Eurasia and North America.

The woolly mammoth coexisted with early humans, who hunted the species for food, and used its bones and tusks for making art, tools, and dwellings. The population of woolly mammoths declined at the end of the Late Pleistocene, with the last populations on mainland Siberia persisting until around 10,000 years ago, although isolated populations survived on [[Saint Paul Island (Alaska)|St. Paul Island]] until 5,600 years ago and on [[Wrangel Island]] until 4,000 years ago. After its extinction, humans continued using its [[ivory]] as a raw material, a tradition that continues today. The completion of the mammoth genome project in 2015 sparked discussion about potentially [[Revival of the woolly mammoth|reviving the woolly mammoth]] through several various methods. However, none of these approaches are currently feasible.

==Taxonomy==

[[File:Boltunov mammoth.jpg|thumb|left|Copy of an interpretation of the "[[Adams mammoth]]" carcass from around 1800, with [[Johann Friedrich Blumenbach]]'s handwriting]]
Remains of woolly mammoths were long known by [[native Siberians]] and [[Indigenous peoples of the Americas|Native Americans]], who had various ways of interpreting them. Remains later reached other parts of Asia and Europe, where they were also interpreted in various ways prior to [[modern science]].<ref name="Lister, 2007. p. 48-55">Lister, 2007. p. 48-55</ref><ref name="Switek"/> The first woolly mammoth remains studied by scientists were examined by the British physician [[Hans Sloane]] in 1728 and consisted of fossilised teeth and tusks from [[Siberia]]. Sloane was the first to recognise that the remains belonged to [[elephant]]s.<ref>{{cite journal | last = Sloane | first = H. | title = An Account of Elephants Teeth and Bones Found under Ground | journal = Philosophical Transactions | volume = 35 | date = 1727–1728 | pages = 457–471|bibcode = 1727RSPT...35..457S | doi = 10.1098/rstl.1727.0042 | issue = 399–406 | doi-access = free }}</ref> Sloane turned to another biblical explanation for the presence of elephants in the Arctic, asserting that they had been buried during the [[Genesis flood narrative|Great Flood]], and that Siberia had previously been tropical before a drastic climate change.<ref>{{Cite journal  | last =  Sloane | first = H.  | title = Of Fossile Teeth and Bones of Elephants. Part the Second  | journal = Philosophical Transactions  | volume = 35  | pages = 497–514 | year = 1727–1728 |bibcode = 1727RSPT...35..497S  | doi =  10.1098/rstl.1727.0048  | issue =  399–406 | doi-access = free}}</ref> Others interpreted Sloane's conclusion slightly differently, arguing the flood had carried elephants from the [[tropics]] to the [[Arctic]]. Sloane's paper was based on travellers' descriptions and a few scattered bones collected in Siberia and Britain. He discussed the question of whether or not the remains were from elephants, but drew no conclusions.<ref name="ans">{{cite web | url = http://www.ansp.org/museum/jefferson/otherFossils/mammuthus.php |archive-url= https://web.archive.org/web/20070927013323/http://www.ansp.org/museum/jefferson/otherFossils/mammuthus.php |archive-date= 27 September 2007 |title=Woolly Mammoth (''Mammuthus primigenius'')|access-date=29 September 2007|publisher=The Academy of Natural Sciences|year=2007|author=The Academy of Natural Sciences}}</ref> In 1738, the German zoologist [[Johann Philipp Breyne]] argued that mammoth fossils represented some kind of elephant. He could not explain why a tropical animal would be found in such a cold area as Siberia, and suggested that they might have been transported there by the Great Flood.<ref name="Breyne1741">{{Cite journal | last1 = Breyne | first1 = J. P. | last2 = s. | first2 = T. | last3 = Wolochowicz | first3 = M. | doi = 10.1098/rstl.1737.0026 | title = A Letter from John Phil. Breyne, M. D. F. R. S. To Sir Hans Sloane, Bart. Pres. R. S. With Observations, and a Description of Some Mammoth's Bones Dug up in Siberia, Proving Them to Have Belonged to Elephants | journal = Philosophical Transactions of the Royal Society of London | volume = 40 | issue = 445–451 | pages = 124–138 | year = 1737 | bibcode = 1737RSPT...40..124P | doi-access = free }}</ref>

In 1796, [[French people|French]] biologist [[Georges Cuvier]] was the first to identify the woolly mammoth remains not as modern elephants transported to the Arctic, but as an entirely new species. He argued this species had gone [[Extinction|extinct]] and no longer existed, a concept that was not widely accepted at the time.<ref name="Switek"/><ref name=cuvier1796>{{Cite journal | last = Cuvier | first = G. | title = Mémoire sur les épèces d'elephans tant vivantes que fossils, lu à la séance publique de l'Institut National le 15 germinal, an IV | journal = Magasin Encyclopédique, 2e Anée | pages = 440–445 | language = fr| year = 1796}}</ref> Following Cuvier's identification, the German naturalist [[Johann Friedrich Blumenbach]] gave the woolly mammoth its scientific name, ''Elephas primigenius'', in 1799, placing it in the same [[genus]] as the [[Asian elephant]] (''Elephas maximus''). This name is Latin for "the first-born elephant". Cuvier coined the name ''Elephas mammonteus'' a few months later, but the former name was subsequently used.<ref name="Lectotypes"/> In 1828, the British naturalist [[Joshua Brookes]] used the name ''Mammuthus borealis'' for woolly mammoth fossils in his collection that he put up for sale, thereby coining a new genus name.<ref>{{cite book | last = Brookes | first = J. | title = A catalogue of the anatomical & zoological museum of Joshua Brookes | publisher = Richard Taylor | volume = 1 | location = London | year = 1828 | page = 73 | url = http://www.rhinoresourcecenter.com/index.php?s=1&act=refs&CODE=ref_detail&id=1165235146 | url-status = live | archive-url = https://web.archive.org/web/20150924090940/http://www.rhinoresourcecenter.com/index.php?s=1&act=refs&CODE=ref_detail&id=1165235146 | archive-date = 24 September 2015}}</ref>

[[File:Mammuthus primigenius lectotypes.jpg|thumb|1930s illustration of the [[lectotype]] [[molars]]; the left one is now lost.]]
Where and how the word "mammoth" originated is unclear. According to the'' Oxford English Dictionary'', it comes from an old [[Vogul]] word ''mēmoŋt'', "earth-horn".<ref>{{cite web|title=Mammoth entry in Oxford English Dictionary|url=http://www.oed.com/view/Entry/113184|year=2000}}</ref> It may be a version of ''mehemot'', the [[Arabic]] version of the biblical word "behemoth". Another possible origin is [[Estonian language|Estonian]], where ''maa'' means "earth", and ''mutt'' means "[[mole (animal)|mole]]". The word was first used in Europe during the early 17th century, when referring to ''maimanto'' tusks discovered in Siberia.<ref name="Lister, 2007. p. 48-55"/> The American president [[Thomas Jefferson]], who had a keen interest in [[palaeontology]], was partially responsible for transforming the word "mammoth" from a noun describing the prehistoric elephant to an adjective describing anything of surprisingly large size. The first recorded use of the word as an adjective was in a description of a wheel of cheese (the "[[Cheshire Mammoth Cheese]]") given to Jefferson in 1802.<ref name=oed>Simpson, J. (2009). "[http://public.oed.com/aspects-of-english/word-stories/mammoth/ Word Stories: Mammoth] {{webarchive|url=https://web.archive.org/web/20130522163541/http://public.oed.com/aspects-of-english/word-stories/mammoth/ |date=22 May 2013 }}." ''Oxford English Dictionary Online'', Oxford University Press. Accessed 5 June 2009.</ref>

By the early 20th century, the taxonomy of extinct elephants was complex. In 1942,  American palaeontologist [[Henry Fairfield Osborn]]'s posthumous monograph on the [[Proboscidea]] was published, wherein he used various taxon names that had previously been proposed for mammoth species, including replacing ''Mammuthus'' with ''Mammonteus'', as he believed the former name to be invalidly published.<ref name="Osborn 1942">{{Cite book
 |url          = https://archive.org/stream/proboscideamonog02osbo#page/1116/mode/2up
 |title        = Proboscidea: A monograph of the discovery, evolution, migration and extinction of the mastodonts and elephants of the world
 |year         = 1942
 |last1        = Osborn
 |first1       = H. F.
 |editor-last  = Percy
 |editor-first = M. R.
 |location     = New York
 |publisher    = J. Pierpont Morgan Fund
 |volume       = 2
 |pages        = 1116–1169
 |url-status      = live
 |archive-url   = https://web.archive.org/web/20160313142753/https://archive.org/stream/proboscideamonog02osbo#page/1116/mode/2up
 |archive-date  = 13 March 2016
}}</ref> Mammoth taxonomy was simplified by various researchers from the 1970s onwards, all species were retained in the genus ''Mammuthus'', and many proposed differences between species were instead interpreted as [[intraspecific]] variation.<ref>{{Cite journal | title = Origin and evolution of the Elephantidae | last1 = Maglio | first1 = V. J. | year = 1973 | volume = 63 | issue = 3 | pages = 1–149 | journal = Transactions of the American Philosophical Society | doi = 10.2307/1379357 | jstor = 1379357| doi-access = free }}</ref>

Osborn chose two molars (found in Siberia and [[Osterode (district)|Osterode]]) from Blumenbach's collection at [[Göttingen University]] as the [[lectotype]] specimens for the woolly mammoth, since [[holotype]] designation was not practised in Blumenbach's time, and one of the specimens Blumenbach had described had been of the unrelated [[straight-tusked elephant]] (''Palaeoloxodon antiquus''). Soviet palaeontologist [[Vera Gromova]] further proposed the former should be considered the lectotype with the latter as paralectotype. Both molars were thought lost by the 1980s, and the more complete "Taimyr mammoth" found in Siberia in 1948 was therefore proposed as the [[neotype]] specimen in 1990. Resolutions to historical issues about the validity of the genus name ''Mammuthus'' and the [[type species]] designation of ''E. primigenius'' were also proposed.<ref>{{cite journal | last1 = Garutt | first1 = W. E. | last2 = Gentry | first2 = A. | last3 = Lister | first3 = A. M. | title = ''Mammuthus'' Brookes, 1828 (Mammalia, Proboscidea): proposed conservation, and ''Elephas primigenius'' Blumenbach, 1799 (currently ''Mammuthus primigenius''): proposed designation as the type species of ''Mammuthus'', and designation of a neotype | journal = Bulletin of Zoological Nomenclature | volume = 47 | pages = 38–44 | url = https://www.biodiversitylibrary.org/page/12230365#page/52/mode/1up | year = 1990 | url-status = live | archive-url = https://web.archive.org/web/20150713065658/http://www.biodiversitylibrary.org/page/12230365#page/52/mode/1up | archive-date = 13 July 2015 | doi = 10.5962/bhl.part.2651| doi-access = free }}</ref><ref>{{Cite journal |last=Lister |first=Adrian M. |date=July 2017 |title=On the type material and evolution of North American mammoths |url=https://linkinghub.elsevier.com/retrieve/pii/S1040618216310576 |journal=Quaternary International |language=en |volume=443 |pages=14–31 |doi=10.1016/j.quaint.2017.02.027|bibcode=2017QuInt.443...14L |doi-access=free }}</ref> The paralectotype molar (specimen GZG.V.010.018) has since been located in the Göttingen University collection, identified by comparing it with Osborn's illustration of a cast.<ref name="Lectotypes">{{cite journal | title= The rediscovery of type material of Mammuthus primigenius (Mammalia: Proboscidea) | last1 = Reich | first1 = M. | last2 = Gehler | first2 = A. | last3= Mohl| first3=D. | last4=van der Plicht | first4=H. | last5=Lister | first5=A. M. | journal=International Mammoth Conference IV (Poster) | year=2007 | page=295}}</ref><ref>{{Cite journal | last1 = Reich | first1 = M. | last2 = Gehler | first2 = A. | title = Giants' Bones and Unicorn Horns Ice Age Elephants Offer 21st Century Insights | journal = Collections – Wisdom, Insight, Innovation | volume = 8 | pages = 44–50 | year = 2008 }}</ref>

===Evolution===

[[File:Cuvier elephant jaw.jpg|thumb|[[Georges Cuvier]]'s 1796 comparison between the mandible of a woolly mammoth (bottom left and top right) and an [[Indian elephant]] (top left and bottom right)]]
The earliest known members of the Proboscidea, the [[clade]], which contains modern elephants, existed about 55 million years ago around the [[Tethys Sea]]. The closest known relatives of the Proboscidea are the [[sirenians]] (dugongs and manatees) and the [[hyraxes]] (an order of small, herbivorous mammals). The family [[Elephantidae]] existed 6 million years ago in Africa and includes the modern elephants and the mammoths. Among many now-extinct clades, the [[mastodon]] (''Mammut'') is only a distant relative of the mammoths and part of the separate family [[Mammutidae]], which diverged 25 million years before the mammoths evolved.<ref>Lister, 2007. pp.&nbsp;18–21</ref> The Asian elephant is the closest extant relative of the mammoths.<ref name=Shoshani_etal_2007>{{Cite journal | last1 = Shoshani | first1 = J. | last2 = Ferretti | first2 = M. P. | last3 = Lister | first3 = A. M. | last4 = Agenbroad | first4 = L. D. | last5 = Saegusa | first5 = H. | last6 = Mol | first6 = D. | last7 = Takahashi | first7 = K. | title = Relationships within the Elephantinae using hyoid characters | doi = 10.1016/j.quaint.2007.02.003 | journal = Quaternary International | volume = 169–170 | pages = 174–185 | year = 2007 |bibcode = 2007QuInt.169..174S }}</ref> The following [[cladogram]] shows the placement of the woolly mammoth among Late Pleistocene and modern proboscideans, based on genetic data:<ref>{{Cite journal |last1=Palkopoulou |first1=Eleftheria |last2=Lipson |first2=Mark |last3=Mallick |first3=Swapan |last4=Nielsen |first4=Svend |last5=Rohland |first5=Nadin |last6=Baleka |first6=Sina |last7=Karpinski |first7=Emil |last8=Ivancevic |first8=Atma M. |last9=To |first9=Thu-Hien |last10=Kortschak |first10=R. Daniel |last11=Raison |first11=Joy M. |date=2018 |title=A comprehensive genomic history of extinct and living elephants |journal=Proceedings of the National Academy of Sciences|volume=115 |issue=11 |pages=E2566–E2574 |bibcode=2018PNAS..115E2566P |doi=10.1073/pnas.1720554115 |issn=0027-8424 |pmc=5856550 |pmid=29483247 |doi-access=free}}</ref><ref>{{Cite journal |last1=Baleka |first1=Sina |last2=Varela |first2=Luciano |last3=Tambusso |first3=P. Sebastián |last4=Paijmans |first4=Johanna L.A. |last5=Mothé |first5=Dimila |last6=Stafford |first6=Thomas W. |last7=Fariña |first7=Richard A. |last8=Hofreiter |first8=Michael |date=January 2022 |title=Revisiting proboscidean phylogeny and evolution through total evidence and palaeogenetic analyses including Notiomastodon ancient DNA |journal=iScience |language=en |volume=25 |issue=1 |pages=103559 |doi=10.1016/j.isci.2021.103559 |pmc=8693454 |pmid=34988402|bibcode=2022iSci...25j3559B }}</ref>

{{clade
|{{clade
 |1={{extinct}}''[[Mammut americanum]]'' (American mastodon)
 |2={{clade
 |1={{extinct}}''[[Notiomastodon platensis]]'' (South American gomphothere)
 |label2=[[Elephantidae]]
 |2={{clade
 |1={{clade
 |1={{clade
  |1={{extinct}}''[[Mammuthus columbi]]'' (Columbian mammoth)
  |2={{extinct}}'''''Mammuthus primigenius''''' ('''woolly mammoth''')}}
  |3={{clade
   |1=''[[Elephas maximus]]'' (Asian elephant)}} }}
   |2={{clade
 |1={{extinct}}''[[Palaeoloxodon antiquus]]'' (European straight-tusked elephant)
 |2={{clade
   |1=''[[Loxodonta cyclotis]]'' (African forest elephant)
   |2=''[[Loxodonta africana]]'' (African bush elephant)
}} }} }} }} }}}}

[[File:Mammoth molars compared.jpg|thumb|left|Comparison between the lower molars of a woolly mammoth (above) and a [[southern mammoth]]; note the lower number of [[Tooth enamel|enamel]] ridges in the older species (below)]] 
Within six weeks from 2005 to 2006, three teams of researchers independently assembled [[Mitochondrial DNA|mitochondrial genome]] profiles of the woolly mammoth from [[ancient DNA]], which allowed them to confirm the close evolutionary relationship between mammoths and Asian elephants (''Elephas maximus'').<ref>{{Cite journal | last1 = Gross | first1 = L. | title = Reading the Evolutionary History of the Woolly Mammoth in Its Mitochondrial Genome | doi = 10.1371/journal.pbio.0040074 | journal = PLOS Biology | volume = 4 | issue = 3 | pages = e74 | year = 2006 | pmid =  20076539| pmc =1360100 | doi-access = free }}</ref><ref name="Year">{{cite journal |last1=Cooper |first1=A. |title=The year of the mammoth |journal=PLOS Biology |date=2006 |volume=4 |issue=3 |pages=e78 |doi=10.1371/journal.pbio.0040078|pmid=16448215 |pmc=1360097 |doi-access=free }}</ref> A 2015 DNA review confirmed Asian elephants as the closest living relative of the woolly mammoth.<ref>{{Cite journal|title = Elephant Natural History: A Genomic Perspective|journal = Annual Review of Animal Biosciences|date = 2015|pmid = 25493538|pages = 139–167|volume = 3|issue = 1|doi = 10.1146/annurev-animal-022114-110838|first1 = Alfred L.|last1 = Roca|first2 = Yasuko|last2 = Ishida|first3 = Adam L.|last3 = Brandt|first4 = Neal R.|last4 = Benjamin|first5 = Kai|last5 = Zhao|first6 = Nicholas J.|last6 = Georgiadis}}</ref> African elephants (''Loxodonta africana'') branched away from this clade around 6 million years ago, close to the time of the similar split between [[Pan (genus)|chimpanzee]]s and humans.<ref>{{Cite journal | last1 = Krause | first1 = J. | last2 = Dear | first2 = P. H. | last3 = Pollack | first3 = J. L. | last4 = Slatkin | first4 = M. | last5 = Spriggs | first5 = H. | last6 = Barnes | first6 = I. | last7 = Lister | first7 = A. M. | last8 = Ebersberger | first8 = I. | last9 = Pääbo | first9 = S. | doi = 10.1038/nature04432 | last10 = Hofreiter | first10 = M. | title = Multiplex amplification of the mammoth mitochondrial genome and the evolution of Elephantidae | journal = Nature | volume = 439 | issue = 7077 | pages = 724–727 | year = 2005 | pmid =  16362058| bibcode = 2006Natur.439..724K | s2cid = 4318327 }}</ref> A 2010 study confirmed these relationships and suggested the mammoth and Asian elephant lineages diverged 5.8–7.8 million years ago, while African elephants diverged from an earlier common ancestor 6.6–8.8 million years ago.<ref>{{Cite journal | last1 = Rohland | first1 = N. | last2 = Reich | first2 = D. | last3 = Mallick | first3 = S. | last4 = Meyer | first4 = M. | last5 = Green | first5 = R. E. | last6 = Georgiadis | first6 = N. J. | last7 = Roca | first7 = A. L. | last8 = Hofreiter | first8 = M. | editor1-last = Penny | editor1-first = David | doi = 10.1371/journal.pbio.1000564 | title = Genomic DNA Sequences from Mastodon and Woolly Mammoth Reveal Deep Speciation of Forest and Savanna Elephants | journal = PLOS Biology | volume = 8 | issue = 12 | pages = e1000564 | year = 2010 | pmid =  21203580| pmc =3006346 | doi-access = free }}</ref>

In 2008, much of the woolly mammoth's [[chromosome|chromosomal]] [[DNA]] was mapped. The analysis showed that the woolly mammoth and the African elephant are 98.55% to 99.40% identical.<ref>[http://www.post-gazette.com/pg/08325/929359-115.stm Will findings recreate the woolly mammoth?] {{webarchive|url=https://web.archive.org/web/20090211051321/http://www.post-gazette.com/pg/08325/929359-115.stm |date=11 February 2009 }}, Pittsburgh Post-Gazette, 20 November 2008</ref> The team mapped the woolly mammoth's nuclear genome sequence by extracting DNA from the hair follicles of both a 20,000-year-old mammoth retrieved from [[permafrost]] and another that died 60,000 years ago.<ref>{{cite web |url=https://www.sciencedaily.com/releases/2008/11/081119140712.htm |title=Woolly-Mammoth Genome Sequenced |website=Science Daily |date=20 November 2008 |access-date=22 June 2010 |url-status=live |archive-url=https://web.archive.org/web/20110111171128/https://www.sciencedaily.com/releases/2008/11/081119140712.htm |archive-date=11 January 2011 }}</ref> In 2012, [[proteins]] were confidently identified for the first time, collected from a 43,000-year-old woolly mammoth.<ref>{{Cite journal | last1 = Cappellini | first1 = E. | last2 = Jensen | first2 = L. J. | last3 = Szklarczyk | first3 = D. | last4 = Ginolhac | first4 = A. L. | last5 = Da Fonseca | first5 = R. A. R. | last6 = Stafford | first6 = T. W. | last7 = Holen | first7 = S. R. | last8 = Collins | first8 = M. J. | last9 = Orlando | first9 = L. | last10 = Willerslev | first10 = E. | last11 = Gilbert | first11 = M. T. P. | last12 = Olsen | first12 = J. V. | doi = 10.1021/pr200721u | title = Proteomic analysis of a Pleistocene mammoth femur reveals more than one hundred ancient bone proteins | journal = Journal of Proteome Research | volume = 11 | issue = 2 | pages = 917–926 | year = 2012 | pmid =  22103443}}</ref>

