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Fig. 7. Leopard mandibles from Equi: IGF6122V, IGF6120V, and IGF6103V in labial (A, B, and C) and lingual (A’, B’, and C’, respectively) views; IGF10037V/1 in lingual (D) and labia (D’) views; IGF10037V/2 in labial (E) and lingual (E’) views; IGF6102V in lingual (F) and labial (F’) views; IGF185V/4 in labial (G) and lingual (G’) views; IGF6104V in lingual (H) anc labial (H’) views.

Figure 7 Leopard mandibles from Equi: IGF6122V, IGF6120V, and IGF6103V in labial (A, B, and C) and lingual (A’, B’, and C’, respectively) views; IGF10037V/1 in lingual (D) and labia (D’) views; IGF10037V/2 in labial (E) and lingual (E’) views; IGF6102V in lingual (F) and labial (F’) views; IGF185V/4 in labial (G) and lingual (G’) views; IGF6104V in lingual (H) anc labial (H’) views.

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Fig. 1. The location of Equi in the Tuscany region (A). Equi is located on the northern side of the Apuane Alps, above the homonymous town (B) (photo by P. Nannini, Soprintendenza per i Beni Archeologici della Toscana). The valley of Fagli has a V-shape and the cave entrance is north-east oriented (C).
Fig. 1. The location of Equi in the Tuscany region (A). Equi is located on the northern side of the Apuane Alps, above the homonymous town (B) (photo by P. Nannini, Soprintendenza per i Beni Archeologici della Toscana). The valley of Fagli has a V-shape and the cave entrance is north-east oriented (C).
tefacts) induced a lengthy harsh scientific debate between the highest authorities in the field (Mochi, 1916, 1920, 1929; Rellini, 1917, 1924, 1929) which was resolved only fifteen years later, when an excavation field season by the Italian Institute of Human Palaeontology (Graziosi, 1934; Rellini, 1935) confirmed the absence of ceramics within Palaeolithic levels. These dates are also consistent with the biochronological interpretation and with the recent radiometric dating of specimens recently excavated from older layers (Bigagli et al., 2013). The Panthera pardus specimen (sample #360175) tentatively referred to layer HB seems to indicate an age inversion from a layer strati- graphcally superimposed to that of samples #370173 and #360176. The degree of uncertainty due to possible bias in the data available for the historical collections (samples were excavated in 1911-1917), do not allow us however to exclude the provenance of the specimen from an older layer. Since the De Stefani excavation times, the evidence of two distinct phases of human presence (ceramics and Palaeolithic ar- During our revision of the De Stefani collection of historical Equi excavations, three bone samples with registered stratigraphic provenance were sent to the Beta Analytic Inc. (Miami, Florida, Usa) for radiometric dating (Accelerator Mass Spectrometry, AMS, analysis). All the three remains contained sufficient collagen and data from these analyses are listed below:
tefacts) induced a lengthy harsh scientific debate between the highest authorities in the field (Mochi, 1916, 1920, 1929; Rellini, 1917, 1924, 1929) which was resolved only fifteen years later, when an excavation field season by the Italian Institute of Human Palaeontology (Graziosi, 1934; Rellini, 1935) confirmed the absence of ceramics within Palaeolithic levels. These dates are also consistent with the biochronological interpretation and with the recent radiometric dating of specimens recently excavated from older layers (Bigagli et al., 2013). The Panthera pardus specimen (sample #360175) tentatively referred to layer HB seems to indicate an age inversion from a layer strati- graphcally superimposed to that of samples #370173 and #360176. The degree of uncertainty due to possible bias in the data available for the historical collections (samples were excavated in 1911-1917), do not allow us however to exclude the provenance of the specimen from an older layer. Since the De Stefani excavation times, the evidence of two distinct phases of human presence (ceramics and Palaeolithic ar- During our revision of the De Stefani collection of historical Equi excavations, three bone samples with registered stratigraphic provenance were sent to the Beta Analytic Inc. (Miami, Florida, Usa) for radiometric dating (Accelerator Mass Spectrometry, AMS, analysis). All the three remains contained sufficient collagen and data from these analyses are listed below:
Fig. 2. Pie chart of leopard bones' age-stage (A), and age-stage frequencies for anatomical regions (B). Pie chart describing metapodial leopard remains from Equi (C). Gener. representation of leopard specimens from Equi (D).
