F R A G M E N T A P A L A E O N T O L O G I C A H U N G A R I C A 24-25, B U D A P E S T , 2007

A revision of three Pleistocene subspecies of Panthera, based on mandible and teeth remains, stored in Hungarian collections

by

Eszter Piroska HANKÓ

Abstract — This paper is the first comperehensive revision of the Pleistocene lion-like cat remains stored in Hungarian collections. The morphological comparison and the cladistical analysis were based on 132 teeth and 9 mandible remains. The fossil cat species, previously described as Leo gombas^pegensis K R E T Z O I , 1938 stands near to the recent jaguar, therefore it must be referred to Panthera onca gombas%oegensis. The Middle Pleistocene lion-like cat is a subspecies of Panthera leo (P. leo fossilis) just as the Late Pleistocene cave lion (P. leo spe/aea), which is not a direct descendant of the former, but represents a separate more advanced offshoot.

Keywords — Panthera, Pleistocene, Hungary, morphology, teeth, mandible, taxonomy.

HANKÓ, E . P.: A revision of three Pleistocene subspecies of Panthera, based on mandible and teeth remains, stored in Hungarian collections. — Fragmenta Palaeontologica Hungarica, 24: 25—43.

Introduction

The territory of Hungary and the surrounding intra-Carpathian areas are mostly covered by Pleistocene sediments. Especially the cave deposits are very rich in vertebrate remains, their palaeontology and stratigraphy were reviewed by JÁNOSSY (1986).

Most of the mammalian groups from the Pleistocene of Hungary have been studied, but little is known about the fossil felids. In his work, published in the 1920s, KRETZOI established the taxonomy of Eelidae (KRETZOI 1929), and later he described the oldest "lion" remains (Leo gombas^oegensis KRETZOI, 1938), but a detailed morphological comparative analysis was not published. JÁNOSSY (1969) contributed a survey on the metacarpals, metatarsals and canines of lions, but could not distinguish different species by this method.

The aim of this paper is a taxonomic revision of the Pleistocene Pantherinae from Hungary on the basis of mandible and teeth remains because these skeletal parts bear the most characteristic morphological features. In addition to the morphological comparison the results of a cladistical analysis will also be discussed.

First, I wished to clarify the taxonomic position of Leo gombas-^oegensis, which is recently considered to be a fossil

subspecies of the genus Panthera. Owing to the priority'" of the species name described by Kretzoi, it was renamed Panthera gombas^oegensis ( K R E T Z O I , 1938) by Hi: MM ER (1971). The latest investigations identify this taxon as the fossil subspecies of the recent jaguar, and it is referred to as Panthera onca gombas^oegensis (HEMMER 2001).

Secondly I examined the problem of other Pleistocene lion-like cats. The "lion-like" term has been used in the last decades only for two groups of Panthera, the older "fossilis" group (Middle Pleistocene), and the vounger "spelaea" group (Late Pleistocene). Some authors consider the two groups as two different species of the genus Panthera (ARGANT 1998, BARYSHNIKOY & BOESKOROV 2001): P . fossilis and P. spelaea. According to others ( R I E D E L 1982, SCHUTT 1969) both groups belong to the same species of the living African and Asian lions (P. leo), merely representing two different subspecies. I have examined the relationship between the two Eurasian lion subspecies Panthera leo fossilis (REICHENAU, 1906) and Panthera leo spelaea (GOLDFUSS, 1810).

Finally, the relation between Panthera leo spe/aea and the recent lion will be also discussed on the basis of their morphology.

Localities (Figure 1)

Püspökfurdö"jBetfia — The Betfia locality-complex is situated in Romania, near the LIungarian border. The finds originate from locality JV° 5. Its age is Early Middle Pleistocene ( K R E T Z O I 1941a).

Gombas^pglGombasek — The limestone quarry of Gom-baszög/Gombasek is situated in Slovakia near the Sajó River. T A S N Á D I - K L ' B A C S K A (1935) determined two fossil assemblages. In one locality only Late Pleistocene faunal

elements were found. From the other five localities Middle Pleistocene fossils were recovered including remains of Felidae. KRETZOI (1941b) considered the age of the older localities as Cromerian Interglacial, and correlated the fauna with the Mosbachian faunal association.

Vértessé/ős II — The fossils originate from a freshwater limestone (travertine) quarry (JÁNOSSY, 1990). According to KORDOS (1994) the fauna can be correlated

26 HANKÓ,

with the Holsteinian Interglacial. Uppony I — The rock-shelter of Uppony I . is the

remnant of a former large cave. The type fauna of "Uppo-nyian phase" within the Middle Pleistocene was revised by K O R D O S (1994), who separated the older (Holsteinian) layers 7—8 from the younger (Rissian) layers 1—6.

Paks — In the loess-quarry of the Paks brickyard 8 paleosoil layers in a 20-25m thick loess sequence were recorded. P É C S I (1993) correlates the loess layers with Rissian and Würmian Glaciations.

S^uhogy-Csorbakő— This is a shelter-cave in the quarry at Szuhogy village. The cave fill was divided into three

E . P.

levels. The two lowTer layers were rich in larger mammal bones. The age of the fauna is Holsteinian-Rissian ( J Á N O S S Y & V Ö R Ö S 1985).

Kövesvárad — The shelter-cave was discovered by D. J Á N O S S Y in 1955. The fauna was described by J Á N O S S Y (1963, 1986) and its age was correlated with the Mimomys savini partial range zone.

Solymár-Ordöglyuk Cave — The fossils came from the infilling sediment of the shaft near the entrance of the cave. The fauna, revised by Jánossy, indicates a late Middle Pleistocene age ( J Á N O S S Y 1986, V Ö R Ö S 1988).

Figure 1 — Sketch map of the localities.

Tokod-Nagyberek — The locality is situated in a freshwater limestone section. The bones came from loess and palaeosoil layers intercalated between thick banks of the limestone. Early collections provided a fauna of the Würmian age ( J Á N O S S Y 1971, 1986). Later (1990-1992) G A S -P A R 1 K made excavations at Tokod-Nagyberek in a deep ravine from deposits of a tetarata basin succession that is probably identical with J Á N O S S Y ' s locality. G A S P A R I K (1993) published two faunas of different ages from the locality. The upper layers provided a cold climate fauna from the Würmian glaciation, but the lower (major) part of the loess intercalation, where the larger mammal bones came from, is older and contained an interglacial fauna. Recently, M E S Z O E L Y & G A S P A R I K (2002) concluded that this older, warm climate fauna represents the Eemian (Rissian-Würmian) interglacial (-100-110 ky BP). According to

G A S P A R I K (pers. comm.), after the Eemian the upper freshwater-limestone beds became fragmented because of tectonical movements, and the Würmian fauna was washed into the succession through a fissure opened.

Szelim Cave — GAÁL (1934) distinguished two parts in the upper diluvial level. The fauna is typical of the Toko-dian phase within the Würmian Glacial.

Kiskunfélegyháza — Panthera leo spelaea specimens came from a sandpit and got to the Hungarian Geological Institute. According to the inventory, their age is Late Pleistocene.

Kiskevély Cave — The fauna of this cave was referred to Tokodian/Subalyukian phase within the Würmian Glacial ( J Á N O S S Y 1986).

Igric/Pestere Cave — The fossil assemblage represents a typical glacial (Würmian) fauna with elements as Crocuta spelaea, Panthera leo spelaea, Ursus spelaeaus ( K O R M O S 1914).

Háromkút-Cave — KADIC (KADIC & M O T T L 1944) performed the first excavations in the cave. VERTHS (1965) specified the age of the fauna as Würmian.

htál/óskő Cave — The fauna was first described by JÁNOSSY (1952, 1955) and later revised by VÖRÖS (1984). RlNGF.R (2002) perfomed further excavations in order to

check the stratigraphy; C 1 4 data gave an age between 27— 33 ky BP.

