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Russian J. Theriol. 13(2): 7182

© RUSSIAN JOURNAL OF THERIOLOGY, 2014

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
I.V. Foronova
ABSTRACT. A peculiar form of the thick-enamel mammoth from late Middle Pleistocene of the Kuznetsk Basin (South of Western Siberia) and from the Acheulian-Mousterian site Ust'-Izhul'1 with a unique faunal assemblage (North-Minusinsk Basin, Kurtak, Yenisei River, south of Central Siberia) is described. It is compared to Mammuthus intermedius (Jourdan, 1861) from Western Europe. The stratigraphic position and morphometric features of molars, defining the position of this form in the mammoth lineage, characterize it as an intermediate link in the transition from M. trogontherii to M. primigenius. Mammuthus intermedius inhabited interstadial/interglacial environments of late Middle Pleistocene and had a vast Eurasian range. KEY WORDS: Middle Pleistocene, Early Paleolithic, Mammuthus intermedius, mammoth lineage, south of Siberia, Russia.
Irina V. Foronova [irina_foronova@mail.ru], V.S. Sobolev Institute of Geology and Mineralogy, Siberian Branch of Russian Academy of Science, Pr. Koptiuga, 3, Novosibirsk 630090, Russia.

Mammuthus intermedius (Proboscidea, Elephantidae) ():
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. , ( ) - - 1 (- , , . , ), Mammuthus intermedius (Jourdan, 1861) . , , , , M. trogontherii M. primigenius. Mammuthus intermedius / . : , , Mammuthus intermedius, , , .

Introduction
Proboscideans have a long history of study. The taxonomic definitions based on the traditional methods of dental analysis are, however, strongly impeded by the transgression of teeth characters. The taxonomy of the group still remains unclear and requires revision. This often and inevitably causes problems in biostratigraphic definitions and paleoclimatic reconstructions. In order to obtain more precise taxonomical definitions and override the transgression, a more complex statistical approach was suggested using traditional

morphometric molar features in an untraditional way. It was a method of so called "optimal parameters" (Garutt & Foronova, 1976), and a multiple stage method of cluster analysis of large series of M3. It allowed constructing multidimensional models of the Archidiskodon-Mammuthus lineage in Northern Eurasia (Foronova & Zudin, 1986, 1995, 1999, 2001; Foronova, 2001a, b, 2007, 2014) (see Figs. 4, 5). This method clearly shows the structure of the mammoth lineage to be more complex than a simple gradualistic sequence presumed before. It illustrates evolutionary trends and discreteness of processes in the mam-

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I.V. Foronova

Fig. 1. Location of the Kuznetsk Basin (Southwestern Siberia) and North-Minusinsk Basin, Kurtak archaeological region (South-Central Siberia). A: Kuznetsk Basin. Locations of coal-mining pits (fossil mammal localities): 1 -- Kedrovka, 2 -- Chernigovo, 3 -- Mokhovo, 4 -- Gramoteino, 5 -- Bachatsk, 6 -- Novosergeevo, 7 -- Krasnobrodsk. B: North-Minusinsk Basin. Kurtak archaeological region.

moth lineage. Moreover, the method may provide a solution to many pending problems, including the main ones: revealing intermediate elements (Garutt, 1971), revision of old taxa, and determining new ones within the lineage. The present paper is devoted to one of such taxa, a peculiar Mammuthus intermedius (Jourdan, 1861) widely spread in Northern Eurasia at the end of Middle Pleistocene. The structure of its teeth gives it a definite position in the mammoth lineage among other intermediate forms between Mammuthus trogontherii (Pohlig, 1885) and Mammuthus primigenius (Blumenbach, 1799). Originally, this form was described in Western Europe (Lyon Plateau in the RhТne Valley, France) as Elephas intermedius (Jourdan, 1861; Lortet & Chantre, 1872). For a long time it was recognized as valid (Osborn, 1942; Aguirre, 1969; Beden & GuИrin, 1975). But later the validity of this species was questioned (Lister, 1996). Recently French scientists revived M. intermedius and supplemented its diagnostics with molars from AbНmes de La Fage (CorrХze, France) (Beden & GuИrin, 1975; Labe & GuИrin, 2005). In Russia, it was Baygusheva (1980) who described specific features of this unusual mammoth based on a record from the left bank of Severskiy Donets River (Kamensk, Rostov Region). She defined this form to be very close to Mammuthus chosaricus Dubrovo, 1966. In her opinion some of its distinctive features support establishing a separate subspecies within M. primige-

