Early and Middle Pleistocene of Northern Eurasia

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Vislobokova I., and Tesakov A. (2013) Early and Middle Pleistocene of Northern Eurasia. In: Elias S.A. (ed.) The Encyclopedia of Quaternary Science, vol. 4, pp. 605-614. Amsterdam: Elsevier.

Early and Middle Pleistocene of Northern Eurasia I Vislobokova , Paleontological Institute of the Russian Academy of Sciences, Moscow, Russia

A Tesakov , Geological Institute of the Russian Academy of Sciences, Moscow, Russia

Introduction Mammal Assemblages of Northern Eurasia

Early Pleistocene: Gelasian, Middle Villafranchian, Late The best represented vertebrate records in the early and middle Villanyian, MN17 Pleistocene of Northern Eurasia are mammalian and they are summarized in this article. The data from Russia and neigh- The Khaprovian (Khapry) mammal assemblage and its boring countries of the former USSR are very important for the analogs understanding of the history of Late Cenozoic biota. The terri- This stage is well documented in the northern Black Sea region, tory was a prime area of origin and evolution of taxa and of peri-Azov area, Siberia, and Central Asia. The mammal com- dispersals of some mammals to Western Europe, Africa, North munities superficially resembled those of modern Africa or America and China, and vice versa. The configuration of Asia southern Asia by the presence of proboscideans, equids, rhi- was close to that of the present day but differed through the noceroses, giraffids, and various antelopes, but differed periodic existence of the Bering land bridge, connecting Siberia considerably from them in species composition. These assem- and Alaska. The reorganization of the biota went through blages are characterized by the first appearance and subseq- climatic and environmental changes; its main trends and uent wide distribution of the elephant Archidiskodon gromovi phases are very pronounced there. (¼ A. meridionalis gromovi in Titov, 2008, ¼ Mammuthus gro- The key stages in the history of Pleistocene mammals in this movi of Lister and Sher, 2001). In contrast to Russian paleon- territory were first recorded by Gromov (1948) who defined tologists, most researchers refer this species to the genus the faunal assemblages for Eastern Europe and named them by Mammuthus, mainly based on the tooth structure (Lister and their type localities or regions. Regional complexes were later Sher, 2001), although Archidiskodon is retained here. Other defined in the Caucasus, Siberia, Transbaikalia, and Central features of the fauna include the presence of the bear Ursus Asia. The data on the ages of the assemblages and their com- etruscus and the rhinoceros Stephanorhinus sp., the rapid positions were considerably enriched thanks to many subse- diversification and wide distribution of stenonid horses, quent researchers and new discoveries. In addition to large the presence of diverse canids (Eucyon, Nyctereutes, Canis), two mammals, a detailed biochronology based on small mammals saber-toothed cats Homotherium crenatidens and Megantereon has appeared in the last decades. At present, early and middle cultridens, the camel Paracamelus, the large-sized comb-antlered Pleistocene mammal remains are known from many localities deer Eucladoceros and the elk (¼ moose) Libralces, and other of Northern Eurasia. The age of the localities has been estab- extinct forms. In Northern Eurasia, there were three paleozoo- lished by a combination of geological and paleontological geographic subareas: European-Siberian, Mediterranean, and methods (including the first appearance of index taxa, as well Central Asian (Vangengeim and Pevzner, 1991; Vislobokova as the evolutionary levels of dominant forms and of some et al., 1995)(Figure 1). The global cooling witnessed at about phyletic lineages), paleoclimatic data and correlation. Paleo- 2.6 Ma (indicated in the territory by the glaciations of the magnetic data also help to distinguish more precisely the age of Caucasus and Pamir, the most ancient loess formation in early Pleistocene sites, whereas the glacial–interglacial succes- Tadjikistan, Uzbekistan, etc.) caused a decrease in humidity sion provides further important markers for age determination. in temperate latitudes and in Central Asia (Dodonov, 2002), The beginning of the Pleistocene is currently set at 2.6 Ma where animals adapted to open woodlands and grasslands (the Gauss–Matuyama magnetic reversal), the early–middle became dominant. Some taxa have North American ancestry, Pleistocene boundary at 0.78 Ma (the Brunhes–Matuyama immigrating during periods of low sea level prior to the start magnetic reversal), and the middle–late Pleistocene boundary of the Pleistocene, such as the camels, Hipparion and Eucyon at 0.126 Ma, according to the International Commission on inthelateMiocene,andEquus, Canis,andMegantereon in the Stratigraphy in 2009. However, the Geological Surveys of Pliocene (Vislobokova et al., 2003). A number of thermophilous Russia and Ukraine have until recently retained the Pliocene– herbivores (mastodons, giraffids, and others), typical of Pliocene Pleistocene boundary in its former position at 1.8 Ma. In 2012 times, were gradually replaced by forms adapted to cooler cli- the Russian Stratigraphic Committee voted to lower the base of matic conditions and possessing more advanced herbivorous the Quaternary to 2.58 Ma. The position of the lower boundary adaptations. The data on herbivorous taxa (elephants, horses, of the Pleistocene and its divisions have thus been repeatedly ruminant Artiodactyla, voles, etc.) are of great value for biochro- changed over the last decades and it must be appreciated that nology and paleoenvironmental reconstruction because their the published names of particular stratigraphical units often tooth structure yields clearly recognizable signals in wear patterns, refer to different time slices. hypsodonty, etc. The increase in diversity of herbivore taxa was

The early and middle Pleistocene in Northern Eurasia accompanied by considerable changes in the carnivore guild. currently include the following mammal assemblages: At this time, an extensive European-Siberian part of the Khaprovian, 2.6–2.2 Ma; Psekupsian, 2.2–1.2 Ma; Tamanian, Palearctic was inhabited by the elephant Archidiskodon gromovi, 1.2–0.8 Ma; Tiraspolian, 0.8–0.4 Ma; Singilian and Khasarian, the rhinoceros Elasmotherium, and large Equus livenzovensis 0.4–0.126 Ma. and small Equus sp., together with other animals adapted to

