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Quaternary International xxx (2013) 1e11

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Quaternary International

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Vertebrates from the Middle Pleistocene locality Lysa Gora 1in Ukraine

Leonid Rekovets a, Stanislav �Cermák b, Olexandr Kovalchuk c, Valentin Prisyazhniuk d,Dariusz Nowakowski a,*aWrocław University of Environmental and Life Sciences, Institute of Biology, Kozuchowska 5b, 51-631 Wroclaw, Polandb Institute of Geology, Academy of Sciences of the Czech Republic, v.v.i., Rozvojová 269, CZ-165 00 Prague 6-Lysolaje, Czech RepubliccNational Museum of Natural History, National Academy of Sciences of Ukraine, B. Khmelnitskogo str. 15, 01-601 Kyiv, Ukrained Institute of Geological Sciences, National Academy of Sciences of Ukraine, Kyiv, Ukraine

a r t i c l e i n f o

Article history:Available online xxx

* Corresponding author.E-mail addresses: leonid.rekovets@up.wroc.pl (L. R

(S. �Cermák), Biologiest@ukr.net (O. Kovalchuk), daridarekn@hot.pl (D. Nowakowski).

1040-6182/$ e see front matter � 2013 Elsevier Ltd ahttp://dx.doi.org/10.1016/j.quaint.2013.10.016

Please cite this article in press as: RekovetInternational (2013), http://dx.doi.org/10.10

a b s t r a c t

A vertebrate fauna from the newMiddle Pleistocene locality Lysa Gora 1 of Southern Ukraine (Zaporizhyaregion) is described. The locality has yielded an assemblage of at least 11 taxa of bony fishes (Cypri-niformes, Siluriformes, Esociformes, Perciformes) and small mammals (Lagomorpha e 2 taxa, Rodentiae 11 species). The species composition of Arvicolidae, particularly the presence of Lagurus transiens,Eolagurus luteus, Arvicola mosbachensis, Microtus (Terricola) arvalidens, and Microtus (Stenocranius) gre-galis, as well as the structure of the fauna, suggests Middle Pleistocene age (MQR 4, MIS 13-12, EarlyToringian). New data on the paleoecology of the locality and taxonomy/phylogeny of selected taxa arediscussed.

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1. Introduction

The Miocene, Pliocene, and Pleistocene theriofaunas of theEastern Paratethys are relatively well understood and published bymany authors (cf. Topachevsky, 1973; Alexandrova, 1976; Alexeeva,1977; Markova, 1982; Korotkevich, 1988; Topachevsky and Nesin,1989; Rekovets, 1994; Tesakov, 2004; Agadjanian, 2009;Krokhmal and Rekovets, 2010). A historical review of geology andfauna of Zaporozhye region was recently provided by Manyuk(2009) and Rekovets and Pashkov (2009). In this region, the Sar-matian marine sediments of the Late Miocene basin consist mainlyof limestone, clay, sand and other rocks, locally extending to thesurface. They are covered by Pliocene marine and/or continentaldeposits and also by Pleistocene silty sediments (Semenenko,1987). These deposits contain mainly late Neogene and Pleisto-cene vertebrate remains (particularly mammals). Selected taxa ofthat region were studied in detail by Markova (1982) andKorotkevich (1988).

ekovets), cermaks@gli.cas.czusz.nowakowski@up.wroc.pl,

nd INQUA. All rights reserved.

s, L., et al., Vertebrates from16/j.quaint.2013.10.016

2. Material and methods

The investigatedmaterial was collected in the alluvial sedimentsof Lysa Gora 1 (Zaporozhye region) during three field expeditions(2009e2011). Referred fossil material is housed in the Departmentof Vertebrate Palaeozoology and Palaeontological Museum (Na-tional Museum of Natural History NAS of Ukraine, Kiev).

A direct determination of the fossil material was processed onthe basis of diagnostic cranial and postcranial elements studying,and also sensu Lepiksaar (1994) and Radu (2005). Recent fish boneswere used for comparison. Ichthyologic systematics in this paperfollow Movchan (2011). Paleoecological analysis of freshwater fishassemblage was processed using ecotopic preferences of theirrecent analogues.

The dental terminology and/or metrics used here to describedental structures of Lagomorpha and Rodentia follow Stirton (1935),López-Martínez (1974), Rabeder (1981), Erbajeva (1988), Heinrich(1990). All drawings and measurements were made using a binoc-ular microscope with a camera lucida and ocular micrometer.

Abbreviations used in the text and tables are as follows: HPT e

height of the pharyngeal tooth, WPT e width of the pharyngealtooth, A e anteroconid length, AC e anteroconid, A/L e the ratio ofanteroconid length to the tooth length, AW e anteroconid width,CCD e conid confluence distance, H e height, L e tooth length,Ltrig e trigonid length, PL e posteroconid length, T e triangle vel

the Middle Pleistocene locality Lysa Gora 1 in Ukraine, Quaternary

Ochotona sp. (cf. gr. ?pusilla) 9Ochotona sp. 1Spermophilus cf. suslicus 10Trogontherium minus 2Allactaga cf. ucrainicus 1

L. Rekovets et al. / Quaternary International xxx (2013) 1e112

conid, W e tooth width, Want e anteroloph width, Wpost e

posteroloph width, Wtal e talonid width, Wtrig e trigonid width.Capital and lower-case letters, P/p (premolar) and M/m (molar),refer to upper and lower cheek-teeth, respectively.

Sylvaemus sylvaticus 3Lagurus transiens 18Eolagurus luteus gromovi 2Arvicola mosbachensis 5Microtus (Terricola) arvalidens 14Microtus (Stenocranius) gregalis 3Microtus nivaloides (¼ arvalinus) 2Microtus sp. (cf. Chionomys ex gr. nivalis) 1

3. Geological setting and taphonomy

The sites of Lysa Gora were discovered in 2009 by the Palae-ontological Museum (National Museum of Natural History NAS ofUkraine, Kiev). They are situated near Vasilyevka, Zaporozhye re-gion, Ukraine (47�270N, 33�150E, Fig. 1). The locality is a naturaloutcrop (20 m high) of Miocene, Pliocene, and Pleistocene alluvialsediments of the left bank of Karachekrak River at its confluencewith the Dnieper River (Fig. 2). Two horizons of sands and gravelswith fossil remnants, of different ages, were distinguished withindeposits of clays, limestones, loesses and fossil soils: Lysa Gora 1 e

the Middle Pleistocene (Early Toringian, MQR 4, MIS 13-12) andLysa Gora 2 e upper Miocene (Sarmatian, MN-11, Early Turolian)with remains of Ischymomys ponticus Topachevski, Skorik, Reko-vets, 1978. Numerous bones of freshwater fishes also were found inthe Sarmatian sediments: Leuciscus sp. (4), Rutilus rutilus (11),Rutilus cf. frisii (3), Rutilus sp. (13), Scardinius erythrophthalmus (3),Scardinius sp. (86), Chondrostoma sp. (3), Cyprinus cf. carpio (1),Carassius carassius (1), Carassius sp. (8), Tinca sp. (5), and Silurus sp.(9). The presence of a large number of Scardinius, Rutilus, and Silurusremnants indicates that the climate in the late Miocene of SouthernUkraine was warm and mild, without sharp seasonal temperaturechanges.

