The Mzammuthus Early Pleistocene Nihewan Formation, North

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}til egl.f."･･')', r, 7 #,Earth Science289

-- 298 (2003 ･gl)

(Chikyu Kagaku) vol, tt7, 289-298.2003289

The earliest specimens of the steppe mammoth, Mzammuthus tragontherii,

from the Early Pleistocene Nihewan Formation, North Chinai)

WEI Guangbiao2),TARUNO Hiroyuki3),JIN Changzhu`) and XIE Fei5}

Abstract Here we provide new evidence suggesting that thcNorth Chinais probably the Locus of erigin ol' thc

steppe mamnioth, ?T4, tragontherii, which had first adaptcd to the ceol grassland environment in the mammoth

lineage, The evidence comprises four teeth reeently reeovered from the lacustrine scdiments of the N{hewan

Formation at Majuangeu sjte, Hebei Province, The seciitnents are dated earticr than 1.36 Ma, which predates thc

first appearances of the steppe tnamtnoth not oni}, in Europe, but also in northeasteni Siberia (ca.1.2-O.8 Ma).

Compared with all thc contcmporancous manimoths, the molars of the four specimens has,e more plates(plate nurnber: x17i":'x on M] and x17x en M3>, higher and narrower erown. thinner enamel, largcr Lamellur

frequency and ciuption angle. Thcsc charactcrs can be regarded as advanced in the mammoth lincage, and the

molars are barely indistinguishablc from those uf ,W, tregvntherii t'rom Europe not only in morphology, hut also

in al] the measurin.o data. Thus they are rei'en'ed te i'Vf. tvo.epnthd'rii herein,

From chronological, distributional and morphological viewpoints, the specimens described here bridgc

lhe gaps betweeri the earlier niainrnoth fioTn China probabl>, rei'erab]e to fi4. ineridlonatis arid the later ,el tv'agontherii

1'rom the Europe and nettheasteni Siberia,

Kv, llFbivts : M ajuungou site. Nihewan Forn]ution,NorLh China, steppe mamniotb , Afmo'n niuthfts tragontiJeriiEarlvI']eistoccne u

Introduction

In 2001. tk)ur true elephant teeth werc discovered from the

flui,io-lacust rine sedi ments of thc Early Pleistocene Nihe-,un

Forrnation c] [posed iil the Maj uangou site, Yangyuan County,

Hebei Province, North China, The sediments also yieldcd

paleoljthic a/tifacts which suggest one of the earliest human

activitics in North China, The dctailed examination of the

teeth revealed that the teeth have the characteristics of thc

genus imimett.thus. Furthermore, they are assignable to thc

steppe maminoth, M trqgentherii, on the basis ()f the detailed

charactcristi)s which distinguish the species of Mlxmmuthus.

The ]ayer yielding the teeth is dated earlier than 1 ,36 Ma,

whose date is earlier than that of M/ trogontherii so far rc-

corded, The teeth are, therefore, very important for elucidat-

ing the origin, evo]ution and dispersal not only of M.

tragentherii, but atse ot' the mammoth lineage as a whole in

Eurasia.

In this pa[ er, we describe the teeth systematically and dis-

cuss their si gnificance in reeonstructing thc niammoth ]jn-

eage. We al:;o present a scenario on the cvolution and rnigra-

tion of M, tv 2gontherit' in Eurasia.

Geological Setting

The studies on biostratigraphy. paLeontologts' and archacol-

og>, of thc Nihewan Formation in the Nihewun Basin, North

China (Fig,1) have been done by many Chincse and foreign

scicntists since 1920's. The ibrmation has cN,er been treated

as the standard strata of the Early Pleistocene of North China,

Multidisciplinar>, research projccts have been carried out

mainly in the eastcm part of the N{hewan Busin in the last ten

years, A]ong with the progrcss of the magneto-stratis,raphie

studics, the Nihewan Formation hus becomc onc of the most

important strata of the Late Pliocene-Pieistocene in North

China because it yields abundant rnammalian remains as well

as stone implemcnts. and sei,eral Paleolithic sites have been

dated ma.tTneto-stratis,raphica][y with confidence.

Among these sites. the sediments of the Maj uangou site

have becn considered as the Early Pleistocenc. They include

three distinct artifact layers as MJGI, MJG2 and MJG3 in

descending ordcr. MJG1 is equal to the `Ma.iuangou

site' used

by such authors as Xie and Li (1993), Wei (t994, 1997) and

Wei et al. (]999), The mammoth specimens described here

ure obtained from the [luvio-lacustrine sand and mud of MJG3,

ReceivedAJgust 1, 2003. Accepted Septeniber 24. 2003.

I) Partly pres inted at the 2003 Annual Meeting of Japan Assoeiation for Quaternary Rcsearch.

2) Institute ol VertebrtLte Paleontology and Paleoanthropology, Chinese ,nLeademy of Sciences, R O. Box 643, Beijing, 1OO044, China,

(Present addiess : Department of Geoscienecs, Graduate School of Sciencc, Osaka City University, 3-3-13S. Sugimoto, Sumi>,oshi-ku, Osaka,

558-8S85, Japan). E-mail : "･'eL.uuangbiao@hotmai].com

3) Osaka Museum of Natural History, 1-23, Nagai-koen, Hig.ashi-sumiyoshi-ku. Osaka, 546-OO34, Japan, E-maiL:[email protected]>,,osaka.jp4) Institute ot Vertebrate 1'aleonto]ogy and Paleoanthropolog>,, Chinese Academy of Sciences, P, O. Box 643, Beijing. 1OO044, China.

