Ann. Zool. Fennici 28:361 -369Helsinki 19 February 1992

ISSN 000 3-455X© 1992 Finnish Zoologica l Publishing Board

A new species of Machairodus from the late MioceneKalmakpai locality in eastern Kazakhstan (USSR)

M. V. Sotnikova

Sotnikova, M. v., Geo logica l Institute, USS R Academy oj Sciences. Pysh evskii per. 7,

1090/ 7 Moscow, U.S.S.R.

Recei ved 20 Au gust 1990, accept ed November 1990

A new species, Machairodus kurt eni is described from the late Miocene localityKalmakpai in eastern Kazakhstan (USSR). Other Vallesian and Turolian species ofMa chairodu s in the USSR are discussed. Evolutionary changes appear to have takenplace in the genus Machairodus during the Turolian. In the morphology of the dentition,M. kurteni is intermediate between Ma chairodus and Homotherium .

1. Introduction

The fossil locality of Kalmakpai (Fig. J) wasdiscovered by a geologist, B. A. Borisov in 1960.It is located in the Zaisan depression, 160 km SEof the town Zaisan in Kazakhstan, USSR. TheKarabulak suite on the right bank of the Kal­makpai River is lithologically subdivided intotwo parts: the upper, reddish-brown part, 30 mthick, and the lower, yellowish grey one, 29.4 mthick. The mammalian fauna of Kalmakpai hasbeen found in the upper part (Borisov 1964).

Abundant fossil material has been obtainedfrom the locality during excavations carried outby the Palaeontological Institute (PIN) of theUSSR Academy of Sciences in 1961- 1967. TheKalmakpai fauna has been tentatively dated tothe Pliocene (Ruscinian) on the basis of the geo­logical position of the Karabulak Fm. in theNeogene sequence in the Zaisan depression. TheKalmakpai locality is currently known to contain

the following mammal taxa: Vorm ela sp., Manessp., Plesiogulo crassa Teilhard, Adc rocuta eximia(Roth & Wagner), Hyaenictitherium hyaenoidesorlovi Semenov, Machairodus kurteni Sotnikova,Hipparion hippidiodus Sefve, H. elegans Grornova,Chilot herium sp., S in otherium za isa nens isBayshashov, Cervavitus sp., Pro capreolus sp. ,Samotheriutn sp., Paleotrag us (Yuorlovia ) asia­ticus Godina, Tragocerus sp., and Gazella dorca ­doides Schl.(Dmitrieva 1977, Zhegallo 1966, 1978,Godina 1979, Bayshashov 1986, Sernenov 1989).

An analysis of the Kalmakpai fauna indicatesthat the complex is largely dominated by Turolianforms. Most of the mammalian genera reportedfrom Kalmakpai have a wide stratigraphic rangelimited by the end of the Turolian. These includeChilotherium , Sinotherium, Tragocerus, Adem­cuta and Hya enictitheriuin . The evolutionarylevel of the genera Hvaenictitheriiun and Ma ­chairodu s places the Kalmakpai fauna in the lateTurolian.

362 Sotnikova : A lI ell' sp ecies ofMacltairodus • ANN . ZOOL. FENNICI Vol. 28

<0 E sn

~ '" Q) m £ Mammal locality,c £ Q) E oi E ~ 2 (8 = small , L = large

Q) ·C U'0 '" E Q)

'" 0Ol "f(j ~ mammals), paleo-E

~ tfr '" e-'" e'" '"i= U5 -' E :2: 2 we.. magnetic polarity

5 ' Q) e e.Q c U, '" ' '"- Q) &:§ E ' Ce.. () MN14 ~ ~

Kalmakpay L +6 MN13

c C Pavlodar SL -0

'" o,

7 Q)0 MN12 e

'".!:Q Taraklia L +

-':::J '0

8 I- Q)

'"MN11 :2:

Q)Grebeniki L +c

9 Q) eo '"

Q)

c tii0 sn MN10 ", -'

10 :2: Q)

'""' ~

Q) > E:g11 MN 9 ~ ~ Kalla 8L +

'0'0 (j)

12 :2:

Fig. 1. The strat igraphic and paleomagnetic posit ion ofmiocene mammal localit ies in the USSR (Pevzner &Vang enge im 1990 ).