Since many remains of each species of mammoth are known from several localities,  reconstructing the evolutionary history of the genus through [[morphology (biology)|morphological]] studies is possible. Mammoth species can be identified from the number of enamel ridges (or [[Lamella (anatomy)|lamellar plates]]) on their [[molars]]; primitive species had few ridges, and the number increased gradually as new species evolved to feed on more abrasive food items. The crowns of the teeth became deeper in height and the skulls became taller to accommodate this. At the same time, the skulls became shorter from front to back to minimise the weight of the head.<ref name="Mammoth evolution"/><ref>{{Cite journal | last = Ferretti | first = M. P. | title = Structure and evolution of mammoth molar enamel | journal = Acta Palaeontologica Polonica | volume = 48 | series = 3 | pages = 383–396 | year = 2003 }}</ref> The short and tall skulls of woolly and [[Columbian mammoths]] (''Mammuthus columbi'') were the culmination of this process.<ref name="Lister 2007 12 43"/>

The first known members of the genus ''Mammuthus'' are the African species ''[[Mammuthus subplanifrons]]'' from the [[Pliocene]], and ''[[Mammuthus africanavus|M. africanavus]]'' from the [[Pleistocene]]. The former is thought to be the ancestor of later forms. Mammoths entered Europe around 3 million years ago. The earliest European mammoth has been named ''[[M. rumanus]]''; it spread across Europe and China. Only its molars are known, which show that it had 8–10 enamel ridges. A population evolved 12–14 ridges, splitting off from and replacing the earlier type, becoming the [[southern mammoth]] (''M. meridionalis'') about 2–1.7 million years ago. In turn, this species was replaced by the [[steppe mammoth]] (''M. trogontherii'') with 18–20 ridges, which evolved in eastern Asia around 1 million years ago.<ref name="Mammoth evolution">{{Cite journal |last1=Lister |first1=A. M. |last2=Sher |first2=A. V. |last3=Van Essen |first3=H. |last4=Wei |first4=G. |year=2005 |title=The pattern and process of mammoth evolution in Eurasia |url=http://doc.rero.ch/record/13496/files/PAL_E277.pdf |journal=[[Quaternary International]] |volume=126–128 |pages=49–64 |bibcode=2005QuInt.126...49L |doi=10.1016/j.quaint.2004.04.014}}</ref> Mammoths derived from ''M. trogontherii'' evolved molars with 26 ridges 400,000 years ago in Siberia and became the woolly mammoth. The earliest identified forms of woolly mammoth date to the [[Middle Pleistocene]].<ref name="Mammoth evolution" /> Woolly mammoths entered North America about 100,000 years ago by crossing the [[Bering Strait]].<ref name="Lister 2007 12 43">Lister, 2007. pp.&nbsp;12–43</ref>

===Subspecies and hybridisation===

{{multiple image
|align = right
|total_width = 350
|image1 = Mammuthus primigenius fraasi.JPG
|alt1 = 
|image2 = Mammuthus jeffersonii.jpg
|alt2 = Museum skeleton, with curved tusks crossing at the tips
|footer = Cast of an intermediate form between ''M. trogontherii'' and ''M. primigenius''; ''M. p. fraasi'', [[Staatliches Museum für Naturkunde Stuttgart]] (left), and specimen (formerly assigned to ''M. jeffersonii'') suggested to be a [[Hybrid (biology)|hybrid]] between [[Columbian mammoth|Columbian]] and woolly mammoths at the [[American Museum of Natural History]] (right)
}}

Individuals and populations showing transitional morphologies between each of the mammoth species are known, and primitive and [[Synapomorphy|derived]] species coexisted until the former disappeared. The different species and their intermediate forms have been termed "[[chronospecies]]". Many taxa intermediate between ''M. primigenius'' and other mammoths have been proposed, but their validity is uncertain; depending on author, they are either considered primitive forms of an advanced species or advanced forms of a primitive species.<ref name="Mammoth evolution"/> Distinguishing and determining these intermediate forms has been called one of the most long-lasting and complicated problems in [[Quaternary]] palaeontology. Regional and intermediate species and subspecies such as ''M. intermedius'', ''M. chosaricus'', ''M. p. primigenius'', ''M. p. jatzkovi'', ''M. p. sibiricus'', ''M. p. fraasi'', ''M. p. leith-adamsi'', ''M. p. hydruntinus'', ''M. p. astensis'', ''M. p. americanus'', ''M. p. compressus'', and ''M. p. alaskensis'' have been proposed.<ref name="Osborn 1942"/><ref>{{cite conference | last1 = Foronova | first1 = I. V. | last2 = Zudin | first2 = A. N. | title = Discreteness of evolution and variability in mammoth lineage: method for group study | conference= The World of Elephants – Proceedings of the 1st International Congress| location = Rome | year = 2001 | pages = 540–543 | url = http://www.sovraintendenzaroma.it/content/download/4886/62925/file/540_543.pdf | url-status = live | archive-url = https://web.archive.org/web/20141024131427/http://www.sovraintendenzaroma.it/content/download/4886/62925/file/540_543.pdf | archive-date = 24 October 2014}}</ref><ref>{{cite journal | last =  Foronova | first =  I.V. | title =  ''Mammuthus intermedius'' (Proboscidea, Elephantidae) from the late Middle Pleistocene of the southern Western and Central Siberia, Russia: the problem of intermediate elements in the mammoth lineage | journal =  Russian Journal of Theriology | series =  2 | volume =  13 | issue = 2 | url =  http://zmmu.msu.ru/rjt/articles/ther13_2_071_082_Foronova_for_Inet.pdf | pages =  71–82 | year =  2014 | url-status =  live | archive-url =  https://web.archive.org/web/20160304195548/http://zmmu.msu.ru/rjt/articles/ther13_2_071_082_Foronova_for_Inet.pdf | archive-date =  4 March 2016 | doi = 10.15298/rusjtheriol.13.2.03 | doi-access =  free }}</ref>

A 2011 genetic study showed that two examined specimens of the Columbian mammoth were grouped within a subclade of woolly mammoths. This suggests that the two populations interbred and produced fertile offspring. A North American type formerly referred to as ''M. jeffersonii'' may be a [[Hybrid (biology)|hybrid]] between the two species.<ref>{{Cite journal | last1 = Enk | first1 = J. | last2 = Devault | first2 = A. | last3 = Debruyne | first3 = R. | last4 = King | first4 = C. E. | last5 = Treangen | first5 = T. | last6 = O'Rourke | first6 = D. | last7 = Salzberg | first7 = S. L. | last8 = Fisher | first8 = D. | last9 = MacPhee | first9 = R. | doi = 10.1186/gb-2011-12-5-r51 | last10 = Poinar | first10 = H. | title = Complete Columbian mammoth mitogenome suggests interbreeding with woolly mammoths | journal = Genome Biology | volume = 12 | issue = 5 | pages = R51 | year = 2011 | pmid =  21627792| pmc =3219973 | doi-access = free }}</ref> A 2015 study suggested that the animals in the range where ''M. columbi'' and ''M. primigenius'' overlapped formed a [[metapopulation]] of hybrids with varying morphology. It suggested that Eurasian ''M. primigenius'' had a similar relationship with ''M. trogontherii'' in areas where their range overlapped.<ref name="Lister2015">{{cite journal |last1=Lister |first1=A. M. |last2=Sher |first2=A. V. |date=13 November 2015 |title=Evolution and dispersal of mammoths across the Northern Hemisphere |journal=[[Science (journal)|Science]] |volume=350 |issue=6262 |pages=805–809 |bibcode=2015Sci...350..805L |doi=10.1126/science.aac5660 |pmid=26564853 |s2cid=206639522|doi-access=free }}</ref> According to a 2025 study of mammoth teeth from [[Alberta, Canada]], this area was a zone of [[sympatry]] and hybridisation between woolly and Columbian mammoths.<ref>{{Cite journal |last1=Barrón-Ortiz |first1=Christina I. |last2=Jass |first2=Christopher N. |last3=Cammidge |first3=Tasha S. |date=6 January 2025 |title=Taxonomic, biogeographic, and biological implications of mammoth teeth from a dynamic Pleistocene landscape in Alberta, Canada |url=https://www.cambridge.org/core/journals/quaternary-research/article/taxonomic-biogeographic-and-biological-implications-of-mammoth-teeth-from-a-dynamic-pleistocene-landscape-in-alberta-canada/3D4DA534670DDD88615C1CD0B8562A80 |journal=[[Quaternary Research]] |language=en |volume=123 |pages=41–58 |doi=10.1017/qua.2024.47 |bibcode=2025QuRes.123...41B |issn=0033-5894 |access-date=12 March 2025 |via=Cambridge Core|url-access=subscription }}</ref>

In 2021, DNA older than a million years was sequenced for the first time, from two mammoth teeth of [[Early Pleistocene]] age found in eastern Siberia. One tooth from [[Adycha]] (1–1.3 million years old) belonged to a lineage that was ancestral to later woolly mammoths, whereas the other from [[Krestovka]] (1.1–1.65 million years old) belonged to new lineage. The study found that half of the ancestry of Columbian mammoths came from relatives of the Krestovka lineage (which probably represented the first mammoths that colonised the Americas) and the other half from the lineage of woolly mammoths, with the hybridisation happening more than 420,000 years ago, during the [[Middle Pleistocene]]. Later woolly and Columbian mammoths also interbred occasionally, and mammoth species may have hybridised routinely when brought together by glacial expansion. These findings were the first evidence of [[hybrid speciation]] from ancient DNA. The study also found that genetic adaptations to cold environments, such as hair growth and fat deposits, were already present in the steppe mammoth lineage and were not unique to woolly mammoths.<ref name="OldestDNA">{{cite journal |last1=van der Valk |first1=T. |last2=Pečnerová |first2=P. |last3=Díez-del-Molino |first3=D. |last4=Bergström |first4=A. |last5=Oppenheimer |first5=J. |last6=Hartmann |first6=S. |last7=Xenikoudakis |first7=G. |last8=Thomas |first8=J. A. |last9=Dehasque |first9=M. |last10=Sağlıcan |first10=E. |last11=Fidan |first11=F. Rabia |last12=Barnes |first12=I. |last13=Liu |first13=S. |last14=Somel |first14=M. |last15=Heintzman |first15=P. D. |last16=Nikolskiy |first16=P. |last17=Shapiro |first17=B. |last18=Skoglund |first18=P. |last19=Hofreiter |first19=M. |last20=Lister |first20=A. M. |last21=Götherström |first21=A. |last22=Dalén |first22=L. |title=Million-year-old DNA sheds light on the genomic history of mammoths |journal=Nature |date=2021 |volume=591 |issue=7849 |pages=265–269 |doi=10.1038/s41586-021-03224-9 |pmid=33597750 |pmc=7116897 |bibcode=2021Natur.591..265V |language=en |issn=1476-4687}}</ref><ref>{{cite news |last1=Callaway |first1=E. |title=Million-year-old mammoth genomes shatter record for oldest ancient DNA |work=nature.com |date=2021 |volume=590 |issue=7847 |pages=537–538 |doi=10.1038/d41586-021-00436-x |url=https://www.nature.com/articles/d41586-021-00436-x |access-date=17 February 2021}}</ref>

==Description==
{{multiple image
| align             = right
| total_width       = 425
| image1            = Mammuthus Size comparison.png
| alt1              = Colored silhouette of a woolly mammoth (blue), relative in size to a human and past and present elephants
| image2            = M._primigenius_modified.png
| alt2              = Skeletal diagram of 3.7 metre (12.1 ft) tall Columbian mammoth compared to a human
| footer            = Size of a large woolly mammoth (in red) compared to a human and other mammoths (left) and a skeletal diagram of a {{cvt|3.5|m}} tall large European bull and a smaller {{cvt|2.7|m}} Siberian bull compared to a human (right)
}}

The appearance of the woolly mammoth is probably the best known of any prehistoric animal due to the many frozen specimens with preserved soft tissue and depictions by contemporary humans in their art. The average shoulder height for males of the species has been estimated at {{cvt|2.8|-|3.15|m}} with a weight of {{cvt|4.5|-|6|t|lb}}, with females being smaller like living elephants, with a shoulder height of {{cvt|2.3|-|2.6|m}} and a weight of {{cvt|2.8|-|4|t|lb}}.<ref name="Larramendi-2017">{{Cite journal |last1=Larramendi |first1=Asier |last2=Palombo |first2=Maria Rita |last3=Marano |first3=Federica |date=2017 |title=Reconstructing the life appearance of a Pleistocene giant: size, shape, sexual dimorphism and ontogeny of Palaeoloxodon antiquus (Proboscidea: Elephantidae) from Neumark-Nord 1 (Germany) |url=https://www.paleoitalia.it/wp-content/uploads/2021/08/01_Larramendi_et_al_2017_BSPI_563.pdf |journal=Bollettino della Società Paleontologica Italiana |issue=3 |pages=299–317 |doi=10.4435/BSPI.2017.29 |doi-broken-date=20 November 2024 |issn=0375-7633|archive-url=https://web.archive.org/web/20230930183334/https://www.paleoitalia.it/wp-content/uploads/2021/08/01_Larramendi_et_al_2017_BSPI_563.pdf |archive-date=30 September 2023 }}</ref> This size is comparable to the largest living elephant species, the [[African bush elephant]] (''Loxodonta africana''), but is considerably smaller than the earlier ''Mammuthus meridionalis'' and ''Mammuthus trogontherii'' and the contemporary ''Mammuthus columbi''.<ref name="Larramendi-2017" /><ref name="probos_mass">{{cite journal |last1=Larramendi |first1=Asier |date=2015 |title=Proboscideans: Shoulder Height, Body Mass and Shape |journal=Acta Palaeontologica Polonica |volume=61 |doi=10.4202/app.00136.2014 |doi-access=free}}</ref> The woolly mammoth exhibited size variation throughout its range, with individuals from Western Europe being considerably larger (with adult males estimated to be on average {{cvt|2.99|-|3.31|m}} tall and {{cvt|5.2|–|6.9|t|lb}} in weight) than those found in Siberia (with adult males of this population being estimated on average {{cvt|2.66|-|2.94|m}} tall and {{cvt|3.9|-|5.2|t|lb}} in weight). One of the largest recorded woolly mammoths is the Siegsdorf specimen from Germany, with an estimated shoulder height of {{cvt|3.49|m}} and an estimated body mass of {{cvt|8.2|t|lb}}.<ref name="probos_mass" /> A newborn calf would have weighed about {{cvt|90|kg}}.<ref>Lister, 2007. pp. 82–87</ref>

[[File:Woolly mammoth size comparison.svg|thumb|left|Size comparison of average-sized male and female woolly mammoths]]
Few frozen specimens have preserved genitals, so the sex is usually determined through examination of the skeleton. The best indication of sex is the size of the [[pelvic]] girdle, since the opening that functions as the birth canal is always wider in females than in males.<ref>Lister, 2007. pp.&nbsp;174–175</ref> Though the mammoths on Wrangel Island were smaller than those of the mainland, their size varied, and they were not small enough to be considered "[[Insular dwarfism|island dwarfs]]".<ref>{{Cite journal | last1 = Vartanyan | first1 = S. L. | last2 = Arslanov | first2 = K. A. | last3 = Karhu | first3 = J. A. | last4 = Possnert | first4 = G. R. | last5 = Sulerzhitsky | first5 = L. D. | title = Collection of radiocarbon dates on the mammoths (''Mammuthus primigenius'') and other genera of Wrangel Island, northeast Siberia, Russia | doi = 10.1016/j.yqres.2008.03.005 | journal = Quaternary Research | volume = 70 | issue = 1 | pages = 51–59 | year = 2008 | bibcode = 2008QuRes..70...51V | s2cid = 111383180 | url = http://doc.rero.ch/record/16074/files/PAL_E3865.pdf }}</ref> The last woolly mammoth populations are claimed to have decreased in size and increased their [[sexual dimorphism]], but this was dismissed in a 2012 study.<ref>{{Cite journal | last1 = Den Ouden | first1 = N. | last2 = Reumer | first2 = J. W. F. | last3 = Van Den Hoek Ostende | first3 = L. W. | title = Did mammoth end up a lilliput? Temporal body size trends in Late Pleistocene Mammoths, ''Mammuthus primigenius'' (Blumenbach, 1799) inferred from dental data | doi = 10.1016/j.quaint.2011.07.038 | journal = Quaternary International | volume = 255 | pages = 53–58 | year = 2012 |bibcode = 2012QuInt.255...53D }}</ref>

[[File:Woolly mammoth model Royal BC Museum in Victoria.jpg|thumb|Model at the [[Royal BC Museum]]]]
Woolly mammoths had several adaptations to the cold, most noticeably the layer of fur covering all parts of their  bodies. Other adaptations to cold weather include ears that are far smaller than those of modern elephants; they were about {{cvt|38|cm}} long and {{cvt|18|-|28|cm}} across, and the ear of the 6- to 12-month-old frozen calf "Dima" was under {{cvt|13|cm}} long. The small ears reduced heat loss and [[frostbite]], and the tail was short for the same reason, only {{cvt|36|cm}} long in the "Berezovka mammoth". The tail contained 21 vertebrae, whereas the tails of modern elephants contain 28–33. Their skin was no thicker than that of present-day elephants, between {{cvt|1.25|and|2.5|cm}}. They had a layer of fat up to {{cvt|10|cm}} thick under the skin, which helped to keep them warm. Woolly mammoths had broad flaps of skin under their tails which covered the [[anus]]; this is also seen in modern elephants.<ref name="Lister 2007 82 87">Lister, 2007. pp.&nbsp;82–87</ref>

Other characteristic features depicted in [[cave painting]]s include a large, high, single-domed head and a sloping back with a high shoulder hump; this shape resulted from the [[spinous process]]es of the back vertebrae decreasing in length from front to rear. These features were not present in juveniles, which had convex backs like Asian elephants. Another feature shown in cave paintings was confirmed by the discovery of a frozen specimen in 1924, an adult nicknamed the "Middle Kolyma mammoth", which was preserved with a complete trunk tip. Unlike the trunk lobes of modern elephants, the upper "finger" at the tip of the trunk had a long pointed lobe and was {{cvt|10|cm}} long, while the lower "thumb" was {{cvt|5|cm}} and was broader. The trunk of "Dima" was {{cvt|76|cm}} long, whereas the trunk of the adult "Liakhov mammoth" was {{cvt|2|m}} long.<ref name="Lister 2007 82 87"/> The well-preserved trunk of a juvenile specimen nicknamed "[[Yuka (mammoth)|Yuka]]" was described in 2015, and it was shown to possess a fleshy expansion a third above the tip. Rather than oval as the rest of the trunk, this part was ellipsoidal in cross section, and double the size in diameter. The feature was shown to be present in two other specimens, of different sexes and ages.<ref name="Yuka trunk">{{Cite journal | doi = 10.1134/S0031030115020070| title = New data on trunk morphology in the woolly mammoth, ''Mammuthus primigenius'' (Blumenbach)| journal = Paleontological Journal| volume = 49| issue = 2| pages = 200–210| year = 2015| last1 = Plotnikov | first1 = V. V.| last2 = Maschenko | first2 = E. N.| last3 = Pavlov | first3 = I. S.| last4 = Protopopov | first4 = A. V.| last5 = Boeskorov | first5 = G. G.| last6 = Petrova | first6 = E. A.| bibcode = 2015PalJ...49..200P| s2cid = 84849714}}</ref>

===Coat===

{{multiple image
|align = right
|total_width = 350
|image1 = Mammoth wool.jpg
|alt1 = 
|image2 = Woolly Mammoth Overhair (x250).tif
|alt2 = 
|footer = Fur in [[Naturhistorisches Museum, Vienna]] (left), and [[Scanning electron microscope|SEM]] magnified image of an overhair
}}

The [[Coat (animal)|coat]] consisted of an outer layer of long, coarse "guard hair", which was {{cvt|30|cm}} on the upper part of the body, up to {{cvt|90|cm}} in length on the flanks and underside, and {{cvt|0.5|mm}} in diameter, and a denser inner layer of shorter, slightly curly under-wool, up to {{cvt|8|cm}} long and {{cvt|0.05|mm}} in diameter. The hairs on the upper leg were up to {{cvt|38|cm}} long, and those of the feet were {{cvt|15|cm}} long, reaching the toes. The hairs on the head were relatively short, but longer on the underside and the sides of the trunk. The tail was extended by coarse hairs up to {{cvt|60|cm}} long, which were thicker than the guard hairs. The woolly mammoth likely [[moulting|moulted]] seasonally, and the heaviest fur was shed during spring.<ref name="Lister, 2007. pp. 83–84">Lister, 2007. pp.&nbsp;83–84</ref>

Since mammoth carcasses were more likely to be preserved, possibly only the winter coat has been preserved in frozen specimens. Modern elephants have much less hair, though juveniles have a more extensive covering of hair than adults.<ref name="Lister, 2007. pp. 83–84"/> This is thought to be for [[thermoregulation]], helping them lose heat in their hot environments.<ref name="Elephanthair">{{cite journal |last1=Myhrvold |first1=C. L. |last2=Stone |first2=H. A. |last3=Bou-Zeid |first3=E. |title=What is the use of elephant hair? |journal=PLOS ONE |date=10 October 2012 |volume=7 |issue=10 |pages=e47018 |doi=10.1371/journal.pone.0047018|pmid=23071700 |pmc=3468452 |bibcode=2012PLoSO...747018M |doi-access=free }}</ref> Comparison between the over-hairs of woolly mammoths and extant elephants show that they did not differ much in overall morphology.<ref>{{Cite journal | last1 = Valente | first1 = A. | title = Hair structure of the woolly mammoth, ''Mammuthus primigenius'' and the modern elephants, ''Elephas maximus'' and ''Loxodonta africana'' | doi = 10.1111/j.1469-7998.1983.tb02095.x | journal = Journal of Zoology | volume = 199 | issue = 2 | pages = 271–274 | year = 1983}}</ref> Woolly mammoths had numerous [[sebaceous gland]]s in their skin, which secreted oils into their hair; this would have improved the wool's insulation, repelled water, and given the fur a glossy sheen.<ref>{{Cite journal | last1 = Repin | first1 = V. E. | last2 = Taranov | first2 = O. S. | last3 = Ryabchikova | first3 = E. I. | last4 = Tikhonov | first4 = A. N. | last5 = Pugachev | first5 = V. G. | title = Sebaceous Glands of the Woolly Mammoth, ''Mammothus primigenius'' Blum.: Histological Evidence | doi = 10.1023/B:DOBS.0000046662.43270.66 | journal = Doklady Biological Sciences | volume = 398 | issue = 1–6 | pages = 382–384 | year = 2004 | pmid =  15587793| s2cid = 6401669 }}</ref>

