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New postcranial bones of elasmotherium peii from shanshenmiaozui in Nihewan basin, Northern China

Nouveaux os post-crâniens d’elasmotherium peii de shanshenmiaozui du bassin de Nihewan, Chine du Nord
Hao‑wen Tong, Xi Chen et Bei Zhang
p. 195-204


Elasmotherium ou la Licorne géante de Sibérie est assez commune en Sibérie et dans les régions adjacentes pendant le Quaternaire, mais elle est rarement observée dans le Pléistocène inférieur de Chine du Nord. Au siècle dernier, un certain nombre d’ossements d’Elasmotherium ont été récoltés dans le bassin de Nihewan, qui sont pour la plupart des éléments post-crâniens isolés, parmi lesquels le tibia était absent. Récemment, du matériel dentaire, mais aussi un tibia presque complet et quelques os du tarse ainsi que des os métapodiaux d’Elasmotherium ont été découverts sur le site de Shanshenmiaozui, dans le bassin de Nihewan. La morphologie de ces ossements est tout-à-fait distincte de l’espèce russe, surtout dans la forme du Mc III, de l’astragalus et du cuboïde. Les dimensions restent néanmoins proches ou légèrement inférieures à celles de l’Elasmotherium russe et ces ossements ont été attribués à l’espèce E. peii. Les nouveaux os récupérés sont aussi radicalement différents de ceux d’autres rhinocéros fossiles en particulier par leur très grande taille, la morphologie du troisième métacarpien et de l’astragale. L’espèce E. peii a bien coexisté avec Canis chihliensis, Pachycrocuta sp., Mammuthus trogontherii, Coelodonta nihowanensis, Proboscidipparion sp., Equus sanmeniensis, Sus lydekkeri, Paracamelus sp., Eucladoceros sp., Spirocerus wongi, Gazella sinensis et Bison palaeosinensis, et la localité peut être datée de plus de 1,2 Ma et représente un environnement de steppe à buissons.

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The authors would like to thank the following persons : Prof. Wei Q. for providing information on the SSMZ site ; Prof. Shvyreva A.K. for providing the data of tibial measurements of E. sibiricum; Dr. A.-M. Moigne for translating the abstract into French; Dr. Titov V. V. for providing access to literature; Dr. Argot C. of MNNH and Mrs. Zheng M. of TPN for providing access to the collections. Profs. Qiu Z.-X., Deng T. and Kahlke R.-D. for constructive suggestions and fruitful discussions. Dr. Antoine P.-O. and another anonymous reviewer for manuscript review. Drs. Faure M. and Debard É. for organizing this special issue. This work was supported by the following grants: The National Natural Science Foundation of China (Grant No. 41572003), Key Research Program of the Chinese Academy of Sciences (Grant No : KZZD-EW-15), Special Basic Research Project (Grant No : 2014FY110300) of MST of China. This paper is written in commemoration of Prof. Guérin C.

1 – Introduction

1Elasmotherium is the most specialized taxon among the rhinocerotids, whose fossil remains are very scanty in China as only isolated teeth and a few postcranial bones from some sites in northern China were reported in the past century (Teilhard de Chardin & Piveteau, 1930; Teilhard de Chardin, 1942; Chow, 1958; Chia & Wang, 1978; Huang & Ji, 1984; Deng & Zheng, 2005), most of which lack a definite provenance.

2In the recent years, the excavations at the Shanshenmiaozui (SSMZ) Site and Heitugou Site resulted in some recoveries of Elasmotherium fossils, some of which have been reported by Tong et al. (2014). The more recently recovered specimens from SSMZ are going to be described in this study. The significance of the new finds lies at their definite provenance and the first recovery of the tibia material of Elasmotherium in China.

3The SSMZ site lies at the neighboring hill slope of Xiaochangliang, a well-known Paleolithic site in Nihewan Basin, Hebei Province in northern China (fig. 1). Based on the stratigraphic correlation in the field, the fossil-bearing sand-silt bed at the new locality is a little higher than the cultural layer at Xiaochangliang site (Tong et al., 2011; Liu et al., 2016), whose paleomagnetic age is about 1.36 Ma BP (Zhu et al., 2001).

