2000, Annual Reports of the Zoological Institute RAS.

Geographical variability of mammoths in the Late Pleistocene

Irina E. Kuzmina

Zoological Institute, Russian Academy of Sciences, Universitetskaya nab., 1, St. Petersburg, 199034, Russia

The mammoth Mammuthus primigenius Blumenbach was widely spread in Eurasia in the Late Pleistocene. The most abundant material over the past 30-40 years has been collected in Voronezhskaya Oblast, in Northern and Middle Ural and in Yakutia.

The emphasis was usually on mammoth skulls and teeth. Skeleton bones received much less attention. Although mammoths have been studied for 200 years subspecies systematics of the species has been insufficiently elaborated (Katalog..., 1981).

The prime objective of this study was to establish whether geographic variability of mammoths had existed in the Late Pleistocene. With this aim in mind it was decided to compare sizes of tusks, molars, and particularly limb bones and to process these data statistically, which had not been done previously.

Material and methods

The material examined had been collected by an expedition on Berelekh River in Yakutia with the participation of N.K. Vereshchagin, G.F. Baryshnikov, V.M. Khrabryi and others in 1970-1971. The collections comprised 3 skulls, 164 teeth and nearly 8500 skeleton bones of mammoth. Of these 1400 bones have been measured (Vereshchagin, 1977; Baryshnikov et al., 1977).

As a result of archaeological excavations in Late Palaeolithic sites in Kostenki village in Voronezhskaya Oblast vast paleontological material has been accumulated comprising more than 50 000 bones of different mammal species of which more than 10 000 belong to the mammoth. Fossil bones collected were characterized by poor state of preservation, and therefore only 497 specimens were used for measurements during the period from 1977 through 1989.

In Ural the author worked in Medvezhya Cave on the Upper Pechora River in 1961, on the site Byzovaya in the Middle Pechora in 1965, in Chernye Kosti Cave, in Tain Cave, in Bliznetsov Grotto, in Stolbovoi Grotto and others in 1968 through 1971. Moreover, paleontological collections deposited in museums of Ekaterinburg, Ufa, Nizhni Tagil, Serov and other cities of Middle Ural have been examined. A total of approximately 4500 mammoth bones were collected of which 434 bones were measured. Age of all the paleontological material examined has been determined by C14 dating. Geological age of these finds according to dating is 12 000-35 000 years.

Remains of mammoths of different age were discovered in all the studied regions. Bones of adult animals were measured using the previously adopted procedure (Kuzmina, 1971). For a more detailed analysis the material was divided into male and female bones (Kuzmina, 1982). The procedure of determining sex by adult animal bones involved comparison of any two parameters of bones. The sample falls into two groups. The bones were classified as those of males or females by analogy with sizes of modern animals whose sex can be determined (Kuzmina, 1997).


Vereshchagin and Tikhonov (1986) examined mammoth tusks from Yakutia. They cited data on sizes and weight of the tusks and on age and sex of animals to which they belonged. Kuzmina and Tsyganova (1999) processed statistically the data obtained by the above authors. Tusks of 18-65 year old females from Yakutia were 40-93 mm in diameter, on the average 76*1 mm (n=93). Tusks of 25-73 year old males were 89-180 mm in diameter, on the average 132*2 mm (n=85).

Length of numerous tusks in Kostenki in Voronezhskaya Oblast varied from 18 cm to 3 m. Female tusks were 40-95 mm in diameter, on the average 81*3 mm (n=22). Their circumference varied from 130 to 300 mm, on the average 235*73 mm. Tusks ascribed to males were 97-195 mm in diameter on the average 140*2 mm (n=48) with circumference of 310-615 mm on the average 440*22 mm.

Diameter of tusks in females from Ural varied from 63 to 95 mm, on the average 83*2 mm (n=12). Their circumference varied form 220 to 280 mm and was on the average 257*7 mm. Diameter of tusks in males was 100-187 mm, on the average 147*6 mm (n=16). Their circumference varied from 300 to 560 mm, constituting on the average 456*18 mm.

These figures suggest that mammoths in the centre of the Russian Plain were somewhat smaller than in Ural, whereas animals in the Russian Plain and in Ural were notably larger than mammoths in Yakutia.

Comparison of samples from the Russian Plain and Yakutia and also from Ural and Yakutia has revealed statistically significant differences for both females and males, at significance level of 5%. Differences in diameter of female tusks from the Russian Plain and Ural were not statistically significant.

A study of molars of Berelekh mammoths was conducted by Zherekhova (1977). Teeth of Kostenki mammoths were described by Urbanas (1980). Data on teeth of Ural mammoths have been obtained by the authors. Comparison of length and width of the last upper and lower molars shows that these teeth were the largest in mammoths of Ural. Thus, length of the upper M3 varied from 230 to 274 mm and was on the average 259*8 mm (n=5). Width varied from 86 to 110 mm, on the average 100*4 mm. Length of M3 was 245-326 mm, on the average 277*5 mm (n=3). Width was equal to 85-110 mm, on the average to 98*7 mm.

In mammoths from Kostenki length of the upper molar M3 was 250-269 mm, at a width of 77-91 mm, on the average 83.8*3 mm (n=5). Length of the lower molar M3 varied from 220 to 309 mm, on the average 271.5*13 mm (n=6), at a width of 80-100 mm, on the average 87*2 mm (n=10).

