Likhvin Interglacial Small Mammal Faunas of Eastern Europe

ARTICLE IN PRESS

Quaternary International 149 (2006) 67–79

Likhvin Interglacial small mammal faunas of Eastern Europe

Anastasia K. Markova

Institute of Geography Russian Academy of Sciences, Staromonetny per., 29, Moscow 109017, Russia

Available online 30 January 2006

Abstract

Investigations of Middle Pleistocene deposits over the past several decades produced a significant mammal database that can be used to evaluate the complicated history of this interval in Eastern Europe. The most significant late Middle Pleistocene warm interval was the Likhvin Interglacial; among the stratigraphic horizons attributed to this interval are the Inzhavino fossil soil, alluvium of the fourth terraces of the main Eastern European rivers, and the Early Euxinian marine deposits of the Black Sea. Fifteen small mammal localities of Likhvin age have been recovered in different regions of the Russian Plain. The stratigraphical position of the faunas from key sections is quite definite: they occur above the Oka Glacial deposits and under the Kamenka and Romny late Middle Pleistocene fossil soils and Dnieper glacial deposits. These faunas are characterized by Arvicola cantianus, Lagurus transiens, Microtus (Pallasiinus) oeconomus, M. (Microtus) ex gr. arvalis, M. (Microtus) agrestis, and M. (Stenocranius) gregalis. Neither the rhizodont voles Mimomys and Pliomys, nor Microtus (Terricola) arvalidens and Microtus (Stenocranius) gregaloides have been recovered from these localities. According to recent data from Western Europe, the similar species composition and stratigraphical position are typical for the Holsteinian Interglacial mammal faunas. These faunas come from deposits covered by late Middle Pleistocene sediments and are correlated to OIS Stage 11. r 2005 Elsevier Ltd and INQUA. All rights reserved.

1. Middle Pleistocene small mammal faunas of the Likhvin Picea. Such a composition suggests warm interglacial Interglacial conditions at the time of formation of the lake deposits. Later several sections with a similar plant species composi- 1.1. The history of investigation of the Likhvin deposits tion were found on the Russian Plain. Integrated studies of the Likhvin section were performed during the past Likhvin Interglacial deposits were first described in the decades by stratigraphers and paleontologists (Sudakova, Likhvin stratotype section located in the Oka drainage 1975; Agadjanian, 1977; Bolikhovskaya and Sudakova, basin near Chekalin town (formerly Likhvin town). The 1996). Recently, Oka Glacial deposits have been found not section, which is unique in many respects, has been under in the main section but near the stratotype. Deposits of the investigation since the end of the 19th century. Its main earlier Don Glaciation have been found in the base of the stratigraphical features were described by Bogolubov Likhvin section (Figs. 1 and 3, Likhvin). Agadjanian and (1904), Moskvitin (1931), Markov (1939) and Ushko Alexandrova described the fauna of small mammals, which (1959). These authors established two horizons of glacial were found in lake sapropel deposits with an interglacial deposits, an interlaced complicated suite with ancient flora (Likhvin Interglacial deposits) and also in other interglacial lake deposits, and overlaying stratified and layers of this stratotype area (from Dnieper Glacial unstratified deposits. The lower glacial horizon was deposits and from Oka Glacial strata) (Agadjanian, 1977; correlated with the Oka Glaciation and the upper till was Alexandrova, 1982). referred to the Dnieper Glaciation (Markov et al., 1965). Grichuk (1961, 1989) described the pollen spectra derived 1.2. Geographical distribution of the Likhvin small mammal from the interglacial lacustrine deposits and established the localities specific species composition, including broadleaved plants (Quercus, Tilia, Ulmus, Carpinus), as well as Abies and Likhvin faunas were recovered from various parts of the Russian Plain, including the Oka, Don, Dnieper, E-mail address: [email protected]. Danube, Prut, Kama, and Kuma river drainage basins.

Fig. 1. The geology of Likhvin (Chekalin) stratotype section (after Bolikhovskaya and Sudakova, 1996).

The northernmost localities were found at approximately The fauna from the Likhvin stratotype section (sapropel 571N and the southernmost on the Black Sea coast deposits) includes Desmana sp., Apodemus cf. sylvaticus, (451N) and in the Kuma drainage basin in the Northern Clethrionomys glareolus, Microtus malei and water vole Caucasus (441N) (Fig. 3). Arvicola cantianus ( ¼ mosbachesis) remains, which is characterized by a Mimomys—like enamel structure of 1.3. Small mammal species composition the teeth and by small sizes (Agadjanian, 1977). The vole described by A. Agadjanian as Microtus malei should, in The Likhvin small mammal faunas have been studied by the opinion of the author, be referred to the Microtus several paleontologists (Agadjanian, 1977; Markova, 1982, arvalis group. Rhizodont voles Mimomys and Pliomys, 1992, 1998; Motuzko, 1985; Rekovets, 1994). and also microtines Microtus (Terricola) arvalidens and ARTICLE IN PRESS A.K. Markova / Quaternary International 149 (2006) 67–79 69