In the Kal makpai section the bone-bearingbeds are con fined to a zone of norm al magneti­za tion (Jakhimov ich, pel's. corn rn.) and cor relateswith the lower part of the palaeom agnetic epoc h5. Co nsequently, the Kalmakpai fauna may beallocated to the zo ne MN 13.

Th e Kalrnakpai Carnivora ha ve not beenstudied in detail except Hyaenictitherium liya e­noides orlovi, described by Se menov ( 1989).Among the materi al obtained by exc avation at

Table 1. Measurements (mm) of Machairodus skulls .

Kalm akpai , is a Mach airodus skull, exce llentlypreser ved , and three lower jaws. Th is materi al isdescribed as a new species of Maclut irodu s.

2. Systematic description

Order CA RN IVO RA Bowdich, 1821Famil y FELI DAE Gray, 1821Subfamily MACH AIRODONTI NA E G ill, 1872Genu s Mach airodus Kaup , 1833

Machairodus kurteni sp. 11.1

Holotype: Skull with Iq \ C , P"-M I (nasal,frontal and bullae region s cru shed ), PIN 2433/287 .

Oth er materi al : Pair of mandibles with 11_." C,P' ..j, M I, broken behind the tooth row, PIN 2433/524; right mandible with C, PrM" broken be­hind the tooth row, PIN 2433/287.

Type locality: Kalmakpai, USSR, Kazakhstan,Za isa n depression , Karabulak Fm. (upper part ).

Age : Turol ian (Late Mioce ne), MN 13.

Diagnosis: A large Maclutirodus. P" present.Diastema I'- C small. The incisors are close to eachother and form an arc. P~ narrow. Lower canine

I The species is named in honour of Dr. Bj orn Kurten. whohas made a significant contribution to the study of fossilCarnivora .

Length of sku llCondylobasal lengthBasal lengthPalatal lengthZygomatic widthRostral width across cs_csWidth across p4_p4

Width across p 2_p2

Postorbital widthCondylar width

Machairoduskutteni sp. n.

308 .0271.0251.0133 .0165.0

66.2101 .461.488 .053.8

M. giganteustaracl iensis Riab.

310 .5275.0255 .0140 .0168.0

67.8110 .0

96.062 .7

M. palanderi Zd.(Chang 1957)

363.0a

327 .0a

297a

158 .0, 159 .0a

204a

113.070.0

114.0a

72.0a

a Measurements made on Plate I fig. 1 of Chang 1957.

AN N. ZOOL. FENN ICI Vo l. 28 • Sotnikova: A nelt' species of Mach airodus 363

Fig. 2. Machairodus kurteni , Kalmakpai (PIN-2433/287), skull , ventral and lateral view .

closer to the incisor row than in other species ofMacha irodus. Lower prem olars shorte r than inother Macha irodus spec ies . Metaconid-talonidco mplex abse nt from lower ca rnassi al.

DescriptionA large Mach airodu s (Tables 1- 3).Skull (Fig. 2) about the size of that in an

Afr ican lion. Th e skull height in the bullae re­gion is hal f as great as its basal length. Pre­maxillary massive, inci sor row slightly protrud­ing in re lation to cani ne. Diastema I-'-C' small (5.5

mm ). Infraorbital foramen above the paracone ofP' large, vertical diameter 15 mm . Postorbitalprocesses of frontal massive and short. The bullaewe re presumably large, with an ave rage lengthof about 39 mm. Ex te rna l aud itory meatusrou nded and large , located be tween masto idproc ess and glenoid fossa. Mastoid process we ll­deve loped, its lower dorsal edge nearly reachingthe level of the glenoid fossa . Sagittal cres t high .Occipital cres t also we ll developed .

