Cenozoic Primates of Eastern Eurasia (Russia and Adjacent Areas) EVGENY N

ANTHROPOLOGICAL SCIENCE Vol. 113, 103–115, 2005

Cenozoic Primates of eastern Eurasia (Russia and adjacent areas) EVGENY N. MASCHENKO1*

1Paleontological Institute, Russian Academy of Sciences, Profsouyznaya Street 123, 117995 Moscow, Russia

Received 19 May 2003; accepted 7 May 2004

Abstract In the Eocene, distribution of the order Primates in the northern part of eastern Eurasia was confined to Mongolia. A form of Omomyidae (Altanius orlovi) is represented. Northern Eurasian primates attributed to later times cover the interval between the Late Miocene (Late Turolian) to the Mid- dle Pleistocene (Mindel–Riss). Primates are distributed in the western part of eastern Eurasia (Moldavia, Ukraine), Transcaucasus (Georgia, Iranian Azerbaijan) and Central Asia (Tadjikistan, Afghanistan, Transbaikalian, Mongolia). The total number of known primate taxa is not large: seven genera and eleven species in three families (Omomyidae, Hominidae, Cercopithecidae). The Neogene and Pleistocene representatives of the order Primates comprise either widely distributed Eurasian forms or endemic taxa. The distribution pattern of primates in the western and eastern part of eastern Eurasia can be interpreted in relation to links with African and East Asian faunal provinces. By the Late Pleistocene all non-human representatives of the order Primates in the northern part of eastern Eurasia became extinct.

Key words: Eocene, late Cenozoic, eastern Eurasia, Cercopithecoidea, Hominoidea

Introduction Paleontological Institute, Russian Academy of Sciences, Moscow, Russia; GIN, Geological Institute, Russian Acad- The early history of the order Primates from the eastern emy of Sciences, Moscow, Russia; GIN U, Geological Insti- part of Eurasia reflects the restructuring of the mammalian tute Siberian Branch, Russian Academy of Sciences, Ulan- faunas of Eurasia and North America that occurred at the Ude, Russia; ZIN, Zoological Institute, Russian Academy of Paleocene–Eocene boundary at about 57 Ma. The appear- Sciences, St Petersburg, Russia. ance of the order Primates in Asia is interrelated with the advent of certain groups of American mammals in Asia. The most ancient finds of primates from Asia are dated to the Eocene Primates of the Northern Part Early Eocene (Tsagan Khushu, Mongolia in the southern of Eastern Eurasia Gobi) (Dashzeveg and McKenna, 1977; Gingerich et al., Family Omomyidae Trouessart, 1879 1991; Maschenko, 2002). Altanius Dashzeveg and McKenna, 1977 Primates became a frequent component of the mammalian Altanius orlovi Dashzeveg and McKenna, 1977 faunas of the Late Miocene of Russia and adjacent areas (Late Turolian, MN 11–13, about 8 Ma). In terms of zooge- This is the most ancient and the only Asian representative ography, the southern part of eastern Europe, theTranscauca- of the family Omomyidae. Altanius orlovi is one of the sus, and Central Asia were part of the Mediterranean smallest and most specialized representatives of the order province. The taxonomic composition of primates of this Primates. Morphologically it is close to North American area during this time span does not differ much from that of anaptomorphines. It is distinguished by a reduced P4 talonid. typical Mediterranean faunas, except in the presence of Metaconid is well expressed in the P4. The M1 trigonid is some endemic species. Primates of this area are represented open, and the paraconid is cone-shaped and not reduced. The by one genus (and species) of Hominoidea, Dryopithecus, M3 has a large paraconid displaced lingually and an elon- and by one genus and two? species of the subfamily Colobi- gated talonid. nae (Table 1). During the Late Miocene, the apes apparently Additional studies of A. orlovi revealed two forms (spe- became extinct. Later, in the Early Pliocene and Pleistocene, cies?) with similar dental morphology that differ only in Central Asia was most probably linked zoogeographically size. The status of the larger form of Altanius sp. is unclear. with Africa. Such a link would explain the distribution of The larger form has a 30% deeper mandibular corpus than some cercopithecid genera of African affinity. the smaller one. Differences between the two forms in post- Abbreviations in the text and figures are as follows: PIN, canine tooth size are insignificant. A more precise evalua- tion may be possible when more data are available on * Corresponding author. e-mail: [email protected] variability and sexual dimorphism of the species phone: 7-095-116-49-71; fax: 7-095-339-06-22 (Maschenko, 2001, 2002). Published online 10 December 2004 Distribution. Late Paleocene(?)–Early Eocene, Tsagan in J-STAGE (www.jstage.jst.go.jp) DOI: 10.1537/ase.04S015 Khushu locality, Gashatan-Bumbanian, southern Mongolia,