Ancient alluvial sediments (Lysa Gora 1) represent a terrace,which is embedded (or inwashed) in deposits of dense dark clays ofthe upper Miocene. Younger sediments of Lysa Gora 1 lie hypso-metrically 7e8 m below the older sands of Lysa Gora 2. The sectionis composed of the following layers (Fig. 3):

1. Modern soil 0.5 m2. Pale yellow, dusty loess 1.5 m3. Loess-like fawn loams with traces of soil formation 5.0 m4. Terrace clays 1.0 m5. Gravel limestone with sand, and fish and small mammal remnants

(Lysa Gora 1)1.5 m

6. Conglomerate of Sarmatian limestones gravel and pebbles overlyingclays, with deep erosion (terrace)

1.5 m

7. Dark gray sticky ferruginated clays with gypsum crystal inclusions(late Miocene deposits e base of the Pleistocene terrace)

15.0 m

Fig. 1. Lysa Gora locality on the map of Ukraine.

Ancient alluvial deposits with bones are very dense, sometimescemented, coarse- and fine-grained sands and grits with minorlayers of clay, which often have redeposited clamshells of Sarma-tian deposits (Late Miocene). There are many fish remnants, somebone fragments of large mammals, and also teeth and bones ofsmall mammals in the third alluvial layer of the ancient Pleistoceneterrace (Lysa Gora 1). The faunal record of freshwater fishes waspublished earlier (Kovalchuk and Rekovets, 2012). Identified spe-cies are represented by elements of viscerocranial skeleton, frag-ments of branchial arches and isolated pharyngeal teeth, and alsovertebrae, fragments of ribs, and flipper rays. Preservation of theosteological material resulted from their alluvial genesis. Theboniferous horizon of Lysa Gora 1 differs considerably, with fewerfish remnants (123 bone fragments, including 95 diagnostic),commonly broken.

Qualitative and quantitative composition of theriofauna issignificantly augmented compared with the list in Krokhmal andRekovets (2010):

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The taxonomic composition of this association and morpho-metric parameters of Arvicolidae suggest late Tiraspolian age,Middle Pleistocene, Early Toringian, MQR 4 (Vangenheim et al.,2001).

4. Systematic paleontology

CLASSIS ACTINOPTERYGII KLEIN, 1885Order Cypriniformes Goodrich, 1909Family Cyprinidae Fleming, 1822Genus Leuciscus Cuvier, 1816Leuciscus sp.Referred material e 4 pharyngeal teeth.Measurements e HPT ¼ 4.0e4.9e5.4 mm, WPT ¼ 1.4e1.6e

1.8 mm.Remarks e Remnants of Leuciscus cf. idus are known from the

Middle Pleistocene sediments of England, the West Runton Fresh-water Bed near Cromer, Norfolk (Böhme, 2010), and also from theLower Pleistocene of Germany, Stuttgart Wilhelma Bärengehege(Böttcher, 1994). Some bones of Leuciscus sp. were identified inmaterials of the Middle Pleistocene localities of Bobkovo in Russia(Sytchevskaya, 1989), Bobovek in Slovenia (Gaudant, 1978) andLower Pleistocene deposits of Spain: Barranco León 5, Dépressionde Guadix, Baza, Granada (Agustí et al., 2010). On the territory ofUkraine fossil remnants of this species are known from the MiddlePleistocene of Medzhybozh (Kovalchuk, 2012) and also on theNovgorod-Siversky Palaeolithic camp on the Desna River (Nikolsky,1945, 1952; Lebedev, 1960).

Genus Rutilus Rafinesque, 1820Rutilus cf. frisii (Nordmann, 1840)Referred material e 4 pharyngeal teeth.Measurements e HPT ¼ 1.7e2.8e3.2 mm, WPT ¼ 2.2e2.8e

3.1 mm.Remarks e Isolated pharyngeal teeth, fragments of pharyn-

geal bones, and caudal fin bones of this species were found in

the Middle Pleistocene locality Lysa Gora 1 in Ukraine, Quaternary

Fig. 2. Geological profile of the Lysa Gora 1 locality.

L. Rekovets et al. / Quaternary International xxx (2013) 1e11 3

excavations of the Late Palaeolithic campMurzak-Koba (Crimea) bythe expedition under the guidance of S.M. Bibikov (Lebedev, 1960).Fossil remnants of R. frisii also are known from the Pleistocenesediments of the Middle Dnieper (Shpet, 1949; Tarashchuk, 1962)and Medzhybozh (Kovalchuk, 2012).

Rutilus sp.Referred material e 2 pharyngeal teeth.Measurements e HPT ¼ 2.4e4.1e5.7 mm, WPT ¼ 1.8e2.2e

2.5 mm.Remarks e Remnants of the Rutilus are known from the West

Runton in England (Böhme, 2010), StuttgartWilhelma Bärengehegein Germany (Böttcher, 1994), and Barkowice near Sulejów in Poland

Fig. 3. Scheme of the geological cross-section of Lysa Gora.

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(Paw1owska, 1963). Bones of Rutilus rutilus pleistocenicus weredescribed from the Middle Pleistocene sediments of Bobkovo,Russia (Sytchevskaya, 1989). On the territory of Ukraine fossilremnants of this species are known from the Middle Pleistocene ofMedzhybozh (Kovalchuk, 2012) and also on the Novgorod-SiverskyPalaeolithic camp on the Desna River (Nikolsky, 1945, 1952;Lebedev, 1960).

Genus Scardinius Bonaparte, 1837Scardinius erythrophthalmus (Linnaeus, 1758)Referred material e 2 pharyngeal teeth.Measurements e HPT ¼ 7.6e8.0e8.3 mm, WPT ¼ 2.2e2.6e

3.0 mm.Remarks e Remnants of the Scardinius genus are numerous in

the Pleistocene sediments of Europe. Particularly, isolated bones of S.erythrophthalmus have been found on the locality West RuntonFreshwater Bed near Cromer, England (Böhme, 2010), Stuttgart Wil-helma Bärengehege, Germany (Böttcher, 1994), and also in the LatePleistocene of Poland, Barkowice near Sulejow (Paw1owska, 1963).

Scardinius sp.Referred material e 37 pharyngeal teeth.Measurements e HPT ¼ 2.3e4.2e7.5 mm, WPT ¼ 0.8e2.2e

2.4 mm.Remarks e The conical pharyngeal teeth are hooked at the

tips. The grinding surface is very narrowwith a higher edge that has5e7 robust pointed jags and a lower edge without jags. The hook isrobust, and the angle of grinding surface is 20�.

Genus Chondrostoma Agassiz, 1832Chondrostoma cf. nasus (Linnaeus, 1758)Referred material e 2 pharyngeal teeth.Measurements e HPT ¼ 2.4e2.8e3.2 mm, WPT ¼ 1.4e1.45e

1.5 mm.Remarkse This species is identified in materials of the Middle

Pleistocene locality of Medzhybozh (Kovalchuk, 2012).

Genus Cyprinus Linnaeus, 1758Cyprinus cf. carpio Linnaeus, 1758Referred material e 5 pharyngeal teeth.Measurements e HPT ¼ 2.0e4.0e7.5 mm, WPT ¼ 2.6e4.3e

9.1 mm.Remarks e Fossil carp scale was found in the Lower Pleisto-

cene sediments of Germany (Böttcher, 1994). Tarashchuk (1962)writes about the find of isolated pharyngeal teeth of C. carpio,dated to Early Anthropogene, on the right bank of Arabka gulchnear Tichonovka (Melitopol’sky district, Zaporizhia region,Ukraine). Fossil remains of carp fishes also are known from theMezhybozh locality (Kovalchuk, 2012).

