FRAGMENTA MINERALOGICA ET PALAEONTOLOGICA 17. BUDAPEST 1994 p. 71-89.

Cricetinus beremendensis sp. n. (Rodentia, Mammalia) from the Pliocene fauna of Beremend 15. (S )

by

J. HÍR

Hír. J.: Cricetinus beremendensis sp. n. (Rodentia, Mammalia) from the Pliocene fauna of Beremend 15. (S Hungary). - Fragm. Min. et Pal, 17: 71-89.

Abstract: The detailed description of the new species is given with the discussion of the systematic and stratigraphie relationships.

INTRODUCTION

Beremend is situated in the southernmost point of Hungary () 9 km from Villány. The limestone quarry of the village yielded the classical vertebrate faunae studied for more than a century (see in Jánossy, 1986). The fauna of the locality Beremend 15. was collected by Jánossy (1987, 1990, 1992). He gave the review of the material and the special study of the bird remains. From biochronological point of view the most important species are Estramomys simplex Jánossy, Mimomys cf. hajnac kens is Fejfar, Mimomys cf. hungaricus Kormos, Mimomys cf. pusillus Méhely, Mimomys cf. reidi Hinton, Mimomys cf. pitymyoides Jánossy & Meulen and Dolomys milieu Nehring. The aim of this paper is to give the detailed analysis of the small-sized cricetid finds of the locality Beremend 15.

METHODS

The measurements [LM 1-3 = length of upper toothrow; Lm 1-3 = length of lower toothrow; L = length of the toothcrown; Wa = anterior width of the toothcrown at the protocon(-id) - paracon(-id) or anterocon(-id) (on Ml, ml) height; Wp = posterior width of the toothcrown at the hypocon(-id) - metacon(-id) height] were taken by the ocularmic- rometer of a stereomicroscope following the method by Pradel (1981, 1988) and worked out by the basic statistic parameters (N = sample size, min.-max. = observation range, X - arithmetic mean, SD = standard deviation). The dimensions are given in mm. The morphological investigation is based on the nomenclature of Fahlbusch (1964) and Mein & Freudenthal (1971). The strongly worn teeth were not counted into the morphological examination. In the morphological distributions "+" means the presence, and "-" means the absence of the investigated element. SYSTEMATIC DESCRIPTION

Order: RODENTIA Bowdich, 1821 Family: CRICETIDAE Rochebrune, 1883 Subfamily: Cricetinae Murray, 1866

Genus: Cricetinus Zdansky, 1928

Cricetinus beremendensis sp. n. (Figs. 1-3, 5, 8, 10, 15-17)

Holotype: mandibula fr. dext. with ml-m3 (Figs. 1,3.). Paratype: maxilla fr. sin. with M1-M3 (Fig. 2.). Type-locality: Beremend 15, southern Hungary. Stratum typicum: terra rossa, karstic fissure filling.

Age: MN unit 16a, Lower Villafranchian, Villanyian in the system of Mein (1975) and De Bruijn et al. (1992). In the zonation Fig. 1. Toothrow ofFejfar et Heinrich (1986) it refers to the Mimomys hajnackensis of the holotype of zone of the Borsodia-Dolomys stage in the Villafranchian. After Cricetinus beremen- the system of Jánossy (1979, 1986) it refers to the Beremendian densis sp. n. phase of the Lower Pleistocene. Derivatio nominis: after the name of the locality. Diagnosis: small sized Cricetinus species, smaller than Cricetinus europaeus Kretzoi, 1959 írom Csarnóta 2. In the ml the anteroconid is undivided or weakly divided, the anterolophulid is mainly underdeveloped, mesolophid does not exist.

Measurements of the holotype and paratype:

Lml-3: 5.0L Ml-3: 46

Ml M2 M3 ml m2 m3 L: 2.02 1.43 1.22 1.96 1.64 1.48 Wa: 0.98 1.29 1.11 0.73 1.26 1.34 Wp: 1.29 1.25 1.19 1.26

Preliminary determinations: "Allocricetus éhiki" (Jánossy 1987, 1990)

Material: 1 maxilla with complete toothrow, 10 maxillae fr. with incomplete toothrows, 6 mandibulae with complete tooth- rows, 20 mandibulae fr. with incomplete toothrows. 35M1, 39M2, 13M3, 54ml, 38m2, 40m3 Deposition: Department of Geol. and Pal, Hungarian Natural History Museum. Fig. 2. Toothrow of the paratype of Cricetinus beremen­ densis sp. n. Ml (Figs. 5, 19)

On the mesial (anterior) surface of the ante- rocone a protrusion of the enamel (pre-antero- cone cingulum) or small conulet is frequent. The anterocone is always divided. The antero- lophule connects the two cones of the antero­ cone with the protocone in "Y" shape. The an- terolophule has no spurs. The parastyle and the entostyle are rare. In 20 molars the paracone Fig. 3. Lingual profile of the has anterior connection towards the anterior holotype Cricetinus beremendensis margin of the protocone (in this case there is a sp. n, fragmentary mandible closed basin between the protocone and para­ cone). In 24 molars the paracone has not anterior connection. Mesolophe does not exist. The metacone is always connected with the posterolophe without metalophule.

