Mammoth: Difference between revisions

| fossil_range = Late [[Miocene]] to Late [[Holocene]], {{fossilrange|6.2|0.0037004}}

A '''mammoth''' is any [[species]] of the [[extinct]] [[elephantidae|elephantid]] [[genus]] '''''Mammuthus'''.'' They lived from the late [[Miocene]] epoch (from around 6.2 million years ago) into the [[Holocene]] about 4,000 years ago, and various species existed in Africa, Europe, Asia, and North America. Mammoths are distinguished from living [[elephants]] by their (typically large) spirally twisted tusks and in at least some later species, the development of numerous adaptions to living in cold environments, including a thick layer of fur.

Mammoths and [[Asian elephant]]s are more closely related to [[Asianeach elephant]]sother than either of them are to [[African elephant]]s. The oldest mammoth representative, ''[[Mammuthus subplanifrons]]'', appeared around 6 million years ago during the late Miocene in what is now southern and Eastern Africa.''<ref name=":22">{{Cite book |last=Sanders |first=William J. |url=https://www.taylorfrancis.com/books/9781315118918 |title=Evolution and Fossil Record of African Proboscidea |date=2023-07-07 |publisher=CRC Press |isbn=978-1-315-11891-8 |edition=1 |location=Boca Raton |pages=155, 208–212 |language=en |doi=10.1201/b20016|s2cid=259625811 }}</ref>'' Later in the [[Pliocene]], by about three million years ago, mammoths dispersed into Eurasia, eventually covering most of Eurasia before migrating into North America around 1.5–1.3&nbsp;million years ago, becoming ancestral to the [[Columbian mammoth]] (''M. columbi''). The [[woolly mammoth]] (''M. primigenius''), evolved about 700–400,000 years ago in Siberia, with some surviving on Russia's [[Wrangel Island]] in the [[Arctic Ocean]] until as recently as roughly 3,700 to 4,000 years ago, still extant during the existence of the earliest civilisations in [[ancient Egypt]] and [[Mesopotamia]].

According to the ''[[The American Heritage Dictionary]],'' the word "mammoth" likely originates from *mān-oŋt, a word in the [[Mansi languages|Mansi language]] wordof *mān-oŋtwestern Siberia meaning "earth horn", in reference to mammoth tusks.<ref>“[https://www.ahdictionary.com/word/search.html?q=mammoth mammoth]”, in ''[[The American Heritage Dictionary of the English Language]]'', 5th edition, Boston, Mass.: [[Houghton Mifflin Harcourt]], 2016, <small>→ISBN</small>.</ref> Mammoths appear in the folkore of the indigenous people of Siberia, who were impressed by the great size of their remains. In the mythology of the [[Evenks|Evenk]] people, mammoths were responsible for the creation of the world, 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 [[Nenets]] and the [[Mansi people|Mansi]] (the latter of whom, along with the [[Khanty]], conceived mammoths as giant birds) believed that mammoths were responsible for the creation of mountains and lakes, while the [[Yakuts]] regarded mammoths as water spirits.<ref name=":10">{{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}}</ref>

The word ''mammoth'' was first used in Europe during the early 17th century, when referring to ''maimanto'' tusks discovered in Siberia,<ref>{{cite book |last1=Lister |first1=A. |title=Mammoths – Giants of the Ice Age |last2=Bahn |first2=P. |date=2007 |publisher=Frances Lincoln |isbn=978-0-520-26160-0 |edition=3rd |location=London |page=49 |ref=Lister}}</ref> as recorded in the 1618 edition of the ''Dictionariolum Russico-Anglicum.''<ref>''"[https://www.oed.com/dictionary/mammoth_n Mammoth]" Oxford English Dictionary'' 2000</ref> The earliest scientific research paper on mammoths was by [[Vasily Tatishchev]] in 1725.<ref name=":10" /> [[John Bell (traveller)|John Bell]], who was on the [[Ob River]] in 1722, said that mammoth tusks were well known in the area. They were called "mammon's horn" and were often found in washed-out river banks. Some local people claimed to have seen a living mammoth, but they came out only at night and always disappeared under water when detected. Bell bought one and presented it to [[Hans Sloan]] who pronounced it an elephant's tooth.<ref>[[John Bell (traveller)|John Bell]], Travels from St Petersburg in Russia to diverse parts of Asia, Edinburgh, 1806, pages 383-386</ref>