Preserved woolly mammoth fur is orange-brown, but this is believed to be an artefact from the bleaching of pigment during burial. The amount of [[pigment]]ation varied from hair to hair and within each hair.<ref name="Lister 2007 82 87" /> A 2006 study sequenced the [[Melanocortin 1 receptor|''Mc1r'' gene]] (which influences hair colour in mammals) from woolly mammoth bones. Two [[allele]]s were found: a [[dominance (genetics)|dominant]] (fully active) and a [[recessive trait|recessive]] (partially active) one. In mammals, recessive'' Mc1r'' alleles result in light hair. Mammoths born with at least one copy of the dominant allele would have had dark coats, while those with two copies of the recessive allele would have had light coats.<ref name="Römpler">{{Cite journal | last1 = Rompler | first1 = H. | last2 = Rohland | first2 = N. | last3 = Lalueza-Fox | first3 = C. | last4 = Willerslev | first4 = E. | last5 = Kuznetsova | first5 = T. | last6 = Rabeder | first6 = G. | last7 = Bertranpetit | first7 = J. | last8 = Schöneberg | first8 = T. | last9 = Hofreiter | first9 = M. | title = Nuclear Gene Indicates Coat-Color Polymorphism in Mammoths | doi = 10.1126/science.1128994 | journal = Science | volume = 313 | issue = 5783 | pages = 62 | year = 2006 | pmid =  16825562| s2cid = 20153467 | url = http://oceanrep.geomar.de/28050/1/2006_Roempler-etal-Gne_Science-313-5783.pdf }}</ref> A 2011 study showed that light individuals would have been rare.<ref>{{Cite journal |last1=Workman |first1=C. |last2=Dalen |first2=L. |last3=Vartanyan |first3=S. |last4=Shapiro |first4=B. |last5=Kosintsev |first5=P. |last6=Sher |first6=A. |last7=Gotherstrom |first7=A. |last8=Barnes |first8=I. |year=2011 |title=Population-level genotyping of coat colour polymorphism in woolly mammoth (''Mammuthus primigenius'') |journal=[[Quaternary Science Reviews]] |volume=30 |issue=17–18 |pages=2304–2308 |bibcode=2011QSRv...30.2304W |doi=10.1016/j.quascirev.2010.08.020}}</ref> A 2014 study instead indicated that the colouration of an individual varied from nonpigmented on the overhairs, bicoloured, nonpigmented and mixed red-brown guard hairs, and nonpigmented underhairs, which would give a light overall appearance.<ref>{{cite journal|last1=Tridico|first1=Silvana R.|last2=Rigby|first2=Paul|last3=Kirkbride|first3=K. Paul|last4=Haile|first4=James|last5=Bunce|first5=Michael|title=Megafaunal split ends: microscopical characterisation of hair structure and function in extinct woolly mammoth and woolly rhino|url=https://www.researchgate.net/publication/259123707|journal=Quaternary Science Reviews|volume=83|year=2014|pages=68–75|doi=10.1016/j.quascirev.2013.10.032|bibcode=2014QSRv...83...68T|url-status=live|archive-url=https://web.archive.org/web/20171102052328/https://www.researchgate.net/publication/259123707_Megafaunal_split_ends_microscopical_characterisation_of_hair_structure_and_function_in_extinct_woolly_mammoth_and_woolly_rhino|archive-date=2 November 2017}}</ref>

===Dentition===
[[File:Woolly mammoth with downward spiralling tusk.jpg|left|thumb|Skull from Poland with one broken and one downward spiralled tusk]] 
Woolly mammoths had very long tusks (modified [[incisor]] teeth), which were more curved than those of modern elephants. The longest known male tusk is {{cvt|4.05|m}} long (measured along the outside curve) and weighs {{cvt|115.5|kg}}, with a historical report of a {{cvt|4.30|m}} long tusk found in Siberia, while the heaviest tusk is {{cvt|121|kg}}, suggested to have been {{cvt|125|-|130|kg}} when complete;<ref name="Tuskers" /><ref name="Lister, 2007. pp. 94–95"/> {{cvt|2.4|-|2.7|m}} and {{cvt|45|kg}} was a more typical size. Female tusks were smaller and thinner, {{cvt|1.5|-|1.8|m}} and weighing {{cvt|9|kg}}. For comparison, the record for longest tusks of the [[African bush elephant]] is {{cvt|3.4|m}}. The sheaths of the tusks were parallel and spaced closely. About a quarter of the length was inside the sockets. The tusks grew spirally in opposite directions from the base and continued in a curve until the tips pointed towards each other, sometimes crossing. In this way, most of the weight would have been close to the skull, and less [[torque]] would occur than with straight tusks.<ref name="Lister, 2007. pp. 94–95">Lister, 2007. pp.&nbsp;94–95</ref><ref name="Kurten & Anderson"/><ref name="Tuskers">{{cite journal |last1=Boeskorov |first1=G. |last2=Tikhonov |first2=A. |last3=Shchelchkova |first3=M. |last4=Ballard |first4=J. P. |last5=Mol |first5=D. |date=2020 |title=Big tuskers: Maximum sizes of tusks in woolly mammoths - Mammuthus primigenius (Blumenbach) - from East Siberia |journal=[[Quaternary International]] |volume=537 |pages=88–96 |bibcode=2020QuInt.537...88B |doi=10.1016/j.quaint.2019.12.023 |s2cid=213262363}}</ref>

The tusks were usually asymmetrical and showed considerable variation, with some tusks curving down instead of outwards and some being shorter due to breakage. Calves developed small [[milk tusks]] a few centimetres long at six months old, which were replaced by permanent tusks a year later. Tusk growth continued throughout life, but became slower as the animal reached adulthood. The tusks grew by {{cvt|2.5|-|15|cm}} each year. Some cave paintings show woolly mammoths with small or no tusks, but whether this reflected reality or was artistic license is unknown. Female Asian elephants have no tusks, but no fossil evidence indicates that any adult woolly mammoths lacked them.<ref name="Lister, 2007. pp. 94–95"/><ref name="Kurten & Anderson">{{cite book | last1 = Kurten | first1 = B. | last2 = Anderson | first2 = E. |year= 1980 | pages = 348–354|title= Pleistocene Mammals of North America |publisher=Columbia University Press |isbn= 978-0-231-03733-4 |location=New York}}</ref>

[[File:Molaire de mammouth - font de Champdamoy.jpg|thumb|Molar from Font de Champdamoy, France, [[Musée Georges-Garret]]]]
Woolly mammoths had four functional [[molar (tooth)|molar]] teeth at a time—two in the upper jaw and two in the lower. About {{cvt|23|cm}} of the crown was within the jaw, and {{cvt|2.5|cm}} was above. The crown was continually pushed forwards and up as it wore down, comparable to a [[conveyor belt]]. The teeth had up to 26 separated ridges of [[Tooth enamel|enamel]], which were themselves covered in "prisms" that were directed towards the chewing surface. These were quite wear-resistant and kept together by [[cementum]] and [[dentine]]. A mammoth had six sets of molars throughout a lifetime, which were replaced five times, though a few specimens with a seventh set are known. The latter condition could extend the lifespan of the individual, unless the tooth consisted of only a few plates. The first molars were about the size of those of a human, {{cvt|1.3|cm}}, the third were {{cvt|15|cm}} long, and the sixth were about {{cvt|30|cm}} long and weighed {{cvt|1.8|kg}}. The molars grew larger and contained more ridges with each replacement.<ref>Lister, 2007. pp. 92–93</ref> The woolly mammoth is considered to have had the most complex molars of any elephant.<ref name="Kurten & Anderson"/>

==Palaeobiology==

[[File:Ice age fauna of northern Spain - Mauricio Antón.jpg|thumb|left|[[Life restoration]] of fauna during the Pleistocene epoch in northern Spain, by [[Mauricio Antón]], 2004]]
Adult woolly mammoths could effectively defend themselves from predators with their tusks, trunks, and size; however, juveniles and weakened adults were vulnerable to pack hunters such as [[wolves]], [[cave hyena]]s, and large [[felidae|felines]]. The tusks may have been used in intraspecies fighting, such as fights over [[Territory (animal)|territory]] or mates. Display of the large tusks of males could have been used to attract females and intimidate rivals. Because of their curvature, the tusks were unsuitable for stabbing but may have been used for hitting, as indicated by injuries to some fossil shoulder blades.

The very long hairs on the tail probably compensated for the shortness of the tail, enabling its use as a [[flyswatter]], similar to the tail on modern elephants. As in modern elephants, the sensitive and muscular trunk worked as a limb-like organ with many functions. It was used for manipulating objects and in social interactions.<ref>Lister, 2007. pp.&nbsp;95–105</ref> The well-preserved foot of the adult male "[[Yukagir mammoth]]" shows that the soles of the feet contained many cracks that would have helped in gripping surfaces during locomotion. Like modern elephants, woolly mammoths walked on their toes and had large, fleshy pads behind the toes.<ref name="Lister 2007 82 87" />

Like modern elephants, woolly mammoths were likely very social and lived in [[matriarchal]] (female-led) family groups. This is supported by fossil assemblages and cave paintings showing groups, implying that most of their other social behaviours were likely similar to those of modern elephants. How many mammoths lived at one location at a time is unknown, as fossil deposits are often accumulations of individuals who died over long periods of time. The numbers likely varied by season and lifecycle events. Modern elephants can form large herds, sometimes consisting of multiple family groups, which can include thousands of animals migrating together. Mammoths may have formed large herds more often since animals that live in open areas are more likely to do this than those in forested areas.<ref>Lister, 2007. pp.&nbsp;62–63</ref> Trackways made by a woolly mammoth herd 11,300–11,000 years ago have been found in the [[St. Mary Reservoir]] in Canada, showing that in this case, almost equal numbers of adults, subadults, and juveniles were found. The adults had a stride of {{convert|2|m|ft|abbr=on}}, and the juveniles ran to keep up.<ref name="Trackway"/>

Woolly mammoth dental enamel from Poland has demonstrated that woolly mammoths were seasonally migratory. Recurring shifts in [[Δ18O|δ<sup>18</sup>O]] and <sup>87</sup>Sr/<sup>86</sup>Sr found in layers of the enamel correspond to seasonal variations and indicate that Polish woolly mammoths inhabited southern Poland during winter but grazed the Polish midlands during summer.<ref>{{Cite journal |last1=Kowalik |first1=Nina |last2=Anczkiewicz |first2=Robert |last3=Müller |first3=Wolfgang |last4=Spötl |first4=Christoph |last5=Bondioli |first5=Luca |last6=Nava |first6=Alessia |last7=Wojtal |first7=Piotr |last8=Wilczyński |first8=Jarosław |last9=Koziarska |first9=Marta |last10=Matyszczak |first10=Milena |date=15 April 2023 |title=Revealing seasonal woolly mammoth migration with spatially-resolved trace element, Sr and O isotopic records of molar enamel |url=https://linkinghub.elsevier.com/retrieve/pii/S0277379123000847 |journal=[[Quaternary Science Reviews]] |language=en |volume=306 |pages=108036 |doi=10.1016/j.quascirev.2023.108036 |bibcode=2023QSRv..30608036K |access-date=3 May 2024 |via=Elsevier Science Direct|url-access=subscription }}</ref>

===Adaptations to cold===

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|footer = Head and leg of the adult male "Yukagir mammoth" (the trunk is not preserved); note fur and small ears
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The woolly mammoth was probably the most specialised member of the family Elephantidae. In addition to their fur, they had lipoplexes (fat storage) in their neck and [[withers]] for times when food availability was insufficient during winter, and their first three molars grew more quickly than in the calves of modern elephants. The expansion identified on the trunk of "Yuka" and other specimens was suggested to function as a "fur mitten"; the trunk tip was not covered in fur but was used for foraging during winter and could have been heated by curling it into the expansion. The expansion could be used to melt snow if a shortage of water to drink existed, as melting it directly inside the mouth could disturb the thermal balance of the animal.<ref name="Yuka trunk"/><ref name="YukaHistory"/> As in [[reindeer]] and [[musk oxen]], the [[haemoglobin]] of the woolly mammoth was adapted to the cold, with three mutations to improve oxygen delivery around the body and prevent freezing. This feature may have helped the mammoths to live at high latitudes.<ref>{{Cite journal | last1 = Campbell | first1 = K. L. | last2 = Roberts | first2 = J. E. E. | last3 = Watson | first3 = L. N. | last4 = Stetefeld | first4 = J. R. | last5 = Sloan | first5 = A. M. | last6 = Signore | first6 = A. V. | last7 = Howatt | first7 = J. W. | last8 = Tame | first8 = J. R. H. | last9 = Rohland | first9 = N. | doi = 10.1038/ng.574 | last10 = Shen | first10 = T. J. | last11 = Austin | first11 = J. J. | last12 = Hofreiter | first12 = M. | last13 = Ho | first13 = C. | last14 = Weber | first14 = R. E. | last15 = Cooper | first15 = A. | title = Substitutions in woolly mammoth hemoglobin confer biochemical properties adaptive for cold tolerance | journal = Nature Genetics | volume = 42 | issue = 6 | pages = 536–540 | year = 2010 | pmid =  20436470| s2cid = 9670466 }}</ref>

In a 2015 study, high-quality [[genome]] sequences from three Asian elephants and two woolly mammoths were compared. About 1.4 million DNA nucleotide differences were found between mammoths and elephants, which affect the sequence of more than 1,600 proteins. Differences were noted in genes for a number of aspects of physiology and biology that would be relevant to Arctic survival, including the development of skin and hair, storage and metabolism of adipose tissue, and perceiving temperature. Genes related to both sensing temperature and transmitting that sensation to the brain were altered. One of the heat-sensing genes encodes a protein, TRPV3, found in the skin, which affects hair growth. When inserted into human cells, the mammoth's version of the protein was found to be less sensitive to heat than the elephant's. This is consistent with a previous observation that mice lacking active TRPV3 are likely to spend more time in cooler cage locations than wild-type mice and have wavier hair. Several alterations in circadian clock genes were found, perhaps needed to cope with the extreme polar variation in the length of daylight. Similar mutations are known in other Arctic mammals, such as reindeer.<ref name="Lynch2015">{{Cite journal | doi = 10.1016/j.celrep.2015.06.027| pmid = 26146078| title = Elephantid genomes reveal the molecular bases of Woolly Mammoth adaptations to the arctic| year = 2015| last1 = Lynch | first1 = V. | last2 = Bedoya-Reina | first2 = O. C. | last3 = Ratan | first3 = A. | last4 = Sulak | first4 = M. | last5 = Drautz-Moses | first5 = D. I. | last6 = Perry | first6 = G. H. | last7 = Miller | first7 = W. | last8 = Schuster | first8 = S. C.|journal=Cell Reports|volume=12|issue=2|pages=217–228| doi-access = free | hdl = 10220/38768 | hdl-access = free }}</ref><ref name="Callaway2015">{{cite web|url=http://www.scientificamerican.com/article/mammoth-genomes-provide-recipe-for-creating-arctic-elephants/|title=Mammoth Genomes Provide Recipe for Creating Arctic Elephants|date=4 May 2015|author=Ewen Callaway, Nature magazine|website=scientificamerican.com|url-status=live|archive-url=https://web.archive.org/web/20150505135155/http://www.scientificamerican.com/article/mammoth-genomes-provide-recipe-for-creating-arctic-elephants/|archive-date=5 May 2015}}</ref>

A 2019 study of the woolly mammoth [[mitogenome]] suggests that these had metabolic adaptations related to extreme environments.<ref name="Signals">{{cite journal |last1=Ngatia |first1=J. N. |last2=Lan |first2=T. M. |last3=Dinh |first3=T. D. |last4=Zhang |first4=L. |last5=Ahmed |first5=Ahmed Khalid |last6=Xu |first6=Yan Chun |title=Signals of positive selection in mitochondrial protein-coding genes of woolly mammoth: Adaptation to extreme environments? |journal=Ecology and Evolution |date=2019 |volume=9 |issue=12 |pages=6821–6832 |doi=10.1002/ece3.5250|pmid=31380018 |pmc=6662336 |doi-access=free |bibcode=2019EcoEv...9.6821N }}</ref> A genetic study from 2023 found that the woolly mammoth had already acquired a broad range of genes associated with the development of skin and hair, fat storage, metabolism, and the immune system by the time the species appeared and that these continued to evolve within the last 700,000 years, including a gene that resulted in mammoths of the Late Quaternary having small ears.<ref name="Genomics">{{cite journal |last1=Díez-del-Molino |first1=David |last2=Dehasque |first2=Marianne |last3=Chacón-Duque |first3=J. Camilo |last4=Pečnerová |first4=Patrícia |last5=Tikhonov |first5=Alexei |last6=Protopopov |first6=Albert |last7=Plotnikov |first7=Valeri |last8=Kanellidou |first8=Foteini |last9=Nikolskiy |first9=Pavel |last10=Mortensen |first10=Peter |last11=Danilov |first11=Gleb K. |last12=Vartanyan |first12=Sergey |last13=Gilbert |first13=M. Thomas P. |last14=Lister |first14=Adrian M. |last15=Heintzman |first15=Peter D. |last16=van der Valk |first16=Tom |last17=Dalén |first17=Love |title=Genomics of adaptive evolution in the woolly mammoth |journal=Current Biology |date=2023 |volume=33 |issue=9 |pages=1753–1764.e4 |doi=10.1016/j.cub.2023.03.084|pmid=37030294 |doi-access=free |hdl=11250/3145739 |hdl-access=free }}</ref>

===Diet===
[[File:Woolly mammoth jaw.jpg|thumb|left|upright|Mandibles and lower molars, Barcelona]]
Food at various stages of digestion has been found in the intestines of several woolly mammoths, giving a good picture of their diet. Woolly mammoths sustained themselves on plant food, mainly grasses and sedges, which were supplemented with [[herbaceous plant]]s, [[flowering plant]]s, [[shrub]]s, [[moss]]es, and tree matter. The composition and exact varieties differed from location to location. Woolly mammoths needed a varied diet to support their growth, similar to modern elephants. An adult of 6 tonnes would need to eat {{cvt|180|kg}} daily and may have [[foraged]] as long as 20 hours every day. The two-fingered tip of the trunk was probably adapted for picking up the short grasses of the last [[ice age]] ([[Quaternary glaciation]], 2.58 million years ago to present) by wrapping around them. In contrast, modern elephants curl their trunks around the longer grass of their tropical environments. The trunk could be used for pulling off large grass tufts, delicately picking [[bud]]s and flowers, and tearing off leaves and branches where trees and shrubs were present. The "Yukagir mammoth" had ingested plant matter that contained spores of [[coprophilous fungi|dung fungus]].<ref>Lister, 2007. pp.&nbsp;88–91</ref> [[Isotope analysis]] shows that woolly mammoths fed mainly on [[C3 plants]], unlike horses and rhinos.<ref>{{Cite journal | last1 = Bocherens | first1 = H. | last2 = Fizet | first2 = M. | last3 = Mariotti | first3 = A. | last4 = Gangloff | first4 = R. A. | last5 = Burns | first5 = J. A. | title = Contribution of isotopic biogeochemistry (13C,15N,18O) to the paleoecology of mammoths (''Mammuthus primigenius'') | doi = 10.1080/10292389409380453 | journal = Historical Biology | volume = 7 | issue = 3 | pages = 187–202 | year = 1994 | bibcode = 1994HBio....7..187B }}</ref>

Scientists identified milk in the stomach and [[faecal matter]] in the intestines of the mammoth calf "[[Lyuba (mammoth)|Lyuba]]".<ref name="van Geel et al., 2011">{{Cite journal | last1 = Van Geel | first1 = B. | last2 = Fisher | first2 = D. C. | last3 = Rountrey | first3 = A. N. | last4 = Van Arkel | first4 = J. | last5 = Duivenvoorden | first5 = J. F. | last6 = Nieman | first6 = A. M. | last7 = Van Reenen | first7 = G. B. A. | last8 = Tikhonov | first8 = A. N. | last9 = Buigues | first9 = B. | doi = 10.1016/j.quascirev.2011.10.009 | last10 = Gravendeel | first10 = B. | title = Palaeo-environmental and dietary analysis of intestinal contents of a mammoth calf (Yamal Peninsula, northwest Siberia) | journal = Quaternary Science Reviews | volume = 30 | issue = 27–28 | pages = 3935–3946 | year = 2011 |bibcode = 2011QSRv...30.3935V }}</ref> The faecal matter may have been eaten by "Lyuba" to promote development of the [[gut flora|intestinal microbes]] necessary for digestion of vegetation, as is the case in modern elephants.<ref name="Fisher et al., 2012" /> An isotope analysis of woolly mammoths from Yukon showed that the young nursed for at least 3 years and were [[weaned]] and gradually changed to a diet of plants when they were 2–3 years old. This is later than in modern elephants and may be due to a higher risk of predator attack or difficulty obtaining food during the long winter darkness at high latitudes.<ref>{{Cite journal | last1 = Metcalfe | first1 = J. Z. | last2 = Longstaffe | first2 = F. J. | last3 = Zazula | first3 = G. D. | doi = 10.1016/j.palaeo.2010.09.032 | title = Nursing, weaning, and tooth development in woolly mammoths from Old Crow, Yukon, Canada: Implications for Pleistocene extinctions | journal = Palaeogeography, Palaeoclimatology, Palaeoecology | volume = 298 | issue = 3–4 | pages = 257–270 | year = 2010 | bibcode = 2010PPP...298..257M }}</ref>

