Quaternary International xxx (2014) 1e10

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The molluscs record: A tool for reconstruction of the Late Pleistocene (MIS 3) palaeoenvironment of the Bol'shoj Naryn site area (Fore-Baikal region, Eastern , )

* Guzel Danukalova a, b, , Eugeniya Osipova a, Fedora Khenzykhenova c, Takao Sato d a Institute of Geology, Ufa Scientific Center RAS, Ufa, Russia b Kazan Federal University, Kazan, Russia c Geological Institute, Siberian Branch RAS, Ulan-Ude, Russia d Department of Archaeology and Ethnology, Faculty of Letters, Keio University, Tokyo, Japan article info abstract

Article history: A representative mollusc fauna attributed to the late phase of the Karginian Interstadial (MIS 3) has been Available online xxx found in the Bol'shoj Naryn Palaeolithic site (Fore-Baikal region). The general organization of the strata at the Bol'shoj Naryn site has been established through excavations realized during the previous field Keywords: seasons. It shows a modern soil made of sandy loess deposits 1 m thick, dated from the Sartan glacial Terrestrial molluscs stage, and underlined by a high viscosity paleosol layers which is up to 1 m thick developed during the Late Pleistocene Karginian Interstadial. The “cultural layer” has been correlated with the upper Karginian soil contains Karginian Interstadial numerous stone tools and fossils. This paper focus on the mollusc assemblage attributed to the Fore-Baikal region upper Karginian sediment. The mollusc assemblage (2460 determined specimens) consists of six and five genera of terrestrial molluscs. oblonga, Pupilla muscorum and tenuilabris are the best represented species. The molluscs suggest the existence of landscapes corresponding with humid meadows and forests located in the relief depressions or along banks of the river. Molluscs of the Bol'shoj Naryn site have been compared with equivalent mollusc complexes from Siberian and Southern Fore-Uralian lo- calities. They display a distinctive poverty in the species composition and show similarity with the complex of the Gornovo locality (Southern Fore-Urals region). Comparison of the mollusc complex from the Bol'shoj Naryn locality with molluscs from the last glacial period also showed similarity with the complex of the Gornovo locality (Kudashevo period in the Southern Urals e last phase of the Late Valdai equivalent MIS 2). The climatic conditions which prevailed in the surroundings of the Bol'shoj Naryn locality during the late phase of the Karginian interglacial period (32e25 ka BP) of the Fore-Baikal region were cooler than those existing at the same time in Europe and Southern Urals. Correlation between the data obtained after malacological investigations and other paleontological data obtained from paly- nology and vertebrate records suggest a moderate cold and humid climate in the Fore-Baikal region during the second part of MIS 3, with predominance of open landscapes of steppes and relatively limited taiga forest, tundra, and wetlands. © 2014 Elsevier Ltd and INQUA. All rights reserved.

1. Introduction and Quaternary (Florensov, 1960). Cenozoic deposits filled this wide depression (for example the Baikalian rift zone) where they The Fore-Baikal region is located mainly inside the Baikalian are several kilometers thick. Other smaller depressions (for structural zone, which was tectonically active during the Neogene example the Fore-Baikalian foredeep) display only thicknesses of several hundred meters. The sedimentation started in this region in the Oligocene. Quaternary deposits close to the mountains are * Corresponding author. Institute of Geology, Ufa Scientific Center RAS, Ufa, represented by conglomerate, gravel, and sand. Large depressions Russia. are occupied by lakes where thin lacustrine sediments were E-mail addresses: [email protected], [email protected] (G. Danukalova), deposited. The Fore-Baikalian and the Fore-Sayanian foredeeps are [email protected] (E. Osipova), [email protected] (F. Khenzykhenova), sato@flet.keio. filled by alluvial deposits. Glacial, deluvial, and other types of ac.jp (T. Sato). http://dx.doi.org/10.1016/j.quaint.2014.08.034 1040-6182/© 2014 Elsevier Ltd and INQUA. All rights reserved.

Please cite this article in press as: Danukalova, G., et al., The molluscs record: A tool for reconstruction of the Late Pleistocene (MIS 3) palaeoenvironment of the Bol'shoj Naryn site area (Fore-Baikal region, Eastern Siberia, Russia), Quaternary International (2014), http:// dx.doi.org/10.1016/j.quaint.2014.08.034 2 G. Danukalova et al. / Quaternary International xxx (2014) 1e10

Pleistocene deposits are also known in the Fore-Baikalian area and high viscosity paleosol layers up to 1 m in thickness which (Zeitlin et al., 1984). developed during the Karginian Interstadial (Fig. 2). The “cultural Bol'shoj Naryn site is a Palaeolithic settlement (53 33016.9900N, layer” correlated to the upper Karginian soil contains numerous 1033407.8800E) which is located 185 km north of Irkutsk city stone tools and animal fossils (Sato et al., 2014). The general (Fig. 1). This settlement was situated on the left bank of the Osa thickness of the Karginian deposits is 0.05 m. The radiocarbon age River, 10 km east of the main bed of the River. The Osa obtained on charcoal samples taken in the upper layer was River is a right tributary of the Angara River, now flooded by the 25,000e26,500 BP. Samples from the lower part are dated at Bratsk Reservoir. 27,000e31,500 BP (Sato et al., 2014). This locality is cut by a small gully and can be divided into This is the first mollusc assemblage of the Karginian paleosol western (Bosl'shoj Naryn I) and eastern (Bol'shoj Naryn II) parts. noted in the Fore-Baikalian region. The main aim of this paper is to The western and southwestern surroundings of the locality are focus on the characteristics of the mollusc assemblage attributed to 500 m above sea level and form an amphitheater, which is open to the upper Karginian sediment and to use it as a tool for the the northeast towards the steep Bilchir slope Reservoir. A hill 450 m reconstruction of the Late Pleistocene (MIS 3) palaeoenvironment high protected the Bol'shoj Naryn settlement to the south. The of the surroundings of the Bol'shoj Naryn site. amphitheater and the hill correspond with remnants of an ancient large karst depression. Loose deposits up to 30 m thick cover the 2. Materials and methods slopes and are overlapped by Karginian soil, Sartan glacial deposits and Holocene soil. These deposits were tectonically uplifted in the A total of 199 samples containing mollusc shells were examined. east and downlifted in the western parts of the settlement during All the samples were taken during 2010 and 2011 from the “cultural the late Karginian. Suffusional and cryogenic processes increased layer” of the archaeological excavation 1 of the Bol'shoj Naryn the deformation of the Karginian paleosol (Yoshida et al., 2007; settlement, which was attributed to the upper Karginian soil (loam) Sato et al., 2014). and dated between 32,000 and 25,000 BP (Sato et al., 2008). The Field work (excavations and water-sieving of the soil samples) studied samples were taken from squares (1 1 m) made in the involved geologists, archaeologists and palaeontologists of Irkutsk, excavation at a depth of 0.6 m with an interval of 5 cm. The datum Tokyo, Keio, and Hokkaido universities and the Geological Institute of sample collection is considered to represent the upper stratum of (Siberian branch of the Russian academy of sciences) in 2004e2005 the “cultural layer”. It is located at 406e407 m a.s.l. The rinsed and 2009e2011. The general structure of the Bol'shoj Naryn site material from all samples was equal to 0.05 1 1 m, 0.05 m3.As was established after various excavations made during previous the density of loam was 1500e1600 kg/m3, each sample corre- field seasons. It shows a modern soil, underlain succesively by sponds to 75e80 kg. Conversion to 1 L indicates that the samples sandy loess deposits 1 m thick (dated from the Sartan glacial stage), were rich in mollusc shells.

