FRAGMENTA MINERALOGICA ET PALAEONTOLOGICA. Allocricetus ehiki Schaub, 1930 (Rodentia, Mammalia) finds from Villány 3 and Esztramos 3 (Hungary)

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1 FRAGMENTA MINERALOGICA ET PALAEONTOLOGICA 16. BUDAPEST p Allocricetus ehiki Schaub, 1930 (Rodentia, Mammalia) finds from Villány 3 and Esztramos 3 (Hungary) by J. HÍR Abstract: The metrical and morphological analyses of two Pliocene Allocricetus populations are given. The ancestral morphotypes in these assemblages are more frequent than in the Pleistocene Allocricetus materials. On the basis of the metrical features we presume the beginning of the separation the Allocricetus bursae from the, Allocricetus ehiki in the fauna of Esztramos 3. INTRODUCTION The purpose of this paper is to give the metrical and morphological analysis of the small-sized hamster material of two Hungarian Pliocene paleovertebrate localities. These localities are well known in the Hungarian, and in the international vertebrate paleontological literature, but the standard of the elaboration and documentation of the finds is fallen behind the recent requirements. The cricetids have a general importance in the biostratigraphy of the Late Neogene. The author studied fossil non arvicoloid cricetids (Hír 1989, 1992) and in the course of this he effected the description of the material of Villány 3 and Esztramos 3. LOCALITIES The classical locality of Villány 3 is situated in Southern Hungary in the limestone quarry of Villány, Templom Hill (Kormos 1937, Kretzoi 1956, Jánossy 1979, 1986). The elaborated material of this article was collected by D. Jánossy in 1979 with washing and sieving. (This method was not applied by the former researchers.) Esztramos is another prominent group of the Hungarian Plio - Pleistocene paleovertebrate localities. It is situated near the northernmost point of Hungary in the Aggtelek Karstic Region (Jánossy 1972a, 1972b, 1973, 1974, 1979, 1986, Jánossy & Kordos 1976, Jánossy & Van der Meulen 1975). Villány 3 and Esztramos 3 were described by Jánossy (1979, 1986) as Lower Pleistocene localities. After the acceptance of the lower chronostratigraphic boundary of the Pleistocene at 1.8 MY, (Zagwijn 1985) these faunas have fallen to the Pliocene, MN 17. (De Bruijn et al. 1990).

2 METHODS The morphological examination follows the nomenclature of Mein & Freudenthal (1971). The creation of the different morphotypes is based on the presence or absence of certain elements of the toothcrown. The measurements were taken after Pradel (1981,1988): L M 1-3, Lm 1-3: length of upper and lower tooth-rows. L: length of the toothcrown. Wa: anterior width of the toothcrown measured across the protocone-paracone on the upper teeth and across the protoconid-paraconid on the lower teeth. Wp: posterior width of the toothcrown measured across the hypocone-metacone on the upper teeth and across the hypoconid-metaconid on the lower teeth. The measurements were taken by the ocularmicrometer of a stereomicroscope to an accuracy of 0.01 mm. The data are elaborated by the basic statistic parameters and listed in the appendix. PRESENTATION OF DATA Allocricetus ehiki SCHAUB, 1930 Material: Esztramos 3: Villány 3: 3 complete maxillae, 14 Ml, 8 M2, 1 M3, 7 complete mandibulae, 12 ml, 16 m2,13 m3; 2 complete maxillae, 9 Ml, 5 M2, 2 M3, 17 complete mandibulae, 6 ml, 14 m2, 11 m3. Sum total: 198 teeth. Measurements: LM1-3 no. min. Esztramos Villány Vil.3.-Esz Lml-3 no. min. Esztramos Villány Vil.3.-Esz Ml Esztramos 3. no.17, min. L Wa Wp Villány 3. no. 11, min. L Wa Wp

3 Vil.3.-Esz.3. no.28, min. L 2.00 Wa 0.99 Wp 1.25 M2 Esztramos 3. no.ll, min. L 1.54 Wa 1.26 Wp 1.13 Villány 3. no.7, min. L 1.67 Wa 1.33 Wp 1.26 Vil.3.-Esz.3. no. 18, min. L 1.54 Wa 1.26 Wp 1.13 M3 Esztramos 3. no.4, min. L 1.26 Wa 1.15 Villány 3. no.4, min. L 1.44 Wa 1.19 Vil.3.-Esz.3. no.8, min. L 1.26 Wa 1.15 m 1 Esztramos 3. no. 19, min. L 1.75 Wa 0.62 Wp 1.06 Villány 3. no.23, min. L 1.95 Wa 0.74 Wp 1.18 Vil.3.-Esz.3. no.42, min. L 1.75 Wa 0.70 Wp

