Pliocene faunas with Proboscideans of the Former Soviet Union

I.A. Vislobokova1, M.V. Sotnikova2 1Paleontological Institute of the Russian Academy of Science, Moscow, Russia - [email protected] 2Geological Institute of the Russian Academy of Science, Moscow, Russia - [email protected]

SUMMARY: In the territory of the former Soviet Union, remains of proboscideans are known in faunas from Moldova to the Transbaikal region. They belong to four genera and seven species of four fam- ilies: , , Gomphotheriidae and . During the early Pliocene, mammu- tid and mastodonts prevailed. The first elephantids of the Archidiskodon – Mammuthus lineage dispersed together with other inhabitants of open landscapes in the south of that territory in the middle Pliocene (MN 16). The members of that lineage became widespread in the south of Northern Eurasia begin- ning from the late Pliocene (MN17) after 2,5 Ma.

1. INTRODUCTION 2. EARLY PLIOCENE

Proboscideans were among the dominant The early Pliocene (Ruscinian) faunas of the groups in the Pliocene and mam- FSU lived in a more continental climate than malian faunas of Northern Eurasia. Data on that of Western Europe. Despite this, the their occurrences are traditionally used in the Ruscinian faunas of the European part of the biostratigraphy of these stages. FSU contained many inhabitants includ- The Pliocene was a time when the first mem- ing proboscideans, cercopithecid primates bers of the Archidiskodon-Mammuthus lineage (Macaca sp. and Dolichopithecus cf. ruscinen- dispersed into Eurasia. During a part of that sis), various muntiacines, pliocervines and oth- epoch they co-existed there with other pro- ers (Vangengeim et al. 1998). boscidean groups. The evolution of pro- Proboscideans were represented by three boscideans as well as of other herbivores was families: Deinotheriidae, Mammutidae and closely related to palaeoenvironments and Gomphotheriidae (Fig. 1). They were found in coincided with the large environmental the European part of the FSU and belonged to changes. The history of proboscideans in the three genera, , and Pliocene of Northern Eurasia reflects the , which appeared in the . increase of global cooling accompanied by the Remains of Deinotherium have been found enlargement of open woodlands in the temper- in the northern Ciscaucasus: D. cf. gigantissi- ate latitudes. mum near the Armavir town and Deinotherium In the territory of the former Soviet Union sp. in Kosyakino near the Stavropol town (FSU), remains of Pliocene proboscideans (Alekseeva 1977). are known from the numerous well-dated In the northern Black Sea area, a mammutid localities of Moldova, Ukraine, southern Zygolophodon borsoni (= Mammut borsoni) Russia, Georgia, Kazakhstan, Uzbekistan, was most abundant in the first part of the and Tadzhikistan. The data on proboscideans Ruscinian. The remains of this species are from the FSU offer important insights into the recorded in Ukraine (Novopetrovka and else- history of this group, its occurrences and dis- where) in the Kuchurgan Beds (MN 14 and ini- persals. tial part of MN 15) and in Moldova in the

157 The World of - International Congress, Rome 2001

Fig.1 - The main localities and occurrences of proboscideans in the Pliocene of the FSU.