Fig. 1. Location of the Bol'shoj Naryn site (1) in the surroundings of the Baikal area.

Please cite this article in press as: Danukalova, G., et al., The molluscs record: A tool for reconstruction of the Late Pleistocene (MIS 3) palaeoenvironment of the Bol'shoj Naryn site area (Fore-Baikal region, Eastern Siberia, Russia), Quaternary International (2014), http:// dx.doi.org/10.1016/j.quaint.2014.08.034 G. Danukalova et al. / Quaternary International xxx (2014) 1e10 3

Fig. 2. East section of the Bol'shoj Naryn site I and radiocarbon dates of charcoals from paleosol layer.

Samples were rinsed in field conditions using 1 mm sieves. The In the laboratory, the samples were sorted out according to the rinsed remainder was dried, labeled, packed into bags (1e4 bags/ squares where they came from and depths (Tables 1 and 2). Each samples from each level) and sent for examination to the Institute sample was calibrated according to the species structure with the of Geology of the Ufimian Scientific Centre of the Russian Academy use of brushes and MBS 10 binocular microscope with different of Sciences (Ufa). All the samples were registered at the Laboratory amplification ( 4 e 10). Mollusc species were identified ac- of the Cenozoic Geology and labeled with an index number. Visual cording to Kerney and Cameron (1999) and Likharev and inspection of the samples revealed unbroken mollusc shells, large Rammelmeier (1952). For the malacological taxonomical nomen- fragments of shells, and fine detritus. clature, we used the publication of Falkner et al. (2002).

Table 1 Molluscs species composition from the Bol'shoj Naryn site upper Karginian soil (MIS 3) in the squares 54 and 505.

Taxons Interval of depth of upper Karginian soil (MIS 3), m Total

00e0.05 0.05e0.10 0.10e0.15 0.15e0.20 0.20e0.25 0.25e0.30 0.30e0.35 0.35e0.40 0.40e0.45 0.45e0.50 0.50e0.55 0.55e0.60

Square 54 1. 27913633321514122 (Draparnaud) 2. Vallonia tenuilabris 13 7 26431 2 83 (Braun) 3. Pupilla muscorum 3 13 17 7 44 106 70 1 12 274 (Linnaeus) 4. Vertigo alpestris Alder 1 1 Total 6 23 26 8 88 3 165 134 7 1 18 479

Square 505 1. Succinella oblonga 2 1 18 3 1 25 (Draparnaud) 2. Vallonia tenuilabris 13 3 7 (Braun) 3. Pupilla muscorum 11 2 22 35 (Linnaeus) 4. Vertigo alpestris Alder 11 Total 13 4 43 4 4 68

Table 2 Molluscs species composition from the Bol'shoj Naryn site upper Karginian soil (MIS 3).

Taxons Interval of depth of upper Karginian soil (MIS 3), m Total

00e0.05 0.05e0.10 0.10e0.15 0.15e0.20 0.20e0.25 0.25e0.30 0.30e0.35 0.35e0.40 0.40e0.45 0.45e0.50 0.50e0.55 0.55e0.60

1. Succinella oblonga 134 90 96 94 34 156 53 82 103 54 7 3 4 910 (Draparnaud) 2. þ (Succinella sp.) 3. Vallonia tenuilabris 27 15 53 11 1 12 7 12 32 53 3 5 2 233 (Braun) 4. Pupilla muscorum 230 256 196 95 42 138 32 76 150 91 1 12 1319 (Linnaeus) 5. Pupilla sp. (cf. Pupilla þ muscorum) 6. Vertigo alpestris Alder 1 1 1 3 7. Vertigo cf. modesta 1 1 (Say) 8. Limacidae 1 2 3 Total 392 361 346 200 77 308 94 170 285 199 11 8 18 2469

þ small fragments of the whorls (middle part of the shell) with size 1mm.

Please cite this article in press as: Danukalova, G., et al., The molluscs record: A tool for reconstruction of the Late Pleistocene (MIS 3) palaeoenvironment of the Bol'shoj Naryn site area (Fore-Baikal region, Eastern Siberia, Russia), Quaternary International (2014), http:// dx.doi.org/10.1016/j.quaint.2014.08.034 4 G. Danukalova et al. / Quaternary International xxx (2014) 1e10

In total, 2469 mollusc shells were processed. The amount of shells which were counted was based on the following method. All samples from one level were summarized. For gastropods, the total number of unbroken fully-grown and juvenile shells was counted (Lozek, 1986). Afterwards, the apices and apertures, which were considered as equivalent to one shell when taken together, were also counted. The resulting amount was added to unbroken spec- imens. The species percentage in each sample was not counted, because the method needs more than 200 specimens of shells in each sample to be valid (White et al., 2008). The shells were pho- tographed at the laboratory of Archeosciences of the University of Rennes 1 (Rennes, France) on a stereomicroscope Leica M205С with a camera Leica DFC295 (Figs 3e7).