Dimensions: N: 44

min.-max. X SD L: 1.90-2.21 2.06 0.06475 Wa: 0.95-1.12 1.03 0.05152 Wp: 1.20-1.40 1.30 0.04904

a b c d

e f

Fig. 4. Mesial (anteriro) profiles of ml molars, (a, b: Allocricetus bursae, Tarkő, layer 16, V. 67. 97. Hung. Nat. Hist. Mus.; c, d: Cricetinus beremendensis, Beremend 15 (c=holotype); e, f: Cricetinus europaeus, Csarnóta 2 (e=holotype) Morphological distribution

A preanterocone cingulum paracone anterior connection 7 %

B preanterocone cingulum paracone anterior connection 19 43.%

C preanterocone cingulum paracone anterior connection parastyle 2 4.5% Dpreanterocone cingulum paracone anterior connection 17 38.7% Fig. 5. Occlusal surface of a E preanterocone cingulum Cricetinus paracone anterior connection beremendensis Ml parastyle molar entostyle 1 2.3% (D-morphotype) Fpreanterocone cingulum paracone anterior connection 2 4.5% total 44 100.0%

M2 (Figs. 8, 20)

The 4 main cusps and the anterior cingula are the constant elements of the teeth. Relatively rare accessory elements are the small remnant mesolophe, the mesostyle, the entostyle and the "parametalophule" which is an enamel crest between the metalophule and the posteroloph trimming the metacone on the lingual side (Fig. 26).

Dimensions: N: 43

min.-max. X SD Fig. 6. Cricetinus L 1.39-1.71 1.55 0.06356 europaeus Ml Wa 1.20-1.40 1.29 0.06367 molar (holotype, Wp 1.11-1.29 1.21 0.04567 Hung. Geol. Inst, Morphological distribution V. 12769)

A mesolophe -, parametalophule 28 67% B mesolophe +, parametalophule 4 9% C mesolophe +, parametalophule I 2%

D mesolophe -, parametalophule mesostyle +, entostyle 5%

E mesolophe -, parametalophule mesostyle -, entostyle F mesolophe -, parametalophule mesostyle +, entostyle 10%

G mesolophe +, parametalophule mesostyle +, entostyle 1 2% total 42 100%

M3 (Figs. 17, 21)

On the crowns of the molars there are not any accessory elements with statistic occurrence Fig. 7.' Cricetinus Dimensions: N: 24 europaeus Ml min - max. X SD molar (no. 10). L 1.15-1.4 1.27 0.06546 with small, Wa 1.04-1.19 1.11 0.04947 remnant mesolophe

ml (Figs. 4, 10, 13, 22)

The mesial (anterior) surface of the teeth is smooth. The anteroco- nid is mainly undivided (69 %) or weakly divided. In the latter case the division is visible only on the distal (posterior) surface of the anteroconids. The well developed anterolophulid is rare. In the most cases it is indistinct (does not emerge from the level of the anterosi- nusid) or missing. If the anterolophulid is developed it can be con­ nected with the labial or with the lingual part of the anteroconid and in one case it is doubled. The mesolophid and mesostylid are missing. The terminal part of the posterolophuhd is slightly broadening out.

Fig. 8. Dimensions: N: 67 Cricetinus min.-max. X SD beremendensis L 1.76-2.10 1.89 0.07225 Ml molar (no. Wa 0.59-0.84 0.70 0.06383 3/2) Wp 1.05-1.21 1.13 0.03671

Morphological distribution

A ACD undivided, ALPLD weak B ACD weakly divided, ALPLD weak C ACD undivided, ALPLD- D ACD weakly divided, ALPLD well developed 13 18% E ACD weakly divided, ALPLD - 2 2% Fi8- 9- F ACD undivided, ALPLD well developed 4 5% Cricetinus Total 72 100% europaeus M2 molar (Holotypus) Fig. 10. Cricetinus beremendensis ml molars (a: no. 19, b: no. 20, c: no. 31)