4The mammalian taxa associated with Elasmotherium peii at SSMZ are as follows: Lepus sp., Ochotona sp., Canis chihliensis, Pantherinae gen. et sp. indet., Pachycrocuta sp., Mammuthus trogontherii, Coelodonta nihowanensis, Proboscidipparion sp., Equus sanmeniensis, Sus sp., Eucladoceros sp., Spirocerus wongi, Gazella sinensis and Bison palaeosinensis (Tong et al., 2011), which represents a shrub-steppe environment; furthermore, a metatarsal bone of Paracamelus was recovered in 2015 and a metacarpal bone of Alactaga was recovered in the 2016 season.

5The SSMZ site bears quite rich and well-preserved fossils, such as Canis chihliensis (Tong et al., 2012), Coelodonta nihowanensis (Tong & Wang, 2014), M. trogontherii (Tong, 2012; Tong & Chen, 2016; Chen & Tong, 2016) and B. palaeosinensis (Tong et al., 2016), which formed the early assemblage of the Mammuthus-Coelodonta faunal complex.

Fig. 1: Spatial (A) and temporal (B) distributions of Elasmotherium, with details of the Chinese sites (C) of E. peii.

Fig. 1: Spatial (A) and temporal (B) distributions of Elasmotherium, with details of the Chinese sites (C) of E. peii.

1/ Xiashagou; 2/ SSMZ; 3/ Heitugou; 4/ Xihoudu; 5/ Sanmenxia; 6/ Shanxi but without location details.

Data for Elasmotherium sites outside China are after Shvyreva (2016)

2 - Fossil materials

6The fossil materials are given in table 1.

Tab. 1: Recently recovered fossils of Elasmotherium peii from Nihewan Basin.

Tab. 1: Recently recovered fossils of Elasmotherium peii from Nihewan Basin.

3 - Methods, terminology and abbreviations

7The classification system above the species level is after Antoine (2002). The terminology is mainly after the following references: Guérin (1980), Antoine (2002) and Prothero (2005). The method of measurements is according to the following authors: Guérin (1980), Mazza (1988), Antoine (2002) and Prothero (2005). All the dimensions are in millimeter.

8The abbreviations are given below:

9- Institutional abbreviations―IVPP, Institute of Vertebrate Paleontology and Paleoanthropology; MNHN, Muséum National d’Histoire Naturelle; NHW, Nihewan (= Nihowan); SSMZ, Shanshenmiaozui; TNP, Tianjin Natural History Museum (= Musée Hoang ho Pai ho).

10- Anatomical abbreviations―H, height; L, length; W, width; DT prox, proximal transverse diameter; DAP prox, anteroposterior diameter of the proximal end; DT dia, transverse diameter of the shaft; DAP dia, anteroposterior diameter of the shaft; DT dist, distal transverse diameter; DAP dist, anteroposterior diameter of the distal end; DT artic dist, transverse diameter of the distal articular surface; DAP arti dist, anteroposterior diameter of the distal articular surface.

