New Early and Middle Pleistocene of Northern Eurasia · 2019. 3. 13. · Encyclopedia of Quaternary...

This article was originally published in the Encyclopedia of Quaternary Science published by Elsevier, and the attached copy is provided by Elsevier for the author's benefit and for the benefit of

the author's institution, for non-commercial research and educational use including without limitation use in instruction at your institution, sending it to specific colleagues who you know, and

providing a copy to your institution’s administrator.

All other uses, reproduction and distribution, including without limitation commercial reprints, selling or licensing copies or access, or posting on open internet sites, your personal or institution’s

website or repository, are prohibited. For exceptions, permission may be sought for such use through Elsevier's permissions site at:

http://www.elsevier.com/locate/permissionusematerial Vislobokova I., and Tesakov A. (2013) Early and Middle Pleistocene of Northern Eurasia. In: Elias

S.A. (ed.) The Encyclopedia of Quaternary Science, vol. 4, pp. 605-614. Amsterdam: Elsevier.

© 2013 Elsevier Inc. All rights reserved.

Author's personal copy

Early and Middle Pleistocene of Northern EurasiaI Vislobokova, Paleontological Institute of the Russian Academy of Sciences, Moscow, RussiaA Tesakov, Geological Institute of the Russian Academy of Sciences, Moscow, Russia

ã 2013 Elsevier B.V. All rights reserved.

Introduction

The best represented vertebrate records in the early and middle

Pleistocene of Northern Eurasia are mammalian and they are

summarized in this article. The data from Russia and neigh-

boring countries of the former USSR are very important for the

understanding of the history of Late Cenozoic biota. The terri-

tory was a prime area of origin and evolution of taxa and of

dispersals of some mammals to Western Europe, Africa, North

America and China, and vice versa. The configuration of Asia

was close to that of the present day but differed through the

periodic existence of the Bering land bridge, connecting Siberia

and Alaska. The reorganization of the biota went through

climatic and environmental changes; its main trends and

phases are very pronounced there.

The key stages in the history of Pleistocene mammals in this

territory were first recorded by Gromov (1948) who defined

the faunal assemblages for Eastern Europe and named them by

their type localities or regions. Regional complexes were later

defined in the Caucasus, Siberia, Transbaikalia, and Central

Asia. The data on the ages of the assemblages and their com-

positions were considerably enriched thanks to many subse-

quent researchers and new discoveries. In addition to large

mammals, a detailed biochronology based on small mammals

has appeared in the last decades. At present, early and middle

Pleistocene mammal remains are known from many localities

of Northern Eurasia. The age of the localities has been estab-

lished by a combination of geological and paleontological

methods (including the first appearance of index taxa, as well

as the evolutionary levels of dominant forms and of some

phyletic lineages), paleoclimatic data and correlation. Paleo-

magnetic data also help to distinguish more precisely the age of

early Pleistocene sites, whereas the glacial–interglacial succes-

sion provides further important markers for age determination.

The beginning of the Pleistocene is currently set at �2.6 Ma(the Gauss–Matuyama magnetic reversal), the early–middle

Pleistocene boundary at �0.78 Ma (the Brunhes–Matuyamamagnetic reversal), and the middle–late Pleistocene boundary

at 0.126 Ma, according to the International Commission on

Stratigraphy in 2009. However, the Geological Surveys of

Russia and Ukraine have until recently retained the Pliocene–

Pleistocene boundary in its former position at 1.8 Ma. In 2012

the Russian Stratigraphic Committee voted to lower the base of

the Quaternary to 2.58 Ma. The position of the lower boundary

of the Pleistocene and its divisions have thus been repeatedly

changed over the last decades and it must be appreciated that

the published names of particular stratigraphical units often

refer to different time slices.

The early and middle Pleistocene in Northern Eurasia

currently include the following mammal assemblages:

Khaprovian, 2.6–2.2 Ma; Psekupsian, 2.2–1.2 Ma; Tamanian,

1.2–0.8 Ma; Tiraspolian, 0.8–0.4 Ma; Singilian and Khasarian,

0.4–0.126 Ma.

Encyclopedia of Quaternary Scien

Mammal Assemblages of Northern Eurasia

Early Pleistocene: Gelasian, Middle Villafranchian, LateVillanyian, MN17

The Khaprovian (Khapry) mammal assemblage and itsanalogsThis stage is well documented in the northern Black Sea region,

peri-Azov area, Siberia, and Central Asia. The mammal com-

munities superficially resembled those of modern Africa or

southern Asia by the presence of proboscideans, equids, rhi-

noceroses, giraffids, and various antelopes, but differed

considerably from them in species composition. These assem-

blages are characterized by the first appearance and subseq-

uent wide distribution of the elephant Archidiskodon gromovi

(¼ A. meridionalis gromovi in Titov, 2008, ¼ Mammuthus gro-movi of Lister and Sher, 2001). In contrast to Russian paleon-

tologists, most researchers refer this species to the genus

Mammuthus, mainly based on the tooth structure (Lister and

Sher, 2001), although Archidiskodon is retained here. Other

features of the fauna include the presence of the bear Ursus

etruscus and the rhinoceros Stephanorhinus sp., the rapid

diversification and wide distribution of stenonid horses,

the presence of diverse canids (Eucyon, Nyctereutes, Canis), two

saber-toothed cats Homotherium crenatidens and Megantereon

cultridens, the camel Paracamelus, the large-sized comb-antlered

deer Eucladoceros and the elk (¼ moose) Libralces, and otherextinct forms. In Northern Eurasia, there were three paleozoo-

geographic subareas: European-Siberian, Mediterranean, and

Central Asian (Vangengeim and Pevzner, 1991; Vislobokova

et al., 1995) (Figure 1). The global cooling witnessed at about

2.6 Ma (indicated in the territory by the glaciations of the

Caucasus and Pamir, the most ancient loess formation in

Tadjikistan, Uzbekistan, etc.) caused a decrease in humidity

in temperate latitudes and in Central Asia (Dodonov, 2002),

where animals adapted to open woodlands and grasslands

became dominant. Some taxa have North American ancestry,

immigrating during periods of low sea level prior to the start

of the Pleistocene, such as the camels, Hipparion and Eucyon

in the late Miocene, and Equus, Canis, and Megantereon in the

Pliocene (Vislobokova et al., 2003). A number of thermophilous

herbivores (mastodons, giraffids, and others), typical of Pliocene

times, were gradually replaced by forms adapted to cooler cli-

matic conditions and possessing more advanced herbivorous

adaptations. The data on herbivorous taxa (elephants, horses,

ruminant Artiodactyla, voles, etc.) are of great value for biochro-

nology and paleoenvironmental reconstruction because their

tooth structure yields clearly recognizable signals inwear patterns,

hypsodonty, etc. The increase in diversity of herbivore taxa was

accompanied by considerable changes in the carnivore guild.