3. Results

3.1. distribution according to depth

The squares, containing representative material throughout the e Fig. 4. Succinella oblonga (Drap.) found in Bol'shoj Naryn site, Fore-Baikal region, Eastern studied depth, were analyzed (sq. 54, depth 0 0.6 m; sq. 60, depth Siberia, Russia (Karginian horizon, Upper Pleistocene): 1 e IG N 224/3604/3, ex. 1, sq. 0e0.45 m; sq. 66, depth 0e0.5 m; sq. 72, depth 0e0.1 m; sq. 77, 54, d. 0.05e0.10 m; 2 e IG N 224/3619/1, ex. 1, sq. 60, d. 0.15e0.2 m; 3 e IG N 224/ e e e depth 0 0.3 m; sq. 78, depth 0 0.15 m; sq. 504, depth 0 0.55 m; 3631/1, ex. 1, sq. 66, d. 0.25e0.3 m; 4 e IG N 224/3609/3, ex. 1, sq. 54, d. 0.35e0.4 m; sq. 505, depth 15e0.55 m; sq. 604, depth 0e0.45 m; sq. 605, depth 5 e IG N 224/3611/1, ex. 1, sq. 54, d. 0.45e0.5 m; 6 e IG N 224/3612/1, ex. 1, sq. 54, d. 15e0.45 m; sq. 704, depth 0e40 cm; sq. 705, depth 5e0.4 m; sq. 0.5e0.55 m; a e apertural view; b e abapertural view (view from the opposite aper- e e e e 804, depth 0e0.3 m; sq. 805, depth 0e0.25 m; sq. 904, depth ture's side); c lateral view (top right); ex. excavation, sq. square, d. depth. 0e0.15 m; sq. 905, depth 0e0.15 m). The analysis of the mollusc distribution as a function of the depth (for the most representative squares) showed that the mollusc shells are irregularly located in the soil (eg. squares 54 and 505, Table 1). The largest amount of shells is concentrated at a depth of 0.5e0.45 m. The upper part of the investigated soil (0e0.05 m) and the lower interval (0.5e0.6 m) contain fewer mollusc shells. It is likely that the part of the soil located in 0.35e0.45 m interval was formed in the most favourable climatic conditions. The end of the soil accumulation interval could have been influenced by a changing environment, lowering of the air temperature as a result of the coldest phase of the last cold stage (glacial) (25e10 ka).

Fig. 3. Pupilla muscorum (L.) found in Bol'shoj Naryn site, Fore-Baikal region, Eastern Siberia, Russia (Karginian horizon, Upper Pleistocene): 1 e IG N 224/3604/1, ex. 1, sq. 54, d. 0.05e0.1 m; 2 e IG N 224/3606/1, ex. 1, sq. 54, d. 0.15e0.2 m; 3 e IG N 224/ Fig. 5. Vallonia tenuilabris (Braun) found in Bol'shoj Naryn site, Fore-Baikal region, 3671/1, ex. 1, sq. 505, d. 0.25e0.3 m; 4 e IG N 224/3609/1, ex. 1, sq. 54, d. 0.35e0.4 m; Eastern Siberia, Russia (Karginian horizon, Upper Pleistocene): 1 e IG N 224/3703/2, 5 e IG N 224/3610/1, ex. 1, sq. 54, d. 0.4e0.45 m; 6 e IG N 224/3613/1, ex. 1, sq. 54, d. ex. 1, sq. 804, d. 0e0.05 m; 2 e IG N 224/3628/2, ex. 1, sq. 66, d. 0.1e0.15 m; 3 e IG N 0.55e0.6 m; a e apertural view; b e abapertural view (view from the opposite aper- 224/3671/2, ex. 1, sq. 505, d. 0.25e0.3 m; 4 e IG N 224/3609/2, ex. 1, sq. 54, d. ture's side); c e lateral view (top right); IG N 224/3604/1 e specimen number in the 0.35e0.4 m; 5 e IG N 224/3668/1, ex. 1, sq. 504, d. 0.45e0.5 m; 6 e IG N 224/3613/2, mollusc base of the Institute of Geology USC RAS (Ufa); ex. e excavation, sq. e square, ex. 1, sq. 54, d. 0.55e0.6 m; a e apertural view; c e lateral view (top right); d e um- d. e depth. bilical view; e apical view; ex. e excavation, sq. e square, d. e depth.

Please cite this article in press as: Danukalova, G., et al., The molluscs record: A tool for reconstruction of the Late Pleistocene (MIS 3) palaeoenvironment of the Bol'shoj Naryn site area (Fore-Baikal region, Eastern Siberia, Russia), Quaternary International (2014), http:// dx.doi.org/10.1016/j.quaint.2014.08.034 G. Danukalova et al. / Quaternary International xxx (2014) 1e10 5

Fig. 6. Vertigo found in Bol'shoj Naryn site, Fore-Baikal region, Eastern Siberia, Russia (Karginian horizon, Upper Pleistocene): Vertigo cf. modesta (Say): 1 e IG N 224/3679/ 1, ex. 1, sq. 604, d. 0.2e0.25 m; Vertigo alpestris Alder: 2 e IG N 224/3676/1, ex. 1, sq. 604, d. 0.05e0.1 m; 3 e IG N 224/3607/4, ex. 1, sq. 54, d. 0.2e0.25 m; 4 e IG N 224/ 3673/1, ex. 1, sq. 505, d. 0.45e0.5 m; a e apertural view; b e abapertural view (view from the opposite aperture's side); c e lateral view (top right); ex. e excavation, sq. e square, d. e depth.