Genus Carassius Jarocki, 1822Carassius carassius (Linnaeus, 1758)Referred material e 10 pharyngeal teeth.Measurements e HPT ¼ 1.2e1.9e2.2 mm, WPT ¼ 2.5e2.7e

5.1 mm.Remarks e Remnants of C. carassius are known from the Up-

per Pleistocene of Poland, Barkowice near Sulejów (Paw1owska,1963), and also from Medzhybozh in Ukraine (Kovalchuk, 2012).

Carassius sp.Referred material e 6 pharyngeal teeth.Measurements e HPT ¼ 1.5e2.2e3.0 mm, WPT ¼ 2.2e3.1e

4.0 mm.Remarks e The teeth are submolar and compressed with a

broad grinding surface, drawn to width. The grinding surface is

the Middle Pleistocene locality Lysa Gora 1 in Ukraine, Quaternary

Fig. 4. Morphology of p3 in ochotonids from Lysa Gora 1. 1-2, 2a e Ochotona sp. (cf. gr.?pusilla), 1-2 e occlusal view, 2a e root view; 3 e Ochotona sp., occlusal view. Scale bar:1 mm.

L. Rekovets et al. / Quaternary International xxx (2013) 1e114

obliquewith a distinct sharp frontal edge. Sometimes there is a verysmall marginal tip on the grinding surface, at the base of the teeth isbroken. The material contains both smaller and larger teeth.

Genus Tinca Cuvier, 1816Tinca sp.Referred material e 9 pharyngeal teeth.Measurements e HPT ¼ 2.1e3.6e8.8 mm, WPT ¼ 2.4e4.5e

7.0 mm.Remarks e The teeth of Tinca sp. are subconical and com-

pressed, drawn to width. The crown is bluntly rounded at thebroad grinding surface is oval. All teeth have distinct, blunt andhooked tips. Tench remnants are very numerous in the Pleistocenesediments of Europe, described from the Middle Pleistocene ofEngland (Böhme, 2010) and Germany, Stuttgart quarries Hass undLauster (Böttcher, 1994). It occurs also at Medzhybozh (Kovalchuk,2012).

Family Cobitidae Swainson, 1839Genus Cobitis Linnaeus, 1758Cobitis sp.Referred material e Broken lateral ethmoid.Measurements e H ¼ 5.0 mm, W ¼ 1.5 mm.Remarks e The posteromedial spine of the lateral ethmoid is

long, slender and slightly curved, the posterolateral spine is shortand stump-shaped, the medial projection is not prominent, thelateral projection is not preserved. The base of the lateral ethmoid isworn.

Order Siluriformes Cuvier, 1816Family Siluridae Cuvier, 1816Genus Silurus Linnaeus, 1758Silurus glanis Linnaeus, 1758Referred material e 1 tooth.Measurements e H ¼ 3.7 mm, W ¼ 0.8 mm.Remarks e Cross-sections of the small, pointed, more or less

curved teeth are nearly circular, decreasing regularly in diametertowards the tip. The morphology of the teeth corresponds to thoseof modern S. glanis. Fossil remnants of this species are known fromthe Middle Pleistocene Bilzingsleben locality, north of Erfurt,Thuringia, Germany (Böhme, 1998) and also in materials of Medz-hybozh (Kovalchuk, 2012).

Order Esociformes Bleeker, 1858Family Esocidae Cuvier, 1816Genus Esox Linnaeus, 1758Esox lucius Linnaeus, 1758Referred material e fragment of dentale with tooth, 4 isolated

teeth.Measurements e Total height of the bone is 4.1 mm, width e

3.3 mm, tooth height ¼ 4.4e6.3e11.1 mm, tooth width ¼ 1.8e2.3e3.8 mm.

Remarks e The long, slender, pointed teeth have sharp edges.Cross-sections of the teeth are interiorly smooth and exteriorlyconvex. This species occurs in the Lower Pleistocene of Germany(Böttcher, 1994). Esox cf. lucius is described from the Middle Pleis-tocene locality of Bobkovo, Russia (Sytchevskaya, 1989) andMedzhybozh, Ukraine (Kovalchuk, 2012).

Order Perciformes Bleeker, 1859Family Percidae Cuvier, 1816Genus Perca Linnaeus, 1758Perca sp. nov.Referred material e broken os praeoperculare.Measurements e L ¼ 10.4 mm, W ¼ 4.4 mm.

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Comparison and remarks e New fossil Perca species charac-terized by broadest praeoperculum with smaller, frequent massiverounded cogs. Total length of the complete bone is near 4 cm. Bonesurface is raised, slightly bulging. Angulus superior is destroyed,angulus anterior is partially breaked. Pori canales lineae lateralisare well-preserved. Medial part of the os praeoperculare is sharplydefined, with developed relief. Praeopercular limbus posterior isjagged, similar to the all species of Perca Linnaeus, 1758.

Cogs of the os praeoperculare should be uniform in size,whereas Gymnocephalus Bloch, 1793, the nearest to Perca genus ofthe family Percidae based on mitochondrial DNA sequencing (Songet al., 1998; Sloss et al., 2004) has 3 long large cogs at the point ofpraeopercular angle. Contrary to the praeoperculum of Perca flu-viatiliswith flat saw-like infrequent cogs, praeopercular cogs of thenew species are more frequent (cogs quantity proportion betweenP. fluviatilis and Perca neopleistocenica is 1:2), massive, adjoined faceto face and rounded in the cross section. Judging from the pre-served lamina praeopercularis, the angle among two ends of boneis 120� (praeopercular angle of P. fluviatilis is 90�). Plate of thepraeoperculum is broad, maximum width is 0.64 cm.

CLASSIS MAMMALIA LINNAEUS, 1758Order Lagomorpha Brandt, 1855Family Ochotonidae Thomas, 1897Genus Ochotona Link, 1795Ochotona sp. (cf. gr. ?pusilla)Fig. 4: 1-2, 2aReferred material e Lysa Gora 1: p3 dex (LG2010/O1), frag-

ment of p3 sin (LG2011/O1), anteroloph of M2 sin (LG2009/O2), M2dex, subadult (LG2009/O3). Lysa Gora 1 (clays): p4 dex (LG2010/O2), P4 or M1 sin (LG2010/O3), P4 or M1 dex (LG2010/O4), brokentalonid of m1, juvenile (LG2010/O5), fragment of trigonid of sinlower molariform (LG2010/O6).

Measurementse p3 (LG2010/O1): L¼ 1.48 mm,W¼ 1.52mm,A ¼ 0.52 mm, AW ¼ 0.53 mm, PL ¼ 1.03 mm, CCD ¼ 0.27 mm; p3(LG2011/O1, measured at root side): L ¼ 1.43 mm, W ¼ 1.47 mm,A ¼ 0.50 mm, AW ¼ 0.52 mm, PL ¼ 1.03 mm, CCD ¼ 0.20 mm; M2(LG2009/O2): Wpost ¼ 2.03 mm, M2 (LG2009/O3): L ¼ 1.33 mm,Want ¼ 1.83 mm,Wpost¼ 1.57 mm; p4 (LG2010/O2): L¼ 1.57 mm,Wtrig ¼ 1.47, Wtal ¼ 1.58; P4/M1 (LG2010/O3): L ¼ 1.63 mm,Want ¼ 2.73 mm, Wpost ¼ 2.63 mm; P4/M1 (LG2010/O4)L ¼ 1.40 mm, Want ¼ 2.30 mm, Wpost ¼ 2.53 mm.