The approximate stratigraphie positions of the localities are shown in Table 1. The ages of the chronostrati-graphic units were taken from the updated Geological Time Scale (GRADSTHIN et al. 2004).

Table 1 — The stratigraphie position of localities and the ranges of taxa studied.

Absolute age (ky)

Chrono-stratigraphy

Glacial chronology

Stages Localities Panthera ssp.

Holocene

isto

cen

e

W ü r m i a n Istállóskő, Igric/Pestere, H á r o m - k ú t

Szelim, Kiskevély, Kiskunfé legyháza a le

o sp

elae

a

1 0 0

ate

Pie

1 ' S *•*

ate

Pie

E e m i a n Tokod-Nagyberek SS

G

ing

2 0 0 Tor

S o lymár ?

R i s s i a n

Szuhogy-Csorbakő leo

foss

ilis

3 0 0 Paks ys

sto

cen

e - Uppony I. layer 4. a í

sto

cen

e

H o l s t e i n i a n Uppony I . layers 7, 8

e P

lei

Vértesszőlős I I .

4 0 0 e P

lei

Mid

dl

M i n d e l i a n

G t K ö v e s vár ad zoeg

ensi

s

ihar

i;

'nba

s.

5 0 0 C r o m e r i a n

CQ G o m b a s z ö g / G o m b a s e k ag

oi

C r o m e r i a n

Be t f i a /Püspökfürdő

1 thera

one

6 0 0 G ü n z

Pan

Material

All fossil specimens were studied that are stored in the Department of Geology and Palaeontology of the Hungarian Natural History Museum and in the Geological Institute of Hungary (Table 2). Comparaüve investigations were based on 18 individuals of recent Panthera ko,

(mainly from East-Africa) and 5 individuals of recent Panthera onca in the Mammalian Collection of the Hungarian Natural History Museum. Further 5 individuals of Panthera leo and 2 individuals of Panthera onca were examined in the Naturhistorisches Museum,Vienna.

Table 2 — A catalogue of the studied speciemens. — V 59.1039; G 57.60 and Gyn440 inventory number formulae, and the specimens without number indicate Dept.. of Geolog)' and Palaeontology, Hungarian Natural 1 listory Museum as depositor}'; V 24062 and Ob. 2978 specimens are from the Hungarian Geological Institute.

Inventory NQ

Species Locality Age Skeletal element Specimen No

V 59.1039 Panfhem onca gombasyoegensis C lombaszög/( iombasek Middle Pleistocene P4 fr. 1 V 59.1044 Panthera onca gombasyoegensis C jombaszög/Gombasek Middle Pleistocene Ci„f

C fr. V 59.1064 Panthera onca gombasyoegensis Gombaszög/Gombasek Middle Pleisti icene 1 sin. 1 V 59.1084 Pan'thera onca gombasyoegensis Gombaszög/Gombasek Middle Pleistocene Mi sin. 1 V 59.1085 Panthera onca gombasyoegensis Gombaszög/( iombasek Middle Pleistocene Mi sin. 1 V 59.1041 Pan thera onca gombasyoegensis ( Ii imbaszög ( ii imbasek Middle Pleistocene P3 dext. 1 V 60.1185 Panthera onca gombasyoegensis Uppony I . Layer 4. Middle Pleistocene Maxilla fr. +

P-]

V 60.1251 Panthera onca gombasyoegensis Uppony I . Layer 7. Middle Pleistocene Mandibula sin. fr. Mi P4 fr.

1 Mandibula sin. fr. Mi P4 fr. !

V 63.246 Panthera onca gombasyoegensis Kövesvárad Middle Pleistocene Mi 1 V 69.642 Panthera onca gombasyoegensis Vértess2Őlős 11 Middle Pleistocene Qnf

C fr. \

V 69.643 Panthera onca gombasyoegensis Vértesszőlős I I : Middle Pleistocene Mandibula sin. fr. + P3 C fr.

j V 69.646 Panthera onca gombasyoegensis Vértesszőlős I i. Middle Pleistocene I " dext. fr. 1 Y 10726 Panthera onca gombasyoegensis Püspökfürdő/Betfia Middle Pleistocene P4 1 V 10717 Panthera onca gombasyoegensis Püspökfürdő/Betfia Middle Pleistocene P4 fr.

P'1 fr. P4 fr. M, fr.

1 V 24062 Panthera onca gombas-ypegensis Gombaszög/Gombasek Middle Pleistocene Mandibula sin. fr.

Mi P4 C

1

V 24063 Panthera onca gombasyoegensis Gombaszög/Gombasek Middle Pleistocene Maxilla fr. P4

P3

C fr. Mandibula fr.

; V 69.647 Panthera onca gombasyoegensis Vértesszőlős I I . Middle Pleistocene P4 sin. 1 Gyn489 holotype

Panthera onca gombas-ypegensis Gombaszög/Gombasek Middle Pleistocene P4 1

Gyn440 Panthera onca gombasyoegensis Gombaszög/C iombasek Middle Pleistocene P4 fr. 1 V 59.1044 Panthera onca gombas-ypegensis Gombaszög/Gombasek Middle Pleistocene C inf. sin.

C fr. ! V 60.1185 Panthera onca gombasyoegensis Upponv I . 1 .aver 4. Middle Pleistocene Maxilla fr. +

C. fr. P- fr.

1 Maxilla fr. + C. fr. P- fr. !

V 59.1064 Panthera onca gombasyoegensis Gombaszög/Gombasek Middle Pleistocene I 1 \ (Ö.159 Panthera onca gombssyoegensis I 'pponv I . Laver 8. Middle Pleistocene Mi dext. 1 V 65.258 Panthera onca gombasyoegensis 1 "pponv 1. 1 .aver 7. Middle Pleistocene P3 sin. 1 V10717 Panthera onca gombasyoegensis Püspökfürdő/Betfia Middle Pleistocene P1 fr.

P3 fr. P4 fr. Mi fr.

1 V 69.672 Panthera leo fossilis Vértesszőlős 11. Middle Pleistocene L

I2 I 2

I 1

2

2

2

L I2 I 2

I 1

2

2

2

V 69.651 Panthera leo fossilis Vértesszőlős I I . Middle Pleistocene Mandibula sin. fr P4 fr. Mi fr.

1 V 69.652 Panthera leo fossilis Vértesszőlős I I . Middle Pleistocene I 3

I fr. V 69.675 Panthera leo fossilis Vértesszőlős 11. Middle Pleistocene Mi

Mi fr. ! V 69.689 Panthera leo fossilis Vértesszőlős I I . Middle Pleistocene P' sin. fr. 1 V 69.681 Panthera leo fossilis Vértesszőlős I I . Middle Pleistocene ( : fr.

C sup. 'I

V 69.678 Panthera leo fossilis Vértesszőlős I I . Middle Pleistocene c: fr. 1

Table 2 — continued.

Inventory NQ

Species Locality Age Skeletal element Specimen NQ

V 69.650 Panthera leo fossilis Vértesszőlős I I . Middle Pleistocene P 4 fr. L I 3 fr.

j

- Panthera leo fossilis Vértesszőlős I I . Middle Pleistocene I ' fr. - Panthera leo fossilis Vértesszőlős I I . Middle Pleistocene L 1 V 69.676 Panthera leo fossilis Vértesszőlős I I . Middle Pleistocene C fr. 1 V 69.286 Panthera leo fossilis Solymár Middle Pleistocene Mandibula fr. + M,

fr. P 4fr.

j

Gyn.496 Panthera leo fossilis Vértesszőlős Middle Pleistocene P*fr. p3 C fr.

