The World of Elephants - International Congress, Rome 2001

Elasmotherians - evolution, distribution and ecology

N.G. Noskova Moscow State University, Moscow, Russia - [email protected]

SUMMARY: Elasmotherians from an Early , Tamanian assemblage at Sinaya Balka ( area, Russia) were studied in order to (1) assess the evolution and distribution of this , and (2) evaluate the likelihood that caucasicum is a bona fide species. In this region, Elasmotherium caucasicum Borissjak, 1914, is commonly associated to the Tamanian elephant (Archidiskodon meridionalis tamanensis). It has a more primitive dental morphology than E. sibiricum Fisher, 1808, with which it was formerly considered to be synonymous, and therefore constitutes a separate species. Ecological peculiarities and paleoenvironmental conditions, proposed for E. caucasicum, could be useful for understanding the biogeography of the region at the time of Archidiskodon meridionalis tamanensis.

1. INTRODUCTION basin is typical for slightly worn teeth only. Two more species, which represent the earli- Elasmotheriums are giant est record of the genus, were described from which lived in the of by Chow (1959): E. inexpectatum and E. Southern Russia. After the Tamanian elephant peii (Shansi). Chinese species were considered (Archidiskodon meridionalis tamanensis), by Chow (1959) to be ancestral for E. cauca- Elasmotherium caucasicum is the most fre- sicum and E. sibiricum, which were widely dis- quently found mammal in the Tamanian assem- tributed in the Eastern Europe (e.g., Russia), blage. They form the most typical assamblage there being only two questionable records of of large for the interval 1.1-0.8 My. Elasmotherium sp. thus far recorded in western This assemblage is found in Sinaya Balka and Europe (Shvireva 1995). Tsymbal localities. Moreover, Elasmotherium Investigation of new material from an Early is known from Akhtanisovskaya and Pleistocene Tamanian mammal assemblage at Fontalovskaya localities and Archidiskodon Sinaya Balka, near the Black Sea, shows that E. from Kuchuguri locality (Fig. 1). caucasicum differs from E. sibiricum by its rel- atively longer teeth-row and P4, bigger teeth, 2. DISCUSSION and an s-shaped metaconid. It is likely that E.caucasicum is ancestral to E. sibiricum as it A new genus and species, Elasmotherium retains some primitive dental features, notably sibiricum, was established by Fisher von (1) a postfossette basin on the upper teeth, (2) a Valdgeim (1808). A second species, E. cauca- less sinuous enamel layer of the external wall sicum was distinguished by A. Borissjak (1914) of ectoloph, (3) irregular fold of the enamel on the basis of its large and morphologically- (the peaks of the fold have the different height). distinct teeth from Sinaya Balka. E. cauca- The genus Elasmotherium appeared in sicum was considered to have more primitive during the Late . Its ori- “-like” teeth morphology, due to gin appears to be connected to the genus presence of the postfossette basin on the upper (Upper – Lower teeth. The validity of this species, however, was Pliocene), although certain species of subsequently challenged by V.A. Terjaev Sinotherium, which could be ancestral to (1948), who equated it with E. sibiricum on the Elasmotherium, is unknown. E. inexpectatum grounds that the presence of the postfossette and E. peii inhabited Eastern China during the 126 Elasmotherians - evolution, distribution and ecology

Fig.1 - The distribution with the localities of Tamanian mammal assemblages.

Upper Pliocene - Lower Pleistocene. They dis- view portrays them as large wooley appeared c. 1.6 Ma ago (Qui & Qui 1995). The with a large forehead horn that thrived on an earliest records of Elasmotherium sp. in Russia open . The other view, proposed by V.A. are known from the Upper Pliocene assem- Terjaev (1948), assigned elasmotherians to blages near the Black Sea. This area evidently riparian biotopes. became a second center of their radiation. This Our work indicates that elasmotherians dwelt suggests that Elasmotherium sp. may have in both riparian and steppe biotopes. The ripar- developed separately in Russia and China. ian biotope is suggested by dental and skull E. caucasicum was widely distributed in this morphology. The combination of such charac- area 1.1 Ma and 0.8 Ma. The more advanced E. ters as the absence of canines and strongly sibiricum appeared in the Middle Pleistocene. developed lateral processes of the atlas implies It occupied all of the southwestern part of lateral movements of the head, presumably for Russia, reaching eastward to western Siberia. grasping grass. The hypsodont dentition indi- Elasmotherians persisted in the eastern Europe cates presence of mineral grains in the food. untill the end of the Middle Pleistocene. The Such food could be obtained by pulling out records of elasmotherians in the western dense plants from the moist soil. These condi- Europe, however, is problematic, thus the west- tions are typical for riparian biotopes. On the ern limit of their migration remains to be docu- other hand, a steppe biotope is indicated by mented. their rather long and slender limbs, which Morphological peculiarities of elasmotheri- would have served well for creatures ans have generated two main hypotheses con- over vast areas. Thus, the available data suggest cerning their appearance and the character of that the habitats of elasmotherians could their habitat. The first, most widely accepted haveincluded both riparian biotopes and the

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adjacent high grass steppe. Given that elas- 4. REFERENCES motherians coexisted with Archidiskodon, the same ecological conditions can be inferred for Borissjak, A.A. 1914. The teeth structure of the latter. Elasmotherium caucasicum n. sp. Izvest. Akad. Nauk SSSR, 6(9): 555-584. [in 3. CONCLUSION Russian]. Chow, M. 1958. New elasmotherine rhinoceros Sinaya Balka (Early Pleistocene, Black Sea from Shansi. Vertebrata Palasiatica 2(2-3): area, Russia) is the type locality of Tamanian 135-142. mammal assemblage as well as of Elasmo- Fischer von Waldheim, G. 1808. La notice d'un therium caucasicum and Archidiskodon merid- fossile de Siberie inconnu aux natu- ionalis tamanensis. The Early Pleistocene E. ralistes. Programme d'invitation a la seance caucasicum is confirmed as a valid species. It publique de la Societe imperiale des natu- may be regarded as a likely morphological ralistes: 23-28. ancestor for E. sibiricum due to its primitive Qiu, Z. & Qiu, Z. 1995. Chronological sequence dental characteristics. E. caucasicum was and subdivision of Chinese Neogene dispersed in the upper part of the Early Mammalian faunas. Palaeogeography, Pleistocene. Its upeer stratigraphical range is Palaeclimatology, Palaeoecology 116: 41-70. limited by Matuyama chron. The assemblage Shvireva, A.K. 1995. History of the rhinoceros E. caucasicum and A. meridionalis tamanensis Elasmotherium: 103 pp. Stavropol. [in is of characteristic this time interval in south- Russian] western Russia. Terjaev, V.A. 1948. Geological position of the Habitats of elasmotherians included both forehaed-crested rhinoceros (Elasmotherium). riparian biotopes and contiguous high grass Sovetskaya geologiya 34: 81-87. [in Russian] .

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