MINERALOGICA ET PALAEONTOLOGICA. Hamster remains (Rodentia, Mammalia) from the Lower Pleistocene of the Hajnóczy Cave, Hungary

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1 FRAGMENTA MINERALOGICA ET PALAEONTOLOGICA 15. BUDAPEST, 1992 p Hamster remains (Rodentia, Mammalia) from the Lower Pleistocene of the Hajnóczy Cave, Hungary by J. HÍR Abstract: A vertebrate fauna was collected from the Hajnóczy Cave (Bükk Mountains) between 1976 and The description of the locality along with a systematic study of the cricetid material is given here. Four hamster taxa (Cricetus cricetus praeglacialis, Cricetus cricetus nanus, Allocricetus ehiki and Allocricetus bursae) were identified. The assemblage is assigned to the Betfia Phase of the Early Pleistocene. INTRODUCTION The Hajnóczy Cave is located in the southern part of the Bükk Mountains (northern Hungary), on the Odorvár Hill, north of the town of Mezőkövesd. The cave was formed in the Upper Triassic grey limestone of the Felsőtárkány Formation. It was discovered by the speleological club of the Hajnóczy Secondary School of Tiszaföldvár (Varga, Cs. 1978, Varga, L. 1978). The genesis of the cave is complex. The system was originated at the limestone/slate (Kisgyőr Formation) lithological boundary (Text-fig. 1). The layout of the cave is maze-like (Hír 1982) (Text-fig. 2). Text-fig. 1 - Simplified geological section of the Hór Valley. 1. Kisgyőr Slate Formation. 2. Felsőtárkány Limestone Formation In the eastern part of the Galéria-terem (Gallery Chamber) a red clay layer containing bones was found. It covers a dripstone slope, filling its tetarata-like depressions by a thickness of cm (Text-fig. 3). The first paleontological collection was carried out by L. Kordos (1974). The present author made further collections between 1978 and Two reports were published on the speleology and sedimentology of the locality, supplemented by a preliminary faunal list (Hír

2 1982, 1985). In this paper a detailed study of the cricetids of the Lower Pleistocene of the Hajnóczy Cave is given. The peculiarity of this fauna is its mixed composition: the co-occurrence of older and younger elements (e. g. Mimomys pusillus, M. savini and a probably Late Pleistocene Microtus species). LOCALITIES The following are the main fossil localities in the cave (Text-fig. 3): Hajnóczy 1 It is a small hole situated close to the topographical reference point No. 17 between the Galéria- and Óriás-terem (Giant Chamber) Galéria 2 It is a locality found at the time of the first collection yielding bones of a specimen of Ursus deningeri Galéria 2 This locality is situated 3 m above Galéria 2 Galéria J It is a narrow pass below the locality Galéria 1, leading to the topographical reference point No. 7 Text-fig. 2 - Lower Pleistocene vertebrate localities of the Hajnóczy Cave. B: Entrance; Hi: Hajnóczy 1; 1: Galéria 1; 2: Galéria 2; 3: Galéria 3; 7: Galéria 7; 12: Óriás-terem 12; KG: Kis-galéria

3 All of the above localities are situated below a filled chimney system connected with the surface. Apart from these, there are 3 minor localities which yielded some additional material: Galéria At topographical reference point No. 7 Oriás-terem At topographical reference point No. 12 Kis Galéria (Small Gallery) (Text-figs 2-3) During the collections practically all of the fossiliferous clay layers have been excavated. METHODS OF STUDY MORPHOLOGICAL EXAMINATION The molars of Allocricetus were particularly suitable for this type of study. The basic pattern of tooth crowns of Pleistocene cricetids is the same on all species but there are several variable morphological elements as well. Some of them are treated in the terminology of Fahlbusch (1964) and Mein and Freudenthal (1971). Additional characteristics are described by Malez and Rabeder (1984) and in the present paper. In this study the presence or absence of the following elements were recorded (Text-fig. 4): Text-fig. 3 - Lower Pleistocene vertebrate localities of the section of Galéria and Óriás-terem (not to scale). Hi: Hajnóczy 1; Gi: Galéria 1; G2: Galéria 2; G3: Galéria 3.