The earliest known [[proboscideans]], the [[clade]] that contains the elephants, existedarose about 55 million years ago aroundon the [[Tethyslandmass Sea]]of areaAfro-Arabia. The closest relatives of the Proboscidea are the [[sirenians]] and the [[hyrax]]es. The family [[Elephantidae]] is known to have existedarose sixby million years ago in Africa, and includes the living elephants and the mammoths. Among many now extinct clades, the [[mastodon]] is only a distant relative of the mammoths, and part of the separate [[Mammutidae]] family, which diverged 25 million years before the mammoths evolved.<ref name="Lister 2007">{{cite book |ref=Lister|last1=Lister |first1=A. |last2=Bahn |first2=P. |date=2007 |title=Mammoths – Giants of the Ice Age |edition=3rd |publisher=Frances Lincoln |location=London |isbn=978-0-520-26160-0}}</ref>

Following the publication of the woolly mammoths [[mitochondrial genome]] sequence in 1997, it has since become widely accepted that mammoths and [[Asian elephants]] share a closer relationship to each other than either do to [[African elephant|African elephants]].<ref>{{Cite journal |last=Ozawa |first=Tomowo |last2=Hayashi |first2=Seiji |last3=Mikhelson |first3=Victor M. |date=April 1997 |title=Phylogenetic Position of Mammoth and Steller's Sea Cow Within Tethytheria Demonstrated by Mitochondrial DNA Sequences |url=http://link.springer.com/10.1007/PL00006160 |journal=Journal of Molecular Evolution |language=en |volume=44 |issue=4 |pages=406–413 |doi=10.1007/PL00006160 |issn=0022-2844}}</ref><ref name=":52" />

The following [[cladogram]] shows the placement of the genus ''Mammuthus'' among other proboscideans, based on [[hyoid]] characteristics and genetics:<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><ref name=":52">{{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=2018-03-13 |title=A comprehensive genomic history of extinct and living elephants |journal=Proceedings of the National Academy of Sciences |language=en |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>

The earliest mammoths, assigned to the species ''[[Mammuthus subplanifrons]]'', are known from southern and eastern Africa, with the earliest records dating to the Late [[Miocene]], around 6.2–5.3 million years ago.<ref name=":22" /> By the Late [[Pliocene]], mammoths had become confined to the northern portions of the African continent with remains from this time assigned to ''[[Mammuthus africanavus]].<ref name=":2">{{Cite book |last=Sanders |first=William J. |url=https://www.taylorfrancis.com/books/9781315118918 |title=Evolution and Fossil Record of African Proboscidea |date=2023-07-07 |publisher=CRC Press |isbn=978-1-315-11891-8 |edition=1 |location=Boca Raton |pages=245, 252, 263–266 |language=en |doi=10.1201/b20016|s2cid=259625811 }}</ref>'' During the Late Pliocene, by 3.2 million years ago, mammoths dispersed into Eurasia via the Sinai Peninsula. The earliest mammoths in Eurasia are assigned to the species ''[[Mammuthus rumanus]]''.<ref>{{Cite journal |last1=Iannucci |first1=Alessio |last2=Sardella |first2=Raffaele |date=2023-02-28 |title=What Does the "Elephant-Equus" Event Mean Today? Reflections on Mammal Dispersal Events around the Pliocene-Pleistocene Boundary and the Flexible Ambiguity of Biochronology |journal=Quaternary |language=en |volume=6 |issue=1 |pages=16 |doi=10.3390/quat6010016 |doi-access=free |issn=2571-550X|hdl=11573/1680082 |hdl-access=free }}</ref> The youngest remains of mammoths in Africa are from Aïn Boucherit, Algeria dating to the Early Pleistocene, around 2.3–2 million years ago (with a possible later record from Aïn Hanech, Algeria, dating to 1.95–1.78 million years ago).''<ref name=":2" />''