nius. Later, while developing the method and constructing diagrams based on the material from Europe, Eastern and Western Siberia (Foronova & Zudin, 1986), a prominent adaptive peak was recognized on each of the diagrams. Its center nearly matched the parameters of M. intermedius from France (according to data of Osborn, 1942) (see Figs 4, 5). Mammoths of similar morphological structure and geological age were also found in the Volga region (Kamskoe Ust'e and Mysy-Mansurovo) and described as a part of the Volga fauna (Averianov et al., 1992). Based on cranial and postcranial material Averianov attributed them to the so-called `post-Khozar mammoth', following Baygusheva (1980) who placed the mammoth from Kamensk to this form. Thus, the analysis of extensive Eurasian materials using the above mentioned method confirmed the existence of the specific mammoth form and showed its distribution from Western and Eastern Europe to Asia (in Western and Central Siberia) (Foronova & Zudin, 1986, 1995, 1999, 2001; Drozdov et al., 1990; Foronova, 2001a, b). Despite the early discovery of this mammoth form in Siberia, the cited publications refer to it using the open nomenclature: M. primigenius ssp., thick-enamel form, intermediate between M. chosaricus and early M. primigenius; M. ex gr. trogontheriiintermedius and M. cf. intermedius, etc. We follow Baygusheva (1980, 1999) in considering this peculiar form of "post-chosaricus" thick-enamel early mammoth as a separate taxon.

Mammuthus intermedius from Middle Pleistocene of Siberia

73

Fig. 2. Mammuthus intermedius (Jourdan, 1861): M3 sin, IGM SB RAS #755. Krasogolovo layers of Kedrovka Formation, Bachatsk pit, Kuznetsk Basin. Occlusal view.

Below we provide a systematical description of this mammoth's molars from two major stratigraphic regions in southern Siberia, the Kuznetsk and NorthMinusinsk Basins (Fig. 1), and justify its attribution to M. intermedius (Jourdan, 1861). Abbreviations. IGM SB RAS -- V.S. Sobolev Institute of Geology and Mineralogy, Siberian Branch of the Russian Academy of Sciences, Novosibirsk; IAE SB RAS -- Institute of Archaeology and Ethnography, Siberian Branch of the Russian Academy of Sciences, Novosibirsk.

Localities and material
The Kuznetsk Basin and Kurtak archaeological district are the largest South-Siberian stratigraphic regions that represent almost continuous sequence of Quaternary sediments and faunas (Fig. 1). The examination of rich paleontological collections from these regions allowed us to trace the development of Proboscidea through the entire Pleistocene (Foronova, 1986, 1998, 2001a, b, 2010). Kuznetsk Basin (54°13 N, 86°20 E) is a vast intermountain depression situated in the South-East of Western Siberia. It is surrounded by Kuznetsk Alatau Ridge from the East, Mountainous Shoriya from the South, Salair Ridge in the West and Kolyvan-Tomsk Upland from the North-East (Fig. 1A). Eight of Russia's largest coal-mining pits located here expose a thick (over 100 m) layer of Neogene-Quaternary sediments. This creates unique possibilities for collecting abundant paleontological material characterizing almost every period of Pleistocene. The coal-mine exposures also facilitate complex geological and taphonomical observations. A detailed biostratigraphic subdivi-

sion of the Quaternary in the region was performed on the basis of the multimethod study of the large mammal fauna. It resulted in a refined regional stratigraphic scale of the Kuznetsk Basin, including important refinement of stratigraphical ranges of the Kedrovka and Bachatsk Formations, and establishing the Berezovo and Chernigovo Formations (Foronova, 2001b). The material under study (18 last mammoth molars) was excavated in situ in Bachatsk, Novosergeevo, Mokhovo, and Kedrovka coal pits, from Krasogolovo beds in the upper part of fluviatile Kedrovka Formation. Molars of the peculiar form under study (Fig. 2) constitute the majority of mammoth teeth found in the entire Kedrovka Formation. The Krasogolovo beds also yielded remains of Ursus rossicus Borissiak, 1930, Panthera spelaea Goldfuss, 1810, Equus cf. germanicus Nehring, 1884, Cervus elaphus Linnaeus, 1758, Megaloceros giganteus Blumenbach, 1803, Alces cf. alces Linnaeus, 1758, and Bison priscus Bojanus, 1827. Two additional teeth of similar structure were found in the Mokhovo pit, in loess-like loams of the overlying Berezovo Formation, particulary, in its lower part with a paleosol. These deposits also yielded remains of Ursus cf. arctos Linnaeus, 1758, Panthera spelaea, Stephanorhinus kirchbergensis (JДger, 1839) (=Dicerorhinus mercki (JДger, 1839)), Equus sp., Cervus elaphus, Megalocerus giganteus, Bison priscus. Paleontological characteristic of this series can be supplemented with a complex of small mammals. According to Galkina (Zudin et al., 1982), it is represented by: Lagurus cf. lagurus, Microtus (Stenocranius) cf. gregalis, Myospalax cf. myospalax, Dicrostonyx simplicior-henseli. The species composition of large and small mammals and a clear geological position within a nearly continuous sequence of Quaternary deposits in the Kuznetsk Basin, indicate late Middle Pleistocene and the

74

I.V. Foronova constrained by the author using its faunal composition. Broad-legged horse attributed to E. ex gr. mosbachensis-germanicus occurred in Siberia in the Middle Pleistocene. Mammoths, similar to M. intermedius, are represented by a thick-enamel form transitional between Middle Pleistocene M. chosaricus and the typical Late Pleistocene M. primigenius (Drozdov et al., 1990; Foronova & Zudin, 1995). These data were later used for refining the age of the site (Ovodov, 1995; Laukhin et al., 1999; Foronova, 2000).

