Panthera fossilis ( REICHENAU , 1906)
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publication ID |
https://doi.org/ 10.14446/AMNP.2014.59 |
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https://treatment.plazi.org/id/03988789-FFE7-FFA7-26D7-6EF42B6FFE8A |
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treatment provided by |
Felipe |
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scientific name |
Panthera fossilis ( REICHENAU , 1906) |
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Panthera fossilis ( REICHENAU, 1906)
Text-figs 2–4 View Text-fig View Text-fig
1906 Reichenau, W. von, Beiträge zur näheren Kenntnis der Carnivoren aus den Sanden von Mauer und Mosbach. Abh. Grossherzogl. Hess. Geol. Landesanst. 4, Darmstadt, p. 301.
2005 P. cf. mosbachensis (DIETRICH) – Ábelová: 172, 174–175, Pl. I/1-2.
2005 P. cf. spelaea (GOLDFUSS) – Ábelová: 172, 174–177.
L o c u s Ty p i c u s. Mauer, Germany (MIS 15 or MIS 13).
S t r a t i g r a p h i c a n d g e o g r a p h i c d i s t r i b u t i o n.
Middle Pleistocene (MIS 17 – MIS 9/6?), Europe and Asia?
S p e c i e s c h a r a c t e r i s t i c s. Large extinct representative of Panthera genus with less specialized lower teeth and less reduced lower premolars. The skull postorbital process is situated more anteriorly and bullae are relatively small and less protuberant. In comparison to P. spelaea , P. fossilis possesses smaller incisors; less flattened, but proportionally narrower canines; narrower P2; somewhat narrower P4 with shorter metastyle and smaller parastyle; proportionally shorter main cusps in P3, p3, and p4; less reduced anterior cusps in premolars; wider m1; and proportionally longer p4 from the viewpoint of the Lm1/Lp4 index. Its skull is slightly wider in the muzzle part and narrower in the postorbital and mastoid part, orbits are relatively smaller, nasals are relatively wider, and bullae are less inflated. Compared to P. leo , the species shows a greater overall size, shorter P4 metastyle, and smaller P2 (modified according to Kurtén 1965, Dietrich 1968, Schütt 1969, Schütt and Hemmer 1978, Argant 1988, Baryshnikov and Boeskorov 2001, Sotnikova and Nikolskyi 2006, Barycka 2008, and Marciszak et al. 2014).
M a t e r i a l. Vykopaná chodba (left side; age unknown, probably ≤ MIS 9) – thoracic vertebra, 2 fragments of radius, Mc III dext., metacarpal fragment, fragment of calcaneus, Mt III sin. fragment, Mt IV sin.? fragment;
Vykopaná chodba (Profile ZH P-3, layers 3 (MIS 9) and 4 (> MIS 9, cold phase) – left astragalus, 2 medial phalanges;
Chodba naděje (Profile ZH P-7b, layer 3; age unknown, probably MIS 9) – fragment of P4 sin.?, m1 dext. fragment, fragment of left ulna, 2 right and 2 left calcanei, Mt IV sin., proximal phalanx, medial phalanx;
Chodba naděje (Profile ZH P-7b, layers 3–4, depth 80–140 cm; age unknown, probably ~ MIS 9) – P4 dext., proximal phalanx;
Chodba naděje (Profile ZH P-7b, layer 4, depth 180–200 cm; age unknown, probably ≥ MIS 9) – right astragalus;
Chodba naděje (Profile ZH P-7b, layer 6; age unknown, probably> MIS 9) – m1 dext.;
Kostnice I (Profile ZH P-6, depth to 4 m; age unknown, probably ≥ MIS 9) – Mc IV sin.;
Kostnice II (Profile ZH P-5, layer 2b; MIS 9) – fragment of P4 sin.?, left astragalus;
Kostnice II (Profile ZH P-5, layers 2b-3b; MIS 9) – fragment of P4 dext.?;
Kostnice II (Profile ZH P-5, brown loam complex; probably MIS 9) – 2 fragments of right mandible with p4 and m1, Mt IV sin.;
Narozeninová chodba (Profile ZH P-9, layer 5;> MIS 9) – C dext., fragment of left mandible with m1 alveolus, 5 caudal vertebrae, fragment of right scapula, fragment of right humerus, fragment of left humerus, right capitatum, left capitatum, Mc I dext., left patella, fragment of right fibula, fragment of left fibula, right calcaneus, Mt IV dext., fragment of metapodial distal epiphysis, 5 proximal phalanges, 4 medial phalanges, fragments of 2 distal phalanges, sesamoid bone;
Non-stratified finds (exception) – fragment of left mandible with p4 and m1.
