First finding of a partially articulated elephant skeleton from a Late Pleistocene hyena den in Sicily (San Teodoro Cave, North Eastern Sicily, Italy)

First finding of a partially articulated elephant skeleton from a Late Pleistocene hyena den in Sicily (San Teodoro Cave, North Eastern Sicily, Italy)

Quaternary International 276-277 (2012) 53e60 Contents lists available at SciVerse ScienceDirect Quaternary International journal homepage: www.else...

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Quaternary International 276-277 (2012) 53e60

Contents lists available at SciVerse ScienceDirect

Quaternary International journal homepage: www.elsevier.com/locate/quaint

First finding of a partially articulated elephant skeleton from a Late Pleistocene hyena den in Sicily (San Teodoro Cave, North Eastern Sicily, Italy) Gabriella Mangano*, Laura Bonfiglio Department of Earth Sciences, University of Messina, Viale Ferdinando Stagno d’Alcontres 31, I-98166, Messina-S. Agata, Italy

a r t i c l e i n f o

a b s t r a c t

Article history: Available online 27 August 2011

Associated and partially articulated fossil remains of the endemic reduced size elephant Palaeoloxodon mnaidriensis have been recently discovered at San Teodoro Cave, a large Late Pleistocene hyena den in Sicily. The skeletal elements belong to both cranial and post-cranial portions, and are represented by a semi-complete mandible, a tusk fragment, a cervical vertebra, three thoracic vertebrae, a rib fragment, a scapula, a distal epiphysis of radius, a pyramidal bone, a III metacarpal bone, a coxal bone fragment, a femur shaft, two symmetrical pairs of tibias and fibulas, a patella and an astragalus. The bones were recovered mixed with remains belonging to other taxa, numerous hyena coprolites and with a juvenile elephant mandible in the perinatal stage. On the basis of anatomical representation, spatial distribution and development stages, the partially articulated elephant bones from the San Teodoro Cave belong to a single adult individual, perhaps a female, which probably entered the cave before its death, and was afterwards scavenged and disarticulated by hyenas. This is the first finding of a partially articulated elephant skeleton from a fossil hyena den, and may attest to the intermittent use of the cave by hyenas and elephants. Ó 2011 Elsevier Ltd and INQUA. All rights reserved.

1. Introduction A rich Pleistocene vertebrate fossil assemblage has been highlighted during recent excavations in the San Teodoro Cave at Acquedolci (Messina, Sicily, Italy) (Bonfiglio et al., 2001). The San Teodoro Cave deposits have been known since the second half of the 19th century, mainly for the important discovery of Upper Palaeolithic human burials (Anca, 1860; Vaufrey, 1929; Maviglia, 1941; Graziosi, 1943, 1947; Graziosi and Maviglia, 1946; Vigliardi, 1968). At the San Teodoro Cave, the Upper Late Glacial sedimentary unit (unit A; in Bonfiglio et al., 2001) containing humans’ feeding remains (mammal bones) associated with Late Upper Palaeolithic (Epigravettian) stone artefacts, overlies a lower sedimentary unit (unit B; in Bonfiglio et al., 2001) containing Late Pleistocene endemic mammal remains associated with remains of non-endemic mammals, as well as unequivocal evidence of cave frequentation by spotted hyena populations, from which the cave has been identified as a Pleistocene hyena den (Bonfiglio et al., 1999, 2001, 2008; Mangano et al., 2005; Mangano and Bonfiglio, 2005a, b). The continuous and intense human frequentation of * Corresponding author. E-mail addresses: [email protected] (G. Mangano), lbonfi[email protected] (L. Bonfiglio). 1040-6182/$ e see front matter Ó 2011 Elsevier Ltd and INQUA. All rights reserved. doi:10.1016/j.quaint.2011.08.034