The generalized mollusc distribution throughout the depth for Fig. 7. Limacidae found in Bol'shoj Naryn site, Fore-Baikal region, Eastern Siberia, all squares (Table 2) is less informative, the amount of shells found Russia (Karginian horizon, Upper Pleistocene): 1 e IG N 224/3615/1, ex. 1, sq. 60, d. e e e in the upper and middle parts of the soil showing little difference. 0m;2 IG N 224/3621/1, ex. 1, sq. 60, d. 0.25 0.3 m; 3 IG N 224/3621/2, ex. 1, sq. 60, d. 0.25e0.3 m; a e apical view; b e basal view (view from the opposite apical side); However, the lower part of the section has fewer mollusc shells. ex. e excavation, sq. e square, d. e depth.

3.2. Systematic composition analysis swamps) can be affected by the reduction of the tooth in the All the examined shells belong to one class of . Our aperture. A. Shileyko also noted that the reduction of the teeth in research indicated that the samples include shells of the terrestrial the aperture of P. muscorum is usual (Shileyko, 1984,p.185e186; mollusca (6 species, 5 genera), and also numerous fine shell Sysoev and Shileyko, 2009, Fig. 18 I). There is a possibility that the detritus associated with small mammal bones. The mollusc species specimen from Bol'shoj Naryn is an ecophaenotype of P. muscorum composition is shown in Table 2. (Ecophaenotypus, 1965). The dominant species is Pupilla muscorum (Linnaeus, 1758) Succinella (Fig. 4). The genus is represented by S. oblonga. Shells (53.3%), and the second prevailing species is Succinella oblonga in most cases have medium preservation. The shell is conispiral (Draparnaud, 1801) (37%). Vallonia tenuilabris (Braun, 1843) repre- with a relatively rapid whorl increase; marked with thin and even sents 9.5% of the total amount of all the examined shell specimens. striae. The shell has 3.5 convex round whorls, which are separated Species of Vertigo and slugs occur rarely. with a distinctive suture. The last whorl is almost 2 times larger Pupilla (Fig. 3). Shells are represented by P. muscorum which than the first one. The aperture is oval, arrow-headed at the top; lip has an oval-cylindrical form shell which is finely, evenly striated. of the aperture is thin; inner edge of the aperture borders the shell, The shell has 5¾ e 6 convex well defined whorls, which are covering the umbo. In comparison with the other representatives of separated with a distinctive but not very deep suture. The sizes the Succineidae family (Succinea and Oxyloma) the shells of Succi- (average) are: shell height e 3.2 mm, width e 1.8 mm. The nella genus are different in the shell form (more convex round aperture is semicircular with slightly turned away well developed whorls) (Likharev and Rammelmeier, 1952,p.121e123, Figs 28e30, lip; the palatal edge has a distinctive “constriction”; the aperture 32; Danilovskyi, 1955,p.86e87, Table 5, Figs 83e90, 93, 96e99, 101, lacks teeth. Umbo is small and narrow. P. muscorum of Bol'shoj 103e109; Lozek, 1964, p. 229e232, Table 12, Figs 1e4, 7e9; Naryn site is similar to the type specimen of P. muscorum known in Shileyko and Likharev, 1986,p.200e202, Fig. 1; p. 204e207, Fig. 5; the Linnaeus collection because of the absence of teeth in the p. 223e225, Fig. 23; Sysoev and Shileyko, 2009, Figs 5, 6). aperture (Linnaeus, 1758, p. 767; Hanley, 1855, p. 352, pl. 4, Fig. 6). Vallonia (Fig. 5). The genus is represented by the species V. The morphological peculiarities of the examined specimen are tenuilabris. The shell is spiral, thin evenly striated, pateliate with most likely connected with the ecological conditions where the slow whorl increase, rounded from umbo (or apex) side. The shell mollusca were living. According to R. Preece's data (White et al., has 3.5 convex whorls, separated by a deep suture. The last whorl is 2008), P. muscorum living in wet environments (flood meadows, lowered to aperture. The average sizes of the shells are: height e

Please cite this article in press as: Danukalova, G., et al., The molluscs record: A tool for reconstruction of the Late Pleistocene (MIS 3) palaeoenvironment of the Bol'shoj Naryn site area (Fore-Baikal region, Eastern Siberia, Russia), Quaternary International (2014), http:// dx.doi.org/10.1016/j.quaint.2014.08.034 6 G. Danukalova et al. / Quaternary International xxx (2014) 1e10

1.5 mm, width e 3.1 mm. The aperture is rounded (average diam- 3.3. Palaeoecological interpretations eter e 1.1 mm), the outer edge of the aperture is thin, slightly turned out, not thickened; the inner edge is thin, attached to the The ecological characteristics of the identified species of mol- shell surface. The umbo is open and wide. In the Quaternary de- lusca have been analyzed and particularly the relationships of the posits of various locations V. tenuilabris differs from the other to the air temperature and humidity and the characteristics species of this genus by its large shells (Likharev and Rammelmeier, of their habitat (Table 3). During the upper Karginian, the climate 1952, p. 166, Fig. 78; Danilovskyi, 1955, p. 88, Table 6, Figs. 125e127; surrounding the site was temperate and wet.

Table 3 Ecological characteristics of the mollusc species identified in Bol'shoj Naryn site according to Lozek (1964), Likharev and Rammelmeier (1952), Shileyko (1984), Sysoev and Shileyko (2009), Puissegur (1976), Sümegi and Krolopp (2002), Willis et al. (2000), Kerney and Cameron (1999), Kerney et al. (1983), Germain (1930), Adam (1960), Zilch and Jaeckel (1962), Animal base (n.d).