D e s c r i p t i o n. The right upper canine (C dext.) belonged to an adult specimen ( Text-fig. 2a View Text-fig ), according to measurements of the preserved crown part and root probably to a male ( Tab. 1). A slightly worn distinct lingual as well as serrated aboral crest is present on the crown. There are also distinguishable grooves on the flattened (lesser than in cave lions) lateral crown sides. In spite of this, the basal crown shape is still oval in cross-section. No basal cingulum is developed.
The crown of the isolated right upper carnassial (P4 dext.) is partly covered by a very thin surface sinter on the protocone lobe ( Text-fig. 2b View Text-fig ). The dominant robust paracone possesses distinct anterior and posterior crests as well as a distinctly rounded blunt crest on the anterolingual side, running from the cusp tip probably as far as the protocone lobe. In front of the paracone, the crest-shaped parastyle with rounded anteroposterior cutting edge is separated by a notch. The cingular preparastyle (a typical character for P. fossilis ) is absent, although the anterior cingular bulge is cuspletshaped (i.e. with the indication of tiny accessory cusps). The slightly damaged metacone is smaller than the parastyle, separated from the paracone by a prominent notch, continuing on the buccal side into a shallow, buccally opening depression with a flat bottom. The metacone continues in a reverse direction via its crest-shaped cutting edge into a rounded metastyle at the posterior crown margin. The metacone – metastyle length is proportionally longer than in the upper carnassials of cave lions ( Tab. 1). The cingulum is moderately developed only on the anterobuccal crown margin and indicated on the posterolingual side. From three roots, the posterior root is the most robust and the lingual root under the protocone lobe is the smallest. The rest of the isolated upper fourth premolars, referred to from the site by Ábelová (2005) as 2 P4 sin. and 1 P4 dext., are preserved only as fragments according to the original description. During revision, the presence of these teeth within the fossil record was not validated and also an illustration of one of these specimens ( Ábelová 2005: tab. I.1) indicates a rather different tooth type than P4 (P3?).
The anterior cusp (paraconid) of a double-rooted p4, still situated in a left mandible fragment, is smaller than the posterior one (hypoconid). It possesses a distinctly developed posterior crest and the cusp is separated from the main one (protoconid) by a notch. The main cusp is moderately sloping posteriorly, with a distinctly longer anterior as well as posterior crest. It is more flattened on the buccal side and triangular on the lingual side with a blunt rounded crest-shaped margin. The posterior cusp is also separated from the main cusp by a notch. The cusp possesses a distinct longitudinal crest and it is separated posteriorly by a weak notch from the basal cingulum. This is developed mainly on the posterolingual crown margin, where it is bordering the talonid area. Both notches, separating the main cusp from the paraconid and the hypoconid, pass into shallow valleys on the lingual side. Similar morphology can also be observed in somewhat smaller p4s in two right mandible fragments ( Tab. 1), differing more or less only in the similar size of the anterior and the posterior cusp, or the slightly smaller hypoconid respectively (see Ábelová 2005: tab. I.2).
The paraconid of an isolated right lower carnassial (m1 dext.; Text-fig. 2c View Text-fig ) is smaller, but more robust than the protoconid, whereas it is optically lower than the protoconid of m1 sin. from the mandible fragment, similarly in a further two m1s from right mandible fragments. The paraconid anterior margin is either rounded (isolated m1 dext.) or oblique and straight, rounding only near the crown base (m1 dext. et sin. in situ). The paraconid cutting edge is shorter than the protoconid one. The paraconid is separated from the protoconid by a distinct, relatively deep notch that passes into a wide shallow lingual valley without an accessory cusp at the bottom, or with only an indication of it near the crown base respectively (isolated m1 dext.). The protoconid possesses distinguishable anterior and posterior margins. The posterior one is nearly vertical, moderately deviated medially at the base (isolated m1 dext.) or it is sharp, passing into the basal cingulum (or cingular talonid resp.) and bordered by a groove on the inner side (m1 sin. in situ). In the posterior part of the cingular talonid, a tiny cingular cusp may be developed. The cingulum is present only in the posterior crown part or also on the buccal side (isolated m1 dext.). In the occlusal view, the crown is convexly curved – it is halfmoon-shaped with a distinct cutting edge which forms a relatively wide-open V in m1 sin. On the lingual side of all m1s, a typical bulge can be observed ( Text-fig. 2e View Text-fig ). A zigzag enamel structure is also present, although this character is variable and its classification value (cf. Schütt and Hemmer 1978 or Barycka 2008) is questionable. Both roots of the isolated right lower carnassial are entirely preserved and the anterior one is more robust as well as larger than the posterior root. A fragment of another isolated m1 dext. with preserved paraconid and protoconid parts is also known from the site ( Ábelová 2005) ( Tab. 1).