5) Hebei Institute of Cultural Relics, Shijiazhuang, 050000, China.(37)

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290 WEI Guuntsbiao, TilRL'NO Hirovuki, JINChe'ins.zhLL and XIE Fc･i

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4ti 16 N

40

30

20

x:#･

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tu

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40TiN

AiPaiaeolithicsite l:Town&village

Fig, 1. Location of the Mtu'uangou siLe and ether pu] eo]ithic sitcs iTithe eastei'n part ol' lhe NLhe",an Basin. Redrawn frem XVej c1994),

which is about 8m lower than MJG [ (Fi.o. 2), Althuu.gh the

NT aj uangou site is about 600m away from the Xiaochangliang

site and the sediments ol' the former site are not ovcrlain bx,

those of the latter directly, MJGHs considered te bc about -] 2in or 20m lower than rhe aiti fact 1aver o(' {he Xiaochangliang J LL

sitejudging fi'oni lithostratigraphy (Xie and I.i 1993 : XNJei et

al, 1 999). Ii'urthermore, MJG1 is positioned about 24m lc)wer

Lhan the Banshan site in the same continuous scdiment se-

quence, and Lhe age of the Banshan site is conciderd to be thc

same as the Xiaochangliang sitc ("bi l994, 1997). which is

datcd as 1.36 Ma on the basis of palaeomagnelism (Zhu ct al.2001;Zhu ct al. 2002). Therefore, the geological agc of the

Majuun.gou sitc js older than l.36 Ma. Cai and Li (2003)proposed that the age of MJG3 is probabl}, oldcr than 1.8 Ma,

and possibly reaches 2.0 )LCa on the basis of t/ossil micromatn-

mals. Cai and Li's conc]usion on the agc ot' MJG3 is still

requircd to be verified by mu]tidisciplinary mcthods includ-

ing pa]aeomu.[Tnctism in the f'uturc, ln any case, the age of

MJG3 is older than l.36 Ma as stated abex,c. Before further

dating work is donc. wc rcgarded thc age ol' MJG] as Early

Pleistocene provisionall>, in thc present paper.

C38)

Tlerminology and measuring method

Terininology and measuring method i'or cheek teeth of true

elephants (Elephantinae) are given in Fig. 3. Elcphantinae is

charactcrized by c]asrnodent cheek teeth, naniely their crowns

arc composed of distinct plates (or lame]]ae) and U-shaped

transverse valle>,s bctween them, Half plates (or talons) are

usuallx, attachecl on the tnes{al and disTal cnds ofthc cro",n.

They beur incomplete plate f'eatures, and arc smaller than all

of eomplete plates, Senii-pLutcs sometbnes appear on the ia-

bial or lins,ual sidc oi.' the middlc part of the crown. All the

eomplete and incompletc plates are eovered by cementum.

On the occlusa] suri'ace, enamel l'igures of each plate are rep-

resented by u ruw ol' cnalncl rings in the earliest stage of abra-

sion. "Jith the adx,ance of abrasion. thcsc cnamel rings are

connected to eachothcr and then form somewhat ¢ omplicated

enamel loops, The enaniel ]oops sometimes hnve median si-

nus.cs projecting distally andfor mesial]y.

'1'hc

numbcr of thc plates are expressed as fo11ows. : The

nuinbcr of Ihc completc platcs is addcd b}, "x".

"112'T,

`'

oo "

and 'C+".

Among them, "`x"

and "112'7

mcan thc half-platcs

and seini-plates respectively, '` ov

"

indicatcs that thc nicsial

part of the ci'own has been svorn out, and "+"

nieans that the

niesial andfor distal parts hav'e been lost by some othcr rcu-

sons except 1'or normal abrasion.

The measuring methods descri bed as follc)ws and illustrated

in Fig. 3, tLnd ure modil'ied ai'ter those of Maglio <1973> and

Noj'iriko Excavation Research Group C1975).L (Length of crown) : )Ccasured parallel to the ]ong ax{s of

thc crown. In thc uppcr check tceth, the measuretnent shou]d

be taken vertical to the average plate plane. Ill the lower cheek

teeth. it is measured from the mesial end ot' the occlusul sur-

1'ace ut' thc mesial half platc Cor mcsial end of the abrasion

surfacc) to the distal cnd ot' the occlusal surfacc of thc distal

half plate. It should be taken parallel to the abrasLon surfacc

of the tooth.

Lmax (rv'Taxitnu-i length ofcrown) : In thc upper cheek teeth.

Lmax is equal to L, In the tower cheek tecth. howcver, it is

thc mesio-distal maxi mum lcngth of the crown which is mea-

sured parallel to the abraston surface or the tooth,

W (Width ol' ero",n) : Labio-[ingual maximum width oi' the

widest plate. including cementum.

H (Hei.oht oferown) : Occluso-cervical maximum height of

the unworn highest plate, measured along the plate.Hmax (Maximum hight of' crown) i ln the upper cheek teeth,

Hinax is equal to H, tn thelower cheek teeth. it is measured

vei'tica] to the abrasion surface,

E (Enainel thickness) : It is measured vcrtical to the outer

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Earliest steppe inainnnoth froin NorLl) China Lt.9l

Altitude (m)

970

950

930

910

890

870

850

830

810Fig,

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Paleolithic Site

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Sand and Gravel

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Majuangou3 (MJG3)

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Sanggan River

Synthetic geological section of thc Nihcwan Fermation in the easrcrn part of thc Nihcwan Basm,

Layer 3-22 represent the Nihewan Formation. Redrawn froni "rci (1994).