Central Asia. Superfamily Hominoidea Gray, 1825 Dryopithecus Lartet, 1856 Late Miocene Primates of Russia and Dryopithecus (Udabnopithecus) garedzianus the Adjacent Areas (Burchak-Abramovich and Gabashvili, 1945) In 1908, Professor V.D. Laskarev reported finds of Oreo- This taxon is represented by a right P4 and M1 in a small pithecus sp. from Kalfa, Moldavia (Late Turolian) upper jaw fragment. The fossil is housed in the State (Laskarev, 1908) (Figure 1). The material, on which the Museum of Georgia, Tbilisi. This is a small-sized Dryopith- communication was based, was later lost. However, given ecus (Figure 2), the only representative of the genus from the that the tribe Oreopithecini is endemic to central Italy, the geographical area considered in the present review. It indi- report may by unreliable. The taxonomy of primates from cates a possible link with Dryopithecus of eastern Europe. the Kalfa locality remains uncertain. So, by the beginning of This demonstrates that links between the Mediterranean and the Ruscinian, no hominoid representatives are known from Asia were maintained through the Transcaucasus, hence the the geographic area considered in this review, whereas Cer- zoogeographic significance of this specimen. copithecidae became more abundant. Several finds from Both teeth are little worn. Tooth dimensions are: P4 southern Russia were known from the beginning of the 20th length, 6.9 mm; P4 width, 10.1 mm; M1 length, 9.0 mm; M1 century (Homenko, 1915). At the end of the Miocene, cerco- width, 10.3 mm. The P4 crown is bicuspid and has three pithecids became a common component of the Hipparion roots. The protocone is the largest cusp. The mesiodistal faunas of Moldavia, Ukraine, Georgia, northern Iran and crown diameter is greater lingually than buccally. A cingu- Afghanistan. Out of the five known Miocene primate locali- lum is present on the lingual side of the crown. The M1 ties, Dryopithecidae is present in one locality and Colobinae crown is four-cusped and has three roots. The paracone and in four localities (Gremiatskiy, 1958, 1962; Burchak-Abra- metacone are larger than the protocone and hypocone. The movich, 1967, 1973; Maschenko, 1989). cingulum is weak and present at the buccal side of the crown (Burchak-Abramovich and Gabashvili, 1945; Gabunia et al., Order Primates 2001). Gen. indet.? Distribution. Late Miocene (Upper Maeotian, MN 11– 12?), Udabno locality, Sagaredgi district, eastern Georgia The list of mammals of the Hipparion fauna from the (Figure 1B). Eldar locality, eastern Georgia, contains an entry “the skull of monkey.” No systematic position was determined for this Subfamily Colobinae Blyth, 1875 specimen, and no description is known (Gremiatskiy, 1957; Mesopithecus Wagner, 1839 Burchak-Abramovich, 1967). All Late Miocene mammalian Mesopithecus ucrainicus Gremiatskiy, 1953 materials from this locality were collected before 1917 and lost in 1918. This taxon is represented by PIN 1997-4, a left maxillary Distribution. Late Miocene (Upper Sarmatian, MN 12?), fragment with P4–M3 (Gremiatskiy, 1957, 1961; Eldar locality, eastern Georgia (Figure 1B). Maschenko, 1989) (Figure 3). It is a small-sized species of Mesopithecus. The largest tooth is the M2. The M3 does not

Figure 1. Major primate localities. (A) Map of the southern part of eastern Europe. (B) Map of the Transcaucasus area. 1, Kalfa, Moldavia; 2, Eldar, Georgia; 3, Udabno, Georgia; 4, Grebeniki 1 and 2, Ukraine; 5, Petroverovka (Novopetrovka), Ukraine; 6, Budey, Moldavia; 7, Gavanosa, Moldavia; 8, Maragha, Iran; 9, Zotven, Ukraine; 10, Novopetrovka, Ukraine; 11, Voynichevo, Ukraine; 12, Kuchurgan (Voynichevo), Ukraine; 13, Kotlovina, Ukraine; 14, Kudaro 1, Georgia. Vol. 113, 2005 CENOZOIC PRIMATES OF EASTERN EURASIA 105

Figure 2. Right upper P4 and M1 of Dryopithecus garedzianus from the Udabno locality, Georgia, Late Miocene (from Burchak- Abramovich and Gabashvili, 1945). (A) P4 occlusal view; (B) P4 mesial view; (C) M1 occlusal view; (D) M1 mesial view.