Mandibles are represented within the fossil record under study only as an unmeasurable left mandible fragment with m1 alveoli and a left mandible fragment with p4 and m1 (described above) and with the alveoli of a canine and two-rooted p3. Both fragments probably belonged to adult males ( Text-fig. 2d View Text-fig ; Tab. 2). Another two fragments of right mandibles with p4 and m1 are figured in the paper by Ábelová (2005: tab. I.2A-B), although only the mandible fragment from tab. I.2B is also briefly described there. The chodba, left side, ≤ MIS 9), lateral view; b – fragment of left ulna (Chodba naděje, layer 3, MIS 9), lateral view; c – fragment of right humerus (Narozeninová chodba, layer 5,> MIS 9), caudal view; d – right capitatum (Narozeninová chodba, layer 5,> MIS 9), lateral view; e – fragment of right Mc III (Vykopaná chodba, left side, ≤ MIS 9), dorsal view.
mandible ramus of the adult male ( Text-fig. 2d View Text-fig ) is relatively high, straight to slightly convex at its base, with three foramina mentalia, the largest of which is situated under the anterior alveolus of p3 (the remaining two foramens are situated under the diastema between the canine and p3). The preserved anterior part of the masseteric fossa is relatively deep in all mandible finds, extending to under the m1 protoconid. The diameter of the p3 anterior alveolus in the adult male mandible fragment is 11.47 mm x 8.72 mm and the diameter of the p3 posterior alveolus in the same fragment is 11.52 mm x 11.65 mm.
Based on the so far studied lion-like fossils from Za Hájovnou Cave, the axial skeleton is represented by only a fragment of thoracic vertebra and five caudal vertebrae from minimally two adults. Interestingly, the body of the thoracic vertebra fragment shows some biting marks of a larger predator/scavenger (Text-fig. 3a), probably a hyena ( Crocuta crocuta ssp.) or wolf ( Canis lupus ssp.), fossils of which were also found at the site ( Ábelová 2005). It can not be excluded that this bone damage could have been caused by another carnivore, such as a bear or specimen of the same species (interspecific or intraspecific conflict?).
C dext. P4 dext. p4 dext. p4 sin. m1 dext. m1 dext. m1 sin.
Narozeni- Chodba Kostnice II non- Chodba Kostnice II non- Panthera fossilis, Za Hájovnou Cave nová chodba naděje brown stratified naděje brown stratified layer 5 layers 3–4 loam com. finds layer 6 loam com. finds
> MIS 9 ~ MIS 9 MIS 9> MIS 9 MIS 9
total tooth length – 41.70 31.00 32.86 32.45 29.61 34.33 longitudinal diameter of crown base 31.10 – – – – – – transversal diameter of crown base 22.80 – – – – – – longitudinal diameter of crown base in %
73.31 – – – – – –
of transversal diameter of crown base paracone length – 15.90 – – – – – paracone length _ 100 / total tooth length – 38.13 – – – – – metacone – metastyle length – 16.90 – – – – – metacone – metastyle length 100 / total tooth length – 40.53 – – – – – metacone – metastyle length 100 / paracone length – 106.29 – – – – – paracone – metastyle length – 30.50 – – – – – paracone – metastyle length 100 / total tooth length – 73.14 – – – – – paraconid length – – – 5.61 16.50 – 16.99 protoconid length – – – 15.53 15.50 – 17.73 hypoconid length – – – 6.81 – – – maximum tooth width – – 15.92 16.90 17.20 16.80 17.23 maximum tooth width 100 / total tooth length – – 51.36 51.43 53.01 56.74 50.19 maximum anterior tooth width – – – 14.52 – – – maximum anterior tooth width 100 / total tooth length – – – 44.19 – – – maximum tooth width in the middle – – – 16.47 – – – maximum posterior tooth width – 13.70 – 15.65 – – – maximum posterior tooth width 100 / total tooth length – 32.85 – 47.63 – – – paracone height / paraconid height – 23.00 – 15.01 18.40 – 19.82 protoconid height – – 17.00 18.24 16.30 16.69 19.68 hypoconid height – – – 17.24 – – – lower height of the notch in m1 – – – – 10.50 – 11.91 upper height of the notch in m1 – – – – 13.40 – 15.46
(= crown height in the middle)
length of the paraconid cutting edge – – – – 11.00 – 10.30 length of the protoconid cutting edge – – – – 13.80 – 15.01 length between tips of the paraconid and the protoconid – – – – 21.50 – 20.54 maximum longitudinal diameter of the root 34.30 – – – – – – maximum transversal diameter of the root 24.40 – – – – – –
max. longitudinal diameter of the root in %
71.14 – – – – – –
of max. longitudinal diameter mandibula sin.