the cave until the present time almost totally destroyed the Epigravettian anthropic deposit and partially removed the uppermost portions of the older Late Pleistocene deposit. Charcoal accumulation from Palaeolithic levels, as well as from recent surface deposits, often directly overlies the unit B deposit. Recent systematic excavations have been carried out from 1998 to 2006, and two trenches have been excavated: the a trench, located close to the cave entrance, and the b trench, located further inside the cave (Fig. 1a). As only scattered traces of the anthropic Epigravettian soil (unit A) were preserved (Bonfiglio et al., 2006), the trenches have been mainly excavated in the older deposits (unit B). A third sterile sedimentary unit (unit C), probably older than unit B, has been unearthed in the south-eastern sector of the b trench (Mangano and Bonfiglio, 2005a). During recent excavations, a total volume of approximately 25 m3 has been excavated from the two trenches, and about 5000 fossil bone remains and 10000 hyena coprolites have been recovered. No evidence of human frequentation has been detected from the unit B deposits. The rich fossil material recovered is a wonderful example of an exclusively hyena-collected fossil assemblage and the results of the taphonomic analysis are going to be published. This paper deals with the discovery of partially articulated elephant bones from the b trench of the San Teodoro Cave, which represents the first finding of an elephant skeleton from a fossil hyena den.

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Fig. 1. a) Plan of the San Teodoro Cave with location of the a trench (squares E-I/9-13) and b trench (squares A-G/29-34). The arrow indicates the entrance to the cave. b), c) Plan of the elephant bones grouping from b trench. Dark grey: elephant bone remains. Light grey: other taxa bone remains. Black circles: coprolites.

2. Sediments, faunal assemblage and dating of the San Teodoro Cave deposits The San Teodoro Cave has very large dimensions (about 60 m long, 20 m wide, up to 20 m high) and occupies a total area of about 1000 m2. It is composed of a single large chamber and one very small lateral chamber. Some vertical conduits are present in the cave ceiling, but their dimensions are too small to act as a morphological trap for large mammals. The largest non-carbonate detrital elements found in the cave, coming from the sedimentary cover of a terrace located above the cave also containing very large boulders, have dimensions smaller than 4e5 cm.

The floor of the cave ascends along its major axis for about 15 m from the entrance to the southern end of the cave. The central part of the floor is a detrital fan which slopes down laterally towards the eastern and western walls of the cave. The detrital fan is made up of fine gravels, sands and silt incorporating very large carbonate boulders. The b trench is located on the eastern edge of the detrital fan. Sediments from both trenches are made up of non-carbonate fine gravels, sands and silt, including carbonate blocks of different sizes which have fallen from the cave ceiling. Vertebrate remains are scattered within all excavated levels. Skeletal remains of large mammals are fragmented and not articulated. The numerous and diverse evidence of cave frequentation

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by spotted hyena populations, represented by the occurrence of several Crocuta crocuta spelaea skeletal elements, an impressive quantity of coprolites and ubiquitous traces of crushing, gnawing, chewing and digestion occurring on almost all large mammal remains, are the most prominent taphonomic feature of the deposit. Hyena-damaged bones and hyena coprolites have also been retrieved in several small pits excavated at different points along the cave floor, including the central fan, thus indicating that Crocuta crocuta spelaea extensively inhabited the cave. The mollusc fauna is represented by poor to quite rich species assemblages of land and freshwater gastropods and bivalves with MediterraneanEuropean character (Esu et al., 2007). Pollen analysis from coprolite samples coming from the a trench depicts a glacial landscape. Lower percentages of pollen of mesophilous taxa suggest the existence of nearby refugia of temperate and Mediterranean vegetation (Yll et al., 2006). The large mammal assemblage from the unit B deposit of the San Teodoro Cave includes Palaeoloxodon mnaidriensis, Bos primigenius siciliae, Bison priscus siciliae, Cervus elaphus siciliae, Sus scrofa, Equus hydruntinus, Crocuta crocuta spelaea, and very few remains of Canis lupus and Vulpes vulpes, while small mammals are represented by Microtus (Terricola) ex gr. Savii, Apodemus cf. sylvaticus, Erinaceus cf. europaeus, Crocidura cf. sicula and Chiroptera. These taxa represent the type of assemblage of a Pleistocene Faunal Complex of Sicily, named the “Grotta di San Teodoro e Pianetti” F. C. (Bonfiglio et al., 2001). A 230Th/234U dating on a concretion intercalated with two clayey levels within the b trench (Fig. 2) yielded a date of 32000  4000 a (Bonfiglio et al., 2008), so discrediting a previous AAR dating of 455000  90000 a by Bada et al. (1991), based on an elephant molar from the “Anca” collection stored at the “G.G. Gemmellaro” Museum (Palermo). According to Herridge (2010), this AAR date is recalculated and the amended AAR dating is 370000  74000 a. AAR dates associated with Mediterranean fauna should be treated with caution: the AAR method is based on a chemical reaction as the racemisation rate is a function of temperature as well as of time, and the depositional environment (e.g. pH, water circulation, temperature) can impact on amino acid diagenesis and leaching, violating closed system assumptions (Walker, 2005; Clarke and Murray-Wallace, 2006). On the basis of recently collected data, the 0.21 alle/ile value by Bada et al. (1991) for the San Teodoro Cave sample may be attributed to a possible heating of the dated material, whose precise stratigraphic provenance is unknown, but probably comes from a superficial level, heated by old or recent fireplaces. However, the 230 Th/234U geochronometric date, which will be tested by further dating on other carbonate concretions, is consistent with the faunal association collected from the unit B deposit of the San Teodoro