Description e The available two p3s belongs to a mediumsized Ochotona. The occlusal surface, preserved only in one p3,possesses a relatively small and simple anteroconid with roundedtop. Non-plicated para- and protoflexid are short and nearly of thesame depth. The confluence between anteroconid and poster-oconid is wide. The lingual margin of posteroconid is relativelystraight. In the second p3, the occlusal surface is damaged in itsanterior and lingual part. Nevertheless, from the root view allmorphostructures are preserved. With exception of concave ante-rior walls of anteroconid (very common shape in root end of p3),

the Middle Pleistocene locality Lysa Gora 1 in Ukraine, Quaternary

Fig. 5. Morphology of teeth of rodents from Lysa Gora 1. 1 e P3; 2 e P4; 3,6 e M1 e 2;4,5 e M3; 7 e p4; 8e10 e m1e2 e Spermophilus cf. suslicus.

L. Rekovets et al. / Quaternary International xxx (2013) 1e11 5

the other important features are compatible with those preservedon the occlusal surface of the former p3. The available upper andlower molariforms do not differ morphologically from the corre-sponding teeth of Ochotona species.

Comparison and remarks e In its general occlusal pattern,taking into account the accompanying faunal assemblages,the both teeth are somewhat similar to Ochotona pusilla; partic-ularly by small rounded anteroconid and wide anteroconid-posteroconid junction. On the other hand, the available teethstrongly differ from this species, both fossil and recent, in itsmuch larger size. Moreover, the typical O. pusilla has also moretriangular appearance of p3 (i.e., its occlusal outline is generallyshorter than wide and metaconid is less developed). The small-sized form sharing general features of p3 fairly typical forO. pusilla were found in the close localities Popovo 1 and 2(correlated by Rekovets and Pashkov (2009) with the Middle-LateKhaprovian faunal complex; i.e, cf. late MN 16 sensu Nesin andNadachowski, 2001 or cf. early MN 17 (ca 2.4 Ma) sensuTesakov, 2004). These ochotonids were tentatively determined by�Cermák (2010) as Ochotona sp. (ex gr. pusilla). Unfortunately, thegeological settings together with relatively fragmentary fossilrecord do not allow excluding the presence of redeposited olderelements in these localities. Within Ukraine (and also in Europe),the fossil record of undoubted well-dated O. pusilla is known onlyfrom the Late Pleistocene. One of the most extensive records ofO. pusilla in Ukraine comes from the Late Pleistocene locality ofNovgorod-Severskiy (Rekovets, 1985). This population, in additionto the above features typical for O. pusilla, is characterized by veryshort para- and protoflexid in p3 (�Cermák, unpubl. data). Withinthe Early Pleistocene (VillányianeBiharian), the other phenotypicgroup of Europe is represented by ochotonids morphodimen-sionally close to Ochotona polonica. All these ochotonids, recordedfrom many localities of Central and Eastern Europe (see �Cermák,2009 for details), differ from the p3s under comparison in theirlarger p3 anteroconid and narrower p3 anteroconideposter-oconid junction, or smaller size. The fragmentary teeth undercomparison are also similar, particularly by relatively smallanteroconid and wide anteroconideposteroconid junction, withsome large pikas reported from several Pleistocene localities ofTranscaucasia (see �Cermák et al., 2006 for details). These formswere mostly referred to Ochotona transcaucasica (cf. Markova,1982; Erbajeva, 1988; Averianov and Baryshnikov, 1992), thespecies described by Vekua (1967) from Copi, Georgia. Unfortu-nately, for these findings any detailed characteristics are not givenin the related literature, and a relevant comparison is difficult.Nevertheless, it seems that ochotonids from Transcaucasia arelarger than those available from Lysa Gora. The teeth tentativelydetermined here as Ochotona sp. (cf. gr. ?pusilla) have somemorphological features similar to some extreme morphotypes ofOchotona recorded in several Late Miocene localities of Ukraine(�Cermák, 2010).

Ochotona sp.Fig. 4: 3Referred material e Lysa Gora 1: broken p3 dex (LG2009/O1).Measurements e W ¼ 1.47 mm, PL ¼ 0.97 mm,

CCD ¼ 0.07 mm.Descriptione One fragment of p3 (with damaged anteroconid

in the all tooth shaft) of medium sized Ochotona is stronglydifferent from the two other p3s of Lysa Gora. The tooth possesses arelatively large and wide anteroconid. The non-plicated proto- andparaflexid are long, approximately of the same length. Both arebent slightly posteriorly. The anteroconideposteroconid junction isnarrow and placed symmetrically to the tooth. The lingual marginof posteroconid is straightened.

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Comparison and remarks e The only available fragment of p3differs from the above discussed teeth of Lysa Gora in its muchnarrower anteroconideposteroconid junction and larger ante-roconid. In its size and morphology, the p3 specimen is close tosome findings, morphologically close to O. polonica, recorded fromseveral localities of Central and Eastern Europe (see �Cermák, 2009for details). Within Ukraine, the available fragment is most similarto Ochotona from the Lower Pleistocene locality of Nogaisk(�Cermák, unpubl. data), but it differs in its larger size. This stillpoorly understood morphological group of Ochotona is relativelycommon in the fossil record of Ukraine (cf. �Cermák, 2010; �Cermákand Rekovets, 2010). It is also present in some Late Miocene lo-calities in the vicinity of Lysa Gora (Rekovets and Pashkov, 2009;�Cermák, 2010). Nevertheless, without an additional fossil materialfrom this locality and detailed revision of material from other lo-calities, it seems quite premature to provide more detailed com-parisons or species attributions.

Order Rodentia Bowdich, 1821Family Sciuridae Gray, 1821Genus Spermophilus Cuvier, 1825Spermophilus cf. suslicus Guldenstaedt, 1770Fig. 5: 1e10The remains of the genus are ubiquitous in the Eastern

Europe since the Late Khaprovian faunas, MN 17 (Kotlovina 2).They belong to Spermophilus (Urocitellus) nogaici Topachevski,1965, and another species e Spermophilus pygmaeus Pallas, 1778 e

appeared in the early Pleistocene (Tiraspol fauna, MQR 7), andquite later (MQR 3, 4) e Spermophilus suslicus. All these species inthe Pleistocene are consecutive subspecies in time. The Pleistocenehistory of evolution of the Spermophilus genus of Eastern Europe,including fossil and karyological data, is well understood.

Referred material e 1 P3, 1 P4, 2 M1-2, 2 M3, 1 p4, 3 m1-2.Measurements e P3: L ¼ 1.5 mm, W ¼ 1.6 mm, H ¼ 3.8 mm;

P4: H ¼ 3.5 mm; M3: L ¼ 3.0 mm; W ¼ 2.65 mm; H ¼ 4.5 mm; p4:L ¼ 2.5 mm, W ¼ 2.6 mm; m1-2: L ¼ 1.7e2.0e2.2 mm, W ¼ 2.5e2.7e2.9 mm, H ¼ 4.7 mm.

