Szabó János szerk.: Fragmenta Mineralogica Et Palaentologica 24-25. 2007. (Budapest, 2007)
Characters, distinguishing the Recent P. leo, from P. I. fossilis: - the protocone in P 4 can be totally absent; - degree of constriction in P 4 is greater than in "fossilis"; - margin of enamel on P 4 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 P 4 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 relationship 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 P 3 . However the separation from the Recent Panthera leo species is clearly demonstrated by the morphology of the Mi and P 4 . The characters, as the absence of thickening of the crown on the buccal side of P 4 , 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 (GOLDFUSS, 1810) — Istállóskő, Late Pleistocene, V 59.245, Mi; a: buccal view, b: occlusal surface; xl. 2 a-b Panthera leo Spelaea (GOLDFUSS, 1810) — Kiskevély, I .ate Pleistocene, Ob.2977, mandible; a: lingual view, b: view from above; x0.77. 3 a-b Panthera leo Spelaea (GOLDFUSS, 1810) — Igric, Late Pleistocene, V 60.1785 mandible, Mi, P 4 , P3, C lnl ; a: buccal view, b: Mi and P4 buccal view; x0.65.