N Species Mollusc groups Mollusc groups Ecology (habitat) (temperature) (humidity)

1. Succinella oblonga Cold resistant and Hygrophilous Open habitat preference. (Draparnaud, 1801) eurytherms Wet environments with sparse vegetation (humid meadows, forests, bushes, swamps and stream banks). European and West-Asiatic areas. 2. Vallonia tenuilabris Cold resistant and Mesophilous Intermediate. (Braun, 1843) eurytherms Wet environments (under wet fallen leaves in mixed and leaved forests, among moss). Palaeoarctic area. 3. Pupilla muscorum Cold resistant and Mesophilous Open habitat. (Linnaeus, 1758) eurytherms Dry or slightly wet and sunny calcareous and open environments (meadows, under stones, under fallen leaves, in mosses, littoral dunes). Plains and mountains (up to 2400 m). Holarctic area. 4. Vertigo alpestris Mesophilous Mesophilous Intermediate. Alder, 1838 Wet environments (under wet fallen leaves in leaved forests, among moss, bushes, flood plain meadows). Palaeoarctic area. 5. Vertigo cf. modesta Mesophilous Mesophilous Intermediate. (Say, 1824) Wet environments (under wet fallen leaves in leaved forests, under stones, in mosses, bushes, flood plain meadows). Holarctic area. 6. Limacidae Mesophilous Subhygrophilous Intermediate. Warm and wet environments with developed vegetation cover or open areas (humid meadows, forests). Holarctic area.

Lozek, 1964; с. 223, Table 10, Fig. 1 aec; Shileyko, 1984, p. 169; The examined species of molluscs are characteristic of biotopes Sysoev and Shileyko, 2009, Fig. 16 E). showing an increasing humidity. They probably corresponded with Vertigo (Fig. 6). Two species were examined: Vertigo alpestris flooded meadows or with flooded plains near a water reservoir. The Alder, 1838 and Vertigo cf. modesta (Say, 1824). The species are S. oblonga species is hydrophilic and could live directly near the represented by conispiral shells with even whorls increase. water on the plants. Molluscs from the genus Vertigo, representa- V. alpestris has an oval e subcylindrical shell, which is marked by tives of the Limacidae family and V. tenuilabris lived in leaf litter, thin and even striae, 4¾e5 convex whorls. The average size is: moss, wood rot, which indicates the presence of a forest. width e 2.1 mm, height e 1.1 mm. The aperture is weakly semi- circular with slightly turned away well developed lip, the palatal 4. Discussion edge either may have a depression; there are 4 teeth in the aper- ture: one parietal plate, one collumellarl plate, two palatal (upper Saks (1948) described the Karginian horizon for the first time. and lower) pleats (Figs 2e4, 6). V. cf. modesta has an oval shell, The deposits of the Karginian horizon are widespread in Siberia which is marked with thin and even striae, 5 convex whorls. The including the Eastern Siberia area (the Siberian Platform) and are sizes of V. cf. modesta are width e 2.1 mm, height e 1.2 mm. The known in the glacial and periglacial areas (Laukhin et al., 1971; aperture is slightly hemispheric with turned away well developed Lazukov, 1981; Laukhin, 1982, 2001; Zeitlin et al., 1984; Medvedev lip, the palatal edge may have a depression; there are 3 teeth in the et al., 1990; Volkova, 2001; Volkova et al., 2003, 2005). We will aperture: one parietal plate, one columellar plate, and one palatal concentrate first on the area showing strong continental influences (lower) fold (Figs 6 and 1). A.A. Shileyko (1984,p.211;Sysoev and similar to the analyzed area and will subsequently discuss the Shileyko, 2009, Fig. 20 G, H) considers V. alpestris to be a subspecies southern part of the Siberia where deposits are attributed to the of V. modesta (¼V. modesta alpestris Alder, 1838). Karginian horizon. They are represented by alluvial (stream and Limacidae (Fig. 7). There are three reduced shells in our collec- floodplain) facies and by paleosols (Chistyakov et al., 2000)are tion. They display the shape of an oval plate with included off- mainly developed in the river valleys. center apex. The length of the plate is 3.1 mm, width 1.8 mm According to the radiocarbon data, the Karginian Interstadial (average). After the specified degree of probability, these plates lasted around 50e24 ka. Several cooling and warming episodes are could belong to genus Limax (Steklov, 1966, p. 219, tab. 9, known during this period, which is generally considered as Inter- Fig 175e177). glacial (Zeitlin et al., 1984; Volkova et al., 2005).

Please cite this article in press as: Danukalova, G., et al., The molluscs record: A tool for reconstruction of the Late Pleistocene (MIS 3) palaeoenvironment of the Bol'shoj Naryn site area (Fore-Baikal region, Eastern Siberia, Russia), Quaternary International (2014), http:// dx.doi.org/10.1016/j.quaint.2014.08.034 G. Danukalova et al. / Quaternary International xxx (2014) 1e10 7

The transitional zone between the Atlantic and the Pacific cli- Krasnoyarsk) accumulated in climatic conditions close to modern matic influence is located in Western Siberia. The Karginian episode or even cooler (Lazukov, 1981) when the taiga landscapes were is characterized by a “European” (Interstadial) character in the widespread. Atlantic zone and by an Interglacial character in the Pacificarea (Laukhin and Firsov, 2011). 4.3. Lower Tunguska basin