“Moldavian Russilion” (MN 15) (Vangengeim is also indicated at this time. et al. 1998). A gomphothere Anancus was more common 3. MIDDLE-LATE PLIOCENE in the south of European part of the FSU dur- ing the late Ruscinian. Its dispersal in this ter- 3.1 Early Villafranchian ritory coincided with the appearance of some inhabitants of open landscapes due to more arid At the Early/Middle Pliocene (Ruscinian/ climatic conditions than in the first half of the Villafranchian) transition, the number of inhab- Ruscinian. The remains of Anancus arvernen- itants of open landscapes markedly increased. sis occurred in Moldova (Etuliya, Luchesty During the middle and late Pliocene, and others) in the Karboliya Beds (late part of mastodonts were gradually replaced by ele- MN 15) and in the northern Ciscaucasus phantids. (Kosyakino) (Vangengeim et al. 1998). In the The early Villafranchian faunas of the FSU northern Black Sea area, that species co-exist- are characterised by the last occurrence of ed with camels Paracamelus, which dispersed some warmth-requiring forest elements includ- there again after its first appearance there in the ing Zygolophodon, a procyonid (Parailurus), a Messinian time (Vislobokova et al. 2001). The large badger (Parameles), and a further radia- appearance of a small canid Eucyon odessanus tion of Eucyon-like canids; and by the first 158 Pliocene faunas with proboscideans of the Former Soviet Union appearance of a number of boreal forms. 1988). In Europe, Anancus persisted until the The most north-eastern finding of early Pleistocene (Göhlich 1999). Zygolophodon occurred in Transbaikal region The elephantids of the Archidiskodon- in the Udunga fauna (MN 16), one of the most Mammuthus lineage became widespread representative faunas of this age in the Asian together with other adapted to savanna- part of the FSU. The Udunga fauna also con- like conditions: Canis etruscus group, tains a cercopethecid primate Parapresbytis, a Pliocrocuta, Homotherium, Acinonyx, Equus, small bear Ursus cf. minimus, and the first Paracamelus, Elasmotherium and others. A. boreal forms (Gulo, Capreolus and others) gromovi was found in a number of localities in (Kalmykov 1992, Sotnikova & Kalmykov the south of European Russia (Khapry, 1991, Vislobokova et al. 1993, 1995; Liventzovka and others), in Kazakhstan Vislobokova et al. 2001). (Podpusk-Lebyazh’e, Kopaly, Tadzhikistan Anancus continued to exist in some refugia in (Kuruksay) and in Uzbekistan (Kairakkum)) the south of Northern Eurasia. The was (Baigusheva 1971; Sotnikova et al. 1997; represented by A. arvernensis in the Caucasus Vangengeim et al. 1988; Vislobokova 1996). (Kvabeby) and by A. kazakhstanensis in the In the faunas of the terminal Pliocene of the Tekess Depression in southern Kazakhstan FSU, mastodonts are unknown and A. gromovi (Esekartkan and Ajgyrzhal). Mastodonts is replaced by A. meridionalis. occurred there together with progressive hip- parions. 4. FINAL REMARKS The first records of the Archidiskodon– Mammuthus lineage, of African origin, are A large diversity of proboscideans was typi- observed in the south-western FSU in middle cal of late Ruscinian faunas in the territory Akchagylian times. But Archidiskodon could of the FSU, characterized by the presence have arrived there before that time. Alekseeva of deinotheres, and mammutid (1977) supposed that A. cf. meridionalis from mastodonts and, possibly, by the first appear- Kosyakino could be A. gromovi but this pre- ance of the Archidiskodon-Mammuthus line- sumption needs to be checked. In Italy, the age. entry of Archidiskodon occurred at the second The main turnovers of proboscideans in this half of the early Villafranchian (Montopoli) territory coincided with a wide spread of open (Azzaroli et al. 1988). In Roumania, Anancus landscapes at the early Pliocene/middle arvernensis and a primitive Archidiskodon are Pliocene transition (about 3,5 Ma) and at the reported from the Dacic Bassin ( MN 16b) middle Pliocene/late Pliocene transition (about (Tersea et al. 1997). In China, a record of 2,5 Ma). In the middle Villafranchian a primi- Archidiskodon-Mammuthus lineage in the tive Archidiskodon - A .gromovi - became dom- Yushe Basin is referred to MN 15 zone inant; in some regions these elephants coexist- (Tedford 1995). ed with Anancus. At the end of Pliocene proboscideans in the 3.2 Middle Villafranchian FSU were represented by the single family Elephantidae (A. meridionalis). The middle Villafranchian faunas of the FSU are characterized by the diversity and last 5. ACKNOWLEDGEMENTS occurrence of Anancus, and a wide distribution of elephantids. A. arvernensis is reported from We thank Dr. Adrian Lister for reading the Ukraine (Zhevakhova gora) (Alekseeva 1977). text and for his comments. A progressive A. alexeevae was discovered in The study was supported by the Russian the south of European Russia (Liventsovka) Foundation for Fundamental Research (proj- (Baigusheva 1971). A. kazakhstanensis is pres- ect 99-04-48636, 99 –05-64150 and 00-15- ent in Kazakhstan (Adyrgan) (Tleuberdina 97754). 159 The World of Elephants - International Congress, Rome 2001