Teeth possibly initia lly serrated, but serrationpreserved only on upper and lower ca nines . Inci-

364 Sotnikova: A new species of Machairodus • AN N. ZOOL. FENNICI Vol. 28

Table 2. Measurements (mm) of upper dentit ion ofMachairodus kurteni (PIN 2433-287).

posterior width 9.4 mm. Th e blade of P! (ca r­nassial ) co ns ists of fo ur cu sp s, prep ar astyl epresent , protocone wea k with a n at tip , inner roo tclosely pressed to the other roo ts. P" in M. kurteniis prop orti onally nar rower than that in other spe­cies of the ge nus Machairodus (Fig. 3) . M I sma ll,sing le-rooted, situated at right angles to P~.

Mandible (Fig. 4) high (sy mphys ial height72 .5 mm). Incisors and canine situated high abovethe level of the cheek teeth . Incisors c lose toeach othe r, their tips directed upw ard to form asingle co ntac t sys tem with the low er canine (Fig.SA ). In Homotherium the incisors are morpho­logically similar. Early Turolian Machairodus, i.e.M. gig ante us (Wagner) from Taraklia (Fig. 5B) ,has -a broader symphysis, inci sors set in a broadarc , their tips dir ected forward, the canin e sepa­rated from the incisor row by a sma ll diastem a.There are two large mental foramina, one underthe first root of P , the other under the middl e partof the diastem a Ci-P~. Mandible flange for pro­tection of upper ca nine sabre present and bound edby distin ct ment al cres ts trending along the dor­sa l and posterior borde r of the flan ge. Massetericfossa relative ly deep , its anterio r margin reachingthe level of the posterior roo t of MI. P.1 four­cusped, the fourth posterior cusp weak, formedby the basal cingulum. Blade of P, is also four­cu sped . Anterior cusp in clo se co ntac t with themain one, with a slightly backward inclination;two posterior cu sps also inclined. Anterior edge

50

H1

H2

Mp Mp

Mk

40

p 4 length (mm)

30

12 +------ --,--- - -------,--

16

E.s..c:-0;:

~o..

14

H2

Fig. 3. Relationship between width and length of p 4 inMachairodus and Homotherium. - Me: Machairoduscopei (Grebeniki), Mp:·Machairodus palanderi (China,)Mg : Machairodus giganteus (Tarakl ia), Mk:Machairodus kurteni (Kalmakpai) , H1: Homotherium(Upper Valdarno and Olivola), H2: Homotherium(Kuruksay).

H1

Mp

18

Me

Mg

9.0 5.610.5 8.014.1 11.5

13.05.5 3.0

9.238.2 14.8

sors large, se t in an arc with width (P-P) 49 mrn,and closer to canine than in other species ofMachairo dus. The third inci sor in clo se contactwith F. The upper canines broken and worn,possibly during the animal' s life, large, with ser­rated anterior and posteri or edges. The diastemabetween the ca nine and P:' is 17 mm . Left p2present, sing le-rooted and unicusped , possiblynon-functional , not wo rn. Diastem a C'_p2 is 9 .5mrn, p2_p_~ is 3.5 mm . P' wit h four cusps . Ante­rior cusp large and se t we ll apart from the mainone. Th ere are two distinct posterior cusps on P 'behind the main one. Anter ior width of p.1 7.7 rnrn,

11

12

/3

CS

p2

p 3p4M1

13_M1

CS_M 1

p 3_M1

Diastema 13_cs

Diastema Cs_ p 3

Rightlength width

134.1

61.0

17.1

Leftlength width

8.9 6.011.014.533.3 13.0

absent21.0 9.138.5

5.0 7.5131 .0110.059 .2

5.517.0

ANN. ZOOL. FENNICI Vol. 28 • Sotnik ova : A new spec ies ofMacha irodus 365

o~I _~~--JI 3 eM

Fig. 4. Machairodus kurteni, Kalmakpa i (PIN-2433/287) , right mandib le, external and dorsal views.

of M, overlapping posterior edge of P4 in all thethree specimens. Similar (though more pro­noun ced) overlapping occurs in Homotherium . In

other spec ies of Machairodus, P4 and M I overlapto a lesser degree. M , is relatively narrow, notraces of metaconid or talonid present.

Table 3. Measurements of lower dent ition and mandib le of Machairodus kurteni.