4 - Systematic paleontology

4.1 – Systematics

11Mammalia Linnaeus, 1758

12Perissodactyla Owen, 1848

13Rhinocerotidae Owen, 1840

14Subfamily Elasmotheriinae Bonaparte, 1845

15Genus Elasmotherium Fischer von Waldheim, 1808

16Elasmotherium peii Chow, 1958

171930. Elasmotherium sp.–Teilhard de Chardin & Piveteau, p.29-30.

181958. Elasmotherium inexpectatum–Chow, p.139, Pl I-B.

191958. Elasmotherium cf. inexpectatum–Chia & Wang, p. 35–37, Pls. XIV-1 and XV-1.

202002. Elasmotherium caucasicum–Antoine, p.43-45. Partial.

212005. Elasmotherium caucasicum–Deng & Zheng, p.110-121.

4.2 - Comparative descriptions

4.2.1 - Ulna

22The distal end and the olecranon are not preserved, and the anconeal process is partially broken; the trochlear notch and most part of the shaft are preserved. It distinguishes itself from other Quaternary rhinos by a larger size. In general, the shaft is not so strongly bent backward and the mid-shaft has a triangular cross section. The anterior surface of the proximal part is strongly flared medially, which makes the articulation facet much wider (fig. 2: 1a; tab. 2). The surface of the semi-lunar notch is bifid at the lower part, and the medial part is oblique and strongly projecting medially, while the external part runs parallel to the sagittal plane of the shaft. Between the two branches of the trochlear facet, there exists the pit-like radial notch. There is a notch at each edge of the anterior face just below the trochlear surface, while the lateral one has a higher positioned than the medial one, and both of them should be corresponding with the interosseous space of forearm (spatium interosseum antebrachii proximale). Just below the medial notch, there exists a prominent nutrient foramen. The antero‑medial edge is forming a sharp ridge. Both the medial and lateral surfaces are quite smooth and without any prominent structures, except the longitudinal trench along the antero‑medial ridge at the medial surface.

Fig. 2: Forelimb bones of E. peii from Shanshenmiaozui

Fig. 2: Forelimb bones of E. peii from Shanshenmiaozui

1/ Left ulna (V23590.1); 2/ left magnum (V23590.4); 3/ right Mc III (V23590.2); 4/ right Mc IV (V23590.3). 1a, 2a, 3a and 4a, anterior views; 1c, 2d and 4c, lateral views; 1b, 2b, 3b and 4b, posterior views; 1d, 2c, 3c and 4d, medial views; 3e and 4e, proximal views.

4.2.2 - Magnum

23It resembles the magnum of other Quaternary rhinos except its larger size (fig. 2: 2a-2d; tab. 2). In anterior view, the upper and lateral margins of the front face are relatively straight, while the inferior margin is roundish. The whole surface is convex and with some nutrient foramina. In dorsal view, three facets can be observed, scaphoid and unciform facets in the front, and the lunar facet is located higher and more backward, after which is the elongated posterior tubercle (or volar process), and it is quite straight rather than bended as in other Quaternary rhino taxa. In lateral view, the unciform and the Mc III as well as the lunar facets can be observed; the unciform facet is slope-like, while the Mc III is deeply concave and the lunar facet is convex and roundish; the volar process is beak-like and its inferior border is deeply concave. In medial view, the trapezoideum facet can be observed, which is continuous with the Mc III facet; the concavity of the Mc III facet is much shallower than those of the primitive elasmothere taxa (Antoine, 2002: p.200-201). In ventral view, only the concave Mc III facet and the volar process can be observed, the antero-medial corner of the facet is roundish rather than rectangular as in E. sibiricum. Between the facets, the surface is rough and with some nutrient foraminae. It differs from that of E. sibiricum in the invisibility of the Mc III facet in lateral view, and the more deeply concave Mc III facet and the inferior notch of the volar process.

Tab. 2: Measurements of postcranials of Elasmotherium peii, compared with related taxa (in mm).

Tab. 2: Measurements of postcranials of Elasmotherium peii, compared with related taxa (in mm).

4.2.3 - Metacarpal III

24It is somewhat larger than those of other Quaternary rhinos and is also slightly different in morphology (fig. 2: 3a-3e; tab. 2). It is broad, flat and straight, and with expanded extremities. There are five facets at the proximal end: the medial banded facet is tiny and in contact with Mc II, the biggest facet contacts magnum, the lateral top facet contacts with unciform, the lateral and posterior facets articulate with Mc IV. In dorsal view, the magnum facet is deeply concave and the unciform (uncinate) facet prominently elevated; the distal articular surface is lower than that of E. sibiricum. In posterior view, the outline of the distal articular surface is pointed rather than roundish as in E. sibiricum. The specialties of the Mc III of E. peii lie at the antero-posteriorly expanded distal end and the much lower situated Mc IV lateral facet as well as the upward tapering of the postero-distal trochlea.