4 m 2 Esztramos 3. no.23, min. L 1.47 Wa 1.18 Wp 1.12 Villány 3. no.31, min. L 1.50 Wa 1.29 Wp 1.29 Vil.3.-Esz.3. no.54, min. L 1.47 Wa 1.18 Wp m 3 Esztramos 3. no.20, min. L 1.39 Wa 1.09 Villány 3. no.28, min. L 1.48 Wa 1.18 Vil.3.-Esz.3. no.48, min. L 1.39 Wa M 1 Morphological distribution: DESCRIPTION AND REMARKS A: preanterocone cingulum (PAC)+, Parastyle (PAST) +: 12 B: PAC -, PAST +: 1 C: PAC 4-, PAST -: 9 D: PAC -, PAST -: 3 E: PAC+,PAST+,mesolophe(ML)orentostyle(ENST) + : 3 The frequency of the PAC and PAST is rather high. The mesolophe on the E- type molars is a feature of the Miocene cricetids (Kordos 1987). In the Upper Biharian Allocricetus bursae populations of Hungary (Hír 1989, 1992) the PAC and the PAST have lower frequency and the E-type is absolutely missing. On the scatter diagram of the M 1 molars the metrical bimodality is clearly visible (Fig. 1) and it is confirmed by other Allocricetus ehiki materials. The previous authors (Fahlbusch 1969, Pradel 1988) have not commented this phenomenon, because the teeth were very few in the material of them. The bimodal distribution is characteristic at the Ml, M2 and m2 molars (Figs. 1, 2, 5) but it was not recorded at the M3, ml and m3 molars (Figs. 3, 4, 6). Among the few complete

5 maxillae there are two specimens, where the large sized character is consequent on Ml and M2 molars (Villány 3 SN max. no. 3, Villány 3 F max. no. 1). In the A. éhiki Ml material the metrical difference is remarkable, but not sufficient for establish a distinct subspecies. Another interesting feature is the considerable difference between the anterior and posterior widths Wa-Wp (Fig. 8). The examination of this character was initiated by Pradel (1981,1988). He found very small difference between the fossil A, bursae material of Poland and a recent Cricetulus migratorius population from Syria. This parameter is definitely larger in the Hungarian Allocricetus populations than in the Polish Allocricetus materials. M2 Morphological distribution: A: B: C: D: metalophule (MTL) +, MTL -, MTL +, MTL -, mesolophe (ML) ML ML ML 1 5 (Fig. 9, c) 12 In the morphological composition there is no remarkable difference compared to the Hungarian Pleistocene populations. The scatter diagram of these teeth is bimodal similarly to the Ml molars. M3 All the 4 primary cusps are well developed. The reduction of the para-, hypo-, and metacone is not so characteristic than at Middle- and Upper Pleistocene Allocricetus and recent Cricetulus. The only peculiarity on one tooth is an extra enamel branch between the metacone and the anterocingulum (Fig. 9, d). m 1 Morphological distribution: A: PAC-, ML- B: PAC-, ML -, anteroconid undivided C: PAC-, ML+, D: PAC-, ML -, anteroconid 3-parted CD: PAC-, ML+, anteroconid 3-parted CB: PAC-, ML+, anteroconid undivided The preanteroconid cingulum (PAC) is equal to the anterolingual conulid in Reig (1977), and to the additional furrows on the front wall in Pradel (1982). In the investigated Pleistocene Allocricetus populations (Hír 1989, 1992) the PAC is occurred on 2-20% of the ml molars. In the material of Villány 3 and Esztramos 3 it is completely missing. On the majority of the teeth the anteroconid is divided in two parts by a depression according to the descriptions of Fahlbusch (1969) and Pradel (1981, 1988). On one molar the anteroconid is undivided (Fig. 10, b) and another two specimens have tripartite anteroconid (Fig. 10, d-e). These rare types are unknown in the Pleistocene Allocricetus materials, but occurred in the dental