4.1. Western Siberia In this area the Karginian horizon is represented by a paleosol which covers the alluvial deposits of the III terrace and by the al- The pedocomplex which formed during the Karginian Inter- luvium and peat (28,800 ± 120 BP) of the lower part of the II stadial in the southern part of Western Siberia consists of two or terrace. This paleosol is underlain by alluvium of the Zyrian horizon three soils intercalated with thin loess layers. The thin fossil soil is and covered by the deposits of the Sartan horizon (Zeitlin et al., represented by a poorly developed chernozem, which suggests the 1984). In the lower stream of the Lower Tunguska River valley, existence of a forest-steppe and steppe environment (Dobretsov the Karginian deposits are subdivided in three parts by episodes of et al., 2003; Volkova et al., 2005; Zykina, 2012). The accumula- incision and cooling as shown by cryoturbation. These features tion of humus and the carbonatization of the paleosols dominated could be correlated to the similar structure of the Karginian Inter- during this period. Two soils of the Karginian Interstadial (MIS 3) stadial complex of the Yenisei valley. differ from the Interglacial and from the modern soils because of shorter period of accumulation (horizon A e 0.25e0.3 m thick; 4.4. Tunka valley horizon BC e 0.7 m thick). According to the geochronological dating, each soil of this pedocomplex formed over 6 to 7 ky In the southwest Fore-Baikalian area, the Late Pleistocene is (Zander et al., 2003; Frechen et al., 2005). The lower Karginian soil represented by alluvial sediments (Golubeva and Ravskyi, 1962; is better developed than the upper soil, which shows that the Adamenko et al., 1975; Ufimtcev et al., 2002; Shchetnikov and Karginian Interstadial climatic optimum developed at the same Filinov, 2007; Shchetnikov et al., 2009) (Karginian and Sartan time as the lower soil. The climate which developed during the horizons; Shabartai, Elovka, Tuyana, Belyi Yar, Slavin Yar, Zaktui, early Karginian warming, was wetter and warmer. Arkhipov and Bol'shoj Zangisan and other sites). Bones of large mammals and Volkova (1994) suggest that the boreal-cold climate which exis- radiocarbon dates support the attribution of these deposits to the ted during Karginian time was characterized by the following Late Pleistocene (Shchetnikov et al., 2010). The pedocomplex temperatures: July temperature þ15 C and was 3 C lower than located at the base of the Bol'shoj Zangisan site is attributed to the the modern temperature; winter temperatures were around Karginian age (MIS 3) according to the 14C AMS dating 25С and also 3e4C lower than the modern one. These data (32,000e35,000 BP). These deposits contain bones of large and suggest that the climate was cooler than during the Interglacials. small mammals of the Palaeolithic complex (Sizov, 2014). The Today, the investigated area, which is located inside the Eurasian Zaktui site contains a horizon with bones incorporated in a sandy continent, is characterized by a severe continental climate with loam at a depth of 2.3e2.6 m. The bones were dated by 14C AMS long and cold winters and short and warm summers. The method. All the data (33,010e36,800 BP) indicate a Karginian maximum differences in the annual air temperature can exceed Interstadial age (MIS 3) (Sizov, 2014). The alluvial deposits that 80 C and daily 30 C. The difference between the summer and the constrain the Belyi Yar I and II sites accumulated during the Kar- winter day and night temperature increases from the south to the ginian and Sartanian times according to geochronological dating. north in the region. The average annual temperature is negative in The Karginian clayish sandy loam and the loam and peat of the this area (Irkutsk: 1 C). layer 9 (Belyi Yar II site) contain molluscs (determination of S.M. The comparison of the loess-paleosol sequence of the southern Popova and Ya.I. Starobogatov: S. ex gr. oblonga, Valvata (Sibir- part of Western Siberia (Dobretsov et al., 2003; Volkova et al., 2005; ovalvata) ssorensis W. Dybowsky, 1886, Lymnaea aff. viridis Quoy Zykina, 2012) with the Baikalian biogenic silica sequence shows and Gaimard, 1832, Lymnaea (Radix) auricularia torquilla West- that the loess-paleosol record better reflects the climatic changes erlund, 1876, Lymnaea (Galba) truncatula (Müller, 1774), Gyraulus that occurred during the Late Pleistocene. Two soils, which can be acronicus Ferussac, 1807, Helicorbis sujfunensis Starobogatov, 1957, correlated with MIS 3, are well represented in the loess and soil Helicorbis sp., Unio? sp., Sphaerium levinodis Westerlund, 1876, sequence of Siberia, in contrast to the Baikalian sequence which Pisidium sp. (Popova, 1981; Sizov, 2014). The sandy loam and the displays only one weak peak at the same time (Kuzmin et al., 2001). sand showing a deformed paleosol (layer 6) and mammal bones of the Tuyana site was attributed to the Karginian Interstadial ac- 4.2. Yenisei Siberia cording to the radiocarbon date (35,900 ± 750 BP) (Sizov, 2014). The deposits of the Shabartai site are represented by alluvial sands In this area, the Karginian Interglacial has been subdivided and gravels; where the interval spanning between 4.5 and 10 m into three warm and two cool episodes on the base of the could be attributed to the Karginian (Sizov, 2014) according to the palynological data (Kind, 1974; Lazukov, 1981; Zeitlin et al., TL and 14C data (50.4 ± 3 ka and 46,600 ± 900 BP). The Slavin Yar 1984). By comparison with the Late Karginian, the Early Kargi- site (Zun-Murin River) is a key-site where the floodplain facies nian warming was characterized by a better development of the contain peat, a thin paleosol layer, wood coal, mollusc and landscape-climatic conditions according to S.A. Laukhin (Zeitlin mammal remains (Shchetnikov et al., 2009; Sizov, 2014). The et al., 1984), as the deposits of the right tributaries of the lower part of these deposits has been dated at 45,810 ± 4070 BP Angara River contain shells of Unio annulatus (determination by (IGAN 3133) and the upper part at 37,790 þ 310 BP (TO-13270). Ya.I. Starobogatov in: Popova, 1981), and some plants which do The lower part of the layer (level 11 m below the terrace surface) not exist today in this region. At that time, the climate was contains freshwater species Aplexa hipnorum (Linnaeus, 1758), warmer and wetter than now. According to the palynological Limnaea (Galba) truncatula sibirica (Westerlund, 1885), and data, the climate was more or less similar to the modern one terrestrial molluscs Succinea putris (Linnaeus, 1758), Vallonia cos- during the Late Karginian warming (30,000e25,000 BP) (Laukhin tata (Müller, 1774), P. muscorum, V. modesta, Nesovitrea hammonis et al., 1971; Laukhin, 1982, 2001; Zeitlin et al., 1984). The Kar- (Strom,€ 1765), Fruticola schrencki (Middendorff, 1851), and Euco- ginian alluvium of the Yenisei River (in the surroundings of nulus fulvus (Müller, 1774).