Fig. 2. Schematic section of the sedimentary sequence of the unit B deposit out cropping in the b trench at the eastern edge of the detrital fan. R: recent. NCUB: nonconcretioned unit B. CUB: concretioned unit B. CLUB: clayey unit B. CL: radiometrically dated concretion. Grey: elephant bone remains (after Bonfiglio et al., 2008, modified).

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Cave. Bone samples from unit B could not be radiocarbon-dated due to the lack of collagen (Bonfiglio et al., 2008). 3. Elephant bones from the unit B deposit of the San Teodoro Cave The existence of dwarf elephants in the Mediterranean islands was recognised for the first time during the early 19th century, after excavations at the S. Ciro Cave (Palermo) by Scinà (1830). The Sicilian elephants have been attributed to different species (see, among others, Anca and Gemmellaro, 1867; Pohlig, 1888e1892, 1893; Vaufrey, 1929; Aguirre, 1968e69; Ambrosetti, 1968). Since the late 19th century, a flurry of palaeontological explorations in Malta and Sicily had resulted in the description of three species of dwarf elephants: ‘Elephas’ melitensis Falconer, in Busk (1867), ‘Elephas’ falconeri (Busk, 1867) and ‘Elephas’ mnaidriensis (Adams, 1874). According to Vaufrey (1929), the remains of E. melitensis from the Luparello cave (Palermo) underlie the remains of E. falconeri. Vaufrey drew the conclusion that the three species of elephants of different sizes, E. mnaidriensis, E. melitensis and E. falconeri, represent three successive phases of size reduction. According to the literature, Pleistocene vertebrate faunas of Sicily and Malta are closely linked, but conspecificity of taxa has been assumed without recourse to detailed taxonomic verification. The synonymy of dwarf elephants, or other endemic fossil taxa, on Sicily and Malta has never been systematically verified (Herridge, 2010). Researchers, however, have tended to follow Vaufrey (1929) and Osborn (1942) in attributing each taxon from Sicily and Malta to a different size class: ‘E.’ mnaidriensis (considered a large-sized dwarf), ‘E.’ melitensis (considered a medium-sized dwarf) and ‘E. falconeri’ (considered the smallest dwarf taxon). ‘E. melitensis’ has fallen out of use after Ambrosetti (1968) recognised E. melitensis from the Spinagallo cave (Siracusa, Sicily) as the masculine morphotype of E. falconeri. The biochronological scheme of Sicily actually recognises five Pleistocene Faunal Complexes (Bonfiglio et al., 2001): Mount Pellegrino F.C., E. falconeri F.C., E. mnaidriensis F.C., S.Teodoro Cave e Pianetti F.C., Castello F. C. (from the oldest to the youngest). The oldest and the youngest Faunal Complexes (Mount Pellegrino and Castello) do not contain elephant remains. The current attribution of Sicilian and Maltese dwarf elephant taxa is to the genus Palaeoloxodon, and Palaeoloxodon antiquus has been thought to be the probable ancestor of the dwarf elephants of Sicily and Malta (Ferretti, 2008). There are no species described from Sicilian material; material from the Spinagallo and Luparello Caves is currently referred to Palaeoloxodon falconeri, while material from the Puntali, Za Minica and San Teodoro Caves is referred to Palaeoloxodon mnaidriensis (Ambrosetti, 1968; Bonfiglio et al., 2008). The only available morphological and biometric data are provided by Ambrosetti (1968) for P. falconeri (the pygmy elephant, about 1.10 m tall) and by Bonfiglio and Berdar (1980) and Ferretti (1998) for P. mnaidriensis (the little reduced-sized elephant, about 2 m tall). Herridge (2010) provides the first pan-Mediterranean study that incorporates taxonomic and allometric approaches to the evolution of dwarf elephants. At the species level, named taxa were shown to be valid by Herridge (2010), but the taxonomic integrity of the P. mnaidriensis hypodigm was rejected and a new species of ‘largesized’ dwarf elephant on Sicily and Malta was identified. As a consequence, Herridge (2010) groups the dwarf elephant species of Sicily and Malta into three broad size-classes: ‘smallsized’ (P. falconeri), ‘medium-sized’ (P. mnaidriensis) and ‘largesized’ (Palaeoloxodon sp. nov.). Elephant remains from the Puntali Cave, San Teodoro Cave and Za Minica Cave are referred by Herridge (2010) to Palaeoloxodon sp. nov., that cannot be considered part of