4.2.4 - Metacarpal IV

25It is obviously smaller than Mc III; the body is obviously bent, with a pronounced notch on the lateral side (fig. 2: 4a-4e; tab. 2). There are four facets at the proximal end: the biggest one is in contact with unciform, the two medial facets articulate with Mc III, but the lateral facet is in contact with the vestigial Mc V. This Mc IV specimen matches well with the Mc III specimen described above, and it is likely that they belong to the same individual. The broken shaft shows that there is no medullary cavity in Mc IV. The lateral Mc V facet is quite large, which means the Mc V is quite big, compare to the rudimentary ones in the extant rhino species and in the woolly rhinos (Coelodonta spp.).

4.2.5 - Tibia

26The nearly complete tibia only with partial proximal end (cnemial crest and the lateral condyle) broken off (fig. 3: 1a-1e; tab. 2) represents the only tibia specimen of Elasmotherium ever discovered in China. The tibia is obviously larger than those of other Quaternary rhinos. The shaft is not a trihedral form as in the ordinary rhinos, but with a nearly roundish cross section, and it becomes broad and antero-posteriorly compressed near the distal end.

27The anterior part of the proximal end is nearly completely broken off. It seems that the cnemial crest only occurs at the upper third part, but their details are uncertain; the situation of the tuberosity groove (for patellar ligament) is not clear. In proximal view, only the articular facet of the medial condyle remains, which surface is not convex or slope-like as in the ordinary rhinos, but slightly concave as in the elephant. In posterior view of the proximal end, there exists a small isolated facet behind the medial condyle, which is facing to the popliteal notch and hanging above the popliteal fossa (as indicated by the arrow in fig. 3:1b), but its function is still uncertain. The popliteal fossa is slightly concave, the popliteal line and the muscular lines are not too prominent, and the nutrient foramen is not detected. The medial outline is not as straight as in Coelodonta, but slightly concave instead. At the distal end, the inferior margin is wave-like in anterior view; at the medial-posterior side, a tongue-shaped process erupts and extends downward, but not as far as in E. sibiricum, and it is housed in the saddled groove of astragalus. In distal view, the outline is almost rectangular and with the posterior edge slightly shorter; the articular surface is composed of two longitudinal grooves and an intermediate ridge, all of which contact with the astragalus; the medial groove is relatively narrow and deep; the intermediate ridge is nearly anteroposteriorly oriented. In lateral view, both the proximal and the distal articular surfaces incline anteriorly; most part of the lateral face, except the upper third, has rough surface, which means the fibula once co-ossified with the tibia, and partial fibula still remains on the tibia near the distal end. In medial view, the surface is smooth and without any special structure.

Fig. 3: Hindlimb bones of E. peii from Shanshenmiaozui

Fig. 3: Hindlimb bones of E. peii from Shanshenmiaozui

1/ Right tibia with partial fibula (V23590.6); 2/ left astragalus (V23590.7); 3/ right cuboid (V23590.8); 4/ partial left Mt IV (V23590.9); 5/ the distal phalanx of toe III of the pes (V23590.11). 1a, 3a and 4a, anterior views; 1b, posterior view; 1c, 3e and 4c, medial views; 1d, 3d and 4d, lateral views; 2a, 3a and 5a, dorsal views; 2b, 4b and 5b, plantar views; 1e, 2c and 3c, distal views; 3b, 4e and 5c, proximal views.

4.2.6 - Astragalus

28The astragalus is almost intact, which is prominently larger than those of the usual rhinocerotids (fig. 3: 2a-2c; tab. 2). In dorsal view, both of the superior and inferior borders are saddle-like, but the saddles are less proununced as in E. sibiricum; the trochlea is not symmetric with the lateral ridge being much broader and slightly higher than the medial one. The medial and lateral ridges are parallel to each other, and both of them are parallel to the long axis of the limb bone; the fibular facet on the extremely lateral part is moderately developed. The neck of the astragalus is shorter than those of other Quaternary rhinocerotids and higher medially than laterally, and the medial side bears a prominent tuberosity.