Shirta Interglacial/Interstadial (inter-Saalian) as the time of sedimentation of the upper part of Kedrovka Formation (Krasogolovo layers) and Berezovo Formation (Foronova, 2001b). North-Minusinsk Basin (55°09 N, 91°32 E) is situated in the south of Central Siberia and represents one of intermountain depressions in the eastern part of Altai-Sayan mountainous region (Fig. 1B). From east and west it is bounded by the ridges of Kuznetsk Alatau and Eastern Sayan. The main water artery of the Basin is the Yenisei River. Its valley is presently filled by the Krasnoyarsk Reservoir. The erosional coastal cliffs of the reservoir hosted the most important archaeological sites of Siberia. 20-km long coastal zone is known as the Kurtak archeological district with numerous sites and localities of Quaternary mammals (Drozdov et al., 1990). The local sections of the Quaternary are among the best in south Siberia. They comprise the 60100 m thick loess-paleosol sequence and alluvial sediments of high terraces (Krukover & Chekha, 1999; Haesaerts et al., 2009; etc.). In this south Siberian region, mammoths were found in the Ust'-Izhul'1 locality. It is one of the well-known Paleolithic sites occurring in the eastern part of Kurtak archaeological district, at the left bank of the Krasnoyarsk Reservoir. The locality is famous for the immense amount of animal remains and human artifacts that have been described in numerous publications (Drozdov et al., 1990; Ovodov, 1995; Laukhin et al., 1999; etc.). The mass burial of mammal bones (beds 25 and 26) occurred at the depth of 1820 m at the base of mostly loamy member overlying the eroded terrace alluvium. The fossiliferous beds are represented by cherry-terracotta colored clays of bed 25 (with interbeds of loamy sand) changing to dark nonstratified humus-rich clays of bed 26. The major part of the fossil accumulation is represented by hundreds of mammoth bones and more than 40 teeth of various generations, from dP4 to M3. It was established that these remains belonged to approximately 16 individuals of presumed age from 2 to 40 60 years. Other finds included badger, marmot, mole vole, broad-toed horse, rhinoceros, red deer, and small bison. The fact that some bones were found in anatomical order implies in situ accumulation (Ovodov, 1995). Eight molars of last generation (4 uppers and 4 lowers) are used in the present study (Fig. 3). These faunal remains co-occurred with AcheulianMousterian implements. Numerous artifacts with characters of the Levallois technique occurred among the animal remains. The composition of rock material used for these tools is identical to Early-Paleolithic and Mousterian assemblages of the Kutrak archaeological district. Human activity is also evidenced by fire spots and abundant charcoal pieces on the surface of a paleosol in the layer 26. In addition, some bones have cut-marks and signs of breaking easily distinguishable from bites of carnivores (Laukhin et al., 1999). Along with archaeological data and other dating information, the age Ust'-Izhul' 1 site was significantly

Methods
The methodology used in this contribution has been elaborated and described in detail by Foronova & Zudin (1986, 1999) and discussed in subsequent publications (Foronova & Zudin, 1995, 2001; Foronova, 2001a, b, 2003, 2007, 2014). It was based on plotting and analyzing multi-dimensional diagrams based on values of enamel thickness (E), plate frequency (PF) per 100 mm of crown length and average plate length (PL) of several thousands of molars (M3) representing the Archidiskodon-Mammuthus lineage from numerous Quaternary localities and archaeological sites in Northern Eurasia (Western and Eastern Europe, Western, Central, and Eastern Siberia) (Figs. 4, 5). These features show directed evolution and are traditionally regarded as the most important for defining taxonomic position of mammothoid elephants. The isolines of abundance plotted against bivariate graphs revealed a complex and nonlinear morphospace. It is believed that in the giant data set as we used the major peaks of the morphospace correspond to non-random adaptive zones of dental evolution in the mammoth lineage. The detailed description of the methods is provided in above mentioned publications. In the course of analysis of two-dimensional diagrams, additional dental parameters were also used for comparison of individual clusters and evaluation of distribution pattern of molars. Crown width, total number of plates, hypsodonty index, angle of molar eruption, and angle of abrasive wear were analyzed using Kolmogorov-Smirnov's criterion (Miller & Kahn, 1963). Shape of enamel loops on the masticatory surface and ratios of plate cement, dentine, and enamel in molars were also included in the analysis (Foronova & Zudin, 1986).