Fig. 11. Cricetinus europaeus ml molar (Holotypus) Fig. 12. Cricetinus europaeus ml molar from Csarnóta 2 (with remnant mesolophid) Fig. 13. Cricetinus beremendensis ml from Csarnóta 4

Fig. 14. Cricetinus europaeus m2 molar from Csarnóta 2 with extremely developed mesolophid and separated posterolophuhd Fig. 15. Cricetinus beremendensis ml molar (no. 26, morphotype C) Fig. 16. Cricetinus beremendensis ml molar (no. 47 morphotype F) Fig. 17. Cricetinus beremendensis M3 molar (no. 17) Fig. 18. Cricetinus europaeus M3 molar (no. 18), with mesolophe ni2 (Figs. 15, 16, 23)

By the side of the constant antero-buccal cingulum a weakly developed antero-labial cingulum is relatively frequent. In some cases a short remnant mesolophid exists. The broadened terminal part of the posterolophuhd is sometimes separated from the hypoconid by a narrow part of the posterolophuhd. In 4 cases the posterolophuhd forked in Y-shape. The extra branch is called "posterolophuhd".

Dimensions: N:64 min.-max. X SD L 1.40-1.65 1.53 0.053031 Wa 1.13-1.34 1.21 0.046987 Wp 1.08-1.36 1.21 0.056314

Morphological distribution

A ALC +, ML -, PLD non separated 11 18% B ALC -, ML -, PLD non separated 20 33% C ALC +, ML -, PLD separated 8 13% D ALC -, ML -, PLD separated 11 18% E ALC +, ML -, PLD non sep, PLLD + 1 1.6% F ALC +, ML +, PLD non separated 4 6.6% G ALC -, ML -, PLD separated, PLLD + 1 1.6% H ALC -, ML +, PLD non separated 2 3.4% 1 ALC +, ML +, PLD separated, PLLD + 1 1.6% .1 ALC -, ML +, PLD separated 1 1.6% K ALC -, ML +, PLD non separated, PLLD + 1 1.6% Total: 61 100%

m3 (Figs. 24, 27)

Antero-lingual cingulum exists in 20 teeth. The mesolophid is present in 100%, but the morphology of the mesolophid region is very variable. We could distinct 9 morphotypes (Fig. 28.).

Dimensions: N: 53

min - max. X SD L 1.33-1.57 1.46 0.05389 Wa 1.06-1.34 1.15 0.05398 Morphological distribution: Fig. 28

Length of the complete toothrows N: min - max. X: SD

LM1-3 2 4.60-4.75 4.67 Lml-3 7 4.55-5. 04.80 0.1728 Proportions in the toothrows

N: min.-max. X: SD LM1/LM2 2 1.28-1.41 1.34 L M3/LM2 3 0.77-0.85 0.81 L ml/Lm21 2 1.16-1.29 1.23 0.0424 51 L m3/Lm2 13 0.88-0.98 0.93 0.0335 31 DIFFERENTIAL DIAGNOSIS

1. Separation from Cricetinus europaeus KRETZOI, 1959

The available material of Cricetinus europaeus is scarce. Only the holotype (3 molars) were described (Kretzoi 1959, 1962 Kordos 1987). The present author checked the material of Csarnóta 2 stored in the collection of the Hungarian Geological Institute. During the course of this he found a few additional Cricetinus teeth. Based on the knowledge of this material we can slightly change the description of this species.

Csarnóta 2. Cricetinus europaeus material and measurements L Wa Wp 5M1 2.10 1.02 1.40 holotypus 2.10 1.11 1.43 2.18 1.12 1.42 2.20 1.05 1.40 2.18 1.05 1.45

4M2 2.0 1.50 1.20 C holotypus 1.68 1.30 1.40 B 1.60 1.19 1.30 A 1.68 1.26 1.40 ML -, PML +

4M3 1.40 1.30 — ML + 1.48 1.30 - ML + 1.44 1.33 - ML + 1.36 1.20 ML- 6ml 1.90 0.70 1.10 F holotypus 1.96 0.70 1.19 F 1.89 0.70 1.16 A 1.96 0.77 1.18 F, ML + 1.89 0.70 1.11 F 1.93 0.63 1.12 F, ML +

7m2 1.75 1.29 1.26 F 1.69 1.36 1.33 C 1.67 1.33 1.27 F, MSSTD + 1.67 1.23 1.30 A, ECSD + 1.68 1.26 1.27 J 1.79 1.30 1.27 J