24l//

'e. 2S

.o.=t==i 26-･--- .L 1-s･

Labia]

Mcsial"

xtJ y?irdf-<")/ Occlusal

,,,

ng

tc

Enamel leop

Fig, 3, Diagram e]' u'ue elephant moLurs,

In occlusal and lateral views, showing thc tern)inolo.crv. and

measurernentstaken. Lcft/right )vli. Right/right),{i. L:length;Lmax:maximumlcngth;H:heighr:Hmax:maxgmuinheight;Vif' : width ; a : eruprion amg]e.

C39)

f'aee of the enamel surface,

F (Lamel]ur frequency) : Average number ot' plates in a dis-

tancc of 1 Ocm aiong the mesio-distal axis of thc cheek tooth,

Thc mosL accurate method ot' measurernent is to calculatc the

F from the number of comp]ete plates (cxc]udjng the termi-

na] half plates) and the length of the series ()f plates.

Hypsodonty index : HfW x i OO.

Eruption angle : Acute ung[e betwecn thc abrasion surface

und the ccrvical ]{ne or crown base in earLy stage of wear. It

is measured t'or the upper cheek teeth.

Systematic description

Elephantidae 6ray, 182 I

EIephantinae Gray, 1 821

Mkmmuthtss Brookcs. 1828

"'agonthen'i Pohlig, 1885

Selccted s>,nollynis :

Lkphas t;rn2enhaeus Falconer, 1857:p.319:1863:p.74, pl.2,

I'ig. 2,Elk7,has t7ztgontherii Pohlig., ]8g5 :p.]027: 1888:p,]93. fig,79;

p. 195. fig82.

klepthas interinentus Jourdan, 1891: p,1Ol3,

E7aph`zs nestii Pohlig, 1891i p.303 ;Osborn, l942: p,1059,



TheAssociation for the Geological Collaboration in Japan{AGCJ)

L92 11'EI GuEmgbiar).TAI< L/NC) IlirovLiki, ,JIN Chang'zhLL tmd XIE Fei

1'ig.94].EZtphas

wtistii Pavlow, 1910:p,6. pl.1, Iigs.1, 2.

EZL'phas antiettus "zLgontheroideg Zutfardi, 1 9 1 3: p. 1 30, pl.9 (III),figs.6a and 6b.

Iberffl4phas trogontherii Guenther, 1969 : p,71 1-734, TAFEL.

L. ilgs. 4, 5, 6 ; TAFEL LI-LXI.

M}im,nuihus (Rirek?)has) trqgontherii Zhou and Zhan.oJ974 :

p,57-58, pl. XXV, figs.1, 2.

ildanzmuthus armeniacus Maglio, 1973 : p.S7-60. pl, XVIII.

fig.1.Ibet}teofoxoth)n

sp. WOi ct al., 1985 : p,227, pl,], fig.4.

Rilaeoloscodthn sp, Tang et al,, 199S : p.79, pl,1. fi.o,r,,

Material. An isolatccl ]eft pt{S (V136lO). stored in the Insti-

tute of Vertebrate Palconto]ogy and Paleoanthropolot,>,, Chi-nese Academy of Scicnces (Bcijing>, An isolated right Tya]

<unnumbercd), an iso]ated left N{2 (unnumbcrcd) and a tusk

fragment (unnumbcrcd), storcd in thc Hebei Institute ofCul-

tural Relics (Shijiazhuaag, Hebei). All the specimens prob-ably belong to the saine individuul,

Locality and horizon. Third artil'act layer ut the Mtviuan.gou

site (40'13'N, -4'39'E), Yang>,uan County, Hcbei Provinee,

China (Figs, 1. 2).

Geolog{cal age. Early PIcistocene (earlier than 1.36 )vla).

Description

Third left upper molar {Pl. I, Fis,s, 1, 2)i

Thc motar is ",ell preserved andjust in an early worn stagc.

In occtusa] view, the mcsial part of thc crown is nearly straight.

and the dista] part of the crown s}ightly concave lingually.

The p]ate numbcrs ure x17x and x18x countcd in thc labial

and lingua] sides rcspectivcly. A semi-plate is well dcvel-

opcd between the t3th and 14th plales in the Iins,ual side, but

it is abscnt between the corresponding plates in the labial side.The p]ate numbcr can bc recorded as x]7]]2'x, it' we follow

Gueltther { t969) who describcd thc cheek tceth of At

"･agoniherii J'rem SUssenborn, German.v.

L and W arc 264mm and 89nim respectively; H is 1 63mm,

which is about 83 percent greater than W: F is 7.3. and the

enurnel is quitc thin (2.0mm in thickness).

Other characters of this molar are us follows : The plates

ure narrow mcsio-distaliy, which are parailel to cach other ;

Mesial ten plates are worn and form continuous and parallel-sided loops or rings on the abrasion suri'aee, In the first plate,two laLt'ge central loops ['use with each other, and form a dumb-

bell-like [oop, while a mcsio-distall>, elongatcd small ring isebserved near the ]ingual mar.g.in of the plate. The second

plate is subdivided into twu elengated irregular loops, of which(,To)

the lin.guul one is roughl.v fo]decl distally and is targer than

the labia] onc with sniooth enamel. The lin.o.ua] loop has a

promincnt round distul inec{ian sinus. The third to fifth platcsare divided into thrcc enasnel loops. The ccntral one ot' thc

three loops, is re]atively -･ider mesio-distaliy and somewhat

irregu]ar in shape. It is neuriy equal tu two ]ateral ones in

Lubio-lingual length. The distal inedian sinus of the central

loop is small, but dear in the third and fifth platcs. The cen-

tral loop is separatcd fi/om the two parallel-sided lateral loops

with smooth enamel by two longituclinal lateral grooves. The

sixth plate js divided into two parts by a relatively narrow

mediun gi'oove, The ling, ual one of the two parts consists of

t",o lateral loops and one round central rin.g., while the labial

one eonsists or two loops. or which the inner one is wider

rnesio-distall>, than the outer one. The seventh p]ate is mor-

ph()]o.yieal]y similar to the sixth plate, Its lingual part con-

sists ot' an eiongatcd large inncr loop and three sina]l round

rings. Its labiaL one consists of an inncr siiiall round ring and

two largcr os,al outer rings. Thc lingual part somewhat stcps

in front of thc labial one in the sixth and sevefith plates. Thc

eighth plate consists of four ox'al loops with nearly the saune

size. Thc central two loopg closely connect to each other.