At this time, an extensive European-Siberian part of the

Palearctic was inhabited by the elephant Archidiskodon gromovi,

the rhinoceros Elasmotherium, and large Equus livenzovensis

and small Equus sp., together with other animals adapted to

605

ce, (2013), vol. 4, pp. 605-614

60� 70� 80� 80� 70� 60�

50�

40�

30�

130�120�110�100�90�80�

1 2 3

70�60�50�

0 1250 2500 km

30�

40�

50�

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

606 VERTEBRATE RECORDS | Early and Middle Pleistocene of Northern Eurasia


savanna-like conditions (the wolves of the Canis etruscus group,

the hyena Pliocrocuta, the saber-toothed cat Homotherium cre-

natidens, the cheetah Acinonyx, the camel Paracamelus, the an-

telope Gazellospira, the bison-like Leptobos, and others), along

with forest and ecotonal animals (the bear Ursus etruscus and

the deer Eucladoceros and Libralces). Most of these mammals

were found in common with those from the Middle Villafran-

chian faunas of Western Europe (Kahlke et al., 2011; Palombo

et al., 2006). Early Villafranchian elements (the mastodon

Anancus arvernensis, the archaic horse Hipparion, and giraffids)

became very rare. Among the small mammals, the bank voles

Clethrionomys made their first appearance, the water vole Mim-

omys polonicus was replaced by M. pliocaenicus in the Northern

Black Sea area, and the pikas Pliolagomys and Ochotona and

jerboas Allactaga, more typical of Asian faunas, were present.

Fossil remains of Archidiskodon gromovi, the main indicator

species of the Khaprovian mammal assemblage, have been

found in a number of localities in the European part of

Russia (Khapry, Liventzovka, and others), in the Ukraine

(Zhevakhova Gora), in the south of Western Siberia (Podpusk-

Lebyazh’e), and inCentral Asia (Adyrgan, Kuruksay) (Bajgusheva,

1971; Garutt and Bajgusheva, 1981; Vangengeim et al.,

1988; Vislobokova, 1996, 2005; and others). A. gromovi

has also been found in Italy (Montopoli) (Gliozzi et al.,

1997).

Remains of other members of the Khaprovian assemblage

are recorded from many European–Russian sites along the

northern coast of the Sea of Azov and the right bank of the

Encyclopedia of Quaternary Scienc

Don River near Rostov-on-Don and Taganrog. They come from

the lower part of the Khapry alluvium recorded in sand pits

and natural outcrops. The large mammals were represented by

Anancus arvernensis alexeevae, the horses Hipparion moritorum,

Equus (Allohippus) livenzovensis, and a small Equus sp., the rhi-

noceroses Stephanorhinus ex gr. megarhinus-kirchbergensis and

Elasmotherium chaprovicum (¼ E. cf. caucasicum), the wild boarSus strozzi, the camels Paracamelus alutensis and Paracamelus cf.

gigas, the deer Cervus (Rusa) philisi, Eucladoceros cf. dicranios,

Arvernoceros sp., and Libralces gallicus, the giraffid Palaeotragus

(Yuorlovia) priasovicus, and the bovids Leptobos sp., Gazellospira

gromovae, Tragelaphini gen. indet., Gazella cf. subgutturosa, and

others (Alexeeva, 1977; Bajgusheva, 1971; Bajgusheva et al.,

2001; Gromov, 1948; Titov, 2008; and others) (Figure 2). The

Khapry carnivores included the raccoon-dog Nyctereutes mega-

mastoides, Canis sp. (C. cf. senezensis : Sotnikova et al., 2002),

the mustelids Lutra sp. and Pannonictis nesti, the medium-sized

hyena Pliocrocuta perrieri and the dominant larger-sized

short-faced hyena Pachycrocuta brevirostris, the felids Lynx issio-

dorensis, the saber-toothed catHomotherium crenatidens, and the

cheetah Acinonys pardinensis. Most of these animals were in-

habitants of open savannah-like landscapes, with a part of the

fauna adapted more to forested or ecotonal zones (mustelids,

lynx, wild boar, and deer). Among the small mammals, the

archaic lagurine Borsodia praehungarica, the water volesMimomys

praepliocaenicus and Mimomys ex gr. reidi, and the primitive red-

backed voles Clethrionomys kretzoii were also present (Tesakov,

2004).

e, (2013), vol. 4, pp. 605-614

Figure 2 Anancus arvernensis, C. Flerov painting exhibited in thePaleontological Museum of the Paleontological Institute, RussianAcademy of Sciences (PIN) in Moscow, with PIN permission.

Figure 3 Podpusk–Lebyazh’e locality at the right bank of the IrtyshRiver, Western Siberia.

VERTEBRATE RECORDS | Early and Middle Pleistocene of Northern Eurasia 607


In the south of Western Siberia, faunal assemblages

described by Vangengeim and Zazhigin have been reported

from Podpusk-Lebyazh’e, from the lower part of the thick

sandy alluvial deposits exposed on the right bank of

the Irtysh River, south of Pavlodar (Vangengeim, 1977;

Vislobokova, 1996; Figure 3). Typical members include A.

gromovi, S. cf. etruscus, Elasmotherium sp., Equus livenzovensis,

Equus sp. (small), Paracamelus cf. gigas, Eucladoceros sp., Lepto-

bos, and the antelopes Gazella cf. sinensis and Antilospira

cf. gracilis (Vislobokova, 1996). The faunal assemblage con-

sisted mainly of forms widespread in the temperate zone of the

Palearctic region (Ursus cf. etruscus, Homotherium, Pachycrocuta,

and others), with some Central Asian elements (Ochotonoides,

Elasmotherium, Paracamelus, Antilospira cf. gracilis, Gazella

cf. sinensis), some of which (e.g., Paracamelus) reached Eastern

Europe. Among the small mammals, voles with rooted denti-

tion of the genus Mimomys (M. pliocaenicus and M. reidi) and

Borsodia (B. praehungarica-petenyii) were predominant and

Clethrionomys appeared for the first time in Siberia (Zazhigin,

1980). The mammalian assemblages indicate the prevalence

of steppes with woodland developed along river valleys.


In northeastern Asia, the association of Synaptomys-like

lemmings Plioctomys and mimomyian voles Cromeromys ex gr.

irtyshensis-hordijki and smaller Mimomys sp. were found in the

lower member of the Kutuyakh Formation at the Krestovka

River in the Kolyma Lowland (Sher, 1987; Tesakov and van

Kolfschoten, 2011). The assemblage is dominated by lem-

mings and ‘Cromeromys’ voles, with Ochotona sp. and Mimomys

sp. being less abundant.

Several faunas of this age have been referred to the Central

Asian subarea (Erbaeva and Alexeeva, 2000; Sotnikova et al.,

1997; Vangengeim, 1977; Vislobokova et al., 1995). These

include the Itanza fauna of Klochnevo I, II (Transbaikalia)

with Ochotona intermedia, the ground squirrel Spermophilus

itancinicus, Allactaga sp., the hamster Cricetinus varians, the

voles Mimomys pseudintermedius, Clethrionomys sp., and

‘Villanyia’ klochnevi, the archaic mole rat Prosiphneus cf.

paratingi, the first representative of the giant deer genus

Praemegaceros and others, as well as the Kiikbai and Andyrgan

faunas (southern Kazakhstan) with Ochotonoides complicidens,

Mimomys pliocaenicus, the gerbil Meriones cf. meridianus,

Equus stenonis, the camel Gigantocamelus longipes, and other

forms.

