A N N A L E S H I S T O R I C O - N A T U R A L E S M U S E I N A T I O N A L I S H U N G A R I C I Volume 89. Budapest, 1997 pp. 23-42.

Cricetus runtonensis solymarensis ssp. n. (Mammalia, Rodentia) from the Late Middle Pleistocene fauna of So lymár

J. H Í R

Municipal Museum H-3060 Pásztó, Pf. 15, Hungary

HÍR , J. (1997): Cricetus runtonensis solymarensis ssp. n. (Mammalia, Rodentia) from the Late Middle Pleistocene fauna of Solymár. - Annls hist.-nat. Mus. natn. hung. 89: 23^42.

Abstract - Detailed metrical and morphological analysis of the "large sized" hamster material from Solymár is given with the systematical and biostratigraphical relations. With 18 figures and 14 tables.

I N T R O D U C T I O N

The cave of the Ördög- lyuk is situated at 325 m height above sea level on the NE slope of the Sziklás-hill , W of village Solymár, near Budapest. The complicated cave system developed hydrothermally in Triassic (Dachstein) limestone ( V É R T E S 1950). The total length of the cave is 2000 m. Paleontological excavations and collections were made here between 1939 and 1951. The field activities were managed by V É R T E S (1950) the later well known paleolithic archaeologist. The detailed history of the research activ­ity and the list of the different localities is given by V Ö R Ö S (1988).

A rich microvertebrate fauna was found in a small hall of the cave named as "Kis ­körút" .

The later history of this material was rather unfortunate because it was partially de­stroyed during the second world war and during the 1956 revolution. The first study o f the fauna ( K R E T Z O I 1946) was never published, it was left in manuscript. Other special studies were published on the land snail fauna ( R O T A R I D E S 1944) and on the charcoal material (STIEBER 1952).

After a long break, the enamel thickness o f the Arvicola molars was studied by JÁNOSSY (1976a) and H E I N R I C H (1982). The investigation o f the large mammalian finds was made by V Ö R Ö S (1985, 1988). The revision o f the mollusc fauna was published by K R O L O P P ( 1 9 9 2 ) .

New studies on the available Solymár fauna are important, because biochronologi-cal units as "third faunal wave" K R E T Z O I (1953) "Solymár faunal stage" JÁNOSSY (1986) "So lymár ium" K R E T Z O I & P É C S I (1979) are based on the material.

The aim o f this paper is to give the accurate analysis of the fossil hamster material from Solymár, Ördög-lyuk.

M A T E R I A L A N D METHODS

The following hamster material is housed in the Paleontological Collection of the Hungarian Natural History Museum from Solymár.

9 viscerocrania, 15 maxillae with complete toothrows, 18 maxillae with incomplete toothrows, 48 mandibulae with complete toothrows, 40 mandibulae with incomplete toothrows.

The dental measurements were taken using the ocularmicrometer of a stereomicroscope to an accuracy of 0.01 mm. The following dimensions were measured (Fig. 1):

L M1-M3 - length of the upper row of molars; L ml-m3 - length of the lower row of mo­lars; L - length of tooth crown; Wa - anterior width of the toothcrown. In M l molars it was measured across the anterocone, in ml molars it was measured across the anteroconid. In M2, M3 molars it is the width across the protocone-paracone. In m2, m3 molars it is the width across the protoconid-paraconid. Wp - posterior width of the toothcrown. In M l , M2 molars it was measured across the hypocone-metacone, or across the hypoconid-metaconid in m l . m2 molars. This measurement was not taken in M3, m3 molars.

During the statistic elaboration of the data the next parameters were computed: n - sample size; min. - the minimal measurement; max. - the maximal measurement; X. -

arithmetic mean; Med. - median; SD - standard deviation; CV - coefficient of variation; K - 95% confident interval of the arithmetic mean(+ -); V - 100 R/M. In which R is the difference between max.and min., and M is the midpoint between max. and min. (FREUDENTHAL & CUENCA BESCOS 1984).