Figure 3. Fragment of left maxilla of Mesopithecus ucrainicus have a markedly reduced distal part of the crown, although from the Grebeniki 1 locality, Ukraine, Late Turolian. PIN 1997-4: (A) its mesial transverse diameter is 25% larger than the distal occlusal view; (B) lingual view. transverse diameter. Relief of the molar crowns is moder- ately high. The P4 is bicuspid, and the cingulum is well developed on the distolingual surface of the crown. The length of the P4–M3 tooth row is 25.1 mm. mandible is housed in the Natural History Museum, Paris, Distribution. Late Miocene (Late Turolian/Upper Maeo- collection number MRG 1905-10; the location of the other tian, MN 13?), Grebeniki 1 locality, Velikomichailovskiy specimens is unknown. district, Odessa region, Ukraine (Figure 1A). Distribution. Late Miocene (MN 12?), Maragha locality (Figure 1B), northern Iran (Iranian Azerbaijan), 50 km from Subfamily Colobinae Blyth, 1875 the southeast coast of Lake Urmia. Mesopithecus Wagner, 1839 Mesopithecus pentelicus Wagner, 1839 Pliocene and Pleistocene Primates of Russia and Adjacent Areas This taxon is represented by a male mandibular corpus with complete tooth row and missing ascending ramus From the Pliocene (Ruscinian) and Pleistocene of Russia (Heintz et al., 1981). This is the only find of Colobinae from (Transbaikalian province), northern Mongolia, Ukraine, the Miocene of Central Asia (collection number MOL 001, Moldavia, and Tadjikistan, representatives of Cercopitheci- housed in the Paris Natural History Museum). dae are known, including two genera and four? species of Distribution. Late Miocene (MN 12?), Molayan locality, Cercopithecinae, and two genera and three species of Colo- central Afghanistan, Kabul province, Central Asia. binae. From the territories of Russia and the adjacent countries, 11 localities of this age are known that have produced Mesopithecus sp. (Mesopithecus pentelicus?) remains of cercopithecid monkeys. The chronological distribution of Cercopithecidae from these localities spans the end A complete male mandibular corpus, isolated teeth, lower of the Early Pliocene (MN 14, 4.5 Ma) to the Middle Pleis- jaw fragments, and limb bones of different individuals of tocene (0.35 Ma) (Table 1). In Moldavia and Ukraine, cerco- Mesopithecus are known from Maragha in northern Iran. pithecids make up part of the Moldavian (Ruscinian) and The mammalian fauna of Maragha is similar to the Pikermi Chaprovian (Early Villafranchian) mammalian assemblages. fauna (de Mecquenem, 1925; Bernor et al., 1996). The male One cercopithecid genus present in the Late Pliocene of Tad- 106 E.N. MASCHENKO ANTHROPOLOGICAL SCIENCE

Table 1. Chronological distribution of primates in the Miocene–Pleistocene of Russia and adjacent countries Ages Ma Country and Mammal ages Taxonomic groups localities Cercopithecinae Colobinae Hominoidea Pleistocene Late 0.35 Georgia, — Mindel M. ex gr. Kudaro 1 anderssoni–arctoides Middle Early 1.85 Pliocene Late 2.2 Tadjikistan, Villafranchian MN 16 Papio sushkini Kuruk-Say 2.2–2.4 (?) Ukraine, MN 16 (?) Dolichopithecus cf. ruscinensis Kotlovina 2.4–2.6 Mongolia, MN 16 (?) Parapresbytis eohanuman Shamar Middle 2.9–3.2 (?) Transbaykalian, MN 16 (?) Parapresbytis eohanuman Udunga Early 2.6–3.4 Moldavia, Ruscinian MN 15 Macaca sp. Gavanosa Moldavia, MN 15 Macaca sp. Tcebricovo Ukraine, MN 15 Dolichopithecus hyspsulophus Voynichevo 3.5 (?) Moldavia, MN 14/15 Dolichopithecus ruscinensis Budey Ukraine, MN 14/15 Macaca sp. Grebeniki 2 4.5 Ukraine, MN 14 Macaca sp. Dolichopitehcus ruscinensis Novopetrovka Iran, Maragha Mesopithecus pentelicus Miocene Late 5.3 Ukraine, Turolian MN 13 Mesopithecus ucrainicus Grebeniki 1 Afghanistan, Mesopithecus sp. Molayan Georgia, Eldar MN 12 primate gen. indet. Georgia, MN 11 Dryopithecus garedzianus Udabno