Microtus (Stenocranius) gregaloides have not been found Several sites with Likhvin-like fauna were found in the in this locality (Tables 1and 2; Fig. 2). Don drainage basin (Verkhnaya Emancha, Strelitsa, In the Dnieper drainage basin, a similar fauna is known Vladimirovka, Mikhailovka 3) (Table 1, Figs. 2 and 3) from the Gunki locality. It occurs in fluvial deposits of the (Agadjanian, 1977; Markova, 1982, 1992; Agadjanian and fourth terrace of the Psel River, the left tributary of the Erbaeva, 1983; Agadjanian and Glushankova, 1986). The Dnieper (Fig. 3). The layer with bones is overlain by a thick fossiliferous layers occur under the Kamenka and Romny loess–paleosol sequence and also by glacial deposits of paleosols (Table 2). However, Dnieper till has not been Dnieper age. Between the Dnieper till and the fossiliferous recorded in these sites, as the Dnieper ice-sheet did not layer, there are two well-pronounced paleosols and two penetrate to the Don drainage basin. The species composi- loess horizons. These soils have been described as tion of these localities is very close to the Likvin faunas Kamenka (lower) and Romny (upper) soils (Velichko situated in the Dnieper and Oka basins (Table 1). et al., 1997). Sapropel, containing small mammal remains, Three localities of similar species composition were yielded pollen spectra typical for the Likhvin Interglacial found on the southwestern part of the Russian Plain (Gubonina, 1980). Malacofauna recovered from the same outside the Dnieper ice-sheet (Tiraspol, Inzhavino paleosol horizon was correlated with the Early Euxinian transgres- layer, Dniester basin; Uzmari, Prut basin; Ozernoe, sion of the Black Sea by Chepalyga (Velichko et al., 1997). Danube basin) (Tables 1 and 2; Figs. 2 and 3)(Mikhailesku The Gunki small mammal fauna is significantly richer than and Markova, 1992). In the Tiraspol locality the small the fauna from the Likhvin section and contains about mammal bones came from mole burrows of the Inzhavino 1500 identified bones of small mammals, mostly rodents fossil soil horizon, which is correlated with the Likhvin (Markova, 1982). This fauna includes abundant remains of Interglacial. The full Pleistocene loess–paleosol sequence is the water vole Arvicola cantianus. Lagurids are represented represented in the Tiraspol section. Here the Inzhavino soil by the advanced type of steppe lemming Lagurus transiens. is overlain by four paleosols alternating with loess horizons Rhizodont voles of Mimomys and Pliomys genera and also (Briansk, Mikulino, Romny, and Kamenka fossil soils) and Terricola-like microtines such as Microtus (Terricola) overlies the loess horizon, the Vorona fossil soil and fluvial arvalidens and Microtus (Stenocranius) gregaloides are deposits of the Dniester river (Mikhailesku and Markova, lacking here (Tables 1 and 2). 1992; Dodonov et al., this volume). The Ozernoe and The fauna of this evolutionary level was suggested to be Uzmari fluvial deposits with small mammal remains lie described as the Gunkovian assemblage (Markova, 1990), under the complicated loess–paleosol series with a number which is presumably synchronous with the Singilian large of fossil soils. Thus, in the Ozernoe locality, the alluvium of mammalian age, determined by Gromov (1948) and the fourth Danube terrace with bone remains is covered Alexeeva (1977). Unfortunately, the stratotype of the by a loess–paleosol sequence, including four fossil soils Singilian fauna in the Volga drainage basin near the town (the Late Pleistocene Briansk and Mikulino soils, and of Raigorod contains only few small mammal remains, also the late Middle Pleistocene Romny and Kamenka collected by L. Alexandrova in 1965 (Number 824 of the soils) (Mikhailesku and Markova, 1992; Markova, 1996) Geological Institute RAS collections). Among these are a (Tables 1 and 2). transitional form of Lagurus transiens—L. lagurus, Eola- The most southeastern locality, Otkaznoe, was found in gurus luteus, Microtus (Stenocranius) gregalis, Microtus mole burrows of the Inzhavino paleosol in the middle arvalinus, Microtus oeconomus, Ellobius sp. and Spermo- Kuma river drainage basin, near Georgievsk town, North- philus sp. (Tesakov, pers. comm.). Unfortunately, Arvicola ern Caucasus (Fig. 3). Its fauna includes only species remains are absent in this fauna. The evolutionary level of remains of open landscapes (ground squirrels, steppe and lagurids and microtines would indicate that the Raigorod yellow lemmings, mole rats, etc.). The taphonomy of the fauna is comparable with that from the Gunki section. It fossiliferous layer (fossil soil) excludes the possibility of was the latter, however, that gave its name to the small finding the remains of water voles (Table 1). The Otkaznoe mammal assemblage, corresponding to the Likhvin Inter- section is very complicated in structure. The loess–paleo- glacial; such a choice may be explained by the paucity of sol–fluvial sequence is more than 130 m thick. The the Raigorod small mammal fauna compared with the rich Inzhavino soil horizon lies below the Kamenka, Romny, Gunki fauna. Besides, the Gunki locality occupies a very and Mezin (Mikulino) fossil