A New Species of Machairodus from the Late Miocene Kalmakpai Locality in Eastern Kazakhstan (USSR)

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

M. V. Sotnikova

Sotnikova, M. v., Geo logical Institute, USS R Academy ojSciences. Pysh evskii per. 7, 1090/7 Moscow, U.S.S.R.

Received 20 August 1990, accepted November 1990

A new species, Machairodus kurt eni is described from the late Miocene locality Kalmakpai in eastern Kazakhstan (USSR). Other Vallesian and Turolian species of Ma chairodu s in the USSR are discussed. Evolutionary changes appear to have taken place 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 following mammal taxa: Vorm ela sp., Manes sp., Plesiogulo crassa Teilhard, Adcrocuta eximia The fossil locality of Kalmakpai (Fig. J) was (Roth & Wagner), Hyaenictitherium hyaenoides discovered by a geologist, B. A. Borisov in 1960. orlovi Semenov, Machairodus kurteni Sotnikova, It is located in the Zaisan depression, 160 km SE Hipparion hippidiodus Sefve, H. elegans Grornova, of the town Zaisan in Kazakhstan, USSR. The Chilot herium sp., Sin otherium zaisa nens is Karabulak suite on the right bank of the Kal Bayshashov, Cervavitus sp., Pro capreolus sp., makpai River is lithologically subdivided into Samotheriutn sp., Paleotrag us (Yuorlovia ) asia two parts: the upper, reddish-brown part, 30 m ticus Godina, Tragocerus sp., and Gazella dorca thick, and the lower, yellowish grey one, 29.4 m doides Schl.(Dmitrieva 1977, Zhegallo 1966, 1978, thick. The mammalian fauna of Kalmakpai has Godina 1979, Bayshashov 1986, Sernenov 1989). been found in the upper part (Borisov 1964). An analysis of the Kalmakpai fauna indicates Abundant fossil material has been obtained that the complex is largely dominated by Turolian from the locality during excavations carried out forms. Most of the mammalian genera reported by the Palaeontological Institute (PIN) of the from Kalmakpai have a wide stratigraphic range USSR Academy of Sciences in 1961- 1967. The limited by the end of the Turolian. These include Kalmakpai fauna has been tentatively dated to Chilotherium , Sinotherium, Tragocerus, Adem the Pliocene (Ruscinian) on the basis of the geo cuta and Hya enictitheriuin. The evolutionary logical position of the Karabulak Fm. in the level of the genera Hvaenictitheriiun and Ma Neogene sequence in the Zaisan depression. The chairodu s places the Kalmakpai fauna in the late Kalmakpai locality is currently known to contain Turolian. 362 Sotnikova : A lI ell' sp ecies ofMacltairodus • ANN. ZOOL. FENNICI Vol. 28

sn <0 E £ Mammal locality, Kalm akpai, is a Mach airodus skull, excellently ~ '" Q) m E oi ~ ·cC £U Q) E 2 (8 = small, L = large Q) Ol E Q) preser ved , and three lower jaws. Th is material is '0e-'" "f(j ~ mammals), paleo- E ~'" tfr0 '" e described as a new species of Maclut irodu s. i= U5'" -''" E'" :2: 2 we.. magnetic polarity e e 5 ' Q) .Q c ' - Q) U, '" E ''"C e.. () &:§ MN14 ~ ~ Kalmakpay L + 6 MN13 C c 0 Pavlodar SL - 2. Systematic description '" o, 7 Q) 0 MN12 e .!:Q Taraklia L + :::J '0 Order CA RN IVO RA Bowdich, 1821 -''" Q) 8 I- Famil y FELIDAE Gray, 1821 :2:'" Q) MN11 Subfamily MACH AIRODONTI NA E G ill, 1872 c Grebeniki L + 9 Q) e Q) Genu s Mach airodus Kaup , 1833 o tii 0 '" c sn MN10 ", -' 10 :2: Q) Machairodus kurteni sp. 11.1 "' ~ E:g Q) >'" I 11 MN 9 ~ ~ Kalla 8L + Holotype: Skull with Iq\ C , P"-M (nasal, '0 '0 (j) frontal and bullae region s cru shed ), PIN 2433/ 12 :2: 287.