Variability
The samples included into the analysis were selected on the basis of their relative uniformity: different types of variability, except intraspecific variability, were excluded. The age variability of teeth was excluded because only the molars of last generation (M3) were analyzed. The dentition asymmetry was analyzed using the subset of European mammoths. Separate diagrams for lower, upper, right and left M3 are virtually indistinguishable, the number, position and parameters of clus-

Mammuthus intermedius from Middle Pleistocene of Siberia

75

Fig. 3. Mammuthus intermedius (Jourdan, 1861): M3 sin, IAE SB RAS #25, Ust'-Izhul'1 site, Kurtak archaeological region. A -- occlusal view; B -- lingual view.

ters do not change (Foronova & Zudin, 1986, 1999, 2001; Foronova, 2007). Variations of features from different stages of abrasive wear of a crown were excluded due to technique for measuring enamel thickness (along the entire chewing surface), described above. Pathological teeth were extremely rare. However, their key features appeared to be similar to those of normal teeth. Geographical variability was excluded by plotting diagrams separately for Europe, Western and Eastern Siberia. These graphs showed both similarity of the general structure and the presence of the overwhelming majority of elements, analogous in position and structure.

Analysis of distribution structure
Separate diagrams were built for Europe, Western and Eastern Siberia. They appeared to be extremely

informative and clearly demonstrated the lineage structure to be far more complex than a traditional gradualistic sequence still in use. Generally, the variability area (from archaic forms to the latest mammoth) reflects canalizing selection in the lineage due to global natural changes of the Quaternary (Figs. 4, 5). However, the most important and innovative result is that selection of combinations of the features under study (i.e. of levels of dental system specialization, most optimal for different stages of lineage development) was observed to be discrete. The structure consists of subordinated "adaptive peaks" (zones of high distribution density) and depressions, and resembles the "Wright's symbolic picture" (in terminology of Dobzhansky, 1951). The peaks group into ensembles, so a hierarchy of marginal depressions can be seen. One of the large

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I.V. Foronova

Fig. 4. Variability of characters of M3 in elephants of the mammoth lineage in Europe (by materials from Western and Eastern Europe) (modified after Foronova & Zudin, 1999).
Coordinate axes: E -- enamel thickness; PF -- plate frequency per 100 mm; PL -- length of one plate. Continuous isolines of distribution density are drawn with increment of 0.5 Uniform Density Units, punctuated isolines, 0.25 Units, outer isoline corresponds to 0.25. Punctuated lines show assumed direction of lineage development due to selection pressure. Rectangles show variability limits for features of early and late form of M. primigenius (according to Vangengeim, 1961). Dots are the coordinates of typical specimens of taxa distinguished in the lineage and some peculiar forms: 1 -- M. primigenius sibiricus; 2 -- M. primigenius primigenius, neotype; 3 -- M. primigenius jatzkovi, holotype; 4 -- M. primigenius fraasi, holotype; 5 -- M. primigenius, early form, average parameter values; 6 -- M. primigenius (Chokurcha site); 7 -- M. primigenius, lectotype; 8 -- M. intermedius (after Osborn, 1942); 9 -- M. trogontherii chosaricus, holotype; 10 -- M. trogontherii chosaricus, holotype (authors' measurements); 11 -- M. trogontherii trogontherii, lectotype; 12 -- M. trogontherii (Azov Museum; authors' measurements) = A. wuesti; 13 -- A. meridionalis cromerensis, holotype; 14 -- A. m. voigtstedtensis, holotype; 15 -- A. m. tamanensis, holotype; 16 -- A. meridionalis; 17 -- A. m. meridionalis, holotype; 18 -- A. gromovi; 19 -- A. m. taribanensis. Color dots stand for mean values: red -- M. intermedius (Jourdan, 1861), RhТne, France, after Labe & GuИrin (2005); green -- M. primigenius from Kamensk town, Rostov Region, after Baygusheva (1980, 1999); blue -- M. primigenius from Kamensk town, Rostov Region, measurements of I. Foronova and A. Zudin.

ensembles corresponds to the final stage of lineage development within the genus Mammuthus. In addition to adaptive peaks of axial zone, we pioneered to find series of peaks in "thick-enamel" and "thin-enamel" areas of distribution. They are oppositely oriented and clinally linked with the axial zone peaks. These peaks are entirely new elements, significantly differing the structure from traditional gradualistic model. Thick-enamel peaks are represented by the forms with thickened folded enamel, medial sinuses, and low hypsodonty of a crown, whereas high hypsodonty, rare narrow plates with thin and weakly folded enamel are typical of thin-enamel forms. The comparison of regional diagrams shows transcontinental distribution of the majority of phenotypes

and chiefly autochthonous development at the continental scale. Slight difference between the parameters of analogous phenotypes from various regions implies geographical clinal variability due to different responses of regional environments to global climatic changes. Thick-enamel and thin-enamel adaptive peaks are regarded as forms with different ecological adaptations. Morpho-functional differences, accompanying fauna and flora, different habitats, physical parameters, and certain agreement between the sequence of these forms and marine oxygen-isotope stages provide grounds to link thick-enamel and thin-enamel adaptations with interglacial and periglacial environments respectively. It makes forms of the mammoth lineage significant for paleoecologic reconstructions.