The fauna of Southern Tadzhikistan has affinities with the

Eastern Mediterranean subarea and is characterized by a com-

bination of Mediterranean and Central Asian forms. The rich

assemblage from Kuruksay contained, along with A. gromovi

and Mastodontoidea fam. et gen. indet., the baboon Papio

sushkini (¼ Paradolichopitecus sushkini), the porcupine Hystrixsp., diverse carnivores including Nyctereutes megamastoides,

eucyon ‘Canis’ kuruksaensis, Ursus cf. etruscus, the cursorial

hyena Chasmaporthetes lunensis kani, Pliocrocuta perrieri, Lynx ex

gr. issidorensis, Acinonys cf. pardinensis,Megantereon megantereon,

and Homotherium crenatidens, the rhinoceros Stephanorhinus

sp., the horse Equus stenonis pamirensis (¼ E. s. bactrianus),the camel Paracamelus praebactrianus, the deer Axis flerovi

and Elaphurus eleonorae, the first representative of the giant

deer genus Sinomegaceros (S. tadzhikistanis), the elk Libralces

cf. gallicus, the giraffids Sogdianotherium kuruksaense and

Sivatherium, and the antelopes Gazella parasinensis, Protoryx

paralaticeps, Damalops palaeindicus, Gazellospira gromovae, and

Antilospira sp., amongst others (Dmitrieva, 1977; Vangengeim

et al., 1988). In this region, a large number of Asian elements

occurred. Animals of dry, open landscapes (Ellobius, Pliocrocuta,

Chasmaporthetes, Paracamelus, Equus) were accompanied by

forest animals (bear, lynx, monkey, the deer Axis and Libralces)

and savannah inhabitants (Sivatherium, Protoryx, Damalops,

Gazellospira). Such a mixed composition reflected the mosaic

landscapes and vertical zonality typical of mountainous

regions. In Tadzhikistan, the local faunas attributed to this

assemblage are known also from Obigarm, Karamaidan,

Tutak, and Zil’fi localities, while faunas close to them in age

were found in Kyrgyzstan (Akterek, Dzylgyndykoo) (Dmitrieva

and Nesmeyanov, 1982; Sotnikova et al., 1997).

The reversed polarity of the fossiliferous deposits in Khapry

indicates that these sediments should be attributed to the lower

part of the Matuyama Chron, together with analogous sites

in Siberia and Central Asia (Podpusk-Lebyazh’e, Krestovka,

Kuruksay) (Sotnikova et al., 1997; Tesakov et al., 2007;

Vangengeim and Pevzner, 1991; Vangengeim et al., 1988;

Vislobokova, 1996).

ce, (2013), vol. 4, pp. 605-614



Late Early Pleistocene: Late Villafranchian, MN18–MQ19, ¼Tiglian–Menapian

Psekups mammal assemblage and its analogsThis stage is characterized by significant changes in the diver-

sity of mammal communities (Figure 4). In Northern Eurasia,

a cold-climate event around 2.2–1.9 Ma resulted in the appear-

ance of new boreal elements, including some modern genera.

In the European-Siberian subarea, Archidiskodon gromovi was

replaced by the more progressive A. meridionalis (¼ Mam-muthus meridionalis), close to the type of the species from

the Upper Valdarno, Italy. Voles of the genus Allophaiomys,

the giant deer Praemegaceros, the elk Alces, the large bovids

of the genus Bos and the Bison (Eobison) – B. (Bison) phyletic

lineage, and the musk-ox Soergelia first appeared in this stage.

Ma

0.5

1.0

1.5

2.0

2.5

Reu

nion

Old

uvai

Mat

uyam

a

Ple

isto

cene

Gel

asia

n

Kha

pro

vian

Pod

pus

k-Le

bya

zhia

n

Itanz

inia

n

Kre

stov

kian

faun

a

Mim

omys

plio

caen

icus

Mim

omys

sav

ini

Arv

icol

a

Mim

omys

(Tch

aryn

omys

) sp

p.

Vily

uisk

ian

faun

aO

lyor

ian

Dod

ogol

ian

faun

aZ

asuk

hino

faun

aTo

logo

ian

Ivol

gini

an-

Ust

’kire

nski

an

Kiz

ikhi

anR

azd

olia

nV

yatk

inia

n

Pse

kup

sian

Tam

ania

nTi

rasp

olia

nS

ingi

lian-

Cha

saria

n

Mid

dle

Late

Ear

ly

20

19

18

17

Mid

dle

–lat

eV

illaf

ranc

hian

Gal

eria

n

Ear

lyM

idd

le

Bru

nhes

Jara

mill

o

Ep

och

Age

Mam

mal

age

Mammalassemblage

Mam

mal

zon

e

Eas

tern

Eur

ope

Wes

tern

Sib

eria

Eas

tern

Sib

eria

Tran

sbai

kalia

Mag

netic

pol

arity

tim

esc

ale

(Can

de

and

Ken

t, 1

995)

Figure 4 Mammal assemblages and faunas of Northern Eurasia and the ma


In Psekups (Northern Ciscaucasia), Archidiskodon meridio-

nalis, Eucladoceros orientalis (¼ Psekupsoceros orientalis, first rec-ognized as Cervus pliotarandoides), Stephanorhinus etruscus, and

Equus stenonis were present (Alexeeva, 1977; Gromov, 1948;

Vislobokova, 1990; and others). The small mammals included

Borsodia newtoni-arankoides, Pitymimomys pitymyoides, Mimomys

cf. pliocaenicus, Mimomys reidi, and Clethrionomys kretzoii

(Alexandrova, 1976; Tesakov, 2004). These reversed polarity

deposits correspond to the Matuyama Chron, between the

Reunion and Olduvai Subchrons (Tesakov, 2004; Vangengeim

and Pevzner, 1991). Slightly younger assemblages from the

Odessa small mammal complex contain hypsodont Mimomys

forms (dominant M. reidi and M. ex gr. savini), the last repre-

sentatives of Borsodia, the first occurrence of lagurines with

Allo

pha

iom

ys d

euca

lion

A. p

lioca

enic

us

Mic

rotu

s (s

teno

cran

ius)

hin

toni

Bor

sod

ia s

pp

.P

rola

guru

s te

rnop

olita

nus

Lagu

rod

on a

rank

ae

Eol

agur

us a

rgyr

opul

oi

Can

is e

trus

cus

Urs

us e

trus

cus

Hom

othe

rium

cre

natid

ens

U. d

enin

geri H

omot

heriu

m s

p.

Gul

o cf

. sch

loss

eri

G. g

ulo

U. s

pel

aeus

C. m

osb

ache

nsis

–C. v

aria

bili

s

Pal

aeol

oxod

on s

p.

P. a

ntiq

uus

C. l

upus

Arc

hid

isko

don

grom

ovi

Arc

hid

isko

don

mer

idio

nalis

mer

idio

nalis

Arc

hid

isko

don

m. t

aman

ensi

sM

amm

uthu

str

ogon

ther

iiM

amm

uthu

sch

osar

icus

E. l

uteu

s

P. p

anno

nicu

sLa

guru

s tr

ansi

ens

L. la

guru

s

M. g

rega

loid

esM

. gre

galis

Evolutionary lineage

in lineages.

e, (2013), vol. 4, pp. 605-614

Ma

Ep

och

Age

Mam

mal

age

Mammalassemblage

Evolutionary lineage

Mam

mal

zon

e

Eas

tern

Eur

ope

Wes

tern

Sib

eria

Eas

tern

Sib

eria

Tran

sbai

kalia

Mag

netic

pol

arity

tim

esc

ale

(Can

de

and

Ken

t, 1

995)

Reu

nion

Old

uvai

Mat

uyam

aB

runh

es

0.5

1.0

1.5

2.0

2.5

Jara

mill

o

Ple

isto

cene

Gel

asia

nE

arly

Mid

dle

Mid

dle

Late

Ear

lyM

idd

le–l

ate

Vill

afra

nchi

anG

aler

ian

20

19

18

17

Kha

pro

vian

Pse

kup

sian

Tam

ania

nTi

rasp

olia

nS

ingi

lian-

Cha

saria

n

Pod

pus

k-Le

bya

zhia

nK

izik

hian

Raz

dol

ian

Vya

tkin

ian

Itanz

inia

nD

odog

olia

nfa

una

Zas

ukhi

nofa

una

Tolo

goia

nIv

olgi

nian

-U

st’k

irens

kian

Kre

stov

kian

faun

aV

ilyui

skia

n fa

una

Oly

oria

n

Ste

pha

norh

inus

etr

uscu

sS

. kirc

hber

gens

is

Eq

uus

liven

zove

nsis

Eq

uus

sues

senb

orne

nsis

Eq

uus

mos

bac

hens

is

Eq

uus

sten

onis

Par

acam

elus

gig

as

E. c

abal

lus

P. a

lute

nsis

Cam

elus

Ran

gife

r

Cer

vus

abes

alom

i

Arv

eroc

eros

sp

.M

egal

ocer

os s

p.