Fig. 2. Pie chart of leopard bones' age-stage (A), and age-stage frequencies for anatomical regions (B). Pie chart describing metapodial leopard remains from Equi (C). Gener. representation of leopard specimens from Equi (D).
Fig. 3. Fossil of perinatal cub of leopard (IGF6120V) in lingual (A), labial (B) and ventral (C) views. D: putative larval holes on diaphysis surface of a young humerus (IGF6141V).
Fig. 3. Fossil of perinatal cub of leopard (IGF6120V) in lingual (A), labial (B) and ventral (C) views. D: putative larval holes on diaphysis surface of a young humerus (IGF6141V).
Fig. 4. Cranial leopard remains from Equi in top view: IGF15111V/1 (A); IGF15110V (B); IGF10036V (C); IGF185V/1 (D); IGF184V (E). IGF10036V, IGF185V/1, and IGF184V are also represented in lateral views (F, G, H, respectively).
Fig. 4. Cranial leopard remains from Equi in top view: IGF15111V/1 (A); IGF15110V (B); IGF10036V (C); IGF185V/1 (D); IGF184V (E). IGF10036V, IGF185V/1, and IGF184V are also represented in lateral views (F, G, H, respectively).
Fig. 5. IGF10036V and IGF15111V/1 in detail. IGF10036V in dorsal (A), ventral (B), caudal (C), left (D) and right (E) views. Cranium IGF15111V/1 in dorsal view (G) and with mandible IGF15111V/2 in left side view (F).
Fig. 5. IGF10036V and IGF15111V/1 in detail. IGF10036V in dorsal (A), ventral (B), caudal (C), left (D) and right (E) views. Cranium IGF15111V/1 in dorsal view (G) and with mandible IGF15111V/2 in left side view (F).
Morphological characters of the specimens of P. pardus from Equi compared to other Pleistocene and Holocene skulls. OC-MZF: osteological collection of the Natural History Museum of the University of Florence. two McV. The proximal epiphysis is deeply convex and the anterior limit ends with an acute angle. Laterally, it develops a bulge and forms a semilunar convex surface. The diaphysis has a rectangular transversal section. trochanter ventral outline is rounded and curves from the lateral to the medial side. IGF5341V (Fig. 8B) is from a very young animal and does not preserve the proximal epiphysis (un-ossified); the same specimen instead preserves the distal epiphysis showing the ossi- fication line at the contact with the diaphysis.
Morphological characters of the specimens of P. pardus from Equi compared to other Pleistocene and Holocene skulls. OC-MZF: osteological collection of the Natural History Museum of the University of Florence. two McV. The proximal epiphysis is deeply convex and the anterior limit ends with an acute angle. Laterally, it develops a bulge and forms a semilunar convex surface. The diaphysis has a rectangular transversal section. trochanter ventral outline is rounded and curves from the lateral to the medial side. IGF5341V (Fig. 8B) is from a very young animal and does not preserve the proximal epiphysis (un-ossified); the same specimen instead preserves the distal epiphysis showing the ossi- fication line at the contact with the diaphysis.
Fig. 6. Premaxillar IGF185V/2 with incisive serie in frontal (A) ventral (B) and dorsa (C) views.
Fig. 6. Premaxillar IGF185V/2 with incisive serie in frontal (A) ventral (B) and dorsa (C) views.
Fig. 11. Left McII (IGF5303V) in dorsal (A), proximal (C — not in scale), and medial (B) views. Fig. 10. Radious of leopards: IGF5337V is from a left paw of an adult animal (A) and IGF5340V is a left radious from a younger one (B).
Fig. 11. Left McII (IGF5303V) in dorsal (A), proximal (C — not in scale), and medial (B) views. Fig. 10. Radious of leopards: IGF5337V is from a left paw of an adult animal (A) and IGF5340V is a left radious from a younger one (B).
Fig. 8. A) right leopard humerus (IGF5342V) with evidence of pectoralis muscle stress on the proximal diaphysis; B) young right humerus of leopard (IGF5341V).
Fig. 8. A) right leopard humerus (IGF5342V) with evidence of pectoralis muscle stress on the proximal diaphysis; B) young right humerus of leopard (IGF5341V).
Fig. 9. Left ulna of leopard in lateral view: from an adult animal (IGF5355V) (A) and a cub (IGF5354V) without the proximal and distal epiphysis (B); specimen IGF5402V (C) is massive and probably from a male or old animal.
Fig. 9. Left ulna of leopard in lateral view: from an adult animal (IGF5355V) (A) and a cub (IGF5354V) without the proximal and distal epiphysis (B); specimen IGF5402V (C) is massive and probably from a male or old animal.