I Gyn.495 Panthera leo fossilis Solymár Middle Pleistocene P4 fr. 1 V 10822 Panthera leo fossilis Paks Middle Pleistocene Mi

O P \

V 59.245 Panthera leo spelaea Istállóskő Late Pleistocene M I sin. 1 G 57.60 Panthera leo spelaea Szelim Late Pleistocene Mi dext. fr. 1 V 59.242 Pa ni fh ra leo spelaea Istállóskő Late Pleistocene Mi dext. 1 V 59.244 Panthera leo spelaea Istállóskő Late Pleistocene ( : fr. 1 V 59.261 Panthera leo spelaea Istállóskő Late Pleistocene P3 dext. 1 V 60.1092 Panthera leo spelaea Szuhogy Middle Pleistocene C inf. 2 Ob.2978 Panthera leo spelaea Igric/Pestere Late Pleistocene Cranium +

P4

P3

CSUP fr. M 1

1 2 2 2 1

Ob. 29 83 Panthera leo spelaea Igric/Pestere Late Pleistocene Mandibula L fr. Ci„f fr. P3 Pt M,

1 2 2 2 2 2

V 60.1785 Panthera leo spelaea Igric/Pestere Late Pleistocene Cranium + maxilla P4 fr. P3 C fr. Mandibula Mi Mi fr. P4 fr. P 3 P3 fr. Cfr.

1 2 2

V 63.1480 Panthera leo spelaea Három-kút Late Pleistocene Mi dext. 1 V 63.1621 Panthera leo spelaea Szuhogy Middle Pleistocene C fr. 1 V 64.798 Panthera leo spelaea Tokod-Nagyberek Late Pleistocene p3 1 Ob.2977 Panthera leo spelaea Kiskevély Late Pleistocene Mandibula sin.

M, P4 C fr. Maxilla fr. P4

C fr.

j

V11433 Panthera leo spelaea Kiskunfélegyháza Late Pleistocene Mandibula fr. + Mi dext. P4 dext. M| dext. C dext. 2

H A N K Ó , E . P .

Methods

The examinations of teeth and mandibles were based on both qualitative features and measurements. In the morphological description of bones the works of S C H M I D (1940), S C H U T T (1969), V A U G H N et al. (2000) and Z B O R A Y

(2001) were taken into consideration. For comparison, skeletal elements of Recent lions (mainly Panthera leo nubica) and jaguars were studied. The most important and diagnostic features for morphological analyses were found in the lower molar, the upper and lower premolars and the mandible. Only these elements have been measured and described morphologically in details, but the incisivi and the canines have been also taken into consideration.

The measurements on the teeth and mandibles were taken in mm as shown in Figures 2—6.

The results of the morphological studies were evaluated by the cladistical method of Hennig 86 Version 1.5 [ K O R S Ó S (1999) after F A R R I S (1988)].

Par

Par

Par

dist

Figure 2 — Measurements of P4. — A lateral (buccal) view, B: lateral (palatal) view, C: occlusal view, dist: distal, M: metacone, mes: mesial, Ms: metastyle, Par: paracone, Pr: protocone, Ps: parastyle. — 1: maximum length, 2: maximum height, 3: length of paracone, 4: length of metacone, 5: length at the constriction, 6: length at the protocone, 7: width at the protocone, 8: width behind the protocone, 9: the rear maximum width (after SCHMID 1940).

pal lbucc

A B Figure 3 — Measurements of P3. — A: Lateral view, B: Occlusal

view, bucc: buccal, C: cingulum, dist: distal, H: hypocone, pal: palatal (for other abbreviations, see Figure 2). — 1: maximum length, 2: maximum height, 3: height of paracone, 4: length of paracone, 5: maximum width (after SCHMID 1940).

t l i s i

Ling

Mes

Bucc

Figure 4 — Measurements of P 4 . — A: lateral (buccal) view, B: occlusal view, H: hypoconid, ling: lingual, Par: paraconid, Pr: protoconid (for other abbreviations, see Figure 3). — 1: maximum height, 2: height of protocone, 3: maximum length, 4: maximum width (after SCHMID 1940).

A

Figure S — Measurements of Mi. — A; Lateral view, B: Occlusal view, I: incisor, T: talonide, (for other abbreviations see Figure 4). — 1: maximum height, 2: height of protoconid, 3: maximum length, 4: height of crown at the incisor, 5: height of paraconid, 6: maximum width (after Schmid 1940).

Figure 6 — Measurements of the mandible in lateral view. — 1: length of mandible (from premaxillary bones to the end of Mi), 2: height of corpus mandibulae in front of P3,3: height of corpus mandibulae behind Mi.

Pleistocene Panthera in Hungarian collections

Systematics

Family Felidae F I S C H E R D E W A L D H E I M , 1817 Subfamily Pantherinae P O C O C K , 1917

Genus Panthera O K E N , 1816 Panthera onca L I N N É , 1758

Panthera onca gombaszoegensis ( K R E T Z O I , 1 9 3 8 ) (Plate I : 1-5; Figure 7: A)

1938: I mgombasypgensis n. sp. — K R E T Z O I , p. 100, pl. 1, figs 1-7. 1963: Leo ci. gombasyoegensis K R E T Z O I — J Á N O S S Y , p. 117, pl. 1, fig. 20. 1971: Panthera gombasyoegensis ( K R E T Z O I ) — H E M M E R , p. 702. 2001: Panthera onca gombasyoegensis ( K R E T Z O I ) — H E M M E R , p. 708, Pis 132, 133, 134.

Table 3 — Measurements of P 4 of Panthera onca gombaszoegensis.

V 69.647 G vn489 Gyn440 V 24063 Maximum length 32.0 33.0 32.0 SIM Length at the constriction 30.0 32.5 30.5 29.5 Length at the protocone - 32.0 31.0 ? Length of paracone 12.0 13.0 12.0 12.li J .ength of metacone 12.0 13.0 12.0 12.0 Para- and metacone length 24.0 25.0 23.5 23.0 Width at the protocone - 18.0 17.0 -Rear biggest width - 12.0 11.5 10.5 Width behind the protocone 12.(1 12.5 10.5 Height of paracone - 17.0 - 1 5.(1 Height of protocone 6.0 5.5 Maximum height 18.0 - 16.0

P 4 — The degree of the constriction between the protocone and parastyle is negligible (in occlusal view). The well-developed enamel crest runs from the paracone to the protocone. The protocone is medium developed in most cases, except in specimen Gyn 489, where it is strongly developed. The parastyle is strongly expressed. On the buccal side of two of the four teeth a developed prepara-style (secondary cone) can be recognized. The tooth is not thickened on the buccal side. On the buccal side the margin of enamel rises steeply up toward the metastyle.

Table 4 — Measurements of P 3 of Panthera onca gombaszoegensis.

V 24063 Maximum length 21.0 Length of paracone 9.0 Height of paracone 13.0 Maximum width 9.5 Width of front part 8.0 Maximum height 13.0

P 3 — The protocone is slightly developed, slightly extends to the palatal side. The paracone and the hypocone are strongly developed. The tooth has a cingulum on its distal side. The tooth has no protostyle in the mesial part, what in other case is only a very strong cingulum instead of a cone. (The protostyle is a typical feature of the jaguar. The development of the pro to style varies in the Recent jaguar wilirin the species and in one individual, too. In the same individual the left tooth seems to have a protostyle, while there is only a strong cingulum on the right tooth instead of protostyle.) The crown does not form an extension on the palatal side. The crown forms a little

arch on the buccal and palatal side towards the occlusal surface.

P 2— This is one of the less typical types of teeth. It is separated from the upper canine by a usually short diastema.

