A Revision of Late Oligocene Associations of Small Mammals from the Aral Formation (Kazakhstan) in the National Museum of Georgia, Tbilissi

Palaeodiversity 2: 343–377; Stuttgart, 30.12.2009. 343

A revision of Late Oligocene associations of small mammals from the Aral Formation (Kazakhstan) in the National Museum of Georgia, Tbilissi

Ol e g G. Be n d u k i d z e , Ha n s d e Br u i j n & La r s W. v a n d e n Ho e k Os t e n d e

Abstract The taxonomy and age of the small mammals from the type area of the Aral Formation have been disputed ever since the first specimens from Akespe were published by Ar g y r o p u l o in 1939. The material from Altyn Schokysu, Sayaken, Akespe and Akotau dicussed below was collected by the first author (OGB) during the second half of the twentieth century and published in Russian in 1993. The revision of the fauna from the Aral Formation suggests a biostratigraphical correlation with the radiometrically calibrated Oligocene zone C/C` in Mongolia defined by Da x n e r -Hö c k . Hence the original assignment of the Aral fauna to the Late Oligocene (Tabenbulukian) is considered to be correct. K e y w o r d s : Kazakhstan, Rodents, Lagomorphs, Insectivores, Oligocene.

Zusammenfassung Die Taxonomie und das Alter der Kleinsäuger aus dem Typusgebiet der Aral-Formation werden seit der Be schreibung der ersten Funde von Akespe durch Ar g y r o p u l o (1939) diskutiert. Das hier besprochene Material von Altyn Schokysu, Sayaken, Akespe und Akotau wurde vom Erstautor (OGB) während der zweiten Hälfte des zwan- zigsten Jahrhunderts gesammelt und 1993 in Russisch publiziert. Die Revision der Fauna der Aral-Formation deutet auf eine biostratigraphische Korrelierung mit der radiometrisch kalibrierten und von Da x n e r -Hö c k definierten Oligozän-Zone C/C’ in der Mongolei hin. Daher wird die ursprünglich angenommene Einstufung in das späte Oli- gozän (Tabenbulukium) als richtig betrachtet.

Contents 1. Introduction ...... 343 2. Methods ...... 344 3. Systematic paleontology ...... 345 Erinaceomorpha Gr e g o r y , 1910 ...... 345 Erinaceidae Fi s c h e r , 1814...... 345 Soricomorpha Gr e g o r y , 1910...... 345 Talpidae Fi s c h e r , 1814 ...... 345 Heterosoricidae Vi r e t & Za p f e , 1951...... 346 Lagomorpha Br a n d t , 1855...... 347 Leporidae Fi s c h e r v o n Wa l d h e i m , 1817...... 347 Ochotonidae Th o m a s , 1897...... 349 Rodentia Bo w d i c h , 1821...... 349 Castoridae He m p r i c h , 1820...... 349 Ctenodactylidae Ge r v a i s , 1853...... 351 Dipodidae Fi s c h e r v o n Wa l d h e i m , 1817...... 352 Muridae Il l i g e r , 1811...... 352 4. List of the small mammals of the Aral local fauna represented in the Be n d u k i d z e collection...... 356 5. The age of the assemblage...... 356 6. References ...... 356

1. Introduction mammals from the type area of the Aral Formation have been the subject of a series of detailed studies by Lo p a t i n , The decision to give an annotated series of figures of who greatly enlarged the collections from Altyn Schokysu the specimens from the North Aral area in western Ka- and increased the knowledge of its interesting fauna (Lo- zakhstan described and discussed in the monograph of p a t i n 1996, 1998, 1999a, 1999b, 2003, 2004). However, it Be n d u k i d z e (1993) was taken shortly after the interna- is of interest to figure and update the identification of the tional meeting on “The Oligo/Miocene boundary in Ka- specimens in the Be n d u k i d z e collection because it con- zakhstan,” Aktyubinsk, 1994. Since that time the small tains many types, and to discuss the age of the fauna in the 344 PALAEODIVERSITY 2, 2009

collecting bias because a sample taken by D. KÄLIN, T. BOLLIGER, G. DAXNER-HÖCK and the second author (HdB) from level 2 at Altyn Schokysu in 1994 (n = about 400) contained only about 5 % of the larger rodent species that dominate the BENDUKIDZE collection. For the geographical position of the localities and detailed descriptions the reader is referred to LOPATIN (2004), BENDUKIDZE (1993) and Fig. 1. Figures of the more interesting specimens as well as updated identifications will be given below. The material discussed below is housed in the collections of the Institute of Paleobiology, Georgian National Museum, Tbilissi.

Acknowledgements We thank Mr. W. DEN HARTOG for making the SEM photographs and Mr. J. LUTEIJN for retouching these photographs. Mr. F. TRAPPENBURG made the plates and Mr. R. RABBERS made the sketch map of the study area (both Geomedia Department, Utrecht). Mr. H. BRINKERINK made the casts that served as the basis for the photographs, so covering the original specimens with gold could be avoided. Ms. J. RICHTER assisted in finding literature references for supraspecific taxa. The expert advice of our colleagues MARGUERITE HUGUENEY (Lyon), MARY R. DAWSON (Pittsburgh) and MARGARITA ERBAJEVA (Ulan Ude) on the Castoridae respectively the Lagomorpha in our collection is much appreciated. The second author (HdB) gratefully acknowledges his son in law, MARC TER HAAR for translating some Russian texts and KEES HORDIJK for reading the Fig. 1. Sketch map of Kazakhstan and the study area showing manuscript critically. the geographical position of the localities that yielded the small GUDRUN DAXNER-HÖCK made an essential contribution to this mammals discussed below (modified after LOPATIN 2004). paper by showing her collection of rodents from the Valley of Lakes, Mongolia to the second author (HdB). light of new information on the, radiometrically calibrat- 2. Methods ed, fauna succession from the Valley of Lakes in Mongolia (DAXNER-HÖCK et al. 1997). Measurements of non-lagomorph teeth represent max- The associations of fossil small mammal remains from imal length × width of the occlusal surface and are given the North Aral area, collected by the first author (OGB) in 0.1 mm units unless indicated otherwise. The nomen- during the eighties of the last century (BENDUKIDZE 1993), clature of parts of the cheek teeth follows LÓPEZ-MARTÍNEZ can be divided into a Late Oligocene and a Miocene group (1977) for the lagomorphs and HUGUENEY (1999) for the on the basis of biostratigraphical criteria. The material castorids. Three measurements are given for the molari- from the localities from the Aral Formation (Fig. 1): Altyn form lower cheek teeth of lagomophs: the first applies to Schokysu (levels 1–4), Sayaken, Akespe and Akotau in- the length, the second to the width of the trigonid and the cluded in the first group will be discussed below. The col- third to the width over the talonid. lections from the four different fossiliferous levels in the The specimens on the plates are all figured as if they Altyn Schokysu escarpment that have been recognised by are from the left side. If the original is from the right side some authors are united into one local fauna, because they the relevant number has been underlined on the plate. come from lenses that are situated at considerable lateral The specimens from the northern Aral area housed in distances. Their relative stratigraphical position is there- the Institute of Paleobiology, Tbilissi carry a locality as fore not clear. For a discussion of the litho- and chrono- well as a specimen number. Nine stands for the locality logical position of the Aral Formation we refer to LUCAS et Sayaken, eleven for Akespe, fifteen for Altyn Schokysu al. (1998). and sixteen for Akotau. The collection studied seems to suffer from a strong BENDUKIDZE ET AL., OLIGOCENE MAMMALS FROM KAZAKHSTAN 345

3. Systematic paleontology tion (LOPATIN 1999a). The material therefore clearly represents a different and possibly new species. As the taxono- Erinaceomorpha GREGORY, 1910 my of Amphechinus is in mayhem as it is, we refrain from Erinaceidae FISCHER, 1814 describing yet another species based on scanty material. Galericinae POMEL, 1848

Galericinae gen. et sp. indet. BOHLIN, Pl. 1, Fig. 1 Amphechinus cf. minimus ( 1942) Pl. 1, Figs. 6–8 Locality: Altyn Schokysu. BENDUKIDZE Material and measurements: 1993 Amphechinus minimus. – , p. 10, pl. 2, figs. 1–4. No. Fig. Tooth Length × Width 1999 Amphechinus microdus LOPATIN, sp. nov. – LOPATIN, pp. 15/1 Pl. 1, Fig. 1 m1 sin. 27.1 × 18.4 185–187, fig. 3. [1999a] 2004 Amphechinus microdus LOPATIN, 1999. – LOPATIN, pp. S323–S234, fig. 8, pl. 6, figs. 6–8. R e m a r k s . – The presence of galericines in the fauna had already been noted by LOPATIN (2004), who assigned a L o c a l i t y : Altyn Schokysu. Material and measurements: p4 to Galerix sp. However, the specimen he illustrates seems to be an m3 of Amphechinus sp., fitting well in No. Fig. Tooth Length × Width 15/7 Pl. 1, Fig. 6 M1 sin. – × – morphology and dimensions with the m3 in the BEN- Fragment of a man- DUKIDZE collection. Nevertheless, one m1 in our collection 15/8 Pl. 1, Fig. 8 dible sin. with m2 18.2 × 12.6 strongly resembles Galerix in having a conical paraconid m3 7.8 × 4.2 at the end of a long paralophid. Galerix has its first occur- 15/8a Pl. 1, Fig. 7 m1 sin. 22.8 × 13.1 rence in the uppermost Oligocene of Anatolia (DE BRUIJN et al. 1992). Metrically, our m1 falls just outside the range of R e m a r k s . – The smaller of the two Amphechinus VAN DEN HOEK the oldest species of Galerix, G. saratji species from Altyn Schokysu was originally described as OSTENDE, 1992 , being slightly larger. Since we just have Amphechinus minimus by BENDUKIDZE (1993). LOPATIN one m1, and the Oligocene history of the Asian galericines (1999a) named the smaller species A. microdus. This new is poorly known, this tooth cannot be identified beyond species was distinguished mainly on the basis of the stron- the subfamily level. ger reduction of the m3 relative to the m2 (40 % vs 55 %). Although this feature is also observed in our material we believe that, without knowing the variation of this charac- F , 1814 Erinaceinae ISCHER ter, it is insufficient to warrant distinction at the species A Amphechinus YMARD, 1850 level. Therefore, we maintain the original classification as A. minimus. Amphechinus sp. Pl. 1, Figs. 2–5