Descriptionand comparisoneMedium-sized form, its coronarylength of the upper and lower teeth is in the range 10.0e10.1 mm,

the Middle Pleistocene locality Lysa Gora 1 in Ukraine, Quaternary

L. Rekovets et al. / Quaternary International xxx (2013) 1e116

which is somewhat higher than in the Spermophilus citelloides Kor-mos, 1916 (8.4e9.2e10.1 mm) and recent S. suslicus (7.8e8.4e8,8 mm). However, these indexes are much smaller than in the EarlyPleistocene species e Spermophilus primigenius Kormos, 1934, Sper-mophilus polonius Gromov, 1965, Spermophilus dietrichi Kretzoi, 1965(Maul, 2001). Two-top P3 has two well-developed roots. P4 is rela-tively wellmolarized, theW/A ratio is 123.5% (in S. suslicuse 125.0%),anterior ridge is well developed, mesostylus in M1-2 is almost notdeveloped. Posterior crest of M3 is well developed and divided intotwo parts,W/A¼ 88.3%. E.g.,W/A in the S. suslicus¼ 105.0%, while inthe S. nogaici e 107.0%.

p4 is square shaped, W/L index is 104% (110.0% in S. suslicus,115.0% in S. citelloides, 115.0% in Spermophilus severskensis Gromov,1965), its anterior tops are almost identical in shape, hypoconid andprotoconulid are well developed, tooth has two roots; m1-2 arealmost rectangular in shape with well-developed hypoconid, W/Lindex is 138.6%, i.e. teeth are well molarized. Mesostylid on m1-2 ispresent, metalofid is well developed. In the above features, theform under study is closer to recent S. suslicus than S. citelloides, it ismarkedly different from the known Early Pleistocene species.

Family Castoridae Gray, 1821Genus Trogontherium Fischer von Waldheim, 1809Trogontherium minus Newton, 1890Fig. 6: 1e2Remains of the Trogontherium genus are authentically known

in Eastern Europe from the Late Miocene (MN 9) to the MiddlePleistocene (MQR 3). They are represented by at least three species(Trogontherium minutum Meyer, 1838, T. minus Newton, 1890 andTrogontherium cuvieri F. von Waldheim, 1809) that alternated intime (Rekovets et al., 2009). Taxonomic status of T. cuvieri as thelargest cementless form is beyond any doubt. However, the firsttwo species are smaller, integrated in the subgenus EuroxenomysSamson et Radulesco, 1973, and their taxonomy is much morequestionable (Franzen and Storch, 1975; Hugeney, 1999; Stefen andRummel, 2003). T. minus is known in Eastern Europe from thePliocene localities (fromMN 14 toMN 17), and the presence of theirremnants in more recent deposits of this region may be a result ofredeposition. This species is almost always found together withT. cuvieri to Holsteinian time.

Referred material e 1 p4, 1 m. Teeth are rounded and some-what damaged.

Measurements e L ¼ 4.4 mm,W ¼ 3.3 mm,W/L ratio ¼ 75.0%.

Fig. 6. Morphology of teeth of rodents from Lysa Gora 1. 1,2 e p4 e Trogontheriumminus; 3,4 e m3 - Allactaga cf. ucrainica; 5,6 e M1; 7 e M2; 8 e m1 e Sylvaemussylvaticus.

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Description e Relatively small sized species. Teeth withoutcement, rather worn, their height of the average is 4.2 mm, heightof the roots is almost equal to the crown height. Lower p4 has threeclosed fossettes, and much curved parafossettid (crescent) form.Well defined brand structure as a result of the chewing surfaceabrasion of the tooth is in the front of paraconid. Elongated ellipsoidmesofossettid is located parallel to the hypoflexid and occupies twothirds of the tooth width. Metafossettid also has elipsoid form,located against hypoflexid, and occupies half of the tooth crownwidth. Hypostriid is very weak due to the tooth abrasion, its heightis about 1 mm.

Comparison and remarks e Such a late finding of this form,typical for the Pliocene of Europe, seems to be doubtful (Rekovetset al., 2009). There is a reason to believe that in Eastern Europe,in contrast to Western Europe, Trogontherium disappeared duringthe Early Pleistocene, near the Villányian/Biharian boundary, i.e. caMQR 11.

Family Dipodidae Waterhouse, 1842Genus Allactaga Cuvier, 1836Allactaga cf. ucrainica Gromov et Schevtschenko, 1961Fig. 6: 3e4The history (from the late Miocene) of five-fingered gerbils in

southern Eastern Europe is relatively well understood. Themorphologically primitive genus Paralactaga is known throughoutthe Pliocene. The earliest members of Allactaga genus are knownsince the Khaprovian faunas (ca. Villányian), analyzed in detail byTopachevsky (1973). They are represented by separate species:Allactaga ucrainica, Allactaga nogaiskiensis Topachevski, 1965,Allactaga praejaculus Topachevski, 1965.

Referred material e 1 m3.Measurements e L ¼ 1.8 mm, W ¼ 1.5 mm, W/L ratio ¼ 83.3%.Description and comparison e The similar W/L ratio is also

available in the form from Kotlovina 1. However in the recentspecies Allactaga major Kerr, 1792 this ratio is an average of 115.0%.In addition, m3 of this form has developed and forked mesoconid.Tooth height is 1.2mm, it has two roots. By its proportions, the formunder study is also close to A. nogaiskiensis from Cherevychne 1,although it differs by its strongly developed mesoconid.

Family Muridae Illiger, 1811Genus Sylvaemus Ognev, 1924Sylvaemus sylvaticus Linnaeus, 1758 (¼Apodemus cf. sylvaticus)Fig. 6: 5e8Representatives of the genus Apodemus are known from the

Late Miocene of Europe represented by nearly ten species, whichrequire a revision (Maul, 1990; Nesin and Storch, 2004; Nesin,2013). Their remains are known in Eastern Europe from the LatePliocene as Apodemus dominans Kretzoi, 1959 and Apodemusargyropuloi Topachevsky, 1973 (Topachevsky and Nesin, 1989). Thefindings, probably belonging to Sylvaemus, were also reported insome Pleistocene localities of Ukraine.

Referred material e 1 m1, 1 M1, 1 M2.Measurements e M1: L ¼ 2.0 mm, W ¼ 1.3 mm, H ¼ 1.8 mm;

M2: L ¼ 1.4 mm, W ¼ 1.2 mm.Description and comparison e All teeth are quite worn

(belong to old individuals) and poorly preserved. M1 has 4 roots,occlusal surface of the tooth has three tubercles in anterior part,including T1 e moved slightly backward. Six tubercles in posteriorpart are connected by a crest and form an almost closed circle witha little additional (seventh) tubercle on the buccal side. Two tu-bercles are distinctively viewed in front of the M2, in the back ofprobably five, tooth has 4 roots. m1 has only the front part withwell-developed five contiguous tubercles; the anterior one (medialanteroconid) is very poorly developed and merged with two

the Middle Pleistocene locality Lysa Gora 1 in Ukraine, Quaternary

L. Rekovets et al. / Quaternary International xxx (2013) 1e11 7

tubercles of anteroconid. In its morphology and size, the materialunder study is very similar to those fromMedzhybozh in Ukraine. Itdiffers from recent S. sylvaticus by smaller size, backward shifted T1on M1 and poorly developed medial anteroconid.

Family Arvicolidae Gray, 1821Arvicola Lacépedè, 1799Arvicola mosbachensis Schmidtgen, 1911Fig. 7: 1e2The evolutionary history of the genus Arvicola (including

Mimomys) in the Palearctic has intensively been studied(Agadjanian, 2009), but in many respects many questions stillremain unclear or controversial (cf. Rekovets, 1989, 1990; Heinrich,1990; Maul et al., 2000). Members of the Arvicola genus are indis-pensable elements for stratigraphic correlations in almost all Tor-ingian localities of Europe (Krokhmal and Rekovets, 2010).