[[File:Worn Woolly mammoth tusk.png|thumb|Male tusk with signs of wear]] 
The molars were adapted to their diet of coarse tundra grasses, with more enamel plates and a higher crown than their earlier southern relatives. The woolly mammoth chewed its food using its powerful jaw muscles to move the mandible forwards and close the mouth, then backward while opening; the sharp enamel ridges cut across each other, grinding the food. The ridges were wear-resistant, enabling the animal to chew large quantities of food, often containing grit. Woolly mammoths may have used their tusks as shovels to clear snow from the ground, reach the vegetation buried below, and break ice to drink. This is indicated on many preserved tusks by flat, polished sections up to {{cvt|30|cm}} long, as well as scratches on the part of the surface that would have reached the ground (especially at their outer curvature). The tusks were used for obtaining food in other ways, such as digging up plants and stripping off bark.<ref>Lister, 2007. pp.&nbsp;92–95</ref><ref name="Projectiles"/>

===Life history===

[[File:Cross sections of African elephant and woolly mammoth tusks.png|thumb|left|Cross sections of African elephant and woolly mammoth tusks; [[growth rings]] can be used to determine age]] 
The lifespan of mammals is related to their size. Since modern elephants can reach the age of 60 years, the same is thought to be true for woolly mammoths, which were of a similar size. The age of a mammoth can be roughly determined by counting the [[growth rings]] of its tusks when viewed in cross-section, but this does not account for its early years, as these are represented by the tips of the tusks, which are usually worn away. In the remaining part of the tusk, each major line represents a year, and weekly and daily ones can be found in between. Dark bands correspond to summers, so determining the season in which a mammoth died is possible. The growth of the tusks slowed when foraging became harder, for example, during winter, during disease, or when a male was banished from the herd (male elephants live with their herds until about the age of 10). Mammoth tusks dating to the harshest period of the last glaciation 25–20,000 years ago show slower growth rates.<ref name="Lister, 2007. pp. 83–107">Lister, 2007. pp.&nbsp;83–107.</ref><ref name="Herbert2010">{{cite web | url = https://helix.northwestern.edu/article/mammoth-find-clues-past-present-and-future | title = A Mammoth Find: Clues to the Past, Present and Future | last1 = Herbert | first1 = B. | last2 = Fisher | first2 = D. | date = 5 May 2010 | website = Helix | publisher = [[Northwestern University]] | access-date = 1 August 2016 | url-status = live | archive-url = https://web.archive.org/web/20160818074348/https://helix.northwestern.edu/article/mammoth-find-clues-past-present-and-future | archive-date = 18 August 2016}}</ref>

Woolly mammoths continued growing past adulthood, like other elephants. Unfused limb bones show that males grew until they reached the age of 40, and females grew until they were 25. The frozen calf "Dima" was {{cvt|90|cm}} tall when it died at the age of 6–12 months. At this age, the second set of molars would erupt, and the first set would be worn out at 18 months of age. The third set of molars lasted for 10 years, and this process was repeated until the final, sixth set emerged when the animal was 30 years old. When the last set of molars was worn out, the animal would be unable to chew and feed, and it would die of starvation. A study of North American mammoths found that they often died during winter or spring, the hardest times for northern animals to survive.<ref name="Lister, 2007. pp. 83–107"/>

Examination of preserved calves shows that they were all born during spring and summer, and since modern elephants have [[gestation periods]] of 21–22 months,  the [[mating season]] probably was from summer to autumn.<ref>Lister, 2007. pp.&nbsp;104–105</ref> [[δ15N]] isotopic analysis of the teeth of "Lyuba" has demonstrated their [[prenatal development]] and indicates its gestation period was similar to that of a modern elephant and that it was born in spring.<ref name="Rountrey et al., 2012" />

The best-preserved head of a frozen adult specimen, that of a male nicknamed the "Yukagir mammoth", shows that woolly mammoths had temporal glands between the ear and the eye.<ref name="Yukagir">{{Cite journal | last1 = Mol | first1 = D.| last2 = Shoshani| first2 = J. | last3 = Tikhonov | first3 = A. | last4 = van Geel | first4 = B. | last5 = Sano | first5 = S. | last6 = Lasarev | first6 = P.| last7 = Agenbroad| first7 = L. | title = The Yukagir mammoth: brief history, 14c dates, individual age, gender, size, physical and environmental conditions and storage | journal = Scientific Annals, School of Geology Aristotle University of Thessaloniki | volume = 98 | pages = 299–314 | year = 2006}}</ref> This feature indicates that, like bull elephants, male woolly mammoths entered "[[musth]]", a period of heightened aggressiveness. The glands are used especially by males to produce an oily substance with a strong smell called [[temporin]]. Their fur may have helped spread the scent further.<ref>Lister, 2007. pp.&nbsp;102–103</ref> This was confirmed by a 2023 study that compared the [[testosterone]] level in the dentine of an adult African elephant tusk with that of a male woolly mammoth.<ref name="Musth">{{cite journal |last1=Cherney |first1=Michael D. |last2=Fisher |first2=Daniel C. |last3=Auchus |first3=Richard J. |last4=Rountrey |first4=Adam N. |last5=Selcer |first5=Perrin |last6=Shirley |first6=Ethan A. |last7=Beld |first7=Scott G. |last8=Buigues |first8=Bernard |last9=Mol |first9=Dick |last10=Boeskorov |first10=Gennady G. |last11=Vartanyan |first11=Sergey L. |last12=Tikhonov |first12=Alexei N. |title=Testosterone histories from tusks reveal woolly mammoth musth episodes |journal=Nature |date=2023 |volume=617 |issue=7961 |pages=533–539 |doi=10.1038/s41586-023-06020-9|pmid=37138076 |bibcode=2023Natur.617..533C |s2cid=258485513 }}</ref>

===Palaeopathology===
Evidence of several different [[bone disease]]s has been found in woolly mammoths. The most common of these was [[osteoarthritis]], found in 2% of specimens. One specimen from Switzerland had several fused [[vertebra]]e due to this condition. The "Yukagir mammoth" had suffered from [[spondylitis]] in two vertebrae, and [[osteomyelitis]] is known from some specimens. Several specimens have healed [[bone fracture]]s, showing that the animals had survived these injuries.<ref>Lister, 2007. pp.&nbsp;108–111</ref> Likewise, [[spondyloarthropathy]] has also been identified in woolly mammoth remains.<ref name="SpondyloarthropathyInProboscideans">{{cite journal |last1=Rothschild |first1=Bruce M. |last2=Wang |first2=Xiaoming |last3=Shoshani |first3=Jeheskel |date=September 1994 |title=Spondyloarthropathy in Proboscideans |url=https://www.jstor.org/stable/20095389 |language=en |journal=Journal of Zoo and Wildlife Medicine |volume=25 |issue=3 |pages=360–366 |jstor=20095389 |access-date=17 April 2024}}</ref> An extra number of cervical vertebrae has been found in 33% of specimens from the North Sea region, probably due to a drop in numbers and subsequent inbreeding.<ref>{{Cite journal | doi = 10.7717/peerj.318| pmid = 24711969| pmc = 3970796| title = Extraordinary incidence of cervical ribs indicates vulnerable condition in Late Pleistocene mammoths| journal = [[PeerJ]]| volume = 2| pages = e318| year = 2014| last1 = Reumer | first1 = J. W. F. | last2 = Ten Broek | first2 = C. M. A. | last3 = Galis | first3 = F. | doi-access = free}}</ref> Vertebral lesions in woolly mammoths have been speculated to have resulted from nutritional stress.<ref>{{Cite journal |last1=Haynes |first1=Gary |last2=Klimowicz |first2=Janis |date=27 August 2015 |title=A preliminary review of bone and teeth abnormalities seen in recent Loxodonta and extinct Mammuthus and Mammut, and suggested implications |url=https://linkinghub.elsevier.com/retrieve/pii/S1040618215002499 |journal=[[Quaternary International]] |language=en |volume=379 |pages=135–146 |doi=10.1016/j.quaint.2015.04.001 |bibcode=2015QuInt.379..135H |access-date=19 April 2024 |via=Elsevier Science Direct|url-access=subscription }}</ref> Woolly mammoth ribs have also been shown to contain pathologies, including healed fractures and [[osteoporosis]].<ref>{{Cite journal |last1=Krzemińska |first1=Alina |last2=Wędzicha |first2=Sylwia |date=2 March 2015 |title=Pathological changes on the ribs of woolly mammoths (Mammuthus primigenius) |url=https://www.sciencedirect.com/science/article/pii/S1040618214002833 |journal=[[Quaternary International]] |language=en |volume=359-360 |pages=186–194 |doi=10.1016/j.quaint.2014.04.059 |bibcode=2015QuInt.359..186K |access-date=24 March 2025 |via=Elsevier Science Direct|url-access=subscription }}</ref> [[Parasitism|Parasitic]] flies and [[protozoa]] were identified in the gut of the calf "Dima".<ref>Lister, 2007. p. 87</ref>

Distortion in the molars is the most common health problem found in woolly mammoth fossils. Sometimes, the replacement was disrupted, and the molars were pushed into abnormal positions, but some animals are known to have survived this. Teeth from Britain showed that 2% of specimens had [[periodontal disease]], with half of these containing [[caries]].<ref name="Lister, 2007. pp. 88–89" /> The teeth sometimes had [[cancerous]] growths.<ref>Lister, 2007. pp.&nbsp;108–109</ref>

==Distribution and habitat==

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|footer = The [[Altai-Sayan assemblages]], such as [[Khar-Us Nuur]] and [[Ukok-Sailiugem]], are similar to the "[[mammoth steppe]]"
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The habitat of the woolly mammoth is known as "[[mammoth steppe]]" or "tundra steppe". This environment stretched across northern Asia, many parts of Europe, and the northern part of North America during the last ice age. It was similar to the grassy [[steppes]] of modern Russia, but the [[flora]] was more diverse, abundant, and grew faster. Grasses, sedges, shrubs, and herbaceous plants were present, and scattered trees were mainly found in southern regions. This habitat was not dominated by ice and snow, as is popularly believed, since these regions are thought to have been [[high-pressure area]]s at the time. The habitat of the woolly mammoth supported other grazing herbivores such as the [[woolly rhinoceros]], [[Equus (genus)|wild horses]], and [[bison]].<ref name="Lister, 2007. pp. 88–89">Lister, 2007. pp.&nbsp;88–89</ref> The [[Altai-Sayan assemblages]] are the modern [[biomes]] most similar to the "mammoth steppe".<ref name="Altai-Sayan">{{cite journal |last1=Pavelková Řičánková |first1=V. |last2=Robovský |first2=J. |last3=Riegert |first3=J. |title=Ecological Structure of Recent and Last Glacial Mammalian Faunas in Northern Eurasia: The Case of Altai-Sayan Refugium |journal=PLOS ONE |date=13 January 2014 |volume=9 |issue=1 |pages=e85056 |doi=10.1371/journal.pone.0085056|pmid=24454791 |pmc=3890305 |bibcode=2014PLoSO...985056P |doi-access=free }}</ref> A 2014 study concluded that [[forb]]s (a group of herbaceous plants) were more important in the steppe-tundra than previously acknowledged, and that it was a primary food source for the ice-age megafauna.<ref>{{Cite journal | doi = 10.1038/nature12921| title = Fifty thousand years of Arctic vegetation and megafaunal diet| journal = Nature| volume = 506| issue = 7486| pages = 47–51| year = 2014| last1 = Willerslev | first1 = E. | last2 = Davison | first2 = J. | last3 = Moora | first3 = M. | last4 = Zobel | first4 = M. | last5 = Coissac | first5 = E. | last6 = Edwards | first6 = M. E. | last7 = Lorenzen | first7 = E. D. | last8 = Vestergård | first8 = M. | last9 = Gussarova | first9 = G. | last10 = Haile | first10 = J. | last11 = Craine | first11 = J. | last12 = Gielly | first12 = L. | last13 = Boessenkool | first13 = S. | last14 = Epp | first14 = L. S. | last15 = Pearman | first15 = P. B. | last16 = Cheddadi | first16 = R. | last17 = Murray | first17 = D. | last18 = Bråthen | first18 = K. A. | last19 = Yoccoz | first19 = N. | last20 = Binney | first20 = H. | last21 = Cruaud | first21 = C. | last22 = Wincker | first22 = P. | last23 = Goslar | first23 = T. | last24 = Alsos | first24 = I. G. | last25 = Bellemain | first25 = E. | last26 = Brysting | first26 = A. K. | last27 = Elven | first27 = R. | last28 = Sønstebø | first28 = J. R. H. | last29 = Murton | first29 = J. | last30 = Sher | first30 = A. | display-authors = 29 | pmid=24499916| bibcode = 2014Natur.506...47W| s2cid = 4461741| url = https://eprints.soton.ac.uk/377215/3/Willerslev_Fig2.pdf}}</ref>

[[File:Wooly Mammoths.jpg|thumb|Mural depicting a herd walking near the [[Somme River]] in France, by [[Charles R. Knight]], 1916]]
The southernmost woolly mammoth specimen known is from the [[Shandong province]] of China and is 33,000 years old.<ref>{{Cite journal | last1 = Takahashi | first1 = K. | last2 = Wei | first2 = G. | last3 = Uno | first3 = H. | last4 = Yoneda | first4 = M. | last5 = Jin | first5 = C. | last6 = Sun | first6 = C. | last7 = Zhang | first7 = S. | last8 = Zhong | first8 = B. | doi = 10.1016/j.quascirev.2006.12.001 | title = AMS 14C chronology of the world's southernmost woolly mammoth (''Mammuthus primigenius'' Blum.) | journal = Quaternary Science Reviews | volume = 26 | issue = 7–8 | pages = 954–957 | year = 2007 |bibcode = 2007QSRv...26..954T }}</ref> The southernmost European remains are from the [[Depression of Granada]] in Spain and are of roughly the same age.<ref>{{Cite journal | last1 = Álvarez-Lao | first1 = D. J. | last2 = García | first2 = N. | doi = 10.1016/j.quascirev.2011.11.004 | title = Comparative revision of the Iberian woolly mammoth (''Mammuthus primigenius'') record into a European context | journal = Quaternary Science Reviews | volume = 32 | pages = 64–74 | year = 2012 | bibcode = 2012QSRv...32...64A | hdl = 10651/9463 | hdl-access = free }}</ref><ref>{{Citation |title=The Padul mammoth finds — On the southernmost record of ''Mammuthus primigenius'' in Europe and its southern spread during the Late Pleistocene | author=Diego J. Alvarez-Lao|display-authors=etal | journal=Palaeogeography, Palaeoclimatology, Palaeoecology | year=2009 | volume=278 | issue=1–4 | pages=57–70 | doi=10.1016/j.palaeo.2009.04.011 | bibcode=2009PPP...278...57A| url=http://doc.rero.ch/record/209767/files/PAL_E4203.pdf }}</ref> DNA studies have helped determine the [[phylogeography]] of the woolly mammoth. A 2008 DNA study showed two distinct groups of woolly mammoths: one that became extinct 45,000 years ago and another one that became extinct 12,000 years ago. The two groups are speculated to be divergent enough to be characterised as [[subspecies]]. The group that became extinct earlier stayed in the middle of the high Arctic, while the group with the later extinction had a much wider range.<ref>{{Cite journal | last1 = Gilbert | first1 = M. T. P. | last2 = Drautz | first2 = D. I. | last3 = Lesk | first3 = A. M. | last4 = Ho | first4 = S. Y. W. | last5 = Qi | first5 = J. | last6 = Ratan | first6 = A. | last7 = Hsu | first7 = C. -H. | last8 = Sher | first8 = A. | last9 = Dalen | first9 = L. | last10 = Gotherstrom | doi = 10.1073/pnas.0802315105 | first10 = A. | last11 = Tomsho | first11 = L. P. | last12 = Rendulic | first12 = S. | last13 = Packard | first13 = M. | last14 = Campos | first14 = P. F. | last15 = Kuznetsova | first15 = T. V. | last16 = Shidlovskiy | first16 = F. | last17 = Tikhonov | first17 = A. | last18 = Willerslev | first18 = E. | last19 = Iacumin | first19 = P. | last20 = Buigues | first20 = B. | last21 = Ericson | first21 = P. G. P. | last22 = Germonpre | first22 = M. | last23 = Kosintsev | first23 = P. | last24 = Nikolaev | first24 = V. | last25 = Nowak-Kemp | first25 = M. | last26 = Knight | first26 = J. R. | last27 = Irzyk | first27 = G. P. | last28 = Perbost | first28 = C. S. | last29 = Fredrikson | first29 = K. M. | last30 = Harkins | first30 = T. T. | title = Intraspecific phylogenetic analysis of Siberian woolly mammoths using complete mitochondrial genomes | journal = Proceedings of the National Academy of Sciences | volume = 105 | issue = 24 | pages = 8327–8332 | year = 2008 | pmid =  18541911| pmc = 2423413| bibcode = 2008PNAS..105.8327G | doi-access = free }}</ref> Recent stable isotope studies of Siberian and New World mammoths have shown there were differences in climatic conditions on either side of the [[Bering land bridge]] (Beringia), with Siberia being more uniformly cold and dry throughout the Late Pleistocene.<ref>{{Cite journal | last1 = Szpak | first1 = P. | last2 = Gröcke | first2 = D. R. | last3 = Debruyne | first3 = R. | last4 = MacPhee | first4 = R. D. E. | last5 = Guthrie | first5 = R. D. | last6 = Froese | first6 = D. | last7 = Zazula | first7 = G. D. | last8 = Patterson | first8 = W. P. | last9 = Poinar | first9 = H. N. | title = Regional differences in bone collagen δ13C and δ15N of Pleistocene mammoths: Implications for paleoecology of the mammoth steppe | doi = 10.1016/j.palaeo.2009.12.009 | journal = Palaeogeography, Palaeoclimatology, Palaeoecology | volume = 286 | issue = 1–2 | pages = 88–96 | year = 2010 | bibcode = 2010PPP...286...88S | url = https://durham-repository.worktribe.com/output/1543860 }}</ref> During the [[Younger Dryas]] age, woolly mammoths briefly expanded into north-east Europe, whereafter the mainland populations became extinct.<ref>{{Cite journal | last1 = Stuart | first1 = A. J. | doi = 10.1016/j.quaint.2004.04.021 | title = The extinction of woolly mammoth (''Mammuthus primigenius'') and straight-tusked elephant (''Palaeoloxodon antiquus'') in Europe | journal = Quaternary International | volume = 126–128 | pages = 171–177 | year = 2005 |bibcode = 2005QuInt.126..171S | url = http://doc.rero.ch/record/13497/files/PAL_E278.pdf }}</ref>

A 2008 genetic study showed that some of the woolly mammoths that entered North America through the Bering land bridge from Asia migrated back about 300,000 years ago and had replaced the previous Asian population by about 40,000 years ago, not long before the entire species became extinct.<ref>{{Cite journal | last1 = Debruyne | first1 = R. | last2 = Chu | first2 = G. | last3 = King | first3 = C. E. | last4 = Bos | first4 = K. | last5 = Kuch | first5 = M. | last6 = Schwarz | first6 = C. | last7 = Szpak | first7 = P. | last8 = Gröcke | first8 = D. R. | last9 = Matheus | first9 = P. | last10 = Zazula | doi = 10.1016/j.cub.2008.07.061 | first10 = G. | last11 = Guthrie | first11 = D. | last12 = Froese | first12 = D. | last13 = Buigues | first13 = B. | last14 = De Marliave | first14 = C. | last15 = Flemming | first15 = C. | last16 = Poinar | first16 = D. | last17 = Fisher | first17 = D. | last18 = Southon | first18 = J. | last19 = Tikhonov | first19 = A. N. | last20 = MacPhee | first20 = R. D. E. | last21 = Poinar | first21 = H. N. | title = Out of America: Ancient DNA Evidence for a New World Origin of Late Quaternary Woolly Mammoths | journal = Current Biology | volume = 18 | issue = 17 | pages = 1320–1326 | year = 2008 | pmid =  18771918| s2cid = 18663366 | doi-access = free | bibcode = 2008CBio...18.1320D }}</ref> Fossils of woolly mammoths and Columbian mammoths have been found together in a few localities of North America, including the [[Hot Springs sinkhole]] of [[South Dakota]] where their regions overlapped. It is unknown whether the two species were sympatric and lived there simultaneously, or if the woolly mammoths may have entered these southern areas during times when Columbian mammoth populations were absent there.<ref name="Lister, 2007. pp. 88–89"/>

==Relationship with humans==
[[File:Woolly mammoth carved in ivory.jpg|thumb|left|Woolly mammoth engraved on ivory found in 1864, the first known contemporary depiction of a prehistoric animal]]
Modern humans coexisted with woolly mammoths during the [[Upper Palaeolithic]] period when humans entered Europe from Africa between 30,000 and 40,000 years ago. Before this, [[Neanderthals]] had coexisted with mammoths during the [[Middle Palaeolithic]] and already used mammoth bones for tool-making and building materials. Woolly mammoths were very important to ice age humans, and human survival may have depended on the mammoth in some areas. Evidence for such coexistence was not recognised until the 19th century. [[William Buckland]] published his discovery of the [[Red Lady of Paviland]] skeleton in 1823, which was found in a cave alongside woolly mammoth bones, but he mistakenly denied that these were contemporaries. In 1864, [[Édouard Lartet]] found an engraving of a woolly mammoth on a piece of mammoth ivory in the [[Abri de la Madeleine]] cave in [[Dordogne]], France. The engraving was the first widely accepted evidence for the coexistence of humans with prehistoric extinct animals and is the first contemporary depiction of such a creature known to modern science.<ref>Lister, 2007. pp.&nbsp;116–117</ref>