Mammuthus intermedius from Middle Pleistocene of Siberia

77

Fig. 5. Variability of characters of M3 in elephants of the mammoth lineage from Kuznetsk Basin (south of Western Siberia). See Fig. 4 for details.
Color dots stand for mean values of M. intermedius: red -- after Labe & Guerin (2005); yellow -- Kuznetsk Basin, after Foronova (2001b); violet -- Ust'-Izhul' 1, Kurtak, after Zudin & Foronova, 1995; Foronova, 2000; green -- Oyash River, Novosibirsk Region, measurements of V.E. Garutt and I. Foronova.

Results Systematic Paleontology
Order Proboscidea Illyger, 1811 Family Elephantidae Gray, 1821 Genus Mammuthus Brookes, 1828 Mammuthus intermedius (Jourdan, 1861) Figs. 2, 3; Tab.1.
1861: Elephas intermedius: C. Jourdan, Des terrains sidИrolitiques: 1013. 1990: Mammuthus primigenius ssp. (thick-enamel form, intermediate between M. chosaricus and early form of M. primigenius): Drozdov et al., Chronostratigraphy of the Paleolithic Sites...: 111, Table 5. 1995: Mammuthus ex gr. trogontherii intermedius: I.V. Foronova & A.N. Zudin, O novom metode izucheniya...: 132, fig. 1. 1999: Mammuthus cf. intermedius: I.V. Foronova, Quaternary mammals....: 86, fig. 9(2), # 575. 2001a: Mammuthus cf. intermedius: I.V. Foronova, History of Quaternary proboscideans of south...: 110111, figs.1, 2. 2001b: Mammuthus cf. intermedius: I.V. Foronova, Quaternary mammals of the South-East...: 8687, 208209, Table X, fig. 2.

TYPE MATERIAL. The species was originally based on check teeth found in the vicinity of Lyon, France (Jourdan, 1861). Neotype: right hemimandible No.42341 (Beden & GuИrin, 1975, pl. 8). Paratypes: four third molars from AbНmes de La Fage (CorrХze, France): lower M3 No.42303 and 42329, upper M3 No.42302 and No.42312 (Labe & Guerin, 2005). TYPE LOCALITY. Karst filling of Aven I des AbНmes de La Fage, at Noailles (CorrХze, France), upper Middle Pleistocene (Labe & Guerin, 2005). MATERIAL Kuznetsk Basin, Southeast of Western Siberia. Upper molars: M3 dex and M3 sin, paired, complete, #6240; M3 dex, #6007; M3 dex, #6021; M3 dex, #5042, Mokhovo coal-mining pit, Krasogolovo layers of Kedrovka Formation, basal part of Berezovo Formation. M3 dex, #4038; M3 sin, #766, Kedrovka coalmining pit. M3 dex, #11; M3 dex, #7000, Novosergeevo coal-mining pit. M3 sin #755, Bachatsk coal-mining pit, Krasogolovo layers of Kedrovka Formation. Lower molars: M3 sin, complete, #9075; M3 sin, #6258, Novosergeevo coal-mining pit, Krasogolovo layers of Kedrovka Formation. M3 dex, complete, ##

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I.V. Foronova
Table 1. Molars (M3) of mammoths from the late Middle Pleistocene of Europe, Western and Central Siberia. M. intermedius Kuznetsk Basin, IGM SB RAS, (Foronova & Zudin, 1986, 1995, 1999; Foronova, 1990, 2001) M3 n=6 240285 90106 180186 23+ 7.08.0 13.114.6 1.82.3 M3 n=9 245365 80108 130144 2425 6.58.0 13.014.4 1.72.3

M. intermedius (Jourdan, 1861) Measurements, mm RhТne, Lion, France, (Labe B., GuИrin C., 2005) M3 n=2 1. Crown length 2. Crown width 3. Crown height 4. Plate number 5. Plate frequency / 100 mm 6. Average length of a plate 7. Enamel thickness 300305 96102 158 24 6.46.7 1.72.0 (mean 6.75) 1.92.4 M3 n=10 +138305 8096 122154 6.57.5

M. intermedius Ust'-Izhul' 1, Kurtak, IAE SB RAS (Foronova & Zudin, 1995; Foronova, 2000, 2001) M3 n=4 280300 8994 160190 2224 7.58.5 12.513.5 2,02,2 M3 n=4 257290 8188 130140 2023 6.58.0 12.514.8 2,02,2

(mean 7.3) (mean 13.6)

(mean 7.25) (mean 13.4) (mean 2.1) M. primigenius Kamensk, Rostov Region, Collection of Novocherkassk Regional Studies Museum (measurements by I. Foronova and A. Zudin) M n=3 245310 9398 175 23+ 7.08.0 15.015.9 1.92.3
3