The apex of the ninth plate is broken, and covered with gyp-sum. so that it is invisible in occiusat view. Two slightly worn

mammillae of the Lcnth plate are visible on the abrasion sur-

faee,Third

right lower molar <Pt. I. Figs. 3, 4) :

In occlusa] view. the cro-･n is slightiy twistcd ctockwisc

and moderatcly concave labially. The mesial part of the abra-

sion sui'facc is rclative]y well prcscrved. whilc its distal partis somewhat brokcn and covered with g.vpsum. The p]atesare stron.uly convergent toward the occlusal surface in ]abial

and lin.fiual vje",s. The plate number is x17x. and Lrnax is

306mm. W is 87mm, which is measured on the second plate.E and F measure 2,8mm and 6.4 respectivcly. The lower partof the erown is covered with gypsum, so that thc height of the

visible part is 1 12nun, but H probab]y attains to l60mm,

The plates are rather narrow niesio-cfistalt}, and run straight,

They are separated b>, re]atively ",ide valleys. which extend

nearl}, para[lcl to cach othcr in occlusal view, This molar is

vcry sinii1ar to the abovc-mentioned Mr' (V1361O) in the ba-sic structure oi' thc occlusal surface, cspecial1y in the form of

the plate subdivjsion. For example, the third plate of this

mo]ur is subdivided into three widely separated, parallel-sidcd,nearly equui-sjzed enumet loops, as V]361O.

Second left upper molar <Pl. T, Fig. 5) :

This specimen is strongly worn, so that several mesial platesare worn out. In occlusal vie",, the crown is somcwhat coi]-

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Earlieststel)pe n]aminoth froinNorth China 293

cave ]ingual}y, and lhe plates are slightly convergent iingually

and closcly spaeed. The ptatc number is oo 9x. Thc prc-scrved part ot' thc crown is 124mm in length, W, E und F

measure 79mm. 2,8 mm and g.O respcctivel>,, The tncsial

mtLrgin oi' the abrasion surf;Lce becomes a dentine areu. The

disto-labial part ot' the ['irst ptaLe is preserx･ed, ",hile the mesio-

lingual part ot' the sccond plate is absent. The mesial three

platcs are connccted tightly in the midline ef the crown, The

fourth to eighth plutes are also strongly ",orn, and cach plate

formsalurge enamcl loop en Lhe abrasion surface, The ine-

sial and distal sides of Lach plate are smeoth, -･eakly folded.

and nearly paral]el to each other. 'I'hc

central parL of cach

plate is slightl>･ expunded mcsio-dista]1>,. The distal median

sinuses of lhe fi fth, sixth and seventh ptates and the mcsial

median sinuses of the seventh platc arc sniall but c]ear, The

ninth platc is open distal]y and connected with the distal hnlf

plate.Right tusk (Pl. I, Fig, 6}:

This specimeii is Cragmental, but attributable vo a tugk frag-

ment, The prescnt part is slender, somewhat curved upwards

and strongly t",istcd spirally anti-clc)ckwise. Its lcngth mca-

sures about 750mm, Thc transyerse ancl longitudinal diam-

eters at one end are SOtnm and g3mm respectively, while the

corrcsponding diamcters ut another end are 76mm and 107mm

respectively,

Discussion

The teeth described abovc can be rcfcrred to the genus

it4Lxsnmuthus by the l'ollowin.o characters : They have thc par-allel-sidcci arrangemcnt of plales and ename] loops, white do

not have a prominent mesio-dista] expansion in thc median

part ol' the plates and cnamcl loops, Some ol' the moderately

worn plates are subdivided into three widely separated. par-ailel-sided. near]y equal-sizcd enamel loops on thc abrasion

surl'ace. The tusk is strongly twistcd spifally.

The genus ,lfummuthus has convcntionally been divided into

three chronospecies in Europe. They are i';ti, i'nerith'onak's rang-

ins. frorn 2.6 to O.7 Ma, M. tragontherii rangin.o from O.7 to

O.5Ma and M/ pri7nigenius ranging from O.3S to O,OIMa (Lister1996 ; Lister and Sher 2001), Amon.o them, M. "'agontheFii is

treated s>,nonymous to it4, izrJ'neniacus (Falconer), 1857

(Aguirre 1969; Maglio 1973;Beden and GuErin ]975;