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Figure 1 Early Pleistocene paleozoogeographic subareas and main localities: (1) European-Siberian subarea; (2) Central Asian subarea; (3) Mediterranean subarea.

savanna-like conditions (the wolves of the Canis etruscus group, Don River near Rostov-on-Don and Taganrog. They come from the hyena Pliocrocuta, the saber-toothed cat Homotherium cre- the lower part of the Khapry alluvium recorded in sand pits natidens , the cheetah Acinonyx, the camel Paracamelus, the an- and natural outcrops. The large mammals were represented by telope Gazellospira, the bison-like Leptobos, and others), along Anancus arvernensis alexeevae, the horses Hipparion moritorum, with forest and ecotonal animals (the bear Ursus etruscus and Equus (Allohippus) livenzovensis, and a small Equus sp., the rhi- the deer Eucladoceros and Libralces). Most of these mammals noceroses Stephanorhinus ex gr. megarhinus-kirchbergensis and were found in common with those from the Middle Villafran- Elasmotherium chaprovicum (¼ E. cf. caucasicum), the wild boar chian faunas of Western Europe (Kahlke et al., 2011; Palombo Sus strozzi, the camels Paracamelus alutensis and Paracamelus cf. et al., 2006). Early Villafranchian elements (the mastodon gigas, the deer Cervus (Rusa) philisi, Eucladoceros cf. dicranios,

Anancus arvernensis, the archaic horse Hipparion, and giraffids) Arvernoceros sp., and Libralces gallicus, the giraffid Palaeotragus became very rare. Among the small mammals, the bank voles (Yuorlovia) priasovicus, and the bovids Leptobos sp., Gazellospira Clethrionomys made their first appearance, the water vole Mim- gromovae, Tragelaphini gen. indet., Gazella cf. subgutturosa, and omys polonicus was replaced by M. pliocaenicus in the Northern others (Alexeeva, 1977; Bajgusheva, 1971; Bajgusheva et al., Black Sea area, and the pikas Pliolagomys and Ochotona and 2001; Gromov, 1948; Titov, 2008; and others) (Figure 2). The jerboas Allactaga, more typical of Asian faunas, were present. Khapry carnivores included the raccoon-dog Nyctereutes mega-

Fossil remains of Archidiskodon gromovi, the main indicator mastoides, Canis sp. (C. cf. senezensis: Sotnikova et al., 2002), species of the Khaprovian mammal assemblage, have been the mustelids Lutra sp. and Pannonictis nesti, the medium-sized found in a number of localities in the European part of hyena Pliocrocuta perrieri and the dominant larger-sized Russia (Khapry, Liventzovka, and others), in the Ukraine short-faced hyena Pachycrocuta brevirostris, the felids Lynx issio- (Zhevakhova Gora), in the south of Western Siberia (Podpusk- dorensis, the saber-toothed cat Homotherium crenatidens, and the

Lebyazh’e), and in Central Asia (Adyrgan, Kuruksay) (Bajgusheva, cheetah Acinonys pardinensis. Most of these animals were in- 1971; Garutt and Bajgusheva, 1981; Vangengeim et al., habitants of open savannah-like landscapes, with a part of the 1988; Vislobokova, 1996, 2005; and others). A. gromovi fauna adapted more to forested or ecotonal zones (mustelids, has also been found in Italy (Montopoli) (Gliozzi et al., lynx, wild boar, and deer). Among the small mammals, the 1997). archaic lagurine Borsodia praehungarica, the water voles Mimomys

Remains of other members of the Khaprovian assemblage praepliocaenicus and Mimomys ex gr. reidi, and the primitive red- are recorded from many European–Russian sites along the backed voles Clethrionomys kretzoii were also present (Tesakov, northern coast of the Sea of Azov and the right bank of the 2004).

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In northeastern Asia, the association of Synaptomys-like lemmings Plioctomys and mimomyian voles Cromeromys ex gr. irtyshensis-hordijki and smaller Mimomys sp. were found in the lower member of the Kutuyakh Formation at the Krestovka River in the Kolyma Lowland (Sher, 1987; Tesakov and van

Kolfschoten, 2011). The assemblage is dominated by lemmings and ‘Cromeromys’ voles, with Ochotona sp. and Mimomys sp. being less abundant. Several faunas of this age have been referred to the Central Asian subarea (Erbaeva and Alexeeva, 2000; Sotnikova et al.,

1997; Vangengeim, 1977; Vislobokova et al., 1995). These include the Itanza fauna of Klochnevo I, II (Transbaikalia) with Ochotona intermedia, the ground squirrel Spermophilus itancinicus, Allactaga sp., the hamster Cricetinus varians,the voles Mimomys pseudintermedius, Clethrionomys sp., and

‘Villanyia’ klochnevi, the archaic mole rat Prosiphneus cf. Figure 2 Anancus arvernensis, C. Flerov painting exhibited in the paratingi,thefirstrepresentativeofthegiantdeergenus Paleontological Museum of the Paleontological Institute, Russian Praemegaceros and others, as well as the Kiikbai and Andyrgan Academy of Sciences (PIN) in Moscow, with PIN permission. faunas (southern Kazakhstan) with Ochotonoides complicidens, Mimomys pliocaenicus,thegerbilMeriones cf. meridianus, Equus stenonis,thecamelGigantocamelus longipes, and other

forms. The fauna of Southern Tadzhikistan has affinities with the Eastern Mediterranean subarea and is characterized by a combination of Mediterranean and Central Asian forms. The rich assemblage from Kuruksay contained, along with A. gromovi