The results are given in the Tables 1-8. The statistic morphological investigation is based on the nomenclature of M E I N & FREUDEN-

THAL (1971«, b). The creation of the different morphotypes is demonstrated on the Figs 8, 9, 10, 12-17. The numerical data of the morphotypes are given in Tables 9-14. For comparison I quote the metrical data of the Pleistocene Cricetus major WOLDRICH, 1880 and Recent Cricetus cricetus (LINNAEUS, 1758) from Germany after FAHLBUSCH (1976). The data of the Pleistocene Cricetus runtonensis NEWTON, 1909 and Recent C. cricetus from Poland are after PRADEL (1985, 1988). I quote as well the results of my previous investigations (HÍR 1996, 1997,) on the Recent C. cricetus from Hungary and Early Pleistocene Cricetus praeglacialis SCHAUB, 1930 from Villány 8 (V8) and on C. runtonensis from Somssich-hegy 2 (SS2).

S Y S T E M A T I C DESCRIPTION

Order: Rodcntia BOWDICH, 1821 Family: Cricetidae ROCHEBRUNE, 1883 Subfamily: Cricetinae MURRAY, 1866 Genus: Cricetus LESKE, 1779 Species: Cricetus runtonensis NEWTON, 1909

Cricetus runtonensis solymarensis ssp. n.

Holotype. Viscerocranium No. 2. Paratype. Mandibula No. 1. Type locality. Kiskörút Hall in the Ördög-lyuk Cave at Solymár, Buda Mountains, NW from

Budapest. Type level. Red coloured argillaceous cave sediment. Age. Lower part of the Toringium after the system of FFJFAR & HEINRICH ( 1 9 8 1 , 1986, 1990) .

In the traditional Hungarian vertebrate paleontological stratigraphy (JÁNOSSY 1986) the Solymár phase is the oldest part of the Late Middle Pleistocene. Up to the present the exact correlation of this period with the standard Pleistocene chronostratigraphy (ZAGWIJN 1985) is impossible, but we presume that the Solymár phase refers to the Holsteinian or an other mild climatic period within the Saalien.

Etymology. After the name of the settlement, which is close to the locality. Synonyms. Cricetus cricetus inter glaciális KRETZOI, 1946; Cricetus cricetus ssp. JÁNOSSY,

1986.

Diagnosis. Large sized hamster subspecies, different from the Lower Pleistocene Cricetus runtonensis runtonensis population of Somssich-hegy 2 in the fol lowing statistic morphological features, — the higher frequency of the conelet entostyle in the upper M 1 , M 2 molars, and of the

conelet ectostylid in the lower m2 , m3 molars; — the frequent duplication of the mesolophid in the m2, m3 molars.

Among the studied dimensions the L M 2 , L M 3 , WaM3, L m l - m 3 , W a m l , L m 2 are significantly larger, the L M 1 , W p M l , W p M 2 , L M 1 / L M 2 are significantly smaller in C. runtonensis solymarensis than in C. r. runtonensis.

Description. The measurements of the molars are definitely larger, than the corre­sponding dimensions of the Recent C. cricetus from Hungary ( H Í R 1997), Poland (PRA-D E L 1985) and Germany ( F A H L B U S C H 1976), but similar to the metrical data of the C. runtonensis material from Poland ( P R A D E L 1988) and of the C. runtonensis population of Somssich-hegy 2 in Hungary (Figs 2-7, Tables 1-8). During the detailed comparison o f 22 metrical characters of the C. runtonensis materials from Solymár and from Somssich-hegy 2 localities 6 characters are significantly larger in the Solymár material ( L M 2 , L M 3 , W a M 3 , L m l - m 3 , W a m l , Lm2), 4 characters are significantly larger in the SS2 material

. ( L M 1, W p M 1, W p M 2 , L M 1 / L M 2 ) . No significant differnce in the other 12 characters. I n the morphological description we w i l l not reinterprete the general odontomorpho-

logical picture of the hamsters. We want to give the special features o f our material. M l . There are no remarkable differences in the frequency o f the morphotypes

among the investigated C. cricetus, C. praeglacialis materials. A O B Y O B O is the domi­nant morphotype everywhere, but in the C. runtonensis populations of Solymár and Somssich-hegy 2, the type A O B Y O B 1 is subdominant because the ramification o f the posterolophule is more frequent(Fig. 8, Table 9).