1.68 1.26 1.29 ALC +, ML +; 5m3 1.68 1.26 B15/42 1.69 1.19 B15/42 1.75 1.33 B15/33 1.75 1.30 B15/44 1.71 1.18 B15/46

Cricetinus beremendensis differs from C. europaeus by its smaller size (except the ml molars which have equal size: Fig. 22). All the five Ml molars of C. europaeus have metalophule and 2 of them have remnant mesolophe too (Figs. 6, 7) In three M3 molars of C. europaeus have mesolophe which is absolutely missing in the M3 teeth of C. beremendensis (Figs. 17, 18). In the ml molars of C. europaeus the anterolophulid is generally better developed (Figs. 11, 12). In two mis small, remnant mesolophids exist. The labial part of the anterosinusid is definitely deeper, than the lingual part. In the ml molars of C. beremendensis the two parts have the same depth. Among the seven m2 molars of C. europaeus available, five specimens bear mesolophid. In one of them the mesolophid reaches the lingual margin of the tooth (Fig. 14).

2. Separation from Allocricetus ehiki Schaub, 1930 and Allocricetus bursae Schaub, 1930

In this respect the measurements have subordinate importance. Generally the averages of the measurements of Cricetinus beremendensis are between the averages of the corres­ ponding measurements of A. ehiki and A. bursae, but the overlappings are considerable (Hír 1993 b) In the Ml molars of Allocricetus species the frequency of the parastyle is higher. Likewise, in the M2 molars of Allocricetus the "parametalophule" is more frequent (Fig. 25). In the ml molars of the Allocricetus species the undivided anteroconid is an extremely rare curiosity (Hír 1993a). In the majority of cases it is well divided (Fig. 4) and the anterolophulid is well developed. In the m2 molars of the Allocricetus species the short mesolophid occurs more frequently (Fig. 27). In the m3 series of the Allocricetus material the frequency of the B15/34, B15/38 morphotypes is lower than in C. beremendensis and the morphotype B 14/33 occurs only in the material of Beremend 15 (Fig. 28.). The distribution of the other morphotypes seems to be random.

3. Separation from Cricetulus migratorius (Pallas, 1773)

The size measurements of the recent Cricetulus migratorius from Syria (Pradel 1981) and (Hír 1993) are remarkably smaller. In these assemblages the anteroconid of the ml molars is always divided and the anterolophulid is well developed. Fig. 19. Scatter diagram of Ml molars. 1: Cri­ Fig. 20. Scatter diagram of M2 molars. 1: Cri­ cetinus europaeus, Csarnóta 2, 2: Cricetinus cetinus europaeus, Csarnóta 2, 2: Cricetinus beremendensis, Beremend 15, 5: Cricetinae in- beremendensis, Beremend 15, 4: Allocricetus det, Osztramos 7 cf. bursae, Ivanovce b (Fejfar 1970), 5: Crice­ tinae indel, Osztramos 7

Fig. 21. Scatter diagram of M3 molars. 1 : Cri­ Fig. 22. Scatter diagram of ml molars. 1: Cri­ cetinus europaeus, Csarnóta 2, 2: Cricetinus cetinus europaeus, Csarnóta 2, 2: Cricetinus beremendensis, Beremend 15, 3: Cricetinus beremendensis, Beremend 15, 3: Cricetinus beremendensis, Csarnóta 4, 5: Cricetinae in­ beremendensis, Csarnóta 4, 4: Allocricetus cf. del., Osztramos 7 bursae, Ivanovce b (Fejfar 1970) u-

u 13 u Wmax. 12 13 mm

Fig. 23. Scatter diagram of m2 molars. 1 : Cri­ Fig. 24. Scatter diagram of m3 molars 1 : Cri­ cetinus europaeus, Csarnóta 2, 2: Cricetinus cetinus europaeus, Csarnóta 2, 2: Cricetinus beremendensis. Beremend 15, 3: Cricetinus beremendensis, Beremend 15, 3: Cricetinus beremendensis, Csarnóta 4, 4: Allocricetus cf. beremendensis, Csarnóta 4, 4: Allocricetus cf. bursae, Ivanovce b (Fejfar 1970), 5: Criceti­ bursae, Ivanovce b (Fejfar 1970), 5: Criceti­ nae indet, Osztramos 7 nae indet,Osztramos 7

THE MATERIALS OF CSARNÓTA 4 AND OSZTRAMOS 7

Csarnóta 4. Cricetinus beremendensis = Allocricetus (?) sp. indet in Kretzoi (1962)

Material: L Wa Wp morph. 1 M3 1.26 1.06 - ML- 1 ml 1.89 0.71 1.15 E 2 m2 1.51 1.20 1.22 E 1.47 1.19 1.19 C 1 m3 1.43 1.22 — TK 7/4 The material is scarce, but the only ml clearly shows the characteristic features of C. beremendensis (Fig. 13).

