(Mammalia, Perissodactyla) from the Paleogene of Kazakhstan

Home , Amynodontidae, Entelodont, Metamynodon

J. Paleont.,70(4), 1996, pp. 691-696 Copyright? 1996, The PaleontologicalSociety 0022-3360/96/0070-0691$03.00

BIOCHRONOLOGICALSIGNIFICANCE OF AMYNODON'I'I AE (MAMMALIA, PERISSODACTYLA)FROM THE PALEOGENE OF KAZAKHSTAN

SPENCER G. LUCAS AND ROBERT J. EMRY New Mexico Museum of Natural History and Science, 1801 MountainRoad N. W., Albuquerque87104, and Departmentof Paleobiology,National Museum of NaturalHistory, SmithsonianInstitution, Washington,D.C. 20560

ABsTRAcr-Fivetaxa of amynodontidrhinoceroses have been named from Kazakhstan.We consideronly two of these valid, and recognize three amynodontid taxa from Paleogene deposits in Kazakhstan:Cadurcodon ardynensis (=Cadurcodon zaisanensis; =Amynodontuskabakensis) from the upperEocene (Ergilian)-lower Oligocene (Shandgolian) of the Zaysanbasin, Zaisanamynodon borisovi,from the Ergilianof the Zaysanbasin and Cadurcodonkazakademius, from Shandgolianstrata in the vicinity of the Chelkar- Teniz lake basin. Gigantamynodonakespensis, from the upperOligocene north of the Aral Sea is a rhinocerotid,not an amynodontid. Close similarity of the amynodontidsof Kazakhstanto those of Mongolia and China supportsdirect correlationof Ergilianand Shandgolianstrata across Asia.