Oth er material: Pair of mandibles with 11_." C, Fig. 1. The strat igraphic and paleomagnetic position of P' ..j, M I, broken behind the tooth row, PIN 2433/ miocene mammal localities in the USSR (Pevzner & 524; right mandible with C, PrM" broken be Vang enge im 1990 ). hind the tooth row, PIN 2433/287. Type locality: Kalmakpai,USSR, Kazakhstan, In the Kal makpai section the bone-bearing Za isa n depression , Karabulak Fm. (upper part ). beds are con fined to a zone of norm al magneti Age : Turol ian (Late Mioce ne), MN 13. zation (Jakhimov ich, pel's. cornrn.) and correlates with the lower part of the palaeom agnetic epoc h Diagnosis: A large Maclutirodus. P" present. 5. Co nsequently, the Kalmakpai fauna may be Diastema I'-C small. The incisors are close to each allocated to the zo ne MN 13. other and form an arc. P ~ narrow. Lower canine Th e Kalrnakpai Carnivora have not been

studied in detail except Hyaenictitherium liyae I The species is named in honour of Dr. Bj orn Kurten. who noides orlovi, described by Se menov ( 1989). has made a significant contribution to the study of fossil Among the material obtained by exc avation at Carnivora.

Table 1. Measurements (mm) of Machairodus skulls .

Machairodus M. giganteus M. palanderi Zd. kutteni sp. n. taracliensis Riab. (Chang 1957)

Length of skull 308 .0 310 .5 363.0a Condylobasal length 271.0 275.0 327 .0a Basal length 251.0 255 .0 297a Palatal length 133.0 140 .0 158.0, 159.0a Zygomatic width 165.0 168.0 204a Rostral width across cs_cs 66.2 67.8 Width across p4_p4 101.4 110.0 113.0 Width across p 2_p2 61.4 70.0 Postorbital width 88 .0 96.0 114.0a Condylar width 53.8 62 .7 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 of mm ). Infraorbital foramen above the paracone of Macha irodus. Lower premolars shorter than in P' large, vertical diameter 15 mm . Postorbital other Macha irodus spec ies . Metaconid-talonid processes of frontal massive and short. The bullae co mplex abse nt from lower ca rnassi al. we re presumably large, with an ave rage length of about 39 mm. Ex ternal auditory meatus Description rou nded and large, located be tween mastoid A large Machairodu s (Tables 1- 3). proc ess and glenoid fossa. Mastoid process well Skull (Fig. 2) about the size of that in an developed, its lower dorsal edge nearly reaching Afr ican lion. Th e skull height in the bullae re the level of the glenoid fossa.Sagittal cres t high. gion is half as great as its basal length. Pre Occipital crest also well developed . maxillary massive, incisor row slightly protrud Teeth possibly initially serrated, but serration ing in relation to cani ne. Diastema I-'-C' small (5.5 preserved only on upper and lower canines . Inci- 364 Sotnikova: A new species of Machairodus • AN N. ZOOL. FENNICI Vol. 28