non-stratified finds depth ramus at diastema 57.98
depth ramus at p3 anterior margin 57.42
depth ramus at p4 anterior margin 56.53
depth ramus at m1 anterior margin 62.65
depth ramus at m1 posterior margin 60.49 p3–m1 length (alveolar) 87.16
p4 length 32.86
m1 length 34.33
The preserved scapula fragment of an adult is relatively damaged, e.g. in the middle characteristically cut out glenoid cavity with a transverse dimension (the height) of 55 mm.
The forelimb stylopodium is represented within the fossil record under study by only a fragment of right humerus distal part (Text-fig. 3c; Tab. 3) with the possible marks of biotic agent activity as well as a fragment of left humerus distal epiphysis. Both stylopodial fragments belonged to adult animals. The olecranon fossa of the right humerus is lengthwise, relatively small, but deep and sharply bordered at its base.
From the forelimb zeugopodial region, only a left ulna fragment and fragments of the proximal part of a right and left radius were found within the stratified collecting areas until 2007. The left ulna fragment of an adult specimen (Text-fig. 3b; Tab. 3) consists of the proximal part, mostly covered by a thin sinter layer. A preserved irregular to stepped fracture indicates bone disintegration “after its drying” during the fossilization process, whereas tiny gnawing marks indicate the activity of lesser carnivores (mustelids?) after the animal´s death. The oval-shaped heads of both preserved proximal parts of adult radii (caput radii) are slightly narrowed posteriorly and without a sagittal crest. A tuberositas radii on the posterolateral side is less distinguishable than the distinct oval to longitudinal one on humerus radius os carpale os carpale Mc III
ulna sin. radius sin. Mc I dext.
dext. dext. III dext. III sin. dext.
Narozeni- Chodba Narozeninová chodba Narozeni- Vykopaná Panthera fossilis, Za Hájovnou Cave Vykopaná chodba
nová ch. naděje nová ch. chodba layer 5 layer 3 left side layer 5 layer 5 left side
> MIS 9 MIS 9? MIS 9> MIS 9> MIS 9? MIS 9 height of the sigmoid cavity – 47.4 – – – – – – maximum width of the sigmoid cavity – 49.0 – – – – – – maximum anteroposterior diameter
– 68.4 50.1 50.5 – – – –
of the olecranon maximum transversal diameter of the olecranon – 34.2 – – – – – – maximum anteroposterior diameter
– – – – – – – 38.7
of the proximal epiphysis maximum transversal diameter
– – – – – – – 35.2
of the proximal epiphysis height of the anterior apophysis of the olecranon – 69.5 – – – – – – anteroposterior diameter of the neck – – 34.9 35.0 – – – – anteroposterior diameter of the proximal
– – 39.4 47.3 – – – – articular surface anteroposterior diameter of the diaphysis – – 41.8 – – – 17.6 19.1 transversal diameter of the diaphysis – – – – – – 21.5 21.0 transversal diameter of the diaphysis (distal) 57.8 – – – – – – – anteroposterior diameter of the diaphysis (distal) 44.4 – – – – – – – upper transversal diameter of the trochlea 69.2 – – – – – – – lower transversal diameter of the trochlea 75.2 – – – – – – – maximum height of the trochlea 43.7 – – – – – – – maximum width of the olecranon fossa 44.8 – – – – – – – anteroposterior diameter of the distal epiphysis
34.9 – – – – – – –
at the place of the trochlea maximum anteroposterior diameter
– – – – – – 21.4 –
of the distal epiphysis maximum transversal diameter
– – – – – – 22.5 –
of the distal epiphysis maximum transversal diameter – – – – 29.0 28.6 – – maximum anteroposterior diameter – – – – 30.2 33.3 – – maximum vertical diameter – – – – 48.6 46.6 – –
the medial (inner) side. The inner surface of the diaphysis is flat to convex, bordered posteriorly by a distinctly rough crest and by a smaller oblique one in the middle. The diaphyseal lateral surface is moderately concave and the diaphysis is enlarged in a downwards direction.