{{multiple image
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|footer = Various prehistoric depictions of woolly mammoths, including [[cave paintings]] (above) and sculptures
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The woolly mammoth is the third-most depicted animal in [[ice age art]], after horses and bison, and these images were produced between 35,000 and 11,500 years ago. Today, more than 500 depictions of woolly mammoths have been discovered in media ranging from cave paintings and engravings on the walls of 46 caves in Russia, France, and Spain to engravings and sculptures (termed "[[portable art]]") made from ivory, antler, stone, and bone. Cave paintings of woolly mammoths exist in several styles and sizes. The French [[Rouffignac Cave]] has the most depictions, 159, and some of the drawings are more than {{cvt|2|m}} in length. Other notable caves with mammoth depictions are the [[Chauvet Cave]], [[Les Combarelles Cave]], and [[Font-de-Gaume]].<ref name="Lister 2007 118 125">Lister, 2007. pp.&nbsp;118–125</ref> A depiction in the [[Cave of El Castillo]] may instead show ''Palaeoloxodon'', the "straight-tusked elephant".<ref name=art>{{Cite journal | last1 = Braun | first1 = I. M. | last2 = Palombo | first2 = M. R. | doi = 10.1016/j.quaint.2012.07.010 | title = ''Mammuthus primigenius'' in the cave and portable art: An overview with a short account on the elephant fossil record in Southern Europe during the last glacial | journal = Quaternary International | volume = 276–277 | pages = 61–76 | year = 2012 | bibcode = 2012QuInt.276...61B }}</ref>

"Portable art" can be more accurately dated than cave art since it is found in the same deposits as tools and other ice age artifacts. The largest collection of portable mammoth art, consisting of 62 depictions on 47 plaques, was found in the 1960s at an excavated open-air camp near [[Gönnersdorf]] in Germany. A correlation between the number of mammoths depicted and the most often hunted species does not seem to exist since reindeer bones are the most frequently found animal remains at the site. Two [[spear throwers]] shaped as woolly mammoths have been found in France.<ref name="Lister 2007 118 125"/> Some portable mammoth depictions may not have been produced where they were discovered but could have moved around by ancient trading.<ref name=art/>

===Exploitation===
[[File:Mammoth House (Replica).JPG|left|thumb|Reconstructed bone hut based on finds in [[Mezhyrich, Cherkasy Oblast|Mezhyrich]], exhibited in Japan]]
Woolly mammoth bones were used as construction material for dwellings by both Neanderthals and modern humans during the ice age.<ref>{{cite journal|last1=Demay|first1=L.|last2=Péan|first2=S.|last3=Patou-Mathis|first3=M.|title=Mammoths used as food and building resources by Neanderthals: Zooarchaeological study applied to layer 4, Molodova I (Ukraine)|journal=Quaternary International|date=October 2012|volume=276–277|pages=212–226|doi=10.1016/j.quaint.2011.11.019|bibcode=2012QuInt.276..212D|hdl=2268/190618|url=http://orbi.ulg.ac.be/bitstream/2268/190618/1/Demay%20et%20al%2c%202012%20Molodova.pdf}}</ref> More than 70  such dwellings are known, mainly from the [[East European Plain]]. The huts' bases were circular, ranging from {{cvt|8|to|24|m2}}. The arrangement of dwellings varied and ranged from {{cvt|1|to|20|m|ft}} apart, depending on location. Large bones were used as foundations for the huts, tusks for the entrances, and the roofs were probably skins held in place by bones or tusks. Some huts had floors that extended {{cvt|40|cm}} below ground. Some of the bones used for materials may have come from mammoths killed by humans, but the state of the bones and the fact that bones used to build a single dwelling varied by several thousands of years in age suggests that they were collected remains of long-dead animals. Woolly mammoth bones were made into various tools, furniture, and musical instruments. Large bones, such as shoulder blades, were used to cover dead human bodies during burial.<ref>Lister, 2007. pp.&nbsp;128–132</ref>

Woolly mammoth ivory was used to create art objects. Several [[Venus figurines]], including the [[Venus of Brassempouy]] and the [[Venus of Lespugue]], were made from this material. Weapons made from ivory, such as daggers, spears, and a [[boomerang]], are known. A 2019 study found that woolly mammoth ivory was the most suitable bony material for producing big game projectile points during the Late Pleistocene. To process the ivory, the large tusks had to be chopped, chiseled, and split into smaller, more manageable pieces. Some ivory artifacts show that tusks had been straightened, and how this was achieved is unknown.<ref>Lister, 2007. pp.&nbsp;131–137</ref><ref name="Projectiles">{{cite journal |last1=Pfeifer |first1=S. J. |last2=Hartramph |first2=W. L. |last3=Kahlke |first3=R.-D. |last4=Müller |first4=F. A. |title=Mammoth ivory was the most suitable osseous raw material for the production of Late Pleistocene big game projectile points |journal=Scientific Reports |date=2019 |volume=9 |issue=1 |pages=2303 |doi=10.1038/s41598-019-38779-1|pmid=30783179 |pmc=6381109 |doi-access=free }}</ref>

{{multiple image
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|footer = Artifacts made from woolly mammoth ivory; The [[Venus of Brassempouy]], the [[Venus of Moravany]], and the [[Lion-Man]]

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Woolly mammoths were an important food source for both modern humans and Neanderthals.<ref>{{Cite journal |last=Metcalfe |first=Jessica Z. |date=2 July 2017 |title=Proboscidean isotopic compositions provide insight into ancient humans and their environments |url=https://www.sciencedirect.com/science/article/abs/pii/S1040618216302300 |journal=[[Quaternary International]] |language=en |volume=443 |pages=147–159 |doi=10.1016/j.quaint.2016.12.003 |bibcode=2017QuInt.443..147M |access-date=8 September 2024 |via=Elsevier Science Direct|url-access=subscription }}</ref> Several woolly mammoth specimens show evidence of being butchered by humans, which is indicated by breaks, cut marks, and associated stone tools. How much prehistoric humans relied on woolly mammoth meat is unknown since many other large herbivores were available. Many mammoth carcasses may have been scavenged by humans rather than hunted. Some cave paintings show woolly mammoths in structures interpreted as pitfall traps. Few specimens show direct, unambiguous evidence of having been hunted by humans. A Siberian specimen with a spearhead embedded in its shoulder blade shows that a spear had been thrown at it with great force.<ref>Lister, 2007. pp.&nbsp;151–155</ref>

At a site in southern Poland that contains bones from over 100 mammoths, stone spear tips have been found embedded in bones, and many stone spear points in the site were damaged from impact against mammoth bones, indicating that mammoths were the major prey for people at the time.<ref>{{Cite journal|last=Wojtal|first=Piotr|date=2019|title=The earliest direct evidence of mammoth hunting in Central Europe|url=https://doi.org/10.1016/j.quascirev.2019.04.004|journal=Quaternary Science Reviews|volume=213|pages=162–166|doi=10.1016/j.quascirev.2019.04.004 |s2cid=149647112 |url-access=subscription}}</ref> A specimen from the [[Mousterian]] age of Italy shows evidence of spear hunting by Neanderthals.<ref>{{Cite journal | last1 = Mussi | first1 = M. | last2 = Villa | first2 = P. | doi = 10.1016/j.jas.2008.04.014 | title = Single carcass of ''Mammuthus primigenius'' with lithic artifacts in the Upper Pleistocene of northern Italy | journal = Journal of Archaeological Science | volume = 35 | issue = 9 | pages = 2606–2613 | year = 2008 | bibcode = 2008JArSc..35.2606M | url = http://doc.rero.ch/record/15932/files/PAL_E3343.pdf }}</ref>  The juvenile specimen nicknamed "Yuka" is the first frozen mammoth with evidence of human interaction. It shows evidence of having been killed by a large predator and scavenged by humans shortly after. Some of its bones had been removed and were found nearby.<ref name="bbcyuka" /> A site near the [[Yana River]] in Siberia has revealed several specimens with evidence of human hunting, but the finds were interpreted to show that the animals were not hunted intensively, but perhaps mainly when ivory was needed.<ref>{{Cite journal | last1 = Nikolskiy | first1 = P. | last2 = Pitulko | first2 = V. | doi = 10.1016/j.jas.2013.05.020 | title = Evidence from the Yana Palaeolithic site, Arctic Siberia, yields clues to the riddle of mammoth hunting | journal = Journal of Archaeological Science | volume = 40 | issue = 12 | pages = 4189–4197 | year = 2013 | bibcode = 2013JArSc..40.4189N }}</ref> Two woolly mammoths from [[Wisconsin]], the "Schaefer" and "Hebior mammoths", show evidence of having been butchered by [[Paleo-Indians]].<ref>{{Cite journal | doi = 10.1002/gea.10052| title = Geoarchaeological contexts for Late Pleistocene archaeological sites with human-modified woolly mammoth remains in southeastern Wisconsin, U.S.A| journal = Geoarchaeology| volume = 18| pages = 91–114| year = 2003| last1 = Overstreet | first1 = D. F. | last2 = Kolb | first2 = M. F. | issue = 1| bibcode = 2003Gearc..18...91O| s2cid = 129431648}}</ref><ref>{{Cite journal | doi = 10.1016/j.quaint.2005.03.004| title = Chronology and new research on the Schaefer mammoth (?''Mammuthus primigenius'') site, Kenosha County, Wisconsin, USA| journal = Quaternary International| volume = 142–143| pages = 44–57| year = 2006| last1 = Joyce | first1 = D. J. | bibcode = 2006QuInt.142...44J}}</ref>

==Extinction==

Most woolly mammoth populations disappeared during the late Pleistocene and mid-[[Holocene]],<ref name="Yukon20212">{{cite journal |last=Murchie |first=Tyler J. |author2=Monteath, Alistair J. |author3=Mahony, Matthew E. |author4=Long, George S. |author5=Cocker, Scott |author6=Sadoway, Tara |author7=Karpinski, Emil |author8=Zazula, Grant |author9=MacPhee, Ross D. E. |author10=Froese, Duane |author11=Poinar, Hendrik N. |year=2021 |title=Collapse of the mammoth-steppe in central Yukon as revealed by ancient environmental DNA |journal=[[Nature Communications]] |volume=12 |issue=7120 (2021) |pages=2031 |bibcode=2007QSRv...26.2031B |doi=10.1038/s41467-021-27439-6|pmid=34880234 |pmc=8654998 }}</ref> coinciding with the extinction of most North American [[Pleistocene megafauna]]  (including the Columbian mammoth) as well as the extinctions or extirpations of steppe-associated fauna of Eurasia that coexisted with the mammoth species (such as the woolly rhinoceros, the [[Panthera spelaea|cave lion]], reindeer, [[saiga]], the [[Arctic fox]], and the [[steppe lemming]]). This extinction formed part of the [[Late Pleistocene extinctions]], which began 40,000 years ago and peaked between 14,000 and 11,500 years ago. Scientists are divided over whether hunting or climate change, which led to the shrinkage of its habitat, was the main factor that contributed to the extinction of the woolly mammoth, or whether it was due to a combination of the two.<ref>{{Cite journal |last1=Nogués-Bravo |first1=David |last2=Rodríguez |first2=Jesús |last3=Hortal |first3=Joaquín |last4=Batra |first4=Persaram |last5=Araújo |first5=Miguel B. |date=2008-04-01 |title=Climate Change, Humans, and the Extinction of the Woolly Mammoth |journal=PLOS Biology |language=en |volume=6 |issue=4 |pages=e79 |doi=10.1371/journal.pbio.0060079 |pmid=18384234 |pmc=2276529 |issn=1545-7885 |doi-access=free }}</ref><ref>{{cite journal|last1=Stuart|first1=Anthony J.|last2=Lister|first2=Adrian M.|year=2007|title=Patterns of Late Quaternary megafaunal extinctions in Europe and northern Asia|journal=CFS Courier Forschungsinstitut Senckenberg|volume=259|pages=289–299}}</ref> Evidence from tusk-derived [[Δ18O|''δ''<sup>18</sup>O]] values suggests that climate change was not the direct cause of Eurasian woolly mammoths, as ''δ''<sup>18</sup>O did not significantly vary in areas where woolly mammoths died out and where they persisted for longer into the Holocene.<ref>{{Cite journal |last1=Fox |first1=David L. |last2=Fisher |first2=Daniel C. |last3=Vartanyan |first3=Sergey |last4=Tikhonov |first4=Alexei N. |last5=Mol |first5=Dick |last6=Buigues |first6=Bernard |date=July 2007 |title=Paleoclimatic implications of oxygen isotopic variation in late Pleistocene and Holocene tusks of Mammuthus primigenius from northern Eurasia |url=https://linkinghub.elsevier.com/retrieve/pii/S1040618206002096 |journal=[[Quaternary International]] |language=en |volume=169-170 |pages=154–165 |doi=10.1016/j.quaint.2006.09.001 |bibcode=2007QuInt.169..154F |access-date=24 September 2024 |via=Elsevier Science Direct|url-access=subscription }}</ref>

[[File:Projectile points made from mammoth ivory.png|thumb|right|[[Palaeolithic]] projectile points made from mammoth ivory, Pekárna cave]]
Whatever the cause, large mammals are generally more vulnerable than smaller ones due to their smaller population size and low reproduction rates.<ref name="Lister, 2007. pp. 146–148">Lister, 2007. pp.&nbsp;146–148</ref> Climatic patterns during the [[Last Interglacial]] (130–116 kyr BP) suggest that woolly mammoths and associated steppe faunas were sensitive to contractions of steppe-tundra habitats since they were adapted to cold, dry, and open environments. Genetic results and climatic models both indicate that habitats suitable for the woolly mammoth in Eurasia contracted during the interglacial period, which would have caused population [[bottleneck effects]] that restricted its range to a few northern areas. As the climate favoured colder environments, however, woolly mammoth populations rebounded during later glacial periods.<ref>{{cite journal|last1=Palkopoulou|first1=Eleftheria|last2=Dalén|first2=Love|last3=Lister|first3=Adrian M.|last4=Vartanyan|first4=Sergey|last5=Sablin|first5=Mikhail|last6=Sher|first6=Andrei|last7=Edmark|first7=Veronica Nyström|last8=Brandström|first8=Mikael D.|last9=Germonpré|first9=Mietje|last10=Barnes|first10=Ian|last11=Thomas|first11=Jessica A.|year=2013|title=Holarctic genetic structure and range dynamics in the woolly mammoth|journal=Proceedings of the Royal Society B|volume=280|issue=1770 |pages=1–10|doi=10.1098/rspb.2013.1910|pmid=24026825 |pmc=3779339}}</ref>

The Last Glacial Period of the late Pleistocene is considered that of the maximum geographic distribution of the woolly mammoth, occupying most of Europe, northern Asia, and northern North America, although several barriers such as ice sheets, high mountain chains, deserts, year-round water surfaces, and other grasslands prevented them from spreading farther.<ref>{{cite journal|last=Kahlke|first=Ralk-Dietrich|year=2015|title=The maximum geographic extension of Late Pleistocene Mammuthus primigenius (Proboscidea, Mammalia) and its limiting factors|journal=Quaternary International|volume=379|pages=147–154|doi=10.1016/j.quaint.2015.03.023|bibcode=2015QuInt.379..147K }}</ref> Towards the end of the Last Glacial period, from around 15,000 years ago, the mammoth steppe that the woolly mammoth inhabited was gradually replaced across most of Siberia with wet tundra and boreal and temperate forest, which the woolly mammoth would have found to be unfavourable habitat.<ref name="Dehasque-2021">{{Cite journal |last1=Dehasque |first1=Marianne |last2=Pečnerová |first2=Patrícia |last3=Muller |first3=Héloïse |last4=Tikhonov |first4=Alexei |last5=Nikolskiy |first5=Pavel |last6=Tsigankova |first6=Valeriya I. |last7=Danilov |first7=Gleb K. |last8=Díez-del-Molino |first8=David |last9=Vartanyan |first9=Sergey |last10=Dalén |first10=Love |last11=Lister |first11=Adrian M. |date=May 2021 |title=Combining Bayesian age models and genetics to investigate population dynamics and extinction of the last mammoths in northern Siberia |url=https://linkinghub.elsevier.com/retrieve/pii/S0277379121001207 |journal=Quaternary Science Reviews |volume=259 |pages=106913 |bibcode=2021QSRv..25906913D |doi=10.1016/j.quascirev.2021.106913}}</ref>

Different woolly mammoth populations did not die out simultaneously across their range, but gradually became extinct over time.<ref>{{Cite journal |last1=Murchie |first1=Tyler J. |last2=Monteath |first2=Alistair J. |last3=Mahony |first3=Matthew E. |last4=Long |first4=George S. |last5=Cocker |first5=Scott |last6=Sadoway |first6=Tara |last7=Karpinski |first7=Emil |last8=Zazula |first8=Grant |last9=MacPhee |first9=Ross D. E. |last10=Froese |first10=Duane |last11=Poinar |first11=Hendrik N. |date=2021-12-08 |title=Collapse of the mammoth-steppe in central Yukon as revealed by ancient environmental DNA |journal=Nature Communications |language=en |volume=12 |issue=1 |pages=7120 |doi=10.1038/s41467-021-27439-6 |pmid=34880234 |pmc=8654998 |bibcode=2021NatCo..12.7120M |issn=2041-1723}}</ref> The dynamics of different woolly mammoth populations varied as they experienced very different magnitudes of climatic and human impacts over time, suggesting that extinction causes would have varied by population.<ref name="populations">{{Cite journal |last1=Fordham |first1=Damien A. |last2=Brown |first2=Stuart C. |last3=Akçakaya |first3=H. Reşit |last4=Brook |first4=Barry W. |last5=Haythorne |first5=Sean |last6=Manica |first6=Andrea |last7=Shoemaker |first7=Kevin T. |last8=Austin |first8=Jeremy J. |last9=Blonder |first9=Benjamin |last10=Pilowsky |first10=Julia |last11=Rahbek |first11=Carsten |date=2022 |title=Process-explicit models reveal pathway to extinction for woolly mammoth using pattern-oriented validation |url=https://onlinelibrary.wiley.com/doi/abs/10.1111/ele.13911 |journal=Ecology Letters |language=en |volume=25 |issue=1 |pages=125–137 |doi=10.1111/ele.13911 |pmid=34738712 |bibcode=2022EcolL..25..125F |issn=1461-0248|hdl=11343/299174 |s2cid=243762825 |hdl-access=free }}</ref> Most populations disappeared between 14,000 and 10,000 years ago.<ref name="TheLatestWoollyMammoths" /> In Britain, woolly mammoths were still present between 14,500 and 14,000 BP.<ref>{{Cite journal |last=Lister |first=Adrian M. |date=18 June 2009 |title=Late-glacial mammoth skeletons ( Mammuthus primigenius ) from Condover (Shropshire, UK): anatomy, pathology, taphonomy and chronological significance |url=https://onlinelibrary.wiley.com/doi/10.1002/gj.1162 |journal=[[Geological Journal]] |language=en |volume=44 |issue=4 |pages=447–479 |doi=10.1002/gj.1162 |bibcode=2009GeolJ..44..447L |issn=0072-1050 |access-date=10 June 2024 |via=Wiley Online Library|url-access=subscription }}</ref> The youngest fossils of the mainland population are from the [[Kyttyk Peninsula]] of Siberia and date to 9,650 years ago.<ref name="Lister, 2007. pp. 146–148" /><ref name="TheLatestWoollyMammoths">{{Cite journal | last1 = Stuart | first1 = A. J. | last2 = Sulerzhitsky | first2 = L. D. | last3 = Orlova | first3 = L. A. | last4 = Kuzmin | first4 = Y. V. | last5 = Lister | first5 = A. M. | title = The latest woolly mammoths (''Mammuthus primigenius'' Blumenbach) in Europe and Asia: A review of the current evidence | doi = 10.1016/S0277-3791(02)00026-4 | journal = Quaternary Science Reviews | volume = 21 | issue = 14–15 | pages = 1559–1569 | year = 2002 | bibcode = 2002QSRv...21.1559S | url = http://doc.rero.ch/record/13491/files/PAL_E271.pdf }}</ref>