posterior width 9.4 mm. Th e blade of P! (car nassial ) co ns ists of fo ur cu sp s, prep ar astyle Me present , protocone wea k with a n at tip , inner roo t 18 closely pressed to the other roo ts. P" in M. kurteni is prop orti onally narrower than that in other spe cies of the ge nus Machairodus (Fig. 3).M I sma ll, single-rooted, situated at right angles to P ~. Mp Mandible (Fig. 4) high (sy mphys ial height 72.5 mm). Incisors and canine situated high above Mg the level of the cheek teeth . Incisors close to each other, their tips directed upw ard to form a 16 Mp Mp single co ntact sys tem with the low er canine (Fig. E SA). In Homotherium the incisors are morpho .s logically similar. Early Turolian Machairodus, i.e. ..c: -0 M. gig ante us (Wagner) from Taraklia (Fig. 5B) , ;: Mk has -a broader symphysis, incisors set in a broad ~o.. arc , their tips directed forward, the canin e sepa H1 14 rated from the incisor row by a sma ll diastem a. There are two large mental foramina, one under H1 the first root of P, the other under the middl e part of the diastem a C i - P~. Mandible flange for pro H2 tection of upper ca nine sabre present and bound ed by distin ct ment al crests trending along the dor sal and posterior borde r of the flange. Masseteric H2 12 +------,------,-- fossa relative ly deep, its anterior margin reaching 30 40 50 the level of the posterior roo t of MI. P.1 four cusped, the fourth posterior cusp weak, formed p 4 length (mm) by the basal cingulum. Blade of P, is also four Fig. 3. Relationship between width and length of p 4 in cu sped . Anterior cusp in clo se co ntact with the Machairodus and Homotherium. - Me: Machairodus main one, with a slightly backward inclination; copei (Grebeniki), Mp:·Machairodus palanderi (China,) two posterior cu sps also inclined. Anterior edge Mg : Machairodus giganteus (Taraklia), Mk: Machairodus kurteni (Kalmakpai), H1: Homotherium (Upper Valdarno and Olivola), H2: Homotherium (Kuruksay). Table 2. Measurements (mm) of upper dentit ion of Machairodus kurteni (PIN 2433-287).

Right Left sors large, set in an arc with width (P-P) 49 mrn, length width length width and closer to canine than in other species of 11 9.0 5.6 8.9 6.0 Machairo dus. The third inci sor in clo se contact 12 10.5 8.0 11.0 with F. The upper canines broken and worn, /3 14.1 11.5 14.5 possibly during the animal's life, large, with ser CS 13.0 33.3 13.0 2 rated anterior and posteri or edges. The diastema p 5.5 3.0 absent p 3 between the ca nine and P:' is 17 mm . Left p2 9.2 21.0 9.1 p4 38.2 14.8 38.5 present, single-rooted and unicusped, possibly M1 5.0 7.5 non-functional, not wo rn. Diastem a C'_p2is 9.5 13_M1 134.1 131.0 mrn, p 2 _p _~ is 3.5 mm . P' wit h four cusps . Ante CS_M1 110.0 rior cusp large and set we ll apart from the main p 3_M1 61.0 59.2 3 s one. Th ere are two distinct posterior cusps on P' Diastema 1 _c 5.5 s behind the main one. Anterior width of p .1 7.7 rnrn, Diastema C _ p 3 17.1 17.0 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 the other spec ies of Machairodus, P4 and M I overlap three specimens. Similar (though more pro to a lesser degree. M , is relatively narrow, no noun ced) overlapping occurs in Homotherium . In traces of metaconid or talonid present.

Table 3. Measurements of lower dentition and mandible of Machairodus kurteni.

PIN 2433-287 PIN 2433-524 left right length width length width length width

I, 7.3 7.3 3.6 12 8.3 6.1 8.2 5.8 13 10.1 8.5 10.4 8.6 C, 16.2 11.6 15.2 12.2 P3 17.5 al. 17.0 17.5 7.9 P4 26.8 11.5 25.0 25.5 11.1 M, 31.2 13.0 31.5 14.5 31.3 13.5 Cj-M, . 128.2 136.0 135.0 PrP4· 46.1 43.5 42.0 P3-M,. 76.4 73.2 73.0 Diastema Cj-P3 37.7 46.9 Ramus depth behind M, 40.5 46.1 45.0 before P3 39.0 41.1 41.5 366 Sotnikova : A nell' species ofMachairodus • ANN. ZOOL. FEN NICI Vol. 28

JY A .i.,-----;«:

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

3. Discussion Th e spec imens referred to Machairodus fro m Machairodus appears in Euras ia at the end of the the Valles ian of Euro pe form a relati vely hom o middl e Miocene and is reliably traced up to the ge neo us gro up, which is currently treated as a close of the late Miocene . Remains of machai si ng le spe cies, Mach ai rodus t Machairodusv rodonts are very rare in the early Pliocene. At the aphanistus (Kaup). Amo ng the main charac ter end of the early Pliocene a new , scimitar-toothed, istics of this group are: sma ll inciso rs in the fclid , Hotnotherium, appears in Africa and Eurasia. lower jaw forming a straight row , a large lower Th e ea rliest Macha irodu s have been reported canine, a sma ll diastema between C; and Pl. large from North Africa and Asia. Th ey arc know n prem olars with a co mplete set of add itional cusps, fro m asse mblages of an age transitional bet ween a lower carnassial with a well-developed meta the Astarac ian and Valles ian or ea rly Vallesian co nid-talonid co mplex and a strong protoconid. and include Mac hairodus robinson ! Kurten fro m Th e lower jaw is massive and high , while the the Beglia Formation of Bled Douarah, Tunisia, mandibular !lange is practically undeveloped. The and M. pseudailuroides Schmidt-Kittler fro m the upp er ca rnassial has a distin ct pro tocone . A locality Yen i Eski hisar in Tu rkey (Kurten 1976, prcparasty lc is present on P ~. All the teeth were Schmidt- Kittler 1976). Th e mos t primitive forms prob ably serrated. were referred to the subgenus Miomachairodus In Western Europe M. aphanistus is reported Sc hmidt-Kittler. while the later species from the from suc h locali ties as Eppe lsheirn, Charrnoi lle, European Vallesia n were grouped in the subge nus Mo ntrcdo n, So blay and Zillingdorf (Beaumo nt Macliairodus (Bea umo nt 1978 ). 1975). ANN. ZOOL. FENNICI Vol. 28 • Sotniko va: A Ill'\\' species ojMacliairodus 367