Table 5 — Measurements of P3 of Panthera onca gombaszoegensis.

V 69.643 V 59.1041 Maximum length 19.0 16.5 Length of protoconid 9.5 8 Height of protoconid 11.0 10 Maximum width 10.0 8 Width at the protoconid - 7

Maximum height 12.0 11

P3 — The paraconid is weakly developed and extends slightly to the lingual side. The protoconid is well-developed, the hypoconid is weakly developed. Cingulum is slightly developed. The tooth is relatively long.

Figure 7 — T h e typical enamel arch on the palatal side.

— A: The upper, rear premolar (P4) of Panthera onca gombasyoegensis Gyn489 from Gombaszög/Gombasek in lateral (palatal) view, B : The upper, rear premolar (P4) of the lion Panthera leo fossilis V 69.673 Vértesszőlős in lateral (palatal) view.

P4 — The paraconid and the protoconid are very strongly developed. The hypoconid is medium developed. The paraconid is higher than the hypoconid. The protoconid gendy leans to distal direction. The cingulum is well developed. The cingulum and the hypoconid region is very strongly developed in the case of V 59.1039 from Gombaszög/Gombasek.

Table 6 — Measurements of P 4 of Panthera onca gombaszoegensis.

V 60.1251 V 24062 V10726 Maximum length 22.5 21.5 -Length of protoconid 9.5 10.0 11.5 Height of protoconid - -Maximum width 10.5 10.0 Width in front part 9.0 8.0

34

Par Par

HANKÓ, E . P.

strongly extends to palatal direction. The protostyle does not appear. The hypocone is strongly developed and the cingulum is well visible and developed on the distal side.

P4 — The paraconid is moderately developed and extends to lingual direction. The protoconid is strong and slightly leans to distal direcdon. The hypoconid is moderately developed. The metaconide is missing. The cingulum is poorly developed. The enamel of the crown is thickened on the Ungual side, under the protoconid in the case of the specimen from Solymár.

Table 11 — Measurements of P4 of Panthera leo fossilis.

Figure 8 — A : upper, rear premolar (P 4 ) of Panthera leo

spelaea; B: upper, rear premolar (P 4 ) of the Panthera leo fossilis (after F R E U D E N B E R G 1914).

Table 10 — Measurements of P 3 of Panthera leo fossilis.

V 69.689 V 69.682 Gyn496 Maximum length 24.5 24.0 18.0 Length of paracone 13.0 i l j i 1 5.0 1 ieight of paracone 14.5 18.5 Maximum width ! U) l l .o 12.0 Width of front part : 1.5 11.0 12.0 Maximum height 15.5 - 18.5

P 3 — The crown does not form an extension to palatal direction. The protocone is underdeveloped and

V 69.648 V 69.651 V 62.286 Maximum length 26.5 H.O 31.0 Length of protoconid 14.0 - -Height of protoconid 16.5 Maximum width 13.0 15.5 Width of front part 11.5 Rear biggest width 13.0 -Maximum height 19.5 -

M i — The paraconid is developed, the bottom of the mesial edge is rounded. The protoconid is developed, straight and rises higher than the paraconid. The talonide is very developed. The talonide of Mi of Panthera leo fossilis is the most developed of all examined fossil species. The crown is thickened on the lingual side, under the incisor. The margin of enamel rises considerably to occlusal direction.

Table 12 — Individual measurements of Mi of Panthera leo fossilis.

V 69.674/1 V 69.674/2 V 69.674/3 V 69.675/1 V 69.675/2 V 69.651 V 62.286 V10822 Maximum length 30.0 31.0 27.5 32.0 - 31.0 Length of protoconid 18.5 18.5 17.0 19.5 18.0 19.5 1 .ength of paraconid 16.0 15.0 14.0 17.5 15.1 LS.5 Maximum width 16.0 16.5 15.0 17.0 16.0 17.0 15.5 I leight of protoconid 18.0 17.0 15.0 17.5 - 1 S.i 1 Height of paraconid 15.5 15.5 14.5 17.5 -I leight at the incisor 10.0 10.0 10.0 11.5 12.0 -Maximum height 20.0 24.0 24.0 26.5

Mandible — The corpus mandibulae in front of P3 is lower than behind Mi. The fossa masseterica extends below the paraconid of the Mi. The mandible has two foramina mentale. The symphysis mandibulae is very massive, that can be seen mostly in lateral view. The diastema is moderately long, and only slightly arched.

Table 13 -

fossilis. Measurements of mandible of Panthera leo

V 62.286 V 69.651 Maximum length 151.0 138.0 Height of coipus mandibulae in front of P3 61.0 ! leight of corpus mandibulae behind ,\L 63.0 -Length of diastema 25.0 25.li Length of symphysis mandibulae 1.0 -

Pr

Par

Pr

Occurrence — Vértesszőlős, Paks, Szuhogy-Csorbakő, Solymár.

Stratigraphical distribution: Middle Pleistocene, Holsteinian Interglacial, Rissian Glacial.

Figure 9 — A : lower molar (Mi) of Panthera leo fossilis V 69.674 from Vértesszőlős; B: lower molar (Mi) of

Panthera leo spelaea V 59.245 from Istállóskő.

Panthera leo spelaea ( G O L D F U S S , 1 8 1 0 ) (Plate I I I : 1-3, Plate IV: 1-3; Figure 8: A, Figure 9: B)

1810 Felis spelaea — GOLDFUSS p. 277, PI. 5, fig. 1. 1969 Panthera leo spelaea — SCHUTT p. 213, PI. 23, fig. 4, PI. 24. 1969 Leo spelaeus — JÁNOSSY p. 588. 1971 Panthera (leo) spelaea — VERESHCHAGIN p. 181, figs 6:3, 9:2, 13:3, 14:2, 16:1, 18:2, 19:2, 20:2, 27:1,2. 1971 Panthera leo spelaea — THENIUS p. 124, f. 9-14. 2001 Panthera spelaea — BARYSHNIKOY & BOESKOROY p. 21, figs 2-3. 2006 Panthera leo spelaea — BONA p. 165, fig. 5, Pl. 1.

Table 14 — Measurements of P 4 of Panthera leo spelaea.

V 60.1785 Ob.2977 Ob.2978 Maximum length - 34.0 36.5 Length at the constriction - 32.5 34.5 Length at the protocone - 33.5 36.5 Length of paracone 14.0 12.5 14.0 Length of metacone - 13.5 15.0 Length ot para-, and metacone - 25.0 27.0 Width at lire protocone - 17.0 18.5 Rear biggest width 12.0 13.0 Width behind the protocone 12.5 12.5 1 leight of paracone 17.5 -Height of protocone - 6.5 4.5 Maximum height 19.0 -

P 4 — The degree of the constriction (in occlusal view) between the protocone and parastyle is considerable. The form of the frontal (mesial) part of tooth is similar to that of the Recent lion. The protocone is slightiy developed while the paracone is strong. The enamel crest running from paracone to protocone is well-developed. The metacone is strongly developed, and the metastyle rises over the metacone. The parastyle is strongly developed. The preparastyle is not present. The teeth are thickened on the buccal side. On the buccal side the margin of enamel rises steeply up toward the metastyle.

Table 15 — Measurements of P 3 Panthera leo spelaea.

V 59.261 V 64.798 V 60.1785 Ob.2978 Maximum length - 26.5 28.0 24.5 Length of paracone 13.1) 12.0 13.5 13. i Height of paracone - 16.0 17.0 16.5 Maximum width 12.5 14.0 13.5 11.0 Width of front part - 10.0 12.0 10.5 Maximum height - 16.0 17.0 14.0

P 3 — The crown does not form a palatal extension below the paracone. The protocone is underdeveloped and it extends strongly to palatal direction. The paracone is developed. The lower rim of the cown is arched in occlusal direction on both the palatal and the buccal side.