6 pattern of Miocene cricetids (Daxner-Höck 1972). The presence of rudimentary mesolophids is ancestral feature too (Fig. 10, c). On the scatter diagram of the ml molars a part of the teeth from Esztramos 3 is smaller sized but not separated from the mass of the other molars. m 2 Morphological distribution: A: ALC+, ML+ 12 B: ALC-, ML+ 16 C: ALC+, ML- 19 D: ALC-, ML- 7 In the dental pattern the presence of mesolophid and the antero-lingual cingulum (ALC) is much more frequent than in the Pleistocene materials (Fig. 11). On the scatter diagram (Fig. 5) a weakly separated large sized group is visible. In the distribution of the Wa-Wp there is not any remarkable difference from the cited Polish materials contrast with the Ml - ml molars. m 3 Morphological distribution: A: ALC+, ML+ 29 G: ALC+, ML- 1 X: ALC- ML+ 15 E: ALC- ML- 3 The ALC is frequent and well developed (Fig. 12). The antero-buccal cingulum is constant. There is only one molar in the Esztramos 3/8 sample with week ABC (Fig. 12, d). The building up of the mesolophid region is very variable (Fig. 12). CONCLUSIONS The Allocricetus ehiki populations of Villány 3 and Esztramos 3 have some ancestral features in the morphology (mesolophe on Ml, mesolophid on ml, undivided or 3-parted anteroconid, frequent antero-lingual cingula on m2-3) differentiate it from Pleistocene Allocricetus. The most important metrical features are the next. The Ml and ml teeth are more trapezoidal than the same molars in the Polish Pleistocene and Pliocene and in the recent Cricetulus material. The assemblage of Villány 3 is rather homogeneous but the size distribution of teeth from Esztramos 3 is wider or bimodal (e.g. m2, Fig. 5). The two-grouped distribution is visible on the diagram of the Lml-Lm3 relation too (Fig. 7). As it appears to me the tendency of the separation of Allocricetus bursae from Allocricetus ehiki begins in the fauna of Esztramos 3. On this ground the slightly younger age of Esztramos 3 is possible. It was supposed also by Jánossy (1979,1986) on the basis of the faunistical composition of the two localities.

ACKNOWLEDGEMENTS The author would like to express his sincere thanks to Dénes JÁNOSSY retired director of the Paleontological Collection of the Hungarian Natural History Museum for loaning the

7 ACKNOWLEDGEMENTS The author would like to express his sincere thanks to Dénes JÁNOSSY retired director of the Paleontological Collection of the Hungarian Natural History Museum for loaning the material and to Attila VÖRÖS the present director for the possibility of the publication. REFERENCES DAXNER-HÖCK, G. (1972): Cricetinae aus dem Alt-Pliozän vom Eichkogel bei Mödling (Niederösterreich) und von Vösendorf bei Wien. - Paläont. Z., 46: DE BRUIJN, H. et al. (1990): Report of the RCMNS working group on fossil mammals (five appendices). Reisenburg pp. (manuscript) FAHLBUSCH, V. (1969): Pliozäne und Pleistozäne Cricetinae (Rodentia, Mammalia) aus Polen. - Acta Zool. Cracov., 14: HÍR, J. (1989): Revised investigation of the Allocricetus material of the Tarkő rock-shelter. - Fol. hist.-nat. Mus. Matr., 14: HÍR, J. (1992): Hamster remains (Rodentia, Mammalia) from the Lower Pleistocene of the Hajnóczy Cave. - Fragm. Min. et Pal., 15: JÁNOSSY, D. (1972a): Middle Pliocene microvertebrate Fauna from Osztramos Loc. 1. (Northern Hungary). - Annales Hist.-nat. Mus. Nat. Hung., 64: JÁNOSSY, D. (1972b): Ein kleiner Hystrix aus dem Altpleistozän der Fundstelle Osztramos 7. (Nordungarn). - Vertebr. Hung., 13: JÁNOSSY, D. (1973): The boundary of the Plio-Pleistocene based on the Microfauna in North Hungary (Osztramos, 7). - Vertebr. Hung., 14: JÁNOSSY, D. (1974): New "Middle Pliocene" Microvertebrate fauna from Northern Hungary (Osztramos, Loc.9.). - Fragm. Min. et Pal., 5: JÁNOSSY, D. (1979): A magyarországi pleisztocén tagolása gerinces faunák alapján (Subdivision of the Pleistocene of Hungary on the basis of the vertebrate faunas). - Akadémiai Kiadó, 207 pp. (in Hungarian). JÁNOSSY, D. (1986): Pleistocene Vertebrate Faunas of Hungary. - Akadémiai Kiadó - Elsevier, 209 pp. JÁNOSSY, D. & KORDOS, L. (1977): Az Osztramos gerinces lelőhelyeinek faunisztikai és karsztmorfológiai áttekintése (1975-ig) (Faunistical and karst-morphological review of the vertebrate localities at Osztramos (until 1975)). - Fragm. Min. et Pal., 8: (in Hungarian). JÁNOSSY, D. & VAN DER MEULEN, A. (1975): On Mimomys (Rodentia) from Osztramos 3, North Hungary. - Proc. Kon. Nederl. Akad. Wet., Ser. B, 78: KORDOS, L. (1987): Karstocricetus skofleki gen. n., sp. n. and the evolution of Late Neogene Cricetidae in the Carpathian Basin. - Fragm. Min. et Pal., 13: KRETZOI, M. (1956): Die altpleistozänen Wirbeltierfaunen des Villányer Gebirges. - Geol. Hung., Ser. Paleont., 27: KORMOS, T. (1937): Zur Geschichte und Geologie der ober-pliozänen Knochenbreccien des Villányer Gebirges. - Math. Term. Tud. Ért., 56: PRADEL, A. (1981): Biometrical remarks on the hamster Cricetulus migratorius (Pallas, 1773)(Rodentia, Mammalia) from Krak des Chevaliers (Syria). - Acta Zool. Cracov., 25:

8 PRADEL, A. (1988): Fossil hamsters (Cricetinae, Rodentia) from the Pliocene and Quaternary of Poland. - Acta Zool. Cracov., 31: REIG, O. (1977): A proposed unified nomenclature for the enamelled components of the molar teeth of the Cricetidae (Rodentia). - J. Zool., 181: ZAGWIJN, W. (1985): An outline of the Quaternary stratigraphy of the Netherlands. - Geologie en Mijnbow, 64: Author's address: Dr. János HÍR Pásztó, P.O.B Hungary Appendix (data in mm) 1. LM 1-3 Esztramos 3/3: Esztramos 3/8: Villány 3 SN: Villány 3 F: 2. Lm 1-3 Esztramos 3/3: Esztramos 3/3b: Esztramos 3/6: Villány 3 SN: Villány 3 F: mand. 5.: 5.70, max. 1.: 4.75, max. 2.: 5.0 max. 1.: 4.95 max. 1.: 5.25 max. 2.: 5.50 mand. 1.: 4.95, mand. 2.: 5.15, mand. 3.: 5.0, mand. 4.: 4.90 mand. 1.: 4.90 mand. 2.: 5.75 mand. 1.: 5.70, mand. 2 mand. 2.: 5.60, mand. 3.: 5.35, mand. 4 mand. 6.: 5.50, mand. 7.: 5.70, mand , mand. 3.: 5.40, mand , M 1 L max. 1. max. 2. max. 1. Wa Wp morph. Esztramos 3/ A D Esztramos 3/ C A A E Esztramos 3/ C A C c D D Esztramos 3/ B C

9 L Wa Wp morph. Esztramos 3/ A A A Villány 3 SN max A C C A C A A Villány 3 K A Villány 3 F max A max A C 4. M 2 Esztramos 3/3 max D max D Esztramos 3/ C D Esztramos 3/8 max C D Esztramos 3/ A Esztramos 3/ D D D C Villány 3 SN max D max D max D

10 L Wa Wp morph. Villány 3 F max D max C c D 5. M 3 Esztramos 3/3 max max Esztramos 3/8 max Villány 3 SN max max Villány 3 F max m 1 Esztramos 3/3 mand A mand A mand A mand A mand A A A Esztramos 3/6 mand A mand A mand B A A Esztramos 3/3b mand A Esztramos 3/8 mand A A A A A

11 L Wa Wp mor Esztramos 3/ A mand. 1. Villány 3 SN A mand C mand A mand A mand A CD CB Villány 3 K mand A mand A mand. 2. Villány 3 F D mand A mand C mand A mand A mand A mand A mand A mand A mand A mand A mand A mand A A 7. m 2 Esztramos 3/3 mand C mand C mand D mand C mand c mand D Esztramos 3/3b mand D Esztramos 3/6 mand A mand A mand A B