(mean 2.1) M. intermedius Novosibirsk Region, Oyash river, Novosibirsk Regional Studies Museum (measurements by V. Garutt, I. Foronova; Foronova, 2001) M n=2 1. Crown length 2. Crown width 3. Crown height 4. Plate number 5. Plate frequency / 100 mm 6. Average length of a plate 7. Enamel thickness 195+ 9394 190+ 2324 7.08.0 (mean 7.5) 13.013,5 (mean 13.8) 1.82.0 1.852.2
3

(mean 2.0) M. primigenius Kamensk, Rostov Region, Collection of Novocherkassk Regional Studies Museum (Baygusheva, 1980, 1999) M n=4 303310 9597.6 170 2425 7.357.75
3

M3 n=2 225+ 88 15+ 7.5

M3 n=3 280 91.8 18+ 6.757.25 1.613.6 1.72.2

M3 n=2 260265 9294 18+ 6.06.5 13.013,5 1.82.0

(mean 7.25)

(mean 6.9) (mean 14.3)

2.12.2

(mean 2.0)

(mean 2.1)

(mean 2.0)

3186, 8016; M3 dex, #8254, Mokhovo coal-mining pit. M3 dex, #3432; M3 dex, #604. Bachatsk coal-mining pit. M3 dex, #8009, Sartaki Krasogolovo layers of Kedrovka Formation. Collection of IGM SB RAS.

North-Minusinsk Basin, Kurtak archaeological district, South of Central Siberia. Upper molars: M3, ## 10, 11, 12, 13. Lower molars: M3, ## 25, 26, 27, 28.

Mammuthus intermedius from Middle Pleistocene of Siberia Ust'-Izhul'1 Paleolithic site, beds 2526, left bank of Krasnoyarsk Reservoir. Collection of IAE SB RAS. Geological age. Late Middle Pleistocene, Shirta horizon of Western Siberia, Odintsovo horizon of Eastern Europe, second half of Saalian of Western Europe, ca. 200-160 ka.

79

Description and comparison
Molars (M3) of this Siberian mammoth are relatively large: upper molars are 240300 mm long, 89106 mm wide, and 160190 mm high; lower molars are 245365 mm long, 80108 mm wide and 130144 mm high. Mean plate length (PL), 13.413.8 mm; mean plate frequency (PF), 7.257.5; mean enamel thickness (E), 2.02.1 mm (calculated for upper and lower molars) (Tab. 1). In values of PF, PL and E the mammoth under study significantly differs from the neotype of M. primigenius (Blumenbach, 1799) corresponding to the late form of the species (Figs. 4, 5). On the basis of traditional morphometric analysis this mammoth could be regarded as one of the earliest representatives of M. primigenius of the early type, since the provided values partly correspond to extreme "archaic" values of the early form of mammoth (according to the variability range given by Vangengeim, 1961). Alternatively, diagrams based on our method (Figs. 4, 5) demonstrate that a significantly compact group (a distinct adaptive peak near the point 8) can be observed at this stage of mammoth lineage development. Partly, it is situated in "archaic" upper right corner of M. primigenius (see the right rectangle on the diagrams). However it can be clearly seen that variability limits of this form are much broader and overlap the conventional limits proposed for early M. primigenius by Vangengeim (1961) reaching even more "archaic" values (Tab. 1). Obviously this form has an intermediate position between so-called "Khosar" and "early" mammoths. To be more precise, it differs from M. chosaricus Dubrovo in higher (more progressive) plate frequency and from M. primigenius of early type in somewhat thicker enamel. Dental parameters of the Siberian mammoths attributed to this form are rather similar (Tab. 1). Apart from the materials from Kuznetsk and North-Minusinsk Basins, this group of finds includes a full skeleton of female mammoth, found in blue clays of a coastal exposure of Oyash River near the city of Novosibirsk (exhibited in Novosibirsk Regional Studies Museum). Measurements of its skull and post-cranial skeleton were published by V.E. Garutt (Averianov et al., 1992: 3154). This material corresponds to all morphological features of the genus Mammuthus and differs only in small dimensions. Dots corresponding to average values of PF and E of these Siberian forms are densely spaced within the mentioned adaptive peak (Fig. 5). The only exception is slightly increased enamel thickness of mammoths from Ust'-Izhul'1 locality, which is more similar to the values of European representatives