soils and intermediate loess definite stratigraphical position and has been thoroughly layers (Bolikhovskaya, 1995). studied by a variety of geological and paleontological The northernmost localities are situated at the mouth of methods. the Kama river (Fig. 3, Rybnaya Sloboda) (Markova, Two other localities, similar in species composition of 1992), in the Niamunus drainage basin (Fig. 3, Niariavai-2) small mammals, have been found near the Gunki locality (Vozniachuk et al., 1984) and in the Zapadnaya Dvina in the Middle Dnieper drainage basin: Chigirin and Pivikha drainage basin (Fig. 3, Smolenskii Brod) (Vozniachuk (Gradizhsk) (Tables 1 and 2; Fig. 3). These localities are et al., 1979). The core of all of these faunas includes also overlain by Dnieper till and underlain by a loess–pa- Arvicola cantianus and abundant Microtus oeconomus, leosol series, and are referred to the fluvial deposits of the M. ex gr. arvalis and M. agrestis (Tables 1 and 2; Fig. 2). fourth terrace of Dnieper (Markova, 1982). In the opinion of Motuzko (1985), the Smolenskii Brod ARTICLE IN PRESS 70 A.K. Markova / Quaternary International 149 (2006) 67–79 a Otkaznoe a Uzmari a + + Tiraspol (Inzhavino soil) a +++ + Ozernoe +++ a ++ Pivikha a + Gunki I a +++ +++++ + Chigirin + c + Vladimirovka 2 c +++++++ Strelitsa a . ++++ + + Verkhnaya Emancha c + Mikhailovka 3 + a . ++ +++ +++ +++++ + Rybnaya Sloboda + + b Neravai Agadjanian and Glushankova (1986) ; b Vozniachuk et al. (1984) + Smolenskii Brod ; c Likhvin (gittia) al+ +Pall+++ ++++ Selys- . Kretzoi Agadjanian and Erbaeva (1983) sp. Poljak. ; al ++ +++++++Pall+ + Vozniachuk et al. (1979) Pall +++ +++++ Alexandrova + + + + + + + ; ldenstaedt Fischer + ¨ Schreber + + + + + + L. + + + oeconomus Pall Kerr. Gu Pall arvalis sp. + Kerr. itn+++++++ Hinton++++++++++++ cuvieri Janossy + Schaub ex gr. Pall L. ++ ++ ++ + Pall major L. + middendorffi L. lagurus ex gr. cf. L. Sundervall + sp. L. + Lemmus sp. + + + + s Vinogr. sp. ex gr. sp. + + + + + ex gr. ex. gr. vel sp. sp. + sp. sp. + ++++++ + Motuzko (1985) Markova (1982, 1998) Agadjanian (1977) a b c crotus Ochotona Ochotona pusilla Rodentia Sciurus vulgaris Spermophilus (Spermophilus) M. (Microtus) agrestis Spermophilus Spalax Spalax microphtalmus Lagurus Eolagurus luteus volgensis Arvicola cantianus Microtus (Terricola) subterraneus Longchamps M. (Stenocranius) gregalis M. (Pallasiinus) Trogontherium Castor fiber Castor Alactagulus pumilio Clethrionomys glareolus Apodemus Cricetulus migratorius Allocricetus ehiki Cricetus cricetus Myopus Cl. rufocanus Lagurus transiens Ellobius talpinus Lagomorpha Lepus Sorex araneus Sorex Table 1 Species composition of Likhvin small mammal faunasSpecies of Eastern Europe Insectivora Desmana Talpa europaea Localities Allactaga M. (Microtus) Microtus (Terricola) arvalidens hyperboreu Mi ARTICLE IN PRESS A.K. Markova / Quaternary International 149 (2006) 67–79 71 2 Suvorovo, up. Tiraspol/ middle layers (Danube), Eastern Europe small mammal localities (with indication of river drainage basins) & Alpat’evo (Oka) Igorevka (Seim) Plavni (Danube), Priluki (Dnieper), Rasskazovo (Oka-Don Plain), Uzunlar (Black Sea coast) Rybnaya Sloboda (Kama ), Verkhnia Emancha (Don) Likhvin (sapropel) (Oka), Uzmari (Danube), Ozernoe (Danube), Inzhavino paleosol/ (Dniester), Chigirin (Dnieper), Pivikha (Dnieper), Gunki 1 & (Dnieper) Chekalin (Oka Glac. deposits) (Oka) Posevkino, Perevoz (Don) Troitsa 1 (Oka), Bogdanovka (Don) Malakhovka (L. 8), Konakhovka 2 (Oster) (Depth 35.0–38.0 m) (Depth 38–39.5 m) (Depth 39.3–39.6 m); Konakhovka 2 (Depth 39.5–40.5 m), Matveevka (Dnieper) mammal species in Eastern Europe Dicrostonyx simplicior Microtus (Stenocra-nius) gregalis Arvicola cantianus Dicrostonyx simplicior okaensis Arvicola chosaricus Lagurus lagurus Dicrostonyx sp., Microtus hyperboreus Gunkovian ¼ Mammalian ages The first appearance of small Khozarian ) O 18 O/ stages 6 16 9 11 Singilian 13 Late Tiraspolian Shik, 1993 a a Isotope Stages (after and alluvial horizons Interglacial Konakhovka Optimum Podrudniansk cooling 14 Konakhovka 2 (Oster) Interglacial ) Stratigraphical horizons and Oxygen Mertzalovsk loess Moscow Glaciation Kursk paleosol Tsninsk loess Breslav et al., 1992 The author of a recent paper has a different opinion about the stratigraphical position of these suites. a Dnieper (Moskva) Glaciation Table 2 Stratigraphical position of Middle Pleistocene small mammalStratigraphy sites (after in Eastern Europe The underlined localities have been studied by the author of this paper. Glaciations, Interglacials Paleosol–loess deposits Glacial, Interglacial Romny warming Romny paleosolBorisoglebsk CoolingLikhvin Interglacial Borisoglebsk loess Inzhavino paleosolOka GlaciationMuchkap Interglacial Odintsovo Interglacial 7 Korystylevo loess Cooling Smolenski Brod Don Glaciation 10 Don loess Oka Glaciation 12 Don Glaciation — Topka (Don) 16 Mikhailovka 2 (Svapa), Roslavl Seria Vorona paleosol Muchkap Interglacial Glazovo optimum 15 Konakhovka 1 (Oster) Orchik coolingKamenka Interglacial Kamenka paleosol Orchik loess Likhvin s.s. Cooling 8 ARTICLE IN PRESS 72 A.K. Markova / Quaternary International 149 (2006) 67–79