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the same dwarfing lineage (i.e. is not ancestral) to the Sicilian P. falconeri. At the San Teodoro Cave, 74 elephant remains have been collected from the two excavated trenches. The remains come from all excavated levels of the unit B deposits, including those underlying the dated concretion, and are prevalently represented by isolated small fragments, heavily damaged and scattered by hyenas, as well as all the other large mammal remains. Several juvenile molars have been collected. The excavation data suggest that P. mnaidriensis is the only elephant species in the San Teodoro Cave faunal assemblage. The “Anca” fossil collection stored at the “G.G. Gemmellaro” Museum in Palermo contains small elephant molars from previous excavations in the San Teodoro cave, already recognised as juvenile molars (Anca, 1860; Anca and Gemmellaro, 1867), and wrongly attributed by Burgio and Di Patti (1990) to P. falconeri, although the same authors had realized that the attribution to P. falconeri had to be revised. According to Herridge (2010), some of the large-sized elephants from Sicily and Malta, including the San Teodoro Cave elephant, are to be referred to the new species which is temporary named as “Sicily 3” or “Palaeoloxodon sp.”. In this paper, the elephant from San Teodoro Cave is considered as P. mnaidriensis, taking into account that it is probably to be attributed to the new Sicilian species, as suggested by Herridge (2010). 4. The grouping of elephant bones from the b trench During the 2003 and 2004 excavations of the b trench, a densely accumulated bone pile composed of partially articulated and disarticulated skeletal elements of adult elephant mixed with disarticulated bones belonging to other taxa and with numerous hyena coprolites, was found located close to the eastern wall of the cave, in the uppermost levels of the deposit, at depths between þ0.13 and 0.40 m (relative to the height “0”) (Fig. 3) (Mangano and Bonfiglio, 2010). Damage by hyena activity are present on all the bones belonging to the other taxa, while several elephant bones are not damaged. All the elephant bones are represented by post-cranial portions: a cervical vertebra, a thoracic vertebra, a left scapula, a rib fragment, an unfused right distal epiphysis of radius, a left metacarpal bone, a left coxal bone, two

semi-articulated and symmetrical pairs of tibias and fibulas and a right astragalus. During the same excavations, some post-cranial elements of an adult elephant, represented by two thoracic vertebrae in anatomical connection, a left pyramidal bone, a right femur shaft and a right patella, have been found very close to the pile. During the 2005 excavation, two cranial remains of adult elephants, represented by a semi-complete mandible and a tusk fragment, were recovered in the western sector of the same trench, at a distance of about 5 m from the bone pile. In addition to these adult remains, a small elephant mandible, having the posterior portions of the two hemimandibles damaged by hyenas and preserving the left first deciduous molar totally unworn, indicating a perinatal stage, has been recovered very close to the bone pile (Fig. 1b,c). The list of the adult elephant bones, with indication of the squares and inventory numbers, is presented in Table 1. 4.1. Taphonomic observations The grouping of elephant bones includes partially articulated (two pairs of tibias/fibulas) and anatomically connected (two thoracic vertebrae) elements associated with disarticulated elements. All the skeletal portions are represented: cranial, axial, scapular and pelvic girdle, fore-limb and hind-limb (with the semicomplete right hind-limb) (Fig. 4). The grouping includes both hyena-damaged and non-damaged bones. The mandible is a quite complete specimen, partially broken in the posterior portions of the two hemimandibles; no damage by hyenas has been detected (Fig. 5). The tusk fragment is represented by a fractured apical portion showing several teeth grooves on the tip. All the recovered vertebrae and the rib fragment are hyena-damaged; in particular, traces of intense chewing are visible on the body of the cervical vertebra, while teeth traces are present on the bodies and neural spines of the thoracic vertebrae, including those recovered in anatomical connection (Fig. 6). The scapula, very fractured by carbonate blocks in the flat portion, is represented by a large fragment preserving the articular portion. The poor preservation of this remain largely affects teeth mark observations. The coxal bone is fractured in the wing of ilium and lacks both the pubis and ischium shaft; typical crenulated edges produced by hyena teeth are visible (Fig. 7). The pyramidal bone is complete and not damaged. The III metacarpal bone is represented by a medio-