29In plantar view, three distinct articular surfaces can be observed: the proximal calcaneal, sustentacular, and distal calcaneal facets, and there exists a narrow groove between the former two facets, which is different from that of the E. sibiricum. The proximal calcaneal facet is the largest and with a transverse trench. The sustentacular facet is a small oval facet, but is not as flat as in other rhinos. Whereas the distal calcaneal facet looks like a drop of water and is confluent with the cuboid facet of the distal end. The two calcaneal facets are not so distantly separated as in other rhinocerotids. In lateral view, lateral part is strongly antero-posteriorly compressed, and the fibular facet extends along the anterior border. In distal view, two facets can be distinguished, a much larger medial one for the navicular and a narrower lateral one for the cuboid. The former is mainly transversely convex but antero-posteriorly concave at the anterior part; the lateral facet is flat to slightly convex. The two facets consist of a large irregular four-sided polygonal continuous surface, and the anterior edge of which is the longest and wave-like, the latero-posterior edge is the second longest and nearly straight. Concerning the astragali of E. caucasicum, Schvyreva (2016) proposed that they are larger and slenderer, and the distal ends are narrower than those of E. sibiricum and E. chaprovicum.

4.2.7 - Cuboid

30The cuboid’s dorsal face looks cubical and full of nutrient foramens (fig. 3: 3a-3e; tab. 2). The proximal facet is slightly transversely concave and composed of two facets : the lateral calcaneum facet and the medial astragalus facet. In medial view, the larger upper facet is in contact with the navicular, and the bottom facet which is confluent with the distal Mt IV facet is articulating with the ectocuneiform. The volar process is quite pronounced. In distal view, the only articular facet with Mt IV and the volar process can be observed; the facet has a general triangular outline but with a bulge at the medial edge, which is different from that of E. sibiricum.

4.2.8 - Mt IV

31The available Mt IV specimen is obviously smaller than Mt III and only with the proximal part preserved (fig. 3: 4a-4e; tab. 2). The dorsal face is rough and with nutrient foramens at the upper part. In proximal view, the saddled facet for cuboid is antero-posteriorly concave and with a triangular outline. In medial view, there are two roundish facets for Mt III, the upper one is larger and confluent with the cuboid facet; there is a groove between the two facets. The posterior part of the proximal end is protruded prominently. The broken shaft shows that there is no medullary cavity in Mt IV.

4.2.9 - Ungual (third) phalanx of digit III

32Nearly intact except but broken at two sides (fig. 3: 5a-5c; tab. 2). In proximal view, the articular facet for the 2nd phalanx is slightly convex in the middle part and concave at two sides, and the outline is a narrow rectangle, which means the anterior and posterior surface are nearly parallel. The anterior surface is very rugose, and full of grooves at the distal part and tubercles at the proximal part, between which there exists a concave belt. In posterior view, the superior edge is smooth and elevated in the middle point; the distal part is full of nutrient foramens, and the distal edge is uneven.

5 - Comparisons and discussions

33The genus Elasmotherium was established by Fischer von Waldheim in 1808, and the type species is Elasmotherium sibiricum. Borissiak (1914) erected the other species Elasmotherium caucasicum which is characterized by the existence of postfossette on the upper teeth and was regarded as more primitive (Noskova, 2001); but other scholars thought that postfossette will definitely appear in the early stage of abrasion of the tooth and the latter is a synonym of the former (Teryaev, 1948). Investigation of new material from an Early Pleistocene Tamanian mammal assemblage at Sinaya Balka, near the Black Sea, shows that E. caucasicum differs from E. sibiricum by its relatively elongated teeth-row and P4, bigger teeth, and an S-shaped metaconid. It is likely that E.caucasicum is ancestral to E. sibiricum as it retains some primitive dental features, notably (1) a postfossette basin on the upper teeth, (2) a less sinuous enamel layer of the external wall of ectoloph, (3) irregular fold of the enamel (the peaks of the fold have the different height) (Noskova, 2001).