''Mammuthus rumanus'' is thought to be the ancestor of ''[[Mammuthus meridionalis]],'' which first appeared at the beginning of the Pleistocene, around 2.6 million years ago.<ref name=":3">{{Cite journal |last1=Lister |first1=Adrian M. |last2=Sher |first2=Andrei V. |last3=van Essen |first3=Hans |last4=Wei |first4=Guangbiao |date=January 2005 |title=The pattern and process of mammoth evolution in Eurasia |journal=Quaternary International |volume=126-128 |pages=49–64 |bibcode=2005QuInt.126...49L |doi=10.1016/j.quaint.2004.04.014 |issn=1040-6182|url=http://doc.rero.ch/record/13496/files/PAL_E277.pdf }}</ref> ''Mammuthus meridionalis'' subsequently gave rise to ''[[Mammuthus trogontherii]]'' (the steppe mammoth) in Eastern Asia around 1.7 million years ago. Around 1.5–1.3 million years ago, ''M. trogontherii'' crossed the [[Bering Land Bridge]] into North America, becoming ancestral to ''[[Mammuthus columbi]]'' (the Columbian mammoth).<ref>{{Cite journal |last1=Lister |first1=A. M. |last2=Sher |first2=A. V. |date=2015-11-13 |title=Evolution and dispersal of mammoths across the Northern Hemisphere |journal=Science |language=en |volume=350 |issue=6262 |pages=805–809 |bibcode=2015Sci...350..805L |doi=10.1126/science.aac5660 |issn=0036-8075 |pmid=26564853 |s2cid=206639522}}</ref> At the end of the Early Pleistocene ''Mammuthus trogontherii'' migrated into Europe, replacing ''M. meridionalis'' around 1–0.8 million years ago.<ref name=":3" /> ''[[Mammuthus primigenius]]'' (the woolly mammoth) had evolved from ''M. trogontherii'' in Siberia by around 600,000–500,000 years ago, replacing ''M. trogontherii'' in Europe by around 200,000 years ago, and migratingmigrated into North America during the Late Pleistocene.<ref name="Lister 107693">{{Cite journal |last=Lister |first=Adrian M. |date=October 2022 |title=Mammoth evolution in the late Middle Pleistocene: The Mammuthus trogontherii-primigenius transition in Europe |url=https://linkinghub.elsevier.com/retrieve/pii/S0277379122003249 |journal=Quaternary Science Reviews |language=en |volume=294 |pages=107693 |doi=10.1016/j.quascirev.2022.107693|bibcode=2022QSRv..29407693L |s2cid=252264887 }}</ref>

Like living elephants, mammoths typically had large body sizes. The largest known species like ''[[Mammuthus meridionalis]]'' and ''[[Mammuthus trogontherii]]'' (the steppe mammoth) were considerably larger than modern elephants, with malesmature reachingadult heightsmales inhaving thean regionaverage height of approximately {{convert|3.8-4.2|m|1|abbr=on}} at the shoulder and weights of {{convert|119.6-12.7|tonne|STlb|lk=on}}, while exceptionally large males may have reached {{convert|4.5|m|1|abbr=on}} at the shoulder and {{convert|14.3|tonne|STlb|1}} in weight.<ref name="probos_mass">{{Cite journal | last1 = Larramendi | first1 = A. | year = 2016 | title = Shoulder height, body mass and shape of proboscideans | journal = Acta Palaeontologica Polonica | volume = 61 | doi = 10.4202/app.00136.2014 | url = https://www.app.pan.pl/archive/published/app61/app001362014.pdf | doi-access = free }}</ref> However, woolly mammoths were considerably smaller, only about as large as modern [[African bush elephant]]s atwith aboutmales around {{convert|2.3 m to 80-3.15 |m|1|abbr=on}} high at the shoulder, and 2{{convert|4.8 to 5-6 tonnes|tonne|lb|lk=on}} in weight on average).<ref name="probos_mass" />,<ref name=":554">{{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 |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=2023-09-30}}</ref> with the largest recorded individuals being around {{convert|3.5|m|1|abbr=on}} tall and {{convert|8.2|tonne|lb|1}} in weight.<ref name="probos_mass" /> The insular dwarf mammoth species were considerably smaller, with the smallest species ''M. creticus'' estimated to have a shoulder height of only around {{Convert|1|m|ft}} and a weight of about {{Convert|180|kg|lb}}, making it one of the smallest elephantids known.<ref name="probos_mass" />