Referred material e 1 M1, 3 m1, 1 m3.MeasurementseM1: L¼ 3.0mm;m1: L¼ 3.5; 3.65; 3.72mm,

A ¼ 1.42; 1.68; 1.8 mm, A/L ¼ 40.5; 43.6; 45.1%; m3: L ¼ 2.2 mm.Description and comparison e The available material is not

numerous. This is not typical for ancient alluvial deposits, but may

Fig. 7. Morphology of teeth of arvicolids from Lysa Gora 1. 1e m1; 2 e M3 e Arvicolamosbachensis; 3 e m1; 4 e M3 e Lagurus transiens; 5 e m1 e Eolagurus luteus gromovi;6 e m1 e Microtus (Stenocranius) gregalis.

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also reflect the relatively dry (steppe) paleoecological conditions, ortaphonomic features. Molars have a typical morphology, not verycomplicatedm1 anteroconid structure andM3 back part. Accordingto M1 size, the population from Lysa Gora is similar to Medzhybozh(L ¼ 3.7 mm, A/L ¼ 44.0%) and differs by slightly smaller size fromthe recent Arvicola amphibius Blazius, 1858 (L ¼ 3.9 mm, A/L ¼ 46.2%). These indexes in evolutionary intermediate speciesArvicola chosaricus Alexandrova, 1976 from Khazar faunas of Volgaare 3.9 mm and 44.1% (Povodyrenko and Rekovets, 2006).

Inversion and differentiation of tooth enamel in Mimomys andArvicola genera is one of the most important indicators for strati-graphic correlations. The ratio of the thickness of posterior andanterior walls at salient angels in % (SDQ index) is a reliable crite-rion for taxonomic differentiation of these forms (Heinrich, 1990).For m1 ofMimomys, the SDQ index is in the range of 140e120%. ForM1 of A. mosbachensis from Lysa Gora this index is 109e110%, whichis very close to those indexes from different horizons of Medzhy-bozh locality (SDQ ¼ 107; 110.3; 116.8%). The population fromGunky has smaller SDQ values (106%). Significantly lower rates(SDQ ¼ 80%) are found in recent populations of A. amphibius, i.e.enamel is much thinner on the back conid walls and thicker in thefront.

Analysis of the M3 enamel inversion data confirms the thesis ofmore archaic M3 morphology compared to m1. The M3 possessesmany Mimomys attributes/features (Maul et al. 2000). The popu-lation of A. mosbachensis from Lysa Gora is geologically morearchaic in comparison to the population ofMedzhybozh and Gunky.It reflects the earliest stages of the formation of A. mosbachensisspecies. It corresponds with the early A. mosbachensis reportedfromMorozovka 2 and Bolshevik 2 (Morozovka association, MQR 4,Oka horizon).

Tribe Lagurini Kretzoi, 1955Genus Lagurus Gloger, 1871Lagurus transiens Jánossy, 1962Fig. 7: 3e4Representatives of the tribe Lagurini are the most common

group in the Pliocene and Pleistocene of Eastern Europe. Theseare different generic, subgeneric and specific taxa with differentways of morphological transformations in time and differentlevels of evolutionary advancement. In Europe the initial groups(CseriaeVillanyiaeBorsodia) and intermediate taxa (Lagurodon,Prolagurus, Lagurus) are represented by w40 species. They formbasic (at least six) lineages within the tribe.

Pleistocene history of the Lagurus in Europe can be seen uponas anagenesis with directional changes in traits over time and highrates of evolutionary change. This phenomenon is typical also in theevolution of the PraedicrostonyxeDicrostonyx lineage, which dis-tinguishes these groups from the morphologically similar trans-formations in AllophaiomyseMicrotus lineage. Transformation ofthe L. transiens Jánossy, 1962 into Lagurus lagurus Pallas, 1773 tookplace during the EarlyeLate Pleistocene (MQR 6-4). Within L.lagurus several independent chronotaxa (subspecies) are reportedin the Pleistocene faunas of Europe: Lagurus lagurus solimarensisJánossy, 1969, Lagurus lagurus pleistocaenicus Alexandrova, 1976,Lagurus lagurus dobrogicus Radulesko and Samson, 1976, Laguruslagurus trachicus Radulesko and Samson, 1976, and Lagurus lagurusmajor Zazhigin and Rekovets, 1985.

Referred material e 16 m1, 2 M3.Measurements e m1: L ¼ 2.2e2.4e2.55 mm, W ¼ 0.8e0.9e

1.0 mm, A ¼ 1.15e1.4e1.5 mm; M3: L ¼ 1.8; 2.0 mm, W ¼ 0.7e0.9 mm.

Description and comparison e Morphology of the m1 istransitional with typical characteristics for transienselagurus group.The main tooth triangles (T1-3) are well separated (with rare

the Middle Pleistocene locality Lysa Gora 1 in Ukraine, Quaternary

Fig. 8. Morphology of teeth of arvicolids from Lysa Gora 1. 7 e m1; 8 e M3 e Microtus(Terricola) arvalidens; 9 e m1 e Microtus (Microtus) nivaloides; 10 e m1 e Microtus (cf.Chionomys ex gr. nivalis).

L. Rekovets et al. / Quaternary International xxx (2013) 1e118

exceptions), triangles T4-5 are contiguous in 30% of m1. AC2 at thebase has a well-developed conids (T6-7), to varying degrees arefused (not separated), both among themselves and with AC2. Thesefeatures, together with the form of AC2, characterize m1 morpho-types, whichmostly are as follow: “transiens”e 53.3%, “posterius”e13.3% and “lagurus” e 33.3%. A morphotype composition is avail-able in the late Tiraspol faunas of Tikhonovka 1 (MQR 5-4, much-kap), where this species is referred to L. transiens (Rekovets, 1994).

A/L index of m1 is 53.5e54.5e58.0%. These indices of the formfrom Lysa Gora are close to those of Tikhonovka 1e53.9% andSemibalki 1 localities e 54.3% (MQR 5). The A/L index of the recentL. lagurus is on average 57.2%. The length of upper M3 is similar tocomparable populations of the species. m1 and M3 characteristicsof the L. transiens from Lysa Gora indicate a certain similarity of thepopulationwith those of Tikhonovka 1 and Semibalki 1 (Muchkap),and the character is more ancient in comparison with the Medz-hybozh fauna.

Genus Eolagurus Argyropulo, 1946Eolagurus luteus gromovi Topachevski, 1963Fig. 7: 5Extinct species of Eolagurus in the Eastern Europe are known

since advanced Taman faunas (Tarkhankut, MQR 9), and they arerepresented by species that are replaced regionally and over time(Eolagurus argyropuloi Gromov and Parfenova, 1951, Eluteus luteusEversmann,1840) and subspecies (Eluteus luteus adventus Rekovets,1994, Eluteus argyropuloi argyropuloi, E. luteus gromovi Topachevski,1963, Eluteus luteus volgensis Alexandrova, 1976, Eluteus luteusantecedens Rekovets, 1985) (Rekovets and Nadachowski, 1995).Their remains are relatively rare in steppe and semi-desert faunalassociations predominantly in Southeast Europe. The history ofgenus in the Pleistocene is relatively well understood, but theancestral forms of the genus have been poorly understood untilnow.