2004 Galerix sp. – LOPATIN, pp. S228–S230, pl. 1, fig. 1. Soricomorpha GREGORY, 1910 L o c a l i t y : Altyn Schokysu. Talpidae FISCHER, 1814 Material and measurements: Uropsilinae DOBSON, 1883 No. Fig. Tooth Length × Width Theratiskos vAN DEN HOEK OSTENDE, 2001 15/2 Pl. 1, Fig. 2 P4 dext. 23.3 × 29.8 15/4 Pl. 1, Fig. 3 fragm. M1 sin. – × – Theratiskos compactus (LOPATIN, 2004) 15/5 Pl. 1, Fig. 4 M2 dext. 22.6 × 27.9 Pl. 1, Fig. 9 Mandible sin. with p4 20.3 × 14.5 1993 Astenoscaptor [sic] sp. – BENDUKIDZE, p. 19, pl. 15/6 Pl. 1, Fig. 5 fragm. m1 – × – 5, figs. 3, 4. m2 27.4 × 18.5 1999 Desmanella sp. – LOPATIN, p. 16. [1999a] m3 12.5 × 8.9 pro parte 1999 Myxomygale sp. nov. – LOPATIN, p. 66. [1999a] 2004 Desmanella compacta LOPATIN, sp. nov. – LOPA- . Remarks. – This Amphechinus material belongs to TIN, pp. S235–S236, pl. 1, fig. 11 a species that remained hitherto unnoticed, except for one L o c a l i t y : Altyn Schokysu. Material and measurements: m3 that was described by LOPATIN (2004) as a p4 of Gal- erix sp. The p4 and m2 are more than 20 % smaller than No. Fig. Tooth Length × Width those of A. akespensis LOPATIN, 1999 from the Aral forma- 15/9 Pl. 1, Fig. 9 m2 sin. 14.1 × 8.7 346 PALAEODIVERSITY 2, 2009

Remarks. – BENDUKIDZE (1993) assigned this small (1999a) correctly indicated that it is certainly not referable talpid to the genus Asthenoscaptor. LOPATIN (1999a) men- to Mygalea and placed the species in his genus Pseudo- tioned the presence of Desmanella rather than As- paratalpa. ZIEGLER (2003) considered the differences be- thenoscaptor, and in 2004 described his material from tween Pseudoparatalpa and Paratalpa too small to war- Altyn Shokysu under the name Desmanella compacta. A rant generic distinction. We agree, but the synonymy with remarkable feature of the holotype M2 of this species is Paratalpa is based on a few shared characters in the lower the relatively straight posterior outline, whereas usually dentition only. The upper molars – which in talpids are uropsiline talpids have an emarginated posterior side of generally more characteristic than the lower dentition – of the M1 and M2 due to the presence of a hypoconal flange. Pseudoparatalpa are unknown, as is the humerus. LOPATIN A very weak to absent posterior emargination is, in the (2004) described humeri from Altyn Schokysu and attrib- subfamily, only known from Theratiskos, a genus from uted these to Hugueneya. No dental elements were associ- the Upper Oligocene and Lower Miocene of Anatolia (vAN ated with this type of humerus. Pseudoparatalpa makes a DEN HOEK OSTENDE 2001). The morphology of the holotype plausible candidate for such an association, in which case of D. compacta fits well with this genus, so this species is the genus is certainly not synonymous with Paratalpa. better assigned to Theratiskos. The fragmentary M1 as- However, the talpid material is too poor to be certain that signed to Myxomygale asiaprima also fits the morphology dentition and humeri indeed belong together. For the time of Theratiskos, and its size seems to fit well with the M2. being we therefore retain Pseudoparatalpa as a separate At the same time, it seems too small to be associated with genus, until further material can help elucidate its affini- the m1 that was selected as the holotype of M. asiaprima. ties. Therefore, it is also referred to Theratiskos compactus. As Two species were described in the genus, Pseudopara- the holotype M2 is clearly smaller than the corresponding talpa lavroi and the type species P. shevyrevae LOPATIN, element in both T. mechteldae and T. rutgeri, the Kazakh- 1999. The differences indicated by LOPATIN (a weaker cin- stan material clearly represents a separate species. gulum development and slightly higher horizontal ramus) are too small to make a specific distinction, and we therefore consider P. shevyrevae to be a junior synonym of P. FISCHER Talpinae , 1814 lavroi. Urotrichini DOBSON, 1883 Pseudoparatalpa LOPATIN, 1999 Heterosoricidae VIRET & ZAPFE, 1951 BENDUKIDZE Pseudoparatalpa lavroi ( , 1993) Gobisorex SULIMSKI, 1970 Pl. 1, Fig. 10 Gobisorex kingae SULIMSKI, 1970 BENDUKIDZE 1993 Migalea [sic] lavroi sp. nov. – , p. 11, pl. 2, fig. Pl. 1, Figs. 11–13 5, pl. 3, figs. 1, 2. LOPATIN . LOPATIN 1999 Pseudoparatalpa shevyrevae , sp. nov – , 1993 Gobiosorex [sic] kingae. – BENDUKIDZE, p. 22, pl. 7, figs. pp. 187–188, fig. 4. [1999a] 2–4. BENDUKIDZE, 1993. LOPATIN, 1999 Pseudoparatalpa lavrovi – 1993 Gobisorex aff. kingae. – BENDUKIDZE p. 23, pl. 7, fig. 5, pp. 188–189, fig. 5. [1999a] pl. 8, fig. 1. BENDUKIDZE, 1993). LOPATIN 2004 Pseudoparatalpa lavrovi ( – , 1999 Gobisorex sp. – LOPATIN, p. 182. [1999a] pp. S236–S237, fig. 11, pl. 1, fig. 12. 1999 Heterosoricidae gen. et sp. indet. – LOPATIN, p. 182. L o c a l i t y : Altyn Schokysu. [1999a] Material and measurements: 2004 Atasorex edax LOPATIN, sp. nov. – LOPATIN, pp. S242– No. Fig. Tooth Length × Width S243, fig. 15, pl. 2, figs. 2–5. 2004 Gobisorex akhmetievi LOPATIN, sp. nov. – LOPATIN, pp. Mandible sin. with S239–S241, fig. 14, pl. 2, fig. 1. 15/10 Pl. 1, Fig. 10 m1 21.9 × 14.9 m2 23.3 × 13.3 L o c a l i t y : Altyn Schokysu. Material and measurements: R e m a r k s . – Based on size and general appearance No. Fig. Tooth Length × Width this mandible fits with the species lavroi. However, there Mandible dext. are some marked differences with the description of the m2 15/11 Pl. 1, Fig. 12 with fragm. m1 – × – m2 19.6 × 12.1 given by LOPATIN (1999a, 2004). The talonid is shorter, 15/12 Pl. 1, Fig. 11 M1 sin. 16.0 × 17.4 rather than wider, than the trigonid, the oblique cristid ends 15/12a M1 sin. 17.9 × 18.5 halfway the protocristid and a metacristid is present. Fur- Mandible sin. with thermore, the trigonid is remarkably short, a feature not m1 20.5 × 11.7 15/13 Pl. 1, Fig. 13 mentioned by LOPATIN. These specimens were originally m2 17.5 × 10.7 described by BENDUKIDZE (1993) as Migalea lavroi. LOPATIN m3 – × – BENDUKIDZE ET AL., OLIGOCENE MAMMALS FROM KAZAKHSTAN 347