Table 1 List of the grouped bone remains of Palaeoloxodon mnaidriensis from San Teodoro Cave, with indication of the inventory numbers, squares and damages by hyena activity. 0 ¼ not damaged; (þ) ¼ slightly damaged; (þþ) ¼ moderately damaged; (þþþ) ¼ highly damaged.

Fig. 3. View of the elephant bone pile from square 29 B. A coxal bone, a broken scapula, a tibia, an astragalus and a cervical vertebra of Palaeoloxodon mnaidriensis mixed with a mandible of Equus hydruntinus and a femur and a metatarsal of Bos primigenius siciliae can be seen.

Inv. N.

Square

Skeletal element

Side

Damages

PL PL PL PL PL PL PL PL PL PL PL PL PL PL PL PL PL PL PL

34F 34G 29A 29A 29C 29C 30A 29A 30B 29C 30B 29A 30C 30B 30A 29A 30B 30B 29A

Tusk Mandible Cervical vertebra Thoracic vertebra Thoracic vertebra Thoracic vertebra Rib Scapula Radius distal epiphysis Pyramidal bone III metacarpal bone Coxal bone Femur Patella Tibia Tibia Fibula Fibula Astragalus

L e e e e e e L R L L L R R R L R L R

(þþ) 0 (þþþ) (þþ) (þ) (þ) (þ) 0 0 0 (þþþ) (þ) (þþþ) 0 0 0 (þ) 0 0

2970 2968 2958 2940 2959 2960 384 2969 2966 2961 226 2924 2962 2099 735 2956 2963 2964 2957

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Fig. 4. Anatomical representation of the grouped elephant bone remains from the b trench of San Teodoro Cave.

proximal fragment showing teeth pits and intense chewing. The unfused distal epiphysis of radius is not damaged. The femur shaft lacks the two ends, totally destroyed by hyenas; typical crenulated edges together with many teeth grooves are visible (Fig. 8). The patella is complete and not damaged. The two symmetrical pairs of tibias and fibulas are complete and very well preserved, with the exception of the left tibia which is fractured on the shaft; no traces of hyena activity have been detected, with the exception of a few teeth marks on the distal end of the right fibula (Fig. 9). The astragalus is complete and not damaged. Almost all damaged bones show teeth marks and chewed areas on both the upper and lower surfaces. 4.2. Development stages All remains are consistent with an adult individual. The mandible has the two permanent second molars in middle wear (see Fig. 5); the third molars are not erupted and their germs have been seen by X-ray analysis. All post-cranial bones are fused, with

Fig. 5. Mandible of Palaeoloxodon mnaidriensis with permanent second molars, occlusal view.

Fig. 6. Two thoracic vertebrae of Palaeoloxodon mnaidriensis in anatomical connection, lateral view. The bodies and the neural spines show hyena damage.

the only exception of the radius, represented by a distal unfused epiphysis; the compact bones are very well ossified. As the femur lacks the two ends, it cannot be determined if the epiphyses were fused. The two symmetrical tibias have the distal epiphyses fused and the proximal ones are almost completely fused with the epiphyseal lines still partially visible (see Fig. 9). 5. Discussion As the two symmetrical pairs of tibias and fibulas included in the bone pile undoubtedly belong to a single elephant, the anatomical representation of the remains, the spatial distribution, the measurements, the state of the epiphyseal fusion and the dental

Fig. 7. Left coxal bone of Palaeoloxodon mnaidriensis, caudal view.