Teeth of Berelekh mammoths were even smaller. Length of the upper M3 varied from 193 to 238 mm, on the average 213*6 mm (n=7), at a width of 60-90 mm, on the average 78*3 mm (n=10); length of the lower M3 varied from 185 to 250 mm, on the average 217*13 mm (n=5) at a width of 72-96 mm, on the average 83*3 mm (n=8).

Comparison of some measurements of skeleton bones from the above regions has revealed the same relationship (Table 1).


Measurements, mm

Russian Plain












Length of scapula


465 - 790

630 10


496 - 850

630 11


440 - 830

589 10

Width of articulation
surface of scapula


73 - 133

96 1


84 - 182

109 7


80 - 125

92 1

Length of humerus


500 - 1070

682 26


590 - 1020

747 28


460 - 940

639 13

Width of distal epiphyses
of humerus


180 - 305

222 4


180 -330

235 7


160 - 275

191 2

Length of ulna


435 - 920

594 18


473 - 900

646 34


430 - 780

570 9

Width of proximal joint
surface of ulna


148 - 260

188 3


150 - 250

200 7


150 - 230

179 2

Length of femur


640 - 1240

815 19


640 - 1130

833 32


580 - 1130

770 14

Width of distal epiphyses
of femur


147 - 260

186 3


150 - 245

185 5


140 - 225

168 3

Length of tibia


430 - 700

526 15


468 - 700

561 24


330 - 570

453 6

Width of distal epiphyses
of tibia


100 - 215

150 5


112 - 195

153 7


110 - 180

133 1


Calculation of confidence factor of differences between these three samples has shown the most significant differences between mammoth bone sizes of the Russian Plain and Yakutia and also of Ural and Yakutia. The differences between sizes of mammoth bones from the Russian Plain and Ural were also statistically significant at 5% significance level.

For a more detailed study mammoth bones from the above regions were divided into male and female bones. Sizes of female bones from three mammoth populations and also sizes of male bones from the same regions were compared.

Although relationship of sizes of mammoth bones from different geographic regions was generally the same, i.e. mammoths from the centre of the Russian Plain were somewhat smaller than animals from Ural, they were larger than mammoths from Yakutia. However confidence level of the differences varied.

Thus, differences in sizes of humerus in females 1-2 and 2-3 were statistically significant at a high confidence level, whereas differences between samples 1 and 3 showed low significance. The same pattern was observed in differences of sizes of the humerus in males. A similar relationship was revealed by comparison of length of ulna and femur ascribed to females.

However, statistically significant differences have been revealed in sizes of femurs of males between all three samples: between samples 1 and 2, 2 and 3, 1 and 3 at 5% significance level.

Length of tibia in females showed statistically significant differences in all three populations. In males the most significant differences in sizes were between samples 1 and 2 as well as between samples 2 and 3. The differences between samples 1 and 3 were less significant.

Thus, geographic variation of mammoths from northern Eurasia in the Late Pleistocene may be thought of as proven: mammoths from the central Russian Plain were somewhat smaller in sizes than animals which inhabited Ural, but were larger than mammoths of Yakutia. This statement may be fixed taxonomically by splitting the species into separate subspecies.

Skeleton of mammoth male found in Taimyr Peninsula on Mamontovaya River, a tributary of Shrenk River in 1948 (No. 27101, collections of Zoological Institute RAS) is regarded on suggestion of Garutt (1989) as a neotype. In publications of the past years the neotype's number was indicated erroneously as 2710 (Averianov & Sablin, 1991; Garutt et al., 1993; Averianov, 1994).

Sizes of bones of Taimyr mammoth fit within the range of bone sizes of Berelekh mammoths. Therefore they should be assigned to the nominative subspecies Mammuthus primigenius primigenius. Descriptions of the other two subspecies are given below.


Mammuthus primigenius rossicus Kuzmina, subsp. nov.


Named - by distribution in the Russian Plain.

Holotype: Zoological Institute RAS, Laboratory of Terrestrial Vertebrates, No. 30908a, right M3, site Kostenki 1, layer 1, Voronezhskaya Oblast, right bank of Don River, Late Pleistocene.

Diagnosis. Large mammoth with relatively large last molars. Length of M3 284 mm, width of tooth 87 mm, density of plates - 8. Average enamel thickness 1.6*0.19 mm (1.1-2.0 mm, n=4)

Distribution. East-European Plain, Late Pleistocene.


Mammuthus primigenius uralensis Kuzmina, subsp. nov.


Named by distribution in Ural.

Holotype: Zoological Institute RAS, Laboratory of Terrestrial Vertebrates, No. 35080, mandible with right M3, site Byzovaya, Komi Republic, right bank of Pechora River, 25 km from Pechora station up the river, Late Pleistocene.

Diagnosis. A large mammoth with large last molars. Length of M3 278 mm, width 86.5 mm with a relative frequency of plates - 11 and small enamel thickness 1.2*0.06 mm (1.1-1.4 mm, n=4).


Comparison. Differs from the previous subspecies by larger elements of skeleton, teeth with a larger number of plates and thinner enamel.


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