Fig. 2. SDQ index of Arvicola m/1 from Western and Eastern European Middle and Late Pleistocene localities: * studied by A. Markova; ** studied by A. Agadjanian; *** studied by L. Rekovets. fauna belongs to an earlier stage than other Likhvin northern part of Eastern Europe and the whole of Siberia. faunas. The mean SDQ index of the Smolenskii Brod This is the only find of this species in the Middle Arvicola is about 124 (n ¼ 16) (Vozniachuk et al., 1984; Pleistocene faunas of the Russian Plain. Table 1). The value of this index falls within the limits typical for Likhvin Arvicola from other sites of Eastern 1.4. Arvicola SDQ index Europe. The author had the opportunity to examine the materials from Smolenskii Brod and did not see a The differentiation of Arvicola enamel is one of the difference between the Arvicola morphology from Smo- important features characterizing the evolutionary level of lenskii Brod (enamel index and sizes of teeth) and the water Arvicola remains. The higher values of this index, together vole remains from other Likhvin localities. with the smaller sizes of the teeth, indicate a more primitive The Rybnaya Sloboda fauna is unique by its north- stage of Arvicola (Markova, 1975, 1981; Heinrich, 1978, eastern location. This geographical position is reflected in 1990). However, modern Arvicola from different regions the specificity of its special composition, which includes not show different SDQ indices, which depend on the only abundant remains of Arvicola cantianus, Clethrion- latitudinal position of the population area. This feature is omys glareolus, Lagurus transiens, Microtus ex gr. arvalis, more primitive in southern regions (Ro¨ttger, 1987). This M. oeconomus,andM. agrestis, typical for this time, but fact must therefore be taken into account when analyzing also Clethrionomys rufocanus (Table 1). The modern fossil Arvicola remains from different regions. The Likhvin distribution area of Clethrionomys rufocanus covers the Arvicola SDQ index from Eastern Europe varies between ARTICLE IN PRESS A.K. Markova / Quaternary International 149 (2006) 67–79 73