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aging of the mandible were used to determine that all the adult bones belong to the same individual. The measurements of the skeletal elements are presented in Table 2; the dimensions of long bones and lower M2 are comparable with those of the adult specimens of “P. mnaidriensis” from the Puntali Cave (Ferretti, 1998, 2008) and those of “Sicily 3” (Herridge, 2010). In elephants, as in other mammals, the elongation of each bone continues until its growth is finished and epiphyses are fused to the shaft (Lister, 1994). Studies on extant and fossil elephants have demonstrated that each elephant species shows a definite order in epiphyseal fusion; the timing of the epiphyseal fusion varies between bones and, often, between epiphyses of the same bone (Roth, 1984; Lister, 1994, 1999; Herridge, 2010). According to Roth (1984) and Haynes (1991), females of each species show the same fusion sequences as males, but fusions occurs at younger ages and are compressed over a shorter period of time. Developmental markers in Loxodonta africana presented by Herridge (2010) show that the female bears its first calf before M2 comes into wear. The lack of both relative and absolute offset in elephants’ fusion timing means that a single epiphyseal fusion is not useful for aging individuals. The aging of an individual from different fusion states of a single bone has been inferred by Herridge (2010, Table A8.9) for Elephas maximus, Loxodonta africana, Mammuthus primigenius and Palaeoloxodon antiquus. In contrast, the rate of tooth progression and wear is fairly constant in extant elephant species, and precise ageing schemes have been developed for L. africana (e.g. Laws, 1966; Jachmann, Table 2 Measurements (mm) of the grouped bone remains of Palaeoloxodon mnaidriensis from San Teodoro Cave. N ¼ lacking plates lost to wear; x ¼ talon.

Fig. 8. Right femur of Palaeoloxodon mnaidriensis, cranial view. The two ends have been totally destroyed by hyenas. Crenulated edges by hyena chewing and numerous teeth traces on the bone surface are visible.

Fig. 9. Two complete symmetrical pairs of tibias and fibulas of Palaeoloxodon mnaidriensis, cranial view.

Lower M2 Plate count Length Width Lamellar frequency (10 cm) Enamel thickness Scapula Glenoid process maximum length Glenoid cavity length Glenoid cavity breadth Pyramidal bone Maximum medio-lateral width Antero-posterior width (excluding lateral apophysis) Ulnar articular surface medio-lateral width Ulnar articular surface antero-posterior width Maximum dorsal length Coxal bone Acetabulum length Acetabulum length includine the lip Femur Minimum diaphysis length Mid-shaft medio-lateral width Mid-shaft antero-posterior width Tibia Total length Mid-shaft medio-lateral width Mid-shaft antero-posterior width Proximal medio-lateral epiphysis width Proximal antero-posterior epiphysis width Distal medio-lateral epiphysis width Distal antero-posterior epiphysis width Astragalus Maximum width Maximum length Tibial articular surface medio-lateral width Tibia articular surface antero-posterior width Navicular articular surface medio-lateral width Navicular articular surface antero-posterior width

R

L

N9x 173 69 5 3.2

N9x 171 70 5 3.1 190 w160 90 87 90 79 75 48 125 136

398 68 55 425 71 64 149 111 125 92 115 111 89 91 89 52

425 71 63 149 111 126 93

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1988) and E. maximus (Roth and Shoshani, 1988). Epiphyseal fusion has been correlated with tooth-ages of E. maximus, L. africana, M. primigenius and P. antiquus (Roth, 1984; Haynes, 1991; Lister, 1999; Herridge, 2010). According to Herridge (2010), the withinbone fusion patterns for dwarf elephant species, based on postcranial material from the Mediterranean islands, indicate that the fusion sequences are generally consistent with the full-sized elephant patterns. However, fusing rate and dental progression are probably not the same in different sized species (Lister, 1994). No post-cranial elements have been found in association with dental remains in the materials studied by Herridge (2010). The associated tibias/fibulas from San Teodoro Cave represent one of the two known examples of associated skeletal elements for any Mediterranean dwarf elephant taxa (Herridge, 2010). According to Herridge (2010), the proximal and distal epiphyses of the tibia have a simultaneous early fusion in most of the dwarf elephant species, while the distal epiphysis of radius has a late fusion. The distal epiphyses of the tibias may be attributed to the stage 3 of the epiphyseal score schemes by Haynes (1991) and Herridge (2010), and to the stage (X) of the epiphyseal score scheme by Lister (1999). According to Haynes (1991), the coxal bone fuses at 8 AEY (African Elephant Years), the proximal and distal tibia, respectively, at 17e24 and 18e20 AEY in the females, and at 28e32 and 32? AEY in the males, while the distal radius fuses at over 24 AEY in the females and over 40 AEY in the males. In the epiphyseal score scheme by Lister (1994), in which this fusion sequence is basically observed, the fusion age of proximal scapula is about 42 AEY in the males, while no data are available for females. The lower M2 of the mandible shows a mid-stage of wear corresponding to the stage XVI by Laws (1966), i.e. 26 LAY (L. africana Years) or 28 EMY (E. maximus Years), according to Laws (1966), and 19-21 AEY, according to Haynes (1991). From these data, all the elephant bone remains have comparable ages, and belong to a single prime-age adult individual, with the only exception of the fused scapula. The latter can probably only pertain to the same carcass if the individual is not a male. The finding of the juvenile mandible in the perinatal stage very close to the adult remains (Fig. 10) could support the hypothesis that the individual was a female (a pregnant female or a female with a calf).