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4.5. Middle Angara region V. alpestris) indicate the existence of cold climate conditions. They are Holarctic and are now common in Central Europe. The slug In this region, the Karginian deposits form the lower parts of the shells (Limacidae) show that the climate around Bol'shoj Naryn site terraces with heights ranging between 14 and 17 m. They are was not extremely cold. The mollusc community (Limacidae, characterized by stream (pebble) and floodplain (sandy loam and P. muscorum, and V. modesta) do not reflect the existence of an sand) alluvial facies (Ravskyi, 1959). These deposits contain Bison extremely cold climate during MIS 3 at Bol'shoj Naryn Palaeolithic priscus aff. longicornis V.Gromova, 1935 remains and wood dated at site, and the cooler environment was only recognized thanks to the 24,800 ± 120 BP. Forest pollen dominated in the palynological presence of V. tenuilabis and V. alpestris. spectra (Pinus, Abies, Picea and Betula)(Zeitlin et al., 1984). Starobogatov (1970), Zhadyn (1952) and Popova (1981) consid- ered that the modern malacofauna of the southern part of Siberia 4.6. Upper Angara area including Altai, Tyva, Fore-Baikalia, Trans-Baikalia, and Mongolia, is a part of the Palaeoarctic zoogeographical region. Likharev and In this area (Irkutsk), the Karginian deposits are represented by a Rammelmeier (1952) subdivided the Palaeoarctic region into paleosol which covers the alluvium of the terrace (15e16 m height) several subregions and provinces. The territory described in this and by alluvium of the lower part of the terrace (12 m height). They paper belongs to the taiga province (zone) of the European-Siberian are known in the Malta and Bol'shoj Naryn Palaeolithic sites. The subregion. The coniferous forests which grow in the cold climate thickness of the paleosol in the Shamot (fireclay) plant quarry have various negative impacts on mollusc development: 1) needles (Belaya River valley) is 0.4 m, and the deposits are deformed are not edible; 2) trees of the forest grow thicker, sunlight pene- because of the cryoturbation processes. Because this soil is tration is limited, temperature is low, spring snow persists for a long considered to be the remnant of a forest soil, throughout the Kar- time; and 3) the soils are poor in organic matter and the grass cover ginian Interstadial the climate was not cooler than now (Zeitlin is poorly developed. The terrestrial malacofauna of the taiga is thus et al., 1984). very poor. According to Likharev and Rammelmeier (1952) it is characterized by 9 boreal species (Succinea chrysis Westerlund, 4.7. Southern Fore-Ural region 1883, V. alpestris, V. modesta, Acanthinula harpa (Say, 1824), Arion subfuscus (Draparnaud, 1805), Agriolimax reticulatus (Müller, 1774), The Gornovo locality is characterized by the Upper Pleistocene A. heperboreus (Westerlund, 1876), Fruticicola schrenckii (Mid- deposits, accumulated during a climate warming episode (MIS 3) dendorf, 1851), Pseudotrichia rubiginosa (A.Schmidt, 1853)) and Si- during the Valdai period. The malacological fauna which contains berian taiga elements such as: V. tenuilabris, Pseudotrichia numerous terrestrial S. oblonga, P. muscorum, V. tenuilabris, and nordenskioldi€ (Westerlund, 1876). The boreal species are common in freshwater mollusca are part of the lacustrine deposits of the Tab- the taiga as well as in the tundra environment. V. alpestris, ulda horizon (bed 2, Gornovo II) (Yakhemovich et al., 1987; V. modesta and A. harpa also inhabit the mountains of Europe, Cri- Danukalova et al., 2002). Spruce wood debris found in these de- mea and Caucasus, which support the existence of ancient faunistic posits are dated at 26,950 ± 560; 26,990 ± 150; 28,800 ± 125; and connections between tundra, taiga and the mountainous regions 29,700 ± 1250 BP. The comparison of the mollusc complexes from and suggest that these species have inhabited this area since the the Bol'shoj Naryn locality with the molluscs from the last glacial Pliocene. However, Likharev and Rammelmeier (1952) suggested period (MIS 2) of Gornovo locality shows strong similarities. S. that the malacofauna of the taiga province is a new type of fauna oblonga, P. muscorum, V. tenuilabris with some freshwater species which did not exist before the Quaternary glaciations. are present in water-slope periglacial loam covering dated lacus- In the collection of the shells extracted from the Karginian trine deposits at Gornovo (Yakhemovich et al., 1987; Danukalova paleosol of Bol'shoj Naryn site, we have studied six species: et al., 2002). S. oblonga, V. tenuilabris, P. muscorum, V. cf. modesta, V. alpestris, and Limacidae. If we take account of the determination problem of the 4.8. Discussion slugs at the base of the fossil remains and the location of the studied site which is situated in a river valley and not in a taiga The mollusc complexes of the Karginian Interstadial of the Fore- forest, the Karginian species show some similarities with the Baikalian area are not numerous, and most were found in the al- modern molluscs. luvial deposits where freshwater gastropods, bivalves and rare terrestrial species are present. Therefore, the terrestrial molluscs 5. Conclusion described in this paper are unique. They were found for the first time in situ in the Karginian paleosol of the Fore-Baikalian region The deposits of the Karginian horizon are widespread in the (as well as of the all-Siberian area). The correlation between the territory of Siberia including the Eastern Siberia area (the Siberian data obtained by malacological investigations on the Bol'shoj Naryn Platform). They are known in the glacial and periglacial areas. In the site and the palynological and mammalian records suggest a southern part of Siberia, the deposits attributed to the Karginian moderately cold and humid climate in the Fore-Baikal region dur- horizon mainly developed in the river valleys, and are represented ing the latter half of MIS 3, with predominant open landscapes of by alluvial (stream and floodplain) facies and by paleosol. The steppes and relatively limited taiga forest, tundra, and wetlands mollusc complexes of the Karginian Interstadial of the Fore- (Sato et al., 2014). Baikalian area are not numerous, and most were found in the al- The malacological results we obtained are relatively poor, which luvial deposits where freshwater gastropods, bivalves and rare can be related with the aggressive acid environment of the soil, terrestrial species are living. The Bol'shoj Naryn I Palaeolithic lo- which dissolves the carbonate, or perhaps because of local living cality incorporates the first late Karginian suc- conditions. Molluscs provide additional information which com- cession known in the Fore-Baikalian area. plemented the palynological data in the vegetation reconstruction. In terms of species composition, the malacocomplex of the They both support the existence of humid grassland with bushes or Bol'shoj Naryn site is the closest to the equivalent complex of parkland environment. Gornovo in the Southern Fore-Urals and resembles mollusc com- However, it is also possible to constrain the climate even if we plexes, known in the periglacial zone of Kudashevo in Southern have only six species of shells. Two species (V. tenuilabris and Urals (ending phase of the Late Valdai, equivalent to MIS 2). The