INTRODUCTION talonids and a long lowerjaw that suggests a dolicocephalic skull. In these features it A MYNODONTIDS WERE middle Eocene-early Miocene rhino- closely resembles Cadurcodon ardynensis cerotoids known from Asia, North America and Europe. (Figure 2.1, 2.2; Osborn, 1924, figure 1; Gromova, 1954, figures Typically considered to have been amphibious, they actually 2-5), but C. kazakademius is 20-30 percent larger in virtually representa rangeof body plans from subcursorialand terrestrial all dental measurements (Table 1). We thus consider C. kaza- to graviportaland amphibious (Wall, 1982, 1989). kademius a valid, large species of Cadurcodon. In eastern Asia (China and Mongolia), amynodontids first Cadurcodon zaisanensis Belyaeva, 1962.-Belyaeva (1962) appearedduring the middle Eocene (Irdinmanhanland-mam- named Cadurcodon zaisanensis for PIN 1979, a left dentary mal "age" [lma] of Russell and Zhai, 1987) and persisteduntil fragment with P4-M3 (Belyaeva, 1962, figure 1). This specimen the end of the Oligocene(Tabenbulukian lma). The latest known is from the Buran svita (Borisov, 1963) at Kalmakpay Mountain amynodontids are two specimens of the derived, hypsodont southeast of Lake Zaysan at UTM 5257260N, 380050E, zone genus Cadurcotheriumfrom the lowerMiocene Bugti Formation 45 (Figure 1). It clearly belongs to Cadurcodon: note especially of Pakistan (Pilgrim, 1912; Raza and Meyer, 1984). The west- the lack of labial folds between the trigonids and talonids and ernmost Asian occurrencesof amynodontidsare in Kazakhstan the apparent presence of only two premolars. Indeed, the ho- (Figure1), wherefive amynodontidtaxa have been named.Here, lotype of C. zaisanensis is indistinguishable from specimens of we review their taxonomic status and discuss their biochron- C. ardynensis except for the length of the M3, which is about ological significance.AMNH refers to the Departmentof Ver- 20 percent longer than the largest specimens of C. ardynensis tebrate Paleontology, American Museum of Natural History, reported by Gromova (1954, table 10) and 30 percent longer New York; KAN to the Institute of Zoology, Academy of Sci- than the largest AMNH specimens (Table 1). We are hesitant ences of the Republic of Kazakhstan,Almaty; and PIN to the to recognize a distinct species of Cadurcodon solely on the basis PaleontologicalInstitute of the Russian Academy of Sciences, of one unusually long M3, so we regard C. zaisanensis as ajunior Moscow. subjective synonym of C. ardynensis. Cadurcodon ardynensis. -In the Kiin Kerish anticline on the north shore of Lake Zaysan at the locality known as "pantsernyy AMYNODONT TAXONOMY sloy" ("turtle plain") in the Kusto svita (UTM 5334660E, Amynodontids are not common in the Paleogene strata of 311667N, zone 45), we collected canines and a left P4 (KAN Kazakhstan,and virtually all published specimens are the ho- 35-18-50: Figure 3.6-3.7) of Cadurcodon ardynensis. With a lotypes of species. Here, we review the taxonomic statusof these length of 32 mm and width of 16 mm, this specimen is well specimens. within the range of measurements of the species, though its Cadurcodonkazakademius Biryukov, 1961.- Biryukov(1961) slightly oblique metalophid is unusual for C. ardynensis. named Cadurcodonkazakademius for KAN 663-160/54-T, a Amynodon tuskabakensis Biryukov, 1963.-Biryukov (1963, lower jaw with right P3-M3 and left M3; the tusks and left P3- p. 34) named Amynodon tuskabakensis for a right M3 (he called M2 have been restoredin plaster(Figure 2.3-2.5). This fossil is it a left M2), KAN Z-K-57-260/1419 (Figure 3.1). He also from the Myneske-Suyeklocality in centralKazakhstan east of referred a left M3 (Figure 3.3) and three canines, one of which the salt lake Chelkar-Teniz(approximately lat. 48?40'N, long. is illustrated here (Figure 3.4, 3.5), to the taxon. These specimens 66?E) (Figure 1). Biryukov (1961) identified the stratigraphic are from the Kusto svita in the Kiin Kerish anticline near the unit here as Kutanbulaksvita, but Lavrov (1959; Lavrov and locality where we collected Cadurcodon ardynensis (see above), Bazhanov, 1948) referredto it as the indricotheresvita (Russell but no more exact record of Biryukov's locality is available. and Zhai, 1987; Kordikova, 1994). A svita is a Soviet strati- Belyaeva (1971) referred Amynodon tuskabakensis to Gro- graphic term that refers to a lithostratigraphicunit with sup- mova's (1954) genus Hypsamynodon as H. tuskabakensis. Wall posed isochronous boundaries. (1982), however, suggested that Hypsamynodon is a junior sub- The holotype of C. kazakademiushas a lower dental formula jective synonym of Cadurcotherium. Thus, Russell and Zhai of 1-1-2-3, cheek teeth that are not very high crowned, little (1987) referred to Biryukov's (1963) taxon as Cadurcotherium or no labial groove between the lower cheek tooth trigonidsand tuskabakensis. 691 692 JOURNAL OF PALEONTOLOGY, V. 70, NO. 4, 1996