Sin

omeg

acer

os t

adzh

ikis

tani

s

Lib

ralc

es g

allic

us

Lep

tob

os s

p.

Bis

on s

ucho

vi

Ovi

bov

ini

Pra

eovi

bos

Ovi

bos

Bis

on s

choe

tens

acki

B. p

riscu

s lo

ngic

orni

s

Alc

es s

p.

Alc

es la

tifro

ns

Sin

omeg

acer

os s

p.

M. g

igan

teus

C. a

coro

natu

sC

. ela

phu

s

Coe

lod

onta

tolo

goije

nis

Coe

lod

onta

antiq

uita

tis

Figure 4 (Continued)



unrooted teeth Prolagurus and Lagurodon, and Allophaiomys deu-

calion (Alexandrova, 1967; Rekovets and Nadachowski, 1995;

Tesakov, 2004). The Odessa assemblage is subdivided into two

phases: (1) an early phase, with the remains of rootless voles

Allophaiomys deucalion and (2) a late phase, with the first cemen-

ted rootless voles of Prolagurus (P. ternopolitanus) and Lagurodon

(L. arankae) (Markova, 1982, 2007).

In the Caucasus, the Dmanisi fauna is famous for the pres-

ence of the first appearance in Eurasia of Homo, following

dispersal from Africa. The remains of the Dmanisi hominins

are attributed to the Homo erectus group and were found in the

layers directly overlying a lava flow with a radiometric date of

1.8�01 Ma, within the Olduvai Subchron. The fauna com-prises inhabitants of dry, open area (the tortoise Testudo, the

ostrich Struthio, the gerbil Parameriones, the pika Ochotona, the

hamster Cricetulus, A. meridionalis, Canis etruscus, Pliocrocuta


cf. perrieri, Megantereon cultridens, Homotherium crenatidens,

Palaeotragus) and forest animals (Bison (Eobison) georgicus, the

deer Cervus abesalomi and Dama nestii, the bear Ursus etruscus)

(Gabunia and Vekua, 1995; Hemmer et al., 2010; Vekua, 1995;

Vekua and Lordkipanidze, 2008). The first occurrences of the

‘jaguar’ Panthera onca georgica and musk ox Soergelia are

recorded there. The contemporaneous Palan-Tyukan fauna

(Azerbaijan) includes Canis etruscus, Panthera ex gr. onca gom-

baszoegensis, and Nyctereutes megamastoides.

In Western Siberia, A. cf. meridionalis, Equus ex gr. stenonis,

Alces sp., and Gazella sp. along with Borsodia ex gr. petenyii-

hungaricus, andMimomys coelodus are known from the Podpusk

fauna (from the uppermost part of the sands in the Irtysch

River Basin) (Vislobokova, 1996). In southern Kazakhstan, the

Aktogai (Kopaly) locality with Mimomys (Tcharynomys) haplo-

dentatus (bed 3) and Allophaiomys deucalion (bed 7) also

ce, (2013), vol. 4, pp. 605-614

Figure 5 A skeleton of Elasmotherium on display in the PaleontologicalMuseum of the Paleontological Institute, the Russian Academy ofSciences (PIN), Moscow, with PIN permission.



corresponds to this age (Sotnikova et al., 1997; Tjutkova and

Kaipova, 1996).

In the Kuznetsk Depression, Allophaiomys pliocaenicus, Pro-

lagurus ex gr. pannonicus-posterius, Eolagurus argyropuloi, Archi-

diskodon cf. meridionalis, and Ovibovini gen. indet. were found

in the Mokhovo Formation, 1.8–1.2 Ma (Foronova, 2001).

In northeastern Asia, a tooth of Archidiskodon meridionalis

meridionalis was found in the Vilyuisk district, Yakutia; the find

confirmed the spread of the subspecies into North America

over the Bering land bridge (Dubrovo, 1990), probably at the

end of the 2.9–2 Ma global sea level fall.

In Transbaikalia, the Dodogol and Zasuchino II faunas

have been referred to the Central Asian subarea. In Dodogol,

Borsodia laguroformes, Allophaiomys cf. pliocaenicus, Prosiphneus

youngi, the first woolly rhino Coelodonta (of Asian origin),

Equus ex gr. sanmeniensis, and others were present (Vangengeim,

1977). The revised data on the Pleistocene small mammals of

the region are presented in Alexeeva et al. (2007) and Alexeeva

and Erbaeva (2008).

Terminal Early Pleistocene: Early Galerian, 1.2–0.8 Ma

The time range from 1.2 to 1.1 Ma (before the Jaramillo paleo-

magnetic reversal) is an important boundary in the history of

the biota, and is reflected in the nomenclature of stratigraphic

schemes (the Villafranchian–Galerian boundary or the Early–

Late Eopleistocene boundary in the scheme of the Russian

Stratigraphic Committee).

Tamanian mammal assemblage and its analogs(Nogaisky–Morozovka, ¼ Menapian–Bavelian)This stage is characterized by the replacement of Archidiskodon

meridionalis meridionalis by the more advanced A. meridionalis

tamanensis, the first appearance of the giant deer Praemegaceros

verticornis and Megaloceros, the red deer Cervus (Cervus) acoro-

natus, the roe deer Capreolus cf. suessenbornensis, the broad-

fronted elk Alces latifrons, the bison Bison (Bison), camels of

the genus Camel, the first occurrence of the woodland elephant

Palaeoloxodon and woodland antelopes Tragelaphus and

Pontoceros, and the presence of Paracamelus (P. kujalnensis)

(Alexeeva, 1977; Gromov, 1948; Vekua, 1962; Vereshchagin,

1957; and others). From the end of the early to the middle

Pleistocene, true wolves attributed to the Canis mosbachensis–

C. variabilis group and large hunting dogs Lycaon lycaonoides

(¼ Canis (Xenocyon) lycaonoides) existed in Eurasia fromWesternEurope to Transbaikalia andChina (Sotnikova and Rook, 2010).

The Tamanian mammal assemblage is based on the fauna

of Synaya Balka (type locality) and Tsymbal in the Taman

Peninsula. A. meridionalis tamanensis coexisted in the North

Black Sea area with Homotherium crenatidens, the short-faced

hyena Pachycrocuta brevirostris, the horse Equus suessenbornensis,

the rhinoceroses Stephanorhinus etruscus and Elasmotherium

caucasicum, and others (Alexeeva, 1977; Sotnikova and Titov,

2009; Vereshchagin, 1959; Figure 5). Vereshchagin (1957)

also identified the potential impact of early hominins on

some of these assemblages. And recently, stone artifacts of the

Oldowan type were reported from the Synyaya Balka site by

Shchelinsky et al. (2010). In the Asian part of the territory, the

first reliable evidence of human activity comes from the stone

tool assemblage of Kuldara (Ranov et al., 1995) dated to


0.8 Ma and broadly synchronous with stone tools from the

Karama site in the Altai region (Derevyanko and Shunkov,

2005).