Osztramos 7. Cricetinae indet. = Cricetinus cf. europaeus in Jánossy (1973) = Cricetinus sp. in Jánossy (1978, 1979, 1986) Material: L Wa Wp morph.

4M1 2.04 1.05 1.29 B 1.93 0.95 1.19 D 1.85 0.94 1.23 B 1.85 0.98 1.18 B BEREMEND­ ENSIS

B15 V3-03 TARKÓ

16-16 12-11 15-13 10-2

• 7 32 25 37 58 15

Fig. 25. The frequency of the parastyle (PAST) in Ml molars in Plio-Pleistocene cricetid popula­ tions of Hungary. B 15 = Beremend 15, V3-03 = Villány 3 - Osztramos 3

c A. BEREMEND­ BURSAE ENSIS

B15 V3-03 TAR KO

16-16 12-11 15-13 10-2

H I L 23 13 25 V. 16 °/o

Fig. 26. The frequency of the "parametalophule" in M2 molars in Plio-Pleisfocene cricetid popu­ lations of Hungary. B 15 = Beremend 15, V3-03 = Villány 3 - Osztramos 3

A BEREMEND­ EHIKI ENSIS

B15 V3-03 TARKÖ

16-16 12-11 1 15-13 10-2

• n 33 26 16 2

Fig. 27. The frequency of the mesolophid in m2 molars in Plio-Pleistocene Hungarian cricetid populations of Hungary. B 15 = Beremend 15, V3-03 = Villány 3 - Osztramos 3 4M2 1.58 1.30 1.26 ML -,PML + 1.47 1.20 1.12 A 1.40 1.26 1.12 A 1.40 1.19 1.12 A

1M3 1.27 1.05 _ ML + lni2 1.40 1.11 1.05 C 4m3 1.37 1.02 - B15/34 1.50 1.22 - B15/44 1.41 1.08 - B15/40 1.41 1.06 - TK7/4

Without ml the well-found distinction between the genera Allocricetus and Cricetinus is impossible. The presence of M3 with mesolophe is remarkable because it never occurs in Allocricetus M3 molars.

CONCLUSIONS

In Hungary the locality Beremend 15. yielded the richest material of small-sized cricetids of MN unit 16. This material belongs to one species which differs from the Allocricetus genus and situated closer to the Cricetinus of Csarnóta 2.on the basis of the morphological characters, especially the morphology of the anteroconid on ml. The type-species of the genus Cricetinus (Cricetinus variáns Zdansky, 1928) was described from the Middle Pleistocene fauna of Chou-kou-tien, China (The material was revised by Shaohua 1984). The great geographic distance and the age difference make the identification of the finds from Csarnóta uncertain in spite of the morphological similarity. We can clear up this problem only with the direct comparison of the Chinese and Hungarian Cricetinus finds. This uncertainty is increased by the absence of Cricetinus in other European faunae. At this point the question of the first appearence of the genus Allocricetus emerges. In the Hungarian faunal sequence, representatives of this genus appeared in large amount first in Villány 3 (Jánossy 1979, 1986, Hír 1993). In the older faunae their presence is uncertain. One of the uncertain appearances is in the above discussed material of Osztramos 7. The other is in Ivanovce B in Slovakia (Fejfar 1970) where Allocricetus cf. bursae was found together with Kowalskia intermedia. This coexistence is unique. The clearing up of the European evolution of small-sized cricetids after the disap­ pearance of the genus Kowalskia needs further investigations. The finds from Beremend 15 proved the existence of another evolutionary lineage which is independent from Allocricetus.

ACKNOWLEDGEMENTS

The author would like to express his sincere thanks to Prof. Dénes JÁNOSSY for his permission to study the materials of Csarnóta 4, Beremend 15 and Osztramos 7. and to Prof. László KORDOS for the possibility to study the material of Csarnóta 2. Fig. 28. The frequency of m3 morphotypes (described in the material of Beremend 15) in Plio-Pleistocene Hungarian cricetid populations of Hungary CD O

Fig. 28. The frequency of m3 morphotypes (described in the material of Beremend 15) in Plio-Pleistocene Hungarian cricetid populations of Hungary Fig. 28. The frequency of m3 morphotypes (described in the material of Beremend 15) in Plio-Pleistocene Hungarian cricetid populations of Hungary Fig. 29. Chronological sketch of certain Plio-Pleistocene cricetid materials of Central REFERENCES

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Author's address: Dr. János HÍR Pásztó, P.O. Box 15. 3060 Hungary