jikistan forms part of an Asian analogy of the Late Chapro- Subfamily Colobinae Blyth, 1875 vian (Middle Villafranchian) mammalian assemblage. Dolichopithecus Deperet, 1889 Cercopithecidae from western Transbaikalian Russia were Dolichopithecus ruscinensis Deperet, 1889 part of the Early–Middle Villafranchian mammal assemblages. The one and only representative of Cercopithecinae This taxon is represented in the Early Pliocene by isolated is known from the Middle Pleistocene of Georgia; genus lower M2 and P4, and a fragmentary right mandible (part of Macaca, forming a part of the Caucasian analogy of the the mandibular corpus and base of the processus coronoides Cromerian (warm stage of Mindel or Interglacial IV). of a juvenile individual). The dp4 is preserved in the latter In the Late Ruscinian of southern Moldavia and Ukraine, specimen from Novopetrovka, the permanent M1 of which is both subfamilies of Cercopithecidae coexisted: Colobinae not yet erupted (Figure 4) (Alexeeva, 1964). Measurements (one genus and two species: Dolichopithecus ruscinensis of the specimen are: crown length of dp4, 9.4 mm; mandibu- and D. hypsilophus) and Cercopithecinae (one genus: lar corpus depth at M1, 17 mm, corpus width at M1 18 mm. Macaca, perhaps including two species). Cercopithecinae The specimen is housed in the Zoological Institute, Acad- and Colobinae are known together from at least two locali- emy of Sciences of Ukraine, Kiev. ties dated to the latest Ruscinian of Ukraine: Novopetrovka A left M2 (M1?), GIN 541-133 from Kotlovina (Figure 5), and Budey (Figure 1A, Table 1). Beginning in the second is attributed to Dolichopithecus ruscinensis as the latest half of the Late Pliocene (from MN 16), the subfamily Cer- example of the genus in southeastern Europe. The associated copithecinae is present only in the southern part of eastern mammalian fauna is Chaprovian (Early Villafranchian) Europe and in Tadjikistan. Stratigraphic distribution of the (Alexeeva, 1977). The specimen was found by I.M. Gromov genus Dolichopithecus in southern Ukraine was most proba- in 1964, and is housed in the Geological Institute, Russian bly limited to between 2.2 and 2.4 Ma. From the Late Rus- Academy of Sciences, Moscow. cinian(?)–Early Villafranchian of Ukraine, D. hypsilophus, a Distribution. Early Pliocene, Ruscinian (MN 14–15?), species smaller than D. ruscinensis, is known. Novopetrovka locality, Velikomichailovskiy district, Odessa region, Ukraine (Figure 1A). Late Pliocene, Late Villa- Vol. 113, 2005 CENOZOIC PRIMATES OF EASTERN EURASIA 107