Fig. 13. Right McIV (IGF5293V) in dorsal (A), proximal (C— not in scale), and lateral (B views.
Fig. 13. Right McIV (IGF5293V) in dorsal (A), proximal (C— not in scale), and lateral (B views.
Fig. 12. Right McllI (IGF5302V) in dorsal (A), proximal (C— not in scale), and lateral (B views.
Fig. 12. Right McllI (IGF5302V) in dorsal (A), proximal (C— not in scale), and lateral (B views.
Fig. 15. Right femurs of leopards in frontal view: adult (IGF5347V) (A) and juvenile (B: IGF5350V; C: IGF5349V) specimens.
Fig. 15. Right femurs of leopards in frontal view: adult (IGF5347V) (A) and juvenile (B: IGF5350V; C: IGF5349V) specimens.
Fig. 14. Left McV (IGF5309V) in dorsal (A), proximal (C— not in scale), and medial (B) views.
Fig. 14. Left McV (IGF5309V) in dorsal (A), proximal (C— not in scale), and medial (B) views.
Fig. 17. Right MtlI (IGF5311V) in dorsal (A), proximal (C— not in scale), and lateral (B) views.
Fig. 17. Right MtlI (IGF5311V) in dorsal (A), proximal (C— not in scale), and lateral (B) views.
Fig. 16. Tibia of leopards in frontal view: A: right tibia of sub-adult (IGF5346V) B: left tibia from a young animal (IGF5344V).
Fig. 16. Tibia of leopards in frontal view: A: right tibia of sub-adult (IGF5346V) B: left tibia from a young animal (IGF5344V).
Fig. 19. Right MtIV (IGF5292V) in dorsal (A), proximal (C— not in scale), and lateral (B’ views. Dimensionally speaking, the crania from Equi fall within the range of extant leopards. They are generally massive, and all larger than extant subspecies for the palatal and neurocranium width (Fig. 23). Comparative material of jaguars (P. onca) and cougars (Puma concolor) are intermediate in size among P. pardus and lions, and within the leopard-lynx lower limit respectively. Leopards from Equi are quite larger than the conspecific extant animals. It is also notable that all leopard crania from Equi had a shorter muzzle
Fig. 19. Right MtIV (IGF5292V) in dorsal (A), proximal (C— not in scale), and lateral (B’ views. Dimensionally speaking, the crania from Equi fall within the range of extant leopards. They are generally massive, and all larger than extant subspecies for the palatal and neurocranium width (Fig. 23). Comparative material of jaguars (P. onca) and cougars (Puma concolor) are intermediate in size among P. pardus and lions, and within the leopard-lynx lower limit respectively. Leopards from Equi are quite larger than the conspecific extant animals. It is also notable that all leopard crania from Equi had a shorter muzzle
Fig. 18. Right Mtlll (IGF5313V) in dorsal (A), proximal (C— not in scale), and lateral (B views.
Fig. 18. Right Mtlll (IGF5313V) in dorsal (A), proximal (C— not in scale), and lateral (B views.
Fig. 21. Female and male comparative morphologies of leopard crania (from Pocock, 1930; modified): major differences are on temporal lines junctions, the development of the sagittal crest, the feature of the braincase, more or less evident postorbital constriction. Morphology is quite variable in leopards (Table 1) and the dimensional analysis must also be taken into account for a better grounded determination.
Fig. 21. Female and male comparative morphologies of leopard crania (from Pocock, 1930; modified): major differences are on temporal lines junctions, the development of the sagittal crest, the feature of the braincase, more or less evident postorbital constriction. Morphology is quite variable in leopards (Table 1) and the dimensional analysis must also be taken into account for a better grounded determination.
Fig. 20. Right MtV (IGF5318V) in medial (A), proximal (C— not in scale), and lateral (B) views. Distal end is broken by gnawing.
Fig. 20. Right MtV (IGF5318V) in medial (A), proximal (C— not in scale), and lateral (B) views. Distal end is broken by gnawing.
Fig. 22. Total length vs. condilus width scatter diagram of fossils and extant species: black dots are from extant animals from different regions (a single specimen has a declared gender); the gray dot is the Monte Sacro Middle Pleistocene cranium (M.P.U.R./V.1191: Kotsakis and Palombo, 1979); the gray-black dot is the cranium from Algar do Manga Larga (Cardoso and Regala, 2006); white dots are fossils crania described by Diedrich (2013); square, triangle and diamond represent the crania from Equi. Genders were defined on the basis of morphological characters.