150–125–100 (Fig. 2). Mean index values are: 130 (Gunk’ki omys glareolus, Arvicola cantianus, Microtus (Microtus) ex 2, Chigirin and Uzmari); 125 (Gunki 1, Ozernoe, Verch- gr. arvalis, Microtus (Microtus)exgr. agrestis, and naya Emancha, and Rybnaya Sloboda). The Strelitsa Microtus (Pallasiinus) oeconomus. The forest biome can remains have an SDQ index between 138–110–100 be reconstructed for this territory. (n ¼ 8). Unfortunately, the SDQ index of Arvicola The forest-steppe assemblage of small mammals was teeth from the Likhvin (Chekalin) section have not placed south of the previous one. A high diversity of published yet. mammals was reconstructed for this assemblage with many Western European Arvicola from Bilzingsleben 2 have an steppe species (Ochotona pusilla, Spermophilus, Marmota SDQ index of teeth very similar to the Likhvin water voles bobac, Allactaga major, Spalax microphtalmus, Cricetus from Eastern Europe (Fig. 2). Lower values of SDQ are cricetus, Cricetulus migratorius, Lagurus transiens, Eola- characteristic for Miesenheim I (155–140–123). These gurus luteus volgensis, Microtus (Stenocranius) gregalis), data are also supported by a species composition in this some forest mammals (Clethrionomys glareolus, Microtus locality that differs from Holsteinian ( ¼ Likhvinian) (Microtus) agrestis), and intrazonal species (Desmana, ones. Mosbach 2 Arvicola have an SDQ limit between Sorex, Lepus europaeus, Arvicola cantianus, Clethrionomys 159–133–117 (n ¼ 45) (Maul et al., 2000). The SDQ glareolus, Microtus (Microtus) ex gr. arvalis, Microtus index of Arvicola teeth from Kamenka Interglacial (Microtus) agrestis, Microtus (Pallasiinus) oeconomus). localities (Khozarian faunas) appears to be different. It is The southern limit of this biome was extended to the Black about 110 in the localities Chernyi Yar, Plavni, and Sea coast in the western part of the Russian Plain, and Uzunlar (Fig. 2), which would suggest another evolution- shifted northward in the eastern part of the plain. Few ary level for these faunas. Stratigraphical data support small mammal Likhvin localities have been found in the their later age. southernmost territories of Eastern Europe. The existence The tendency in SDQ index characteristics is rather clear of the steppe assemblage of small mammals with Spermo- during the Middle and Late Pleistocene. However, it philus, Allactaga major, Pygeretmus pumilio, Sicista subtilis, should be noted that this coefﬁcient cannot be the only Spalax, Ellobius talpinus, Cricetus cricetus, Allocricetus argument in support of a certain age of faunas, because it ehiki, Arvicola cantianus, Eolagurus luteus volgensis, La- varies considerably within one locality and therefore gurus transiens, Microtus socialis, Microtus obscurus can be depends on the quantity of analyzed materials. Besides, it reconstructed with these data. also depends on the geographical position of the site. Only comprehensive information on species composition, the evolutionary level of the small mammals in different 2. Middle Pleistocene small mammal faunas older than the lineages, and other data obtained by geological and Likhvin Interglacial paleontological methods could determine conclusively the stratigraphical position of the fauna. 2.1. Faunas of the Oka Glaciation