of the group and implies mild environmental conditions of these mammoths' habitat in Siberia. In addition, characteristic features of these mammoths' M3 include lower hypsodonty, rare medial widening of plates, in most cases roughly folded and (as mentioned above) thickened enamel. These features are typical of teeth from the so-called "thick-enamel" and partly "axial" zones of distribution. Sequence of forms with such a specialization of molars, distinguishing them from thin-enamel type, has been traced from the archaic to the latest forms of elephants (Foronova & Zudin, 1986, 1999, 2001; Foronova, 2001a, b). The listed morpho-functional dental features, coupled with geological data and other data, indicate a habitat with reltively mild conditions in the late Middle Pleistocene, corresponding to the intra-Riss/intra-Saalian Shirta interstadial horizon of the West Siberian stratigraphic chart (Foronova, 2001b). In Europe, Mammuthus intermedius (Jourdan, 1861) from interglacial loess of Lyon Plateau in France corresponds to the same stratigraphical interval. This species has a long history of definitions, descriptions, denials and later revivals (Osborn, 1942; Aguirre, 1969; Beden & GuИrin, 1975; Labe & GuИrin, 2005). The morphological and chronological characters of this species correspond to the distinct adaptive peak in the diagrams and define its position in the lineage. Previously, we have shown (Foronova & Zudin, 1986, 1999; Foronova, 2001) that the mean values of PF and E in topotypic forms nearly coincide with the center of the distinct adaptive peak in the diagrams (Figs. 4, 5; black dot 8) and are very similar to the available Siberian data. A recent contribution (Labe & GuИrin, 2005) revisiting M. intermedius provided somewhat different values for this form. The diagrams show that revised mean values (red dot) are slightly shifted towards the "archaic" area, but its variability range still closely corresponds to that of the Siberian form under study. In European Russia, Baygusheva (1980, 1999) has long since described similar mammoths in Kamensk town, Rostov Region. They deserve a special attention, because their values of PF, PL and E, similar to M. intermedius from France, closely match the grouping center of the corresponding adaptive peak (Tab. 1; Fig. 4). In course of her study Baygusheva concluded the position of this so-called "early mammoth" to be conterminal with M. chosaricus and proposed to distinguish it as a subspecies within M. primigenius. Our study demonstrates that this mammoth from southern Russia hardly differs in its parameters from the southern Siberian mammoth studied in the present paper. (Tab. 1; Fig. 4). Two mammoth skeletons from Kamskoe Ust'e and Mysy-Mansurovo sites in the Volga region may be conspecific with the form under study. It is important that Averianov, who studied the molars, cranial and post-cranial material of the Volga mammoths, attributed them to the so-called `post-Khozar mammoth'. Moreover, Averianov emphasized that in dental characters the mammoths from Mysy-Mansurovo characterize the

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Fig. 6. Schematic distribution of M. intermedius (Jourdan, 1861) in the late Middle Pleistocene in Eurasia. Range borders are given according to geographical distribution of material used (after Foronova & Zudin, 1986; Foronova, 2007, with modifications). E -- Europe; WS -- Western Siberia; ES -- Eastern Siberia.

same `stage of phenotype stabilization' (in terminology of Foronova & Zudin, 1986) as the Kamensk mammoth described by Baygusheva (1980, 1999) (Averianov et al., 1992: 74). Hence, all provided data and comparison make us refer the late Middle Pleistocene mammoth form of south of Western and Central Siberia to Mammuthus intermedius (Jourdan, 1861).

ature data indicate a fairly large geographic range of M. intermedius (Fig. 6) covering almost the entire northern latitudes of Eurasia (Foronova & Zudin, 1986, 1999).

Discussion
The materials provided and discussed in this paper refer to one of the most long-lasting and complicated problems in Quaternary paleontology, the distinguishing and determining of the so-called "intermediate elements" in the Archidiskodon-Mammuthus lineage (Pohlig, 1885; Dubrovo, 1966; Garutt, 1971, and others). Considerable climatic and environmental changes at the end of Early and especially in Middle Pleistocene have conditioned further development of the mammoth lineage. Periodic changes of landscape and vegetation have caused an increase of plate number and frequency in the tooth crown, and decrease of plate length and enamel thickness. The obtained diagrams clearly show these changes to be successive stages of elephants' adaptation to periodic environmental changes. A number of transitional forms (well-pronounced adaptive peaks) between M. trogontherii and M. primigenius s.l. can be seen on all regional diagrams (including the one for Kuznetsk Basin) in the major part of the late Middle Pleistocene (Tobol, Samarovo, and Shirta horizons of West Siberian stratigraphic scale (= Holstenian and major part of Saalian of Western Europe). One of such intermediate forms is undoubtedly M. intermedius first described by Jourdan (1861) as Elephas intermedius whose numerous cheek teeth were found near Lyons (RhТne, France). Labe & GuИrin (2005) provided a brief history of study of this species, reestablished it as M. intermedius (Jourdan, 1861) and proposed the inclusion of all described forms spanning the interval between M. trogontherii and M. primige-

Comments
Comparison of European and Siberian diagrams demonstrates that the analyzed adaptive peak is more pronounced and advanced in its parameters in the Siberian diagram (Figs. 4, 5). This effect was noted at the beginning of our work with the method (Foronova & Zudin, 1986, 1999, 2001), when a slight displacement of all similar adaptive peaks of mammoth lineage occurred while superimposing the diagrams of Europe and Siberia. We believe that this is a reflection of clinal variability of the lineage, which was caused by different reaction of regional environments (changes in landscape and vegetation zones) to global climatic changes. This is also illustrated by Fig. 6 which shows schematic distribution of M. intermedius (Jourdan, 1861) in late Middle Pleistocene in Eurasia. Range boundaries are given according to geographical distribution of extensive materials from Europe, Western and Eastern Siberia (Foronova & Zudin, 1986).