{{multiple image
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|footer = Woolly mammoth and [[muskox]] remains displayed on [[Wrangel Island]] (left) and a skeleton being excavated on the [[Taymyr Peninsula]] (right), places where mammoths survived until about 4,000 years ago
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A small population of woolly mammoths survived on [[Saint Paul Island (Alaska)|St. Paul Island]], Alaska, well into the Holocene,<ref name="Guthrie2">{{Cite journal |last1=Dale Guthrie |first1=R. |year=2004 |title=Radiocarbon evidence of mid-Holocene mammoths stranded on an Alaskan Bering Sea island |journal=Nature |volume=429 |issue=6993 |pages=746–749 |bibcode=2004Natur.429..746D |doi=10.1038/nature02612 |pmid=15201907 |s2cid=186242235}}</ref><ref>{{Cite journal |last1=Yesner |first1=D. R. |last2=Veltre |first2=D. W. |last3=Crossen |first3=K. J. |last4=Graham |first4=R. W. |title=5,700-year-old Mammoth Remains from Qagnax Cave, Pribilof Islands, Alaska |journal=Second World of Elephants Congress, (Hot Springs: Mammoth Site, 2005) |pages=200–203}}</ref><ref>{{Cite journal |last=Crossen |first=K. S. |year=2005 |title=5,700-Year-Old Mammoth Remains from the Pribilof Islands, Alaska: Last Outpost of North America Megafauna |url=https://gsa.confex.com/gsa/2005AM/finalprogram/abstract_97313.htm |url-status=dead |journal=Geological Society of America |volume=37 |page=463 |archive-url=https://web.archive.org/web/20160303235336/https://gsa.confex.com/gsa/2005AM/finalprogram/abstract_97313.htm |archive-date=3 March 2016 |access-date=13 February 2020}}</ref> with their extinction on the island being tightly constrained to around 5,600 years ago based on direct dating of bones and environmental proxies. This population is suggested to have gone extinct as a result of sea-level rise and increasing dryness of the island reducing freshwater availability, along with mammoth activity degrading the few freshwater sources on the island.<ref name="Graham20162">{{cite journal |last1=Graham |first1=R. W. |last2=Belmecheri |first2=S. |last3=Choy |first3=K. |last4=Culleton |first4=B. J. |last5=Davies |first5=L. J. |last6=Froese |first6=D. |last7=Heintzman |first7=P. D. |last8=Hritz |first8=C. |last9=Kapp |first9=J. D. |last10=Newsom |first10=L. A. |last11=Rawcliffe |first11=R. |date=1 August 2016 |title=Timing and causes of mid-Holocene mammoth extinction on St. Paul Island, Alaska |journal=Proceedings of the National Academy of Sciences |volume=113 |issue=33 |pages=9310–4 |doi=10.1073/pnas.1604903113 |pmc=4995940 |pmid=27482085 |doi-access=free |last12=Saulnier-Talbot |first12=É. |last13=Shapiro |first13=B. |last14=Wang |first14=Y. |last15=Williams |first15=J. W. |last16=Wooller |first16=M. J.|bibcode=2016PNAS..113.9310G }}</ref> The last population known from fossils remained on [[Wrangel Island]] in the [[Arctic Ocean]] until 4,000 years ago, well into the start of human [[civilization]] and several centuries subsequent to the construction of the [[Great Pyramid of Giza|Great Pyramid]] and [[Great Sphinx of Giza|Sphinx]] of [[ancient Egypt]].<ref>{{cite web |author=Markus Milligan |title=Mammoths still walked the earth when the Great Pyramid was being built |url=http://www.heritagedaily.com/2013/11/mammoths-still-walked-the-earth-when-the-great-pyramids-were-being-built/100307 |url-status=dead |archive-url=https://web.archive.org/web/20150630060405/http://www.heritagedaily.com/2013/11/mammoths-still-walked-the-earth-when-the-great-pyramids-were-being-built/100307 |archive-date=30 June 2015 |access-date=5 July 2015 |website=HeritageDaily – Heritage & Archaeology News}}</ref><ref>{{Cite journal |last1=Stuart |first1=A. J. |last2=Kosintsev |first2=P. A. |last3=Higham |first3=T. F. G. |last4=Lister |first4=A. M. |year=2004 |title=Pleistocene to Holocene extinction dynamics in giant deer and woolly mammoth |journal=Nature |volume=431 |issue=7009 |pages=684–689 |bibcode=2004Natur.431..684S |doi=10.1038/nature02890 |pmid=15470427 |s2cid=4415073|url=http://doc.rero.ch/record/13494/files/PAL_E274.pdf }}</ref><ref name="Vartanyan19952">{{cite journal |last=Vartanyan |first=S. L. |display-authors=etal |year=1995 |title=Radiocarbon Dating Evidence for Mammoths on Wrangel Island, Arctic Ocean, until 2000 BC |url=https://journals.uair.arizona.edu/index.php/radiocarbon/article/view/1640/1644 |url-status=live |journal=Radiocarbon |volume=37 |issue=1 |pages=1–6 |doi=10.1017/S0033822200014703 |bibcode=1995Radcb..37....1V |issn=0033-8222 |archive-url=https://web.archive.org/web/20120402103117/https://journals.uair.arizona.edu/index.php/radiocarbon/article/view/1640/1644 |archive-date=2 April 2012 |doi-access=free}}</ref><ref name="Nyström20122">{{Cite journal |last1=Nyström |first1=V. |last2=Humphrey |first2=J. |last3=Skoglund |first3=P. |last4=McKeown |first4=N. J. |last5=Vartanyan |first5=S. |last6=Shaw |first6=P. W. |last7=Lidén |first7=K. |last8=Jakobsson |first8=M. |last9=Barnes |first9=I. A. N. |last10=Angerbjörn |first10=A. |last11=Lister |first11=A. |year=2012 |title=Microsatellite genotyping reveals end-Pleistocene decline in mammoth autosomal genetic variation |url=http://urn.kb.se/resolve?urn=urn:nbn:se:nrm:diva-31 |journal=Molecular Ecology |volume=21 |issue=14 |pages=3391–3402 |doi=10.1111/j.1365-294X.2012.05525.x |pmid=22443459 |last12=Dalén |first12=L.|bibcode=2012MolEc..21.3391N |url-access=subscription }}</ref> However some studies have asserted that [[environmental DNA]] supports the existence of small mainland populations that died out at around the same time as their island counterparts; two studies in 2021 found that based on environmental DNA, mammoths survived in the [[Yukon]] until about 5,700 years ago, roughly concurrent with the St. Paul population and on the [[Taymyr Peninsula]] of Siberia until 3,900 to 4,100 years ago, roughly concurrent with the Wrangel population. The Taymyr Peninsula, with its drier habitat, may have served as a [[Refugium (population biology)|refugium]] for the mammoth steppe, supporting mammoths and other widespread Ice Age mammals such as wild horses (''[[Equus (genus)|Equus]]'' sp.).<ref name="Yukon20212"/><ref>{{cite journal |last1=Wang |first1=Y |last2=Pedersen |first2=M.W. |last3=Alsos |first3=I.g. |display-authors=etal |date=2021 |title=Late Quaternary dynamics of Arctic biota from ancient environmental genomics. |url=https://doi.org/10.1038/s41586-021-04016-x |journal=Nature|volume=600 |issue=7887 |pages=86–92 |doi=10.1038/s41586-021-04016-x |pmid=34671161 |pmc=8636272 |bibcode=2021Natur.600...86W }}</ref> However, ancient environmental DNA in cold environments can be reworked from older sediments into younger sediments that clearly post-date extinction, raising doubt about validity of these dates.<ref>{{Cite report |url=https://elifesciences.org/reviewed-preprints/89992v1 |title=Mitochondrial genomes of Pleistocene megafauna retrieved from recent sediment layers of two Siberian lakes |last1=Seeber |first1=Pa |last2=Batke |first2=L |date=2023-09-01 |publisher=elife |doi=10.7554/elife.89992.1 |last3=Dvornikov |first3=Y |last4=Schmidt |first4=A |last5=Wang |first5=Y |last6=Stoof-Leichsenring |first6=Kr |last7=Moon |first7=Kl |last8=Shapiro |first8=B |last9=Epp |first9=Ls |doi-access=free }}</ref>

DNA sequencing of remains of two mammoths, one from Siberia 44,800 years BP and one from Wrangel Island 4,300 years BP, indicates two major population crashes: one around 280,000 years ago, from which the population recovered, and a second about 12,000 years ago, near the ice age's end, from which it did not.<ref name = "Palkopoulou2015"/> The Wrangel Island mammoths were isolated for 5,000 years by rising post-ice-age sea level, and resultant inbreeding in their small population of about 300 to 1,000 individuals<ref>{{Cite web|url = http://www.abc.net.au/science/articles/2015/04/24/4223129.htm|title = Lonely end for the world's last woolly mammoths|date = 24 April 2015|access-date = 24 April 2015|website = ABC Science|publisher = Reuters|last = Dunham|first = W.|url-status = live|archive-url = https://web.archive.org/web/20150426174847/http://www.abc.net.au/science/articles/2015/04/24/4223129.htm|archive-date = 26 April 2015}}</ref> led to a 20%<ref name = "Palkopoulou2015">{{cite journal
 | last1 = Palkopoulou | first1 = E. | last2 = Mallick | first2 = S. | last3 = Skoglund | first3 = P. | last4 = Enk | first4 = J. | last5 = Rohland | first5 = N. | last6 = Li | first6 = H. | last7 = Omrak | first7 = A. | last8 = Vartanyan | first8 = S. | last9 = Poinar | first9 = H. | last10 = Götherström | first10 = A. | last11 = Reich | first11 = D. | last12 = Dalén | first12 = L. | title = Complete Genomes Reveal Signatures of Demographic and Genetic Declines in the Woolly Mammoth | url= | journal = [[Current Biology]] | volume =  25| issue =  10| pages = 1395–1400 | date = 23 April 2015 | doi = 10.1016/j.cub.2015.04.007 | pmid=25913407 | pmc=4439331| bibcode = 2015CBio...25.1395P }}</ref> to 30%<ref name="Nyström20122"/> loss of heterozygosity and a 65% loss in mitochondrial DNA diversity.<ref name="Nyström20122"/> The population seems to have subsequently been stable, without suffering further significant loss of genetic diversity.<ref name="Nyström20122"/><ref name = "Nyström2010">{{Cite journal | last1 = Nystrom | first1 = V. | last2 = Dalen | first2 = L. | last3 = Vartanyan | first3 = S. | last4 = Liden | first4 = K. | last5 = Ryman | first5 = N. | last6 = Angerbjorn | first6 = A. | doi = 10.1098/rspb.2010.0301 | title = Temporal genetic change in the last remaining population of woolly mammoth | journal = Proceedings of the Royal Society B: Biological Sciences | volume = 277 | issue = 1692 | pages = 2331–2337 | year = 2010 | pmid =  20356891| pmc = 2894910}}</ref> Genetic evidence thus implies the extinction of this final population was sudden, rather than the culmination of a gradual decline.<ref name="Nyström20122"/>

[[File:Woolly Mammoth Climatic Suitability - Nogués-Bravo 2008.png|thumb|right|Map showing climatic suitability for woolly mammoths in the Late Pleistocene and [[Holocene]] of [[Eurasia]]: red is increasing suitability, green is decreasing suitability. Black points are records of mammoths, black lines are the northern limit of humans]]
Before their extinction, the Wrangel Island mammoths had accumulated numerous genetic defects due to their small population; in particular, a number of genes for olfactory receptors and urinary proteins became nonfunctional, possibly because they had lost their selective value on the island environment.<ref name="Barsh2017">{{cite journal|last1= Barsh|first1=G. S.|last2= Rogers|first2=R. L.|last3= Slatkin |first3= M.|title= Excess of genomic defects in a woolly mammoth on Wrangel Island| journal= PLOS Genetics|volume= 13|issue= 3|date= 2017|page= e1006601|doi= 10.1371/journal.pgen.1006601|pmid=28253255|pmc=5333797 |doi-access=free }}</ref> It is not clear whether these genetic changes contributed to their extinction.<ref name="Switek2017">{{cite journal |last1= Switek|first1= B.|title= Dying woolly mammoths were in 'genetic meltdown' |journal= Nature|date=2017|doi= 10.1038/nature.2017.21575|s2cid= 184732688}}</ref> It has been proposed that these changes are consistent with the concept of [[genomic meltdown]],<ref name="Barsh2017" /> however, this has been contested by later analysis of the genomes of some of the last mammoths on Wrangel Island, which suggests that highly deleterious mutations had been significantly purged to levels lower than that in mainland populations, though the level of moderately deleterious mutations was elevated.<ref name="Dehasque-2024">{{Cite journal |last1=Dehasque |first1=Marianne |last2=Morales |first2=Hernán E. |last3=Díez-del-Molino |first3=David |last4=Pečnerová |first4=Patrícia |last5=Chacón-Duque |first5=J. Camilo |last6=Kanellidou |first6=Foteini |last7=Muller |first7=Héloïse |last8=Plotnikov |first8=Valerii |last9=Protopopov |first9=Albert |last10=Tikhonov |first10=Alexei |last11=Nikolskiy |first11=Pavel |last12=Danilov |first12=Gleb K. |last13=Giannì |first13=Maddalena |last14=van der Sluis |first14=Laura |last15=Higham |first15=Tom |date=June 2024 |title=Temporal dynamics of woolly mammoth genome erosion prior to extinction |journal=Cell |volume=187 |issue=14 |pages=3531–3540.e13 |language=en |doi=10.1016/j.cell.2024.05.033|pmid=38942016 |doi-access=free }}</ref> The sudden disappearance of an apparently stable population may be more consistent with a catastrophic event, possibly related to climate (such as icing of the snowpack), disease, or a human hunting expedition.<ref name="Dehasque-2024" /><ref name="Arppe2019">{{cite journal|last1= Arppe|first1= L.|last2= Karhu|first2=J. A.|last3= Vartanyan|first3= S.|last4= Drucker|first4=D. G.|last5=Etu-Sihvola|first5= H.|last6= Bocherens|first6= H.|title= Thriving or surviving? The isotopic record of the Wrangel Island woolly mammoth population|journal= Quaternary Science Reviews|volume= 222|year= 2019|pages= 105884|doi= 10.1016/j.quascirev.2019.105884|bibcode= 2019QSRv..22205884A|doi-access= free|hdl= 10138/309133|hdl-access= free}}</ref>

The disappearance is relatively close in time with the first evidence of humans on the island,<ref name="Ackerman1998">{{cite journal | last = Ackerman | first = R. E. | title = Early maritime traditions in the Bering, Chukchi, and East Siberian seas | journal = [[Arctic Anthropology]] | volume = 35 | issue = 1 | pages = 247–262 | date = 1998 | jstor = 40316468 }}</ref> though other authors have suggested that woolly mammoths were almost certainly extinct for several centuries prior to the presence of humans on Wrangel Island (which dates to around 3,600 years ago).<ref name="Dehasque-2021" /><ref name="Dehasque-2024" /> The woolly mammoths of eastern Beringia (modern [[Alaska]] and [[Yukon]]) had similarly died out about 13,300 years ago, soon (roughly 1,000 years) after the first appearance of humans in the area, which parallels the fate of all the other late Pleistocene proboscideans (mammoths, [[gomphothere]]s, and mastodons), as well as most of the rest of the [[megafauna]], of the [[Americas]].<ref name="Fiedel">{{Cite book  | first = Stuart  | last = Fiedel  | editor-last = Haynes  | editor-first = G. | contribution = Sudden Deaths: The Chronology of Terminal Pleistocene Megafaunal Extinction  | title = American Megafaunal Extinctions at the End of the Pleistocene  | year = 2009 | pages = 21–37 | publisher = [[Springer Science+Business Media|Springer]] | doi = 10.1007/978-1-4020-8793-6_2  | isbn = 978-1-4020-8792-9| series = Vertebrate Paleobiology and Paleoanthropology  }}</ref> In contrast, the St. Paul Island mammoth population apparently died out significantly before human arrival.<ref name="Graham20162" />

Changes in climate shrank suitable mammoth habitat from {{cvt|7700000|km2}} 42,000 years ago to {{cvt|800000|km2}}, a roughly 90% decrease, by 6,000 years ago.<ref name="Nogues-Bravo2008">{{Cite journal | last1 = Nogués-Bravo | first1 = D. | last2 = Rodríguez | first2 = J. S. | last3 = Hortal | first3 = J. N. | last4 = Batra | first4 = P. | last5 = Araújo | first5 = M. B. | editor1-last = Barnosky | editor1-first = Anthony | title = Climate Change, Humans, and the Extinction of the Woolly Mammoth | doi = 10.1371/journal.pbio.0060079 | journal = PLOS Biology | volume = 6 | issue = 4 | pages = e79 | year = 2008 | pmid =  18384234| pmc =2276529 | doi-access = free }}</ref><ref>{{Cite journal | last1 = Sedwick | first1 = C. | title = What Killed the Woolly Mammoth? | doi = 10.1371/journal.pbio.0060099 | journal = PLOS Biology | volume = 6 | issue = 4 | pages = e99 | year = 2008 | pmid =  20076709| pmc =2276526 | doi-access = free }}</ref> Woolly mammoths survived an even greater loss of habitat at the end of the [[Penultimate Glacial Period]] and the onset of the Last Interglacial, approximately 125,000 years ago.<ref>{{cite book | last = Martin | first = P. S | author-link = Paul Schultz Martin | title = Twilight of the Mammoths: Ice Age Extinctions and the Rewilding of America | publisher = [[University of California Press]] | year = 2005 | pages = [https://archive.org/details/twilightofmammot00paul/page/165 165–173] | isbn = 978-0-520-23141-2 | url = https://archive.org/details/twilightofmammot00paul/page/165 }}</ref><ref>{{Cite journal | last1 = Burney | first1 = D. | last2 = Flannery | first2 = T. | doi = 10.1016/j.tree.2005.04.022 | title = Fifty millennia of catastrophic extinctions after human contact | journal = Trends in Ecology & Evolution | volume = 20 | issue = 7 | pages = 395–401 | year = 2005 | pmid = 16701402 | url = http://www.anthropology.hawaii.edu/Fieldschools/Kauai/Publications/Publication%204.pdf | url-status = dead | archive-url = https://web.archive.org/web/20100610061434/http://www.anthropology.hawaii.edu/Fieldschools/Kauai/Publications/Publication%204.pdf | archive-date = 10 June 2010}}</ref> Studies of an 11,300–11,000-year-old [[trackway]] in south-western Canada showed that ''M. primigenius'' was in decline while coexisting with humans, since far fewer tracks of juveniles were identified than would be expected in a normal herd.<ref name="Trackway">{{Cite journal |last1=McNeil |first1=P. |last2=Hills |first2=L. |last3=Kooyman |first3=B. |last4=Tolman |first4=S. |year=2005 |title=Mammoth tracks indicate a declining Late Pleistocene population in southwestern Alberta, Canada |journal=[[Quaternary Science Reviews]] |volume=24 |issue=10–11 |pages=1253–1259 |bibcode=2005QSRv...24.1253M |doi=10.1016/j.quascirev.2004.08.019}}</ref> It has been suggested that human hunting exerted significant pressure on woolly mammoth populations for thousands of years across their range, making the population abundance of woolly mammoths considerably lower than it would have been otherwise even prior to their range decline, and likely hastened the range collapse of woolly mammoths in response to climate change.<ref name="populations" />

===Revival efforts===
{{Further|Revival of the woolly mammoth}}

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|footer = Models of an adult and the calf "Dima" in [[State Museum of Natural History Stuttgart]]
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The existence of preserved soft tissue remains and DNA of woolly mammoths has led to the idea that the species could be resurrected by scientific means. Several methods have been proposed to achieve this. [[Cloning]] would involve removal of the DNA-containing [[cell nucleus|nucleus]] of the [[Ovum|egg cell]] of a female elephant and replacement with a nucleus from woolly mammoth tissue. The cell would then be stimulated into dividing and inserted back into a female elephant. The resulting calf would have the genes of the woolly mammoth, although its fetal environment would be different. Most intact mammoths have had little usable DNA because of their conditions of preservation. There is not enough to guide the production of an embryo.<ref name='Zimmer 2013'>[http://www.nationalgeographic.com/magazine/2013/04/species-revival-bringing-back-extinct-animals/ Bringing them Back to Life] {{webarchive|url=https://web.archive.org/web/20170329051157/http://www.nationalgeographic.com/magazine/2013/04/species-revival-bringing-back-extinct-animals/ |date=2017 }}. Carl Zimmer, ''National Geographic''. April 2013.</ref><ref name="Lister 2007 42 43">Lister, 2007. pp.&nbsp;42–43</ref>

A second method involves [[artificially inseminating]] an elephant egg cell with [[sperm]] cells from a frozen woolly mammoth carcass. The resulting offspring would be an elephant–mammoth hybrid, and the process would have to be repeated so more hybrids could be used in breeding. After several generations of cross-breeding these hybrids, an almost pure woolly mammoth would be produced. The fact that sperm cells of modern mammals are viable for 15 years at most after deepfreezing makes this method unfeasible.<ref name="Lister 2007 42 43"/>

[[File:Elephants by the water (7568684536).jpg|thumb|Elephants are highly [[gregarious]], as shown by these [[Sri Lankan elephants]]]]
Several projects are working on gradually replacing the genes in elephant cells with mammoth genes.<ref name='genome 2015'>{{cite web |last1=Ghosh |first1=Pallab |title=Mammoth genome sequence completed |url=https://www.bbc.co.uk/news/science-environment-32432693 |website=[[BBC News]]|archive-url= https://web.archive.org/web/20150424120045/http://www.bbc.com/news/science-environment-32432693 |date=23 April 2015| archive-date= 24 April 2015}}</ref><ref name="Long Now">[http://longnow.org/revive/ The Long Now Foundation – Revive and Restore] {{webarchive|url=https://web.archive.org/web/20150424111602/http://longnow.org/revive/ |date=24 April 2015 }}.</ref> By 2015 and using the new [[CRISPR gene editing|CRISPR]] DNA editing technique, one team, led by [[George Church (geneticist)|George Church]], had some woolly mammoth genes edited into the genome of an Asian elephant; focusing on cold-resistance initially,<ref name='Landers 2015'>[https://www.washingtonpost.com/national/health-science/can-scientists-bring-mammoths-back-to-life-by-cloning/2015/02/06/2a825c8c-80ae-11e4-81fd-8c4814dfa9d7_story.html Can scientists bring mammoths back to life by cloning?] {{webarchive|url=https://web.archive.org/web/20171008082010/https://www.washingtonpost.com/national/health-science/can-scientists-bring-mammoths-back-to-life-by-cloning/2015/02/06/2a825c8c-80ae-11e4-81fd-8c4814dfa9d7_story.html?utm_term=.797eebd5da4f |date=8 October 2017 }} Jackson Landers. 9 February 2015. ''The Washington Post''.</ref> the target genes are for the external ear size, subcutaneous fat, [[hemoglobin]], and hair attributes.<ref name='Webster'>{{cite news |last=Webster |first=Ben |date=23 March 2015 |title=Scientist takes mammoth-cloning a step closer |url=https://www.thetimes.com/uk/science/article/scientist-takes-mammoth-cloning-a-step-closer-h3vqsqcl920 |newspaper=[[The Sunday Times]]}}</ref><ref>{{citation |url=http://www.popsci.com/woolly-mammoth-dna-brought-life-elephant-cells |title=Woolly Mammoth DNA Successfully Spliced Into Elephant Cells |author=Sarah Fecht |date=24 March 2014 |publisher=[[Popular Science]] |url-status=live |archive-url=https://web.archive.org/web/20150326161051/http://www.popsci.com/woolly-mammoth-dna-brought-life-elephant-cells |archive-date=26 March 2015  }}</ref> If any method is ever successful, a suggestion has been made to introduce the hybrids to a wildlife reserve in Siberia called the [[Pleistocene Park]].<ref>{{cite journal| last1=Zimov|first1=S. A.| title=Essays on Science and Society: Pleistocene Park: Return of the Mammoth's Ecosystem| journal=Science|volume=308|issue=5723|year=2005|pages=796–798|doi=10.1126/science.1113442|pmid=15879196|doi-access=free}}</ref>