M2. In the Recent hamster from Hungary and in C. praeglacialis the dominant mor­photype is the A O O , but in the C. runtonensis materials (SS 2 and Solymár) A O l is dominant and A O O is only subdominant. In the latter samples the ramification o f the ter­minal part of the posterolophule is much more frequent (Figs 9-10, Table 10).

Fig. 1. Sketch of the investigated measurements on Cricetus molars

Figs 2 -5 . Scatter diagrams on the mean values of molars of some European fossil and recent Crice­tus populations: 2 = M l , 3 = M2, 4 = M3, 5 = m l . Explanation: A: C. major, B : C. runtonensis group, C: C praeglacialis, D: C. cricetus group, P B : Petersbuch ( F A H L B U S C H 1976), R: Poland, different localities ( P R A D E L 1988), L : Solymár, S: Somssich-hegy (HÍR 1996), P: Recent hamster from Poland ( P R A D E L 1988), H: Recent hamster from Hungary (HÍR 1997), E: Recent hamster from Germany ( F A H L B U S C H 1976), V8: Villány 8 ( H Í R 1997)

Table 1. Metrical data of upper toothrows Table 2. Metrical data of lower toothrows

Length of LM1/LM2 LM3/LM2 Length of LM1/LM2 LM3/LM2 complete relation relation complete relation relation toothrows toothrows

No. 28 46 30 No. 48 66 58

Min. 8.15 1.053846 0.769912 Min 8.5 1.076923 0.995357

Max. 9.0 1.277778 0.909091 Max 9.6 1.245136 1.175438

X 8.587679 1.175092 0.863654 X. 9.021042 1.151896 1.074635

Med. 8.6 1.178996 0.877872 Med. 9.0125 1.155878 1.065575

SD 0.211995 0.042155 0.037243 SD 0.237454 0.029293 0.041412

CV 2.468596 3.587414 4.312226 CV 2.632222 2.543023 3.85357

V 9.912536 19.20824 16.57878 V 11.11111 14.48826 16.59124

K 0.017731 0.000535 0.000528 K 0.016613 0.000213 0.000454

Figs 6 - 7 . Scatter diagrams on the mean values of the molars of some European fossil and recent Cricetus populations: 6= m2, 1 - m3. Explanation as in Figs 2-5