PIN 2433-287

I,1213

C,P3P4

M,Cj-M, .PrP4 ·

P3-M, .Diastema Cj-P3Ramus depth behind M,

before P3

length width

16.2 11.617.5 al.26.8 11.531.2 13.0

128.246.176.437.740.539.0

PIN 2433-524left right

length width length width

7.3 7.3 3.68.3 6.1 8.2 5.8

10.1 8.5 10.4 8.615.2 12.2

17.0 17.5 7.925.0 25.5 11.131.5 14.5 31.3 13.5

136.0 135.043.5 42.073.2 73.0

46.946.1 45.041.1 41.5

366

A

Sotnikova : A nell' species ofMachairodus • ANN. ZOOL. FEN NICI Vol. 28

JY.i.;«:

,-----

3. Discussion

Fig. 5. Mecneiiodus mandib les,anterior part in occlusal view. - A.Machairodus kurteni, Kalmakpai(PIN-2433/524). - B. Machairo­dus giganteus taraclie nsis,Taraklia (PIN-1256/3084)

Machairodus appears in Euras ia at the end of themiddl e Miocene and is reliably traced up to theclose of the late Miocene . Remains of machai­rodonts are very rare in the early Pliocene. At theend of the early Pliocene a new , scimitar-toothed,fclid , Hotnotherium, appears in Africa and Eurasia.

Th e ea rlies t Macha irodu s have been reportedfro m North Africa and Asia. Th ey arc know nfro m asse mblages of an age transitional bet weenthe As tarac ian and Valles ian or ea rly Vallesianand include Mac ha irodus robinson ! Kurten fro mthe Begli a Formation of Bled Douarah , Tunisia,and M. pseudailuroides Schmidt-Kittler fro m thelocal ity Yen i Eski hisar in Tu rkey (Kurten 1976,Schmidt- Kittler 1976). Th e mos t primi tive for mswere referred to the subgenus MiomachairodusSc hmidt- Kittler. while the later spec ies from theEuropean Vallesia n were grouped in the subge nusMacliairodus (Bea umo nt 1978 ).

Th e spec imens referred to Macha irodus fro mthe Valles ian of Euro pe form a relati vely hom o­ge neo us gro up, which is currently treated as asi ng le spe cies , Mach ai rodus t Macha irodusvaphanistus (Kaup). Amo ng the main charac ter­istics of this group are: sma ll inciso rs in thelower jaw forming a straight row , a large lowercanine, a sma ll dias tema between C; and Pl. largeprem olars with a co mplete se t of add itional cusps,a lower carnassial with a well -developed meta­co nid-ta lonid co mplex and a strong protoconid.Th e lower jaw is massive and high , while themandibular !lange is practically undeveloped. Theupp er ca rnass ia l has a d istin ct pro tocone . Aprcparasty lc is present on P~. All the teeth wereprob ably serrated.

In Western Europe M. aphanistus is reportedfro m suc h locali ties as Eppe lsheirn, Charrnoi l le,Mo ntrcdo n, So blay and Zillin gdor f (Beaumo nt1975).

ANN. ZOOL. FENNICI Vol. 28 • Sotniko va: A Il l'\\' species ojMacliairodus 367

between C, and P,. The presence of P2 in theMachairodontinae is regarded as a primit ive cha­racter. The Kalfa Machairodus may confidentlybe considered the earliest form of the VallesianMacha irodus group in Europe.

Turolian species of Machairodu s arc referredto the subgenus Amphimacha irodus Kretzoi(Beaumont 1978). The latter includes the major­ity of sabre-toothed Iclids reported from the LateMiocene of Eurasia. The Turolian sabre-toothsfrom Eurasia arc characterized by curved. largeand strong upper sabres. a more prominent man­dibular flange than in Machairodus aphanistus, allteeth serrated, large incisors set in a weak arc. p.with a weaker and lingually less protrudin gprotocone than in Maclia irodu s aphanistus, thelower carnassial with weak or absent metaconid­talonid complex. The above characters allow usto differentiate two distinct groups within theVallesian and Turolian species of Machairo dus.