and Mastodontoidea fam. et gen. indet., the baboon Papio sushkini (¼ Paradolichopitecus sushkini), the porcupine Hystrix sp., diverse carnivores including Nyctereutes megamastoides, eucyon ‘Canis’ kuruksaensis, Ursus cf. etruscus, the cursorial hyena Chasmaporthetes lunensis kani, Pliocrocuta perrieri, Lynx ex

gr. issidorensis, Acinonys cf. pardinensis, Megantereon megantereon, and Homotherium crenatidens, the rhinoceros Stephanorhinus sp., the horse Equus stenonis pamirensis (¼ E. s. bactrianus), the camel Paracamelus praebactrianus, the deer Axis flerovi Figure 3 Podpusk–Lebyazh’e locality at the right bank of the Irtysh and Elaphurus eleonorae, the first representative of the giant River, Western Siberia. deer genus Sinomegaceros (S. tadzhikistanis), the elk Libralces

cf. gallicus, the giraffids Sogdianotherium kuruksaense and Sivatherium, and the antelopes Gazella parasinensis, Protoryx In the south of Western Siberia, faunal assemblages paralaticeps, Damalops palaeindicus, Gazellospira gromovae, and described by Vangengeim and Zazhigin have been reported Antilospira sp., amongst others (Dmitrieva, 1977; Vangengeim from Podpusk-Lebyazh’e, from the lower part of the thick et al., 1988). In this region, a large number of Asian elements sandy alluvial deposits exposed on the right bank of occurred. Animals of dry, open landscapes (Ellobius, Pliocrocuta, the Irtysh River, south of Pavlodar (Vangengeim, 1977; Chasmaporthetes, Paracamelus, Equus) were accompanied by Vislobokova, 1996; Figure 3). Typical members include A. forest animals (bear, lynx, monkey, the deer Axis and Libralces) gromovi, S. cf. etruscus, Elasmotherium sp., Equus livenzovensis, and savannah inhabitants (Sivatherium, Protoryx, Damalops, Equus sp. (small), Paracamelus cf. gigas, Eucladoceros sp., Lepto- Gazellospira). Such a mixed composition reflected the mosaic bos, and the antelopes Gazella cf. sinensis and Antilospira landscapes and vertical zonality typical of mountainous cf. gracilis (Vislobokova, 1996). The faunal assemblage con- regions. In Tadzhikistan, the local faunas attributed to this sisted mainly of forms widespread in the temperate zone of the assemblage are known also from Obigarm, Karamaidan, Palearctic region (Ursus cf. etruscus, Homotherium, Pachycrocuta, Tutak, and Zil’fi localities, while faunas close to them in age and others), with some Central Asian elements (Ochotonoides, were found in Kyrgyzstan (Akterek, Dzylgyndykoo) (Dmitrieva

Elasmotherium, Paracamelus, Antilospira cf. gracilis, Gazella and Nesmeyanov, 1982; Sotnikova et al., 1997). cf. sinensis), some of which (e.g., Paracamelus) reached Eastern The reversed polarity of the fossiliferous deposits in Khapry Europe. Among the small mammals, voles with rooted denti- indicates that these sediments should be attributed to the lower tion of the genus Mimomys (M. pliocaenicus and M. reidi) and part of the Matuyama Chron, together with analogous sites Borsodia (B. praehungarica-petenyii) were predominant and in Siberia and Central Asia (Podpusk-Lebyazh’e, Krestovka,

Clethrionomys appeared for the first time in Siberia (Zazhigin, Kuruksay) (Sotnikova et al., 1997; Tesakov et al., 2007; 1980). The mammalian assemblages indicate the prevalence Vangengeim and Pevzner, 1991; Vangengeim et al., 1988; of steppes with woodland developed along river valleys. Vislobokova, 1996).

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¼ Late Early Pleistocene: Late Villafranchian, MN18–MQ19, In Psekups (Northern Ciscaucasia), Archidiskodon meridio- Tiglian–Menapian nalis, Eucladoceros orientalis (¼ Psekupsoceros orientalis, first recognized as Cervus pliotarandoides), Stephanorhinus etruscus, and Psekups mammal assemblage and its analogs Equus stenonis were present (Alexeeva, 1977; Gromov, 1948; This stage is characterized by significant changes in the diver- Vislobokova, 1990; and others). The small mammals included sity ofmammal communities (Figure 4). In Northern Eurasia, Borsodia newtoni-arankoides, Pitymimomys pitymyoides, Mimomys a cold-climate event around 2.2–1.9 Ma resulted in the appear- cf. pliocaenicus, Mimomys reidi, and Clethrionomys kretzoii ance of new boreal elements, including some modern genera. (Alexandrova, 1976; Tesakov, 2004). These reversed polarity In the European-Siberian subarea, Archidiskodon gromovi was deposits correspond to the Matuyama Chron, between the replaced by the more progressive A. meridionalis (¼ Mam- Reunion and Olduvai Subchrons (Tesakov, 2004; Vangengeim muthus meridionalis), close to the type of the species from and Pevzner, 1991). Slightly younger assemblages from the the Upper Valdarno, Italy. Voles of the genus Allophaiomys, Odessa small mammal complex contain hypsodont Mimomys the giant deer Praemegaceros, the elk Alces, the large bovids forms (dominant M. reidi and M. ex gr. savini), the last repre- of the genus Bos and the Bison (Eobison)–B.(Bison) phyletic sentatives of Borsodia, the first occurrence of lagurines with lineage, and the musk-ox Soergelia first appeared in this stage.