{{gallery|M._meridionalis_skeletal.png|Skeleton of a ''[[Mammuthus meridionalis]]'' bull, around {{convert|4|m|ft}} tall|M. trogontherii skeletal (cropped).png|Skeleton of a [[steppeSteppe mammoth]] (''MammuthusM. trogontherii'') around {{convert|3.9|m|ft}} tall in front-on (without head) side-on and top-down views|M._columbi_skeletals_(cropped).png|Skeleton of a [[Columbian mammoth]] (''MammuthusM. columbi'') bull around {{convert|3.7|m|ft}} tall|M. primigenius modified.png|Skeletons of two [[woollyWoolly mammoths]] (''M. primigenius''), including one of the largest, the Siegsdorf mammoth (left, around {{convert|3.5|m|ft}} tall), one of the largest woolly mammoth specimens, and a smaller mature Siberian bull (around {{convert|2.7|m|ft}} metres tall)|||||width=180185|height=|lines=|align=center|title=Gallery of mammoth skeletons}}[[File:Mammuthus meridionalis molar - Cleveland Museum of Natural History - 2014-12-26 (20267609313).jpg|thumb|Molar of the primitive mammoth ''[[Mammuthus meridionalis]]'']]

[[File:Woolly mammoth molar - Cleveland Museum of Natural History - 2014-12-26 (20859260938).jpg|thumb|Molar of a woolly mammoth]]The number of [[Elephantidae#Description|lamellae]] (ridge-like structures) on the molars, particularly on the third molars, substantially increased over the course of mammoth evolution. The earliest Eurasian species ''M. rumanus'' have around 8-10 lamellae on the third molars,<ref>{{Cite journal |last=Markov |first=Georgi N. |date=October 2012 |title=Mammuthus rumanus, early mammoths, and migration out of Africa: Some interrelated problems |url=https://linkinghub.elsevier.com/retrieve/pii/S1040618211003144 |journal=Quaternary International |language=en |volume=276-277 |pages=23–26 |bibcode=2012QuInt.276...23M |doi=10.1016/j.quaint.2011.05.041}}</ref> while Late Pleistocene woolly mammoths have 20-28 lamellae on the third molars. These changes also corresponded with reduced enamel thickness and increasing tooth height ([[hypsodont]]y).<ref name="Lister 107693" /> These changes are thought to be adaptations to increasing abrasion resulting from the shift in the diet of mammoths from a [[Browsing (herbivory)|browsing]] based diet in ''M. rumanus'', towards a [[Grazing (behaviour)|grazing]] diet in later species.<ref>{{Cite journal |last1=Lister |first1=Adrian M. |last2=Sher |first2=Andrei V. |date=2001-11-02 |title=The Origin and Evolution of the Woolly Mammoth |url=https://www.science.org/doi/10.1126/science.1056370 |journal=Science |language=en |volume=294 |issue=5544 |pages=1094–1097 |bibcode=2001Sci...294.1094L |doi=10.1126/science.1056370 |issn=0036-8075 |pmid=11691991 |s2cid=10662205}}</ref><ref name=":4">{{Cite journal |last1=Rivals |first1=Florent |last2=Semprebon |first2=Gina M. |last3=Lister |first3=Adrian M. |date=September 2019 |title=Feeding traits and dietary variation in Pleistocene proboscideans: A tooth microwear review |url=https://linkinghub.elsevier.com/retrieve/pii/S0277379119302641 |journal=Quaternary Science Reviews |language=en |volume=219 |pages=145–153 |bibcode=2019QSRv..219..145R |doi=10.1016/j.quascirev.2019.06.027 |s2cid=200073388}}</ref>

Both sexes bore tusks. A first, small set appeared at about the age of six months, and these were replaced at about 18 months by the permanent set. Growth of the permanent set was at a rate of about {{convert|2.5|to|15.2|cm|abbr=on|0}} per year.<ref>{{cite book|last1=Agenbroad|first1=Larry|last2=Nelson|first2=Lisa|title=Mammoths|publisher=Lerner|location=Minneapolis|isbn=978-0-8225-2862-3|page=[https://archive.org/details/mammothsiceagegi00larr/page/34 34]|year=2002|url=https://archive.org/details/mammothsiceagegi00larr/page/34}}</ref> The tusks display a strong spiral twisting.<ref>{{Cite journal |last1=Rabinovich |first1=Rivka |last2=Lister |first2=Adrian M. |date=July 2017 |title=The earliest elephants out of Africa: Taxonomy and taphonomy of proboscidean remains from Bethlehem |url=https://linkinghub.elsevier.com/retrieve/pii/S1040618216308370 |journal=Quaternary International |language=en |volume=445 |pages=23–42 |doi=10.1016/j.quaint.2016.07.010|bibcode=2017QuInt.445...23R }}</ref> Mammoth tusks are among the largest known among proboscideans with some specimens over {{convert|4|m|1|abbr=on}} in length and likely {{convert|200|kg|1|abbr=on}} in weight with some historical reports suggesting tusks of [[Columbian mammoths]] could reach lengths of around {{convert|5|m|1|abbr=on}} substantially surpassing the largest known modern elephant tusks.<ref>{{Cite journal |last=Larramendi |first=Asier |date=2023-12-10 |title=Estimating tusk masses in proboscideans: a comprehensive analysis and predictive model |url=https://www.tandfonline.com/doi/full/10.1080/08912963.2023.2286272 |journal=Historical Biology |language=en |pages=1–14 |doi=10.1080/08912963.2023.2286272 |s2cid=266182491 |issn=0891-2963}}</ref>[[File:Mammuthus meridionalis reconstruction 2.jpg|thumb|Life restoration of ''Mammuthus meridionalis''|left]]