from the lower Aksyir svita in the Kiin Kerish anticline north of Lake Zaysan at UTM 45312592E, 5333951N. Lucas et al. (in press) redescribe the holotype as well as referred specimens of Z. borisovi from Ergilian strata in Inner Mongolia, China. This allows us to rediagnose Zaisanamynodon as a distinct genus of metamynodontinine (sensu Wall, 1989) amynodontid as follows: Zaisanamynodon is a large (LM1-3 = 205 mm) metamynodontinine amynodontid (sensu Wall, 1989) distinguished from all other members of the tribe by its third loph on P4. Zaisanamynodon is much larger than Paramynodon, has a more posteriorly positioned orbit, a relatively shorter rostrum, relatively shorter postcanine diastemata and lacks the strongly bowed out zygomatic arches and glenoid shelf of Paramynodon. Unlike Metamynodon, Zaisanamynodon has relatively long diastemata, low crowned cheek teeth, a preorbital fossa that is tightly constricted, a large I3, a relatively small infraorbital fo- ramen, three lower incisors, canines that are not extremely large, curved tusks, an orbit relatively low on the skull, a relatively slender mandibular symphysis and a less massive zygomatic arch. Zaisanamynodon differs from Megalamynodon in lacking a glenoid shelf and having a short preorbital portion of the skull and three lower incisors. Zaisanamynodon is distinguished from Cadurcotherium in that the latter has only two upper incisors and one lower incisor, very hypsodont cheek teeth, very narrow lower molars, anterior ribs and parastyles confluent on the upper molars and a reduced M3 metastyle. Z. borisovi is known in FIGURE I -Amynodontid localitiesin Kazakhstandiscussed in the text. Kazakhstan only from its holotype. 1, Kalmakpay. 2, Kiin Kerish. 3, Kur-Say. 4, Myneske-Suyek.5, Gromova (1954, p. 161) coined the name Gigantamynodon Akespe. cessator, new genus and species, for a left dentary fragment with M3 from the Ergilin-Dzo svita at Khoer-Dzan, Mongolia. We agree with Wall (1989) that G. cessator is a nomen dubium because its holotype is not sufficient to diagnose a taxon; it could The holotype M3 of A. tuskabakensis is not particularly high pertain to any of sEveral large amynodontid genera, including crowned, its parastyle and anterior rib are not confluent and it Zaisanamynodon. The Chinese species of "Gigantamynodon" has a large (not reduced) metastyle, quite different from Cad- named by Xu (1961, 1966) thus are of uncertain generic as- urcotherium (=Hypsamynodon) (see Roman and Joleaud, 1908, signment. plate 1, figures 1, 7, 9A; Gromova, 1954, figure 10a). Size of "Gigantamynodon" promissus Xu, 1966, from the Shara- the tooth (M3 length-51 mm, M3 width-43 mm) is within the murun Formation of Nei Monggol, is much smaller than Zais- range for C. ardynensis reported by Gromova (1954, table 9). anamynodon borisovi and has the large, curved tusks charac- Indeed, the holotype of A. tuskabakensis closely resembles teristic of other genera, such as Metamynodon. It does not belong AMNH 19154, the holotype of Cadurcodon ardynensis (Figure to Zaisanamynodon, but we are currently uncertain to which 3.2), in its very long ectoloph, antecrochet, the anteriorly in- genus "G." promissus should be referred. "G." giganteus Xu, clined cross lophs and size (AMNH 19154, length-49 mm, width- 1961, from the Ergilian Caijiachong Formation of Yunnan, is 44 mm). Therefore, we consider Amynodon tuskabakensis to be about the same size and morphology as Z. borisovi. However, a junior subjective synonym of Cadurcodon ardynensis. The its holotype is a left dentary fragment with M,_3 and could other M3 referred by Biryukov, KAN Z-K-57 260/1419, be- represent any of several large amynodontid genera. Therefore, longs to a rhinocerotid, not an amynodontid (compare Brunet, we consider "G."giganteus to also be a nomen dubium. 1979, plates 13, 14). Two of the canines, including the one Gigantamynodon akespensis Bayshashov, 1993.-Bayshash- illustrated here (Figure 3.4, 3.5), belong to a brontotheriid, not ov (1993) based the new species Gigantamynodon akespensis an amynodontid (compare Osborn, 1929, plates 67, 68, 101- on KAN PK 11/6, a left dentary fragment with a damaged M3 103). The other canine, KAN Z-K-57 233/1418, illustrated by (Figure 3.8, 3.9). The specimen is from the lower part of the Biryukov (1963, figure 3a) but not seen by us, is of an amy- Aral Formation near Akespe (Agyspe) on the northern shore of nodontid, and cannot be distinguished from the canine of C. the Aral Sea at UTM 41315113E, 5186634N (Figure 1). This ardynensis. locality is 3.4 m above the base of the Aral Formation and at Cadurcodon cf. C. kazakademius- Belyaeva (1970, figure 1) approximately the same stratigraphic level as the Paracerath- reported a single, well worn right M2, PIN 478-393, from the erium quarry of Oriov (1939), which is to the southeast at UTM Chilikty svita at Kur-Say gulley near the Chelkar-Teniz salt 41316737E, 5185226N. Mammals associated with the holotype lake (Figure 1). She identified the specimen as "Cadurcodon of Gigantamynodon akespensis are tragulids and Aprotodon sp. (?)". Its size (length M2 = 61 mm, width = 33 mm) and lack of The mammalian fauna of the Aral Formation is either latest a labial groove between the trigonid and the talonid indicate Oligocene (Russell and Zhai, 1987) or earliest Miocene (Akh- close resemblance to Cadurcodon kazakademius (Table 1). Fur- metiev and Sychevskaya, 1994). thermore, the Kur-Say molar is much larger than Cadurcodon Bayshashov (1993) identified KAN PK 11/6 as an amyno- ardynensis. We therefore identify the Kur-Say tooth as Cad- dontid close in morphology to the holotype of Gigantamynodon urcodon cf. C. kazakademius. cessator (Gromova, 1954, p. 162, figure 22). The holotype M3 Zaisanamynodon borisovi Belyaeva, 1971.-Belyayeva (1971) (length = 46 mm, width = 31 mm) of G. akespensis is a bilo- named Zaisanamynodon borisovi for PIN 2761/1-22, a skull, phodont tooth without labial or lingual cingulids. Depth of the lower jaw and partial skeleton (Belyaeva, 1971, figures 2-11) horizontal ramus below the M3 is approximately 74 mm. The LUCAS AND EMR Y-PALEOGENE AMYNODONTID 693

