An exclusively hyena-collected bone assemblage in the Late Pleistocene of Sicily: taphonomy and stratigraphic context of the large mammal remains from San Teodoro Cave (North-Eastern Sicily, Italy)

An exclusively hyena-collected bone assemblage in the Late Pleistocene of Sicily: taphonomy and stratigraphic context of the large mammal remains from San Teodoro Cave (North-Eastern Sicily, Italy)

Journal of Archaeological Science 38 (2011) 3584e3595 Contents lists available at SciVerse ScienceDirect Journal of Archaeological Science journal h...

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Journal of Archaeological Science 38 (2011) 3584e3595

Contents lists available at SciVerse ScienceDirect

Journal of Archaeological Science journal homepage: http://www.elsevier.com/locate/jas

An exclusively hyena-collected bone assemblage in the Late Pleistocene of Sicily: taphonomy and stratigraphic context of the large mammal remains from San Teodoro Cave (North-Eastern Sicily, Italy) Gabriella Mangano 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: Received 7 January 2011 Received in revised form 22 August 2011 Accepted 23 August 2011

A detailed taphonomic analysis of the large mammal assemblage from 1998 to 2006 excavations at San Teodoro Cave is presented, taking into account the stratigraphic context of the deposits. Three not strictly contemporary fossiliferous levels having different lithological features have been detected, here named B-I, B-II, and B-III. Fossil remains are prevalently accumulated in B-I and B-II. The three levels are characterized by evidence of Crocuta crocuta spelaea occupation, represented by their skeletal remains, coprolites, and distinctive damages on the bones, similar to fossil and modern spotted hyena dens from Europe and Africa. A differential distribution of coprolites and small digested bones, probably due to different humidity conditions, has been recognized in B-I and B-II, and can be related to different topographic locations within the cave or to different climate conditions during the sedimentation phases. The very low density of fossil remains in B-III, which is the oldest level, could indicate an area that was less inhabited by hyenas, probably due to geomorphological conditions. Taphonomic comparison of the three fossiliferous levels of the San Teodoro Cave deposits points to a long-term, perhaps cyclic, occupation of the cave by hyenas and confirms the cave as one of the most important Pleistocene hyena dens in Europe. Ó 2011 Elsevier Ltd. All rights reserved.

Keywords: San Teodoro Cave Hyena den Late Pleistocene Taphonomy Cave stratigraphy Sicily

1. Introduction The San Teodoro Cave deposits were discovered in 1859, and several non-stratigraphic excavations were conducted up to the mid-twentieth century by important researchers. The cave is known mainly for its important Upper Palaeolithic human burials and artefacts, referred to the Late Epigravettian, although an older deposit containing Pleistocene mammal remains has been distinguished underneath the cultural level by all authors (Anca, 1860; Fabbri, 1993; Graziosi, 1943, 1947; Graziosi and Maviglia, 1946; Martini, 1997; Maviglia, 1941, 1942; Vaufrey, 1928, 1929; Vigliardi, 1968, 1989). Modern excavations, conducted by Laura Bonfiglio, started in 1998 and went on till 2006 for a total of eight months and six excavation seasons (1998, 2002, 2003, 2004, 2005, 2006). Two trenches have been excavated, named as the “a” trench close to the entrance and the “b” trench inside the cave (Bonfiglio et al., 2008) (Fig. 1). Only the older deposit (named as “unit B” in Bonfiglio et al., 2001) was investigated, as the Upper Palaeolithic one had been