Morozovkian small mammal fauna with more derived

voles Microtus (Stenocranius) hintoni and the pitymyoid Micro-

tus sp. are correlated with these assemblages (Alexandrova,

1976; Rekovets and Nadachowski, 1995). The small mammal

fauna of this stage are also represented in the Dnieper, Dnies-

ter, and Don basins (Agadjanian, 2009; Alexandrova, 1976;

Markova, 2007; Tesakov et al., 2007; Topachevsky et al.,

1987; and others). The assemblages include an advanced

meadow vole Allophaiomys (A. pliocaenicus) and the steppe

lemmings Prolagurus pannonicus, Lagurodon arankae, and Eola-

gurus argyropuloi.

At this time, differentiation of environment sharply

increased. In the Caucasus (Akhalkalaki, Georgia), the most

thermophilous elements continued to exist, including Hippopot-

amus georgicus, Equus suessenbornensis, and Stephanorhinus hund-

sheimensis, the cave or ‘spelaeoid’ bear close to U. deningeri, and

the first jaguar (close to P. onca gomboszoegensis) (Hemmer et al.,

2010; Vekua, 1962, 1986). In Siberia, the diversity ofmusk-oxen

increased. The ovibovine Soergelia occurred in the Razdolie as-

semblage (Western Siberia) with A. meridionalis, Palaeoloxodon

sp., Equus (Allohippus) sp., Bison sp., and Ovibovini (Vangen-

geim, 1977). The remains of Soergelia and the oldest members

of Praeovibos were found in the early stage of the Olyorian land

mammal age (Beringia, Bering Province) (1.4–0.8 Ma) together

with those of the first wolverine Gulo minor (¼ G. cf. schlosseri)(Sher, 1971, 1987; Sher et al., 2011; Vangengeim, 1977). These

co-occurred with Archidiskodon sp., Lycaon lycanoides, Equus

(Plesippus) verae, Alces sp., Bison sp., and others. The northern-

most finding of Homotherium in Asia is associated with this

fauna. In Eastern Siberia, the Aldan fauna included a number

of forest inhabitants (Trogontherium cf. cuvieri, Palaeoloxodon ex

gr. namadicus, Alces latifrons, and Microtus gregaloides). In Trans-

baikalia, the Kudun (Kizhing–Kudun depression) and Zasu-

khino (Itanza River basin) correspond to the youngest part of

the early Pleistocene and are correlated with the uppermost

Matuyama Chron (Erbaeva and Alexeeva, 2000).

The oldest Camelus (C. cf. knoblochi) coexisted with the

giant deer Praemegaceros (close to P. verticornis) in southern

e, (2013), vol. 4, pp. 605-614



Tadjikistan (Lakhuti locality, the uppermost part of the

Matuyama) and members of diverse habitats. They included

inhabitants of open, dry, and savannah-like landscapes, such

as the gerbil Meriones lacutensis and mole vole Ellobius lakhu-

tensis, Archidiskodon sp., Equus cf. namadicus, and Canis cf.

mosbachensis, in association with a large Lycaon, Pachycrocuta

brevirostris,Homotherium sp., the red-backed vole Clethrionomys,

Panthera onca gomboszoegensis, the badgerMeles ex gr.meles, and

the giant deer Sinomegaceros sp. (Sotnikova, 1989; Sotnikova

and Vislobokova, 1990). Small mammals also included white-

toothed shrew Crocidura sp., a hamster Cricetulus sp., the

meadow voles Allophaiomys sp., and Microtus (Phaiomys) lachu-

tensis (data from Zazhigin).

Faunas intermediate in age between the Tamanian and

Tiraspolian faunas are known from the lower course of the

Dnester River (Karai-Dubina) and in the middle Don (Petro-

pavlovka) area and have been studied in detail (Agadjanian,

2009; Markova, 2007). The reversed polarity deposits contain

the remains of Prolagurus and pitymyoid Microtus, the last

representatives of Allophaiomys, and the oldest Microtus with

five closed enamel triangles in the first lower molar.

The dispersal of mammals out of Northern Eurasia in-

creased from about 1.2 to 1.1 Ma; Praemegaceros verticornis

and Soergelia invaded Western Europe and Homo erectus and

Sinomegaceros first occurred in China (Kahlke et al., 2011; Qiu,

2006).

60� 70� 80�

90�80�

1

70�60�

0 1250 2500 km

50�

50�

40�

30�

Figure 6 Zonal occurrence of mammals in the early–middle Pliocene, with sand broadleaved forests; (4) Picea-Pinus and Betula forests, with some broad(7) tundra.


Early Middle Pleistocene: Middle–Late Galerian, ¼Cromerian–Elsterian, 0.8–0.4 Ma

The early–middle Pleistocene boundary of the standard

scheme corresponds to the Eopleistocene–Neopleistocene

boundary of the Russian stratigraphic scheme and the Early–

Middle Galerian boundary in Italy.

Tiraspolian mammal assemblages and its analogsThe considerable reorganization in the composition of mam-

malian communities was related to a drop in temperature

at the early–middle Pleistocene boundary, with a notable

increase in amplitude of climatic oscillations. Along with

progressive cooling, climatic fluctuations changed from 41-ka

low-amplitude cycles to a 100-ka high-amplitude cycle

(Shackleton, 1995). In the Russian Plain, three periods of

glaciation correspond to this stage (Pokrovka, Don, and Oka).

The climatic zonation of the northern Eurasian landscape

began to resemble that of the present day (Figure 6). Archidis-

kodon was replaced by the mammoth Mammuthus trogontherii

(¼ A. wüsti) and the proportion of modern vertebrate generasharply increased.

The representatives of the Tiraspolian mammal assemblage

in southeastern Europe existed under a temperate climatic

regime in a variety of landscapes, dominated by forest-steppes

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

80� 70� 60�

50�

40�

30�

130�120�110�100�

2 3 4 5 6 7

ome data from Markova (2006): (1) steppe; (2) forest-steppe; (3) mixedleaved plants; (5) Picea-Pinus forests and steppe; (6) forest–tundra;

ce, (2013), vol. 4, pp. 605-614

Figure 7 Megaloceros giganteus, C. Flerov’s painting exhibited in thePaleontological Museum of the Paleontological Institute, RussianAcademy of Sciences (PIN), Moscow, with PIN permission.



locality of this assemblage. The fossil remains were found in

the so-called ‘Tiraspolian gravels’ (fluvial deposits of the Dnies-

ter 6th terrace). Horses were represented by both caballoid

(Equus cf. mosbachensis) and stenonis (E. aff. suessenbornensis)

forms. Mammuthus trogontherii, Stephanorhinus etruscus (late

form), the large deer Cervus acoronatus, Praemegaceros verticor-

nis, the fallow deer Praedama and Alces latifrons, and a short-

horned bison Bison schoetensacki are common to both the

Tiraspolian fauna and the fauna of Western Europe (Kahlke

et al., 2011; Nikiforova, 1971). Two skeletons ofM. trogontherii

were found in the Kagalnik sand pit (Azov). Data on small

mammal fauna represent several phases of Tiraspolian mam-

mal assemblages (see Markova, 2007). A considerable cooling

has been recognized in the southern part of European Russia at

the time of the early middle Pleistocene Don glaciations,

highlighted by the presence of subarctic species such as the

lemmings Lemmus ex. gr. Sibiricus and Dicrostonyx sp. and the

North Siberian vole Microtus ex gr. hyperboreus (Agadjanian,

2009).