Referred material e 2 m1.Measurements e L ¼ 2.8; 3.0 mm, W ¼ 1.1; 1.3 mm, A ¼ 1.42;

1.54 mm, A/L ¼ 50.7; 51.3%.Description and comparison e Morphology of the grinding

teeth surface is typical for the species, unpaired loop AC2 is highlydeveloped, with its outer side facing angle is clearly separated (T6conid), T4 and T5 are separated, and the remaining anteroconidelements are contiguous. Enamel is differentiated by Microtus-liketype. Size dimensions are peculiar to remnants of the subspeciesfrom Bol’shevik locality 2, I. This subspecies from other similarlyaged localities (Tikhonovka 1, Krasnoselka etc.) has significantlylarge absolute values (Rekovets, 1994). E. luteus volgensis fromGunky locality is larger (L ¼ 3.23 mm) and has also larger A/Lvalues ¼ 54.8%, which corresponds to the general trend in theevolution of our species, especially on the Pre-Likhvin (Pre-Singi-lian) evolutionary stage (MQR 5, 4).

Genus Microtus Schrank, 1798Stenocranius Kastschenko, 1901Microtus (Stenocranius) gregalis Pallas, 1778Fig. 7: 6Most ancient representatives of the subgenus Stenocranius in

Europe are known from Slivia in Italy (described as Pitymys hintoniKretzoi, 1941). In the Eastern Europe this species appears in thefinal stage of Taman faunas (Cherevychne) e MQR 8-7 andsubstituted in time by transitional taxa in anagenetic trans-formational order of the group hintoniegregalis, that includesMicrotus hintoni-gregaloides, Microtus gregaloides Hinton, 1923,Microtus gregaloides-gregalis and M. gregalis Pallas, 1778, thoseconsistent morphological transformations are well documented bymany researchers (Rekovets, 1994; Tesakov, 2004; Agadjanian,

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2009). Well-established representatives of the M. gregalis firstlyappeared during Muchkap interglacial (MQR 5 zone) in localitiesPosevkino, Perevoz, Krasnoselka, Volnaya Vershina (Markova,1982;Agadjanian, 2009), i.e. on the stratigraphic level of Kozi Grzbiet eSüssenborn (Nadachowski, 1985, 1989; Maul, 1990).

Referred material e 3 m1.Measurements e L ¼ 2.7; 2.74; 2.81 mm, W ¼ 1.0; 1.05;

1.05 mm, A/L ¼ 49.3; 52.2; 53.5%.Description and comparison e Slight remnants of m1 have a

characteristic morphology for this species with an elongated andasymmetrical anteroconid loop e AC2. T7 conid is very welldeveloped, T6 conid is virtually undeveloped, and T4-5 conides arefused (not divided) among themselves, distinguishing this speciesfrom M. gregalis.

According to size features, the material under study is close tothat from Morozovka 2 locality (A/L ¼ 54.0%) and from the fewyounger Medzhybozh (A/L ¼ 52.5%) and Gunky (A/L ¼ 53.5%)populations. Remnants from the more ancient Protopopovka lo-cality yielded M. gregaloides with smaller A/L e 50.0% and theprevalence of hintoni-gregaloides morphotype.

Genus Terricola Fatio, 1867Microtus (Terricola) arvalidens Kretzoi, 1958Fig. 8: 7e8Representatives of theMicrotus and Terricola subgenera of the

Microtus species almost simultaneously appeared in Eastern Europesince the Early Pleistocene (MQR 7 zone) at the Matuyama-Brunhesboundary. They are represented by Microtus nivalinus Hinton, 1923and Microtus arvalidens, which commonly existed, and almostsimultaneously transformed into recent species Microtus arvalisPallas, 1778 and Microtus subterraneus Selys-Longchamps, 1836during the final developmental stages of Singilian faunas (Hol-stein). Europe was characterized by a group of species of the sub-genus Terricola, including taxa Meridiopitymys and Parapitymys,which are morphologically similar to the North American genusPitymys, but phylogenetically distant. M. arvalidens was commonand even dominant in the Likhvin fauna. Later it could transform/evolve into M. subterraneus, but this requires additional evidence(Rekovets, 1994).

Referred material e 10 m1, 4 M3.Measurements e m1: L ¼ 2.4e2.63e2.8 mm, W ¼ 0.8e0.87e

1.0 mm, A ¼ 1.3e1.55e2.0 mm, A/L ¼ 50.9e54.0e55.7%; M3:L ¼ 1.6e2.2 mm, W ¼ 0.7e0.9 mm.

the Middle Pleistocene locality Lysa Gora 1 in Ukraine, Quaternary

L. Rekovets et al. / Quaternary International xxx (2013) 1e11 9

Description and comparison e Relatively small-sized species.Its size is close to that from Bol’shevik 2, I e the average length is2.73 mm, index e 51.1% (Rekovets, 1994). This species in compari-son with other relevant populations (Tikhonovka 1, Morozovka 2,Ozernoje 2, and Medzhybozh) has significantly higher values of theabove metric parameters. Conids in the anteroconid bottom (T4-5)on all samples are commonly fused (C/W index is on average22.0%). The unpaired AC2 loop is also commonly fused with tri-angles at its base (T6-7), the merger (d/e) index is 30.5%. The spe-cies from Lysa Gora in values of indices is closest to the populationsfrom Bol’shevik 2, I and Tikhonovka 1 (Rekovets, 1994), whichcorresponds to the MQR5-4 zone.

Morphological typology of the m1 corresponds with the previ-ously detected morphotypes (Rekovets, 1994, p. 270, Fig. 65, p. 287,Fig. 68), which are distributed as follows: campestris e 12.5%,maskiie 25.0%, dehmie 12.5%, assimilise 37.5%, depressae 12.5%. Themostprimitive “continua” morphotype is missing. The “assimilis” (whichis relatively well represented in recentM. subterraneus) morphotypedominates. Insufficient amount of material does not allow identifi-cation of the main trends in the m1 transformation in the Lysa Gorapopulation. Themajor temporal m1 direction among the extinct andrecent forms of the subgenus is the “maskii” direction, especially inrecent dominant species (Rekovets and Nadachowski, 1995).

Four M3, according to the quantitative ratio of remnants m1 inM. arvalidens (10 m1) andM. nivaloides F. Major, 1902e1 m1, belongtoM. arvalidens. These teeth are noticeably elongated (talonid part).All teeth can be attributed only to the complex (¼variabilis) mor-photype with complicated back structure of the tooth e RS 1. Moreprimitive M3 morphotypes are characteristic for the Microtusnivaloides-arvalis group (Rekovets, 1994, p. 290, Fig. 70). Compli-cated M3 structure and a number of other features question thetransformation along a straight line between M. arvalidens andM. subterraneus.

Microtus (Microtus) nivaloides F. Major, 1902 (¼M. arvalinusHinton, 1923; Rekovets and Nadachowski, 1995)

Fig. 8: 9Referred material e 2 m1.Measurements e L ¼ 2.5 mm, W ¼ 0.9 mm, A ¼ 1.45 mm, A/

L ¼ 58.0%.Description and comparison e m1 has a typical morphology

for the form of that time, well-developed triangle T6-7 in thebottom AC2, which is slightly extended and merged with T6-7conids. The tooth can be assigned to depressa morphotype(Rekovets, 1994, page 287, Fig. 68). Incoming conid corners have ahuge quantity of cement. The species is most similar to those fromSemibalka 1 locality (MQR 5) by the m1 size. Other populationshave considerably larger m1 dimensions.