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almost totally removed by previous excavations and only very small surviving portions (less than 50 cm3) have been recovered within the a trench; other residual portions of the Epigravettian soil encrusting the walls are preserved in the cave. The large mammal assemblage from unit B is composed of 4864 fossil bone remains and 10086 hyena coprolites. The following large mammal taxa have been identified: Palaeoloxodon mnaidriensis, Equus hydruntinus, Bos primigenius siciliae, Bison priscus siciliae, Cervus elaphus siciliae, Sus scrofa, Crocuta crocuta spelaea, Canis lupus, and Vulpes vulpes. Evidence of contemporary human occupation, such as stone artefacts or cut marks on bone surfaces, are lacking. Preliminary taphonomic analysis carried out on part of the materials from the a trench has indicated that the bone remains were collected by hyenas (Bonfiglio et al., 1999; Marra et al., 2004). The unit B deposit has been also retrieved from several test-pits excavated at different points of the cave floor (Fig. 1), testifying that the whole cave area, having a total surface of about 1000 m2, was inhabited by hyenas; for this reason the San Teodoro Cave is the largest Pleistocene hyena den so far known in an insular environment (Bonfiglio et al., 2001, 2008; Mangano and Bonfiglio, 2005a,b; Mangano et al., 2005). The moderate degree of endemism of the large mammal association, also confirmed by the small mammal

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Fig. 1. Plan of the San Teodoro Cave with locations of the two main fossiliferous levels (B-I, B-II; see the text for explanation). Grey squares: excavated area (1998e2006 excavations). Black lines in the a trench: unexcavated area. Black circles: locations of the test-pits.

taxa, attested to a new population phase in Sicily referred to the “San Teodoro Cave-Pianetti” Faunal Complex, which was originated by dispersal events through temporary terrestrial connections (land bridges) presumably located in the Messina Strait area (Bonfiglio et al., 2001, 2002, 2008, 2009; Masini et al., 2008). Deposits from the b trench, unlike those from the a trench, show a quite complex stratigraphy with three recognized fossiliferous levels, whose faunistic features point to different deposition phases which probably occurred during a short time span (Bonfiglio et al., 2008; Esu et al., 2007). Almost all of the collected materials come from the two youngest fossiliferous levels (here named as B-I and B-II; see Section 4). A radiometric dating carried out using the 230 Th/234U method on a concretionary level within the b trench has yielded an age of 32000  4000 yr (Bonfiglio et al., 2008). The dispersal that gave origin to the San Teodoro Cave mammal assemblage most likely took place during the MIS4 stage of the Atlantic Oxygen Isotope curve. Pollen analysis from coprolites of the a trench has attested a steppe-like landscape with a low presence of mesophilous taxa (Yll et al., 2006). Pollen analysis of coprolites from the b trench is in progress. Hyenas are considered as an important taphonomic agent in the formation of many bone assemblages from African and European PlioePleistocene sites and evidence of alternate occupation by humans and hyenas has been frequently recognized (e.g., Arribas and Palmqvist, 1998; Brugal et al., 1997; Diedrich, 2009; Diedrich  and Zák, 2006; Enloe et al., 2000; Fernández Rodríguez et al., 1995; Fosse, 1994, 1996, 1997; Klein, 1975; Klein and Cruz-Uribe, 1984; Klein et al., 1991; Marean et al., 2000; Palmqvist and Arribas, 2001; Palmqvist et al., 1996; Stiner, 1991, 1992; Villa and Bartram, 1996; Villa et al., 2004, 2009). Many actualistic studies have been conducted in order to clarify the role of these carnivores in the formation of the archaeological and/or palaeontological record (e.g., Binford, 1981; Brain, 1981; Bunn, 1983; Capaldo and Blumenschine, 1994; Faith and Behrensmeyer, 2006; Faith et al., 2007; Hill, 1980, 1984, 1989; Holekamp and Smale, 1998; Holekamp et al., 1997; Kruuk, 1972; Kuhn et al., 2010; Lansing et al., 2009; Pickering, 2002; Pokines and Kerbis Peterhans, 2007; Sutcliffe, 1970). From a taphonomic point of view, the bone assemblage from the recent excavations at San Teodoro Cave is particularly important for two reasons: the first is that the hyena is the only agent of bone accumulation; the second is that San Teodoro Cave is one of the few Pleistocene hyena sites in Europe for which not only are detailed