Remarks. – Both BENDUKIDZE (1993) and LOPATIN Lagomorpha BRANDT, 1855 (1999a, 2004) recognised two species of heterosoricid shrew from Altyn Shokysu. BENDUKIDZE assigned the ma- The taxonomy of the diverse lagomorphs from the terial to Gobisorex kingae and Gobisorex aff. kingae. LO- Oligo/Miocene boundary interval in Central Asia is com- PATIN (1999a) classified the material as Gobisorex sp. and plex. Controversies on the status of families, subfamilies, Heterosoricidae gen. et sp. indet. Gobisorex sp. was later genera and species are due to the often insufficiently described as G. akhmetievi LOPATIN, 2004. Gobisorex known intraspecific variation, the uncertainty about the akhemetievi differs according to LOPATIN from G. kingae allocation of some genera to the classical families Ochoto- in the larger size and the presence of a lingual cingulid on nidae and Leporidae, the mosaic type of evolution seen in the m2. This species is represented by the holotype only, lagomorphs and the uncertain age assignment of many as- a mandible fragment with an m2 and the posterior part of sociations. The emphasis in the work of many lagomorph the m1. LOPATIN (2004) assigned the remainder of the ma- specialists is on subdividing the order into a large number terial to a new genus and species, Atasorex edax. The new of families, subfamilies and genera that are each consid- genus was differentiated from Gobisorex on the basis of ered to comprise a clade (BURKE 1941, GUREEV 1960, ER- the weak entocristid of the m1 and m2, the poorly devel- BAJEVA 1994). We think that this practice enhances the in- oped entostylid, the shape of the m2 trigonid and the stability of the nomenclature because new information continuous ectocingulid of the m2. Although such charac- will influence the reconstruction of lineages and thereby ters, when consistent, might be used to distinguish spe- the content of the (sub)families. In sharp contrast to this cies, they seem unsuited for the distinction between gen- practise is the conservative nomenclature that prevails in era. other instances: BOHLIN (1937), TOBIEN (1974) and LOPATIN One of the problems BENDUKIDZE (1993) and LOPATIN (1998) commented on the similarity of Desmatolagus (1999a, 2004) were facing, was that little was known about MATTHEW & GRANGER, 1923 and Amphilagus POMEL, 1853 the variation in Gobisorex kingae, a species mainly known (sensu TOBIEN 1974), but both genera were retained and by the type mandible and a second lower jaw (SULIMSKI have subsequently been subdivided. In my (HdB) opinion 1970). Unfortunately, the illustrations in the original pub- the somewhat deeper hypostriid in the p3 in Amphilagus lication are not very informative. A recent study on Mon- antiquus than in the p3 of the majority of Asiatic species golian insectivores (ZIEGLER et al. 2007) greatly expanded assigned to Desmatolagus is over-valued. Likewise, the our knowledge of this heterosoricid. They noted, for in- difference between Amphilagus and Eurolagus LÓPEZ- stance, that their measurements of the m2 filled the gap MARTÍNEZ, 1977 is one of degree and seems of insufficient between the measurements for the holotypes of Gobisorex importance for generic separation. kingae and G. akhmetievi. Quite another problem is posed by the dental morphol- Like the Mongolian material, the two m2 from Altyn ogy of the Amphilagus/Desmatolagus group of species, Shokysu are intermediate in size between the types of because this shows characteristics of the Leporidae as Gobisorex kingae and G. akhemetievi. Both our speci- well as of the Ochotonidae. The “solution” to unite these mens have a very faint lingual cingulid. The labial cingu- in a separate family Paleolagidae DICE, 1929 seems unsat- lid is interrupted at the base of the protoconid, which isfactory. Although the study of the microstructure of the agrees with the description by LOPATIN (2004), but not with incisor enamel by MARTIN (2004) covers so far only a his illustration (o. c., pl. 2, fig. 1). The material LOPATIN small selection of, mainly primitive, members of the order, (2004) assigned to Atasorex edax also comfortably fits the results suggest that this approach may eventually al- within the measurements given for Gobisorex kingae by low the unambiguous separation of the Leporidae and ZIEGLER et al. (2007, tab. 13). A possible exception is the Ochotonidae. We follow MARTIN (2004) and tentatively M2, which also has a slightly different morphology, with a include all Desmatolagus species into the Leporidae. more reduced posterior side, on which the posterior ridge Since LOPATIN’s (1998) thorough revision of the Lago- does not reach the posterolabial corner of the molar. The morpha from the North Aral area is based on much better m2 also seems a bit small, but this may be partly due to material than is available to us, we refer for detailed de- wear. Wear may also have influenced the assessment of scriptions to that paper. the morphology, as it changes the appearance of the ento- cristids, one of the characters used to distinguish Atasorex edax. Leporidae FISCHER VON WALDHEIM, 1817 All in all, there is insufficient ground to distinguish two species in the heterosoricid assemblage from Altyn Desmatolagus MATTHEW & GRANGER, 1923 Shokysu. As the material falls within the variation given ZIEGLER by et al. (2007) for Gobisorex kingae, we refer the Synonyms: Agispelagus ARGYROPULO, 1940, Procapro- Altyn Shokysu assemblage to that species. lagus GUREEV, 1960. 348 PALAEODIVERSITY 2, 2009

Desmatolagus simplex (ARGYROPULO, 1940) Desmatolagus periaralicus LOPATIN, 1998 Pl. 2, Figs. 1, 2 Pl. 2, Fig. 5

1940 Agispelagus simplex. – ARGYROPULO, p. 76. 1993 Desmatolagus aff. gobiensis MATTHEW & GRANGER, 1923. 1960 Agispelagus simplex. – GUREEV, p. 25, fig. 10a. – BENDUKIDZE, pp. 30–31, pl. 11, figs. 2–5. BOHLIN BEN- 1993 Desmatolagus aff. shargaltensis , 1937. – L o c a l i t y : Altyn Schokysu. DUKIDZE, pp. 28–30, pl. 10, figs. 5, 6, pl. 11, fig. 1. Material and measurements: L o c a l i t y : Altyn Schokysu. No. Fig. Tooth Length × Width Material and measurements: Fragment of a right maxilla with P3–M2, length = 69.7 mm, No. 15/19; Frag- 15/21 Pl. 2, Fig. 5 M2 16.3 × 37.6 ment of a left mandible with p3–m2, length = 57.7 mm, No. 15/20. R e m a r k s . – The only tooth of this species in our No. Fig. Tooth Length × Width collection does not add to the information given in LOPATIN Fragment of a right (1998, 2004). Desmatolagus periaralicus is of about the maxilla with same size as the younger D. kazachstanicus BENDUKIDZE, P3 20.0 × 23.5 15/19 Pl. 2, Fig.1 P4 16.2 × 28.0 1993 from Kentyubek, but more primitive in having a very M1 13.8 × 31.7 shallow mesoflexus in the P3. M2 11.5 × 26.4 Fragment of a left mandible with p3 9.2 × 13.0 15/20 Pl. 2, Fig.2 p4 16.4 × 20.5 (11.0) m1 15.8 × 20.0 (10.3) m2 16.5 × 18.3 (11.3) Desmatolagus veletus LOPATIN, 1998 R e m a r k s . – Small species of Desmatolagus are Pl. 2, Figs. 6–10 well represented in the Oligo/Miocene of Asia and quite a 1993 Amphilagus aff. robustus (MATTHEW & GRANGER, 1923). MAT- number of these have been formally named since – BENDUKIDZE, p. 26, pl. 9, figs. 1–8, pl. 10, figs. 1–4. THEW & GRANGER (1923) described the type species, D. L o c a l i t y : Altyn Schokysu. gobiensis. Some specialists have later interpreted the, of- Material and measurements: ten minor, differences between the type material of some No. Fig. Tooth Length × Width SYCH species as intraspecific variation, e. g. (1975), who 15/14 Pl. 2, Fig. 6 P3 24.6 × 41.2 considered D. pusillus TEILHARD DE CHARDIN, 1926, D. 15/15 Pl. 2, Fig. 7 P4 26.9 × 46.2 radicidens TEILHARD DE CHARDIN, 1926, D. parvidens BOH- 15/16 Pl. 2, Fig. 8 M1 22.5 × 38.0 LIN, 1937, Procaprolagus mongolicus GUREEV, 1960, Pro- 15/18 Pl. 2, Fig. 9 p3 19.0 × 20.4 caprolagus orlovi GUREEV, 1960 and Sinolagomys tatal- 15/18a Pl. 2, Fig. 10 m1-2 31.5 × 31.0 golicus GUREEV, 1960 as junior synonyms of D. gobiensis. 15/18b – m1-2 29.5 × 30.9 ERBAJEVA & SEN (1998), on the other hand, distinguished within material from the type locality of D. pusillus (San- Remarks. – Desmatolagus veletus is quite similar tao-ho, China) two species that they assign to two genera: to D. robustus. This in particular so because the p3 in the Bohlinotona pusilla (TEILHARD DE CHARDIN, 1926) and BENDUKIDZE collection has, in contrast to the specimen Desmatolagus chinensis ERBAJEVA & SEN, 1998. Given the from the same locality described by LOPATIN (1998), no complexity of the matter we are unable to develop an inde- cement. In spite of the overall similarity of the dentitions pendent opinion on the validity of all the formally named of D. robustus and D. veletus we retain these two species species on the basis of literature. because the latter is more evolved in having a P3 with a The material from Altyn Schokysu listed above closely deeper mesoflexus that contains cement and in having resembles Desmatolagus simplex (ARGYROPULO, 1940) as some cement in the hypoflexus of the upper cheek teeth. described by LOPATIN (1998) and we agree with that author Regarding the separation of Desmatolagus from Am- that the dental characteristics of this small Desmatolagus philagus, the species D. veletus is of special interest be- with rather deep hypostriae that contain some cement and cause of its similarity to A. antiquus. In fact, the dentitions with lower cheek teeth devoid of hypoconulids are pro- of these two species are more alike than most Asiatic spe- gressive. cies of Desmatolagus among themselves. BENDUKIDZE ET AL., OLIGOCENE MAMMALS FROM KAZAKHSTAN 349