Kot$akis et al, 1978;LisLer 1996), Thcrefore. in thc present

paper, we follow Pohlig (1885), Beden and Gu6rin (I975).Kotsakis ct al . (l 978) and Lister O 996) to name the Eurasian

steppe mammoth as itl trqgvntherii, The tccth described abos,c

arc undoubtedly referred lo M "'agontherii because their char-

acteristic ['eatures are barely distinguishab]e from lhose of .'V,

C41)

trag'ontherii from Stisscnborn COsborn 1942･ ; Guenther 1969)

and other sites ln Europc (Osborn 1942), All the measare-

mcnts o[' the molars described above fall within tbe ranges of

ill "mpntherii given by Maglio (1 973), Dubrovo ( t 977), Lister

(1996) and Lister ancl Sher (2001), On the other hand. they

are c]carly distin.ouishable from the earlier, more primitive

species. iW/ fineridiona!is. and the later. rnorc advanced spe-

cies, M. primigenius, from inan.v localities orEurasia (Table

1), The platc number of mo]ar has been regardcd as the true

indicator of Chc evolutionarv Iex,el of thmmut]bus (Lister und il

Sher 2001), The present Mt' and M.a havc the nurnber of x17i';x

and x17x rcspectively. These numbers fall within the range

of jW/ "'agontherii (x17x-x22x). but are outside of the ran.ge ol'

iW, prin2igenkts {x20x-x27x) (Lister t996 : Lister and Sher

200] ). Furthcrmore. Aguirre (L969) poinlLd out that the erup-

tion angle ol' upper molars varied from 34" to 580 in M,

txqgvntherii, and firom 25" to 37" in ildZ r"eridionzafis, The erup-

tion angle of the present ps{'i' tneasurcs 42'. which falls wjthin

the ran.g.c of iL4/ "'agonthe,'ii It also precludes the possibility

o(' referrin.q thc M'i to iV, meridionzalis, In addition to these

charactcrs, both o[' the crown hci.ght and hypsodonty index

(163mm and 183 rcspectively) ot' Lhe presLnt ),C" exceed the

range o]' nV, meridionalis (100.2-t41.8mm and 93.8-152,7 re-

speLtively) giyen by Maglio (1973>, whi]e they fall within

the range ofM. tragontherii (1 18.0-21S.{}mm and 145.3-304.9

respcctjve].y) confined by Maglio (]973) (Tablc l). On the

basjs od' ali thc abox,e-mentioned characters, we can rel'er the

prcsent teeth to ilf, trctgvncberii.

Beforc tbe first appearrence of fLI. trogontherii in North

Chtna, thc Chinese mammoth linea.cfe has experienced a long-

tcrm und continuous ci,olution process, We consider that a

serles of retnains t'ound formerly and ncwly t'rQm Shanxi,

Shandong and somc other districts are re ferable to niore pri}ni-

iive mamm()th specics, Among them, the eurliest occurfence

was recorded t'rom the Mazegou Formation at thc Yushe Ba-

sinin Shanxi Provlnce (Flynn et al, 1991:Tedford 1995;Qiu

and Qiu ]995). The Mazegou Formation is dated as 3.4-2.5

Ma (Qiu and Qiu 1995) or 3.4-2,9 Ma (Flynn et al, ]99] :

lbdfOrd 1995) within {he Gauss polarity chron, Thc Mazes,ou

Formation is ox,erlain b.v thc Haiyan Formation, ",hich lics

between 2.5-l.9 Ma within the Tvlatuyama polarity chron

(Flynn et al, 1991). The mammoth molurs from the Loc. 6 in

the Haiyan Forii]ation wcre identificd as 'flrchidiskoden

tokunagai Matsumote' by Tcilhard de Chardin and Trassaert

<1937). Howcver, our observation indicates that the molars

should he referred to M, fineridionafis, bccausc they are indis-

tinguishable from those of the typical fL･L meridionzaii's from

Europe in morphological characters as we]1 as nieasurements.

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The Associationfor the Geological Collaboration in Japan{AGCJ)

L294 WEI ('Tuang. biao, T.'XRLJNO Hiroyuki, JIN Changzhu and XIE Fei

SpeciesLocalitvofsampies

AuthorUpperII.owerP*Lmax W Hmax E L.FHmaxlWXIoo

ff11-142288-31.7.185.6-126.4100.2-141.82.6-4.13.7-6.193.8-152.7EtmasiaMaoiio1973e

M,IO-14212.a3oo.O69.1ffl19.475.a152.02.tl-4.13.5.5.9107.&165.5

Dubrovo1977me11-132`va-2988.S122- p4.5.5 -

M.meridionalisMs11.14220-320 ' - - 45 -

UpperValdame,Italy M312-14220.0-335,O80.0-123.51on.O.147.0253.84.lg6.41113.0N147.0

vanEsseo2oo3M,12-].52so,O-330.075.0-llO,O101,a135.02.2-4,23,83-6,5Sl2o,a13o,o

Eurasiawo1421213.0.358.057,O-107.5118.0-218,O1.5.3,O6.08.21453-304.9

Maglio1on3M,15-21236.0-340.070.0-113.096.0-160.01.8-3.0507.2133.2-206.6

M,trogontherii

Dubrovo1wrif1&as246-40073-I20129r2122-347143-208Slissenborn

,Germany M,l6-2122S3g374114127-167P347,5140-179

Eurasia&N.America ue20-27226,a285.068,O-ll3,O135,O-188,51,3-2,O6,5-11,l164,6211.8M,primigenius Maglio1973M,20-2S2cr7,O-320.265.0.1oo.O1as.O.184.11.3.2.06.8-le,2137.&189.2

sw17.5264 89 1as 2.07.3 183Specimensinthispaper Ms17 306 gy 12(h-2.86.4 '

thble 1. ComparatiN･e measurements 1'or'i'

: 'I'he

nuniber of lhe c{)Tnplete plates.the

third tnoiars of three speeies ol',W}immutht.cs.