stratigraphic data available but also sediment total screening has been carried out. In this contribution, the taphonomic analysis of the entire large mammal assemblage from the unit B deposit of the a and b trenches of San Teodoro Cave (1998e2006 excavations) is presented for the first time and taphonomic relationships between the two main recognized fossiliferous levels are investigated. Data about species abundance, bone and coprolites distribution, bodypart representation, prey age distribution, and bone modification and fragmentation are presented separately for each level in order to detect similarities or differences in deposition and modification modalities of bone remains by hyenas. 2. The site The San Teodoro Cave is a large cavity located at Acquedolci, a small town on the north-eastern coast of Sicily in the province of Messina (Fig. 2a). The cave opens in the northern cliffs of a Jurassic carbonatic massif (San Fratello Massif) at the height of 145 m a.s.l. It was created by karst processes and its opening was due to the intense distensional tectonics which affected this area (Lentini et al., 2000; Robillard, 1975). The San Teodoro Cave is composed of a single large chamber, about 60 m long, 20 m wide, and up to 20 m high, and one much smaller lateral chamber located on the eastern side of the main chamber. In the innermost part of the cave, sub-horizontal narrow conduits are also present. The entrance of the cave, which is triangular, is about 12 m wide and 5 m high at present (Fig. 2b). Currently, the floor of the cave ascends about 15 m from the entrance to the end of the cave along its major axis. The central part of the floor is a detrital fan which slopes down laterally toward the eastern and western walls of the cave. Recent reworked sediments having different thicknesses overlie the unit B deposit (Bonfiglio et al., 2008). 3. Excavation methods A one-metre grid system identified by surface co-ordinates represented by numbers and capital letters was superimposed on the cave floor (Fig. 1). Large sized faunal remains were collected in situ and plotted in three Cartesian coordinates. All sediments have been water-screened with superimposed screens of 5 and 2 mm

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DeEeFeG; Fig. 1) the cave floor was very irregular and the occurrence of very large boulders prevented extensive cuts in all squares. 4. Stratigraphy

Fig. 2. Map of Sicily showing the location of the site of San Teodoro Cave (a). View of the cave entrance (b).

mesh, with total recovery of bone and teeth fragments, small vertebrates, molluscs, vegetable remains, and coprolite fragments. The a trench (1998 excavation) was located on the eastern side of the cave near the entrance over an area of 25 m2 (see Fig. 1) at depths from 2.43 m to 3.85 m with respect to the fixed landmark (height “0”; Fig. 3). Only the eastern squares of the a trench have been excavated (sq. EeFeG; Fig. 1), as the western sector was occupied by reworked sediments from the Graziosi and Maviglia excavations (Bonfiglio et al., 2004). The b trench (2002e2006 excavations) was located on the inner eastern side of the cave at a distance of 28 m from the entrance over an area of 28 m2 (see Fig. 1) at depths from þ1.75 m to 0.67 m with respect to the fixed landmark (Fig. 3). In the western sector of the trench (sq.