FIGURE2-Cadurcodon ardynensisfrom Inner Mongolia,China and Cadurcodonkazakademius from Kazakhstan.1, AMNH 204421, paratype of C. ardynensis,occlusal view of left P3-M3, x 1. 2, AMNH 26054, C. ardynensis,occlusal view of rightP3-M3, x 1. 3-5., KAN 663-160/54- T, holotype of Cadurcodonkazakademius. 3, occlusal view of right P3-M3, x0.75. 4, right lateralview of lowerjaw, x0.25. 5, occlusal view of lowerjaw, x 0.25.

M3 has a short ectoflexid, and its metalophid is transverse and BIOCHRONOLOGY wears transversely, indicating that the specimen is not an amynodontid, rather it is a rhinocerotid such as Aceratherium or Reevaluation of the taxonomy of amynodontids from Ka- Protaceratherium (Belyaeva, 1960, p. 108). zakhstan is of biochronological significance because it helps to 694 JOURNALOF PALEONTOLOGY,V. 70, NO. 4, 1996

TABLEI-Measurements (in mm) of lower dentitions of Cadurcodon. P3L P3W P4L P4W M1L M1W M2L M2W M3L M3W Cadurcodonardynense: AMNH 20441(a) 21 18 37 20 42 22 45 21 AMNH 26135 15 12 22 16 33 18 42 19 43 20 AMNH 26054 19 15 28 19 33 22 45 22 47 21 AMNH 26055 21 15 26 17 39 20 49 20 PIN1979(b) 28 18 38 24 48 29 65 Cadurcodonkazakademius: KAN 663-160/54-T(c) 27 18 38 28 47 30 63 33 68 34 (a) holotype (b) holotypeof Cadurcodonzaisanensis; measurements from Belyaeva (1962) (c) holotype