1.5. Eastern European zoogeographical provinces during the Only few localities have been found in the deposits of the Likhvin Interglacial Oka Glaciation on the Russian Plain, which preceded the Likhvin Interglacial (Table 2). The new species of pied Arboreal faunas of small mammals have been found in lemming Dicrostonyx okaensis was described in the the Smolenskii Brod and Neravai localities, situated in the materials recovered from the Oka glacial deposits in west of the Russian Plain (Table 1). They include Sciurus the Likhvin stratotype region (Alexandrova, 1982). In the vulgaris, Trogontherium cf. cuvieri, Castor fiber, Apodemus opinion of A.K. Agadjanian and the author of this paper, sp., Clethrionomys glareolus, Microtus (Terricola) subter- this taxa has a subspecies level: Dicrostonyx simplicior raneus and Microtus agrestis. Abundant remains of okaensis. Arvicola cantianus make up the core of these faunas. The The locality of Mikhailovka 2 (Svapa drainage basin), absence of lagurids could be explained by the existence of a which is also correlated to the Oka Glaciation, includes forest zone in this geographical belt during the Likhvin Ochotona sp., Spermophilus sp., Clethrionomys cf. glareo- Interglacial. The species composition of the Likhvin lus, Pitymys sp., Lemmus sp., Dicrostonyx simplicior mammal faunas gives us an insight into the main features okaensis, Lagurus transiens, Microtus (Stenocranius) cf. of mammal assemblages and biomes of Eastern Europe gregalis (Agadjanian and Glushankova, 1986). Unfortu- and indicates the existence of natural zonality (Fig. 3). The nately, both localities do not include Mimomys position of all reconstructed biomes was sub-latitudinal. remains, nor those of Arvicola. The core of the Mikhai- The small mammal assemblage of mixed and broadleaved lovka 2 fauna is formed by Dicrostonyx simplicior okaensis forests was located between 60 and 501N and included a and the narrow-skulled vole M. gregalis. The Lagurus sizeable amount of forest species, together with several transiens teeth found in Mikhailovka 2 have a meadow and intrazonal mammals: Desmana cf. moschata, primitive structure. Some of the teeth look like those of Soricidae, Sciurus cf. vulgaris, Trogontherii cf. cuiveri, Prolagurus posterius (Agadjanian and Glushankova, 1986; Castor fiber, Apodemus flavicollis, A. sylvaticus, Clethrion- Fig. 6). ARTICLE IN PRESS 74 A.K. Markova / Quaternary International 149 (2006) 67–79

Fig. 3. Small mammal assemblages of the Likhvin Interglacial and some vegetation communities: (I) Picea–Pinus and Betula forests, with some broadleaved plants in the south (Grichuk, 1989), the mammal data are absent. Mammal assemblages of: (II) mixed and broadleaved forests; (III) forest- steppe; (IV) steppe; (V) mountain forest-steppe; (VI) mammal localities. Mammal localities: (1) Neravai-2; (2) Smolenskii Brod; (3) Rybnaya Sloboda; (4) Likhvin (Chekalin); (5) Mikhailovka 3; (6) Pivikha (Gradizhsk); (7) Gunki; (8) Chigirin; (9) Verkhnaya Emancha; (10) Strelitsa; (11) Vladimirovka 2; (12) Tiraspol (Inzhavino soil ﬁnds); (13) Uzmari; (14) Ozernoe; (15) Otkaznoe.