, between C, and P,. The presence of P2 in the . ~ .... . Machairodontinae is regarded as a primitive cha racter. The Kalfa Machairodus may confidently be considered the earliest form of the Vallesian Macha irodus group in Europe. Turolian species of Machairodu s arc referred to the subgenus Amphimacha irodus Kretzoi (Beaumont 1978). The latter includes the major ity of sabre-toothed Iclids reported from the Late ~ .. ~. - " .·,~~o- ; · ;"..=-- ·::""'~ o I : .. Miocene of Eurasia. The Turolian sabre-tooths from Eurasia arc characterized by curved. large and strong upper sabres. a more prominent man dibular flange than in Machairodus aphanistus, all teeth serrated, large incisors set in a weak arc. p. with a weaker and lingually less protrudin g protocone than in Maclia irodu s aphanistus, the lower carnassial with weak or absent metaconid talonid complex. The above characters allow us to differentiate two distinct groups within the Vallesian and Turolian species of Machairo dus. Beaumont ( 1975) suggested that the Turolian sabre-toothed cats belong to the species Macliai rodus giganteus (= M. leoninusi. Machairodu s Fig. 6. Machairodus laskarevi, Kalla (TGPI-1/2257), gigauteus (Wagner) was described from the mandible, external and occlusal view (Lungu 1978). Turolian fauna of Pikenni (Greece). and a similar form has been reported from the same strati graphic level of Samos. Beaumont (1975) also In the USSR, this group of sabre-tooths in treated the following as synonyms of M. gigan cludes a Machairodu s from the Vallesian local teus: Pogonodon copei Pavlov and Mach airodus ity Kalfa in Moldavia. The combined geological, aphanist us taracliensis Riabinin from Moldavia, palaeontological and palaeomagnetic data allow USSR, and Machairodu s palanderi Zd. and pos us to date the Kalfa fauna to the upper part of sibly M. tingii Zd. from China. zone MN 9 (Lungu 1978). Lungu described a The above forms were referred to a single new species, Machairodus laskarevi, on the ba species on the basis of the characters shared by sis of a well-preserved pair of mandibles (rom all the Turolian sabre-tooths. However, Beaumont Kalfa (Fig. 6). This form shows all the characters ( 1975) indicated a high degree of variability in typical of M. aphanistus: flange virtually absent, some of the characters used for the subdivision lower jaw high, incisors small, canine large and ofmachairodonts, e.g.the height of the mandibular M I with a well-developed metaconid-talonid symphysis region, the extent of the flange devel complex. opment, the length of the diastema between C, The measurements of M. laskarcvi are simi and Pl. and the size of the third lower premolar. lar to those of M. aphanistus from Eppelsheim. Hence, the attempts to recognize evolutionary The length of P" P~ and M,are 19, 25 and 29 mrn, changes in the Turolian Machairodus of Eurasia respectively. while the corresponding measure were unsuccessful . This was partially due to the ments reported for the Machairodu s of Eppels incompleteness of the fossil material available heim arc 21. 27.4 and 30.2 mm. However, P2 is and to the lack of detailed stratigraphic data. generally absent from the M. aphanistus mandi particularly for the Asian find s. For instance. in ble. In M. laskare vi it occurs in the right branch Western Europe the most complete material of of the mandible. P2 is small, unicuspid and single sabre-toothed cats is known from Samos and rooted. and is located in the middle of the diastema Pikermi. but it consists largely of lower jaw rc- 368 Sotn ikova: A new species o]Machairodus • ANN . ZOOL. FENNICI Vol. 28 main s. The fossils from other localities are eve n accessory cusp on its posterior end . The upper more fragmentary. Although the material of Asian carnass ial has an acce ssory cusp on the antero machairodonts is somewhat richer, the North labial side ofthe tooth in front of the preparastyle, Chin ese local ities yielding fossils have not been and the protocone is better develop ed than in dated with sufficient accuracy. The fauna recov othe r Turolian Machairodu s (Fig. 3). ered from these localities is dated to the latest The evolutionary changes trace able in