The southward shifting of ranges, with persistence of some

forms in refugia (including mountain ones), was usual during

continental glaciations. Very diverse mammal faunas are

known from the northern Caucasus. Most are associated with

Early Paleolithic sites and stone artifacts of Acheulian type. The

taxa include the macaque monkey Macaca cf. sylvana, the spot-

ted hyena Crocuta spelaea, diverse bears including cave bear

Ursus deningeri (dominant), black bear Ursus ex gr. thibetanus

probably related toU. etruscus, and the small cave bearU. savini

rossicus (¼ U. aff. spelaeus rossicus), the cave lion Panthera spelaeaand leopard P. pardus, the gray wolf Canis lupus, the fox Vulpes

vulpes, Stephanorhinus,Megaloceros giganteus, Capreolus cf. suessen-

bornensis, Bison cf. schoetensacki, together with typical montane

genera such as the ibex Rupicapra and the goat Capra.

The fauna of the late stage of the Olyorian faunal assem-

blage (West Bering Province) are characteristic of tundra and

forest–tundra environments and cold climatic conditions sim-

ilar to modern ones (Sher, 1971, 1987; Vangengeim, 1977).

This fauna included the first reindeer Rangifer. Alces aff. latifrons

and Bison sp. were also present in the artiodactyl community.

Rangifer is also recorded in the early middle Pleistocene of the

Kuznetsk Depression (Foronova, 2001).

In Transbaikalia, the Tologoi assemblage contained the

woolly rhino Coelodonta tologoijensis, Equus ex gr. sanmeniensis,

Cervus ex gr. elaphus, Spirocerus cf. peii, and Bison sp.

(Vangengeim, 1977). Cryogenic disturbance at the base of the

deposits is associated with remains from the Zasuhino assem-

blages (Zasuhino 3, 1–0.78 Ma) and in Tologoi 2 beneath the

Brunhes–Matuyama magnetic reversal, above the Jaramillo

(Alexeeva, 2005). The fauna is characterized by a large diversity

of carnivores and ungulates: Nyctereutes sp., Canis variabilis,

Lycon cf. lycaonoides (¼ Xenocyon lycaonoides), Ursus sp., Gulosp., Pachycrocuta brevirostris, Homotherium sp., the large-sized

tiger Panthera ex gr. tigris, Coelodonta tologojensis, Equus ex gr.

sanmeniensis, Equus ex gr. suessenbornensis–verae, Alces cf.

latifrons, and others (Vangengeim et al., 1990). Zasuhino 3

locality also has yielded Cervus sp., Megacerini indet, Capreolus

cf. suessenbornensis, Ovibovini indet., and Bison sp.

At the boundary of the early and middle Pleistocene, a

prominent climatic deterioration was marked over northern

and central Eurasia and many immigrants from these regions


are recorded in Western Europe. The Bering land bridge (0.8–

0.011 Ma) further facilitated faunal exchanges between North-

ern Asia and North America.

Late Middle Pleistocene, ¼ Aurelian, ¼ Holsteinian–Saalian,0.4–0.126 Ma

Singilian and Khasarian faunas and their analogsThe Singilian fauna and its analogs are related to the Likhvin

interstadial (¼ Mindel-Riss, ¼ Holsteinian), one of the mostpronounced warm stages in the middle Pleistocene. In Eastern

Europe, the Singilian faunas were represented by the straight-

tusked elephant Palaeoloxodon antiquus and Merck’s rhinoceros

Stephanorhinus kirchbergensis. In the south of Northern Eurasia,

gray wolf (Canis lupus), Camelus knoblochi, Megaloceros gigan-

teus, and Bison priscus longicornis became widespread after

the mid-middle Pleistocene (Alexeeva, 1977; Gromov, 1948).

Typical small mammals include the water vole Arvicola mosba-

chensis, the yellow steppe lemming Eolagurus luteus, the narrow-

skulled vole Microtus gregalis, the northern or root vole

M. oeconomus, Ellobius, and the ground squirrel Spermophilus

(Markova, 2007). In Western Siberia, Palaeoloxodon coexisted

with Equus cf. steinheimensis, a small-sized cave bearUrsus savini

rossicus (¼ Ursus spelaeus rossicus), and the giant deer Megalo-ceros giganteus during the contemporaneous Tobol interstadial

(Vangengeim, 1977; Figure 7). From this time onward

(�400 ka), the giant deer became widespread in the temperatelatitudes of Eurasia, together with modern deer species (red

deer Cervus elaphus and the reindeer Rangifer tarandus).

The Khasarian faunal assemblage (Dnepr glaciation) with

its type locality at Cherny Yar (Volga River Basin) contained

Mammuthus trogontherii chosaricus (ancestor of the woolly

mammoth Mammuthus primigenius), Panthera spelaea, Elas-

motherium sibiricum, the woolly rhinoceros Coelodonta antiqui-

tatis, Equus chosaricus, Camelus knoblochi, the long-horned bison

Bison priscus longicornis, Megaloceros giganteus, and modern

genera and species, including Cervus elaphus, Rangifer tarandus,

the musk-oxen Ovibos, the saiga antelope Saiga, etc., the water

vole Arvicola chosaricus, and a diverse assemblage of steppe-

adapted small mammals (Alexandrova, 1976; Dubrovo, 1985;

Gromov, 1948). The fauna existed in the colder and drier

e, (2013), vol. 4, pp. 605-614



climate of the periglacial forest-steppe along the southernmargin

of the continental Dnieper glaciation (Riss, RI) (Vangengeim,

1977). However, localities rich in large mammal are rather

rare. In the south of the Russian Plain, typical subarctic and

steppe smallmammals occurred (Dicrostonyx simplicior, Lemmus

sibiricus, Lagurus ex gr. lagurus, Microtus gregalis). In the

Caucasus region, the tar pit Binagady fauna near Baky belongs

to the same stage. Under modern conditions, the range of some

animals became completely separated and confined to different

biomes.

Conclusions

The data on the early and middle Pleistocene mammals from

Northern Eurasia offer important insights into our knowledge

of the history of mammals. These assemblages differ consider-

ably in their composition as a result of evolutionary and mi-

gration processes. In the Pleistocene, the general trend toward

global temperature decrease, changes in landscape and climatic

conditions, and differentiated environmental patterns caused

important changes in mammal communities within the terri-

tory at the suprageneric, generic, and species levels. The faunas

demonstrate adaptation from the temperate climatic condi-

tions of the pre-Pleistocene to the cooler and more diverse

conditions at the end of the Pleistocene. A gradual disappear-

ance is therefore noted of the Miocene–Pliocene and ther-

mophilous forms (mastodons, giraffids, hippopotamids,

hyenids) with their extinction or displacement to southern

Asia or Africa, paralleled by the development of Pleistocene

and modern genera and species, and the gradual emergence of

associations typical for modern biomes (tundra, steppes, for-

est, etc.).

The main periods of turnover in mammal communities of

Northern Eurasia coincided approximately with the maximum

drops in global temperature, which were accompanied by

large-scale transformations of the environment. The most sig-

nificant changes occurred at the Pliocene–Pleistocene (2.6 Ma)

and early–middle Pleistocene (0.8 Ma) boundaries. Changes

in faunal composition were also reported at 1.8, 1.2, and

0.4 Ma. Beginning in the middle Pleistocene, the turnover

affected mammal diversity at the generic and species levels.

Immigration to and from adjacent territories played an impor-

tant role in these faunal transformations.