Ochotonidae THOMAS, 1897 specific variation that is enhanced through attrition (LOPA- Sinolagomys BOHLIN, 1937 TIN 2004). In comparing teeth showing different wear stages, homologies of parts of cheek teeth are often not Sinolagomys kansuensis BOHLIN, 1937 clear, because flexi and fossettes may get lost through at- Pl. 2, Figs. 3, 4 trition. As a result, specimens of the same species have often been allocated to different genera and species by dif- 1993 Sinolagomys aff. kansuensis BOHLIN, 1937. – BENDUKIDZE, ferent authors or, more problematically, by the same au- pp. 37–38, pl. 14, figs. 2–7. thor in different publications. An additional problem is 1998 Sinolagomys pachygnatus LI & QIU, 1980. – LOPATIN, pp. 300–301, fig. 4a–l. that much of the older literature dealing with Central Asi- 2004 Sinolagomys pachygnatus LI & QIU, 1980. – LOPATIN, pp. atic fossils has been published in Russian and is not broad- 255–257, fig. 21a–m. ly accessable. The material from the North Aral area avail- L o c a l i t y : Altyn Schokysu (15/23), Akotau (16/6). able here consists at best of dentitions, but more often of Material and measurements: isolated cheek teeth. The identifications given below are No. Fig. Tooth Length × Width therefore necessarily conservative. Fragment of a We agree with LOPATIN (2004), who in his revision of maxilla with the North Aral beavers in the collection of the Paleonto- P3 15.4 × – 15/23 Pl. 2, Fig. 3 P4 15.6 × – logical Institute of the Russian Academy of Sciences, M1 13.7 × – Moscow, concludes that there are three species represent- M2 12.5 × – ed in Altyn Schokysu, while as many as six species were Fragment of a recognised by BENDUKIDZE (1993). The variation in size mandible with and in thickness of the enamel of the beaver cheek teeth in p3 10.9 × 13.2 our sample is obviously too large to include all the speci- 16/6 Pl. 2, Fig. 4 p4 17.2 × 18.9 mens in one species. With the exception of one p4 from m1 16.8 × 19.2 m2 17.3 × 19.5 Altyn Schokysu, which we allocate to Asiacastor (Pl. 5, Fig. 7), the dental pattern of the cheek teeth is, apart from R e m a r k s . – The Sinolagomys specimens from Al- differences due to stage of wear, quite similar to Propa- BORISSOGLEBSKAYA tyn Schokysu originally identified as S. aff. kansuensis by laeocastor , 1967. Our recognition of BENDUKIDZE (1993) were later referred to S. pachygnatus two species in this material is based on a combination of by LOPATIN (1998). Although the difference between these size and enamel thickness. The smaller of the two Propa- two species is subtle, correct identification is of great im- laeocastor species, which has thinner enamel, is assigned LYTSCHEV portance because S. kansuensis is supposed to be restrict- to P. kumbulakensis , 1970, the larger one with BENDUKIDZE ed to the Late Oligocene, while S. pachygnatus is indica- thick enamel to P. schokensis ( , 1993). tive for the Early Miocene. Bearing the dispute about the age of the Aral fauna in mind, we not only compared our material with great care to figures of the type materials, B but also solicited the opinion of MARY R. DAWSON and Propalaeocastor ORISSOGLEBSKAYA, 1967 MARGARITA ERBAJEVA. This inquiry led to the unanimous B conclusion that the specimens from Altyn Schokysu with Propalaeocastor schokensis ( ENDUKIDZE, 1993) rudimentary roots in the upper cheek teeth, relatively Pl. 3, Figs. 1–3, Pl. 4, Figs. 1–4 small rounded talonids in the lower cheek teeth and an 1993 Capatanca schokensis. – BENDUKIDZE, pl. 26, only slightly swollen labial side of the mandible at the figs. 1, 2 (p4–m3 dext., holotype), pl. 26, figs. 3, level of the m1, represent Sinolagomys kansuensis. 4 (P4–M2 sin. from Altyn Schokysu). pro parte 1993 Capacikala sayakensis. – BENDUKIDZE, pl. 27, figs. 1, 2. (p4–m1, holotype from Sayaken). BENDUKIDZE, BOWDICH 1993 Steneofiber aff. kumbulakensis. – Rodentia , 1821 pl. 23, fig. 6; pl. 24, fig. 1 (P4–M3 dext. from Castoridae HEMPRICH, 1820 Altyn Schokysu); pl. 24, figs. 2, 3 (p4–m3 sin. from Altyn Schokysu); pl. 26, figs. 1, 2 (p4–m3 Subfamilial and generic identification of beavers is a dext. from Altyn Schokysu). BENDUKIDZE, complex and difficult affair because their taxonomy is 1993 Palaeocastor sp. – pl. 25, fig. 8 (P4 sin. from Altyn Schokysu). based on a combination of dental, cranial and post-cranial 2004 Steneofiber schokensis. – LOPATIN, fig. 27 (P4 characteristics (KORTH 2002). As a rule, collections of fos- sin. and p4 sin. from Altyn Schokysu). sil beavers do not comprise skull and postcranial material 1994 Propalaeocastor devius. – LYTSCHEV & SHEVY- and the number of dental elements per locality is limited. REVA, figs. 3, 4. Moreover, beaver cheek teeth show considerable intra- H o l o t y p e : Fragment of a right mandible with p4–m3 350 PALAEODIVERSITY 2, 2009 from Sayaken (pl. 26, fig. 1 in BENDUKIDZE 1993), our Pl. 4, R e m a r k s . – The species Propalaeocastor kumbu- Fig. 1. lakensis is based on a fragment of a skull and mandible L o c a l i t y : Altyn Schokysu. Material and measurements: with complete dentition from the Kumbulak cliffs of the northern shore of the Aral Sea at 4 km east of the village No. Fig. Tooth Length (× Width) of Akespe and was originally assigned to the genus 15/43 Pl. 3, Fig. 1 P4–M3 14.1 LYTSCHEV LYTSCHEV 15/49 Pl. 3, Fig. 2 P4–M2 11.8 Propalaeocastor by (1970). Later & SHEVYREVA 15/48 Pl. 4, Fig. 1 p4–m3 15.7 (1984) considered Propalaeocastor a junior 15/45 Pl. 4, Fig. 2 p4–m3 16.4 synonym of Steneofiber GEOFFROY SAINT-HILAIRE, 1833, an 15/47 Pl. 3, Fig. 3 P4 occlusal 4.56 × 5.04 opinion that was shared by LOPATIN (2004). WU et al. (2004) maintain Propalaeocastor for the species kazach- Locality: Sayaken. stanicus (genotype), shevyrevae and zaisanensis, but as- Material and measurements (These two teeth sign the species P. kumbulakensis to Steneofiber. from the same individual are the holotype of Capacikala sajakensis BENDUKIDZE, 1993): Facing these conflicting opinions we compared the dentition of the genotype of Propalaeocastor, P. kazach- No. Fig. Tooth Length × Width stanicus BORISSOGLEBSKAYA, 1967, to that of the type spe- 9/22 Pl. 4, Fig. 3 p4 39.0 × 35.3 VON MEYER 9/22 Pl. 4, Fig. 4 m1 28.8 × 37.4 cies of Steneofiber, S. eseri ( , 1846). The genera Propalaeocastor and Steneofiber share dental characteristics such as the absence of cement and the early closure of BENDUKIDZE Remarks. – Capatanca schokensis , the paraflexus(id) and the metaflexus(id). However, the BENDUKIDZE 1993 and Capacikala sajakensis , 1993 were complexity of the pattern mesial and distal to the meso- published in the same work. Capatanca schokensis has flexus that presumably results from the presence of a page priority over C. sajakensis. double protoloph and a metaloph is much greater in the type species of Propalaeocastor than in that of Steneofi- ber. We therefore maintain these two genera. Propalaeocastor kumbulakensis LYTSCHEV, 1970 Our specimens from Altyn Schokysu assigned to the Pl. 3, Figs. 4–7, Pl. 5, Figs. 1–6 species P. schokensis and P. kumbulakensis show a de- pro parte 1993 Capacikala sajakensis. – BENDUKIDZE, pl. 27, gree of complexity that is much more similar to the con- figs. 3, 4. figuration seen in Propalaeocastor (and Anchitheriomys 1993 Capacikala cf. sciuroides (MATTHEW, 1971). – ROGER, 1898) than to that of Steneofiber. Moreover, the BENDUKIDZE, pl. 27, figs. 5–9. enamel of slightly worn teeth shows the irregular crenula- pro parte 1993 Asiacastor aff. orientalis LYTSCHEV, 1988. – BENDUKIDZE, pl. 28, figs. 1, 2. tions that are characteristic for a number of primitive 2004 Steneofiber kumbulakensis (LYTSCHEV, 1970). – beaver species except Steneofiber. Assignment of our ma- LOPATIN, pp. 259–263, figs. 25–26, pl. 5, figs. terial to Anchitheriomys is excluded, because the few 1–13. mandibles of P. schokensis available are more robust and 2004 Asiacastor sp. – LOPATIN, pp. 266–267, fig. 28. show two mental foramina situated below and slightly in L o c a l i t y : Altyn Schokysu. front of the p4. Material and measurements (crown basis): The material from Altyn Schokysu, Akotau and Saya- No. Fig. Tooth Length × Width ken assigned to P. kumbulakensis agrees in size as well as 15/59 Pl. 3, Fig. 6 P4 38.0 × 47.5 in morphology (Pl. 5, Figs. 1–6) with the type specimen, 15/54 Pl. 5, Fig. 1 M1-2 26.7 × 28.5 15/53 Pl. 5, Fig. 4 m1-2 30.6 × 37.5 but since the species P. schokensis and P. kumbulakensis 15/56 – p4 42.0 × 40.0 have overlapping size ranges and very similar dental pat- 15/55 Pl. 5, Fig. 5 m1-2 34.0 × 36.5 terns, erroneous identification of isolated teeth cannot be 15/58 Pl. 5, Fig. 6 m3 32.0 × 34.5 excluded. In particular the separation of the two species on the basis of figures is uncertain, because the thickness of Locality: Akotau. the enamel is not clear. Material and measurements (crown basis): The two fragments of lower incisor preserved in man- No. Fig. Tooth Length × Width dible fragments show a somewhat concave anterior face 16/50 Pl. 3, Fig. 4 P4 39.0 × 41.3 and no ornamentation of the enamel. The cheek teeth have 16/52 Pl. 3, Fig. 5 P4 39.0 × 45.0 no cement and the interdental wear in adult specimens is strong. The P3 is absent. The hypostria(id) is longer than Locality: Sayaken. the mesostria(id), while the even shorter parastria(id) is Material and measurements: preserved in very fresh teeth only. The metastria(id) is not No. Fig. Tooth Length × Width preserved in any of our specimens, so the metafossette(id) 9/40 – M1-2 28.8 × 41.0 may never have been open. BENDUKIDZE ET AL., OLIGOCENE MAMMALS FROM KAZAKHSTAN 351