The heads of mammoths were prominently domed.<ref name=":53">{{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 |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=2023-09-30 }}</ref> The first several [[thoracic vertebrae]] of mammoths typically had long neural spines.<ref>{{Cite journal |last=Larramendi |first=Asier |date=2014-02-16 |title=Skeleton of a Late Pleistocene steppe mammoth (Mammuthus trogontherii) from Zhalainuoer, Inner Mongolian Autonomous Region, China |url=http://link.springer.com/10.1007/s12542-014-0222-8 |journal=Paläontologische Zeitschrift |volume=89 |issue=2 |pages=229–250 |doi=10.1007/s12542-014-0222-8 |issn=0031-0220}}</ref> The back was typically sloping, with the body being wider than that of African elephants. The tails of mammoths were relatively short compared to living elephants.<ref name=":53" />[[File:Woolly mammoth model Royal BC Museum in Victoria.jpg|thumb|Life restoration of a woolly mammoth at Royal BC Museum]]While early mammoth species like ''M. meridionalis'' were probably relatively hairless, similar to modern elephants,<ref name="Giants">{{Cite book |last=Lister |first=Adrian |url=https://books.google.com/books?id=6bhLvgAACAA |title=Mammoths: giants of the ice age |author2=Bahn, Paul |date=2007 |publisher=Frances Lincoln LTD |isbn=978-0-7112-2801-6 |pages=25–26}}</ref> ''M. primigenius'' and likely ''M. trogontherii'' had a substantial coat of fur, among other physiological adaptations for living in cold environments. Genetic sequencing of ''M. trogontherii''-like mammoths, over 1 million years old from Siberia suggests that they had already developed many of the genetic changes found in woolly mammoths responsible for tolerance of cold conditions.<ref name=":03">{{Cite journal |last1=van der Valk |first1=Tom |last2=Pečnerová |first2=Patrícia |last3=Díez-del-Molino |first3=David |last4=Bergström |first4=Anders |last5=Oppenheimer |first5=Jonas |last6=Hartmann |first6=Stefanie |last7=Xenikoudakis |first7=Georgios |last8=Thomas |first8=Jessica A. |last9=Dehasque |first9=Marianne |last10=Sağlıcan |first10=Ekin |last11=Fidan |first11=Fatma Rabia |date=17 February 2021 |title=Million-year-old DNA sheds light on the genomic history of mammoths |journal=Nature |language=en |volume=591 |issue=7849 |pages=265–269 |bibcode=2021Natur.591..265V |doi=10.1038/s41586-021-03224-9 |issn=1476-4687 |pmc=7116897 |pmid=33597750}}</ref> Scientists discovered and studied the remains of a mammoth calf, and found that fat greatly influenced its form, and enabled it to store large amounts of nutrients necessary for survival in temperatures as low as {{convert|-50|C|F}}.<ref name="Moore2008">{{cite book |author=Peter D. Moore |url=https://archive.org/details/tundra0000moor |title=Tundra |publisher=Facts On File |year=2008 |isbn=978-0-8160-5933-1 |page=[https://archive.org/details/tundra0000moor/page/198 198] |url-access=registration}}</ref> The fat also allowed the mammoths to increase their muscle mass, allowing the mammoths to fight against enemies and live longer.<ref>{{Cite journal |last1=Maschenko |first1=E. N. |last2=Boeskorov |first2=G. G. |last3=Baranov |first3=V. A. |year=2013 |title=Morphology of a mammoth calf (''Mammuthus primigenius'') from Ol'chan (Oimiakon, Yakutia) |journal=Paleontological Journal |volume=47 |issue=4 |pages=425–438 |doi=10.1134/S0031030113040096 |bibcode=2013PalJ...47..425M |s2cid=84317574}}</ref> Woolly mammoths evolved a suite of adaptations for arctic life, including morphological traits such as small ears and tails to minimize heat loss, a thick layer of subcutaneous fat, and numerous sebaceous glands for insulation, as well as a large brown-fat hump like deposit behind the neck that may have functioned as a heat source and fat reservoir during winter.<ref>{{Cite journal |last=Lynch |first=Vincent |date=2 July 2015 |title=Elephantid Genomes Reveal the Molecular Bases of Woolly Mammoth Adaptations to the Arctic |journal=Cell Reports |volume=12 |issue=2 |pages=P217-228 |doi=10.1016/j.celrep.2015.06.027 |pmid=26146078 |doi-access=free |hdl-access=free |hdl=10220/38768}}</ref>