Figure 4. Fragment of the mandible of Dolichopithecus ruscinensis from the Novopetrovka locality, Ukraine, Early Pliocene (from Alexeeva, 1964). (A) Buccal view; (B) occlusal view. Figure 5. Left M2 of Dolichopithecus ruscinensis from the Kot- lovina locality, Ukraine, Late Villafranchian. GIN 541-133: (A) buccal view; (B) occlusal view (stereo pair). franchian (MN 16?), the Kotlovina locality, Odessa region, southern Ukraine (Figure 1A). Dolichopithecus sp. Dolichopithecus hypsilophus (Gremiatskiy, 1958) A fragmentary humerus (Figure 7) and a lower jaw frag- This taxon is represented by two lower jaw fragments ment with P3–M3 lacking the ascending ramus from Budey from a locality near Voynichevo village (Kuchurgan River are attributed to Dolicopithecus. Both specimens are housed valley) (Gremiatskiy, 1958, 1961; Maschenko, 1989, in the Zoological Institute, Academy of Sciences of Molda- 1991a). The specimens are housed in the Paleontological via, Kishinev (Alexeeva, 1964; David and Obade, 1995). Institute, Russian Academy of Sciences: PIN 355-7 is a left Measurements of the humerus are: preserved length (lacking mandibular corpus preserving fragmentary P3–M3; PIN 355- proximal and distal epiphyses), 182 mm; preserved distal 9 is a right mandibular corpus preserving the symphyseal width, 35 mm; minimal diaphyseal width, 16 mm. part, lower canine, and P4–M3 (Figure 6). This is a middle- Distribution. Early Pliocene, Late Ruscinian (MN 14–15), sized colobine species, smaller than D. ruscinensis. The the Budey locality, right bank of the Bolshaia Salcha river, length of the P3–M3 row is 47 mm. The very narrow sym- Porat district, southern Moldavia (Figure 1A). physis is flattened. The incisors are aligned in a straight row and, judging by alveolar size, were very small. In the male Subfamily Cercopithecinae Gray, 1821 D. hypsilophus, the canine is smaller than in the male of D. Macaca Lacepede, 1799 ruscinensis. Molar relief is not as deep as it is in D. ruscin- Macaca sp. ensis. It does not differ significantly from D. ruscinensis in 1 the structure of the C–P4, the morphology of the symphyseal Macaca is represented by an isolated left upper M (GIN part and the relatively small lower incisors. The main cusps 1047/93) (Figure 8) (Tesakov and Maschenko, 1992) from of the molars are not as inclined mesially as they are in D. the Grebeniki 2 locality, the first reliably identified find of ruscinensis. the genus Macaca from the Pliocene of Ukraine. The local- Distribution. Early Pliocene, latest Ruscinian–Early Villa- ity is attributed to the end of the Early Ruscinian to the franchian(?) (MN 15), the Voynichevo locality, Kuchurgan beginning the Late Ruscinian (MN 14/15). The main cusps river, Odessa region, southern Ukraine (Figure 1A). of the molar are bunodont. There is an additional styloid cusp in the middle of the buccal surface. Mesial and distal transverse diameters of the crown are equal. The crown 108 E.N. MASCHENKO ANTHROPOLOGICAL SCIENCE

Figure 7. Fragment of left humerus Dolichopithecus ruscinensis (from Alexeeva, 1964) from the Budey locality, Moldavia, Early Pliocene. (A) Anterior view; (B) posterior view.

Figure 6. Mandibles of Dolichopithecus hypsilophus from the tute, Academy of Sciences of Moldavia, Kishinev (David Voynichevo locality, Ukraine, the latest Ruscinian–Early Villa- and Obade, 1995); and an isolated left male canine from Tce- franchian(?) (MN 15). PIN 355-9: (A) buccal view; (B) occlusal view; PIN 355-7: (C) buccal view; (D) occlusal view. bricovo, Moldavia, housed in the Paleontological Institute, Russian Academy of Sciences, Moscow. Distribution. Early Pliocene, Late Ruscinian (MN 14/15), the Grebeniki 2 locality, Velikomichailovskiy district, relief is similar to the cercopithecine condition, as well as Odessa region, Ukraine (Figure 1A). Early Pliocene, Late the shape of the main cusps and their positions. In features Ruscinian (MN 14–15), the Gavanosa ravine locality, such as the size of the mesial and distal flanks, shape and Bolshaia Salcha river, southern Moldavia (Figure 1A). Early depth of the median-lingual and median-buccal notches and Pliocene, Late Ruscinian (MN 15), the Tcebricovo locality, grooves, the Grebeniki 2 monkey may be assigned to the Kagul district, southern Moldavia (Figure 1A). genus Macaca. The mesial transverse diameter is 6.5 mm; distal transverse diameter 6.5 mm; mesiodistal diameter Colobine Monkey from Transbaikalian and 6.9 mm; length of lingual root 9.5 mm; buccal height of Northern Mongolia crown 4.8 mm. Other specimens attributable to Macaca include a partial In Transbaikalian and Mongolia, from the boundary of the upper jaw with P3–M3 and base of the zygomatic process Early–Middle Pliocene, only Colobinae (Parapresbytis from the Gavanosa ravine, housed in the Zoological Insti- eohanuman) is present and Cercopithecinae is absent. The Vol. 113, 2005 CENOZOIC PRIMATES OF EASTERN EURASIA 109