34The first acknowledgement of the Elasmotherium fossils from Nihowan Basin of Hebei Province in North China was reported by Teilhard de Chardin and Piveteau (1930). In 1958, Chow established two new species: Elasmotherium inexpectatum Chow, 1958 and E. peii Chow, 1958 based on specimens from Shanxi Province in North China. The recently recovered teeth and foot bones of Elasmotherium from Shanshenmiaozui and Heitugou represent the best fossil materials ever recovered in Nihewan Basin. Especially the tooth specimens represent the first discovery in Nihewan Basin. Based on the dental morphology and dimensions, the elasmothere fossils recently discovered can be referred to the species E. peii which was regarded as a valid species name, and all the Early Pleistocene elasmothere fossils in China can be included in this species (Tong et al., 2014). Therefore, the associated skeletal materials of this study also can be referred to the species E. peii, which can be distinguished from the Eurasian species E. caucasicum in quite a number of dental characters (Tong et al., 2014), but the comparisons in postcranial skeletons are not available, because of the shortage of fossil materials; up to date, only the 6th cervical vertebra, scapula, humerus, radius and astragalus have been acknowledged for E. caucasicum (Schvyreva, 2016), and only partial ulna and radius, tibia and the extremities have been known for E. peii.

35Compared with other Quaternary non-Elasmotherium rhinos, the E. peii from Nihewan is different not only in its larger size, but also in the following morphological features: tibia with more straight posterior edge and less developed intercondylar tongue of the distal end and a more roundish cross section of the shaft; the astragalus has a shorter neck with the proximal and the distal calcaneal facets not too widely separated; the Mc III has a much lower positioned lateral facet for Mc IV; the cuboid has a less developed volar process.

36The tibia of Elasmotherium is the longest among all the rhinocerotids, which is only slightly shorter but stouter than that of the lesser giant rhinoceros Juxia sharamurenensis (Qiu & Wang, 2007) and even longer than that of some mammoth (Ziegler, 2001). Therefore, it is certain that Elasmotherium was the largest rhino during the late Cenozoic, whose body size is as large as a male Asian elephant (Prothero et al., 1989). Shvyreva (1995) compared the length of the major segments of the limb bones of Elasmotherium with the counterparts of other rhinos, and the result shows that most of the elements, especially radius, tibia, astragalus and metapodials, are remarkably elongated or enlarged, but the humerus and ulna are only slightly longer than those of the other Quaternary rhinos. Among all the rhinocerotid taxa, the humeral length seems relatively stable.

37Although the existence of Mc V was deduced according to the Mc IV lateral facet in the Nihewan fossils, the real fossil of Mc V was recovered for E. sibiricum (Schvyreva, 2016: Pl-XIV). Moreover, the Mc V is also rudimentary in some taxa of the Paleogene paraceratheres (Turpanotherium elegans) but was pretty well developed in other Paleogene paracerathere taxa (Juxia sharamurenensis) (Qiu & Wang, 2007) and the Miocene aceratherini genus Plesiaceratherium (Young, 1937). In the extant rhinos, all of the four genera also have the vestigial Mc V, but they are functionless (Guérin, 1980); Antoine (2002) once gave a thorough analysis on the rudimentary Mc V.

38The elasmothere is a special group not only for its specialized cranial and dental morphologies, but also for its patched geographical distributions. Up to 1995, about 129 localities of Elasmotherium were known (Shvyreva, 1995), the majority (102) of which are E. sibiricum sites and five more Late Pleistocene localities were added in the recent years (Kosintsev, 2014). Shvyreva’s data only included two Chinese localities. Elasmotherium densely occurred in the western Siberia and the Black-Caspian Seas region, but they also sparsely appeared in western Europe (Fortelius & Heissig, 1989), Mediterranean Islands and Transbaikal region as well as northern China (Schvyreva, 2016) (fig. 1), although the presence of Elasmotherium in the Mediterranean Islands is highly questioned (Guérin, 1980; Antoine, 2002). One of the richest collections of Elasmotherium fossils is in the Vernadsky Museum, which included ca. 70 specimens (Zhegallo et al., 2002).