The teeth have no secondary protostyle. The hypocone is developed. The cingulum is strongly developed on the mesial part, exept the specimen from Igric/Pestere.

Table 16 — Measurements of P 3 of Panthera leo spelaea.

V 60.1785 Ob.2983 Maximum length 17.5 17.5 Length of protoconid 10.0 8.5 Height of protoconid 12.0 11.5 Maximum width 10.0 8.5 Width of front part 7.0 6.5 Maximum height 12.0 11.5

P3 — The paraconid is slightly developed, it leans moderately to Ungual directon. The hypoconid is sUghtly developed. Cingulum is present. The teeth have oval outlines.

Table 17 — Measurements of P4 of Panthera leo spelaea.

V 60.1785 V11433 Ob.2977 Ob.2983 Maximum length 24.5 25.5 25.0 26.0 Length of protoconid 12.0 12.0 11.5 13.5 1 leight of protoconid r . n 15.5 -Maximum width 12.5 12.0 12.0 13.0 Width of front part 12.0 8.5 9.5 11.0 Rear biggest width 11.5 12.0 11.5 12.0 Maximum height 18.0 17.5

P4 — The paraconid is medium developed, it extends sUghtly to Ungual direction. The protoconid is strongly developed and leans to distal direction. The hypoconid is medium developed. The cingulum is weakly developed. The paraconid and hypoconid are equally developed, except the specimen from Kiskevély, where the paraconid is greater than the hypoconid.

M i — The paraconid is weU-developed. The mesial edge of paraconid is Uttle arched except the specimen from Kiskevély and Három-kút, where this edge is straight. The protoconid is strongly developed and rises higher than the paraconid. The apex of protoconid leans

Table 18 — Measurements of Mi Panthera leo spelaea.

V 59.245 V 59.242 G 57.60 V 63.1480 V 60.1785 V 11433/a V11433/b Ob.2977 Ob.2983 Maximum length 27.0 31.0 32.0 27.0 27.0 26.5 25.0 25.5 2S.0 Height of protoconid 17.0 19.0 19.0 17.0 r . n 16.0 16.0 15.5 16.5 Height of paraconid 14.5 17.5 17.0 15.0 14.5 13.5 1 1.0 13.5 14.0 Maximum width 14.5 16.5 16.0 14.5 14.5 13.5 13.0 12.5 i 5.5 Height of protoconid 15.0 16.0 - 15.0 15.5 14.0 13.0 16.0 Height of paraconid 15.0 16.0 - 15.0 16.0 1 5.5 15.0 14.0 -Height at the incisor 9.5 9.0 10.5 8.0 9.0 8.5 8.0 7.0 7.5 Maximum height 21.0 21.0 - 19.0 21.0 18.5 19.0 19.5 18.0

to distal direction. The talonide is very sUghtly developed. The enamel is thickened on the lingual side of the teeth, except the specimen of Igric/Pestere. The lower margin of enamel rises in occlusal direction below the protoconid on the buccal side. This curve is much less developed than in the case of P. /. fossilis.

M a n d i b l e — In buccal view, the end of the symphysis mandibulae lines with the middle oi the diastema. The corpus mandibulae has the same height in front of P3 and behind Mi. The fossa masseterica does not extend under the Mi, it ends at the line of the distal edge of protoconid. The lower rim of the mandible between the diastema and the incisivi is sraight.

O c c u r r e n c e — Tokod-Nagyberek, Szelim, Kiskevély,

Kiskun félegyháza, Istállóskő, Igric/Pestere, Három-kút. Stratigraphical distribution: Late Pleistocene, Eemian

Interglacial, Würmian Glacial.

Table 19 — Measurements of mandible of Panthera leo

spelaea.

Ob.2977 V 60.1785 Ob.2983 Maximum length 104.0 123.0 136 1 leight of the corpus mandibulae in front of P;, 43.0 50.0 46.5 Height of the corpus mandibulae behind the Mi 43.0 50.0 50.5 1 .ength of diastema 15.0 24.0 30 Length of symphysis mandibulae 61.0 79.0 80

C o m p a r i s o n

Comparison of Panthera onca gombasyoegensis to Panthera onca and Panthera leo nubica

jaguar-like characters of P. 0. gombasyoegensis: - protocone of P3 extends only sUghtly in palatal direction; - mesial edge of the paraconid of Mi and the protoconid is straight; - P4 is not thickened on the buccal side;

protocone of P4 is present; - protoconid of Mi stands, not leans to distal direction;

symphysis mandibulae lines with the end of diastema in occlusal view;

cingulum of P4 is strongly developed; paraconid of P4 is strongly developed;

- absolute length of the diastema on mandible is small as compared to the diastema of jaguars and lions, and it is also small as compared to the length of the mandible; - length of mandible (V 24062) falls below the value of the smallest mandible sizes of Panthera onca;

height of the corpus mandibulae is the same behind the Mi and in front of the P3.

Uon-like characters of P. 0. gombasyoegensis: - protostyle is not present on P3; - protoconid on P4 leans sUghtly to distal direction;

on the P3 the crown does not form an extension on the palatal side; - protocone of P3 is slightly developed; - Table 20 shows that Mi is very long as compared to the length of the mandible. This ratio in P. onca gombasyoegensis corresponds to the proportion of the Panthera leo.

Table 20 — Proportion of Mi and mandible length.

Panthera onca gombaszoegensis n = l V 24062

Panthera leo n=16

Panthera onca n=5

Length of M i / mandible 25%

20-25.6% 19-23%

Further particular characters: - degree of the constriction between the protocone and parastyle is negügible (in occlusal view);

preparastyle secondary cone of P4 is strongly developed (where this cone is present), just Uke in the extant jaguar. This character is not typical for the present day lions. When it is present; it is sUghtly developed. The preparastyle can be a plesiomorphic character of Panthera that regressed in Panthera leo during the evolution, while it remained in Panthera onca.

The absolute sizes of aU teeth varied on the lower verge of the absolute teeth-sizes of Panthera leo and on the upper verge of the absolute teeth sizes of Panthera onca, except P3 that corresponds to the mean value of Recent Uons (Table 21).

Table 21 — Measurements of P3 (in mm).

Panthera onca Panthera Panthera gombaszoegensis leo onca

Length of P.i 17.75 17 15.5 Height of P-, 11 1 1.5 in Width ofP 3 10 9 8.3

E x p l a n a t i o n to P l a t e I I

(Teeth in natural size, mandible x0.7)

1 a-b Panthera leo fossilis (REICHENAU, 1906) — Vértesszőlős II, Middle Pleistocene, V 69.689, P3; a: occlusal surface, b: buccal view.

2 a-c Panthera leo fossilis (REICHENAU, 1906) — Vértesszőlős II, Middle Pleistocene, V 69.673, P4; a: occlusal surface, b: paládnál view,

c: buccal view.

3 a-b Panthera leo fossilis (REICHENAU, 1906) — Vértesszőlős II, Mddle Pleistocene, V 69.674, Mi; a: occlusal surface, b: buccal view.

4 a-b Panthera leo fossilis (REICHENAU, 1906) — Solymár-Ördöglyuk, Middle Pleistocene, Gyn495, P4; a: occlusal surface, b: buccal surface.

5 a-b Panthera leo fossilis (REICHENAU, 1906) — Vértesszőlős II, Middle Pleistocene, V 69.648, P4; a: occlusal surface, b: buccal surface.