the Miocene (Baodean ), which corresponds to the teeth of the Turolian Macha irodus suggest that Turolian sensu lato (Li et al. 1984). they may have evolved from form s with large, The richest material of sabre-toothed cats from well develop ed prem ol ar s with complicated a variety of Turolian levels is currently available structures, towards form s with smaller and sim in the USSR . Machairodus has been reported pler premolars. The structure of the premol ars from the localiti es Grebeniki (Ukra ine), Taraklia makes it probable that Machairodus copei from (Molda via), and Kalmakpai (Kazakstan ). The Grebenik i repre sents the earliest stage of Turolian stratigraphic age of the above localities was ten Machairodus in Eura sia. tatively dated as late Vallesian to early Turolian The Ma chairodus from Taraklia is similar to (Gre beniki) and the second half of the Turolian Machairodus giganteus from Samos and Pikermi . (Taraklia). The genus Hipparion provides de It differs from the latter only in the lack of a tailed knowledge of the faunal age. The presence di stinct metaconid and talonid on the low er of Hipparion giganteum Gromova and Hlpparion carnass ial. Thi s Ma cha irodus may be regarded verae Gabunia in Grebeniki, and the occurrence as a separate subspecies, M. giganteus taracliensis of the more progressive H. moldavicum Gromova Riabinin. in Taraklia, allowed Gabunia (1986) to date the Ma chairodus palanderi and Machairodus former fauna as earliest Turolian and the latter as tingii from China were referred by Beaumont Middl e Tu rolian. ( 1975) to the species M. giganteus. Analysis of the On the basis of the literature, Beaumont (1975, materi al showed that the majority of the speci 1978) refe rred the Machairodus from Grebeniki mens descri bed by Zdan sky ( 1924) and Chang and Taraklia to the subgenus Amphimachairodus (1957) from the late Mioc ene of China have Kretzoi and identified them as the species many features in common with Machairodus from Machairodu s giganteus (Wagner) . A study of the Taraklia. Their dentition is characterized by in Ma chairodus rem ains from the se localities cisors set in a weakl y-rounded arc and slightly (Pogo nodon copei' and Machairodu s aphanistus separated from each other. The upper canines are taracliensisii shows that both form s have typical variable in size and separated from the incisors features of the genus Machairodus, but that they by a relatively large dia stema. The second upper must be attributed to different species. premolar is small and single-rooted, if present. Comparison of the upper teeth of these speci The carnassial has a relatively well developed mens yields the best information. The teeth of protocone (Fig. 4) and double parastyle. If present, Ma chairodus cope i, especiall y the premolars, the metaconid-talonid complex on the lower have a more complicated struc ture than those of carnassial is weak. The lower premolars are large the other species of Turolian Machairodus. Thi s in comparison with MI' sabre-tooth has the following characteristics: It When I compared Machairodus kurteni with is large, the third premolar has an extremely machairodonts of the giganteus group, especially stron g basal cingulum, which form s three tran s with Machairodu s from Taraklia and Machai verse plaits on the anterior end of P:' and a fifth rodus irtyschensis Orlov ( 1936), it became clear that it has a different arran gement of the lower I In addi tion to the skull described by Pavlov (1914), incisors (Fig. 5A). In addition to the smaller another bett er pre served speci men from Gr cbcniki wa s premolar length compared with that of M. and the stud ied by the autho r in the Ode ssa State Unive rsity co l reduction of the protocon e on P' , the compl ete loss lec tio n (O GU-2638). ~ Riabini n ( 1929) exami ned the unp repared skull with j aw . of the metaconid and talonid on the lower Th e prese nt author ana lyse d thi s material after the cra nium carnassial are characte ristic. Amon g the scimitar wa s separated from the ma nd ible. tooth s a similar structure of the incisor region of ANN. ZOOL. FENNICI Vol. 28 • Sotn ikova: A new spec ies ofMacha irodus 369