Beaumont ( 1975) suggested that the Turoliansabre-toothed cats belong to the species Macliai­rodus giganteus (= M. leoninusi . Machairodu sgigauteus (Wagner) was described from theTurolian fauna of Pikenni (Greece). and a similarform has been reported from the same strati­graphic level of Samos. Beaumont (1975) alsotreated the following as synonyms of M. gigan­teus: Pogonodon copei Pavlov and Mach airodusaphanist us taracliensis Riabinin from Moldavia,USSR, and Machairodu s palanderi Zd. and pos­sibly M. tingii Zd. from China .

The above forms were referred to a singlespecies on the basis of the characters shared byall the Turolian sabre-tooths. However, Beaumont( 1975) indicated a high degree of variabi lity insome of the characters used for the subdivisionofmachairodonts, e.g.the height of the mandibularsymphysis region, the extent of the flange devel­opment, the length of the diastema between C,and Pl. and the size of the third lower premolar.

Hence, the attempts to recognize evolutionarychanges in the Turolian Macha irodus of Eurasiawere unsuccessful . This was partially due to theincompleteness of the fossil material availableand to the lack of detailed stratigraphic data.particularly for the Asian find s. For instance. inWestern Europe the most complete material ofsabre-toothed cats is known from Samos andPikermi. but it consists largely of lower jaw rc-

Fig. 6. Machairodus laskarevi, Kalla (TGPI-1/2257),mand ible, external and occlusal view (Lungu 1978).

,. ~ .... .

~ ..~.- " .·,~~o- ; · ;"..=--·::""'~

o I : ..

In the USSR, this group of sabre-tooths in­cludes a Machairodu s from the Vallesian local­ity Kalfa in Moldavia. The combined geological,palaeontological and palaeomagnetic data allowus to date the Kalfa fauna to the upper part ofzone MN 9 (Lungu 1978). Lungu described anew species, Machairodus laskarevi, on the ba­sis of a well-preserved pair of mandibles (romKalfa (Fig. 6). This form shows all the characterstypica l of M. aphanistus: flange virtually absen t,lower jaw high, incisors small, canine large andM I with a well-developed metaconid-talonidcomplex.

The measurements of M. laskarcvi are simi­lar to those of M. aphanistus from Eppelsheim.The length of P" P~ and M, are 19, 25 and 29 mrn,respectively. while the corresponding measure­ments reported for the Machairodu s of Eppels­heim arc 21. 27.4 and 30.2 mm. However, P2 isgenerally absent from the M. aphanistus mandi­ble. In M. laskare vi it occurs in the right branchof the mandible. P2 is small, unicuspid and single­rooted. and is located in the middle of the diastema

368 Sotn ikova: A new species o] Machairodus • ANN . ZOOL. FENNICI Vol. 28

main s. The fossils from other localiti es are eve nmore fragmentary. Although the material of Asianmachairodonts is somewhat richer, the NorthChin ese local ities yielding fossils have not beendated with sufficient accuracy. The fauna recov­ered from these localities is dated to the latestMiocene (Baode an ), which corresponds to theTurolian sensu lato (Li et al. 1984).

The richest material of sabre-toothed cats froma variety of Turolian levels is currently availablein the USSR . Machairodus has been reportedfrom the localiti es Grebeniki (Ukra ine), Taraklia(Molda via), and Kalmakpai (Ka zakstan ). Thestratigraphic age of the above localities was ten­tatively dated as late Vallesian to early Turolian(Gre beniki) and the second half of the Turolian(Taraklia). The genus Hipparion provides de­tailed knowledge of the faunal age. The presenceof Hipp arion giganteum Gromova and Hlpparionverae Gabunia in Grebeniki, and the occurrenceof the more progressive H. moldavicum Gromovain Taraklia, allowed Gabunia (1986) to date theformer fauna as earlies t Turolian and the latter asMiddl e Tu rolian .

On the basis of the literature, Beaumont (1975,1978) refe rred the Machairodus from Grebenikiand Taraklia to the subgenus Amphimachairodu sKretzoi and identified them as the speciesMacha irodu s giganteus (Wagner) . A study of theMa chairodus rem ains from the se localities(Pogo nodon copei' and Machairodu s aphanistustaracliensisii shows that both form s have typicalfeatures of the genus Machairodu s, but that theymust be attributed to different species .