Please cite this article in press as: Danukalova, G., et al., The molluscs record: A tool for reconstruction of the Late Pleistocene (MIS 3) palaeoenvironment of the Bol'shoj Naryn site area (Fore-Baikal region, Eastern Siberia, Russia), Quaternary International (2014), http:// dx.doi.org/10.1016/j.quaint.2014.08.034 G. Danukalova et al. / Quaternary International xxx (2014) 1e10 9 climatic conditions which existed during the late phase of the Falkner, G., Ripken, T.E.J., Falkner, M., 2002. Mollusques continentaux de France. e Karginian interglacial period in the surroundings of the Bol'shoj Liste de References annotees et Bibliographie. Patrimoines naturels 52, 1 350. Museum National d'Histoire Naturelle (Paris). France. Naryn locality could have been cooler than in the Southern Urals at Florensov, N.A., 1960. Mesozoic and Cenozoic Depressions of the Fore-Baicalian the same time. Zone. Academy of Sciences of the USSR Press, Moscow-Leningrad, 257 pp. Molluscs provided additional informations which completed (in Russian). Frechen, M., Zander, A., Zykina, V., Boenigk, W., 2005. The loess record from the the palynological data. The examined species of molluscs are section at Kurtak in Middle Siberia. Palaeogeography, Palaeoclimatology, characteristic of biotopes showing an increasing rate of humidity. Palaeoecology, 228e244. These biotopes probably corresponded with floodplain or meadows Germain, L., 1930. Faune de France. Mollusques terrestres et fluviatiles. t. 21 et 22. Lechevalier, Paris, 893 pp. located near a water reservoir. The mollusc community (Limacidae, Golubeva, L.V., Ravskyi, E.I., 1962. Anthropogene of the Tunka Depressions. In: P. muscorum and V. modesta) does not reflect an extremely cold Proceedings of the Commission on the Quaternary Period Investigations, vol. 19. climate during MIS 3 close to the site. The generally cooler envi- Academy of Sciences of the USSR Press, Moscow, pp. 240e259 (in Russian). fi Hanley, S., 1855. Ipsa Linnaei conchylia. Williams and Norgate Press, London, ronment was identi ed by the presence of V. tenuilabis and 556 pp. V. alpestris. These data show the importance of molluscs in the Kerney, M.P., Cameron, R.A.D., 1999. Guide des Escargots et limaces d'Europe. palaeoenvironmental reconstructions. Correlation of the data ob- Delachaux et Niestle S.A. 3, Lausanne, 70 pp. tained on malacological investigations and the other paleontolog- Kerney, M.P., Cameron, R.A., Jungbluth, J.H., 1983. The Land Snails of North and Middle Europe. Hamburg, 384 pp. ical data obtained on palynological and vertebrates record (Sato Kind, N.V., 1974. Geochronology of the Late Anthropogene According to Isotopic et al., 2014) suggests a moderately cold and humid climate in the Data. 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Boundary of the Middle and Late Palaeolithic of the Northern Asia: problem of the Karginian time. Issue 2. In: Problems of a Man and Envi- This work was partly funded by the subsidy of the Russian ronment Interaction. IPOS Siberian Branch of the RAS Press, Tyumen, pp. 11e18 Government to support the Program of Competitive Growth of (in Russian). Laukhin, S.M., Firsov, A.M., 2011. Some distinctions of big and small interglacials (by Kazan Federal University among World's Leading Academic Centers way of example of Western Siberia during Late Pleistocene). In: Korsakova, P., and The JSPS Grant-in-Aid for Science Research (A) (Grant No. Kolka, V.V., Chistyakova, L.D. (Eds), The Quaternary in All of its Variety. Basic 21251009, Title: “Overall Research on the Adaptive Behavior of Late Issues, Results, and Major Trends of Further Research, Proceedings of the VII All- e ” russian Quaternary Conference (Apatity, September 12 17, 2011). Apatity; St Paleolithic Man in Northeastern Eurasia , Head Researcher: T. Sato). Petersburg, vol. 2, pp. 23e25 (in Russian). We heartily thank professors J.-L. Monnier and J.-P. Lefort, labora- Laukhin, S.A., Alekseev, V.A., Mylnikova, Z.K., Chepalyga, A.L., Gracheva, O.E., tory of Archeosciences of Rennes 1 University (France) for opening Smirnov, I.V., 1971. Reports of the Academy of Sciences 196 (1), 172e175 (in Russian). up opportunities to take photos of mollusc shells on their equip- Lazukov, G.I., 1981. Nature and the Ancient Man (main Stages of the Palaeolithic ment. The authors would like also to thank Professor Jean-Pierre Man Nature and Culture Development on the USSR Territory during Pleisto- Lefort for his help during the English translation; Mrs. Natalya cene). Mysl' Press, Moscow, 223 pp. (in Russian). Bogatyreva for her help during the labeling of the collections. The Likharev, I.M., Rammelmeier, E.S., 1952. Land Molluscs of the Fauna of the USSR. 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Please cite this article in press as: Danukalova, G., et al., The molluscs record: A tool for reconstruction of the Late Pleistocene (MIS 3) palaeoenvironment of the Bol'shoj Naryn site area (Fore-Baikal region, Eastern Siberia, Russia), Quaternary International (2014), http:// dx.doi.org/10.1016/j.quaint.2014.08.034