6 Panthera leo fossilis (REICHENAU, 1906) — Solymár-Ördöglyuk, Middle Pleistocene, V 62.286, mandible; buccal surface.

The morphological data show that the "gombasyoegensis" remains have both some jaguar-like and lion-like tooth characters. However, the morphology of the mandible is more similar to P. onca than to P. leo, so the "gombasyoegensis" cannot be included into Panthera leo, but probably stays nearer to the Recent Panthera onca.

A closer relationship to Panthera onca is supported by palaeoecological aspects (HEMMER 2001, HEMMER et.

al. 2001, 2003) according to which P. onca gombasyoegensis probably lived under similar ecological circumstances, in marshlands, gallery forests and alluvial areas of rivers in Akhalkalaki (Transcaucasia) and in Mosbach (Germany) üke the Recent jaguar in (.entrai- and South-America. The faunal association of these localities included taxa that preferred swamp habitats e.g. Hippopotamus and Leutra.

Comparison of Panthera onca gombasyoegensis to Panthera leo fossilis

The remains of Panthera leo fossilis were recorded from Vértesszőlős, Solymár and Paks. At Vérteszőlős additionally P. onca gombasyoegensis was found.

Characters of P. I. fossilis, distinguishing from P. o. gombasyoegensis: - protocone of P3 extends strongly to palatal direction; - in P4, the degree of the constriction between the protocone and parastyle is bigger (in occlusal view) than in the "gombaszoegensis" specimens; - preparastyle in P4 is underdeveloped; - P4 is strongly thickened on the buccal side; - margin of enamel in P4 rises gently to metastyle on the palatal side (Figure 7); - mesial edge of the paraconid of Mi is rounded;

- Mi talonid is very strongly developed; on the buccal side of Mi , the margin of the enamel

forms an abrupt curve to occlusal direction, under the protoconid.

The measurements of the teeth and mandibles are significantly different (Table 22).

Table 22 — The average size of teeth and mandible of the two taxa compared.

Panthera onca gombaszoegensis

Panthera leo fossilis

Mi mean 20.5 29 P+ mean 32.5 37.75 Mandible mean 96 i l l

Comparison between Panthera leo fossilis, Panthera leo spelaea and the Recent Panthera leo

Ri K :i M-\ \ r (1906) did noi separate the "fossilis" re-mams from Panthera leo spelaea. F R E U D E N B E R G (1914) was the first to declare that the "fossilis specimens are not identical with those of the cave lion. According to him the most important character to distinguish "fossilis" is that the temporal region of the skull is more elongated and narrower than at the cave lion.

The "fossilis" deviates from the "spelaea" and the Recent lion in having a more compressed temporal region of its skull and consequently a relatively smaller frontal part of its brain cavity ( F R E U D E N B E R G 1914). The at least sub-specific treatment of the two fossil taxa was supported by other characters, for example body weight. The maximum weight of "fossilis" varied between 350-400 kg ( H E M M E R 2003, G U Z V I C A 1998) and the weight of "spelaea" could have been 10 % greater than that of the recent lion ( R A B E D E R et al. 2000).

Characters, distinguishing P. I. spelaea, from P. I. fossilis: - in P4 the degree of the constriction (in occlusal view) between the protocone and parastyle is considerable; - preparastyle on the P4 is not present;

Characters, distinguishing the Recent P. leo, from P. I. fossilis: - the protocone in P4 can be totally absent; - degree of constriction in P4 is greater than in "fossilis"; - margin of enamel on P4 on the palatal side rises "abrupdy" in some cases to occlusal direction; - talonide on Mi is weakly developed and lacking in some cases; - margin of enamel in Mi does not rise to occlusal direction on the buccal side; - mandible is much less massive in the symphysis region

than at the P. /. fossilis (in particular in the case of the specimen from Solymár).

Characters, distinguishing the Recent P. leo from P. I. spelaea: - the degree of constriction is the greatest of all examined species (it can be seen in occlusal view); - preparastyle in P4 is present in some cases, while this is totally absent at the cave lion; - the talonid on Mi is not present in some cases; - mandible is much less massive in the symphysis region, than that of P. /. spelaea.

Cladistical analysis

The morphological features were also examined by a cladistical analysis using 70 characters. The details of this analysis including the constructed cladograms will be published by HANKÓ & KORSÓS (in press). The outgroup was represented by Crocuta crocuta. In the phylogenetic tree P. o. gombasyoegensis and Panthera onca are grouped together and are separated from the other three subspecies of lion. This confirms the validity of inclusion of the "gombasyoegensis"-hmnch. into Panthera onca. In a previous cladistical analysis HEMMER (1981) presented two trees. In the first one, he used mainly physiological and ethological characters and some osteological features to show the relation

ship between the Recent species. In the rather similar second tree he included P. gombasyoegensis, but considered only one character (the absolute length of P3). Since in that paper he did not examine further characters and other fossil lion species his consideration of the "gombasyoegensis"-branch was not well supported by data.

Our cladogram (HANKÓ & KORSOS in press) shows a close relationship between P. I. fossilis, P. I. spelaea and the Recent P. leo. Based on the phylogenetic tree we suggest at least a subspeciftc treatment of P. I. spelaea and P. I. fossilis within the species of P. leo. The "fossilis"-branch is more distant while die "spelaed'-branch is closer to the Recent lion.

Conclusion

The taxonomic position and relationships of three fossil species, the "gombasyoegensis" KRETZOI and the two lion subspecies of the Middle and Late Pleistocene from Hungary have been clarified. The morphological analysis shows that the "gombasyoegensis" remains display some lion-like characters, as the longer P3 and the absence of the protostylus on the P3. However the separation from the Recent Panthera leo species is clearly demonstrated by the morphology of the Mi and P4. The characters, as the absence of thickening of the crown on the buccal side of P4, or the strong cingulum on P4 but especially the height of corpus mandibulae in front of P3, permit to include this animal in the species Panthera onca. Moreover, the corpus mandibulae is very massively developed in the symphysis region; so it is more similar to Panthera onca than to Panthera leo. Therefore this animal should be regarded as Panthera onca gombasyoegensis as it has been already suggested by HEMMER (2001). It is important to stress, that the morphology is very similar to P. onca, but not identical with that. Also the cladistic results show, that P. 0. gombasyoegensis is situated the nearest to P. onca suggesting a subspeciftc relationship.

Palaeontological data prove, that the ancestor of all Recent species of Panthera came from Africa, where the genus

appeared about 3 million years ago (Laetoli, Tanzania) (TURNER 1990). The Panthera appeared around 1.5 My in Europe.

SCHAU 13 (1949) studied remains of Felis arvernensis CROIZET & JOBERT, 1828 originating from the Upper Villafranchian (1.5 Ma) of Valdarno and Olivola in Italy. As a result, he described a new fossil species, Panthera toscana SCHAUB, 1949. Later FlCCARELLI & TORRE (1968) stated that the features of the skull from this material resemble the modern jaguar (Panthera onca L.), leopard (Panthera pardus L.) and tiger (Panthera tigris L.) rather than the lion (Panthera leo L.). The comparison of the remains of Panthera toscana resulted in a new view, and it was suggested that the fossil species should be included to Panthera onca as Panthera onca toscana (HEMMER 2001). This animal appeared at the end of the Middle Villafranchian (1.9 my) as the first Panthera in the Holartic (HEMMER 2003). According to HEMMER (2001) P. o. toscana is now- considered as an older, and P. o. gombasyoegensis as a younger fossil subspecies of P. onca in Eurasia. The equivalent of these taxa is Panthera o. augusta from the Pleistocene of N America (HEMMER 2003).

Based on the mandible and teeth characters, the Middle and Late Pleistocene lions are included into the frame of the

Explanation to Plate IV

1 a-b Panthera leo Spelaea ( G O L D F U S S , 1810) — Istállóskő, Late Pleistocene, V 59.245, Mi; a: buccal view, b: occlusal surface; x l .