Microtus sp. (cf. Chionomys ex gr. nivalis)Fig. 8: 10Referred material e One fragment m1 with anteroconid.Description and comparison e In general, the occlusal tooth

surface with symmetrical AC2 mushroom shape and pitymys-likeconid merge at its base resembles those in the Chionomys nivalisMartins, 1842. Enamel is poorly differentiated, and there is a sig-nificant amount of cement in the incoming conid angles. Unfortu-nately, a precise taxonomic attribution is impossible.

5. Results and discussion

5.1. Bony fishes

Most identified fish species are common in the present Dnieperriver basin. However, some are less numerous (Leuciscus leuciscus,

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C. nasus, C. carpio) or, generally, did not exist in this river (i.e., R.frisii).

On the basis of taxonomic diversity, taking into accountecological preferences of particular freshwater fish species in-dicates the palaeoecological characteristics. Three ecotypes weredistinguished for Lysa Gora 1: (1) places with standing to slowlyflowing warmwater, silty bottom and low dissolved O2 content; (2)places with moderate flow, sandy or gravelly bottom and coldwater with middle O2 content; and (3) places with stream flowingcoldwater, gravelly bottom and good aeration. All identified speciesindicate well-developed water vegetation. Complete orictocoe-noses, represented by herbivorous, carnivorous, malacophages,omnivorous forms, are available in the materials of Lysa Gora 1. Themajority of identified species are cold-tolerant, indicating consid-erable temperature fluctuations during the Pleistocene.

5.2. Lagomorphs

The taxonomy of the Pleistocene Ochotona from Central andEastern Europe is still poorly understood. The majority of EarlyPleistocene record of that area is almost exclusively limited tosmall-medium sized ochotonids with large anteroconid and narrowanteroconideposteroconid junction in p3 (cf. �Cermák, 2009). Onthe other hand, the European fossil record of O. pusilla, a specieswith small anteroconid and narrow anteroconideposteroconidjunction in p3, is almost exclusively restricted to the Late Pleisto-cene (Late Toringian). Pre-Toringian findings of p3s with morpho-logical affinities to O. pusilla are rare, limited to a few localities fromthe Czech Republic and Hungary (�Cermák, unpubl. data). TheMiddle Pleistocene ochotonids of Ukraine are still poorly docu-mented. The Ukrainian record of Ochotona, close in age to Lysa Gora1, were reported from e.g., Nagornoje 1, Bol’shevik 2, Protopopovka,Kolkotova balka, Shirokino 1, Platovo, Utkonosovka, Tikhonovka 1,Krasnoselka 1, and Sinyakovo 1 (Krokhmal and Rekovets, 2010), butmostly only in faunal lists without detailed descriptions.

The ochotonid material from Lysa Gora 1 does not allow eval-uation of the actual meaning and/or validity of above-mentionedcharacters. Without knowledge of morphotypes proportion in amore significant sample of ochotonids from the locality, it seemsquite premature to provide a more detailed taxonomic attribution.Another complication is the difficult geological setting in the lo-cality. Taking into account the very low abundance and fragmen-tariness of ochotonid material in Lysa Gora 1, in the context of veryhigh abundance of similar ochotonids in neighbouring older lo-calities (cf. Rekovets and Pashkov, 2009; �Cermák, 2010), we cannotexclude the possible presence of re-deposited older ochotonid el-ements in this locality.

At any rate, considering e (1) the morphodimensional featuresfrom the material under studying, (2) the accompanying faunalassemblage of this ochotonid material, (3) possible presence of re-deposited older elements in the locality, (4) the poorly understoodand documented record of the Early Pleistocene (VillányianeEarlyToringian) Ochotona in Central and Eastern Europe, and (5) a rela-tively high phenotypic diversification of discussed ochotonid taxa,two distinct phenotypic entities of Ochotona can be distinguished inthe very fragmentary material from Lysa Gora 1.

5.3. Rodents

Rodents dominate the total quantity of remnants in the locality.M. arvalidens is the dominant species. Qualitative faunal composi-tion suggests a temperate climate with seasonal cooling. Opensteppe-type spaces, inhabited by Spermophilus, Lagurus, and Eola-gurus, were the dominant habitats. Streamside mesophilic habitats

the Middle Pleistocene locality Lysa Gora 1 in Ukraine, Quaternary

L. Rekovets et al. / Quaternary International xxx (2013) 1e1110

were inhabited by species of Trogontherium, Arvicola, and Microtus.The presence of forest habitats was not confirmed.

The faunal association of the Lysa Gora 1 locality strati-graphically corresponds with the final phase of the TiraspolianFaunal complex, with L. transiens and first M. gregalis and A. mos-bachensis. The Morozovka (¼Nagornoe) Faunal association, begin-ning of Middle Pleistocene, corresponds to the Oka stratigraphichorizon scale (Suvorovsky subcomplex), MQR 4. Faunal assem-blages of Posevkino, Perevoz, Volniye Vershiny on the Russian Plainare stratigraphically closest to the fauna of Lysa Gora 1. In WesternEurope, the faunas of MIS 13-12 are equivalent to Cromerian IV andElster, probably somewhat later than faunas fromMauer, Tark}o, andMosbach 2 (Maul and Markova, 2007).

6. Conclusions

The localities of Lysa Gora (situated near Vasilyevka town,Zaporozhye region, Ukraine) were discovered during an expeditionof the Paleontological Museum (National Museum of Natural His-tory NAS of Ukraine, Kiev) in 2009. Two fossiliferous horizons ofdifferent age were distinguished: Lysa Gora 1 (Middle Pleistocene)and Lysa Gora 2 (Late Miocene). In this paper, the fossil material(excavated during 2009e2012) from Lysa Gora 1 including at least11 freshwater fish taxa and 13 species/forms of small mammals(Lagomorpha, Rodentia) was described and analyzed in detail.

The following conclusions can be inferred and briefly summa-rized from the above surveyed facts:

(1) The species composition of Arvicolidae, particularly thepresence of L. transiens, E. luteus, A. mosbachensis, M. (Terri-cola) arvalidens, M. (Stenocranius) gregalis, as well as thestructure of the fauna, suggests the age of the Middle Pleis-tocene (ca. MQR 4, MIS 13-12, Early Toringian).

(2) Lagomorphs are surprisingly rare in the locality. Only 10fragmentary specimens belonging to two forms/taxa ofochotonids were found: (a) the medium sized form referredhere tentatively as Ochotona sp. (cf. gr. ? pusilla) withmorphological similarities to O. pusilla or possibly to somepaleopopulations known from Transcaucasia (cf. �Cermáket al., 2006; �Cermák, 2009); and (b) the medium sizedform referred here as Ochotona sp. with morphodimensionalaffinities to some Pleistocene populations of Central Europeclose to O. polonica (cf. �Cermák, 2009).

(3) From the paleoecological point of view, themammalian fossilrecord in Lysa Gora 1 suggests a presence of open steppe-type spaces (cf. Spermophilus, Lagurus, Eolagurus, or Ocho-tona) or streamside mesophilic habitats (cf. Trogontherium,Arvicola orMicrotus). The assemblage of fish species suggestsa presence of highly diversified freshwater ecotypes, fromstanding to steam flowing water (silty to gravelly bottom)with variable aeration. The situation corresponds with pre-sent view of paleoenvironmental conditions during theMiddle Pleistocene of the area.

Acknowledgements

This study was partly supported by the Academic Project of theInstitute of Geology AS CR, v.v.i. AV0Z30130516 and by theRVO67985831 of the Institute of Geology AS CR, v.v.i.

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