Asiacastor LYTSCHEV, 1971 1997 Yindirtemys sajakensis birgeri. – BENDUKIDZE, p. 207. M a t e r i a l available for study in Utrecht: Asiacastor sp. Altyn Schokysu: P4–M3 of the holotype, No. 15/28, length = Pl. 5, Fig. 7 12.3 mm; p4, No. 15/29. Akotau: m2, No. 16/9; m3, No. 16/9. M e a s u r e m e n t s of all the material housed in the Na- BENDUKIDZE, pro parte 1993 Capacikala aff. sciuroides. – tional Museum of Georgia: pl. 27, fig. 8 ? BENDUKIDZE, Length Width pro parte 1993 Asiacastor aff. orientalis. – pl. 28, Tooth n fig. 3. Range Mean Range Mean L o c a l i t y : Akotau. P4 6 23.0–25.0 24.7 27.5–30.0 29.0 Material and measurements (measured on the M1-2 4 28.7–35.0 30.9 23.7–32.5 26.3 crown basis): M3 10 36.2–41.2 38.8 25.0–35.0 32.4 p4 6 22.0–27.5 24.1 18.5–22.0 20.3 No. Fig. Tooth Length × Width m1 2 31.0–32.5 31.8 20.0–24.0 22.0 16/133 Pl. 5, Fig. 7 p4 39.6 × 32.0 m2 3 39.5–45.0 42.8 26.0–29.0 28.0 m3 5 48.0–50.0 48.8 30.0–32.5 30.9 R e m a r k s . – The part of the BENDUKIDZE collection available for study in Utrecht contains only one p4 dext. of Remarks. – Yindirtemys ambiguus WANG, 1997, a Asiacastor from Akotau (identified as Asiacastor aff. ori- species that is based on a holotype from Shargaltein fig- BENDUKIDZE entalis, pl. 28, fig. 3 in 1993). It is possible the ured by BOHLIN (1937, fig. 87) under the name Tataromys BENDUKIDZE specimen figured in (1993) as Capacikala cf. cf. plicidens, has about the same size and morphology as sciuroides (pl. 27, fig. 8) from Altyn Schokysu belongs to Yindirtemys birgeri. The only difference between these the same taxon, but this cannot be ascertained on the basis two species seems to be that the paracone of the M3 con- of the figure. sistently bears a posterior spur that reaches the metacone The p4 assigned by us to Asiacastor is rather high- (ectoloph) in Y. birgeri, which is absent in Y. ambiguus. crowned. In contrast to the situation in the p4 of Propa- Furthermore, the metalophs of the upper molars are some- laeocastor its hypostriid reaches the basis of the crown. what less posteriorly directed in Y. birgeri than in Y. am- The fossetids are parallel and inclined forwards. biguus. Judging by the figures in BOHLIN (1946) and WANG (1997) these differences seem to be beyond the range of C o n c l u s i o n s . – There are three beaver species individual variation in the associations from Kazakhstan represented in the Late Oligocene localities Altyn Schoky- and China, but are nevertheless a weak basis for distin- su, Akotau and Sayaken. Of these, Propalaeocastor kum- guishing species. For the time being we maintain the sta- bulakensis is common, P. schokensis is rare and Asiacas- tus quo because we have not directly compared specimens tor sp. is very rare. Although this conclusion is in agree- of the two species. ment with LOPATIN (2004), our identifications of individual specimens deviate from LOPATIN’s in many instances. Yindirtemys deflexus (TEILHARD DE CHARDIN, 1926) Pl. 6, Figs. 1–4 Ctenodactylidae GERVAIS, 1853 BENDUKIDZE, Tataromyinae LAVOCAT, 1961 1993 Yindirtemys sajakensis. – pl. 20, figs. 2, 3 (holotype: M2–M3 sin.). Remarks. – We follow WANG (1997) in assigning Locality: Sayaken. LAVOCAT Material and measurements (No. 9/33 is the the subfamily name Tataromyinae to (1961) in- holotype of Y. sajakensis): stead of to BOHLIN (1946) who used this term for the first No. Fig. Tooth Length × Width time, but in an informal sense. We also follow WANG Fragment of a left MATTHEW (1997) in including the genera Tataromys & 9/33 Pl. 6, Fig. 1 maxilla with M2 42.5 × 36.8 GRANGER, 1923, Yindirtemys BOHLIN, 1946 and Bounomys M3 43.5 × 41.0 WANG, 1994 into this subfamily. 9/14 Pl. 6, Fig. 3 p4 26.8 × 24.0 Fragment of a 9/21 Pl. 6, Fig. 2 mandible with m1 36.3 × 27.5 Yindirtemys BOHLIN, 1946 m2 46.8 × 35.3 9/12 Pl. 6, Fig. 4 m3 54.3 × 33.2 Yindirtemys birgeri BENDUKIDZE, 1993 Pl. 5, Figs. 8–11 R e m a r k s . – These Yindirtemys specimens, forming H o l o t y p e : Palate with P4–M3 sin. and P4–M3 dext., the hypodigm of Y. sajakensis, are more robust and a size- pl. 21, fig. 2 in BENDUKIDZE (1993). class larger than Y. birgeri. WANG (1997) correctly ob- 352 PALAEODIVERSITY 2, 2009 served that the teeth from Sayaken are, with exception of species from Altyn Schokysu defined by LOPATIN. In con- the p4, within the lower limit of the size range of Y. de- trast to these, the crowns of the teeth of our specimen are flexus. Although the protolophs of the M2 and M3 from very low, the sinuses shallow and wide and the mesolo- Sayaken are more transverse than in the holotype of Y. de- phids short. In these respects our specimen resembles the flexus (LOPATIN 2004), we follow WANG (1997) and con- dentitions of Allosminthus WANG, 1985 and Litodonomys sider the species Y. sajakensis a junior synonym of Y. de- WANG & QIU, 2000, genera that differ in grade-of-evolu- flexus. The dental morphology of Y. birgeri and the larger tion only. With the exception of the presence of an antero- Y. deflexus from Sayaken is strikingly similar, so it cannot conid, our specimen is very similar to Litodonomys huan- be excluded that these teeth could fit within the unknown gheensis, the poorly documented genotype from the Late size range of that species. For reasons given above, LOPA- Oligocene of the Lanzhou basin, China. Pending the de- TIN (2004) prefers to maintain Y. sayakensis and hypothe- scription of the very good material of Litodonomys from sises a deflexus-sajakensis and an ambiguus-birgeri lin- Mongolia collected by DAXNER-HÖCK we identify our eage. Considering that Y. deflexus from Sayaken is barely specimen as Litodonomys sp. within the lower limit of the size range of the type material of that species, this reconstruction seems improbable. Morphologically, the cheek teeth of the species Y. birgeri, Litodonomys sp. Y. deflexus from Sayaken and the type material of Y. de- Pl. 6, Fig. 5 flexus from Saint Jaques, Nei Mongol, China show a per- 1993 Parasminthus aff. tangingoli BOHLIN, 1946. – BENDUKIDZE, fect cline towards size increase, more robust cusps and pl. 16, figs. 4, 5. more forwards directed protoloph in the upper molars. L o c a l i t y : Altyn Schokysu. LOPATIN’S However, interpretation is in conflict with the Material and measurements: Miocene age supposed for the Aral fauna (LOPATIN 2004) No. Fig. Tooth Length × Width and the Oligocene age of Saint Jaques (RUSSELL & ZHAI Fragment of a right 1987). 15/23 Pl. 6, Fig. 5 mandible with m1 12.4 × 8.8 m2 11.8 × 10.7