2 a-b Panthera leo Spelaea ( G O L D F U S S , 1810) — Kiskevély, I .ate Pleistocene, Ob.2977, mandible; a: lingual view, b: view from above; x0.77. 3 a-b Panthera leo Spelaea ( G O L D F U S S , 1810) — Igric, Late Pleistocene, V 60.1785 mandible, Mi , P4, P3, C l n l; a: buccal view, b: Mi and

P4 buccal view; x0.65.

42 HAN species Panthera leo as two subspecies: Panthera leo fossilis and Panthera leo spelaea. Only the massive corpus mandibulae of some specimens seems to exceed the frame of P. leo, but it probably can be inserted in the range of intraspecific variability. On the other hand the canines, the I 3 , the premolars and molar (Mi) verify the close relationship with the Recent lion.

The separation of Panthera leo fossilis from Panthera leo spelaea is confirmed by the morphology of protoconid of Mi, the development of talonid of Mi , and the presence or absence of the preparastyle on P4.

The cave lion differs from the Recent lion in the height of corpus mandibulae, the absence of the preparastyle on P4

and the absolute sizes of the teeth and mandible. The "fossilis" was the first representative of the lions in

Eurasia. The oldest specimen is known from 700.000 years old deposits of Isernia in Italy ( S A L A 1990). From Hungary, it is known from about 350.000 years old layers of

D, E. P.

Vértesszőlős. According to HEMMER (1974) the lions migrated from Africa to Eurasia during the Middle Pleistocene. This animal could be P. I. fossilis. Its descendant was P. I. spelaea that developed only in Eurasia. The theory, that the P. I. spelaea derived from the P. I. fossilis is neither supported by the present morphological nor the cladistical analysis.

Based on the cladogram, P. I. spelaea is more, while P. I. fossilis is less similar to P. leo. (The cave lion stands nearer to the Recent lion in time, namely separated later from the common ancestor. It is possible, that the "fossilis" represents an earl}- migration of lions to Hurope during the Middle Pleistocene, whereas the cave lions came to Eurasia only in the Late Pleistocene, during a next wave. The cave lion separated later from the common ancestor, because it is more similar to the recent lion. The inferable appearance of P. I. spelaea was in the Eemian interglacial (120 ky ago).

* * *

Acknowledgements — I give my sincere thanks to László KORDOS for his help, and for the possibility to study the vertebrate collecdon stored in the Hungarian Geological Institute. I am indebted to János SZABÓ for the opportunity to carry on my studies in the Geological and Palaeontological Department of the Hungarian Natural History Museum. Many thanks are due to my colleagues, who helped me in preparing this paper, especially to Mihály GASPARIK, Attila VÖRÖS and Erika GÁL. I am grateful to Gábor CSORBA and Barbara H E R Z I G for their kind help in the Mammal Collection of the Hungarian Natural History Museum, and the Naturhistorisches Museum, Vienna, respectively. Lutz Christian MAUL (Senckenberg Research Institute, Research Station of Quaternary Palaeontology, Weimar) critically revised the manuscript, his work is gready acknowledged.

References

ARGANT , A. (1998): Etude de l'exemplaire de Panthera spelaea ( G O L D -FUSS, 1810) (Mammalia, Carnivora, Felidae) du gisement Pleistocene moyen recent de la grotte d'Aze (Saône et Loire) — Revista Paleontologica, 7: 449-466.

BARVSHNIKOV, G. & BOF.SKORov, G. (2001): The Pleistocene cave lion, Panthera spelaea (Carnivora, Felidae) from Yakutia, Russia. — Cranium, 18(1): 9—24.

BONA, F. (2006): Systematic position of a complete lion-like cat skull from the Eemian ossiferous rubble near Zandobbio (Bergamo, North Italy). — Rirista Italiana di Paleontológia e Stratigrafia, 112(1): 157-166. "

FARMS, J. S. (1988): HENNIG. Hennig 86 Reference. Version 1.5 — Manuscript.

FlCCARELLI, G. & TORRE, D. (1968): Upper Villafranchian panthers of Tuscany. — Palaeontographica Italica, 64: 173-184.

FREUDENRF.RG, W. (1914): Die Säugetiere des älteren Quarters von Mttel-europa. — Geologischen, Paleontologischen Abhandlungen, 16: 455-670.

G A Á L , I . (1934): A Gerecse-hegység egyik legérdekesebb barlangcsoportja. (Th e most interesting cave-group in Gerecse Mountain.) — Globus, 5(9): 321-330.

GASPARIK, M. (1993): Late Pleistocene gastropods and vertebrate fauna from Tokod (NE Transdanubia, Hungary). —fragmenta Mineralogica et Palaeontologica, 16: 89-116.

GOLDFUSS, G . (1810): Die Umgebungen von Müggendorf. Ein Taschenbuch fúr Freunde der Natur und Alterthumskunde. — Erlangen, pp. 351.

GRADSTEIN, F., O G G , J. & SMITH, A. (2004): A Geological Time Scale. — Cambridge, University Press, pp. 589

GUZVICA, G . (1998): Panthera spelaea from North-Croatia. — Geológia Croatica, 51(1): 7-14.

HANKÓ, E. P. & K( iRSÖS, Z. (in press): Pleisztocén oroszlánok fog- és állkapocs-morfológiai jellegeinek kladisztikus analízise. (Cladistical analysis of mandible and teeth morphological characters of Pleistocene lions.) — Állattani Közlemények.

HEMMER, H. (1971): Zur Charakterisierung und stratigraphischen Bedeutung von Panthera gombasyoegensis (KRETZOI, 1938). — Neues Jahrbuch für Geologie und Paläontologie Abhandlungen, [Monatsheft], 1971(12): 701—711.

HEMMER, H. (1974): Untersuchungen zur Stammesgeschichte der Pantherkatzen (Pantherinae). Teil I I I . — Veröffentlichungen der Zoolo

gischen Staatssammlung München, 17: 167—280. IILMMER, H . (1981): Die Evolution der Pantherkatzen. Modell zur Über

prüfung der Brauchbarkeit der Hennigschen Prinzipen der phylogenetischen Systematik für wirbeltierpalaeontologische Studien. — Paläontologische Zeitschrift, 55(1): 109-116.

HEMMER, H. (2001): Die Feliden aus dem Ephillafranchium von Untermassfeld. — In: K A H L K E , R.-D. (Ed.). Das Pleistozän von Untermassfeld bei Meiningen (Thüringen). Teil 3. — Monographien des Römisch-Germanischen Zentralmuseums, 40(3): 699—782.

HEMMER, H. (2003): Pleistozäne Katzen Europas-Eine Übersicht. — Cranium, 20(2): 6—22.

HEMMER, H., K A H L K E , R.-D. & K E L L E R , T. (2003): Panthera onca gombasyoegensis (Kretzoi, 1938) aus den frühmittelpleistozänen Mosbach-Sanden (Wiesbaden, Hessen, Deutschland) - 1 'in Beitrag zur Kenntnis der Vanabilität und Verbreitungsgeschichte des Jaguars. — Neues Jahrbuch für Geologie und Paläontologie Abhandlungen, 229(1), 31—60.

HEMMER, H , K A H L K E , R. D. & V E K U A A. K. (2001): The Jaguar - Panthera onca gombasyoegensis (KRETZOI, 1938) (Carnivora: Felidae) in the late Lower Pleistocene of Akhalkalaki (South Georgia; Transcaucasia) and its evolutionary and ecological significance. — Geobios, 34(4): 475-486.

JÁNOSSY, D. (1952): Die aurignacien-Fauna der Höhle von Istállóskő. — Földtani Köylöny, 82(4-6) : 181 -203.