4 Text-fig. 4 - Standard measurements and morphological elements (boxed) of cricetid molars, a: M 1 ; b: M ; c: M ; d: Mi; e: M2; f: M3 (Further symbols are explained in the text.)

5 M 1, Mi - pac ("pre-anterocon (id) cingulum"): It is a small enamel-bulge on the oral surface of the tooth. - past (parastylid). Morphotypes of M : A: pac4-, past 4-; B: pac 4-4-, past-; C: past- past4-; D: pac- past- Morphotypes of Mi: A: pac-h; B: pac- M 2 - mtl ("metalophuloid"): It is a crest between the metalophule and the posteroloph trimming the metacone on the lingual side. - ml (mesolophid) Morphotypes of M 2 : A:ml4-, mth-; B: ml4-, mtl-; C: ml- mtl-l-; D: ml- mtl- M 3 In the studied material there are two morphotypes of this molar. A: this type has no accessory elements. B: there is a transversal crest in the fossa between the hypocone and the metacone. M2, M3 - ale (antero-lingual cingulum): It is the continuation of the persistent antero-labial cingulum before the metaconid. - ml (mesolophid) Morphotypes of M2 and M3: A: alc+, ml4-; B: ale- ml 4-; c: ale4-, ml-; D: ale-, ml- BIOMETRIC METHODS The length and width of molars were measured according to Fahlbusch (1964, 1969). The measurements were taken using the micrometer ocular of an MBS-9 stereomicroscope. The data are presented on scatter diagrams in mm (Text-figs 7-12) and listed in the appendix The following abbreviations are used: Ltr - length of toothrow; 1 - length of molar; w a - anterior width; w p - posterior width; w - maximum width. SYSTEMATIC DESCRIPTIONS Cricetus cricetus praeglacialis SCHAUB, 1930 The few molars assigned to this subspecies constitute the largest hamster remains from the Hajnóczy Cave. On the crown of the upper molars there are no accessory elements. The M2 and M3 molars possess an antero-lingual cingulum. The mesolophid region on the M3 is complex (Text-fig. 5).

6 Text-fig. 5 - Drawings of M3 molars of Cricetus cricetus praeglacialis

7 Cricetus cricetus nanus SCHAUB, 1930 The measurements of this subspecies are generally intermediate between those of Cricetus cricetus praeglacialis and Allocricetus ehiki. On the basis of the few specimens available, the M 3 possesses a basic structure. The M2 and M3 show remnants of mesolophids. On the M3 the hypolophulid is fused with the entoconid forming a ring-like sinus together with the mesolophid (Text-fig. 6). The broadening posterolophid forms a small cone on the Mi and M2. According to Jánossy (1979, 1986), the subspecies occurs in the following localities in Hungary: Nagyharsány-hegy 2, Somssich-hegy 1, Osztramos 2, Osztramos 8, Osztramos 14. All of them represent "Allophaiomys faunas". Allocricetus ehiki SCHAUB, 1930 and Allocricetus bursae SCHAUB, 1930 Scatter diagrams of measurements of Allocricetus molars show bimodal distribution indicating the presence of two distinct species (Text-figs 7-12). Data of the larger one correlate well with homogenous Allocricetus ehiki populations from Villány 3 and Osztramos 3 localities. Also there is partial agreement with Allocric Text-fig. 6 - Drawings of molars of Cricetus cricetus nanus. a: M2; b: Mi; c: M3.

8 Text-fig. 7 - Scatter diagram of the M 1 molars of Allocricetus from the Hajnóczy Cave. Quadrate with solid line: Cluster of A ehiki from Villány 3 and Osztramos 3. Quadrate with broken line: Cluster ol4. ehiki from Poland (Pradel A. 1988)

10 -I » ,4 W m m

cetus ehiki populations from different localities in Poland (Pradel 1988). The smaller species, Allocricetus bursae, is common in the Hungarian Early and Middle Pleistocene faunas (Jánossy 1979,1986).