Distribution
The large factual material (measurements of thousands of M3 from numerous localities of Western and Eastern Europe, and entire Siberia) collected and used for compilation of diagrams, as well as numerous liter-

Mammuthus intermedius from Middle Pleistocene of Siberia nius to this species. These authors estimated a chronological range of M. intermedius within the zone MNQ24 of C. GuИrin biochronological scale, from 305 to 105 ky. All forms more advanced than M. trogonterii, including M. t. chosaricus (Dubrovo, 1966, Garutt, 1972), M. primigenius-trogontherii (Guenther, 1969) and others, are regarded as synonyms to M. intermedius. Another related issue is the status of the species M. chosaricus Dubrovo, 1966. In the stratigraphical scheme, this form spans the interval between 400300 ka and corresponds to Singil/Tobol faunal complexes of the Russian continental scale (Holstenian of Western Europe). Thus the upper boundary of M. trogonterii is restricted to 400 ka or the mid Middle Pleistocene. The recognizing of M. chosaricus is usually a much more difficult problem than in case of other species. As we have already noted (Foronova, 2001b) this form lacks a clear morphological definition. The Khosar mammoth was initially described as a late subspecies of M. trogontherii, M. trogontherii chosaricus (Dubrovo, 1966), but later raised to a species level (Garutt, 1972). Rationalizing the taxonomic validity of theses two subspecies Dubrovo provided the variability range of plate frequency of M. t. trogontherii as 5.07.0, but it appears to be the same for M. t. chosaricus. The enamel thickness varies from 2 to 3.5 in M. t. trogontherii and from 2.0 to 2.5 in M. t. chosaricus. When the mean values of the lectotype of M. trogontherii (PF: 6.25; E: 2.1) are compared to values in M. t. chosaricus (PF: 6, 25, E: 2, 5), it becomes clear that the two forms are hard to distinguish. In other words, when a variability range of a taxon is nested within variability range of another taxon, that Dubrovo's diagnostics is obviously incorrect. The described situation is vividly reflected in the European and Siberian diagrams which demonstrate the proximity of dots 9, 10 and 11 corresponding to the typical specimens of these two forms (Figs. 4, 5). In addition, the diagrams, especially the Siberian one, demonstrate one extra adaptive peak different from M. intermedius that occupies a substantial interval from the "typical" M. trogontherii-chosaricus to M. intermedius. Obviously, it was this form (probably a subspecies), and not a vague M. chosaricus, that could be a predecessor of M. intermedius. Moreover, Labe & GuИrin (2005) spreading the boundaries of M. intermedius to this entire interval (down to M. trogontherii) and including into it "M. chosaricus", date its lower boundary at 305 ka. Our previous studies (Foronova & Zudin, 1986, 1999, 2001; Foronova, 2001a, b, 2007, 2014) have shown that thick- and thin-enamel forms (perhaps, in rank of subspecies) could correspond to the second half of Middle Pleistocene, beginning from 400 ka as shown by provided diagrams and results of other studies. Forms with thin enamel inhabited severe periglacial environments, while thick-enamel forms lived in interstadial and interglacial climatic conditions. Two thin-enamel forms correspond to the time-span of 400130 ka or the late Middle Pleistocene. The first form, identified as

81

Mammuthus sp., occurred in late Middle Pleistocene (Khazar and Samarovo horizons of the Russian timescale, or Early Saalian in Western Europe). The second form, described as M. primigenius cf. fraasi Dietrich 1912 (Figs. 4, 5; dot 4), corresponds to the very end of Middle Pleistocene (Tazovo horizon of the West-Siberian Stratigraphic Scale of Russia, or Late Saalian in Western Europe) (Foronova, 2007, 2014). The form M. intermedius (Jourdan, 1861) described in this paper as judged by its dental characters inhabited mild and relatively warm environment of inter-Saalian, late Middle Pleistocene, Shirta Interglacial/Interstadial of Southern Siberia, ca. 200160 ka. ACKNOWLEDGMENTS I am greatly indebted to Dr. V.S. Baygusheva, Azov Museum-Reserve of History and Archaeology, Azov, Rostov Region for long-lasting help and collaboration; Dr. Alexey S.Tesakov, Geological Institute, RAS, Moscow advised on technical and scientific issues; Prof. Gennady F. Baryshnikov, Zoological Institute, RAS, St. Petersburg; Dr. Vadim V. Titov, Institute of Arid Zones, SSC RAS, Rostov-on-Don, and Dr. Attila VirАg, Hungarian Natural History Museum, Budapest are acknowledged for important advice and comments during the preparation of the manuscript. Drs. N.I. Drozdov, V.P. Chekha and N. D. Ovodov, Institute of Archaeology and Ethnography, SB RAS, Novosibirsk, Krasnoyarsk are thanked for providing the material from Ust'Izhul'1; Igor V. Foronov helped in translation.

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