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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
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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).
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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.
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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
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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
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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.
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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.
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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
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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.
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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