Based on studies of their close relatives, the modern elephants, mammoths probably had a [[gestation]] period of 22 months, resulting in a single calf being born. Their social structure was probably the same as that of [[African elephant|African]] and Asian elephants, with females living in herds headed by a matriarch, whilst bulls lived solitary lives or formed loose groups after sexual maturity,<ref>{{Cite web|url=http://library.sandiegozoo.org/factsheets/_extinct/mammoth/mammoth.htm|title=Columbian Mammoth & Channel Island Mammoth|publisher=[[San Diego Zoo]]|access-date=2010-06-15|archive-url=https://web.archive.org/web/20110727235134/http://library.sandiegozoo.org/factsheets/_extinct/mammoth/mammoth.htm|archive-date=2011-07-27|url-status=dead}}</ref> with adult males experiencing periods of [[musth]].<ref>{{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. |date=2023-05-18 |title=Testosterone histories from tusks reveal woolly mammoth musth episodes |url=https://www.nature.com/articles/s41586-023-06020-9 |journal=Nature |language=en |volume=617 |issue=7961 |pages=533–539 |doi=10.1038/s41586-023-06020-9 |pmid=37138076 |bibcode=2023Natur.617..533C |s2cid=258485513 |issn=0028-0836}}</ref>

The earliest mammoth species like ''M. subplanifrons'' and ''M. rumanus'' were mixed feeders (both browsing and grazing) to browsers. Over the course of mammoth evolution in Eurasia, their diet shifted towards mixed feeding-grazing in ''M. trogontherii,'' culminating in the woolly mammoth, which was largely a grazer, with stomach contents of woolly mammoths suggesting that they largely fed on grass and [[forb]]s. ''M. columbi'' is thought to have been a mixed feeder.<ref name=":4" />

Evidence that humans interacted with mammoths extends back to around 1.8 million years ago, with a number of bones of ''Mammuthus meridionalis'' from the [[Dmanisi hominins|Dmanisi site]] in Georgia having marks suggested to the result of butchery by [[archaic humans]], likely as a result of scavenging.<ref>{{Cite journal |last1=Tappen |first1=Martha |last2=Bukhsianidze |first2=Maia |last3=Ferring |first3=Reid |last4=Coil |first4=Reed |last5=Lordkipanidze |first5=David |date=October 2022 |title=Life and death at Dmanisi, Georgia: Taphonomic signals from the fossil mammals |journal=Journal of Human Evolution |language=en |volume=171 |pages=103249 |doi=10.1016/j.jhevol.2022.103249|pmid=36116366 |doi-access=free |bibcode=2022JHumE.17103249T }}</ref> During the [[Last Glacial Period]], modern humans hunted woolly mammoths,<ref name=":6">{{Cite journal |last1=Wojtal |first1=Piotr |last2=Wilczyński |first2=Jarosław |date=August 2015 |title=Hunters of the giants: Woolly mammoth hunting during the Gravettian in Central Europe |url=https://linkinghub.elsevier.com/retrieve/pii/S1040618215005339 |journal=Quaternary International |language=en |volume=379 |pages=71–81 |doi=10.1016/j.quaint.2015.05.040|bibcode=2015QuInt.379...71W }}</ref> used their remains to create art and tools,<ref name=":7">{{Cite journal |last1=Braun |first1=Ingmar M. |last2=Palombo |first2=Maria Rita |date=October 2012 |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 |url=https://linkinghub.elsevier.com/retrieve/pii/S1040618212004946 |journal=Quaternary International |language=en |volume=276-277 |pages=61–76 |doi=10.1016/j.quaint.2012.07.010|bibcode=2012QuInt.276...61B }}</ref><ref name=":6" /> and depicted them in works of art.<ref name=":7" /> Remains of Columbian mammoths at a number of sites suggest that they were hunted by [[Paleo-Indians|Paleoindians]], the first humans to inhabit the Americas.<ref name=":8" /> A possible bone engraving of ''M.a Columbian mammoth made by columbi''Paleoindians is known from Vero Beach, Florida.<ref>{{Cite journal |last1=Purdy |first1=Barbara A. |last2=Jones |first2=Kevin S. |last3=Mecholsky |first3=John J. |last4=Bourne |first4=Gerald |last5=Hulbert |first5=Richard C. |last6=MacFadden |first6=Bruce J. |last7=Church |first7=Krista L. |last8=Warren |first8=Michael W. |last9=Jorstad |first9=Thomas F. |last10=Stanford |first10=Dennis J. |last11=Wachowiak |first11=Melvin J. |last12=Speakman |first12=Robert J. |date=November 2011 |title=Earliest art in the Americas: incised image of a proboscidean on a mineralized extinct animal bone from Vero Beach, Florida |url=https://linkinghub.elsevier.com/retrieve/pii/S0305440311001828 |journal=Journal of Archaeological Science |language=en |volume=38 |issue=11 |pages=2908–2913 |doi=10.1016/j.jas.2011.05.022|bibcode=2011JArSc..38.2908P }}</ref>