39In China, fossil records of E. peii are limited to Early Pleistocene as known to date and only recovered at six localities. The primitivities and the old geological settings of the Chinese Elasmotherium fossil suggest that it’s very probable that the earliest Elasmotherium animal could have appeared in China.

40Compared with the European and Siberian Elasmotherium fossils, the specimens of E. peii from Nihewan are different in the following aspects: thicker but less crimped enamel layer, metaconid much more developed, upper teeth more elongate, M3 smaller than M2, metastyle less developed and metaloph fused with ectoloph in M3, lophs more developed in upper teeth. In addition, the Chinese material is also different from E. sibiricum with relatively less hypsodont teeth, thicker enamel and prominent root. The recent biostratigraphic study shows that the Elasmotherium-bearing strata are exclusively of an Early Pleistocene age in China (Tong et al., 2014). The differences between the Nihewan specimens and the ones from other regions lie in both dental and postcranial characters.

41E. chaprovicum is characterized by relative massiveness; a low talus (astragalus) with a narrow trochlea of talus and a wide distal portion; with massive metapodial bones; teeth with thick rough enamel and irregularly folded (Titov, 2008; Schvyreva, 2015). The species was put at the root of the phylogenetic lineage of the Elasmotherium species and occupied a time span between 2.6-2.2 Ma.

42The European fossils of E. peii has an early closure of the roots; the prolonged existence of an open postfossette; with presence of a collar on the back wall of the tooth, the lingual parts of the protoloph and metaloph strongly bend backward; a clear differentiation of the crown and root (Schvyreva, 2015). The fossils of E. peii in Europe mainly occurred in the Psekups faunistic complex in Moldova (David & Eremeico, 2003), which geological range is 2.2-1.1 Ma (Schvyreva, 2015).

43The teeth of E. caucasicum have lost the collar and have a more pronounced prismatic shape; with later closure of the pulp cavity and with postfossette; with disorderly growth and greater tortuosity of enamel. The species E. caucasicum appeared in the Taman faunistic complex which geological time interval is 1.1-0.8 Ma (Schvyreva, 2015), and which overlaps with that of E. peii to some extent.

44E. sibiricum displays a shortening of dental series by reduction of the number of premolars; the molars, despite great wear, have open roots; the enamel of the upper and the lower teeth are thinner with strong folding; the early closure and disappearance of the postfossette is typical; all teeth have a slightly curved shape of the prism, without tooth root (Schvyreva, 2015). E. sibiricum appeared in the following faunistic complexes: the Tiraspol (0.8-0.4 Ma) in Moldova, the Syngil (0.4-0.3 Ma) in Ukraine and the Khazar (0.3-0.2 Ma) in the Volga-Ural regions of Russia (Schvyreva, 2015); furthermore, quite a number of Late Pleistocene records of E. sibiricum have also been reported in recent years (Kosintsev, 2014) and some of them even could survive as late as ca. 29 ka BP in Kozhamzhar in Central Asia (Shpansky et al., 2016).

45According to the comparisons on the odontometric data (Titov, 2008; Schvyreva, 2016), it seems that E. sibiricum is prominently odontometrically smaller than the other species of the genus Elasmotherium, but the later three species are not easy to be distinguished among themselves mainly because of the shortage of specimens for E. chaprovicum and E. peii.

46Although the Elasmotherium fossils were mainly recovered in the western Siberia and the Black-Caspian Seas regions, this genus has evolved from Sinotherium in China during the Late Neogene (Chow, 1958; Prothero et al., 1989; Deng et al., 2013) or at least has the closest affinities to Sinotherium (Fortelius & Heissig, 1989; Antoine, 2002, 2003). Cerdeño (1998) even combined the genus Sinotherium into the genus Elasmotherium. Although the morphological evidences support the Sinotherium-deriving of Elasmotherium, the temporal and spatial hiatuses between Sinotherium and Elasmotherium are still existing (fig. 1). In China, cranial material of Sinotherium lagrelii was discovered in recent years, but the latest occurrence of the species was dated back to 7-6 Ma (Xue et al., 2006; Deng et al., 2013). In recent decades, a new species, Sinotherium zaisanensis, and other records of Sinotherium were also reported from northern Tajikistan and southern Kazakhstan respectively, and their geological ages are 7-5 Ma (Bayshashov, 1986 ; Sotnikova et al., 1997), but they still cannot be directly linked to Elasmotherium.