14 cetus ehiki populations from different localities in Poland (Pradel 1988). The smaller species, Allocricetus bursae, is common in the Hungarian Early and Middle Pleistocene faunas (Jánossy 1979,1986). The separation of the two species is based on measurements of the M 2, M 3, M2 and M3 molars. It is less obvious from the M 1 and Mi molars. As shown on Text-fig. 10, a specimen of A. ehiki falls into the cluster of A. bursae teeth. It is an Mi of a complete lower toothrow, whereas M2 and M3 have characteristic A. ehiki measurements. The Mi molars of the two species are less significantly different, only an arbitrary boundary can be drawn. Jánossy (1979,1986) divided the two Allocricetus species on stratigraphie basis. The first appearance of A. bursae was recorded from Osztramos 8 and all other Allocricetus were identified as A, ehiki. Hence the co-occurence of these two species in the Hajnóczy Cave was regarded as a result of the mixed composition of the fauna (D. Jánossy and L. Kordos, pers. comm.). However, A. ehiki and A. bursae were also found together in the Tarkő Rocksheiter (Hír 1989) and there are similar records from Poland. Fahlbusch (1969) reported the two species found together in the faunas of Kadzielnia and Kamyk. Pradel (1988) described 6 localities (Zamkowa Dolna A, Kadzielnia, Kamyk, Kielniki 3a, Zalesiaki la, Kozi Grzbiet) yielding both,4. bursae and A ehiki In Poland the two species range from the Villanyian to the Late Biharian. Generally A bursae is more abundant than A. ehiki In Hungary the last appearance of A. ehiki is in Layer 8 in the section of the Tarkő Rocksheiter (Hír 1989). The faunas of the Tarkő to Varbó Phases contain A. bursae only. The relatively small number of molars recovered from the Hajnóczy Cave allows only preliminary conclusions regarding the tooth morphology of the two species. The following summary of the morphotypes of M, M, M2 and M3 molars can be given: 1. Simple morphology Morphotypes with no accessory elements. M 2 :d; M 3 :a; M2:d; M3:b, d. The mesolophid is frequent on the M3 molars. 2. Complex morphology Morphotypes with accessory elements. M 2 :a, b, c; M 3 :b, c; M2:a, b, c; M3 -.a, c. A. bursae A.ehiki simple complex simple complex M M M Total The simple morphology is more frequent among the A. bursae molars. A similar distribution of morphotypes was observed on the rich material from the Tarkő Rocksheiter (Hír 1989). Further studies of the differences of these two Allocricetus species are planned.

15 CONCLUSIONS The distribution of the cricetid specimens found in different localities of the Hajnóczy Cave is summarized in the following table. Cricetus C. Allo A. cricetus cricetus cricetus bursae praeglaci- nanus ehiki alis Hajnóczy Galéria Galéria Galéria Galéria Óriás-t Kis-galéria Total For the relative age determination of the studied fauna the following stratigraphie evidences were considered: 1. Cricetus cricetus praeglacialis was found in faunas of the Betfia and Templomhegy Phases in Hungary. 2. Cricetus cricetus nanus occurs in the faunas of the Betfia Phase (Jánossy 1979,1986). 3. The oldest arvicolid in the studied material is Mimomys pusillus (MÉHELY) found in localities Hajnóczy 1, Galéria 3 and Kis-galéria. Its last occurrence is known from the Betfia Phase (Jánossy 1979,1986). The material of the main localities (Hajnóczy 1, Galéria 1, 2, 3) and the Kisgaléria represents the Betfia Phase. The age of the other minor localities - yielding Allocricetus bursae only - is between the Betfia and Tarkő Phases but not determinable more precisely. REFERENCES FAHLBUSCH, V. (1964): Die Cricetiden (Mamm.) der Oberen Süsswasser-Molasse Bayerns. - Abh. Bayer. Akad. Wiss., Math.-naturw. Kl., N. F., 118: FAHLBUSCH, V. (1969): Pliozane und pleistozane Cricetinae (Rodentia, Mammalia) aus Polen. - Acta Zool. Cracov., 14: HÍR, J. (1982): Adatok az Odorvár és környékének negyedidőszaki felszín- és karsztfejlődéséhez (Angaben zur quarteren Morpho- und Karstgenetik des Berges Odorvár und seiner Umgebung). - Földr. Ért., 31: HÍR, J. (1985): A Hajnóczy-barlang üledékei [Sediments in the Hajnóczy Cave]. - Karszt és Barlang, (1985) : HÍR, J. (1989): A Tarkői-kőfülke Allocricetus anyagának újravizsgálata (Revision of the Allocricetus material from the Tarkő Rocksheiter. - Folia Hist.-nat. Mus. Matraensis, 14: JÁNOSSY, D. (1979): A magyarországi pleisztocén tagolása gerinces faunák alapján. - Akadémiai Kiadó, Budapest, 207 pp. JÁNOSSY, D. (1986): Pleistocene Vertebrate Faunas of Hungary. - Akadémiai Kiadó and Elsevier, Budapest - Amsterdam, 209 pp.