Mammal assemblage

Evolutionary lineage

scale

Magnetic polarity time (Cande and Kent, 1995) Ma Epoch Age Mammal age Mammal zone Eastern Europe Siberia Western Transbaikalia Eastern Siberia

Arvicola

G. gulo Singilian- E. luteus Ivolginian- Ust’kirenskian Chasarian C. lupus

U. spelaeus chosaricus P. antiquus P. Mammuthus 0.5 Brunhes

Middle

M. gregaloides M. gregalis

Middle–late Tologoian trogontherii Mammuthus

Homotherium sp . Gulo cf. schlosseri

Galerian Olyorian 1.0

20 U. deningeri Early fauna Tamanian Tiraspolian Jaramillo Razdolian Vyatkinian

Zasukhino Archidiskodon m. tamanensis P. pannonicusP. Lagurus transiens L. lagurus Palaeoloxodon sp . C. mosbachensis–C. variabilis

19 Mimomys savini

Early

Lagurodon arankae

1.5 Pleistocene Microtus (stenocranius) hintoni fauna Late Dodogolian Archidiskodon Eolagurus argyropuloi Kizikhian Psekupsian meridionalis 18 Matuyama Vilyuiskian fauna Vilyuiskian Canis etruscus Prolagurus ternopolitanus

Olduvai

2.0 Allophaiomys deucalion A. pliocaenicus Villafranchian

Reunion

Gelasian 17 gromovi Itanzinian Middle

Archidiskodon Khaprovian Borsodia spp. Krestovkian fauna Krestovkian

2.5 Ursus etruscus Homotherium crenatidens Podpusk-Lebyazhian

Mimomys pliocaenicus Mimomys (Tcharynomys) spp. Mimomys (Tcharynomys)

Figure 4 Mammal assemblages and faunas of Northern Eurasia and the main lineages.

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Mammal assemblage

Evolutionary lineage

scale

Magnetic polarity time Eastern Siberia Transbaikalia (Cande and Kent, 1995) Western Siberia Western Eastern Europe Mammal zone Mammal age Epoch Age Ma

Ovibos Singilian- Chasarian Ivolginian- Ust’kirenskian E. caballus Coelodonta antiquitatis C. elaphus

0.5 Brunhes

Rangifer S. kirchbergensis M. giganteus Coelodonta tologoijenis Tologoian Camelus

Galerian

Equus mosbachensis

1.0 C. acoronatus Alces latifrons 20 Early Middle–late Equus suessenbornensis fauna Jaramillo Razdolian Vyatkinian Tamanian Tiraspolian Zasukhino Praeovibos

Sinomegaceros sp . Bison schoetensacki B. priscus longicornis

alutensis P.

1.5 Pleistocene fauna Late Ovibovini Dodogolian

Bison suchovi Alces sp . Kizikhian Vilyuiskian faunaVilyuiskian Olyorian Megaloceros sp .

Psekupsian

Cervus abesalomi Matuyama 18

2.0

Villafranchian

Stephanorhinus etruscus

Reunion Olduvai

Gelasian Early Middle 17 Itanzinian Middle Arveroceros sp . Khaprovian Paracamelus gigas Libralces gallicus Equus stenonis Leptobos sp . 2.5 Equus livenzovensis Krestovkian fauna Krestovkian Podpusk-Lebyazhian Sinomegaceros tadzhikistanis

Figure 4 (Continued)

unrooted teeth Prolagurus and Lagurodon, and Allophaiomys deu- cf. perrieri, Megantereon cultridens, Homotherium crenatidens, calion (Alexandrova, 1967; Rekovets and Nadachowski, 1995; Palaeotragus) and forest animals (Bison (Eobison)georgicus, the Tesakov, 2004). The Odessa assemblage is subdivided into two deer Cervus abesalomi and Dama nestii, the bear Ursus etruscus) phases: (1) an early phase, with the remains of rootless voles (Gabunia and Vekua, 1995; Hemmer et al., 2010; Vekua, 1995; Allophaiomys deucalion and (2) a late phase, with the first cemen- Vekua and Lordkipanidze, 2008). The first occurrences of the ted rootless voles of Prolagurus (P. ternopolitanus) and Lagurodon ‘jaguar’ Panthera onca georgica and musk ox Soergelia are (L. arankae )(Markova, 1982, 2007). recorded there. The contemporaneous Palan-Tyukan fauna In the Caucasus, the Dmanisi fauna is famous for the pres- (Azerbaijan) includes Canis etruscus, Panthera ex gr. onca gom- ence of the first appearance in Eurasia of Homo, following baszoegensis, and Nyctereutes megamastoides. dispersal from Africa. The remains of the Dmanisi hominins In Western Siberia, A. cf. meridionalis, Equus ex gr. stenonis, are attributed to the Homo erectus group and were found in the Alces sp., and Gazella sp. along with Borsodia ex gr. petenyii- layers directly overlying a lava flow with a radiometric date of hungaricus, and Mimomys coelodus are known from the Podpusk 1.801 Ma, within the Olduvai Subchron. The fauna com- fauna (from the uppermost part of the sands in the Irtysch prises inhabitants of dry, open area (the tortoise Testudo, the River Basin) (Vislobokova, 1996). In southern Kazakhstan, the ostrich Struthio, the gerbil Parameriones, the pika Ochotona, the Aktogai (Kopaly) locality with Mimomys (Tcharynomys) haplo- hamster Cricetulus, A. meridionalis, Canis etruscus, Pliocrocuta dentatus (bed 3) and Allophaiomys deucalion (bed 7) also