lafranchian (Kalmykov, 1992). The frontal part of the maxilla is broadened transversely and robust. The upper incisors are relatively large and inclined anteriorly. The labial enamel of the incisors forms vertical folds. The mesiodistal diameter of the incisors is less or equal to the buccolingual diameter. The mandibular incisors are inclined posteriorly. Lateral crests (folds) of enamel and enamel protuberance at the base of the lingual surface of the incisors form pockets. Parapresbytis differs from Doli- chopithecus in the relatively minor development of crown 3 relief of the molars, in C–P and C–P3 complex structures, and in the extremely wide and short lower facial part of the skull. The special adaptation of this Asian colobine is illus- trated by the morphology of the incisors, canines, and premolars (robust and usually deeply worn in comparison with molars), and morphology of the frontal part of the maxilla. These features differentiate Parapresbytis from Rhino- pithecus as well as from other Asian colobines. Two lower jaws (PIN 3381-236, 3381-235) and fragments of the limb, a humerus, and an ulna (PIN 3381-210, 3381- 211) from Shamar, northern Mongolia are also attributable to P. eohanuman (Figure 10). These specimens, found in 1975, were later described as Presbytis eohanuman Boris- soglebskaya, 1981 (Borissoglebskaya, 1981). Re-examina- 1 Figure 8. Isolated upper left M of Macaca sp. from the Grebeniki tion of the Shamar material shows that the Transbaikalian 2 locality, Ukraine, Late Ruscinian. GIN 1074/93: (A) buccal view; colobine bears similarity with recent Rhinopithecus. Parap- (B) lingual view; (C) occlusal view (stereo pair). resbytis is larger in size than recent colobines. The charac- ters of the trochlea of the humerus are indicative of an adaptation to arboreal life rather than to terrestriality (Figure cercopithecines appear in China and Korea in the Middle 10). These specimens are housed in the Paleontological Pleistocene. P. eohanuman is a representative of the ‘rhino- Institute, Russian Academy of Sciences, Moscow. pithecomorph’ colobine group, widely spread in Asia Distribution. Middle Pliocene, latest Ruscinian(?)–Early (Transbaikalian, Mongolia, China, Japan) from the end of Villafranchian (MN 16?), the Udunga locality, southern the Middle Pliocene to the Early Pleistocene. It is a large- Transbaikalian, Russia. Late Pliocene, Early Villafranchian sized representative of Colobinae characterized by robust (MN 16), the Shamar locality, northern Mongolia. upper and lower incisors. Enamel on the labial surface of the incisors has well developed shovelling. The lingual surface Middle Asian Representatives of Papio of the incisors forms an enamel pocket with lateral enamel folds. The M3 has a non-reduced distal pair of main cusps. In the Late Pliocene–Early Pleistocene of Ukraine and The dentition of P. eohanuman is rather close to that of mod- Moldavia the genus Paradolichopithecus is not known. In ern Rhinopithecus roxellana and R. lantianensis from the the Late Pliocene of Central Asia (Tadjikistan, Kuruk-Say) a Early Pleistocene of China. representative of the genus Papio is reliably identified. Papio (Paradolichopithecus) sushkini is a specialized repre- Subfamily Colobinae Blyth, 1875 sentative of the genus. It is medium-sized and differs from Parapresbytis Kalmykov and Maschenko, 1992 other extinct and modern Papio in large bunodont molar Parapresbytis eohanuman (Borissoglebskaya, 1981) teeth and thick enamel. The material from Tadjikistan testifies that Papio was not endemic to Africa, and that in the end The colobine monkey from southern Transbaikalian (the of the Pliocene to the beginning of the Pleistocene the genus Udunga locality, Buryat Republic) is the northernmost pri- was wide-spread in Asia. The time of dispersal of Papio mate ever recorded in Eurasia. It is notably the only primate coincided or was almost synchronous with the adaptive radi- occurrence known from Siberia. Abundant material from the ation of the genus into areas of its basic distribution. Most Udunga locality provided a basis for distinguishing a new likely, the Late Pliocene (Early–Middle Villafranchian) was colobine genus, Parapresbytis eohanuman (Kalmykov and a time of maximal distribution of the genus in the Old World. Maschenko, 1992, 1995; Maschenko, 1994b). The specimens are housed in the Geological Institute, Siberian Branch Subfamily Cercopithecinae Gray, 1821 of the Russian Academy of Sciences, Ulan-Ude. Specimens Papio Muller, 1773 include GIN U 987/445, GIN U 987/493 (1), GIN U 987/878 Papio sushkini (Trofimov, 1977) (1), GIN U 987/878 (2), the upper jaws, and cranial fragment (Figure 9). Dating of the locality is disputable. Most proba- This taxon is represented by crania, lower jaws, and iso- bly, it is at the boundary of Late Ruscinian and Early Vil- lated teeth from Kuruk-Say, Tadjikistan. The specimens are 110 E.N. MASCHENKO ANTHROPOLOGICAL SCIENCE