The Haiyan niammoth i$ a iargc-sized form, and has quitcwidcr molars. Their ptate numbers are x13x on M] and x14x

on M3 respcctiN'ely. The molars also have welS-developed

median sinus, quite thick enamel and small lamel]ar fi'cquency.Bccause of its earlier ag.e, morc primitive characters and

,o,eo-

graphically near occurrence, thc Haiyan mammoth ean be

treated as the direct ancestor of the M, tragontherii froni the

Nihewan Formation described here,

Besides the Majuangou site, sonie other wctl-datcd sites in

the Nihewan Forfnation have also yielded some tnammoth

remains, One brokcn uppcr molar (V1203 l) was fbund fromthe artifact layer of the Xiaochangliung site. and was identi-

fied as 1tzlaeolvxodbn sp. by 'fting

et al. ( ] 995). 1'he age of the

site has been dated as 1,36 Ma (Zhu et al. 2001 ; Zhu et al,

2002). This incomplcte moLat' is almost indistinguishab]e mor-

pholo.oically from the M3 descrlbed hcrc. althou.oh it prescrx,csonly ['our mesiul plates, Another hroken uppcr molar (prob-ably Mi) was recoyered from thc artii'uct layer of the

Donggutuo site, and was idcntified as llalaeotbxotlon sp. by

Wk]i ct a]. (1985). The artifact ]aycr of this site ]iesjust below

the Jaramillo onset, and N-'as dated ear]ier than 1.1 Ma (Li and

Wang 1982 : Zhu et al. 2002), Thig specimefi alse preservesonly four plates, which are strongl.v worn. The mesial and

distal sides of the plales on the abrasion surt'ace are paralleland smeo{h, and do not have prominent mesio-distal cxpan-

sions in the mcdian part of the plates and cnamel loops as in

lltleobuonhn. These are distinctive characteristics ofthe mttm-

moths. Thus the two broken uppci' molars ure undoubtcdly

rel'cJ'able to Mammuthus. Furthcrmorc, it can be stated that

(42)

the thin cnamel (t.5-2,3mm) and high ]amellar frequency (7-g) indicatc that the molar from the Donggutuo site is in the`itti,

trogontherii' leveS.

Owi]ing to the discovery ofM. "'qgvnthen'i 1'rom the Nihewan

Formalion, North China has superseded northeastern Siberia

as the locus of origin of this species, Lister and Sher (2001)considered that the earliest iW/ tragontherii-likc mammoth re-

mai n was discovered t'rom the Ear]y Olyorian of northcastern

Sibcria, spanning from L2 to O.8 Ma. Its a.g.e is younger than

that of the specimens describcd herc <earlier than 1.36 Ma).

Lister and Shcr (2001) aiso proposed that both of M.

tJ'qgvnthgrii and i141 prim4g?nitts had originated in northeastern

Siberia and followed by its southward and westward dispersal

to Eurepe. In morphology, the Early Ol.vor{an remains are

barely distin.guishabie i'rom European MZ trc{gvntherii cxcept

in a smaller tooth size (Lister and Sher 2001). Although avail-

ablc specimcns are too fe", for an assessment of variability

within the Earl}, Pleis(ecene mammoth assemblage of thc

Nihewan Formation, a]] the molavs from the Nihcwan For-

niation are also sniall in s{ze and have rclalively short and

quite narrow crowns, Lister and Sher (2001) presumcd that

the jt4. tragontherii-like mammoth of the Early O]yorian was

derived from an eastern 7L4/ meridl'onall's population, but the

re]iablc remuin of M, merid)'onzaLis has nevcr been reported

from northcastern Sibcria so lar. This imp]ies that the M

trogvntherii-]ike mammoth of the Earl.v Ol.vorian probablymigrated from some arcas outside northeastern Siberia, AI-

though the hypsodonty index of thc present M3 (183) fallswithin the range ef the Ear[y Olyoriun moiars (160-l90) given

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TheAssociationfor the Geological Collaboration in Japan {AGCJ)

Earliest stcppe inanllnothrrom Norlh China 295

by I.i ster and Sher (200 l ), the 1atter molars are considered to

bc inore advanced becausc of their more plate numbers (x19x-x22x) (Lister and Sher 2001). On the basis of their earlier

a.g.e and more primitive characters, the present teeth ot' n4,

tv'ag'ontherii fi'om the Nihewan Formation probably can bc re-

garded as the uncestor ofthe M: trqgentherii-like niammoth of

northeastern Siberia. Morcover. Iarge-scale migration and

dispersal of the Eurasian mammoths probably occuiTed ['rom

south to north in the Ear]>, Plcistocene.

Although il4Z trqgontherii is an usual mcmber of the MiddlePlcistocenc mamma] faunas of Europe. it is absent in wegtern

and central Eufopc beforc O.7 Ma CLister ]996 : Lister and

Sher 2001), The eaTiicstM mogr)ntherii ofEurope was found

in Tanian Peninsu]a. which is gituated ln the eastcrn cxtremi-

tics ol' Europc CLister and Sher 2001). The :`Taman'

niam-

moth assemblagc" is considered as u possible composite of

M merithonalis (primitive form> and M ti'qgvntherii (advancedform) (Listcr and Sher 2001 ; Lister. pers. com,), The platc

number i,arics 1'rom x]4x to x]9x on M3 for the `'Taman'

mammoth assemblage", This number is generall.v less than

that of the Early Olyorian mammoth assemblage. This indi-

cates that the ]atter is mere advanced than thc lbrmer Thcre-

fore, the Early Olyorian mainmoth assemblage can bc pre-cluded l'rorn the ancestor ol' the

"Taman' mammoth asscm-

blage". On the other hand. thc p]ate number of the presentmolars from the Nihe-,an Formation iire slightly less tban

that of the representative molars of the advanced type of the`'Ibman'

mammoth assemblas,e". Furthermore, the geologi-cal age and geographical position oi' the

"`Taman' maminoth

assemblage" are 'intermediate'

bctween M/ trogontherii of

North China and that of central and westcrn Europe, The

advanced form of the "Taman'

matntnoth assemblage" has

transilional murphology. and thus it probably dispcrscd from

North China and followed by its later dispersal to central and

western Europe. The Early PIeistocene M/ tragontherii of thc

Nihewan Furmation js probab]y attributable to thc common

ancestor of itf. "'ag'ontherii from northeastern Siberia as wel!

as Europe. The new discovery of the mammoth I'rem the

Nihewan Formation provides u.g a new viewpoint to undcr-

stand the origin, evolution and dispersal ot' the Eurasian mam-

moth]ineagc.