{{MainFurther|Woolly mammoth#Extinction|Columbian mammoth#Extinction}}

TowardsFollowing the end of the [[Last Glacial PeriodMaximum]], the range of the woolly mammoth began to contract, disappearing from most of Europe by 14,000 years ago.<ref name=":9">{{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=July A. |last11=Rahbek |first11=Carsten |last12=Nogues-Bravo |first12=David |date=January 2022 |editor-last=Coulson |editor-first=Tim |title=Process-explicit models reveal pathway to extinction for woolly mammoth using pattern-oriented validation |url=https://onlinelibrary.wiley.com/doi/10.1111/ele.13911 |journal=Ecology Letters |volume=25 |issue=1 |pages=125–137 |doi=10.1111/ele.13911 |pmid=34738712 |bibcode=2022EcolL..25..125F |issn=1461-023X|hdl=11343/299174 |hdl-access=free }}</ref> By the [[Younger Dryas]] (around 12,900-11,700 years [[Before Present]]), woolly mammoths were confined to the northernmost regions of Siberia. This contraction is suggested to have been caused by the warming induced expansion of forests,unfavourable whichwet were[[tundra]] unfavourableand habitatsforest environments at the expense of the preferred dry open [[mammoth steppe]], with the possible additional pressure of human hunting. An additional issue under serious consideration is the impact of several asteroid or comet fragments impacting 12,800 years ago, evidence of which has been found in recent years, which resulted in forest fires and large dust emissions causing a sudden cooling of the climate ([[YDIH|the Younger Dryas impact hypothesis]]) lasting about 1,400 years<ref name="UofSC-asteroid">{{cite web |last1=Ward |first1=C.J.G. |title=UofSC archaeologist finds evidence of extinction theory |url=https://sc.edu/uofsc/posts/2019/10/10_chris_moore_research.php |publisher=[[University of South Carolina]] |date=22 October 2019 |access-date=21 February 2024 }}</ref><ref name="ComprRefYDIH">{{cite web |last1=Holliday |first1=V.T. |last2=Daulton |first2=T.L |last3=Bartlein |first3=P.J. |last4=Boslough |first4=M.B. |last5=Breslawski |first5=R.P. |last6=Fisher |first6=A.E. |last7=Jorgeson |first7=I.A. |last8=Scott |first8=A.C. |last9=Koeberl |first9=C. |last10=Marlon |first10=J.R. |last11=Severinghaus |first11=J. |last12=Petaev |first12=M.I. |last13=Claeys |first13=P. |title=Comprehensive refutation of the Younger Dryas Impact Hypothesis (YDIH) |url=https://www.sciencedirect.com/science/article/pii/S0012825223001915 |publisher=[[Earth-Science Reviews]] |date=December 2023 |access-date=21 February 2024}}</ref>. The last woolly mammoths in mainland Siberia became extinct around 10,000 years ago, during the early [[Holocene]].<ref name=":0">{{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 |language=en |volume=259 |pages=106913 |doi=10.1016/j.quascirev.2021.106913|bibcode=2021QSRv..25906913D }}</ref> The final extinction of mainland woolly mammoths may have been driven by human hunting.