Figure 9. Upper jaw and cranial fragments of Middle Pliocene Parapresbytis eohanuman from the Udunga locality, Transbaikalian, Russia. GIN U 987/445, right maxilla preserving P3–M3: (A) lingual view; (B) buccal view; (F) occlusal view. GIN U 987/878 (2), premaxilla preserving right I1 and left I1–2: (C) lingual view; (D) labial view. GIN U 987/878 (1), left premaxilla and maxilla preserving I1–P3: (E) lingual view; (G) labial view. GIN U 987/493 (1), cranial fragments: (H) dorsal view; (I) ventral view; (J) frontal view; (K) lateral view. Vol. 113, 2005 CENOZOIC PRIMATES OF EASTERN EURASIA 111

Figure 10. Lower jaws and two limb bone fragments of Parapresbytis eohanuman from Shamar Locality, northern Mongolia (from Boris- soglebskaya, 1981). PIN 3381–236, nearly complete mandibular corpus preserving right and left I1–M3: (A) occlusal view; (B) buccal view. PIN 3381-235, mandible: (C) occlusal view. PIN 3381–210, distal end of left humerus, and PIN 3381-211, right ulna: (D) humerus anterior view; (E) humerus posterior view. (F) ulna lateral view; (G) ulna anterior view. housed in the Paleontological Institute, Russian Academy of 709. Sciences, Moscow (Trofimov, 1977; Maschenko, 1989, Papio sushkini is a medium-sized extinct specialized rep- 1991b, 1994a). Specimens include the holotype female resentative of Papio. It is similar to recent Papio in morphol- skull, PIN 3120-523 (Figure 11A, B); a fragment of the max- ogy of the infraorbital foramina, ratio of lengths of the facial illa of a subadult male, PIN 3120-524; the lower jaw of an and cranial parts of the skull, the short and robust alveolar adult male, PIN 3120-526 (Figure 11C, D); the lower jaw of process of the upper jaw, the large anteroposterior diameter an adult male, PIN 3120-525 (Figure 11E, F); a fragment of of foramen magnum, the shape of the nasal bones, position the skull of a subadult individual, PIN 3120-524; isolated P3 of the fossa glandula lacrimalis (only on os lacrimale), shape 1 2 and M3, PIN 3120-707; isolated I and I , PIN 3120-708, of the cecal foramen, morphology of the CP3 complex, 112 E.N. MASCHENKO ANTHROPOLOGICAL SCIENCE

Figure 11. Skull and lower jaws of the Middle Pleistocene Asian baboon Papio sushkini from the Kuruk-Say locality, southern Tadjikistan. PIN 3120-523, female skull: (A) lateral view; (B) occlusal view. PIN 3120-525, male mandible: (C) lateral view; (D) occlusal view. PIN 3120-526, male mandible: (E) lateral view; (F) occlusal view (E and F are in the same scale).

molar morphology, and relation of lengths of the premolar Subfamily Cercopithecinae Gray, 1821 and molar tooth rows. Specialization of Papio sushkini in Macaca Lacepede, 1799 comparison with modern representatives of the genus is Macaca ex gr. anderssoni–arctoides found in a greater robustness of molars relative to premolars, greater enamel thickness, greater length of the zygomatic A macaque is known from the Middle Pleistocene Kudaro arch relative to total length of the skull, and a more robust 1 locality (Georgia), represented by isolated teeth, astragalus M3 in comparison with M2. (Figure 12), and a phalanx of finger 3 or 4. All macaque Distribution. Late Pliocene, Late Villafranchian (MN 16), remains come from the lower part of cultural layer 5, which the Kuruk-Say locality, southern Tadjikistan, Central Asia. is formed by yellow-brown loam. Layer 5 contains stone artefacts typologically referred to as Acheulean. Layer 5c Vol. 113, 2005 CENOZOIC PRIMATES OF EASTERN EURASIA 113

Figure 12. Middle Pleistocene M. ex gr. anderssoni–arctoides from Kudaro 1 Cave, northern Osetia, Georgia. ZIN 34600. right M2: (A) lingual view; (B) buccal view; (C) occlusal view. ZIN 34605, astragalus: (D) dorsal view.