Some researchers question the ethics of such recreation attempts. In addition to the technical problems, not much habitat is left that would be suitable for elephant-mammoth hybrids. Because the species was social and [[gregarious]], creating a few specimens would not be ideal. The time and resources required would be enormous, and the scientific benefits would be unclear, suggesting these resources should instead be used to preserve extant elephant species which are endangered.<ref name="Lister 2007 42 43"/><ref>{{cite journal |last1=Rohwer |first1=Y. |last2=Marris |first2=E. |title=An analysis of potential ethical justifications for mammoth de-extinction and a call for empirical research |journal=Ethics, Policy & Environment |date=2018 |volume=21 |issue=1 |pages=127–142 |doi=10.1080/21550085.2018.1448043|bibcode=2018EPolE..21..127R |s2cid=158056898 }}</ref><ref name="Griffin2015">{{cite web|url=https://www.independent.co.uk/life-style/gadgets-and-tech/news/woolly-mammoth-could-be-revived-after-scientists-paste-dna-into-elephants-genetic-code-10127125.html|title=Woolly mammoth could be revived after scientists paste DNA into elephant's genetic code|website=The Independent|date=23 March 2015|last=Griffin|first=A.|url-status=live|archive-url=https://web.archive.org/web/20150925044229/http://www.independent.co.uk/life-style/gadgets-and-tech/news/woolly-mammoth-could-be-revived-after-scientists-paste-dna-into-elephants-genetic-code-10127125.html|archive-date=25 September 2015}}</ref> The ethics of using elephants as surrogate mothers in hybridisation attempts has been questioned, as most embryos would not survive, and knowing the exact needs of a hybrid elephant–mammoth calf would be impossible.<ref>{{Cite book| last1=Loi| first1=Pasqualino| last2=Saragusty| first2=Joseph| last3=Ptak| first3=Grazyna| title=Reproductive Sciences in Animal Conservation| chapter=Cloning the Mammoth: A Complicated Task or Just a Dream?|volume=753|year=2014|pages=489–502| pmid=25091921| doi=10.1007/978-1-4939-0820-2_19|series=Advances in Experimental Medicine and Biology|isbn=978-1-4939-0819-6}}</ref> Another concern is the introduction of unknown pathogens if de-extinction efforts were to succeed.<ref>{{Cite web|date=2021-09-17|title=Woolly mammoths are being brought back from extinction by scientists|url=https://www.euronews.com/green/2021/09/17/why-are-scientists-bringing-woolly-mammoths-back-from-extinction|access-date=2021-09-19|website=euronews|language=en}}</ref> In 2021, an Austin-based company raised funds to reintroduce the species in the [[Arctic tundra]].<ref>{{Cite web|last=Carlson|first=Kara|title=Could Austin entrepreneur's company help bring back the woolly mammoth?|url=https://www.statesman.com/story/business/2021/09/17/austin-entrepreneur-aims-bring-back-woolly-mammoths-within-decade/8278799002/|access-date=2021-09-19|website=Austin American-Statesman}}</ref>

==Fossil specimens==
{{Further|List of mammoth specimens}}

[[File:Mammoth family.jpg|thumb|left|Mounted "family group" from [[Tomsk]]]]
Woolly mammoth fossils have been found in many different types of deposits, including former rivers and lakes, and in "[[Doggerland]]" in the [[North Sea]], which was dry at times during the ice age. Such fossils are usually fragmentary and contain no soft tissue. Accumulations of modern elephant remains have been termed "[[elephants' graveyard]]s", as these sites were erroneously thought to be where old elephants went to die. Similar accumulations of woolly mammoth bones have been found;  these are thought to be the result of individuals dying near or in the rivers over thousands of years, and their bones eventually being brought together by the streams. Some accumulations are thought to be the remains of herds that died together at the same time, perhaps due to flooding. Natural traps, such as [[kettle holes]], [[sink holes]], and mud, have trapped mammoths in separate events over time.<ref name="Lister 2007 45 75"/>

[[File:200805231215 Wollhaarmammut Millie Schädel.jpeg|thumb|Skull discovered by fishermen in the [[North Sea]] ("[[Doggerland]]"), at Celtic and Prehistoric Museum, Ireland]]
Apart from frozen remains, the only soft tissue known is from a specimen that was preserved in a [[petroleum seep]] in [[Starunia]], Poland. Frozen remains of woolly mammoths have been found in the northern parts of Siberia and Alaska, with far fewer finds in the latter. Such remains are mostly found above the [[Arctic Circle]], in permafrost. Soft tissue apparently was less likely to be preserved between 30,000 and 15,000 years ago, perhaps because the climate was milder during that period. Most specimens have partially degraded before discovery, due to exposure or to being scavenged. This "[[natural mummification]]" required the animal to have been buried rapidly in liquid or semisolids such as silt, mud, and icy water, which then froze.<ref name="Lister, 2007. pp. 50–53">Lister, 2007. pp.&nbsp;50–53</ref>

The presence of undigested food in the stomach and seed pods still in the mouth of many of the specimens suggests neither starvation nor exposure is likely. The maturity of this ingested vegetation places the time of death in autumn rather than in spring, when flowers would be expected.<ref name="Pfizenmayer 1939">{{cite book |title=Siberian Man and Mammoth |last=Pfizenmayer |first=E. W. |year=1939 | pages= 46–61 | publisher=Blackie and Son |location=London}}</ref> The animals may have fallen through ice into small ponds or potholes, entombing them. Many are certainly known to have been killed in rivers, perhaps through being swept away by floods. In one location, by the [[Byoryolyokh River]] in [[Sakha Republic|Yakutia]] in Siberia, more than 8,000 bones from at least 140 mammoths have been found in a single spot, apparently having been swept there by the current.<ref>{{Cite journal | last1 = Vereshchagin | first1 = N. K. | title = The mammoth "cemeteries" of north-east Siberia | doi = 10.1017/S0032247400031296 | journal = Polar Record | volume = 17 | issue = 106 | pages = 3–12 | year = 2009 | s2cid = 129654739 }}</ref>

===Frozen specimens===

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|image1 = Mammuthus primigenius.jpg
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|image2 = Adams mammoth, Vienna.jpg
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|footer = The "Adams mammoth" as illustrated in the 1800s (left) and on exhibit in Vienna; skin can be seen on its head and feet.
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Between 1692 and 1806, a handful of reports of frozen mammoth remains with soft tissue were published reached Europe, though none were collected during that time.<ref name="Tolmachoff1929"/> While frozen woolly mammoth carcasses had been excavated by Europeans as early as 1728, the first fully documented specimen was discovered near the delta of the [[Lena River]] in 1799 by Ossip Schumachov, a Siberian hunter.<ref>{{Cite journal
 | last = Adams
 | first = M.
 | title = Some Account of a Journey to the Frozen-Sea, and of the Discovery of the Remains of a Mammoth
 | journal = The Philadelphia Medical and Physical Journal
 | volume = 3
 | pages = 120–137
 | year = 1808 }}</ref> While in Yakutsk in 1806, [[Johann Friedrich Adam|Michael Friedrich Adams]] heard about the frozen mammoth. Adams recovered the entire skeleton, apart from the tusks, which Shumachov had already sold, and one foreleg, most of the skin, and nearly {{cvt|18|kg}} of hair. During his return voyage, he purchased a pair of tusks that he believed were the ones that Shumachov had sold. Adams brought all to the [[Zoological Museum of the Zoological Institute of the Russian Academy of Sciences]], and the task of mounting the skeleton was given to [[Wilhelm Gottlieb Tilesius von Tilenau|Wilhelm Gottlieb Tilesius]].<ref name="ans"/><ref>{{Cite journal
 | last = Tilesio
 | first = W. G.
 | title = De skeleto mammonteo Sibirico ad maris glacialis littora anno 1807 effosso, cui praemissae Elephantini generis specierum distinctiones
 | journal = Mémoires de l'Académie Impériale des Sciences de St. Pétersbourg
 | volume = 5
 | pages = 406–514
 | language = la
 | year = 1815 }}</ref> This was one of the first attempts at reconstructing the skeleton of an extinct animal. Most of the reconstruction is correct, but Tilesius placed each tusk in the opposite socket, so that they curved outward instead of inward. The error was not corrected until 1899, and the correct placement of mammoth tusks was still a matter of debate into the 20th century.<ref name = "Cohen">{{cite book
| last = Cohen
| first = C.
| year = 2002
| page = [https://archive.org/details/fateofmammothfos0000cohe/page/113 113]
| title = The Fate of the Mammoth: Fossils, Myth, and History
| publisher = University of Chicago Press
| isbn = 978-0-226-11292-3
| url = https://archive.org/details/fateofmammothfos0000cohe/page/113
}}</ref><ref>{{Cite journal
 | last = Pfizenmayer
 | first = E.
 | title = A Contribution to the Morphology of the Mammoth, ''Elephas Primigenius'' Blumenbach; With an Explanation of My Attempt at a Restoration
 | journal = Annual Report of the Board of Regents of the Smithsonian Institution
 | pages = 326–334
 | year = 1907 }}</ref>

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|image1 = A Berezovka folyó völgye, mamutlelet. Balra a lelet felfedezője, Eugene W. Pfitzenmayer régész. Fortepan 95087.jpg
|alt1 =

|image2 = Мамонт.jpg
|alt2 =

|footer = The "Berezovka mammoth" during excavation in 1901 (left), and a model partially covered by its skin, [[Museum of Zoology in St. Petersburg]]
}}

The 1901 excavation of the "Berezovka mammoth" is the best documented of the early finds. It was discovered at the Siberian [[Berezovka River]] (after a dog had noticed its smell), and the Russian authorities financed its excavation. The entire expedition took 10 months, and the specimen had to be cut to pieces before it could be transported to St. Petersburg. Most of the skin on the head as well as the trunk had been scavenged by predators, and most of the internal organs had rotted away. It was identified as a 35- to 40-year-old male, which had died 35,000 years ago. The animal still had grass between its teeth and on the tongue, showing that it had died suddenly. One of its shoulder blades was broken, which may have happened when it fell into a crevasse. It may have died of [[asphyxiation]], as indicated by its erect penis. One third of a replica of the mammoth in the Museum of Zoology of [[Saint Petersburg|St. Petersburg]] is covered in skin and hair of the "Berezovka mammoth".<ref name="Lister, 2007. pp. 50–53"/><ref name="Pfizenmayer 1939"/>

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|image1 = Mammoth hair and skin, leg.jpg
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|image2 = Mammoth leg.png
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|image3 = Mummified Mammuthus primigenius (wooly mammoth) "Effie" (cropped).jpg
|alt3 =

|footer = Leg with skin and fur at the [[National Museum of Natural History, France]] (left), a stuffed leg (middle), and the calf "Effie" at the [[American Museum of Natural History]] (right)
}}

By 1929, the remains of 34 mammoths with frozen soft tissues (skin, flesh, or organs) had been documented. Only four of them were relatively complete. Since then, about that many more have been found. In most cases, the flesh showed signs of decay before its freezing and later desiccation.<ref>{{Cite journal | last1 = Farrand | first1 = W. R. | title = Frozen Mammoths and Modern Geology: The death of the giants can be explained as a hazard of tundra life, without evoking catastrophic events | doi = 10.1126/science.133.3455.729 | journal = Science | volume = 133 | issue = 3455 | pages = 729–735 | year = 1961 | pmid =  17777646|bibcode = 1961Sci...133..729F }}</ref> Since 1860, Russian authorities have offered rewards of up to {{RUB|1000}} for finds of frozen woolly mammoth carcasses. Often, such finds were kept secret due to superstition. Several carcasses have been lost because they were not reported, and one was fed to dogs.<ref name="Lister 2007 45 75"/> Despite the rewards, native [[Yakuts]] were also reluctant to report mammoth finds to the authorities due to bad treatment of them in the past.<ref name="Tolmachoff1929"/> In more recent years, scientific expeditions have been devoted to finding carcasses instead of relying solely on chance encounters. The most famous frozen specimen from Alaska is a calf nicknamed "Effie", which was found in 1948. It consists of the head, the trunk, and a foreleg and is about 25,000 years old.<ref name="Lister 2007 45 75">Lister, 2007. pp.&nbsp;45–75</ref>

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|image2 = Dima the woolly mammoth calf.jpg
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|footer = "Dima", a frozen calf, during excavation (left), and as exhibited in the Museum of Zoology; note fur on the legs
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In 1977, the well-preserved carcass of a seven- to eight-month-old woolly mammoth calf named "Dima" was discovered. This carcass was recovered near a tributary of the [[Kolyma River]] in northeastern Siberia. This specimen weighed about {{cvt|100|kg}} at death and was {{cvt|104|cm}} high and {{cvt|115|cm}} long. [[Radiocarbon dating]] determined that "Dima" died about 40,000 years ago. Its internal organs are similar to those of modern elephants, but its ears are only one-tenth the size of those of an African elephant of similar age. A less complete juvenile, nicknamed "Mascha", was found on the [[Yamal Peninsula]] in 1988. It was 3–4 months old, and a laceration on its right foot may have been the cause of death. It is the westernmost frozen mammoth found.<ref>Lister, 2007. pp.&nbsp;57–58</ref>

In 1997, a piece of mammoth tusk was discovered protruding from the tundra of the [[Taymyr Peninsula]] in Siberia, Russia. In 1999, this 20,380-year-old carcass and 25 tons of surrounding sediment were transported by an [[Mil Mi-26|Mi-26]] heavy lift helicopter to an ice cave in [[Khatanga, Russia|Khatanga]]. The specimen was nicknamed the "[[Jarkov Mammoth|Jarkov mammoth]]". In October 2000, the careful defrosting operations in this cave began with the use of [[hair dryer]]s to keep the hair and other soft tissues intact.<ref>Mol, D. et al. (2001). "The Jarkov Mammoth: 20,000-Year-Old carcass of a Siberian woolly mammoth ''Mammuthus primigenius'' (Blumenbach, 1799)". ''The World of Elephants, Proceedings of the 1st International Congress'' ( 16–20 October 2001, Rome): 305–309. [https://web.archive.org/web/20060427110530/http://www.cq.rm.cnr.it/elephants2001/pdf/305_309.pdf Full text pdf]</ref><ref>{{Cite journal | last1 = Debruyne | first1 = R. G. | last2 = Barriel | first2 = V. R. | last3 = Tassy | first3 = P. | doi = 10.1016/S1055-7903(02)00292-0 | title = Mitochondrial cytochrome b of the Lyakhov mammoth (Proboscidea, Mammalia): New data and phylogenetic analyses of Elephantidae | journal = Molecular Phylogenetics and Evolution | volume = 26 | issue = 3 | pages = 421–434 | year = 2003 | pmid =  12644401| bibcode = 2003MolPE..26..421D }}</ref>

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|image2 = Picture of Lyuba taken on 3rd May 2012 at the IFC Mall Hong Kong.jpg
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|footer = The calf "Lyuba", in [[Royal BC Museum]] and [[IFC Mall (Hong Kong)|IFC Mall]]
}}

In 2002, a well-preserved carcass was discovered near the Maxunuokha River in northern Yakutia, which was recovered during three excavations. This adult male specimen was called the "Yukagir mammoth" and is estimated to have lived around 18,560 years ago, been {{cvt|2.829|m}} tall at the shoulder, and weighed between 4 and 5 tonnes. It is one of the best-preserved mammoths ever found due to the almost complete head, covered in skin, but without the trunk. Some postcranial remains were found, some with soft tissue.<ref name="Yukagir"/>

In 2007, the carcass of a female calf nicknamed "Lyuba" was discovered near the [[Yuribey Bridge|Yuribey River]], where it had been buried for 41,800 years.<ref name="Fisher et al., 2012" /><ref name="Kosintsev et al., 2012">{{Cite journal | last1 = Kosintsev | first1 = P. A. | last2 = Lapteva | first2 = E. G. | last3 = Trofimova | first3 = S. S. | last4 = Zanina | first4 = O. G. | last5 = Tikhonov | first5 = A. N. | last6 = Van Der Plicht | first6 = J. | doi = 10.1016/j.quaint.2011.03.027 | title = Environmental reconstruction inferred from the intestinal contents of the Yamal baby mammoth Lyuba (''Mammuthus primigenius'' Blumenbach, 1799) | journal = Quaternary International | volume = 255 | pages = 231–238 | year = 2012 |bibcode = 2012QuInt.255..231K | s2cid = 129303118 | url = https://pure.rug.nl/ws/files/6776258/2012QuatIntKosintsev.pdf }}</ref> By cutting a section through a molar and analysing its growth lines, they found that the animal had died at the age of one month.<ref name="Rountrey et al., 2012">{{Cite journal | last1 = Rountrey | first1 = A. N. | last2 = Fisher | first2 = D. C. | last3 = Tikhonov | first3 = A. N. | last4 = Kosintsev | first4 = P. A. | last5 = Lazarev | first5 = P. A. | last6 = Boeskorov | first6 = G. | last7 = Buigues | first7 = B. | title = Early tooth development, gestation, and season of birth in mammoths | doi = 10.1016/j.quaint.2011.06.006 | journal = Quaternary International | volume = 255 | pages = 196–205 | year = 2012 |bibcode = 2012QuInt.255..196R }}</ref> The mummified calf weighed {{cvt|50|kg}}, was {{cvt|85|cm}} high and {{cvt|130|cm}} in length.<ref name="BBC-Mammoth-2007">{{cite news | first=P. | last=Rincon | title=Baby mammoth discovery unveiled | date=10 July 2007 | publisher=BBC News | url=http://news.bbc.co.uk/1/hi/sci/tech/6284214.stm | work=news.bbc.co.uk | access-date=13 July 2007 | url-status=live | archive-url=https://web.archive.org/web/20070811210738/http://news.bbc.co.uk/1/hi/sci/tech/6284214.stm | archive-date=11 August 2007}}</ref><ref name="Reuters-Mammoth-2007">{{cite news | first=D. | last=Solovyov | title=Baby mammoth find promises breakthrough | date=11 July 2007 | publisher=Reuters | url=https://www.reuters.com/article/scienceNews/idUSL1178205120070711 | work=reuters.com | access-date=13 July 2007 | url-status=live | archive-url=https://web.archive.org/web/20070713175239/http://www.reuters.com/article/scienceNews/idUSL1178205120070711 | archive-date=13 July 2007}}</ref> At the time of discovery, its eyes and trunk were intact and some fur remained on its body. Its organs and skin are very well preserved.<ref>{{cite news | url=http://www.nydailynews.com/news/us_world/2009/04/21/2009-04-21_baby_mammoth_lyuba_pristinely_preserved_offers_scientists_rare_look_into_mysteri.html | location=New York | work=Daily News | title=Baby mammoth Lyuba, pristinely preserved, offers scientists rare look into mysteries of Ice Age | first=O. | last=Smith | date=21 April 2009 | url-status=live | archive-url=https://web.archive.org/web/20090815122500/http://www.nydailynews.com/news/us_world/2009/04/21/2009-04-21_baby_mammoth_lyuba_pristinely_preserved_offers_scientists_rare_look_into_mysteri.html | archive-date=15 August 2009}}</ref> "Lyuba" is believed to have been suffocated by mud in a river that its herd was crossing.<ref name="Fisher et al., 2012">{{Cite journal | last1 = Fisher | first1 = D. C. | last2 = Tikhonov | first2 = A. N. | last3 = Kosintsev | first3 = P. A. | last4 = Rountrey | first4 = A. N. | last5 = Buigues | first5 = B. | last6 = Van Der Plicht | first6 = J. | doi = 10.1016/j.quaint.2011.05.040 | title = Anatomy, death, and preservation of a woolly mammoth (''Mammuthus primigenius'') calf, Yamal Peninsula, northwest Siberia | journal = Quaternary International | volume = 255 | pages = 94–105 | year = 2012 |bibcode = 2012QuInt.255...94F | hdl = 11370/a3961dcc-4eaf-47fb-9ad7-904d79a0f4f8 | s2cid = 35667021 | url = https://pure.rug.nl/ws/files/6776236/2012QuatIntFisher.pdf | hdl-access = free }}</ref><ref>{{cite journal | doi = 10.1666/13-092 | title=X-ray computed tomography of two mammoth calf mummies | journal=Journal of Paleontology | date=2014 | volume=88 | issue=4 | pages=664–675 | first=Daniel C. | last=Fisher| bibcode=2014JPal...88..664F | s2cid=28393815 }}</ref> After death, its body may have been colonised by bacteria that produce [[lactic acid]], which "pickled" it, preserving the mammoth in a nearly pristine state.<ref name="Fisher et al., 2012" />

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|image2 = Yuka (Mammuthus primigenius) - Skull and Lower Jaw A.JPG
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|footer = The frozen calf "Yuka" (left), and its skull and jaw which may have been extracted from the carcass by prehistoric humans
}}