Conclusion

On thc basis of the detailed analysis of thc morphologieal

characteristics and measuring data, the mammoth spcciinen

from the Nihewan Formation, North Chjna has been referred

to the steppe mammoth, i'lf. trogoJitherii nt tragontherii ap-

pcared in North China carlier than 1.36 Ma, ahead ofits first

appearance in Europc and northeastern Siberia (ca, 1 .2-O.8

rv･Ia), lt indicutes that North China is more acceptablc than

novthcastern Sibcria as the locus of origin of iLl. "'agontherii.

It alse indicates that thc Eurasian mammoth lineage evolved

more rapid]y in East Asia than in Europc during Earl>, Pteis-

tocene. As a missing link of thc mammoth lineage, the earli-

cst stcppe mammoth 1'rom North China has actecl as an indis-

pensable role in understandin.o the origjn, evolution and dis-

persat of the Eurasian inammoths.

( 4:g )

Acknowledgement

We are very grateful to Prof, Y. Kawamura ofAiehi Unj-

versit,v of Education, who collaborated us in the examination "of

fossit elcphant specimcns in Chinese institutions and mu-

seums, and critieally read the drait of thc present paper. We

gratefully acknowledge Prof. A,M. Li$ter of University Col-

lege of London for reading and cominenting the draft of the

presentpaper and for his hclp['ul discussion, We arc arso grate-

fu] to Prof, H, Kumui. Prof. S, Yk)shikawa andAssociate Prof,

rv{, Mitamura of Osaka City University and Dr. C..H. Chung

ol' Unix,ersity College of Londen, for their manifold hclps.

We thunk Prof, Wei Qi and Mr. Zhang Yingqi ol' the Institute

of Vertebrute Paleontology and Paleoanthropolo.ffy. Chinese

Academy ot' Sciences CIVPP) for providing thc Chinese [it-

cratures. For permission to study the mammoth specimen

(V 1361O), we are much indebtcd to J)n Zhang Zhaoqun, Proll

Qiu Zhuding and Prof, Zhcng Shaohua ot' IVPR ",ho are the

main members of the research project titled `Study

on the

estublishment of the typical Chinese biostratigraphical se-

quences' Cgrants no, 200 lDEA 20020-13). We also thank the

curators of the Tian.jin Museum of Natural History, Museum

ol' Shandong Proi,incc and Shanxi Institute of Cultural Rel-

ics, t'or alEoxN'ing us to stud}, fossil matcriat in their care, This

study was sponsored by China National Kcy-lmportant Ba-

sic Reseurch Project (grants no. PD.980001).

References

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gisemcnr P16istecbnc n]o}･en des Ab)mes de la Fage h Noailles

(Corre7e). Nouvclles Arehivcs du Museum d'Histoire Naturelle

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the Early Pleistecenc in the Nihcwan Basln. Scienee in China, D,

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,n.ikJnnmthus ph},legcnetie 1ine on the rcrriLory of the USSR. Jour-

nal of the PalaeontoLogi¢ al Society of India, 20 : 33-40.

Flynn LJ, Tcdford RH and Qiu ZX <1991) Enricbment and $Labjlity

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TheAssociationfor the Geological Collaboration in Japan{AGCJ)

296 "'EI Guangblao. T.'tRI JNO I-Ilroyuki,JIN Changzhu and XIE Fei

in the Pliocene nianiniulian t'aunu ef North C.hina. Pa]eohielogy,

17l246-265,Gucnther EW (1969) Die Elefantentnelarenaus den Kiesen Non

Stisscnborn bei "'bitnar. Pulaont Abh, Scr, A, 3 / 71 ]-734. pls.50- 61.:,iKetsakis

T, Palombo MR and Petrenje C CI97g) il･fltnemutEnts

chosttriaessedrn,asekLphusdelPleisteecncsuperioredeNiaFlaminia

CRoma). Geologlca Romalla, 17 : 41 1-44S.`'Li HM and Wang JD { ]9S2} Magnetostratigraphical study oi' sev-

cral typical sreological seetions in North China. Quatcrnar>, Ge-

olosy and Enx'ironment of China, 33-38, Ocean Press, Beijing.I.istcr AN･I (1996) Es'olurion a]id Laxononiy efEurasian intuninoths,

In: Shoshani J and Tass>' P {eds.), Tl]e Probosciclea/ Evolution

and ]'alaeoecology of Elcphants and Theh' Rela[ives, 1 e3-213.

Oxford University Press.I.istcr A)vl and Sher AV (2001) The origin and evolurion of the

wool]y niainmoth. Sciencc. 294 : 1094-]097.

)vlaglioV] (1973) Ori,o,ln andEvolutien ofthe Elephantidae. TrtLns

Amer Philos Soc (New Ser), 63 : [-149,Nojiriko ExcavaLion Research Group (L975) Thc Lake Ncljiri exca-

vation 1962-197r,. Kyoritsu Shuppan, Tokyo, Jupan, 278p."

Osborn HF (1942) Proboscidea. Nk)1.II. 80S-1675, iXnier Mus Nat

HisL, New York.