JÁNOSSY, D. (1955): Die Vogel- und Säugetierreste der spätpleistozänen Schichten der Höhle von Istállóskő. — Acta Archaeologica Academiae Sáentiarum Hungaricae, 5: 149—181.

JÁNOSSY, D. (1963): Die altpleistozäne Wirbeltierfauna von Kövesvárad bei Répáshuta (Bükk-Gebirge). — Annales historico-naturales Musei uationalis Hungária, 55: 109-141.

JÁNOSSY, D. (1969): Stratigraphische Auswertung der europäischen mittelpleistozänen Wirbeltierfauna. Teile I.—II. — Berichte der Deutschen Gesellschaft für Geologische Wissenschaften, [A], Geologie und Paläontologie, 14(4-5): 367-438, 573-643.

JÁNOSSY, D. (1971): Der erste Nachweis einer Kalt-Moustérien Verteb-ratenfauna in Ungarn (T°kod-Nagyberek, Kom. Komárom). — Vertebrata Hungarica, 12: 103—110.

JÁNOSSY, D. (1986): Pleistocene Vertebrate Faunas of Hungary. — Akadémiai Kiadó, Budapest, pp. 207.

[ÁNOSSY, D. (1990): Vertebrate fauna of Site I I . — In: K R E T Z O I M. & DOBOSI V. (Eds): I 'értessyőlős wan site and culture. — Akadémiai Kiadó, Budapest, pp. 187-230

JÁNOSSY, D. & VÖRÖS, I . (1985): Revision der rnittelpleistozänen Säuge -üerfauna des "Felsdaches" Szuhogy-Csorbakö (Nordungarn). — Annales historico-naturales Musei nationalis Hungária, 77: 69-95.

KADIC, O. & MOTTL, M. (1944): Die Höhlen des Nordwestlichen Bükkgebirges. — Höhlenforschung, 17(1): 1-1 11.

KORDOS, L. (1994): Revised Biosrratigraphy of the Early Man Site at Vértesszőlős, Hungary. — Courier Forschungs-Institut Seuckenbeig, 171: 225-236.

KORMOS , T. (1914): Az 1913-ban végzett ásatásaim eredményei. [Die Ergebnisse meiner in 1913 geführte Grabungen.] Sonderdruck. — Das Ungarische Königliche Geologische Institute. — A Magyar Királyi Földtani Intérêt 1913. éri jelentéséből: 498-540.

KoRSÖS, Z . (1999): Zoos^sytematikaigyakorlatok. [Zooystemaficai practices]. — Szent István University, Faculty of Veterinär)1 Science, Budapest, pp. 144.

K R E T Z O I , M. (1929): Feliden-Studien. — Budapest, pp. 22. K R E T Z O I , M. (1938): Die Raubtiere von Gombaszög nebst einer Über

sicht der Gesamtfauna. — Annales Musei Nationalis Hungária, Pars Mineralogica, Geologica, Palaeontologica, 31: 87-157.

K R E T Z O I , M. (1941a): Die Unterplcistozäne Säugetierfauna von Betfia bei Nagyvárad. — Földtani Közlöny, 71: 7-12; 308-335.

K R E T Z O I , M. (1941b): Betrachrungen über das Problem der Eiszeiten. (Ein Beitrag zur Gliederung des Jungtertiärs und Quarters). — Annales Musei Nationalis Hungária] Pars Mineralogica, Geologica, Palaeontologica, 34: 80.

MESZ« )ELY, A. M. C H . & GASPARIK, M. (2002): First record of an agamid lizard from the Pleistocene of Hungary. — Fragmenta Palaeontologica Hungarica, 20: 1-2.

PÉCSI, M. (1993): Nejedkor és lösykutás. [Quarter and loess-studies\. — Akadémiai kiadó, Budapest, pp. 376.

RABEDER , G., N A G E L , D. & PACHER, M. (2000): Der Höhlenbär. — Jan

Thorbecke Verlag, pp. 112. REICHENAU, W. (1906): Beitrage zur näheren Kenntniss der Carnivoren

aus den Sanden von Mauer und Mosbach. — Abhandlungen Grossheryoglichen Hessischen Geologischen ÍMndesanstalt, 4: 189—313.

R I E D E L , A. (1982): 11 grande felino della Crotta Tilde (The lion of the Tilde Cave). —Atti Museo Civico di Storia Naturale di Trieste, 34(1): 9—47.

RiNGLR, A. (2002): The new image of Szeleta and Istállós-kő caves in the Bükk Mountains: a revision project between 1999-2002. — Praehistoria, 3: 47—52.

SALA, B. (1990): Panthera leo fossilis (VON REICHENAU, 1906) (Felidae) de Isernia La Pineta (Pleistocene moyen Inférieur d'Italei). — Geobios, 23(2): 189-194.

SCHAUB, S. (1949): Révision de quelques carnassiers villafranchiens du Niveau des F4ouaires (Montagne de Perrier, Puy-de Dome). — FLc/ogae Geologicae Helvetiae, 42(2): 492-506.

SCHMID, E. (1940): Variationsstatistische Untersuchungen am Gebiss pleistozäner und rezenter Leoparden und anderer Feliden. — Zweitschrift für Säugetierkunde, 15(1): 1-178.

SCHUTT, G. (1969): Untersuchungen am Gebiss von Panthera leo fossilis (REICHENAL", 1906) und Panthera leo spelaea (GOLDFUSS, 1810). — Neues Jahrbuch für Geologie und Paläontologie Abhandlungen, 134(2): 192-220.

TASNÁDI-KUBACSKA , A. (1935): Die Mollusken- und Wirbeltier fauna des Plistozän [sie] und Ober-Pliozän von Gombaszög. — Anna/es Musei Nationalis Hungarici. Pars Mineralogica, Geologica, Palaeontologica, 29: 9-20.

THENIUS , E. (1971): Die Feliden (Carnivora) aus dem Pleistozän von Stranska Skala. — //;: Stranska Skala I . 1910-1945. — Anthropos, 20(12): 121-135.

TURNER, A. (1990): The evolution of the guild of larger terrestrial carnivores during the Plio-Pleistocene in Africa. — Geobios, 23(3): 349-368.

VAUGHN, T. A., RYAN,}. & CZAPLEWSKI, M. N.J. (2000): Mammalogy. — Sunders College Publishing, USA. pp. 565.

VERESCHCHAGIN , N . K. (1971): Peshchernyij lev i ego istoryja v golarkrike i v predelah CCCP. [The cave lions of Holarctis.] — Akademie Nauk S SSW, Trudy Zoo/ogicheshkogo Instituta, 49: 123—199.

VÉRTES, L. (1965): Ay őskőkor és a% átmeneti kőkor emlékei Magyamrsyágon. A Magyar Régészet Kéyikönyre I. [The relics of the Paleo- and Mesolithic ages in Hungary. Handbook of Hungarian Archaeology!] — Akadémiai Kiadó, Budapest, pp. 385.

VÖRÖS, I . (1984): Hunted mammals from the Aurignacian cave bear hunter's site in the Istállóskő cave. — Folia Archaeo/ogica, 35: 7—31.

VÖRÖS, I . (1988): Middle Pleistocene mammalian fauna from the Solymár Cave. — Folia Archaeologica, 39: 41—58.

ZBORAY, G. (2001): Összehasonlító anatómiai praktikum II [Practices in comparative anatomy II]. — Nemzeti Tankönvkiadó, Budapest, pp. 444.

Author's address: Eszter Piroska HANKÓ Department of Geology and Palaeontology Hungarian Natural History Museum Budapest VIII, Ludovika tér 2. Mail: 1431 Budapest, pf. 137

Hungary E-mail: osliroda@ nhmus.hu