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Table des illustrations

Titre Fig. 1: Spatial (A) and temporal (B) distributions of Elasmotherium, with details of the Chinese sites (C) of E. peii.
Légende 1/ Xiashagou; 2/ SSMZ; 3/ Heitugou; 4/ Xihoudu; 5/ Sanmenxia; 6/ Shanxi but without location details.
Crédits Data for Elasmotherium sites outside China are after Shvyreva (2016)
Fichier image/jpeg, 56k
Titre Tab. 1: Recently recovered fossils of Elasmotherium peii from Nihewan Basin.
Fichier image/jpeg, 124k
Titre Fig. 2: Forelimb bones of E. peii from Shanshenmiaozui
Légende 1/ Left ulna (V23590.1); 2/ left magnum (V23590.4); 3/ right Mc III (V23590.2); 4/ right Mc IV (V23590.3). 1a, 2a, 3a and 4a, anterior views; 1c, 2d and 4c, lateral views; 1b, 2b, 3b and 4b, posterior views; 1d, 2c, 3c and 4d, medial views; 3e and 4e, proximal views.
Fichier image/jpeg, 96k
Titre Tab. 2: Measurements of postcranials of Elasmotherium peii, compared with related taxa (in mm).
Fichier image/jpeg, 200k
Titre Fig. 3: Hindlimb bones of E. peii from Shanshenmiaozui
Légende 1/ Right tibia with partial fibula (V23590.6); 2/ left astragalus (V23590.7); 3/ right cuboid (V23590.8); 4/ partial left Mt IV (V23590.9); 5/ the distal phalanx of toe III of the pes (V23590.11). 1a, 3a and 4a, anterior views; 1b, posterior view; 1c, 3e and 4c, medial views; 1d, 3d and 4d, lateral views; 2a, 3a and 5a, dorsal views; 2b, 4b and 5b, plantar views; 1e, 2c and 3c, distal views; 3b, 4e and 5c, proximal views.
Fichier image/jpeg, 86k
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Référence papier

Hao‑wen Tong, Xi Chen et Bei Zhang, « New postcranial bones of elasmotherium peii from shanshenmiaozui in Nihewan basin, Northern China », Quaternaire, vol. 29/3 | 2018, 195-204.

Référence électronique

Hao‑wen Tong, Xi Chen et Bei Zhang, « New postcranial bones of elasmotherium peii from shanshenmiaozui in Nihewan basin, Northern China », Quaternaire [En ligne], vol. 29/3 | 2018, mis en ligne le 01 septembre 2020, consulté le 26 novembre 2020. URL : ; DOI :

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Hao‑wen Tong

Key Laboratory of Vertebrate Evolution and Human Origins of Chinese Academy of Sciences, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, CN-100044 BEIJING. Email: ;  CAS Center for Excellence in Life and Palaeoenvironement, CN-100044 BEIJING

Xi Chen

Key Laboratory of Vertebrate Evolution and Human Origins of Chinese Academy of Sciences, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, CN-100044 BEIJING. Email: ; CAS Center for Excellence in Life and Palaeoenvironement, CN-100044 BEIJING; University of the Chinese Academy of Sciences, CN-100049 BEIJING.

Bei Zhang

Key Laboratory of Vertebrate Evolution and Human Origins of Chinese Academy of Sciences, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, CN-100044 BEIJING. Email :; CAS Center for Excellence in Life and Palaeoenvironement, CN-100044 BEIJING;University of the Chinese Academy of Sciences, CN-100049 BEIJING.

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