The right and left capitatum of adult individuals (Text-fig. 3d; Tab. 3) are preserved almost undamaged, without pathological characters and postmortal traces of taphonomic agents.
Metacarpals are represented within the fossil record under study only by a fragment of Mc I dext. distal part and Mc III dext. proximal region (Text-fig. 3e) both from adult animals ( Tab. 3) as well as by a fragment of undetermined metacarpal bone. Ábelová (2005) also mentioned a find of Mc IV sin. distal epiphysis from Middle Pleistocene deposits from this site.
The hindlimb skeleton is represented within the studied sample of fossil remains by a left patella, right and left fibula, as well as by tarsal and metatarsal bones ( Text-fig. 4 View Text-fig ).
The damaged left patella with maximum longitudinal diameter 32.5 mm belonged to an adult, similarly the distal part of a left fibula with preserved distal epiphysis with maximum transversal diameter 35.6 mm and with diaphyseal part, showing a columnar fracture. The unmeasurable fragment of the left fibula consists only of its distal epiphysis.
From tarsal bones, only a right calcaneus and right astragalus have been studied. The damaged right calcaneus from Narozeninová chodba belonged to a smaller adult (female?) ( Tab. 4). From a morphological viewpoint, it shows a greater similarity with the calcaneus of P. spelaea than with that of P. leo . Its anterior and central astragal facets (facies articularis talaris anterior et media) are separated, or are connected only by a narrow crest-shaped element respectively. These astragal facets are frequently separated in calcaneal bones of cave lion, whereas those of modern lion are connected by a distinct longitudinal facet. Ábelová (2005) also mentioned a record of another two right and two left calcaneal bones of adult specimens from layer 3 of the Chodba naděje. She identified these bones with calcaneal bones of P. spelaea from a morphological viewpoint, but attributed them to more robust specimens rather than lion-like felids from the Last Glacial on the basis of their measurements.
The right astragalus of an adult specimen from the Chodba naděje ( Tab. 4) shows marks of taphonomic agent activity (gnawing?) on the bone head. A further two left astragali were found in the Kostnice II (Profile ZH P-5, layer 2b) and the Vykopaná chodba (Profile ZH P-3, layers 4) ( Ábelová 2005).
Only a fragment of the adult left Mt III proximal part with irregular transversal fracture is preserved within the studied sample ( Tab. 4). On the contrary, Mt IV is represented by three specimens (one right and two left) and probably also by a fragment ( Tab. 4). The adult Mt IV dext. is damaged on the surface. Its distal part is broken off (V-shaped fracture, see below). Mt IV sin. from layer 3 of the Chodba naděje, covered by a sinter layer, belonged probably to a relatively large individual (the total bone length is approximatly 151 mm). Also Mt IV sin. from the brown loam complex of the Kostnice II is identically robust. This metatarsal bone is similar to Mt IV of P. spelaea from the morphological viewpoint, but its measurements exceed those of cave lion ( Ábelová 2005). A Mt IV sin. (?) fragment consists of only the diaphysis and the distal part.
Phalanges represent the most frequent element of lion-like fossils within the sample under study. There are 7 proximal (one of these belongs to a first left digit, Text-fig. 4f View Text-fig ), 7 medial (with basal exostosis in one case), and 2 distal phalanges, which are preserved only fragmentarily. All phalanges belonged to adults. Gnawing marks of a rodent are distinguishable on one proximal phalanx ( Text-fig. 4g View Text-fig ).
From other lion-like bones, a nearly undamaged adult sesamoid bone, a calcaneus fragment as well as a fragment of the metapodial distal epiphysis, probably belonging to the above described Mt IV dext., were determined.
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