clarify the age relationshipsof amynodontid-bearingstrata in isovian Ergilianage. However,note that Russelland Zhai (1987) Kazakhstan(Figure 4). These are Paleogenedeposits in the Zay- originally assigned the Ergilianto the early Oligocene, an as- san basin and Turgaydepression. Removal of "Gigantamyno- signmentfollowed by Wang(1992) and Dashzeveg(1993). Des- don" akespensisfrom the Amynodontidaemeans that no amy- ignationof a global stratotypefor the Eocene-Oligocenebound- nodontids are known from the Oligo-Mioceneof the Aral Sea ary in marine strata(Premoli-Silva and Jenkins, 1993) and cor- region of western Kazakhstan. relation of this boundary to nonmarine strata in the western Two amynodontidtaxa, Zaisanamynodonborisovi and Cad- United States using magnetostratigraphyand Ar/Ar geochro- urcodonardynensis are known from the Zaysanbasin of north- nometry indicates the Chadronianland-mammal "age" is late eastern Kazakhstan.Zaisanamynodon occurrences in the Ulan Eocene (Protheroand Swisher, 1992). The Asian correlativeof Gochu, Baron Sog and Houldjin Formations of Nei Monggol, the Chadronian,the Ergilian,thus is late Eocene(Ducrocq, 1993). China are of Ergilianage. The Ulan Gochu Formationcontains Zaisanamynodonborisovi thus is of late Eocene (Ergilian)age an extensive mammalian fauna of lagomorphs,rodents, an an- and may be an index taxon of the Ergilianacross a wide geo- agalid, a mesonychid and brontotheriidand amynodontid pe- graphicarea of Asia. rissodactylscorrelative with the Ergilin-Dzomammal fauna of Stratain the Zaysanbasin that producefossils of Cadurcodon Mongolia (Russell and Zhai, 1987, p. 249-250). The overlying ardynensis-the Kusto svita and overlying Buran svita-are Baron Sog Formation contains the bronthotheriidEmboloth- youngerthan the type localityof Zaisanamynodonborisovi. Fos- erium ultimum and the chalicothereSchizotherium avitum as sil mammalsindicate the Kusto svita is of Ergilianage and that well as Zaisanamynodon borisoviand is also of Ergilian age the Ergilian-Shandgolianboundary lies within the Buran Svita (Russell and Zhai, 1987, p. 248-249). The HouldjinFormation (Russell and Zhai, 1987; Emry et al., 1995). Cadurcodonar- contains a composite mammalian fauna of taxa that range in dynensishas an Ergilian-Shandgoliandistribution in China and age from middleto late Eocene(Irdinmanhan-Ergilian). It seems Mongolia consistent with its Ergilian-Shandgoliandistribution most reasonableto regardthe Zaisanamynodonoccurrence from in the Zaysanbasin (Figure4). the Houldjin Formation at Camp Margettsas of Ergilianage, The mammalianfauna from the indricotheresvita near Lake given that its other Chinese recordsare Ergilian. Chelkar-Tenizassociated with the holotype of Cadurcodonka- Assigning an age of Ergilianto the type locality of Zaisana- zakademiusconsists oftsaganomyid(Cyclomylus turgaicus), cy- mynodonin the Zaysan basin of northeasternKazakhstan thus lindrodontid(Ardynomys kazachstanicus) and castorid (Agno- seems reasonable. Russell and Zhai (1987, p. 231) concluded tocastoraubekerovi) rodents; a hyaenodontidcreodont (Hyaen- that mammalianevidence of the age of the lowerAksyir subsvita odon dubius);a chalicothere(Schizotherium turgaicum); a he- in the Zaysanbasin is "indecisive"because reported (but largely laletid (Colodon orientalis);an indricothere(Paraceratherium unsubstantiated)mammalian taxa are a mixture of middle and transouralicum);another hyracodontid(Ardynia kazachstani- late Eocene forms. Part of this mixture results from including cus); a rhinocerotid(Eggysodon turgaicus); an entelodont (En- taxa found north of Lake Zaysan, such as Z. borisoviwith taxa telodonmajor); and an indeterminatetragulid (Russel and Zhai, found south of the lake,even thoughthickness and facieschanges 1987, p. 338). The fauna is Shandgolian(early Oligocene) in age make exact correlationuncertain across the lake. We, therefore, (Russelland Zhai, 1987). A very similarbut much more diverse apply an Ergilianage to the horizon north of the lake at the mammalian fauna from the nearby Chilikty svita is associated Kiin Kerish anticline where the holotype of Z. borisoviwas with the single molar we identify as Cadurcodoncf. C. kaza- collected. We are less certainif this age also appliesto the strata kademius.Records of Cadurcodonkazakademius in the Chelka- termed lower Aksyir subsvita south of Lake Zaysan. Teniz region thus appear to be of Shandgolianage. Amyno- Belyaeva(1971) originallyassigned Zaisanamynodon borisovi dontids from Kazakhstanthus supportcorrelation of the Ergi- a "late(?) Eocene" age, as did Akhmetiev et al. (1986). We lian and ShandgolianImas from eastern China and Mongolia supportthis conclusion by assigningthe type locality of Z. bor- across Asia.

FIGURE3- Perissodactylsfrom Kazakhstan and Inner Mongolia, China. 1, KANZ-K-57 260/1419, holotype ofAmynodon tuskabakensis, occlusal view of rightM3, x 1. 2, AMNH 19154,holotype of Cadurcodonardynensis, occlusal view of rightM2-3, x 1. 3, KANZ-K-57 260/1419, rhinocerotid,occlusal view of left M3, x 1. 4-5, KANZ-K-57 238/264,brontotheriid, internal (4) andexternal (5) viewsof canine,x0.7. 6- 7, KAN35-18-50, Cadurcodon ardynensis, occlusal (6) and lingual (7) views of leftP4, x 1.5.8-9, KANPK 11/6,holotype of Gigantamynodon akespensis,lingual (8) and occlusal (9) views of left dentaryfragment with M3, x 0.75 (8) and x 0.9 (9). LUCAS AND EMR Y-PALEOGENE AMYNODONTID 695

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