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Author's personal copy 610 VERTEBRATE RECORDS | Early and Middle Pleistocene of Northern Eurasia corresponds to this age (Sotnikova et al., 1997; Tjutkova and Kaipova, 1996). In the Kuznetsk Depression, Allophaiomys pliocaenicus, Pro- lagurus ex gr. pannonicus-posterius, Eolagurus argyropuloi, Archi- diskodon cf. meridionalis, and Ovibovini gen. indet. were found in the Mokhovo Formation, 1.8–1.2 Ma (Foronova, 2001). In northeastern Asia, a tooth of Archidiskodon meridionalis meridionalis was found in the Vilyuisk district, Yakutia; the find confirmed the spread of the subspecies into North America over the Bering land bridge (Dubrovo, 1990), probably at the end of the 2.9–2 Ma global sea level fall. In Transbaikalia, the Dodogol and Zasuchino II faunas have been referred to the Central Asian subarea. In Dodogol, Borsodia laguroformes, Allophaiomys cf. pliocaenicus, Prosiphneus youngi, the first woolly rhino Coelodonta (of Asian origin), Figure 5 A skeleton of Elasmotherium on display in the Paleontological Equus ex gr. sanmeniensis, and others were present (Vangengeim, Museum of the Paleontological Institute, the Russian Academy of 1977). The revised data on the Pleistocene small mammals of Sciences (PIN), Moscow, with PIN permission. the region are presented in Alexeeva et al. (2007) and Alexeeva and Erbaeva (2008). 0.8 Ma and broadly synchronous with stone tools from the

Karama site in the Altai region (Derevyanko and Shunkov, Terminal Early Pleistocene: Early Galerian, 1.2–0.8 Ma 2005). The time range from 1.2 to 1.1 Ma (before the Jaramillo paleo- Morozovkian small mammal fauna with more derived magnetic reversal) is an important boundary in the history of voles Microtus (Stenocranius) hintoni and the pitymyoid Micro- the biota, and is reflected in the nomenclature of stratigraphic tus sp. are correlated with these assemblages (Alexandrova, schemes (the Villafranchian–Galerian boundary or the Early– 1976; Rekovets and Nadachowski, 1995). The small mammal

Late Eopleistocene boundary in the scheme of the Russian fauna of this stage are also represented in the Dnieper, Dnies- Stratigraphic Committee). ter, and Don basins (Agadjanian, 2009; Alexandrova, 1976; Markova, 2007; Tesakov et al., 2007; Topachevsky et al., Tamanian mammal assemblage and its analogs 1987; and others). The assemblages include an advanced (Nogaisky–Morozovka, ¼ Menapian–Bavelian) meadow vole Allophaiomys (A. pliocaenicus) and the steppe

This stage is characterized by the replacement of Archidiskodon lemmings Prolagurus pannonicus, Lagurodon arankae, and Eola- meridionalis meridionalis by the more advanced A. meridionalis gurus argyropuloi. tamanensis , the first appearance of the giant deer Praemegaceros At this time, differentiation of environment sharply verticornis and Megaloceros, the red deer Cervus (Cervus) acoro- increased. In the Caucasus (Akhalkalaki, Georgia), the most natus, the roe deer Capreolus cf. suessenbornensis, the broad- thermophilous elements continued to exist, including Hippopot- fronted elk Alces latifrons, the bison Bison (Bison), camels of amus georgicus, Equus suessenbornensis,andStephanorhinus hund- the genus Camel, the first occurrence of the woodland elephant sheimensis, the cave or ‘spelaeoid’ bear close to U. deningeri,and Palaeoloxodon and woodland antelopes Tragelaphus and the first jaguar (close to P. onca gomboszoegensis)(Hemmer et al., Pontoceros , and the presence of Paracamelus (P. kujalnensis) 2010; Vekua, 1962, 1986). In Siberia, the diversity of musk-oxen (Alexeeva, 1977; Gromov, 1948; Vekua, 1962; Vereshchagin, increased. The ovibovine Soergelia occurred in the Razdolie as-

1957; and others). From the end of the early to the middle semblage (Western Siberia) with A. meridionalis, Palaeoloxodon

Pleistocene, true wolves attributed to the Canis mosbachensis– sp., Equus (Allohippus)sp.,Bison sp., and Ovibovini (Vangen- C. variabilis group and large hunting dogs Lycaon lycaonoides geim, 1977). The remains of Soergelia and the oldest members (¼ Canis (Xenocyon) lycaonoides) existed in Eurasia from Western of Praeovibos were found in the early stage of the Olyorian land Europe to Transbaikalia and China (Sotnikova and Rook, 2010). mammal age (Beringia, Bering Province) (1.4–0.8 Ma) together The Tamanian mammal assemblage is based on the fauna with those of the first wolverine Gulo minor (¼ G. cf. schlosseri) of Synaya Balka (type locality) and Tsymbal in the Taman (Sher, 1971, 1987; Sher et al., 2011; Vangengeim, 1977). These Peninsula. A. meridionalis tamanensis coexisted in the North co-occurred with Archidiskodon sp., Lycaon lycanoides, Equus Black Sea area with Homotherium crenatidens, the short-faced (Plesippus) verae, Alces sp., Bison sp., and others. The northern- hyena Pachycrocuta brevirostris, the horse Equus suessenbornensis, most finding of Homotherium in Asia is associated with this the rhinoceroses Stephanorhinus etruscus and Elasmotherium fauna. In Eastern Siberia, the Aldan fauna included a number caucasicum , and others (Alexeeva, 1977; Sotnikova and Titov, of forest inhabitants (Trogontherium cf. cuvieri, Palaeoloxodon ex 2009; Vereshchagin, 1959; Figure 5). Vereshchagin (1957) gr. namadicus, Alces latifrons,andMicrotus gregaloides). In Trans- also identified the potential impact of early hominins on baikalia, the Kudun (Kizhing–Kudun depression) and Zasu- some of these assemblages. And recently, stone artifacts of the khino (Itanza River basin) correspond to the youngest part of Oldowan type were reported from the Synyaya Balka site by the early Pleistocene and are correlated with the uppermost

Shchelinsky et al. (2010). In the Asian part of the territory, the Matuyama Chron (Erbaeva and Alexeeva, 2000). first reliable evidence of human activity comes from the stone The oldest Camelus (C. cf. knoblochi) coexisted with the tool assemblage of Kuldara (Ranov et al., 1995) dated to giant deer Praemegaceros (close to P. verticornis) in southern