<ref name=":9" /> [[Relict (biology)|Relict]] populations survived on [[St. Paul, Alaska|Saint Paul]] island in the Bering Strait until around 5,600 years ago, with their extinction likely due to the degradation of freshwater sources,<ref>{{Cite journal |last1=Graham |first1=Russell W. |last2=Belmecheri |first2=Soumaya |last3=Choy |first3=Kyungcheol |last4=Culleton |first4=Brendan J. |last5=Davies |first5=Lauren J. |last6=Froese |first6=Duane |last7=Heintzman |first7=Peter D. |last8=Hritz |first8=Carrie |last9=Kapp |first9=Joshua D. |last10=Newsom |first10=Lee A. |last11=Rawcliffe |first11=Ruth |last12=Saulnier-Talbot |first12=Émilie |last13=Shapiro |first13=Beth |last14=Wang |first14=Yue |last15=Williams |first15=John W. |date=2016-08-16 |title=Timing and causes of mid-Holocene mammoth extinction on St. Paul Island, Alaska |journal=Proceedings of the National Academy of Sciences |language=en |volume=113 |issue=33 |pages=9310–9314 |doi=10.1073/pnas.1604903113 |doi-access=free |issn=0027-8424 |pmc=4995940 |pmid=27482085|bibcode=2016PNAS..113.9310G }}</ref> and on [[Wrangel Island]] off the coast of Northeast Siberia until around 4,000 years ago.<ref name=":0" /> The last reliable dates of the Columbian mammoth date to around 12,500 years ago.<ref>{{Cite book |last=Stuart |first=Anthony J. |url=https://press.uchicago.edu/ucp/books/book/chicago/V/bo25538572.html |title=Vanished Giants: The Lost World of the Ice Age |publisher=University of Chicago Press |location=Chicago, IL |pages=97 |language=en |chapter=North America: Mastodon, Ground Sloths, and Sabertooth Cats}}</ref> Columbian mammoths became extinct as part of the [[Late Pleistocene extinctions|Late Pleistocene megafauna extinctions]] of most large mammals across the Americas approximately simultaneously at the end of the Late Pleistocene.<ref>{{Cite journal |last1=Faith |first1=J. Tyler |last2=Surovell |first2=Todd A. |date=2009-12-08 |title=Synchronous extinction of North America's Pleistocene mammals |journal=Proceedings of the National Academy of Sciences |language=en |volume=106 |issue=49 |pages=20641–20645 |doi=10.1073/pnas.0908153106 |doi-access=free |issn=0027-8424 |pmc=2791611 |pmid=19934040|bibcode=2009PNAS..10620641F }}</ref> RemainsHunting of Columbian mammoths have been found at a number of sites with tools and cut marks suggesting that they were hunted by recently arrived [[Paleo-Indians|Paleoindians]], which may have been a contributory factor in their extinction.<ref name=":8">{{Cite journal |last=Haynes |first=Gary |date=2022-07-03 |title=Sites in the Americas with Possible or Probable Evidence for the Butchering of Proboscideans |url=https://www.tandfonline.com/doi/full/10.1080/20555563.2022.2057834 |journal=PaleoAmerica |language=en |volume=8 |issue=3 |pages=187–214 |doi=10.1080/20555563.2022.2057834 |s2cid=251042359 |issn=2055-5563}}</ref>

* {{Cite journal | last1 = Capelli | first1 = C. | last2 = MacPhee | first2 = R. D. E. | last3 = Roca | first3 = A. L. | last4 = Brisighelli | first4 = F. | last5 = Georgiadis | first5 = N. | last6 = O'Brien | first6 = S. J. | last7 = Greenwood | first7 = A. D. | doi = 10.1016/j.ympev.2006.03.015 | title = A nuclear DNA phylogeny of the woolly mammoth (''Mammuthus primigenius'') | journal = Molecular Phylogenetics and Evolution | volume = 40 | issue = 2 | pages = 620–627 | year = 2006 | pmid = 16631387| bibcode = 2006MolPE..40..620C }}