Fig. 22. Total length vs. condilus width scatter diagram of fossils and extant species: black dots are from extant animals from different regions (a single specimen has a declared gender); the gray dot is the Monte Sacro Middle Pleistocene cranium (M.P.U.R./V.1191: Kotsakis and Palombo, 1979); the gray-black dot is the cranium from Algar do Manga Larga (Cardoso and Regala, 2006); white dots are fossils crania described by Diedrich (2013); square, triangle and diamond represent the crania from Equi. Genders were defined on the basis of morphological characters.
Fig. 23. Basion-prostion vs. palatal weight (A) and euryon—euryon (B). In comparison to extant leopards, the fossils from Equi are larger, stoutly built and massive, and clearly distinct from other species of felids.
Fig. 23. Basion-prostion vs. palatal weight (A) and euryon—euryon (B). In comparison to extant leopards, the fossils from Equi are larger, stoutly built and massive, and clearly distinct from other species of felids.
Fig. 24. Dental comparison among upper (A) and lower (B) teeth of leopards from Equi (black squares), extant (white circles) and extinct leopards (Arago — white squares, un- published data; Grotta del Capriolo — gray square, Pitti and Tozzi, 1971; Manga Larga — gray-black circle, Cardoso and Regala, 2006), extant and extinct lions (white triangles) and lynxes (white rhombus), and cougar (gray diamonds).
Fig. 24. Dental comparison among upper (A) and lower (B) teeth of leopards from Equi (black squares), extant (white circles) and extinct leopards (Arago — white squares, un- published data; Grotta del Capriolo — gray square, Pitti and Tozzi, 1971; Manga Larga — gray-black circle, Cardoso and Regala, 2006), extant and extinct lions (white triangles) and lynxes (white rhombus), and cougar (gray diamonds).
Fig. 25. Length and breadth of lower carnassials and respective trendlines of the main leopard collections, namely Equi, Los Rincones and Vraona Cave. The lines are quite parallel to each other and precisely specify for the shape of leopards m1s.
Fig. 25. Length and breadth of lower carnassials and respective trendlines of the main leopard collections, namely Equi, Los Rincones and Vraona Cave. The lines are quite parallel to each other and precisely specify for the shape of leopards m1s.
Fig. 26. Postcranial comparison among different species of Felidae. A: humerus (IGF5342V) B: radius (IGF5339V); total length (Gl) and distal breadth (Bd) ratios.
Fig. 26. Postcranial comparison among different species of Felidae. A: humerus (IGF5342V) B: radius (IGF5339V); total length (Gl) and distal breadth (Bd) ratios.
Fig. 28. Diastema length of different species of Felidae. Modern leopards are partially included to lion's variability but are well-divided to P. uncia. The same is generally for the leopard fossils with the exception of the short mandible of Triagalnata and the problematic Straska Skala specimen.
Fig. 28. Diastema length of different species of Felidae. Modern leopards are partially included to lion's variability but are well-divided to P. uncia. The same is generally for the leopard fossils with the exception of the short mandible of Triagalnata and the problematic Straska Skala specimen.
Fig. 27. Metacarpals (A) and metatarsal (B). Comparative dimensional range of Greatest length (Gl) in Holocene and Late Pleistocene big cats
Fig. 27. Metacarpals (A) and metatarsal (B). Comparative dimensional range of Greatest length (Gl) in Holocene and Late Pleistocene big cats
Fig. 29. Comparison between the types recognized by Spassov and Raychev (1997) and the p4-m1 from Equi. 1) robust type; 2) gracile type. Equi teeth: IGF185V/3 (A); mirrorec IGF6102V (B); IGF6103V (C); mirrored IGF10037V/1 (D); IGF10037V/2 (E). Black lines show the differences between the two types.
Fig. 29. Comparison between the types recognized by Spassov and Raychev (1997) and the p4-m1 from Equi. 1) robust type; 2) gracile type. Equi teeth: IGF185V/3 (A); mirrorec IGF6102V (B); IGF6103V (C); mirrored IGF10037V/1 (D); IGF10037V/2 (E). Black lines show the differences between the two types.
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PaleobiologyPaleontologyPalaeoenvironmentQuaternary GeologyVertebrate PalaeontologyVertebrate EvolutionVertebrate PaleontologyPaleolithic EuropeQuaternary palaeontologyQuaternary environments
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