Table 3 . Metrical data of M l molars

L Wa Wp Wa-Wp

No. 51 50 50 49

Min 3.225 1.77 2.0 -0.4

Max 3.7 2.0 2.35 -0.12

X 3.432216 1.8917 2.1585 -0.26459

Med 3.425 1.875 2.15 -0.27

SD 0.108345 0.064634 0.079531

CV 3.156707 3.416738 4.204231

V" 13.71841 12.20159 16.09195

K 0.003353 0.001206 0.001825

Table 4 . Metrical data of M2 molars

L Wa Wp Wa-Wp

No. 49 49 49 49

Min. 2.70 2.175 2.0 0.1

Max. 3.25 2.50 2.27 0.35

X 2.928714 2.342898 2.123388 0.21961

Med. 2.90 2.35 2.12 0.225

SD 0.126064 0.069879 0.066491

CV 4.304424 2.9826 3.13135

V 18.48739 13.90374 12.64634

K 0.004634 0.001424 0.001289

Table 5. Metrical data of M3 molars Table 6. Metrical data of ml molars

L Wa L Wa Wp Wa-Wp

No. 33 33 No. 73 72 72 72

Min. 2.175 2.0 Min. 3.132 1.15 1.75 -0.85

Max. 2.887 2.337 Max. 3.525 1.5 2.15 -0.40

X 2.527909 2.14497 X 3.303808 1.326097 1.912167 -0.58607

Med. 2.525 2.15 Med. 3.3 1.325 1.92 -0.575

SD. 0.123338 0.08019 SD 0.097297 0.072479 0.075892

CV 4.879043 3.738502 CV 2.944991 5.465555 3.968916

V" 28.13117 15.5407 V 11.80712 26.41509 20.51282

K 0.005486 0.002319 K 0.002231 0.001247 0.001367

Table 7. Metrical data of ml molars Table 8. Metrical data of m3 molars

L Wa Wp Wa-Wp L Wa

N 78 78 78 78 N 59 59

Min. 2.57 2.0 1.97 -0.18 Min. 2.787 2.05

Max. 3.1 2.45 2.45 0.2 Max. 345 2.525

X 2.867513 2.215859 2.222436 -0.00658 X 3.069356 2.263559

Med. 2.87 2.22 2.225 0 Med. 3.05 2.25

SD 0.094252 0.087473 0.105841 SD 0.129278 0.096535

CV 3.286893 3.947597 4.762377 CV 4.211902 4.264748

V 18.69489 20.22472 21.71946 V 21.26022 20.76503

K 0.004389 0.002447 K 0.002025 0.001744 0.002553

Table 9. Morphological data of M l molars

morphotypes individuals

AOBYOBO 25

AOBYOB1 14

AOBY1B1 1

S 40

Table 10. Morphological data of M2 molars

morphotypes individuals

AOO 6

BOO 1

AOl 26

BOl 3

A l l 1

S 37

Table 1 1 . Morphological data of M3 molars

morphotypes individuals

A 3

B 1

C 2

E 2

G 4

11 8

S 20

Table 1 2 . Morphological data of ml molars

morphotypes individuals

Bla 2

Cla 16

Dla 19

Ela 17

Fla 15

Fib 1

S 70

Fig. 8. Morphotype system of the M l molars. Explanation:; A: anterolophule is simple; B: antero-lophule is doubled; 0: labial eperon of the anterolophule is missing; 1: a low developed labial eperon is found; Y: protostyle is missing; X: protostyle is found; b: central ring is missing; a: cen­tral ring is found; 0: mesolophe is missing; 1: mesolophe is found; b: entomesolophe is missing; a: entomesolophe is found; 0: posterolophule is simple; 1: posterolophule is ramified

Table 13. Morphological Table 14. Morphological data of m2 molars data of m3 molars

morphotypes individuals morphotypes individuals

RA 8 00001 1

RD ! 01000 1

RE 8 03000 1

RG 3 04000 1

SA 2 05000 2

SB 8 03010 1

SC 3 04010 2

SD 1 05010 6

SE 11 04011 3

SF 3 04110 1

SG 1 04121 1

SH 1 04210 3

SI 3 04211 2

VE 6 05111 3

VF 1 05210 2

V L 10 05211 2

S 70 05221 1

12110 1

14210 2

14211 1

15010 1

15100 1

15110 1

15120 1

15111 1

15221 1

S 43

M3. In the investigated materials we cannot found characteristic morphological dif­ferences. There are no dominant morphotypes, the dispersion o f them seems to be ran­dom (Fig. 11, Table 11).

m l . The dispersion o f the morphotypes is rather equal in the molars from Solymár (Fig. 12, Table 12). This character is different in the other investigated hamster materials, where the strong dominance o f one type was found ( E l A in V 8 and SS2, F I A in the Re­cent sample (HÍR 1997)

ml. The morphology of this molar is very variable (Table 14). 17 morphotypes were distinguished. 8 o f them are common with the materials (V8, recent C. cricetus) pre­viously studied by the author (Fig. 13). It refers to the 53% of the molars. One morpho­type of this group ( V L ) is subdominant in the Solymár material and dominant in V 8 . A very special morphological feature o f the molar is the duplication of the mesolophid (Fig. 14) in the SC, SD, SE , SH types (37% of the teeth). By the side o f the normal Msld an­other enamel ridge is found starting from the hypolophulid, or from the jo int point o f the ectolophid and the hypolophulid. This element is not named in the nomenclature of M E I N et F R E U D E N T H A L (1971A, b) and I would like to suggest the name "paramesolophid (Pmsld)". In the morphotypes SA and SB only the "paramesolophid" is found and the mesolophid is missing (Fig. 14). This kind of duplication of the mesolophid was not found in the studied C. cricetus and C. praeglacialis materials, only in the C. runtonensis assemblage of Somssich-hegy 2, but only in 2% of the teeth.