2.2. Faunas of the Muchkap (Roslavl) Interglacial Similar faunas with very progressive Mimomys savini (sometimes without roots), Prolagurus posterus, Lagurus Muchkap (Roslavl) Interglacial deposits predate the Oka transiens (early type), Microtus (Stenocranius) gregaloides, strata on the Russian Plain and revealed a complicated M. (S.) gregalis, M. oeconomus, M. (Terricola) arvalidens, history with two warm optimums (Konakhovka and M. ex gr. arvalis were found on the southeast of the Glazovo) and the intermediate Podrudniansk cool phase Russian Plain in the upper and middle layers of Suvorovo (Table 2). The localities of small mammal remains from all locality (Danube basin) (Mikhailesku and Markova, 1992). suites of the Muchkap deposits include very advanced Spermophilus (Citellus) sp., Allactaga ex gr. major, Mimomys intermedius ( ¼ M. savini). Arvicola remains Spalax microphtalmus, Allocricetus ehiki, Cricetus cricetus, have not been recovered from these layers (Agadjanian Lagurus transiens, Eolagurus luteus volgensis, and Microtus et al., 1988; Birukov et al., 1992). Sorex sp., Desmana (Stenocranius) gregalis were recovered from the mole cf. moschata, Spermophilus sp., Cricetus ex gr. cricetus, burrows of the Vorona paleosol in the Tiraspol section, Mimomys intermedius, Clethrionomys sp., Microtus (Ste- Dniester drainage basin (Mikhailesku and Markova, 1992). nocranius) gregaloides, M. oeconomus, M. cf. hyperboreus, Two small mammal localities in the Vorona paleosol mole and Lemmus sp. have been found in the upper part of the burrows were found in the Don basin (Posevkino and Konakhovka suite (the upper warm optimum of the Perevoz); they include few remains of Microtus (Terricola) Muchkap Interglacial) (Birukov et al., 1992). To date, arvalidens and M. (Stenocranius) gregaloides. The core of there are no localities of the Muchkap Interglacial in these faunas consists of Lagurus transiens (early type), Eastern Europe which contain remains of the water vole Microtus (Stenocranius) gregalis, Microtus ex gr. arvalis Arvicola. and Microtus oeconomus (Markova, 1982). All the faunas ARTICLE IN PRESS A.K. Markova / Quaternary International 149 (2006) 67–79 75 of this evolutionary level were correlated with the late Thus, the late Middle Pleistocene small mammal phase of the Tiraspolian mammalian age. Unfortunately materials and also the geological data show a very high the taphonomy of the localities in mole burrows excludes complexity of the sequence between Oka Glacial deposits the possibility of finding there the remains of animals and Dnieper Glacial strata. A minimum of three fossil dwelling near water, such as Mimomys savini and Arvicola. soils, deposited during warm periods, separated by loess Thus, so far we have no evidence that the transformation horizons, were established for this interval. Most of the of the progressive Mimomys savini to Arvicola cantianus strata characterized by the mammalian remains have a took place on the Russian Plain during the Muchkap different evolutionary level (Markova, 1996). Interglacial. Possibly, this evolutionary change occurred at the very end of the Muchkap Interglacial or at the 4. Discussion and correlation beginning of the Oka glaciation or even later, though before the Likhvin Interglacial. Abundant Middle Pleistocene mammal materials were recovered in Western Europe (Chaline, 1972; van Kolfschoten and Roebroeks, 1985; Heinrich, 1990; van 3. Late Middle Pleistocene small mammal faunas younger Kolfschoten, 1990; Schreve, 2001). These data demonstrate than the Likhvin Interglacial that every climatic cycle of the late Middle Pleistocene features different suites of mammalian faunas. In spite of Some localities of small mammal remains were found in disagreement as to the climatic ranks of these cycles, most layers overlying the Likhvin deposits. The fauna from the investigators distinguished at least three well-pronounced locality of Topka in the Don drainage basin likely belongs warm phases during the late Middle Pleistocene. The to a cold interval, possibly corresponding to the Boriso- earliest of these warm phases is correlated with the glebsk loess horizon (Krasnenkov and Kazantseva, 1993). Holsteinian ( ¼ Hoxnian) Interglacial. The localities re- Arvicola chosaricus bones have been found here. ‘‘Terrico- semble most closely Neede, Ka¨rlich H, Petersbuch, la’’—like voles have not been found at this site. Possibly, Bilzingsleben 2, Swanscombe, and Ornac 3 of Western the locality of Matveevka can also be correlated with this Europe (Chaline, 1972; Heinrich, 1990; von Koenigswald cool phase (Rekovets, 1994). and van Kolfschoten, 1996; van Kolfschoten and Turner, Three localities, referred to the Kamenka paleosol mole 1996), which are correlated with the Holsteinian Inter- burrows, have been found in Eastern Europe (Priluki, in glacial on the basis of pollen data. All these sites include the Dnieper drainage basin; Rasskazovo, on the Oka— remains of Arvicola cantianus (A. terrestris cantianus, after Don plain; and Tiraspol, Kamenka paleosol horizon, in the Kolfschoten), Sorex cf. araneus, Microtus agrestis, and Dniester drainage basin). The Plavni fauna of the same M. arvalis. Sorex (Drepanosorex) savini, Pliomys episco- evolutionary level was found in the third terrace alluvium palis, Microtus (Stenocranius) gregaloides, and M. (Terri- in the Danube drainage basin; Uzunlar locality is referred cola) arvalidens are not characteristic to these faunas. This to liman-sea deposits of the late Middle Pleistocene indicates the similarity with the Likhvin faunas of Eastern Uzunlar transgression of the Black Sea (Chepalyga et al., Europe. 1986; Markova, 1992; Mikhailesku and Markova, 1992). A summary of the complex geological, palynological and All these faunas are comparable with materials recovered mammalian data and a correlation with Oxygen Isotope from the Cherny Yar stratotype section (Volga drainage Stages (OIS) was carried out by Turner (1998). All these basin). The Chernyi Yar small mammal assemblage was data show that the Holsteinian ( ¼ Hoxnian) Interglacial is described by Alexandrova (1976) and assigned to the referred to OIS 11 (Table 3). These studies and correlations Khozarian mammalian age, determined by Gromov (1948) demonstrate also the existence of three warm (interglacial) and characterized by Mammuthus trogontherii chosaricus, phases in the different parts of continental Western Europe Camelus knoblochi, Megaloceros euryceros germaniae, during the late Middle Pleistocene (Table 3). Three Late Bison priscus longicornis, Equus caballus chosaricus. Arvi- Middle Pleistocene warm intervals have been revealed in cola chosaricus, Lagurus lagurus pleistocaenicus, and Great Britain, and they were characterized by different Eolagurus volgensis were described from the Chernyi Yar mammal assemblage zones (MAZ): Hoxnian (OIS 11), site (Alexandrova, 1976). Arvicola chosaricus is the Purfleet (OIS 9), and Ponds Farm and Sandy Lane (OIS 7) indicative species for Chosarian small mammal faunas. (Schreve, 2001). Mammal faunas of the Hoxnian Inter- Arvicola remains from the localities, correlated with the glacial (Swanscombe) include Arvicola cantianus remains Kamenka Interglacial, show a little negative enamel ratio with an SDQ140 (n ¼ 4). Water voles from Bilzingsleben on the anterior