The roots of some modern genera are found in the late

Miocene (Canis, Axis, etc.), the Pliocene (Nyctereutes, Ursus,

Lynx, Equus), the Gelasian (Gulo, Cervus, Alces, Capreolus,

Bison), and the middle Pleistocene (Camelus, Rangifer, Ovibos,

Saiga). The Pliocene–Pleistocene boundary is marked by the

disappearance of a number of thermophilous forms. Evolu-

tionary trends can be traced equally in the large herbivorous

animals (elephants, rhinoceroses, horses, artiodactyls), various

carnivores (canids, ursids, hyenids, felids), and small mam-

mals. A clear boreal group, which comprised the ancestral

forms of modern Northern Eurasian species, started to form

actively at the beginning of the late early Pleistocene, around

1.8 Ma. The increasing amplitude of global cooling was then

accompanied by the appearance and widespread distributions

of first boreal and then arctic forms in northern Siberia and the

mountainous regions of Central Asia. Boreal communities


became more widespread in the early middle Pleistocene

(true wolves, tiger Panthera tigris) and at the end of late middle

Pleistocene (Khasarian), with the first appearance of modern

species (Cervus elaphus, Rangifer tarandus, Ovibos, Saiga, etc.).

Zoogeographic provinciality therefore increased from the Early

toward the late middle Pleistocene.

See also: Vertebrate Records: Early Pleistocene; Late PleistoceneMegafaunal Extinctions; Late Pleistocene Mummified Mammals; LatePleistocene of Africa; Late Pleistocene of North America; LatePleistocene of South America; Late Pleistocene of Southeast Asia; Mid-Pleistocene of Africa; Mid-Pleistocene of Europe; Mid-Pleistocene ofSouthern Asia.

References

Agadjanian AK (2009) Small Mammals of the Pliocene–Pleistocene of the RussianPlain. Moscow: Nauka (in Russian).

Alexandrova LP (1976) Rodents of the Anthropogene of the European Part of the USSR.Moscow: Nauka (in Russian).

Alexeeva LI (1977) Early Anthropogene Theriofauna of East Europe. Moscow: Nauka(in Russian).

Alexeeva NV (2005) Environmental Evolution of Late Cenozoic of West Transbaikalia(Based on Small Mammal Fauna). Moscow: GEOS (in Russian).

Alexeeva NV and Erbajeva MA (2008) Diversity of Late Neogene–Pleistocene smallmammals of the Baikalian region and implications for paleoenvironment andbiostratigraphy: An overview. Quaternary International 179: 190–195.

Alexeeva NV, Karasev VV, and Erbajeva MA (2007) Pleistocene biostratigraphy of theTransbaikal area (South East Russia). Courier Forschungsinstitut SenckenbergII: 19–26.

Bajgusheva VS (1971) Fossil theriofauna of Liventzovka sand pit (northeastern of theSea of Azov). In: Bykhovskii BE (ed.) Materials on Faunas of the Anthropogene ofthe USSR, pp. 5–28. Leningrad: Nauka (in Russian).

Bajgusheva VS, Titov VV, and Tesakov AS (2001) The sequence of Plio–Pleistocenemammal faunas from the south Russian Plain (the Azov Region). Bollettino dellaSocieta Paleontoogica Itaiana 40(2): 133–138.

Cande SC and Kent DV (1995) Revised calibration of the geomagnetic polarity timescalefor the Late Cretaceous and Cenozoic. Journal of Geophysical Research, Series B100(4): 6093–6095.

Derevyanko AP and Shunkov MV (2005) Early Paleolithic site Karama in Altai Region:First results. Archaeology, Ethrnography and Anthropology of Eurasia 3(23): 52–69(in Russian).

Dmitrieva EL (1977) Neogene Antelopes of Mongolia and Adjacent Territories. Moscow:Nauka (in Russian).

Dmitrieva EL and Nesmeyanov SA (1982) Mammals and Stratigraphy of ContinentalTertiary Deposits of Southeastern Middle Asia. Moscow: Nauka (in Russian).

Dodonov AE (2002) Quaternary of Middle Asia: Stratigraphy, Correlation,Paleogeography. Moscow: GEOS (in Russian).

Dubrovo IA (1990) The Pleistocene elephants of Siberia. In: Agenbroad LD, Mead JI,and Nelson LW (eds.) Megafauna and Man, vol. 1, pp. 1–8. South Dakota: HotSprings.

Erbaeva MA and Alexeeva NV (2000) Pliocene and Pleistocene biostratigraphicsuccession of Transbaikalia with emphasis on small mammals. QuaternaryInternational 68–71: 67–75.

Foronova IV (2001) Quaternary Mammals of the South-east of Western Siberia(Kuznetsk Basin): Phylogeny, Biostratigraphy, and Paleoecology. Novosibirsk: SBRAS GEO (in Russian).

Gabunia L and Vekua A (1995) A Plio–Pleistocene hominid from Dmanisi, East Georgia,Caucasus. Nature 373: 509–512.

Garutt VE and Bajgusheva VS (1981) Archidiskodon gromovi Garutt etAlexeeva – der altyeste Elephant der Mammutlinie in Eurasien. Quartärpaläontologie4: 7–18.

Gliozzi E, Abbazzi L, Argenti P, et al. (1997) Biochronology of selected mammals,mollusks and ostracods from the Middle Pliocene to the Late Pleistocenein Italy. The state of the art. Rivista Italiana di Paleontologia e Stratigrafia103(3): 369–388.

ce, (2013), vol. 4, pp. 605-614



Gromov VI (1948) Palaeontological and Archaeological Foundation of the Stratigraphyof the Quaternary Continental Deposits in the Territory of the USSR. Moscow:The USSR Academy of Sciences (in Russian).

Hemmer H, Kahlke R-D, and Vekua AK (2010) Panthera onca georgica ssp. nov. fromthe Early Pleistocene of Damnisi (Republic of Georgia) and the phylogeography ofjaguars (Mammalia, Carnivora, Felidae). Neues Jahrbuch für Geologie undPaläontologie Abhandlungen 257(1): 115–127.

Kahlke R-D, Garcı́a N, Kostopoulos DS, et al. (2011) Western Palaearcticpalaeoenvironmental conditions during the Early and early Middle Pleistoceneinferred from large mammal communities, and implications for hominin dispersal inEurope. Quaternary Science Reviews 30(11–12): 1368–1395.

Lister A and Sher A (2001) The origin and evolution of the woolly mammoth. Science294: 1094–1097.

Markova AK (1982) Pleistocene Rodents of the Russian Plain. Moscow: Nauka(in Russian).

Markova AK (2006) Likhvin interglacial small mammal faunas of Eastern Europe.Quaternary International 149(1): 67–79.

Markova A (2007) Pleistocene mammal faunas of Eastern Europe. QuaternaryInternational 160(1): 100–111.

Nikiforova KV (ed.) (1971) Pleistocene of Tiraspol. Kishinev: Shtiintza.Palombo MR, Valli AMF, Kostopoulos DS, Alberti MT, Spassov N, and Vislobokova I

(2006) Similarity relationships between the Pliocene to Middle Pleistocene largemammal faunas of Southern Europe from Spain to the Balkans and the North PonticRegione. Courier Forschungsiinsitut Senckenberg 256: 329–347.

Qiu Zh-X (2006) Quaternary environmental changes and evolution of large mammals inNorthern China. Vertebrata PalAsiatica 44(2): 110–132.

Ranov VA, Carbonell E, and Rodrfguez XP (1995) Kuldara, earliest human occupation inCentral Asia in its Afro-Asian context. Current Anthropology 36: 337–346.

Rekovets LI and Nadachowski A (1995) Pleistocene voles (Arvicolidae) of the Ukraine.Paleontologia i Evolucio 28–29: 145–245.