Fig. 9. Morphotype system of the M2 molars. Explanation:; A: anterolophule is simple; B: antero­lophule is doubled; 0: mesolophe is missing; 1: mesolophe is found; 0: posterolophule is simple; 1: posterolophule is ramified

C. runtonensis, Somssich-hegy 2

C. runtonensis solymárensis, Solymár

C. praeglacialis, Villány 8.

C. cricetus, Recent

Fig . 10. Distribution of the M2 morphotypes in some Hungarian hamster materials (CR: Central Ring)

B C D

E F G H Fig. 1 1 . Morphotypes of the M3 molars. Explanation: A: central ring with mesolophe; B: central ring without mesolophe; C: opened central ring without mesolophe; D: opened central ring with mesolophe H : no central ring, no mesolophe E: reduced central ring with mesolophe; F: reduced central ring without mesolophe; G: mesolophe without central ring

R A R D R E R G

V E V T V L

Fig. 1 3 . Morphotypes of the ml molars. The morphotypes described in the recent Hungarian ma­terial and in the Villány 8 population ( H Í R 1996). Explanation: RA: mesolophid is short; RD: MSLD is long, but interrupted; RE: MSLD is long and horizontal; RG: MSLD is long and touches the metaconid; VE: MSLD is missing, the central ring is opened on the buccal side; VF: MSLD is missing, the CR is opened on the lingual side; VL: no MSLD, no CR

F E D

C B A

Fi«. 1 2 . Morphotype system of the ml molars. Explanation: A: anterolophulid is doubled and com­plicated by accessory ridges; B: doubled anterolophulid without accessory elements; C: anterolo­phulid is Y-shaped and connected to both two conelets of the anteroconid; D: anterolophulid is connected only to the buccal conelet of the anteroconid and has a lingual eperon; E: same as D but without lingual eperon; F: anterolophulid is reduced, it does not emerge from the level of the anterosinusid; 1 : mesolophid is missing; 2: a reduced mesolophid is found; a: posterolophulid is simple; b: posterolophulis is ramified

The applied morphotypes are based on the structure of the central region o f the mo­lars. Other important marker is the high frequency (45%) of the well developed conelet ectostylid, which is very rare in other studied hamster populations (2% in V8 and 4% in recent C. cricetus, 0% in SS 2). In the anterior region of the m2 molar the presence o f the antero-lingual cingulum is 97.5%.

SG SH SI Fig. 14. Morphotypes of" the m2 molars. The morphotypes described in the Solymár material. Ex­planation: SA: central ring is opened on the lingual side, paramesolophid is medium developed; SB: PMSLD is long; SC: mesolophid is long, PMSLD is short; SD: CR is opened on the lingual side, MSLD is long, PMSLD is short; SE: MSLD is long and touches the metaconid, PMSLD is short; SF: CR is pushed to lingual direction, MSLD and PMSLD are short; SG: MSLD is missing, PMSLD is long and ramified; SH: MSLD is short, PMSLD is long; SI: CR is opened on the lingual side, MSLD and PMSLD are not developed, only short irregular enamel ridges are found

C E N T R A L R I N G

M E S O L O P H I D

P A R A M E S O L O P H I D

E C T O M E S O L O P H I D

E C T O S T Y L I D

Fig. 15. Morphotype system of the Cricetus m3 molars from Solymár. Explanation: central ring 0: undeveloped, 1: developed; mesolophid 0: undeveloped, 1: incipient, 2: long diagonal, 3: long hori­zontal, 4: short, but touches the metaconid, 5: long, ramified; paramesolophid 0: undeveloped, 1: short, 2: long; ectomesolophid 0: undeveloped, 1: short, 2: long; ectostylid 0: undeveloped, 1: de­veloped

m3. The morphology of this molar is extremely variable. 25 morphotypes were dis­tinguished, but these are easy to describe on the basis of the variations o f five morpho­logical elements (Fig. 15). 3 morphotypes are common with the morphotypes of the C. praeglacialis-C. cricetus group, which refers to the 9% of the molars of Solymár.