Dipodidae FISCHER VON WALDHEIM, 1817 Muridae ILLIGER, 1811 The only dipodid specimen in the BENDUKIDZE collection is a fragment of a mandible with m1 and m2 from Tachyoryctoidinae SCHAUB, 1958 level four at Altyn Schokysu. This specimen, identified in BENDUKIDZE BOH- (1993) as Parasminthus aff. tangingoli The contents of the Tachyoryctoidinae and their sys- LO- LIN, 1946, was transferred to Parasminthus debruijni tematic position relative to other groups of (? fossorial) LOPATIN PATIN, 1999 by . Muridae have been disputed ever since the genera Tachy- LOPATIN (1999b, 2004) recognises three species of di- oryctoides BOHLIN, 1937 (type species: T. obrutschevi), podids in his material from Altyn Schokysu: Plesiosmin- Aralomys ARGYROPULO, 1939 (type species: A. gigas) and LOPATIN thus tereskentensis , 1999 from level one and Schaubeumys ARGYROPULO, 1939 (type species: S. aralen- Parasminthus debruijni and Bohlinosminthus cubitalus sis) were published. This last species later became the type LOPATIN LOPÉZ ANTOÑAN- , 1999 from level two. We follow of Argyromys SCHAUB, 1958 when he realised that it is a SEN LOPATIN ZAS & (2006) in considering Bohlinosminthus , murid and not a dipodid. The source of the confusion is BOHLIN 1999 a junior synonym of Parasminthus , 1946. that the dentitions of rodents that adapted to a fossorial LOPATIN Judging by the figures in (1999b, 2004) it seems mode of life in different geographical areas tend to devel- that the dental morphology and the size of the cheek teeth op similar dental characteristics regardless, it seems, of of B. cubitalus could fit within the range of variation of P. their phylogenetical relationship. tereskentensis, which has page priority. The difference The Tachyoryctoidinae have been considered to be re- between these species is essentially that P. tereskentensis lated to the Rhizomyinae (BOHLIN 1937, 1946, KOWALSKI has three-rooted M1 and M2 and B. cubitalus has four- 1974, LI & QIU 1980, BENDUKIDZE 1993), to the Spalacinae LOPATIN rooted M1 and M2. Unfortunately, (1999b, 2004) (FLY N N et al. 1985, FEJFAR 1972), to the Anomalomyinae does not report on the shape of the dipodid upper incisors (FEJFAR 1972, FLY N N et al. 1985), or have been regarded as in his sample. This probably means that these were not a separate group of fossorial murids (SCHAUB 1958, KLEIN recognised because they do not have the longitudinal HOFMEIJER & DE BRUIJN 1985, TYUTKOVA 2000). This dis- groove that is characteristic for Plesiosminthus. pute continues to remain unresolved as is demonstrated by It is peculiar that the only dipodid specimen in our col- LOPATIN (2004), who assigns Argyromys to the Spalacinae, lection cannot be referred to any of the well-documented BENDUKIDZE ET AL., OLIGOCENE MAMMALS FROM KAZAKHSTAN 353 but Tachyoryctoides, Aralomys and Eumysodon to the level, and fig. 3d an m2 of Eumysodon sp. If our interpre- Tachyoryctoidinae. tation of the association from the locality Ayaguz is cor- KALTHOFF (2000) has shown that the highly derived rect it contains Ayakosomys, Eumysodon, and Tachyoryc- “Schmelzmuster” (type 10) occurring in a lower incisor of toides, an association of genera that otherwise has never ?Tachyoryctoides from Altyn Schokysu is shared by a been found in one locality. Therefore it seems that the as- large number of murid subfamilies (i. e., the Rhizomyinae, sociation studied by TYUTKOVA (2000) may contain mate- the Spalacinae and the Anomalomyinae). Recent research rial from different time slices. (KALTHOFF pers. comm., 2007) suggests, however, that the Since we have seen only part of the material of genera lower incisor of Tachyoryctoides analysed in 2000 was here included into the Tachyoryctoidinae the classification misidentified, so Tachyoryctoides “Schmelzmuster” suggested below is necessarily tentative. The prevailing should be considered as not known. uncertainty is illustrated by the difference of opinion be- We consider Aralomys to be a junior synonym of tween the first and second author (O B and H d B) on the Tachyoryctoides and tentatively include the genera Argy- synonymy of T. gigas (ARGYROPULO, 1939) and T. obruts- romys, Eumysodon, Aralocricetodon BENDUKIDZE, 1993 chevi BOHLIN, 1937. Although the species T. gigas is listed and Ayakozomys TYUTKOVA, 2000 into the Tachyoryctoidi- below as a junior synonym of T. obrutschevi, BENDUKIDZE nae. considers these species to be different. Our reasons to consider the Rhizomyinae, Spalacinae, Tachyoryctoidinae BOHLIN, 1937 Anomalomyinae and Tachyoryctoidinae to be separate subfamilies are that the oldest representative of each of Tachyoryctoides BOHLIN, 1937 these groups: Prokanisamys arifi DE BRUIJN et al., 1981, Synonyms: Aralomys ARGYROPULO, 1939, Ayako- Debruijnia arpati ÜNAY, 1996, Anomalomys eviensis, somys TYUTKOVA, 2000 pro parte, fig. 3a in TYUTKOVA (2000). KLEIN HOFMEIJER DE BRUIJN & , 1985 and Tachyoryctoides T. obrutschevi BOHLIN, 1937 (genotype) obrutschevi BOHLIN, 1937 differ more from each other Synonyms: Aralomys gigas ARGYROPULO, 1939, than some of the later representatives of these groups do. Aralomys glikmani VORONTSOV, 1963, Ayakosomys sergiopo- TYUTKOVA Other evidence for this interpretation comes from the lis , 2000 pro parte. stratigraphical and geographical ranges of these sub- T. pachygnatus BOHLIN, 1937 BOHLIN families. The early history of the Rhizomyinae seems to Synonyms: T. intermedius , 1937, T. kokono- rensis LI & QIU, 1980. have taken place on the Indian subcontinent during the TYUTKOVA FLY N N R e m a r k . – We consider T. padre , 2000 a Miocene ( 1982), that of the Spalacinae in the Mid- nomen dubium because the hypodigm of that species con- dle East during the Miocene (ÜNAY 1996), that of the sists of two teeth that seem to belong to different species. Anomalomyinae in Central Europe during the Miocene TYUTKOVA (BOLLIGER 1999) and that of the Tachyoryctoidinae in Cen- Ayakosomys , 2000 tral Asia during the Oligo/Miocene (see below). A. sergiopolis TYUTKOVA, 2000 (genotype) Unfortunately, there is no consensus among specialists Eumysodon ARGYROPULO, 1939 about the number and identity of the genera and species of ARGYROPULO the Tachyoryctoididae. The main reason for the widely E. spurius , 1939 (genotype) Synonyms: E. orlovi ARGYROPULO, 1939, diverging opinions is that type species of the genera are ? Protalactaga borissiaki ARGYROPULO, 1939. based on either single specimens or on a very small series. R e m a r k . – LOPATIN (2004) considers the M1, M2 that TYUTKOVA A recent example of this problem is provided by are the holotype of Protalactaga borissiaki as the upper (2000), who defines her genus Ayakosomys (type species dentition of Schaubeumys aralensis ARGYROPULO, 1939, now A. sergiopolis) on an adequate collection, but also names a Argyromys aralensis. At the same time he correctly syn- new species of Aralomys (= Tachyoryctoides) on the basis onymises Schaubeumys woodi ARGYROPULO, 1939 with S. aralensis. Since S. aralensis as well as E. spurius are based of two teeth that do not seem to belong to the same taxon on lower dentitions of roughly the same size from the locali- (TYUTKOVA 2000: fig. 3c, c’, c’’ and fig. 3d, d’, d’’). Other ty Akespe, it remains uncertain whether the upper teeth specimens that may be referable to Ayakosomys have been from Akespe that are the holotype of Protalactaga borissia- published by KORDIKOVA & DE BRUIJN (2001) from the Mi- ki belong to either A. aralensis or E. spurius. ocene of southeast Kazakhstan under the name Tachy- Argyromys SCHAUB, 1958 oryctoidinae gen. A, sp. 1 and by YE et al. (2003) from the ARGYROPULO Early Miocene of Xinjiang (China) under the name Tachy- A. aralensis ( , 1939) (genotype) Synonyms: Schaubeumys woodi ARGYROPULO, 1939, oryctoidinae gen. et sp. nov. Judging by the drawings in ?Protalactaga borissiaki ARGYROPULO, 1939 (see remark TYUTKOVA (2000) fig. 3a seems to represent Tachyoryc- with E. spurius). toides obrutschevi, fig. 3b Ayakosomys sergiopolis, fig. 3c Aralocricetodon BENDUKIDZE, 1993 (the holotype of Aralomys padre) is a tooth that, on the basis of the drawing, cannot be identified at the genus A. schokensis BENDUKIDZE, 1993 (genotype) 354 PALAEODIVERSITY 2, 2009

Tachyoryctoides obrutschevi BOHLIN, 1937 No. Fig. Tooth Length × Width Pl. 7, Figs. 1–10, Pl. 8, Fig. 1 15/37 Pl. 6, Fig. 6 m1 dext. 36.1 × 25.3

For synonymy see above. R e m a r k s . – Among the Tachyoryctoides specimens L o c a l i t y : Altyn Schokysu. from Altyn Schokysu there is one m1 that seems to be Material and measurements: Fragment of a maxilla with M1–M3, No. 15/36; two M1, No. 15/31, 15/40; one somewhat too small to fit the (estimated) size range of the M2, No.15/101; one M3, No. 15/102; three m1, No. 15/32, 15/32a, species T. obrutschevi. Since this tooth is morphologically 15/32b; two m2, No. 15/33, 15/33a; two m3, No. 15/41, 15/100. indistinguishable from the others, we would probably have Length Width included it into that species. However, it appeared that the Tooth n Range Mean Range Mean collections from level C in Mongolia (DAXNER-HÖCK, pers. M1 3 42.0–50.3 47.0 34.7–41.6 39.3 com.) contain a smaller Tachyoryctoides species in asso- M2 1 33.4 34.0 ciation with T. obrutschevi. LOPATIN (2004) reached the M3 1 27.7 31.5 same conclusion and separated one M1, a fragment of an m1 3 36.6–42.4 40.0 29.7–34.3 31.4 M2 and one m2 as Tachyoryctoides sp. from the rest of the m2 2 37.9–38.2 38.1 33.7–35.5 34.6 m3 1/2 33.5 31.2–32.8 32.0 Altyn Schokysu material that he assigned to T. glikmani (VORONTZOV, 1963) and Aralomys gigas (ARGYROPULO, Locality: Sayaken. 1939). M a t e r i a l a n d m e a s u r e m e n t s : One fragment of a mandible with incisor and the m2, No. 9/50; one fragment of a mandible with part of the m2 and the m3, No. 9/31; isolated m2, Aralocricetodon BENDUKIDZE, 1993 No. 9/20; isolated m3, No. 9/20a.