(the southern gallery of the Kudaro cave) was dated by the Conclusions radiothermoluminescence method to 360 ka BP; and layer 5b was dated to 350 ka BP. Thus, layers 5b and 5c can be In the Cenozoic, the eastern Eurasian area under consider- referred to the warm phase of the Mindel interglacial epoch ation in the present paper was not the center of origin of a (interstadial IV of the Cromerian complex) (Vereshchagin, large number of primate taxonomic groups. The western 1959; Vereshchagin and Lubin, 1960; Delson, 1980; Veresh- area was influenced zoogeographically by the Mediterra- chagin and Baryshnikov, 1994). Molar morphology is closer nean; the eastern area was in contact with southern Asia, and to the Asian Macaca condition of the ‘sinica’ species group sometime in the Pliocene, with Africa. This explains the than to that of the European representatives of the genus abundance of either specialized, or widely distributed, forms Macaca (Maschenko and Baryshnikov, 2002). Structural of primates found in the Eurasian faunas. The total number features of the astragalar trochlea and the proportions and of known primate taxa is not great. From the Early Eocene to orientation of the head relative to the axis of the body show Pleistocene there are three families (Omomyidae, Homi- a great similarity mainly to terrestrial species. The macaque nidae, Cercopithecidae), seven genera (Altanius, Dryopithe- from Kudaro is larger than extant M. nemestrina. Among cus, Mesopithecus, Dolichopithecus, Papio, Parapresbytis, extinct species, it is smaller than only M. anderssoni from Macaca), and eleven species. the Middle Pleistocene of China. During the Late Miocene and Early Pliocene, owing to An important common dental feature of Asian macaques links between the Mediterranean and African provinces, rep- and the form from Kudaro is the bottom of the mesial and resentatives of both subfamilies of Cercopithecidae came to distal foveae, which are level with each other while the bot- inhabit the geographic area reviewed in this paper. From the tom of the trigon basin is positioned lower. In some species point of view of general modifications of mammalian fauna from Asia and in M. sylvanus, the bottom of the mesial fovea compositions, the appearance of Omomyidae is due to fau- is positioned higher than with the two other basins. Further- nal exchange between Asia and North America. Other lower more, in the macaque from Kudaro, and in extant M. nemes- primates never existed in this area. In the end of the Miocene trina and M. fuscata and extinct M. anderssoni from China, the extinction of representatives of Hominoidea coincides in the difference of the mesial and distal transverse diameters is time with the distribution of the Hipparion faunas. This may relatively small, and the size difference between M2 and M3 be due to global climatic change (establishment of seasonal is minimal. In these features, the Kudaro form shows a climate) in the end of the Miocene and distribution of open greater similarity to extant Asian macaques than to Pleis- landscapes (disappearance of rainforests). As in western tocene M. sylvanus. This testifies to its affinity with Asian Europe, at the western part of the area considered in this Macaca species. Tooth morphology allows us to propose review, the diversity of Cercopithecidae increased by the end that the sinica group is closer to the Kudaro form than the of the Miocene, overwhelming the diversity of Hominoidea. other Asian groups; and within this group, the closest spe- At the end of the Pliocene to the beginning of the Pleis- cies are M. arctoides and M. anderssoni. Thus, the most tocene, Cercopithecinae came to substitute Colobinae. From likely identification of the macaque from Kudaro is M. ex gr. this time onwards, the latter occurred only in the eastern part anderssoni–arctoides. of the area considered in this review, and later retreated to Distribution. Middle Pleistocene, Mindel interglacial southern Asia. Extinction of the few representatives of the epoch (interstadial IV of the Cromerian complex), the subfamily Cercopithecinae in this area during the Pleis- Kudaro 1 locality, northern Osetia, Georgia (Figure 1B). tocene is connected with the general fall of temperature in the northern hemisphere. During the Pleistocene, however, some species of Macaca existed in the southernmost areas, in a relatively more optimal environment. During the Late 114 E.N. MASCHENKO ANTHROPOLOGICAL SCIENCE

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