The "paramesolophid" is a frequent element of the m3 molars as wel l . I t occurs in 60% of the teeth. The coexistence of the mesolophid - "paramesolophid" is found in 56% of the molars (Figs 16-17). The presence of the ectostylid is 35% .

DISCUSSION

The early scholars of the Solymár fauna described the hamster finds as a subspecies of Cricetus cricetus ( K R E T Z O I 1946, JÁNOSSY 1986) but this determination cannot be confirmed by the results of the present author. The metrical characters o f the material clearly demonstrate the relationship with Cricetus runtonensis (Figs 2-7) . This species was the first large sized cricetid in the Early Pleistocene faunas of Hungary and Poland ( P R A D E L 1988). In Hungary the most abundant material o f this species was found in the fauna of Somssich-hegy 2 (Nagyharsányhegy Phase) accompanied with Cricetus nanus. In the fauna of Villány 8 (Templomhegy Phase) the C. runtonensis disappeared, and it was substituted by the smaller sized Cricetus praeglacialis. The return o f the C. run­tonensis line in Hungary was found in the Late Middle Pleistocene faunas of Solymár , Pongor Hole (HÍR 1989) and Lök-völgyi Cave 8th layer (HÍR 1995) (Fig. 18).

Fig. 16-17. Cricetus runtonensis solymarensis ssp.n.: 16 = m3 molar, morphotype 04210, 17 = m3 molar, morphotype 05221

Fig. 18. Range chart of the evolutionary lines of the "large sized" hamsters in the Hungarian Pleis­tocene

The subspecies of Solymár is different from the Lower Pleistocene C. runtonensis in the more complicated morphology of the M 2 , m2, m3 molars. During the study of the Early Pleistocene C. praeglacialis and Recent C. cricetus HÍR (1997) has found a con­trary trend: the simplification in the morphology. This fact confirms the supposition of the author, that the C. praeglacialis -C. cricetus group and the C. runtonensis group are two independent evolutionary lines of the Middle European Pleistocene hamsters.

After the Late Middle Pleistocene C. runtonensis completely disappeared from the Carpathian Basin. In this moment we can not give an unambiguous explanation of the disappearence and returning of the species. The paleoecological preferences o f C. run­tonensis are not clear. The composition of the fauna Somssich-hegy 2 suggests a mainly continental steppe environment ( JÁNOSSY 1990). The material of Solymár reflects a forest ecosystem and mild climate, but the above mentioned other Late Middle Pleistocene fau­nas (Pongor Hole and Lök-völgyi Cave) are dominated by steppe elements as well .

In the chronological evaluation of the Solymár fauna VÖRÖS (1988) published his opinion, that on the basis of the specific composition of the macromammals the fauna can be put into the Upper Biharian (Early Middle Pleistocene) substage. This is a remarkable contradiction to the relative age determination refers to the micromammals, especially on the Arvicola wi th undifferentiated enamel structure, which establishes the Late Middle Pleistocene age ( H E I N R I C H 1982).

After the detailed study of the hamsters we can confirm the chronological determi­nation based on the micromammals, because in the Hungarian Late Biharian faunas the species Cricetus runtonensis is missing. In the faunas o f Vértesszőlős and Tarkő 1 the C. cricetus is found. In the lower layers (2-16) o f Tarkő an extra large sized hamster was collected. It was determinated as "C. c. runtonensis" by JÁNOSSY (1976b, 1986), but this determination is erroneous, because the dimensions are much larger than the studied C. runtonensis populations and refer to the C. major species after F A H L B U S C H (1976).

Acknowledgements - The author would like to express his sincere thanks to the staff of the Department of Paleontology of the Hungarian Natural History Museum for the kindness in offering the material investigated. The research activity was supported by the Hungarian Scientific Research Fund (OTKA No. 144117).

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