No. Fig. Tooth Length × Width Aralocricetodon schokensis BENDUKIDZE, 1993 9/20 – m2 38.5 × 35.8 Pl. 7, Figs. 11–22 9/20a – m3 34.0 × 29.0 Material and measurements: Measurements Remarks. – Tachyoryctoides obrutschevi has a ro- were taken on A. schokensis casts in the comparative col- bust dentition. The anterocone of the M1 is entirely incor- lection of the department of Earth Sciences Utrecht. The porated into the anteroloph as in fossorial rodents. The originals, collected by KÄLIN, DAXNER-HÖCK, BOLLIGER upper cheek teeth with their more or less transverse pro- and DE BRUIJN from level two at Altyn Schokysu in 1994 toloph and metaloph and the lingually open sinus of the are housed in the Paleontological Institute, Russian Acad- M3 still show primitive cricetid characteristics. The lower emy of Sciences, Moscow. For further measurements and cheek teeth with their deep posteriorly directed sinusid are description see BENDUKIDZE (1993) and LOPATIN (2004). more derived. The anterior part of the m1 shows consider- Length Width Tooth n able individual variation: in some the anterior part of the Range Mean Range Mean longitudinal crest is absent and the metalophid curves M1 5 19.3–24.4 23.0 16.2–19.8 17.7 forwards reaching the anteroconid. In others the metaconid M2 4 16.8–18.4 17.6 14.0–16.6 15.9 is connected to the protoconid and/or the metalophid by M3 4 14.2–15.3 14.8 14.3–15.1 14.8 what seem to be the remnants of the original mesolophid m1 4 20.2–23.9 21.3 13.6–15.5 14.5 and the labial part of the metalophid. These ridges may m2 4 17.4–20.4 18.6 14.2–16.3 15.2 enclose an enamel lake. m3 2 18.2–19.9 19.1 13.5–16.3 14.4 The diastema of the mandible is very long and shallow (Pl. 8, Fig. 1). Apparently the function of the lower incisor R e m a r k s . – The genus Aralocricetodon was origi- (digging tool?) was very different from that of the cheek nally defined on the basis of a single worn M1, but the teeth. KOWALSKI (1974) describes the mandible of T. species is now well represented in collections (LOPATIN obrutschevi as robust. In our opinion it is large relative to 2004). BENDUKIDZE (1993) as well as LOPATIN (2004) con- the size of the cheek teeth rather than robust as in the Rhi- sider the genus Aralocricetodon to be a member of the zomyinae. Cricetodontinae sensu stricto. However, the rather large, robust, semi-hypsodont cheek teeth of Aralocricetodon that show strong resemblance to those of the Middle Mio- Tachyoryctoides sp. cene Rhizomyinae from the Indian subcontinent, suggest Pl. 6, Fig. 6 that this species was adapted to a fossorial life-style. In particular the upper dentition is strikingly similar to that L o c a l i t y : Altyn Schokysu. of the type species of Kanisamys WOOD, 1937 (K. indicus). Material and measurements: The small brachyodont gracile teeth of roughly contempo- BENDUKIDZE ET AL., OLIGOCENE MAMMALS FROM KAZAKHSTAN 355 raneous species of Cricetodon (DE BRUIJN et al. 1993) on MANES, 1995) can only be verified on the basis of better the other hand, are very different. Since the early true material than is available to us. Rhizomyinae (Prokanisamys DE BRUIJN et al. 1981, Kanisamys and Brachyrhizomys FLY N N, 1982) seem to form a clade that developed on the Indian subcontinent Eucricetodon THALER, 1966 during the Miocene, there is good reason to assume that Eucricetodon (Atavocricetodon) FREUDENTHAL, 1996 the similarity of Aralocricetodon and Kanisamys is the result of convergent evolution. For the time being it seems Eucricetodon (Atavocricetodon) occasionalis LOPATIN, therefore best to assign Aralocricetodon to the same clade 1996 as the other Oligocene fossorial murids of Central Asia: Pl. 9, Figs. 1–5 the Tachyoryctoidinae. pro parte 1993 Eucricetodon sajakensis. – BENDUKIDZE, pp. 46–52, pl. 17, figs. 4, 5, pl. 18, figs. 1–4. L o c a l i t y : Altyn Schokysu. Eucricetodontinae MEIN & FREUDENTHAL, 1971 Material and measurements: Fragment of a mandible with m1–m3 (No. 15/25), fragment of a mandible with The few specimens from Altyn Schokysu, Akespe and m1–m2 (No. 15/24), fragment of a mandible with m1–m2 (No. 15/26). Sayaken referable to Eucricetodon THALER, 1966 show all FREU- Length Width the characteristics of the subgenus Atavocricetodon Tooth n DENTHAL, 1996: a short anterocone(id) in the M1 and m1, a Range Mean Range Mean free-ending anterior arm of the protocone in the M1 that is m1 3 16.6–19.0 17.9 11.5–12.5 12.0 m2 3 16.0–18.2 16.8 12.5–14.4 13.5 parallel and subequal in length to the anterior arm of the m3 1 15.6 12.9 hypocone, a forwards directed protoloph and metaloph in the M2. L o c a l i t y : Akespe. The three partial mandibles from Altyn Schokysu in Material and measurements: BENDUKIDZE BENDUKIDZE the collection ( 1993, pl. 17, figs. 2 No. Fig. Tooth Length × Width and 3; pl. 18, figs. 3 and 4) were originally assigned to the 11/9 Pl. 9, Fig. 1 M1 19.9 × 13.3 inadequately known Eucricetodon aff. caducus (SHEVYRE- 11/9a Pl. 9, Fig. 2 M2 15.5 × 14.1 VA, 1967), now E. occasionalis LOPATIN, 1996, and to E. sajakensis BENDUKIDZE, 1993. Since the individual varia- R e m a r k s . – The size of the few molars from Altyn tion within homogeneous Eucricetodon (Atavocricetodon) Schokysu and Akespe, as measured by us, is somewhat associations is, as a rule, considerable, we consider the larger than the measurements given by LOPATIN (2004) for minor differences in size and morphology that induced his E. occasionalis material from Altyn Schokysu, but this BENDUKIDZE (1993) to recognise two species in the mate- may be due to differences in the measuring technique rial from Altyn Schokysu, to be within the range of varia- used. tion of one species. The few specimens from Sayaken that form the hypodigm of Eucricetodon sajakensis BENDUKIDZE, 1993 are, Eucricetodon (Atavocricetodon) sajakensis (BENDUKIDZE, with the exception of the presence of the posterior arm of 1993) the hypoconid in the m2, morphologically very similar to Pl. 9, Figs. 6, 7 the specimens of Eucricetodon occasionalis from Altyn Schokysu and Akespe, but they are a size-class larger. Locality: Sayaken. Material and measurements: Since the number of Eucricetodon (Atavocricetodon) species described from European Oligocene localities on No. Fig. Tooth Length × Width the basis of subtle differences is considerable, it is very Fragment of a 9/32 Pl. 9, Fig. 6 maxilla with M1 23.0 × 16.5 difficult, if not impossible, to evaluate the features of the M2 17.4 × 18.3 few specimens from Kazakhstan. The presence of long Fragment of a mesolophs in the M1, M2 and a strong posterior arm of the 9/34 Pl. 9, Fig. 7 mandible with hypoconid in the m2 in Eucricetodon (Atavocricetodon) m2 18.5 × 16.3 sajakensis differentiates this species from the European members of the subgenus, but the difference between Ata- R e m a r k s . – The specimens from Sayaken are clear- vocricetodon occasionalis on the one hand and species ly larger than the teeth of A. occasionalis from Altyn such as Atavocricetodon atavoides FREUDENTHAL, 1996 Schokysu. Moreover, the posterior arm of the hypoconid (subgenotype), Atavocricetodon hugueneyae FREUDEN- of the m2 is strong in A. sajakensis, but absent in A. oc- THAL, 1996 and Atavocricetodon nanus (PELAEZ CAMPO- casionalis. 356 PALAEODIVERSITY 2, 2009

4. List of the small mammals of the Aral local fauna Aral Formation in this area to the Late Oligocene and the represented in the BENDUKIDZE collection upper part of the formation to the Early Miocene. LUCAS et al. (1998) argue, on the basis of sequence stratigraphy and Erinaceomorpha GREGORY, 1910 mammal biochronology, that all of the Aral Formation is Erinaceidae FISCHER, 1814 of Late Oligocene Age. Part of this discrepancy is due to Galericinae gen. et sp. indet. the lack of possibilities for radiometric and/or magneto- Amphechinus sp. stratigraphic calibration of these deposits. Moreover, the Amphechinus cf. minimus (BOHLIN, 1942) very condensed section of the Aral Formation may be ex- Soricomorpha GREGORY, 1910 pected to be time-transgressive over larger distances. Talpidae FISCHER, 1814 It has been suggested (BENDUKIDZE 1993, LOPATIN 1996, Theratiskos compactus (LOPATIN, 2004) KORDIKOVA et al. 2003) that existing collections from dif- Pseudoparatalpa lavroi (BENDUKIDZE, 1993) ferent bone bearing “levels” allow the distinction of age- Heterosoricidae VIRET & ZAPFE, 1951 determined sub-associations within the Aral local fauna. Gobisorex kingae SULIMSKI, 1970 However, the observed differences may well be due to col- Lagomorpha BRANDT, 1855 lecting bias, local factors that influenced the accumulation Leporidae FISCHER VON WALDHEIM, 1817 of fossils and differences in sample size rather than to dif- Desmatolagus simplex (ARGYROPULO, 1940) ference in age. Moreover, the so-called “levels” at Altyn Desmatolagus periaralicus LOPATIN, 1998 Schokysu are most of the time lenses that are situated at Desmatolagus veletus LOPATIN, 1998 appreciable distances laterally, so their vertical distance Ochotonidae THOMAS, 1897 can only be estimated by measuring their distance relative Sinolagomys kansuensis BOHLIN, 1937 to the base of the formation. We therefore consider the col- Rodentia BOWDICH, 1821 lection from Altyn Schokysu as one assemblage, a “local Castoridae HEMPRICH, 1820 fauna”. Propalaeocastor schokensis (BENDUKIDZE, 1993) Our revised fauna list of the Aral local fauna confirms Propalaeocastor kumbulakensis LYTSCHEV, 1970 the conclusion of BENDUKIDZE (1997) that the differences Asiacastor sp. in composition between the associations from the Aral Ctenodactylidae GERVAIS, 1853 Formation in Kazakhstan and the Hsanda Gol Formation Yindirtemys birgeri BENDUKIDZE, 1993 of Mongolia reflect ecological rather than age differences. Yindirtemys deflexus (TEILHARD DE CHARDIN, 1926) In spite of the absence of some fauna elements that are Dipodidae FISCHER VON WALDHEIM, 1817 characteristic for either the Kazakh or the Mongolian as- Litodonomys sp. semblages, the presence of Aralocricetodon schokensis Muridae ILLIGER, 1811 and Tachyoryctoides obrutchevi in the assemblages from Tachyoryctoides obrutschevi BOHLIN, 1937 biozone C/C’ of HÖCK et al. (1999) in Mongolia and the Tachyoryctoides sp. Aral Formation suggests a correlation of both fauna com- Aralocricetodon schokensis BENDUKIDZE, 1993 plexes with the Chinese Tabenbulukian Mammal Age (= Eucricetodon (Atavocricetodon) occasionalis LO- Late Oligocene). Moreover, our assemblages do not con- PATIN, 1996 tain a single taxon known from the Early Miocene of Eucricetodon (Atavocricetodon) sajakensis (BEN- Mongolia, the Ustyurd area of Kazakhstan and the Aktau DUKIDZE, 1993) Mountains (HÖCK et al. 1999, KORDIKOVA et al. 2003, KORDIKOVA & DE BRUIJN 2001), so we fully agree with LU- CAS et al. (1998) that the Aral Formation at the Kumbulak 5. The age of the assemblage cliffs, the Altyn Schokysu escarpment and Akotau hill is Late Oligocene in age. The geology, sequence stratigraphy, palaeobotany, in- vertebrate palaeontology and vertebrate palaeontology of the Oligo/Miocene marine to continental deposits of the 6. References North Aral area have been intensively studied (AKHMETIEV AKHMETIEV, M. A. (1994): An essay on the geological studies of LUCAS LOPATIN KORDIKOVA 1994, et al. 1998, 2004, et al. the northern Aral region and north-eastern Ustyurt. – Oligo- 2003). cene-Miocene boundary in Kazakhstan: field-excursion Although the majority of Russian scientists consider guide book, Kazakhstan, August 16–28, 1994: 9–11. the Aral Formation as exposed in the Kumbulak cliffs, ARGYROPULO, A. I. (1939): New Cricetidae (Glires, Mammalia) from the Oligocene of middle Asia. – Comptes Rendus along the Altyn Schokysu escarpment and in Akotau (= (Doklady) de l’Academie des Sciences de l’URSS, 1: 111– Aktau) hill to be of Early Miocene age, AKHMETIEV (1994) 114. and KORDIKOVA et al. (2003) attribute the lower part of the ARGYROPULO, A. I. (1940): A survey of the findings of Rodentia BENDUKIDZE ET AL., OLIGOCENE MAMMALS FROM KAZAKHSTAN 357