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Unfortunately, sex determination based on the coxal morphology is very difficult as the coxal bone lacks the diagnostic portions (according to Lister, 1996; Göhlich, 2000). Large-sized remains (skull, mandible) of large mammals (M. primigenius, Coelodonta antiquitatis) have been found in several hyena-collected fossil assemblages from Europe, but grouped or piled bones which could pertain to a single carcass have never been  recovered (Diedrich and Zák, 2006; Fosse, 1997). It is unlikely that hyenas transported whole elephant carcasses to the San Teodoro Cave, but, more likely, the elephant entered the cave, perhaps in search of mineral salts as attested for extant elephants (Redmond, 1982; Bowell et al., 1996; Lundquist and Varnedoe, 2006) or looking for shelter, and then it may have been killed by hyenas, or the carcass may have been scavenged after its natural death. Evidence of scavenging by hyenas is represented by teeth marks and chewed areas on both the upper and lower surfaces of the damaged bones, and by the recovery of a great number of coprolites between the bones. The missing portions of the skeleton, especially the largest bones (skull), may have been removed by hyenas and/or other agents. It cannot be ruled out that other elephants could have partly displaced or removed the largest bones of the carcass, as observed in modern elephants (Douglas-Hamilton and Douglas-Hamilton, 1975) and fossil records (Stuart and Larkin, 2010). Furthermore, it is not unlikely that the subsequent human frequentation of the cave, from the Epigravettian until the present time, could have removed or destroyed the elephant bones, as the remains were located in the uppermost level of the deposit very close to the cave floor. The coexistence of hyena-damaged and non-damaged bones in the elephant bone grouping of the San Teodoro Cave cannot be explained by only the different dimensions of the skeletal elements, as the largest bones recovered are both damaged (coxal bone) and undamaged (mandible, tibias), but could attest to a different level of hyena scavenging intensity, depending on the bone density and the level of competition between hyenas. Faith et al. (2007) observed that the skeletal elements with the highest density, such as the mandible, tibia and compact bones, show a lower probability of being selected for consumption by modern spotted hyenas, especially when the level of competition between hyenas is low, for example in clans composed by few individuals. 6. Conclusion

Fig. 10. Juvenile mandible of Palaeoloxodon mnaidriensis in the perinatal stage, occlusal view.

The following features can be pointed out: a) the elephant bone grouping is composed of non-scattered, non-isolated, partially articulated skeletal elements, which are very unusual in a hyenacollected assemblage; b) the ages of both cranial and post-cranial elements indicate that the bones belong to a single adult individual of P. mnaidriensis, probably about 24e30 years of age; c) the largest elements (mandible, scapula, coxal bone, tibias) are much less damaged or undamaged by hyenas, thus suggesting that hyenas could not have transported these large portions of the carcass but had scavenged them “in situ”; d) the missing elements of the skeleton (skull, fore-limb) may have been destroyed by hyenas and/or the continuous and intense anthropic frequentation of the cave from the Epigravettian until the present time, or may be present but still buried in the sediments. From these data, the elephant (perhaps a pregnant female or a female with a calf) for some reason (perhaps in search of mineral salts or shelter) entered the cave when it was still alive and that hyenas scavenged the carcass after its death due to natural causes or predation by the hyenas. It is not unlikely that other elephants could have displaced the largest elements of the skeleton. Anyway, as it is very unlikely that elephants frequented the cave when it was also frequented by hyenas, these remains may also attest to intermittent use of the cave by Crocuta crocuta spelaea.

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