12 L Wa Wp mc Esztramos 3/8 mand C C c c A Esztramos 3/ C C C Esztramos 3/ D c Villány 3 SN mand c mand A mand D mand B mand A mand D mand C mand B mand C mand B C Villány 3 A mand B Villány 3 F mand B mand C mand A mand B mand A mand A mand C mand D mand B mand B mand B mand C mand A mand B mand B mand A

13 L Wa Wp Villány 3 F mand morph. A B 8. m 3 L Wa morph. Esztramos 3/3 mand A mand X mand A mand X mand A mand X Esztramos 3/3b mand X Esztramos 3/6 mand A A A Esztramos 3/8 mand A A G E A Esztramos 3/ A A Esztramos 3/ A E X Villány 3 SN mand X mand X mand X mand X mand A mand A mand A mand A

14 mand. 10. mand. 1. L Wa morph. Villány 3 SN X A A A A Villány 3 A X mand. 1. Villány 3 F X mand A mand A mand A mand X mand A mand X mand A mand A mand X A A E A

15 Fig. 1. Scatter diagram of Allocricetus ehiki Ml molars 1= Villány 3,2= Esztramos 3,3= Hajnóczy Cave, 4= Tarkő, 5= different localities in Poland (Pradel 1988) Fig. 2. Scatter diagram of Allocricetus ehiki M2 molars 1= Villány 3,2= Esztramos 3,3= Hajnóczy Cave, 4= Tarkő, 5= different localities in Poland (Pradel 1988)

16 Fig. 3. Scatter diagram of Allocncetus ehiki M3 molars 1= Villány 3,2= Esztramos 3, 3= Hajnóczy Cave, 4= Tarkő, 5= different localities in Poland (Pradel 1988) Fig. 4. Scatter diagram of Allocricetus ehiki ml molars 1= Villány 3,2= Esztramos 3,3= Hajnóczy Cave, 4= Tarkő, 5= different localities in Poland (Pradel 1988)

17 Fig. 5. Scatter diagram oïallocricetus ehiki m2 molars 1= Villány 3, 2= Esztramos 3, 3= Hajnóczy Cave, 4= Tarkő, 5= different localities in Poland (Pradel 1988) Fig. 6. Scatter diagram of Allocncetus ehiki m3 molars 1 = Villány 3, 2= Esztramos 3,3= Hajnóczy Cave, 4= Tarkő, 5= different localities in Poland (Pradel 1988)

18 Fig. 7. The relation of the L ml and L m3 in complete Allocricetus mandibles Allocricetus ehiki: 1= Villány 3, 2= Esztramos 3, 3= Hajnóczy Cave, 4= Tarkő Allocricetus bursae: 5= Tarkő layers a Sb Be nid rje mi Fig. 8. The distribution of the differences between the anterior and posterior widths of Ml molar in different cricetid materials a= Cricetulus migratorius, Recent, b=allocricetus bursae, Tarkő 1. layer, c= Allocricetus bursae, Tarkő 14., 16., 17. layers d = Allocricetus bursae, Poland, e=allocricetus ehiki, Poland, f= Allocricetus ehiki, Hungary (a, d, e are after Pradel 1988)

19 Fig. 9. Chewing surface of Allocricetus ehiki molars a = Ml in Esztramos 3/3 max.l., b= Ml in Esztramos 3/6, c= C-type M2 in Villány 3 F, d= M3 in Esztramos 3/8 Fig. 10. Chewing surface of Allocricetus ehiki ml molars a= Esztramos 3/7, b= Esztramos 3/6 mand 3, c= Villány 3 SN (CB-type), d= Villány 3 F (D-type), e= Villány 3 SN (D-type)

20 a b Fig. 11. Chewing surface oîallocricetiis ehiki m2 molars a= Esztramos 3/3 mand 1. (C-type), b= Esztramos 3/8 (A-type) 1 mm Fig. 12. Chewing surface oîallocricetus ehiki m3 molars a= Esztramos 3/3 mand. 4., b= Esztramos 3/3 mand.6., c= Esztramos 3/7, d= Esztramos 3/8, e= Esztramos 3/8, f= Esztramos 3/8

FRAGMENTA MINERALOGICA ET PALAEONTOLOGICA. 18. BUDAPEST, 1996 p

FRAGMENTA MINERALOGICA ET PALAEONTOLOGICA 18. BUDAPEST, 1996 p. 79-90 Cricetinus janossyi sp. n. (Rodentia, Mammalia) from the Pliocene fauna of Osztramos 7. (N Hungary) by J. Hír HÍR, J. (1996): Cricetinus