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Adresses of the authors: OLEG G. BENDUKIDZE, Institute of Paleobiology, Georgian National Museum, Niagvris street, Tbilissi, Georgia HANS DE BRUIJN, Department of Geosciences, P.O. Box 80021, 3508 TA Utrecht, The Netherlands E-mail: [email protected] LARS W. VAN DEN HOEK OSTENDE, Nationaal Natuurhistorisch Museum, Naturalis, P.O. Box 9517, 2300 RA Leiden, The Netherlands E-mail: [email protected]

Manuscript received: 25.2.2009, accepted: 16.7.2009. 360 PALAEODIVERSITY 2, 2009

Plate 1

Fig. 1. Galericinae gen. et sp. indet., m2; No. 15/1; Altyn Schokysu. Figs. 2–5. Amphechinus sp.; Altyn Schokysu. Fig. 2. P4; No. 15/2. Fig. 3. Damaged M1; No. 15/4. Fig. 4. M2; No. 15/5. Fig. 5. Fragment of a mandible with p4–m3 (only the trigonid of the m1 preserved); No.15/6. Figs. 6–8. Amphechinus cf. minimus; Altyn Schokysu. Fig. 6. Damaged M1; No. 15/7. Fig. 7. m1; No. 15/8a. Fig. 8. Fragment of a mandible with m2–m3; No. 15/8b. Fig. 9. Theratiskos compactus, fragment of a mandible with m2; No. 15/9; Altyn Schokysu. Fig. 10. Pseudoparatalpa lavroi, fragment of a mandible with m1–m2; No. 15/10; Altyn Schokysu. Figs. 11–13. Gobisorex kingae; Altyn Schokysu. Fig. 11. M1; No. 15/12. Fig. 12. Fragment of mandible with damaged m1 and m2; No. 15/11. Fig. 13. Fragment of mandible with m1–m3; No. 15/13. BENDUKIDZE ET AL., OLIGOCENE MAMMALS FROM KAZAKHSTAN 361 362 PALAEODIVERSITY 2, 2009

Plate 2

Figs. 1–2. Desmatolagus simplex; Altyn Schokysu. Fig. 1. Fragment of a maxilla with P3–M2; No. 15/19. Fig. 2. Fragment of a mandible with p3–m2; No. 15/20. Fig. 3. Sinolagomys kansuensis, fragment of a maxilla with the alveole of the P2 with damaged P3 and P4–M2; No. 15/23; Altyn Schokysu. Fig. 4. Sinolagomys kansuensis, fragment of a mandible with p3–m2; No. 16/6; Akotau. Fig. 5. Desmatolagus periaralicus, M2; No. 15/21; Altyn Schokysu. Figs. 6–10. Desmatolagus veletus; Altyn Schokysu. Fig. 6. P3; No. 15/14. Fig. 7. P4; No. 15/15. Fig. 8. M1; No. 15/16. Fig. 9. p3; No. 15/18. Fig. 10. m1–2; No. 15/18a. BENDUKIDZE ET AL., OLIGOCENE MAMMALS FROM KAZAKHSTAN 363 364 PALAEODIVERSITY 2, 2009

Plate 3

Figs. 1–3. Propalaeocastor schokensis; Altyn Schokysu. Fig. 1. Fragment of a maxilla with P4–M3 (M3 broken); No. 15/43. Fig. 2. Fragment of a maxilla with P4–M2; No. 15/49. Fig. 3. P4; No. 15/47. Figs. 4–7. Propalaeocastor kumbulakensis, P4;. Fig. 4. No. 16/50; Akotau. Fig. 5. No. 16/52; Akotau. Fig. 6. No. 15/59; Altyn Schokysu. Fig. 7. No. 9/18; Sayaken. BENDUKIDZE ET AL., OLIGOCENE MAMMALS FROM KAZAKHSTAN 365 366 PALAEODIVERSITY 2, 2009

Plate 4

Figs. 1–2. Propalaeocastor schokensis; Altyn Schokysu. Fig. 1. Mandible with p4–m3; No. 15/48. – a. Occlusal view. b. Labial view. Fig. 2. Fragment of a mandible with p4–m3 (p4 damaged); No. 15/45. Figs. 3–4. Propalaeocastor schokensis, two teeth of the same individual; No. 9/22 (Holotype of Capacikala sajakensis BENDUKIDZE, 1993); Sayaken. Fig. 3. p4. Fig. 4. m1. BENDUKIDZE ET AL., OLIGOCENE MAMMALS FROM KAZAKHSTAN 367 368 PALAEODIVERSITY 2, 2009

Plate 5

Figs. 1–6. Propalaeocastor kumbulakensis. Fig. 1. M1-2; No. 15/54; Altyn Schokysu. Fig. 2. M1-2; No. 16/110; Akotau. Fig. 3. p4; No. 9/40; Sayaken. Fig. 4. m1-2; No. 15/53; Altyn Schokysu. Fig. 5. m1-2; No. 15/55; Altyn Schokysu. Fig. 6. m3; No. 15/58; Altyn Schokysu. Fig. 7. Asiacastor sp., p4; No. 16/113; Akotau. Figs. 8–9. Yidirtemys birgeri; Altyn Schokysu. Fig. 8. Palate with left and right P4–M3 (Holotype); No. 15/28. Fig. 9. p4; No. 15/29. Figs. 10–11. Yindirtemys birgeri, two teeth from the same mandible; No. 16/9; Akotau. Fig. 10. m2. Fig. 11. m3. BENDUKIDZE ET AL., OLIGOCENE MAMMALS FROM KAZAKHSTAN 369 370 PALAEODIVERSITY 2, 2009

Plate 6

Figs. 1–4. Yindirtemys deflexus; Sayaken. Fig. 1. Fragment of a maxilla with M2, M3 (Holotype of Y. sajakensis); No. 9/33. Fig. 2. Fragment of a mandible with m1, m2; No. 9/21. Fig. 3. p4; No. 9/14. Fig. 4. m3; No. 9/12. Fig. 5. Litodonomys sp., fragment of a mandible with m1, m2; No. 15/23; Altyn Schokysu (not to scale with the other figures on plate 6). Fig. 6. Tachyoryctoides sp., m1; No. 15/37; Altyn Schokysu. BENDUKIDZE ET AL., OLIGOCENE MAMMALS FROM KAZAKHSTAN 371 372 PALAEODIVERSITY 2, 2009

Plate 7

Figs. 1–10. Tachyoryctoides obrutschevi; Altyn Schokysu. Fig. 1. Fragment of a maxilla with M1–M3; No. 15/36. Fig. 2. M1; No. 15/40. Fig. 3. M1; No. 15/31. Fig. 4. m1; No. 15/32. Fig. 5. m1; No. 15/32a. Fig. 6. m1; No. 15/32b. Fig. 7. m2; No. 15/33. Fig. 8. m2; No. 15/33a. Fig. 9. m3; No. 15/41. Fig. 10. m3; No. 15/100. Figs. 11–22. Aralocricetodon schokensis; Altyn Schokysu (casts of specimens collected by KÄLIN, BOLLIGER, DAXNER-HÖCK & DE BRUIJN housed in the comparative collection of the Department of Earth Sciences of the University of Utrecht; No. U. 2271. Figs. 11–12. M1. Figs. 13–14. M2. Figs. 15–16. M3. Figs. 17–18. m1. Figs. 19–20. m2. Figs. 21–22. m3. BENDUKIDZE ET AL., OLIGOCENE MAMMALS FROM KAZAKHSTAN 373 374 PALAEODIVERSITY 2, 2009

Plate 8

Fig. 1. Tachyoryctoides obrutschevi, fragment of a mandible with the I1 and m2; No. 9/50; Sayaken. – a. Labial view. b. Occlusal view. BENDUKIDZE ET AL., OLIGOCENE MAMMALS FROM KAZAKHSTAN 375 376 PALAEODIVERSITY 2, 2009

Plate 9

Figs. 1–2. Eucricetodon (Atavocricetodon) occasionalis, two teeth from the same individual; Akespe. Fig. 1. M1; No. 11/9. Fig. 2. M2; No. 11/9a. Figs. 3–5. Eucricetodon (Atavocricetodon) occasionalis; Altyn Schokysu. Fig. 3. Fragment of a mandible with m1–m2; No. 15/24. Fig. 4. Fragment of a mandible with m1–m3; No. 15/25. Fig. 5. Fragment of a mandible with m1–m2; No. 15/26. Figs. 6–7. Eucricetodon (Atavocricetodon) sajakensis; Sayaken. Fig. 6. Fragment of a maxilla with M1–M2; No. 9/32. Fig. 7. Fragment of a mandible with m2; No. 9/34. BENDUKIDZE ET AL., OLIGOCENE MAMMALS FROM KAZAKHSTAN 377

A Revision of Late Oligocene Associations of Small Mammals from the Aral Formation (Kazakhstan) in the National Museum of ­Georgia, Tbilissi

A Revision of Late Oligocene Associations of Small Mammals from the Aral Formation (Kazakhstan) in the National Museum of Georgia, Tbilissi