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¼ Tadjikistan (Lakhuti locality, the uppermost part of the Early Middle Pleistocene: Middle–Late Galerian, Matuyama) and members of diverse habitats. They included Cromerian–Elsterian, 0.8–0.4 Ma inhabitants of open, dry, and savannah-like landscapes, such The early–middle Pleistocene boundary of the standard as the gerbil Meriones lacutensis and mole vole Ellobius lakhu- scheme corresponds to the Eopleistocene–Neopleistocene tensis, Archidiskodon sp., Equus cf. namadicus, and Canis cf. boundary of the Russian stratigraphic scheme and the Early– mosbachensis, in association with a large Lycaon, Pachycrocuta Middle Galerian boundary in Italy. brevirostris , Homotherium sp., the red-backed vole Clethrionomys, Panthera onca gomboszoegensis, the badger Meles ex gr. meles, and the giant deer Sinomegaceros sp. (Sotnikova, 1989; Sotnikova Tiraspolian mammal assemblages and its analogs and Vislobokova, 1990). Small mammals also included white- The considerable reorganization in the composition of mam- toothed shrew Crocidura sp., a hamster Cricetulus sp., the malian communities was related to a drop in temperature meadow voles Allophaiomys sp., and Microtus (Phaiomys) lachu- at the early–middle Pleistocene boundary, with a notable tensis (data from Zazhigin). increase in amplitude of climatic oscillations. Along with Faunas intermediate in age between the Tamanian and progressive cooling, climatic fluctuations changed from 41-ka Tiraspolian faunas are known from the lower course of the low-amplitude cycles to a 100-ka high-amplitude cycle

Dnester River (Karai-Dubina) and in the middle Don (Petro- (Shackleton, 1995). In the Russian Plain, three periods of pavlovka) area and have been studied in detail (Agadjanian, glaciation correspond to this stage (Pokrovka, Don, and Oka). 2009; Markova, 2007). The reversed polarity deposits contain The climatic zonation of the northern Eurasian landscape the remains of Prolagurus and pitymyoid Microtus, the last began to resemble that of the present day (Figure 6). Archidis- representatives of Allophaiomys, and the oldest Microtus with kodon was replaced by the mammoth Mammuthus trogontherii five closed enamel triangles in the first lower molar. (¼ A. wu¨sti) and the proportion of modern vertebrate genera

Thedispersal of mammals out of Northern Eurasia in- sharply increased. creased from about 1.2 to 1.1 Ma; Praemegaceros verticornis The representatives of the Tiraspolian mammal assemblage and Soergelia invaded Western Europe and Homo erectus and in southeastern Europe existed under a temperate climatic Sinomegaceros first occurred in China (Kahlke et al., 2011; Qiu, regime in a variety of landscapes, dominated by forest-steppes

2006). (Alexeeva, 1977). Kolkotova Balka (near Tiraspol) is the type

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Figure 6 Zonal occurrence of mammals in the early–middle Pliocene, with some data from Markova (2006): (1) steppe; (2) forest-steppe; (3) mixed and broadleaved forests; (4) Picea-Pinus and Betula forests, with some broadleaved plants; (5) Picea-Pinus forests and steppe; (6) forest–tundra; (7) tundra.

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Author's personal copy 612 VERTEBRATE RECORDS | Early and Middle Pleistocene of Northern Eurasia locality of this assemblage. The fossil remains were found in are recorded in Western Europe. The Bering land bridge (0.8– the so-called ‘Tiraspolian gravels’ (fluvial deposits of the Dnies- 0.011 Ma) further facilitated faunal exchanges between North- ter 6th terrace). Horses were represented by both caballoid ern Asia and North America. (Equus cf. mosbachensis) and stenonis (E. aff. suessenbornensis) forms. Mammuthus trogontherii, Stephanorhinus etruscus (late