In 2010, a juvenile nicknamed it "[[Yuka (mammoth)|Yuka]]" was found in Siberia, the first known adolescent female, estimated to have been 2.5 years. It had man-made cut marks, and its skull and pelvis had been removed prior to discovery, but were found nearby.<ref name=bbcyuka>{{cite news | url = http://www.bbc.co.uk/nature/17525070 | title = Woolly mammoth carcass may have been cut into by humans | first = B. | last = Aviss | publisher = BBC | date = 4 April 2012 | access-date = 9 April 2012 | url-status = live | archive-url = https://web.archive.org/web/20120406185744/http://www.bbc.co.uk/nature/17525070 | archive-date = 6 April 2012}}</ref><ref>{{cite journal|last1=Mashchenko|first1=E. N.|last2=Protopopov|first2=A. V.|last3=Plotnikov|first3=V. V.|last4=Pavlov|first4=I. S.|title=Specific characters of the mammoth calf (''Mammuthus primigenius'') from the Khroma River (Yakutia)|journal=Biology Bulletin|volume=40|issue=7|year=2013|pages=626–641|doi=10.1134/S1062359013070042|bibcode=2013BioBu..40..626M |s2cid=16675371}}</ref><ref name="YukaHistory">{{cite journal |last1=Boeskorov |first1=G. G. |last2=Protopopov |first2=A. V. |last3=Maschenko |first3=E. N. |last4=Potapova |first4=O. R. |last5=Plotnikov |first5=V. V. |last6=Shchelchkova |first6=M. V. |last7=Pavlov |first7=I. S. |last8=Klimovsky |first8=A. I. |last9=Kolesov |first9=S. D. |last10=Gorokhov |first10=G. V. |title=History of Studies of the Female Woolly Mammoth Mummy Yuka (Mammuthus primigenius (Blumenbach, 1799)) |journal=Paleontological Journal |date=2021 |volume=55 |issue=11 |pages=1215–1223 |doi=10.1134/S0031030121110022|bibcode=2021PalJ...55.1215B }}</ref> After being discovered, the skin of "Yuka" was prepared to produce a [[taxidermy]] mount.<ref name="Yuka trunk"/> In 2019, a group of researchers managed to obtain signs of biological activity after transferring nuclei of "Yuka" into mouse [[oocytes]].<ref name="biologicalactivities">{{cite journal |last1=Yamagata |first1=K. |last2=Nagai |first2=K. |last3=Miyamoto |first3=H. |last4=Anzai |first4=M. |last5=Kato |first5=H. |last6=Miyamoto |first6=K. |last7=Kurosaka |first7=S. |last8=Azuma |first8=R. |last9=Kolodeznikov |first9=I. I. |last10=Protopopov |first10=A. V. |last11=Plotnikov |first11=V. V. |last12=Kobayashi |first12=H. |last13=Kawahara-Miki |first13=R. |last14=Kono |first14=T. |last15=Uchida |first15=M. |last16=Shibata |first16=Y. |last17=Handa |first17=T. |last18=Kimura |first18=H. |last19=Hosoi |first19=Y. |last20=Mitani |first20=T. |last21=Matsumoto |first21=K. |last22=Iritani |first22=A. |title=Signs of biological activities of 28,000-year-old mammoth nuclei in mouse oocytes visualized by live-cell imaging |journal=Scientific Reports |date=2019 |volume=9 |issue=1 |pages=4050 |doi=10.1038/s41598-019-40546-1|pmid=30858410 |pmc=6411884 |bibcode=2019NatSR...9.4050Y }}</ref>

In 2013, a well-preserved carcass was found on [[Maly Lyakhovsky Island]], one of the islands in the [[New Siberian Islands]] archipelago, a female between 50 and 60 years old at the time of death. The carcass contained well-preserved muscular tissue. When it was extracted from the ice, liquid blood spilled from the abdominal cavity. The finders interpreted this as indicating woolly mammoth blood possessed antifreezing properties.<ref>{{Cite journal | last1 = Wong | first1 = K. | doi = 10.1038/nature.2013.13103 | title = Can a mammoth carcass really preserve flowing blood and possibly live cells? | journal = Nature | year = 2013 | s2cid = 87298066 }}</ref> In 2022, a complete female baby woolly mammoth was found by a miner in the [[Klondike, Yukon|Klondike]] gold fields of [[Yukon]], Canada. The specimen is estimated to have died 30,000 years ago and was nicknamed "Nun cho ga", meaning "big baby animal" in the local [[Hän language]]. It is the best preserved woolly mammoth mummy found in North America, and was the same size as  Lyuba.<ref name="Nun cho ga">{{cite news |last1=Reardon |first1=Sophie |title=Rare mummified baby woolly mammoth with skin and hair found in Canada |url=https://www.cbsnews.com/news/woolly-mammoth-found-yukon-canada/ |access-date=26 June 2022 |work=www.cbsnews.com |date=2022}}</ref><ref>{{cite news |url=https://www.cbc.ca/news/canada/north/frozen-whole-baby-woolly-mammoth-yukon-gold-fields-1.6501128 | title='She's perfect and she's beautiful': Frozen baby woolly mammoth discovered in Yukon gold fields | work=[[Canadian Broadcasting Corporation]] |access-date=8 November 2022 | first1=Michel | last1=Proulx | date=24 June 2022 }}</ref> In 2025, a 50,000-year-old frozen calf nicknamed "Yana", after the Yana River basin of Yakutia where it was found, was announced. It was found by local residents and was described as the "best preserved" mammoth carcass, since it was found before it could be eaten by animals.<ref name="YanaBBC">{{cite news |last1=Smith |first1=Alex |title=Scientists unveil 50,000-year-old baby mammoth carcass |url=https://www.bbc.com/news/articles/cy47xj4lpyzo |access-date=18 January 2025 |work=www.bbc.com |agency=BBC |date=2025}}</ref>

==Cultural significance==

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|image1 = Carved 19th century Eskimo mammoth tusk De Young Museum 34430 2.JPG
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|image2 = Carved 19th century Eskimo mammoth tusk De Young Museum 34430 detail 1.JPG
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|footer = A mammoth tusk with [[Inuit]] carvings of scenes on the [[Yukon River]], 19th century, [[De Young Museum]]
}}

The woolly mammoth has remained culturally significant long after its extinction. Indigenous peoples of Siberia had long found what is now known to have been woolly mammoth remains, collecting their tusks for the [[ivory trade]], and had a diverse range of mythological interpretations of mammoth remains.<ref name="Serikov-2005">{{Cite journal |last1=Serikov |first1=Iu.B. |last2=Serikova |first2=A.Iu. |date=April 2005 |title=The Mammoth in the Myths, Ethnography, and Archeology of Northern Eurasia |url=https://www.tandfonline.com/doi/full/10.1080/10611959.2005.11029015 |journal=Anthropology & Archeology of Eurasia |language=en |volume=43 |issue=4 |pages=8–18 |doi=10.1080/10611959.2005.11029015 |issn=1061-1959|url-access=subscription }}</ref><ref>Newcomb, Raymond Lee (1888). ''[https://archive.org/details/ourlostexplorers01newc Our lost explorers: the narrative of the Jeannette Arctic Expedition as related by the survivors, and in the records and last journals of Lieutenant De Long] {{webarchive|url=https://web.archive.org/web/20160317100908/https://archive.org/details/ourlostexplorers01newc |date=17 March 2016 }}''. p. 96</ref><ref>{{citation|first=S.|last=Patkanov|title=Die lrtysch-Ostjaken und ihre Volkspoesie|volume=I|pages=123–124|place=St. Petersburg|year=1897|url=http://babel.hathitrust.org/cgi/pt?id=njp.32101073232413;view=1up;seq=137|archive-url=https://web.archive.org/web/20181107011842/https://babel.hathitrust.org/cgi/pt?id=njp.32101073232413;view=1up;seq=137|url-status=dead|archive-date=7 November 2018|publisher=St. Petersburg}}</ref> The [[Mansi people|Mansi]] and the  [[Khanty]] peoples conceived of mammoths as giant birds, with the Mansi (along with the [[Nenets]]) believing that they were responsible for the creation of mountains and lakes. In the mythology of the [[Evenks|Evenk]] people, mammoths were responsible for the world's creation, digging up the land from the ocean floor with their tusks. The [[Selkup people|Selkup]] believed that mammoths lived underground and guarded the [[underworld]], while the [[Yakuts]] regarded mammoths as water spirits.<ref name="Serikov-2005" />

The indigenous peoples of North America used woolly mammoth ivory and bone for tools and art.<ref>{{cite book |last= Cohen|first= C. |date= 2002 |title= The Fate of the Mammoth: Fossils, Myth, and History|url= https://archive.org/details/fateofmammothfos0000cohe |url-access= registration|publisher= University of Chicago Press |pages= [https://archive.org/details/fateofmammothfos0000cohe/page/197 197]–198|isbn= 978-0-226-11292-3 |access-date= 10 August 2015}}</ref> As in Siberia, North American natives had "myths of observation" explaining the remains of woolly mammoths and other elephants; the Bering Strait [[Inupiat]] believed the bones came from burrowing creatures, while other peoples associated them with primordial giants or "great beasts".<ref name=Stone>{{Cite journal | last1 = Strong | first1 = W. D. | doi = 10.1525/aa.1934.36.1.02a00060 | title = North American Indian traditions suggesting a knowledge of the mammoth | journal = American Anthropologist | volume = 36 | pages = 81–88 | year = 1934 | doi-access = free }}</ref><ref>{{cite journal | jstor=540573 | pages=294–296 | last1=Lankford | first1=G. E. | title=Pleistocene Animals in Folk Memory | volume=93 | issue=369 | journal=[[The Journal of American Folklore]] | year=1980 | doi=10.2307/540573 }} {{subscription required}}</ref><ref name=Mayor>{{cite book |title=Fossil Legends of the First Americans | last=Mayor | first=A. | year=2005 | publisher=Princeton University Press | location=Princeton | page=97 | isbn=978-0-691-11345-6}}</ref> Observers have interpreted legends from several Native American peoples as containing [[folk memory]] of extinct elephants, though other scholars are skeptical that folk memory could survive such a long time.<ref name=Stone/><ref name=Mayor/><ref>{{cite journal | jstor=540573 | pages=293–304 | last1=Lankford | first1=G. E. | title=Pleistocene Animals in Folk Memory | volume=93 | issue=369 | journal=[[The Journal of American Folklore]] | year=1980 | doi=10.2307/540573 }} {{subscription required}}</ref>

Woolly mammoth tusks had been articles of trade in Asia long before Europeans became acquainted with them. [[Güyük]], the 13th-century Khan of the Mongols, is reputed to have sat on a throne made from mammoth ivory.<ref name="Tolmachoff1929">{{cite journal | last1 = Tolmachoff | first1 = I. P. | year = 1929 | title = The carcasses of the mammoth and rhinoceros found in the frozen ground of Siberia | journal = Transactions of the American Philosophical Society | volume = 23 | issue = 1 | pages = 11–23 | jstor = 1005437 | doi = 10.2307/1005437 }}</ref> Inspired by the Siberian natives' concept of the mammoth as an underground creature, it was recorded in the 16th-century Chinese pharmaceutical encyclopedia, ''[[Ben Cao Gangmu]]'', as ''yin shu'', "the hidden rodent".<ref>{{citation|title=Arabic and Chinese Trade in Walrus and Narwhal Ivory
|first=Berthold|last= Laufer |author-link=Berthold Laufer |journal=T'oung Pao |series=Second Series |volume= 14|issue= 3 |year=1913  |page=329 |jstor=4526349|doi=10.1163/156853213X00213|hdl=2027/hvd.32044009725912|hdl-access=free}}</ref><ref>{{harvnb|Patkanov|1897|pages=123–124}}</ref> Remains of various extinct elephants were known by Europeans for centuries but were generally interpreted as the remains of [[legendary creatures]] such as [[behemoth]]s or [[Giant (mythology)|giants]], based on [[bible|biblical]] accounts. They were thought to be remains of modern elephants that had been brought to Europe during the [[Roman Republic]], for example the [[war elephants]] of [[Hannibal]] and [[Pyrrhus of Epirus]], or animals that had wandered north.<ref name="Switek">{{cite book | last = Switek | first = B.|year= 2010|title= Written in Stone: Evolution, the Fossil Record, and Our Place in Nature| publisher= Bellevue Literary Press | pages = 174–180 |isbn=978-1-934137-29-1}}</ref>

{{multiple image
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|image1 = Peter III of Russia (mammuthus).jpg
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|image2 = Elizabeth of Russia (mammuthus).jpg
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|footer = [[Peter III of Russia|Peter III]] and [[Elizabeth of Russia]] carved in mammoth ivory
}}

Siberian mammoth ivory is reported to have been exported to Russia and Europe in the 10th century. The first Siberian ivory to reach Western Europe was brought to London in 1611. When Russia occupied Siberia, the ivory trade grew and it became a widely exported commodity, with huge amounts being excavated. From the 19th century onwards, woolly mammoth ivory became a highly prized commodity, used as raw material for many products. Today, it is still in great demand as a replacement for the now-banned export of elephant ivory and has been referred to as "white gold".<ref name="Lister, 2007. pp. 137–139">Lister, 2007. pp.&nbsp;137–139</ref> Local dealers estimate that 10 million mammoths are still frozen in Siberia. Conservationists have proposed that these mammoths could help prevent the extinction of living elephant species. Elephants are often hunted by poachers for their ivory, but if the demand for ivory could be met by sourcing it from the already extinct mammoths, it might reduce the pressure on elephants. Trade in elephant ivory has been forbidden in most places following the 1989 [[Lausanne]] Conference, but dealers have been known to label it as mammoth ivory to get it through customs. Mammoth ivory looks similar to elephant ivory, but the former is browner, and the [[Schreger lines]] are coarser in texture.<ref name="Lister, 2007. pp. 137–139"/> In the 21st century, [[global warming]] has made access to Siberian tusks easier since the permafrost thaws more quickly, exposing the mammoths embedded within it.<ref name="Larmer2015">{{cite web|url=http://ngm.nationalgeographic.com/2013/04/mammoth-tusks/larmer-text|title=Mammoth Tusk Hunters|website=nationalgeographic.com|date=April 2013|last=Larmer|first=B.|url-status=dead|archive-url=https://web.archive.org/web/20130402004219/http://ngm.nationalgeographic.com/2013/04/mammoth-tusks/larmer-text|archive-date=2 April 2013}}</ref>

Stories abound about frozen woolly mammoth meat consumed once defrosted, especially that of the "Berezovka mammoth", but most of these are considered dubious. In most cases, the carcasses were decayed, and the stench so unbearable that only wild scavengers and the dogs accompanying the finders showed any interest in the flesh. Such meat apparently was once recommended against illness in China, and Siberian natives have occasionally cooked the meat of frozen carcasses they discovered.<ref name="Lister, 2007. p. 48-55"/> According to one of the more famous stories, members of the [[The Explorers Club|Explorers Club]] dined on the meat of a frozen mammoth from Alaska in 1951. In 2016, a group of researchers genetically examined a sample of the meal and found it to belong to a [[green sea turtle]] (it had also been claimed to belong to ''[[Megatherium]]''). The researchers concluded that the dinner had been a publicity stunt.<ref name="Dinner">{{cite journal |last1=Glass |first1=J. R. |last2=Davis |first2=M. |last3=Walsh |first3=T. J. |last4=Sargis |first4=E. J. |last5=Caccone |first5=A. |last6=Fiorillo |first6=A. |title=Was Frozen Mammoth or Giant Ground Sloth Served for Dinner at The Explorers Club? |journal=PLOS ONE |date=2016 |volume=11 |issue=2 |pages=e0146825 |doi=10.1371/journal.pone.0146825|pmid=26840445 |pmc=4740485 |bibcode=2016PLoSO..1146825G |doi-access=free }}</ref> In 2011, the Chinese palaeontologist Lida Xing [[livestreamed]] while eating meat from a Siberian mammoth leg (thoroughly cooked and flavoured with salt) and told his audience it tasted bad and like soil. This triggered controversy and gained mixed reactions, but Xing stated he did it to promote science.<ref>{{cite news |last1=Weiyun |first1=T. |title='Lucky Hands' in pursuit of dinosaurs |url=https://www.shine.cn/feature/art-culture/1812126571/ |access-date=4 July 2019 |work=SHINE |date=2011 }}</ref> In 2023, An Australian cultured meat start-up, Vow, revealed a lab-grown "mammoth meatball" produced using a DNA sequence from the woolly mammoth. The meatball sparked conversations about the potential of cultured meat as a sustainable food source, highlighting its environmental benefits compared to traditional agriculture.<ref>{{Cite web |last=Chun |first=Alex |title=This massive meatball was made With woolly mammoth DNA |url=https://www.smithsonianmag.com/smart-news/this-massive-meatball-was-made-with-woolly-mammoth-dna-180981908/ |access-date=2024-03-24 |website=Smithsonian Magazine |date=March 30, 2023}}</ref>

===Alleged survival===

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|image1 = Coat of Arms of Ust-Yansky rayon (Yakutia).png
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|image2 = Mezhirich cher gerb.png
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|image3 = Coat of Arms of Srednekolymsk (Yakutia).png
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|footer = Woolly mammoths represented on the [[coats of arms]] of regions in Russia and Ukraine
}}

There have been occasional claims that the woolly mammoth is not extinct and that small, isolated herds might survive in the vast and sparsely inhabited [[tundra]] of the [[Northern Hemisphere]]. In the 19th century, several reports of "large shaggy beasts" were passed on to the Russian authorities by Siberian tribesmen, but no scientific proof ever surfaced. A French ''[[chargé d'affaires]] ''working in [[Vladivostok]], M. Gallon, said in 1946 that in 1920, he had met a Russian fur-trapper who claimed to have seen living giant, furry "elephants" deep into the [[taiga]].<ref name="Sjögren1962"/> Due to the large area of Siberia, the possibility that woolly mammoths survived into more recent times cannot be completely ruled out, but evidence indicates that they became extinct thousands of years ago. These natives had likely gained their knowledge of woolly mammoths from carcasses they encountered, which is likely the source of their legends of the animal.<ref name="Lister, 2007. p. 48-55"/>

In the late 19th century, rumours existed about surviving mammoths in Alaska.<ref name="Sjögren1962">{{cite book| last1 = Sjögren| first1 = B.|year= 1962|language= sv|page = 168 |title= Farliga djur och djur som inte finns|publisher= Prisma}}</ref> In 1899, Henry Tukeman detailed his killing of a mammoth in Alaska and his subsequent donation of the specimen to the [[Smithsonian Institution]] in [[Washington, DC]]. The museum denied the story.<ref>{{cite web |url=http://www.tacomapubliclibrary.org/morgan/Tukeman.htm |title=Henry Tukeman: Mammoth's Roar was Heard All The Way to the Smithsonian |publisher=Tacoma Public Library |year=1960 |access-date=17 January 2008 |last=Murray |first=M. |url-status=dead |archive-url=https://web.archive.org/web/20120118060303/http://www.tacomapubliclibrary.org/morgan/Tukeman.htm |archive-date=18 January 2012 }}</ref> The Swedish writer Bengt Sjögren suggested in 1962 that the myth began when the American biologist [[Charles Haskins Townsend]] travelled in Alaska, saw [[Inuit]] trading mammoth tusks, asked if mammoths were still living in Alaska, and provided them with a drawing of the animal.<ref name="Sjögren1962"/> [[Bernard Heuvelmans]] included the possibility of residual populations of Siberian mammoths in his 1955 book, ''On The Track Of Unknown Animals''; while his book was a systematic investigation into possible unknown species, it became the basis of the [[cryptozoology]] movement.<ref>{{Cite news |title= The Big Question: Are so-called 'extinct' species really extinct, and will we rediscover any? |first= Michael |last= McCarthy |date= 28 September 2009 |work= The Independent |url= https://www.independent.co.uk/environment/nature/the-big-question-are-so-called-extinct-species-really-extinct-and-will-we-rediscover-any-1794282.html |access-date= 5 August 2019}}</ref>

==References==
{{Reflist|30em}}

===Bibliography===
* {{cite book | last1 = Lister | first1 = A. | last2 = Bahn| first2 = P.| title = Mammoths – Giants of the Ice Age| publisher = Frances Lincoln | year = 2007 | edition = 3 | location = London | isbn = 978-0-520-26160-0 | oclc= 30155747 |ref=none}}
* {{cite book|last= Shapiro|first= Beth|title= How to Clone a Mammoth: The Science of De-Extinction|url= https://archive.org/details/howtoclonemammot0000shap|url-access= registration|date=5 April 2015|publisher= Princeton University Press|isbn=978-1-4008-6548-2|oclc= 965621402|ref=none}}

==External links==
{{Commons category|Mammuthus primigenius|position=right}}
{{Portal|Paleontology}}
* {{Wikispecies-inline|Mammuthus primigenius}}
*[https://www.youtube.com/watch?v=a2744xoqRKo Natural History Museum: "The last of the mammoths"] – three-minute video about the extinction of the woolly mammoth, presented by Adrian Lister
*[https://www.youtube.com/watch?v=GM_W6ncz-R0 National Geographic: "Mammoth tusk treasure hunt"] – two-minute video about mammoth tusk collecting in modern Siberia

{{Taxonbar|from=Q3699044}}
{{Authority control}}

{{DEFAULTSORT:Woolly Mammoth}}
[[Category:Mammoths]]
[[Category:Pleistocene proboscideans]]
[[Category:Pleistocene first appearances]]
[[Category:Pleistocene mammals of North America]]
[[Category:Fauna of the Holarctic realm]]
[[Category:Holocene extinctions]]
[[Category:Extinct animals of Asia]]
[[Category:Extinct animals of the United States]]
[[Category:Extinct animals of Canada]]
[[Category:Extinct mammals of Europe]]
[[Category:Extinct mammals of North America]]
[[Category:Fossil taxa described in 1799]]
[[Category:Cenozoic animals of North America]]
[[Category:Taxa named by Johann Friedrich Blumenbach]]
[[Category:Symbols of Alaska]]
[[Category:Symbols of Vermont]]
[[Category:Taxa with lost type specimens]]
[[Category:Species that are or were threatened by climate change]]
[[Category:Species that are or were threatened by human consumption]]