and posterior sides of the m/1 angles 2 and Petersbuch have an SDQ index between 130–120 (Fig. 2). Among the steppe lemmings of the genus Lagurus, (Fig. 2)(von Koenigswald and Heinrich, 1999). Mimomys the ‘‘lagurus’’ morphotype was dominant at this time, and is absent in these faunas. the ‘‘transiens’’ morphotype is of secondary importance There are a number of West European localities where (Markova, 1996). Unfortunately, small mammal faunas the archaic Arvicola have been found in the deposits of the associated with the Romny fossil soil have not been Cromer Interglacials III and IV. One of the most primitive recovered yet. Arvicola with SDQ values ranging from 155–140–123 has ARTICLE IN PRESS 76 A.K. Markova / Quaternary International 149 (2006) 67–79 Stadial Kursk warming Interstadial Dnieper Stadial Breslav et al., ) Velichko et al., 1992 ; Eastern Europe 1992 Mikulino Interglacial glacial epoch Romny warming Orchik cold interval Borisoglebsk cold interval Likhvin Interglacial Interglacial Interglacial Cool interval with two interstadials Cool interval with two interstadials Praclaux (Holstenian) Interglacial persdorf Interglacial) Landos Interglacial Kamenka Interglacial ¨ mnitz Interglacial Le Bouchet ¨ Fuhne cold ‘‘Stage’’ with an interstadial Saale Till Moscow (Dnieper) (Paludina beds) Elster Till Oka Glaciation ningen NE Germany Massif Central Central Russian Plain (after ningen Interglacial Do ¨ ¨ interstadial Cold interval with interstadials interstadials Elsterian late glacial deposits and Elster till ) Turner, 1998 Saale till Cold interval with two (marine) and Elster tills The Netherlands NW Germany Scho Cold interval with permafrost permafrost Oxygen Isotope Stages Table 3 Correlation of Late Middle Pleistocene stratigraphical divisions inWest Western European and stratigraphical Eastern divisions Europe (after 5e6 Eemian Interglacial Drenthe till Eemian Interglacial7 Eemian Interglacial Warthe till Bantega Interstadial Eemian Interglacial Wacken Interstadial Ribains (Eemian) Saale till Scho Warthe till Costaros glacial Moscow 8910 Cold interval11 Hoogeveen Interstadial Cold Interval with Holsteinian Interglacial Mehlbeck cold ‘‘stage’’ Holsteinian Interglacial Cold interval with Holsteinian an Interglacial Holsteinian Interglacial Reinsdorf Interglacial (Ro 12 ‘‘Potklei’’ (Elsterian) Elsterian Lauenberg Clay ARTICLE IN PRESS A.K. Markova / Quaternary International 149 (2006) 67–79 77 been found in Miesenheim I (von Koenigswald and van sufficient, this form could be described as Arvicola as well Kolfschoten, 1996; van Kolfschoten and Turner, 1996; von as Mimomys. Existing data show that the earliest Arvicola Koenigswald and Heinrich, 1999). This locality contains in Eastern Europe were found in Likhvin Interglacial Sorex (Drepanosorex) savini, Pliomys episcopalis, Microtus deposits (OIS 11), but it is quite possible that they could (Stenocranius) gregaloides, M. (S.) gregalis, M. (Terrico- appear during OIS 12 (the Oka Glaciation). la) arvalidens, M. arvalis, M. agrestis, M. oeconomus and Likhvin ( ¼ Holsteinian) Interglacial faunas of Eastern others. Such a composition differs from Holsteinian faunas Europe are characterized by a definite species composition by persistence of Sorex (Drepanosorex) savini, Pliomys including Arvicola cantianus, Lagurus transiens, Eolagurus episcopalis and ‘‘Terricola’’ like voles. luteus volgensis, Microtus ex gr. arvalis, Microtus agrestis, Ka¨rlich Gb also correlates with the level of an early stage Microtus (Stenocranius) gregalis, and Microtus oeconomus. of Arvicola cantianus faunas and includes primitive water Rhizidont voles Mimomys, Pliomys, Borsodia, and also vole remains. Sorex (Drepanosorex) savini, Pliomys Sorex (Drepanosorex) savini have not been found in the episcopalis and Microtus (Terricola) arvalidens, Microtus Likhvin localities. Microtus (Terricola) arvalidens and (Stenocranius) gregaloides, and Microtus gregalis, M. (Stenocranius) gregaloides are not typical for this M. arvalis/agrestis have been identified from this layer stratigraphical stage. (van Kolfschoten and Turner, 1996). In the opinion of van The stratigraphical position of Likvin localities in Kolfschoten and Turner (1996) and based on complex different parts of the Russian Plain correlates with the geological and paleontological data, this locality is Inzhavino paleosol, the fourth terrace of the main rivers of probably older than Miesenheim I and could be referred Eastern Europe, and the Early Euxinian transgression of to Cromer III. However, the fauna composition of these the Black Sea. These deposits overlie the Oka Glacial strata two localities is very similar to one another. and are covered by Borisoglebsk loess and also Kamenka The Mosbach 2 fauna is correlated with OIS 13 and and Romny paleosols and an intermediate loess horizon. includes archaic Arvicola, recently re-described as This entire late Middle Pleistocene series is covered by A. mosbachensis (partly with rooted teeth and SDQ Dnieper Glacial deposits (moraine, fluvioglacial suite, and coefficients ranging from 159–133–117), Sorex (Drepano- Dnieper loess horizon in the periglacial regions). sorex) savini, Pliomys episcopalis, Microtus nivaloides, Mean values of the Likhvin Arvicola SDQ index vary M. nivalis and even M. hintoni (Maul et al., 2000). The between 135–128–110. The morphotypes of lagurids show persistence of Sorex (Drepanosorex) savini and Pliomys prevalence of ‘‘transiens’’ ones, but ‘‘lagurus’’ morpho- episcopalis suggests an earlier age of this fauna than types also existed in the steppe lemming lineage (Markova, Holsteinian. In my opinion, Microtus hintoni remains must 1996). have been re-deposited from earlier deposits, because this The combination of data shows that the stratigraphical species is characteristic of older stratigraphical levels. position of Likhvin faunas in the late Middle Pleistocene Thus, a comparison between West European and East succession is similar to that of Holsteinian ( ¼ Hoxnian) European faunas with Arvicola remains shows that the ones, and is definitely correlated with OIS-11 (Turner, earliest stage of Arvicola cantianus faunas is absent in 1998; Schreve, 2001). Eastern Europe. There could be two explanations for this fact. Possibly we have not yet found the fauna of this stage. Acknowledgments We do not have sufficient data correlated with the Oka Glaciation. Probably, at this time the earliest Arvicola I should like to thank very much the organizers of the existed on the Russian Plain. On the other hand, there are SEQS conference in Kiev, Ukraine (2001), who initiated numerous data relating to the Muchkap (Roslavl) Inter- this paper. I am very grateful to Dr. Lutz Maul for glacial. Even in the deposits of the upper warm phase of the critically reading and improving this manuscript. Muchkap (Roslavl) Interglacial—Konakhovka optimum (which strictly preceded the Oka Glaciation) Arvicola References remains have not been found. Only very advanced Mimomys, which in many cases have no roots, have been Agadjanian, A.K., 1977. Quarta¨re Kleinsaüger aus der Russischen Ebene. described from this stratum. Agadjanian identified also Quarta¨r 27/28, 111–145. Sorex sp., Desmana cf. moschata, Spermophilus sp., Agadjanian, A.K., Erbaeva, M.A., 1983. 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