the mand ible is seen in the Pliocene-Pleistocene Gabuniya. L. K. rrafiyuu» . JI. 1( .) 1986: [Terrestrial ge nus Homotherium, e.g. in Homotherium ere mamm als.I (In Russian) - In:Muratov, M, V, & natidens (Fabrini) from the Kuruk say locality in Nevesskaja, L. A. (Myparon, M. B. & Heuecckas, JI . 1\. ) (eds.), [Stratigraphy of the USSR. Neoge ne Tadjikistan, described by Sotni kova (1989: table systcrn .] Izd. Nedra, Moscow, 440 pp. 8, fig. I). All the peculiarities of the teeth of Godina, A. Ya. (fonJma, A. ~I. ) 1979: IThc historical Machairodu s kurt eni are present in more spe evo lution of the giraffes (genus Palaeotragus).I (In cialized form in the Plio cene scimitar cats. Russian) - Trud y Paleontol. lnst.. Akad . Nauk USSR Hom otherium always lacks the P2, P' is small and 177. 114 pp. single-rooted and P and P are small compared Kurten, B. 1976: Fossil Carnivora from the late Tertiary of J 4 Bled Douarah and Chcrichira, Tunisia. - Notes Scrv. to MI ' The lower carnass ial lacks the metaconid Geo l. Tunisie 42: 178- 214, and talonid , the prot ocone of the upper carnassial Kurten, B. & Anderson, E, 1980: Pleistocene mammals of is weakly developed and the preparastyle is small North America, - New York. 442 pp. or lacking. Li, Ch., Wu, W. & Qiu, Zh. 1984: Chinese Neogenc: Kurten & Anderson ( 1980) suggested that subdivision and corre lation. - Vertebrata Palasiatica Homoth erium might be derived from Machai 22(3) :163- 178. Lungu. A. N. (Jly ury , A. II .) 1978: [The Hipparion fauna rodus. Among all the known species of Machai of the middle Sarma tian of Moldavia (carnivoro us rodus, M. kurteni is most similar to Homotherium . marnmalsj.] (In Russian) - lzd. Shtiintsa. Kishinev. It is a typical representative of the genus Ma 132 pp. chairodu s, but differs from the other Turolian Mein, P, 1990: Updating of MN zones, - In: Lindsay, E. sabre-too thed cat s in its morpholo gy, which H., Fahlbusch, V. & Mcin, P. (eds.), European Neogene shows an evo lutionary trend leading to Homo mamm al chronology: 73--90. Plenum Press. New York. therium . In the Machairodu s lineage, the sabre Orlov, Yu. A. 1936: Tc rtiarc Raubtierc des Westlichcn toothed cat from Kalm akpai appears to represent Sibiriens.- Trav. Inst. Paleozool. Acad. Sci. URSS the latest evoluti onary stage of this genus. 5: 111-1 52. Pavlow, M. 1914: Mammiferes Tcrtiaircs dc la Nouvelle Russie. - N. Mern. Soc. Nat. Moscou 17:6- 52. Pevzner, M. A. & Vangcngeim, E. A. 1990: The position of the Neoge ne key mamm al localities of the USSR on thc magnctochronological scale. - IX R.C.M .N.S. References Congr.. Barcelona 1990, Abstracts: 267-268. Riabinin , A. 1929: Faunc dc Mammiferes de Taraklia. Baishashov, B. U. (ba ruuanr ou. G. Y.) 19B8: [Stages of de Carnivora vera, Rodenti a, Subungulata. - Trav. Mus. vclopment of some Neoge ne rhinoceroses of Kazakh Gco l. Acad. Sci. URSS 5:127- 134. stan.] (In Russian) - Matcrialy po istorii Iauny i Ilory Schmidt-Kittler, N. 1976: Raubtierc aus dem Jungtcn iar Kazakhstana 10:74- 82. Alma-Ata. Klcinasicns. - Palaeontographica A 155:1-1 31. Beaumont. G. 1975: Recherches sur Ics Fclidcs (Mammi Semcnov, Yu. A. (CCMCIIO II. 10. A.) 1989: Ictithcrcs and feres. Carnivores) du Pliocene infericur des sables it the morphologically close hyaenids of the Neogene of Dinoth crium des en viro ns d' Eppcl shci m (Rhein the USSR. (In Russian) - Izd. Naukova Dumka, Kiev. hesscn ). - Arch. Sci. 28:369-400. 180 pp. 1978: Noles complemcntaircs sur quelques Felidcs Sotnikova, M V. (COT II IIKolla. M. B.) 1989: Late Plioccnc (Carnivores). - Arch. Sci. 3 1:219- 227. early Pleistocene Carnivora. Stratigraphic significance. Borisov, B. A. (EOpIICOIl. I) . 1\.) 1964: [Stratigraphy of (In Russian with English summary) - Izd. Nuuka, Quaternary deposits in Zaisan Basin.] (Summary of Moscow . 123 pp. Master' s thesis, in Russian) - Izd. VSEG EI. Lcnin Zdansky. O. 1924: Jungtcrtiarc Carnivoren Chinas. grad. 20 pp. Palaeontol. Sinica (C)2( I): 1-1 55. Chang. H.-C. 1957: On new material of some Machai Zhegallo, V. I. OKera!l.lo, ll. VI .) 1966: IOn thc history of rodonts of pontian age from Shansi. - Vertebrata Pliocene hipparion faunas of Mongolia and Central Palasiatica 1(3):193- 200. Asia.] (In Russian) - Bull. MOIP. Old. Gcol. 6: 11 5. Dmitrievu. E. L. (1l ~H1T pll clla . E. JI. ) 1977: [Ncogcne an 197B: [The hipparions of Central Asia.] (In Russian ) tclopes of Mongolia and adjacent territories.] (In Rus - Covmestnaya Sovc tsko- Mongo lskaya paleoruo sian) - Izd. Nauka, Moscow. 120 pp. gichcskaya ekspcditsiya. Trudy 7. 151 pp.