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Persicn, Uber 1'ossile Elei'antcnrcste Kaukasiens und Persiens, u]d

tiber dic ResaltaLc einer Monographic der fossilcn Ele['anLcn

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Gesellschaft, 37 : 1022-1027,"i

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YJ, Li Y and Chen VVJY (1995) }v{ammallian fossils and the

age ofXiaochangliang Paleolithic sitc ef Yangyuan, Hebei, Ver- Lebrata PalAsiatica, 33 : 74-83.iiTcdford RH (1995) Neogene mammalian biostratigraphy in China:

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Essen H c2003) 'L'ooth

merphology of ,'T･f}xnxmuthus nieritth'vnak's

i'roin the south bight ofthe North Sca and frotn several localities

in the Nctherlands. Dcinsea, 9/453-511,

"'ei Q "994) Banshan l'aleolithic site from the Lower Pleistoeene

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: 223-238.'iNVei Q (I997) The i'ramework of archaeolo.vical geology of the

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Prof Chun.g.ehien Ybung on the Hundredth Anniversar}, of his birth.

193-207. Chhia Occan Prcss, Bei.iin.e. .!iWei Q, HouY]vt and Feng XW (1999) Stone artit'acts fi'omXujiapo in the Nihcwan Basjn, Longgupo prehistoricculture, L : 119-127,

7honghua Boc'}k Co, Beijin.e.i'

"･'ei Q, rvlcng. H and Chcng SQ (1985) Ncw palaeolithic sitc from

thc Nihewan (Nihowaii) Beds. Acta Anthrop Sinica, 4 / 223-232.i}Xic F and I.i J (1993) On the cu]tural reljcs and archaeo]o.cricat

features of the Lower Paleo]ithLc site at Cen.J'iawan villas,e. Acta

Anthrop Sinica. 12 : 224-234.i',

Zhu RX,Aii ZS, 1'otts R and }Ioffman KA (2002) Magncrostrati- graphic datin.u ot' early humans in China, Earth Science Rcvicws,

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Zhu RX. IIoffman KA, Potts R, Deng CL, Pan YX. Guo B, Shi CD.

Guo ZT, \uan BYI HeuYM ttnd I']uaiig WXaJ <200 b Earlicst prcs- ence of humans in northeast China. Nature, 413:413-4I7,

1.) / in I;rench,

2) / in Chinese with Engtish abstrac.t.

3) : i" Gcrman with English abstract,

4):inItalian,

5):in.lapanese,

6):inGerman.

Explanation of Plate I,

,W}nnmu.thus trogo"therii (Pohlig)

Fig. 1:Left M". T.abial x,icw. Thc scale indicates1Ocm,FiZ., 2 / Ditto, Occlusal vie",.Fis,,3: Right rvl3: Oeeiusal view.Fig. 4:Ditto. Labial view.1-iig. )" : Left M". Occlusa] view,t"i'g. 6:Right Lusk. The scale indicates O-Ocm,

This scalc is also apptied to Figs. 2-S,

(4,1)

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AGCJJapanlnGeologioal 　Collaborationfor 　theThe 　 Assooiation

297Earliest　steppe 　mal τ1moth 　fr〔⊃【n 　N 〔）rth　Chil．la

貯

雨

霾

（45 ）

ServioeLibraryN 工工一



The 　Assooiation 　for 　the 　Geologioal 　Collaboration 　in 　Japan 　（AGCJ ｝

298 WEI 　Gual ／gbia〔

一，，　TARUN （う Ilir⊂〕yuki ，　JIN　Changzhu 　alld　XIE　Fei

魏光彪・樽野博幸

・金昌柱・謝飛，2003．中国北部の前期更新世の泥河湾累層から発見された最古の

Mammuth 〃 s　trogontherii 　地王求手斗学．　57，　289298 ，

WEI 　Guangbiao，　TARUNO 　HiroyukiJIN　Chang∠hu　and 　XIE　Fei．2003．　Tho 　cariicst　speclmens 　of　the

steppe 　mammoth ，ハ4a」’nptzut 乃us 伊 ogo η’加厂ガノ，「roln 　the　Early　Pleistocene　Nihewan　Forlnation，　Nolth　China，

Earth　Sciellce（Chikyu　Kagaku），57，289−298．

要旨

　中国河北省泥河湾盆地に分布する前期更新世の泥河湾累層で発見された．、協燗 ηz班肋 ∫初 go腕加厂〃の臼歯

3 点と切歯 1点の化石を記載した．これらは旧石器を産 1「．Llする馬圏溝遺跡から得られたもので，そのうちの

臼歯は同年代の他産地の M η露加 ∫属の化石と比較すると、咬板数が多いこと，咬板頻度が大きいこと、エ

ナメル質が薄いことから，より進化した形質を持つことがわか一った．これらの臼歯の特徴は，SUsenbornな

どヨーロッパの i「「期更新統から産出する Mtro，qoivtht．rii のそれと区別できない．

　これらの化百の産出層準の年代は，占地磁気層序の研究と地層の対比により、1．36Ma より占く，前期更新

世と考えられる．これまで Mtl’o．

VOIntlJerii あるいはそれに近いもので最古のものは，東シベリアのもので，そ

の年代は O．8−1．2Ma とされてきたが．本報告の臼歯は，それよりさらに古いことになる．他地域のド部更新

統産の痂アη槻〃加嘱の化石の年代および形質．さらに躍狸鯉誑腕の祖先の砿概ア泌 oη4出の分布を考

慮すると，ここで記載した馬圏溝遺跡産の M 脚 go励 8漉は他地域の照 t］’o．・，’ontherii の祖先であった凵∫能性

が非常に高い．

（・16 ）

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The Mzammuthus Early Pleistocene Nihewan Formation, North