16 KORDOS, L. (1974): Jelentés a Hajnóczy-barlangban végzett őslénytani és üledékföldtani vizsgálatokról [Report on the sedimentological and paleontological studies of the Hajnóczy Cave]. - Karszt- és Barlangkutatási Tájékoztató, 5-6: MALEZ, M. and RABEDER, G. (1984): Neues Fundmaterial von Kleinsaugern aus der altpleistozânen Spaltenfüllung Podumci 1 in Norddalmatien (Kroatien, Jugoslawien). - Beitr. Paläont., 11: MEIN, P. and F^EUDENTHAL, M. (1971): Une nouvelle classification des Cricetidae (Rodentia, Mammalia) du Tertiaire de l'europe. - Scripta Geol., 2:1-37. PRADEL, A. (1988): Fossil hamsters (Cricetidae, Rodentia) from the Pliocene and Quaternary of Poland. - Acta Zool. Cracov., 31: SCHAUB, S. (1930): Quartare und jungtertiäre Hamster. - Abh. Schweiz. Pal. Gesellsch., 49:1-49. VARGA, CS. (1978): Az Odor-vári Hajnóczy-barlang [The Hajnóczy Cave of Odor-vár]. - Karszt és Barlang, (1978): VARGA, L. (1978): A dél-bükki Odorvár barlangjai [Caves of the Odorvár of the southern Bükk Mountains]. - Földr. Közlem. 26 (52): Author's address:dr. János HÍR Municipal Museum of Pásztó H-3060 Pásztó Kölcsey u. 8. Hungary

17 APPENDIX Cricetus cricetus praeglacialis SCHAUB, 1930 Hajnóczy 1. Galéria 1. Galéria 3. 1 w w a Wp 1M? IM IM IM M IM IM IM IM Cricetus cricetus nanus SCHAUB, 1930 Galéria 1. Galéria 2. Galéria 3. 2Mi IM2 IM IM3 IM2 1 w W a Wp Allocricetus ehiki SCHAUB, 1930 Hajnóczy 1. Galéria 1. Galéria 2. 2M w w a Wp pac past IM IM b 2 mandibula no.l: ltr: 5.2 Mi M M no.2: ltr:5.0 Mi M M M IM M IM ; - IM b 2M

18 Allocricetus bursae SCHAUB, 1930 Hajnóczy 1. Galéria 1. Galéria 2. 1 w w, Wp pac past ale ml 1 2M IM a IM] lm 3 l UM IM Mi M M a a 1 UM

19 1 w w a Wp pac past alc ml pl M 2 6M 3 8Mi 12M 2 4M 3 1 w mtl a a a a a a 1 w. Wp pac w alc ml w alc ml Galéria 3. 1 w w a a w p p pac past mtl ml alc ÍM ÍM lmi

20 1 w ÍM w a 1.02 ÍM IM Wp pac past alc ml pl Oriás-terem mandibula ltr:4.4 1 w w. Wp pac past alc ml Ml Kis Galéria M 2 M 3 ÍM 1 lmi 2M 2 IM3 2M l _ a a Summary of the measurements of Allocricetus ehiki specimens from Villány 3 and Osztramos 3 min. X max. no. M*l w M w M w Mil w M2I w M w

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

FRAGMENTA MINERALOGICA ET PALAEONTOLOGICA 16. BUDAPEST 1993. p. 61-80 Allocricetus ehiki Schaub, 1930 (Rodentia, Mammalia) finds from Villány 3 and Esztramos 3 (Hungary) by J. HÍR Abstract: The metrical