Late Middle Pleistocene, ¼ Aurelian, ¼ Holsteinian–Saalian, form), the large deer Cervus acoronatus, Praemegaceros verticornis, the fallow deer Praedama and Alces latifrons, and a short- 0.4–0.126 Ma horned bison Bison schoetensacki are common to both the Singilian and Khasarian faunas and their analogs Tiraspolian fauna and the fauna of Western Europe (Kahlke The Singilian fauna and its analogs are related to the Likhvin et al., 2011; Nikiforova, 1971). Two skeletons of M. trogontherii interstadial (¼ Mindel-Riss, ¼ Holsteinian), one of the most were found in the Kagalnik sand pit (Azov). Data on small pronounced warm stages in the middle Pleistocene. In Eastern mammal fauna represent several phases of Tiraspolian mam- Europe, the Singilian faunas were represented by the straight- mal assemblages (see Markova, 2007). A considerable cooling tusked elephant Palaeoloxodon antiquus and Merck’s rhinoceros has been recognized in the southern part of European Russia at Stephanorhinus kirchbergensis. In the south of Northern Eurasia, the time of the early middle Pleistocene Don glaciations, gray wolf (Canis lupus), Camelus knoblochi, Megaloceros gigan- highlighted by the presence of subarctic species such as the teus, and Bison priscus longicornis became widespread after lemmings Lemmus ex. gr. Sibiricus and Dicrostonyx sp. and the the mid-middle Pleistocene (Alexeeva, 1977; Gromov, 1948). North Siberian vole Microtus ex gr. hyperboreus (Agadjanian, Typical small mammals include the water vole Arvicola mosba- 2009). chensis, the yellow steppe lemming Eolagurus luteus, the narrow- The southward shifting of ranges, with persistence of some skulled vole Microtus gregalis, the northern or root vole forms in refugia (including mountain ones), was usual during M. oeconomus, Ellobius, and the ground squirrel Spermophilus continental glaciations. Very diverse mammal faunas are (Markova, 2007). In Western Siberia, Palaeoloxodon coexisted known from the northern Caucasus. Most are associated with with Equus cf. steinheimensis, a small-sized cave bear Ursus savini Early Paleolithic sites and stone artifacts of Acheulian type. The rossicus (¼ Ursus spelaeus rossicus), and the giant deer Megalo- taxa include the macaque monkey Macaca cf. sylvana, the spot- ceros giganteus during the contemporaneous Tobol interstadial ted hyena Crocuta spelaea, diverse bears including cave bear (Vangengeim, 1977; Figure 7). From this time onward Ursus deningeri (dominant), black bear Ursus ex gr. thibetanus (400 ka), the giant deer became widespread in the temperate probably related to U. etruscus, and the small cave bear U. savini latitudes of Eurasia, together with modern deer species (red rossicus (¼ U. aff. spelaeus rossicus), the cave lion Panthera spelaea deer Cervus elaphus and the reindeer Rangifer tarandus). and leopard P. pardus, the gray wolf Canis lupus, the fox Vulpes The Khasarian faunal assemblage (Dnepr glaciation) with vulpes, Stephanorhinus , Megaloceros giganteus, Capreolus cf. suessen- its type locality at Cherny Yar (Volga River Basin) contained bornensis, Bison cf. schoetensacki, together with typical montane Mammuthus trogontherii chosaricus (ancestor of the woolly genera such as the ibex Rupicapra and the goat Capra. mammoth Mammuthus primigenius), Panthera spelaea, Elas- The fauna of the late stage of the Olyorian faunal assem- motherium sibiricum, the woolly rhinoceros Coelodonta antiqui- blage (West Bering Province) are characteristic of tundra and tatis, Equus chosaricus, Camelus knoblochi, the long-horned bison forest–tundra environments and cold climatic conditions sim- Bison priscus longicornis, Megaloceros giganteus, and modern ilar to modern ones (Sher, 1971, 1987; Vangengeim, 1977). genera and species, including Cervus elaphus, Rangifer tarandus, This fauna included the first reindeer Rangifer. Alces aff. latifrons the musk-oxen Ovibos, the saiga antelope Saiga, etc., the water and Bison sp. were also present in the artiodactyl community. vole Arvicola chosaricus, and a diverse assemblage of steppe- Rangifer is also recorded in the early middle Pleistocene of the adapted small mammals (Alexandrova, 1976; Dubrovo, 1985; Kuznetsk Depression (Foronova, 2001). Gromov, 1948). The fauna existed in the colder and drier In Transbaikalia, the Tologoi assemblage contained the woolly rhino Coelodonta tologoijensis, Equus ex gr. sanmeniensis, Cervus ex gr. elaphus, Spirocerus cf. peii, and Bison sp. (Vangengeim, 1977). Cryogenic disturbance at the base of the deposits is associated with remains from the Zasuhino assemblages (Zasuhino 3, 1–0.78 Ma) and in Tologoi 2 beneath the

Brunhes–Matuyama magnetic reversal, above the Jaramillo (Alexeeva, 2005). The fauna is characterized by a large diversity of carnivores and ungulates: Nyctereutes sp., Canis variabilis, Lycon cf. lycaonoides (¼ Xenocyon lycaonoides), Ursus sp., Gulo sp., Pachycrocuta brevirostris, Homotherium sp., the large-sized tiger Panthera ex gr. tigris, Coelodonta tologojensis, Equus ex gr. sanmeniensis, Equus ex gr. suessenbornensis–verae, Alces cf. latifrons, and others (Vangengeim et al., 1990). Zasuhino 3 locality also has yielded Cervus sp., Megacerini indet, Capreolus cf. suessenbornensis, Ovibovini indet., and Bison sp.

At the boundary of the early and middle Pleistocene, a Figure 7 Megaloceros giganteus, C. Flerov’s painting exhibited in the prominent climatic deterioration was marked over northern Paleontological Museum of the Paleontological Institute, Russian and central Eurasia and many immigrants from these regions Academy of Sciences (PIN), Moscow, with PIN permission.

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Author's personal copy VERTEBRATE RECORDS | Early and Middle Pleistocene of Northern Eurasia 613 climate of the periglacial forest-steppe along the southern margin became more widespread in the early middle Pleistocene of the continental Dnieper glaciation (Riss, RI) (Vangengeim, (true wolves, tiger Panthera tigris) and at the end of late middle 1977). However, localities rich in large mammal are rather Pleistocene (Khasarian), with the first appearance of modern rare. In the south of the Russian Plain, typical subarctic and species (Cervus elaphus, Rangifer tarandus, Ovibos, Saiga, etc.). steppe small mammals occurred (Dicrostonyx simplicior, Lemmus Zoogeographic provinciality therefore increased from the Early sibiricus, Lagurus ex gr. lagurus, Microtus gregalis). In the toward the late middle Pleistocene. Caucasus region, the tar pit Binagady fauna near Baky belongs to the same stage. Under modern conditions, the range of some animals became completely separated and confined to different See also: Vertebrate Records: Early Pleistocene; Late Pleistocene biomes. Megafaunal Extinctions; Late Pleistocene Mummified Mammals; Late Pleistocene of Africa; Late Pleistocene of North America; Late

Pleistocene of South America; Late Pleistocene of Southeast Asia; Mid- Conclusions Pleistocene of Africa; Mid-Pleistocene of Europe; Mid-Pleistocene of Southern Asia. The data on the early and middle Pleistocene mammals from

