Editing Mammoth (section)

== Description ==
{{See also|Elephant#Anatomy}}
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 mature adult males having an average height of approximately {{convert|3.8-4.2|m|1|abbr=on}} at the shoulder and weights of {{convert|9.6-12.7|tonne|lb|lk=on}}, while exceptionally large males may have reached {{convert|4.5|m|1|abbr=on}} at the shoulder and {{convert|14.3|tonne|lb|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 with males around {{convert|2.80-3.15|m|1|abbr=on}} high at the shoulder, and {{convert|4.5-6|tonne|lb|lk=on}} in weight on average,<ref name=":54">{{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=2024-11-20 |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 ''[[Mammuthus creticus|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|''[[Mammuthus meridionalis]]'' bull, around {{convert|4|m|ft}} tall|M. trogontherii skeletal (cropped).png|[[Steppe mammoth]] (''M. trogontherii'') around  {{convert|3.9|m|ft}} tall in front-on (without head) side-on and top-down views|M._columbi_skeletals_(cropped).png|[[Columbian mammoth]] (''M. columbi'') bull around  {{convert|3.7|m|ft}} tall|M. primigenius modified.png|[[Woolly mammoths]] (''M. primigenius''), including one of the largest, the Siegsdorf mammoth (left, around {{convert|3.5|m|ft}} tall), and a mature Siberian bull (around {{convert|2.7|m|ft}} metres tall)|||||width=185|height=|lines=|align=center|title=Gallery of mammoth skeletons}}

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 |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 |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 |volume=219 |pages=145–153 |bibcode=2019QSRv..219..145R |doi=10.1016/j.quascirev.2019.06.027 |s2cid=200073388}}</ref>

{{Multiple image
| image1            = Mammuthus sp. sectioned upper and lower molars 1.jpg
| header            = Molars
| align             = center
| image2            = Mammuthus meridionalis molar - Cleveland Museum of Natural History - 2014-12-26 (20267609313).jpg
| image3            = Woolly mammoth molar - Cleveland Museum of Natural History - 2014-12-26 (20859260938).jpg
| total_width       = 600
| caption1          = Cross section through elephantid molars, showing their internal structure
| caption2          = Molar of ''[[Mammuthus meridionalis]]''
| caption3          = Molar of a woolly mammoth (''Mammuthus primigenius'')
}}

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 |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 |volume=37 |pages=45–58 |doi=10.1080/08912963.2023.2286272 |s2cid=266182491 |issn=0891-2963}}</ref>

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 |doi-broken-date=2024-11-20 |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 |bibcode=2015PalZ...89..229L |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 |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=217–228 |doi=10.1016/j.celrep.2015.06.027 |pmid=26146078 |doi-access=free |hdl-access=free |hdl=10220/38768}}</ref>