Shackleton NJ (1995) New data on the evolution of Pliocene climatic variability.In: Vrba ES, Denton GH, Partridge TC, and Burkle LH (eds.) Paleoclimate and Evolutionwith Emphasis on Human Origin, pp. 242–248. New Haven: Yale University Press.

Shchelinsky VE, Dodonov AE, Baigusheva VS, et al. (2010) Early Palaeolithic sites onthe Taman Peninsula (Southern Azov Sea region, Russia): Bogatyri/Sinyaya Balkaand Rodniki. Quaternary International 223–224: 28–35.

Sher AV (1971) Mammals and Stratigraphy of the Pleistocene of Northeast USSR andNorth America. Moscow: Nauka (in Russian). Sher AV (1974) Pleistocene Mammalsand Stratigraphy of the Pleistocene of the Far Northeast USSR and North America.International Geological Review 16(7–10): 1–284 (in English).

Sher AV (1987) Olyorian land mammal age of northeastern Siberia. PalaeontographiaItalica 74: 97–112.

Sher AV, Weinstock J, Baryshnikov GF, et al. (2011) The first record of ‘spelaeoid’ bearin Arctic Siberia. Quaternary Sciences Reviews 30(17–18): 2238–2249.

Sotnikova MV (1989) Late Pliocene–Early Pleistocene Carnivora: StratigraphicSignificance. Moscow: Nauka (in Russian).

Sotnikova MV, Baigusheva VS, and Titov VV (2002) Carnivores of the Khapry faunalassemblage and their stratigraphic implication. Stratigraphic and GeologicalCorrelation 10(4): 375–390.

Sotnikova MV, Dodonov AE, and Pen’kov AV (1997) Upper Cenozoic bio-magneticstratigraphy of Central Asian mammalian localities. Palaeogeography,Palaeoclimatology, Palaeoecology 133: 243–258.

Sotnikova M and Rook L (2010) Dispersal of the Canini (Mammalia, Canidae: caninae)across Eurasia during the Late Miocene to Early Pleistocene. QuaternaryInternational 212: 86–97.


Sotnikova M and Titov V (2009) Carnivora of the Tamanian faunal unit (the Azov Seaarea). Quaternary International 201: 43–52.

Sotnikova MV and Vislobokova IA (1990) Pleistocene mammals from Lakhiti, SouthernTajikistan, USSR. Quartärpaläontologie 8: 237–244.

Tesakov AS (2004) Biostratigraphy of Middle Pliocene–Eopleistocene of Eastern Europe(Based on Small Mammals). Moscow: Nauka (in Russian).

Tesakov AS, Dodonov AE, Titov VV, and Trubikhin VM (2007) Plio–Pleistocenegeological record and small mammal faunas, eastern shore of the Azov Sea,Southern European Russia. Quaternary International 160(1): 57–69.

Tesakov AS and van Kolfschoten T (2011) The Early Pleistocene Mimomys hordijki(Arvicolinae, Rodentia) from Europe and the origin of modern nearctic sagebrushvoles (Lemmiscus). Palaeontologia Electronica 14(3): 1–11.

Titov VV (2008) Late Pliocene Large Mammal from Northeastern Sea of Azov Region.Rostov-on-Don: SSC RAS Publishing (in Russian).

Tjutkova LA and Kaipova GO (1996) Late Pliocene and Eopleistocene micromammalfaunas of southeastern Kazakhstan. Acta Zoologica Cracoviensia 39: 549–557.

Topachevsky VA, Scorik AF, and Rekovets LI (1987) Rodents of the Upper Neogene andEarly Anthropogene Deposits of the Khadjibei Lagoon. Kiev: Naukova Dumka(in Russian).

Vangengeim EA (1977) Paleontological Foundation of the Anthropogene Stratigraphy ofNorthern Asia. Moscow: Nauka (in Russian).

Vangengeim EA, Erbaeva MA, and Sotnikova MV (1990) Pleistocene mammals fromZasuhino, Western Transbaikalia. Quartärpaläontologie 8: 257–264.

Vangengeim EA and Pevzner MA (1991) The Villafranchian of the USSR: Bio- andmagnetostratigraphy. In: Vangengeim EA (ed.) Pliocene and AnthropogenePalaeogeography and Biostratigrapy, pp. 124–145. Moscow: Geological Institute(in Russian).

Vangengeim EA, Sotnikova MV, Alekseeva LI, et al. (1988) Biostratigraphy of LatePliocene–Early Pleistocene of Tadzhikistan. Moscow: Nauka (in Russian).

Vekua AK (1962) Akhalkalaki Lower Pleistocene Mammal Fauna. Tbilisi: The GeorgianNational Academy of Sciences (in Russian).

Vekua AK (1986) The Lower Pleistocene mammalian fauna of Akhalkalaki(Southern Georgia, USSR). Paleontographia Italica 74: 63–96.

Vekua AK (1995) Die Wirbeltierfauna des Villafranchian von Dmanisi und ihrebiostratigraphische Bedeutung. Jahrbuch des Romish-GermanischenZentralmuseums Mainz 42: 77–180.

Vekua A and Lordkipanidze D (2008) The history of vertebrate fauna in Eastern Georgia.Bulletin of the Georgian National Academy of Sciences 2(3): 149–154.

Vereshchagin NK (1957) Remains of Mammals from Lower Quaternary Deposits of theTaman. Leningrad: Nauka (in Russian).

Vereshchagin NK (1959) Mammals of the Caucasus. Moscow: The USSR Academy ofSciences (in Russian).

Vislobokova I (1990) Fossil Deer of Eurasia. Moscow: Nauka (in Russian).Vislobokova I (1996) The Pliocene Podpusk-Lebyazh’e mammalian faunas and

assemblages, Western Siberia. Palaeontographia Italica 83: 1–23.Vislobokova IA (2005) On Pliocene faunas with Proboscideans in the territory of the

former Soviet Union. Quaternary International 126–128: 93–105.Vislobokova IA, Sotnikova MV, and Dodonov AE (2003) Bio-events and diversity of the

Late Miocene–Pliocene mammal faunas of Russia and adjacent areas. Deinsea10: 563–574.

Vislobokova IA, Sotnikova MV, and Erbaeva MF (1995) The Villafranchianmammalian faunas of the Asiatic part of former USSR. II Quaternario8(2): 367–376.

Zazhigin VS (1980) Late Pliocene and Anthropogene Rodents of the South of WesternSiberia. Moscow: Nauka (in Russian).

e, (2013), vol. 4, pp. 605-614

Early and Middle Pleistocene of Northern EurasiaIntroductionMammal Assemblages of Northern EurasiaEarly Pleistocene: Gelasian, Middle Villafranchian, Late Villanyian, MN17The Khaprovian (Khapry) mammal assemblage and its analogs

Late Early Pleistocene: Late Villafranchian, MN18-MQ19, = Tiglian-MenapianPsekups mammal assemblage and its analogs

Terminal Early Pleistocene: Early Galerian, 1.2-0.8 MaTamanian mammal assemblage and its analogs (Nogaisky-Morozovka, = Menapian-Bavelian)

Early Middle Pleistocene: Middle-Late Galerian, = Cromerian-Elsterian, 0.8-0.4MaTiraspolian mammal assemblages and its analogs

Late Middle Pleistocene, = Aurelian, = Holsteinian-Saalian, 0.4-0.126MaSingilian and Khasarian faunas and their analogs

ConclusionsReferences

New Early and Middle Pleistocene of Northern Eurasia · 2019. 3. 13. · Encyclopedia of Quaternary...

Documents

Transcript of New Early and Middle Pleistocene of Northern Eurasia · 2019. 3. 13. · Encyclopedia of Quaternary...