Northern Eurasia offer important insights into our knowledge of the history of mammals. These assemblages differ considerably in their composition as a result of evolutionary and mi- References gration processes. In the Pleistocene, the general trend toward global temperature decrease, changes in landscape and climatic Agadjanian AK (2009) Small Mammals of the Pliocene–Pleistocene of the Russian conditions, and differentiated environmental patterns caused Plain. Moscow: Nauka (in Russian). Alexandrova LP (1976) Rodents of the Anthropogene of the European Part of the USSR. important changes in mammal communities within the terri- Moscow: Nauka (in Russian). tory at the suprageneric, generic, and species levels. The faunas Alexeeva LI (1977) Early Anthropogene Theriofauna of East Europe. Moscow: Nauka demonstrate adaptation from the temperate climatic condi- (in Russian). tions of the pre-Pleistocene to the cooler and more diverse Alexeeva NV (2005) Environmental Evolution of Late Cenozoic of West Transbaikalia conditions at the end of the Pleistocene. A gradual disappear- (Based on Small Mammal Fauna). Moscow: GEOS (in Russian). Alexeeva NV and Erbajeva MA (2008) Diversity of Late Neogene–Pleistocene small ance is therefore noted of the Miocene–Pliocene and ther- mammals of the Baikalian region and implications for paleoenvironment and mophilous forms (mastodons, giraffids, hippopotamids, biostratigraphy: An overview. Quaternary International 179: 190–195. hyenids) with their extinction or displacement to southern Alexeeva NV, Karasev VV, and Erbajeva MA (2007) Pleistocene biostratigraphy of the Asia orAfrica, paralleled by the development of Pleistocene Transbaikal area (South East Russia). Courier Forschungsinstitut Senckenberg II: 19–26. and modern genera and species, and the gradual emergence of Bajgusheva VS (1971) Fossil theriofauna of Liventzovka sand pit (northeastern of the associations typical for modern biomes (tundra, steppes, for- Sea of Azov). In: Bykhovskii BE (ed.) Materials on Faunas of the Anthropogene of est, etc.). the USSR, pp. 5–28. Leningrad: Nauka (in Russian). The main periods of turnover in mammal communities of Bajgusheva VS, Titov VV, and Tesakov AS (2001) The sequence of Plio–Pleistocene Northern Eurasia coincided approximately with the maximum mammal faunas from the south Russian Plain (the Azov Region). Bollettino della Societa Paleontoogica Itaiana 40(2): 133–138. drops in global temperature, which were accompanied by Cande SC and Kent DV (1995) Revised calibration of the geomagnetic polarity timescale large-scale transformations of the environment. The most sig- for the Late Cretaceous and Cenozoic. Journal of Geophysical Research, Series B nificantchanges occurred at the Pliocene–Pleistocene (2.6 Ma) 100(4): 6093–6095. and early–middle Pleistocene (0.8 Ma) boundaries. Changes Derevyanko AP and Shunkov MV (2005) Early Paleolithic site Karama in Altai Region: First results. Archaeology, Ethrnography and Anthropology of Eurasia 3(23): 52–69 in faunal composition were also reported at 1.8, 1.2, and (in Russian).

0.4 Ma. Beginning in the middle Pleistocene, the turnover Dmitrieva EL (1977) Neogene Antelopes of Mongolia and Adjacent Territories. Moscow: affected mammal diversity at the generic and species levels. Nauka (in Russian).

Dmitrieva EL and Nesmeyanov SA (1982) Mammals and Stratigraphy of Continental Immigration to and from adjacent territories played an important role in these faunal transformations. Tertiary Deposits of Southeastern Middle Asia. Moscow: Nauka (in Russian). Dodonov AE (2002) Quaternary of Middle Asia: Stratigraphy, Correlation, The roots of some modern genera are found in the late Paleogeography. Moscow: GEOS (in Russian). Miocene (Canis, Axis, etc.), the Pliocene (Nyctereutes, Ursus, Dubrovo IA (1990) The Pleistocene elephants of Siberia. In: Agenbroad LD, Mead JI, Lynx, Equus ), the Gelasian (Gulo, Cervus, Alces, Capreolus, and Nelson LW (eds.) Megafauna and Man, vol. 1, pp. 1–8. South Dakota: Hot

Bison), and the middle Pleistocene (Camelus, Rangifer, Ovibos, Springs. Erbaeva MA and Alexeeva NV (2000) Pliocene and Pleistocene biostratigraphic Saiga). The Pliocene–Pleistocene boundary is marked by the succession of Transbaikalia with emphasis on small mammals. Quaternary disappearance of a number of thermophilous forms. Evolu- International 68–71: 67–75. tionary trends can be traced equally in the large herbivorous Foronova IV (2001) Quaternary Mammals of the South-east of Western Siberia animals (elephants, rhinoceroses, horses, artiodactyls), various (Kuznetsk Basin): Phylogeny, Biostratigraphy, and Paleoecology. Novosibirsk: SB carnivores (canids, ursids, hyenids, felids), and small mam- RAS GEO (in Russian). Gabunia L and Vekua A (1995) A Plio–Pleistocene hominid from Dmanisi, East Georgia, mals. A clear boreal group, which comprised the ancestral Caucasus. Nature 373: 509–512. forms of modern Northern Eurasian species, started to form Garutt VE and Bajgusheva VS (1981) Archidiskodon gromovi Garutt et actively at the beginning of the late early Pleistocene, around Alexeeva – der altyeste Elephant der Mammutlinie in Eurasien. Quarta¨rpala¨ontologie 1.8 Ma. The increasing amplitude of global cooling was then 4: 7–18. Gliozzi E, Abbazzi L, Argenti P, et al. (1997) Biochronology of selected mammals, accompanied by the appearance and widespread distributions mollusks and ostracods from the Middle Pliocene to the Late Pleistocene of first boreal and then arctic forms in northern Siberia and the in Italy. The state of the art. Rivista Italiana di Paleontologia e Stratigrafia mountainous regions of Central Asia. Boreal communities 103(3): 369–388.

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