6. REFERENCES malienne du Cénozoïque en Europe et domaines reliés: 649-660. Actes du Alekseeva, L.I. 1977. Early Anthropogene the- Congrès BiochroM’97 Montpellier. riofauna of East Europe. Moscow: Science. Tleuberdina, P.A. 1988. Main localities of hip- Azzaroli, A., De Giuli, C., Ficcarelli, G. & parion fauna and their biostratigraphic cor- Torre, D. 1988. Late Pliocene to early mid- relation. In E.I. Gavrilov (ed.), Pleistocene in Eurasia: faunal Interregional comparison of faunas and flo- succession and dispersal events. ras of Mezozoy and Cainozoy of Palaeogeogr. Palaeoclimatol. Palaeoecol. Kazakhstan: 38-73. Alma-Ata: Science. 66: 77-100. Vangengeim, E.A., Sotnikova M.V., Baigusheva, V.S. 1971. Fossil theriofauna of Alekseeva, L.I., Vislobokova, I.A., the Liventzovka sand-pit. Transaction of Zhegallo, V.I., Zazhigin, V.S. & Shevyreva, Zoological Institute, 49: 5-29. N.S. 1988. Biostratigraphy of Late Pliocene Göhlich, U.B. 1999. Order . In – Early Pleistocene of Tadzhikistan (based G.E. Rössner & K. Heissig (eds.), Land on mammalian fauna). Moscow: Science. mammals of Europe: 157-168. München: Vangengeim, E.A., Vislobokova, I.A. & Verlag Dr. F.Pfeil. Sotnikova, M.V. 1998. Large Ruscinian Kalmykov, N.P. 1992. Biostratigraphy and Mammalia in the territory of the former mammalian fauna of the Pliocene of Soviet Union. Stratigraphy and Geological Transbaikal. Novosibirsk: Science. Correlation 6(4): 368-382. Sotnikova, M.V., Dodonov, A.E. & Pen’kov, Vislobokova, I.A. 1996. The Podpusk- A.V. 1997. Upper bio-magnetic Lebyzh’e mammalian faunas and assem- stratigraphy of Central Asian mammalian blage, Western Siberia. Palaeontographia localities. Palaeogeogr. Palaeoclimatol. Italica 83 (1-5): 1-23. Palaeoecol. 133: 243-258. Vislobokova, I., Dmitrieva, E. & Kalmykov, N. Sotnikova, M.V. & Kalmykov, N.P. 1991. 1995. Artiodactyls from the late Pliocene of Pliocene Carnivora from Udunga locality, Udunga, Western Trans-Baikal, Russia. J. Transbaikal, SSSR. In E.A. Vangengeim Vertebrate 15(1): 146-159. (ed.), Pliocene and Anthropogene palaeo- Vislobokova, I.A., Erbaeva, M.A & Sotnikova, geography and biostratigraphy: 146-160. M.V. 1993. The early Villafranchian stage Moscow: Geological Institute. in the development of the mammalian fauna Tedford, R.H. 1995. Neogene mammalian bios- of Northern Eurasia. Stratigraphy and tratigraphy in China: past, present and Geological Correlation 1(5): 555-564. future. Vertebrate Palasiatica, 33 (4): 277- Vislobokova, I.A., Sotnikova, M.V. & 289. Dodonov, A.E. 2001. Late Miocne - Tersea, E. 1997. Biochronologie du Pliocène Pliocene mammalian faunas of Russia and du bord méridional du Bassin Dacique neighbouring countries. Bolletino della So- (Roumanie). In J.P. Aguilar, S. Legendre & cietà Paleontologica Italiana 40: 307-313. J. Michaux (eds.), Biochronologie mam-

160