Comparison of the upper teeth of these speci­mens yields the best information. The teeth ofMa chairodus cope i, especiall y the premolars ,have a more complicated struc ture than those ofthe other species of Turolian Machairodus. Thi ssabre-tooth has the following characteristic s: Itis large, the third premolar has an extremelystron g basal cingulum, which form s three tran s­ver se plait s on the anterior end of P:' and a fifth

I In addi tion to the skull described by Pavlov (1914 ),ano ther bett er pre served speci men from Gr cbcniki wa sstud ied by the autho r in the Ode ssa State Unive rs ity co l­lec tio n (O GU-2638) .~ Riabini n ( 1929) exami ned the unp repared skull with j aw .

Th e prese nt author a na lyse d thi s material afte r the cra niumwa s separated from the ma nd ible .

accessory cusp on its posterior end . The uppercarnass ial has an acce ssory cusp on the antero­labial side of the tooth in front of the preparastyle,and the protocone is better develop ed than inothe r Turolian Machairodu s (Fig. 3).

The evolutionary changes trace able in theteeth of the Turolian Macha irodus sugges t thatthey may have evolved from form s with large,well dev elop ed prem ol ar s with complicatedstructures , towards form s with smaller and sim­pler premolars. The structure of the premol arsmakes it probable that Machairodus copei fromGrebenik i repre sents the earliest stage of TurolianMachairodu s in Eura sia.

The Ma chairodus from Taraklia is similar toMachairodus giganteus from Samos and Pikermi .It differs from the latter only in the lack of adi stinct metaconid and talonid on the low ercarnass ial. Thi s Ma cha irodus may be regardedas a separate subspecies, M. giganteus taracliensisRiabinin .

Ma chairodus palanderi and Machairodustingii from China were referred by Beaumont( 1975) to the species M. giganteus. Analysis of themateri al showed that the majority of the speci­mens descri bed by Zdan sky ( 1924) and Chang(1957) from the late Mioc ene of China havemany features in common with Machairodus fromTaraklia. Their dentition is characterized by in­cisors set in a weakl y-rounded arc and slightlyseparated from each other. The upper canines arevariable in size and separated from the incisorsby a relatively large dia stema. The second upperpremolar is small and single-rooted, if present.The carnassi al has a relatively well developedprotocone (Fig. 4) and doubl e parastyle. If present ,the metaconid-talonid complex on the lowercarnassial is weak. The lower premolars are largein comparison with MI'

When I compared Machairodus kurteni withmachairodonts of the giganteus group, especiallywith Machairodu s from Taraklia and Machai­rodus irtyschensis Orlov ( 1936), it became clearthat it has a different arran gement of the lowerincisors (Fig. 5A). In addition to the smallerpremolar length compared with that of M. and thereduction of the protocon e on P' , the compl ete lossof the metaconid and talonid on the lowercarnassial are characte ristic. Amon g the scimitar­tooth s a similar structure of the inci sor region of

ANN. ZOOL. FENNICI Vol. 28 • Sotn ikova: A new spec ies ofMa cha irodus 369

the mand ible is seen in the Pliocene-Pleistocenege nus Homotherium, e.g. in Homotherium ere­natidens (Fabrini) from the Kuruk say locality inTadjikistan, described by Sotni kova (1989: table8, fig. I). All the peculiarities of the teeth ofMachairodu s kurt eni are present in more spe­cia lize d form in the Plio cen e scimitar cats.Hom otherium always lacks the P2, P' is small andsingle-rooted and PJ and P4 are small comparedto MI ' The lower carnass ial lack s the metaconidand talonid , the prot ocone of the upper carnassialis weakly developed and the preparastyle is smallor lacking.

Kurten & Anderson ( 1980) suggested thatHomoth erium might be derived from Mach ai­rodus. Among all the known species of Machai­rodus, M. kurteni is most similar to Homotherium .It is a typical representative of the genus Ma­chairodu s, but differs from the other Turoliansabre-too thed cat s in its morpholo gy, whichshows an evo lutionary trend leading to Homo­therium . In the Machairodu s lineage, the sabre­toothed cat from Kalm akp ai app ears to representthe latest evoluti onary stage of this genus.

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