The fossiliferous unit B deposit of San Teodoro Cave is composed of grey-green clayey sand including 1e10 cm sized blocks of limestone together with large carbonate boulders (maximum diameter of more than 1 m) (Bonfiglio et al., 2001, 2008). During the 1998e2006 excavations three different levels have been recognized in the unit B deposit, here named as B-I, B-II, and B-III (Fig. 4), characterized by different lithological features. B-I, the richest fossiliferous level, is mainly composed of clayey sand containing very few large carbonate boulders, and no concretionary processes on fossil bones and coprolites or on sediments and boulders have been recognized; this level occupies the whole investigated a trench and the eastern sector of the b trench (sq. AeBeC; Fig. 1). B-II is mainly composed of large concretioned carbonate boulders intercalated with concretioned levels of clayey sand and small lenses of non-concretioned clayey sand; the concretioning process is very intense and also affected bones and coprolites. This level has been highlighted in the western sector of the b trench (sq. DeEeFeG; Fig. 1). B-III has been recognized in the most western sector of the b trench, underlying the B-II deposit (sq. 33-34/FeG; Fig. 1); this level is mainly composed of clays and contains very few fossil remains, but it has not yet been well investigated. Previous analyses carried out on molluscs and small mammals from B-I and B-II pointed to different environmental conditions and perhaps different ages for these deposits (Bonfiglio et al., 2008; Esu et al., 2007). On the basis of the radiometric dating carried out on a concretionary level underlying both B-I and B-II and overlying BIII, the first two levels would be younger and the third one older than 32000  4000 (Bonfiglio et al., 2008). Although further investigations are needed to clarify the rather complex stratigraphy of the western sector of the b trench, the deposition of the B-I level seems to be later than that of B-II, as the B-I sediments are overlying the erosional surfaces of B-II. 5. Materials The faunal assemblage from the two excavated trenches of San Teodoro Cave consists of 14549 objects, including materials from screening. Small vertebrates are not included. If we also include objects from test-pits and surface recovery, the total number of recovered materials is 14950 (Table 1). The number of remains from B-I, B-II, and B-III is shown in Table 2. Coprolites from the three levels are extremely abundant and represent about two thirds of the material (B-I bone/coprolite ratio ¼ 1:2.1; B-II bone/coprolite ratio ¼ 1:2.2; B-III bone/coprolite ratio ¼ 1:1.8). Because of intense fragmentation, bone remains are mostly represented by unidentifiable shaft splinters. About one

Fig. 3. Profile of the San Teodoro Cave, from the entrance to the position of the b trench, and locations of the excavated trenches. The cross indicates the fixed landmark (height ‘0’). The numbers indicate the maximum excavated depth. A: unit A deposit. B: unit B deposit. R: reworked recent level (after Bonfiglio et al., 2008; modified).

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Fig. 4. Schematic section of the sedimentary sequence of the unit B deposits cropping out in the b trench along the segment 29Ae34G. R: recent. B-I: non-concretioned unit B. B-II: concretioned unit B. B-III: clayey unit B. CL: radiometrically dated concretion. Grey: fossil bones (after Bonfiglio et al., 2008; modified).

quarter of the identified bones from B-I and one third from B-II are represented by isolated teeth; if we exclude teeth, 20.7% and 13.1% of bone remains are identifiable in B-I and B-II, respectively. Total percentages of skeletal remains and coprolites from the two main fossiliferous levels are comparable, even though in B-II the bone/ teeth ratio is smaller and bone splinters are slightly better represented (Table 2). Table 3 shows the excavated areas (m2) and volume (m3) for each level, excluding large boulders, and the distribution per cubic metre of the materials. The highest density of fossil remains is observed from B-I, where the number per cubic metre is nearly twice that from B-II. Bone concentration in B-III is very low and fossil remains are decidedly sporadic. Because of this, only data from B-I and B-II will be discussed and compared with the available data from fossil and modern hyena sites. 6. Results 6.1. Large mammal composition A total NISP of 983 large mammal remains from B-I and 130 remains from B-II were recovered (see Table 2). The relative abundance of the large mammal taxa by NISP and by MNI is shown, respectively, in Table 4 and Table 5. The MNI was calculated taking into account the side and the age of the most common skeletal element of a taxon. The faunal assemblage is dominated by herbivores, which represent about 90% of the total NISP in both the levels. The endemic red deer Cervus elaphus siciliae is the most common taxon, with the highest NISP percentage (more than 60%). All the other herbivores (the wild boar S. scrofa, the small horse Equus hydruntinus, the endemic bovids Bos primigenius siciliae and Bison priscus siciliae, and the endemic reduced-size elephant Palaeoloxodon mnaidriensis) show much lower frequencies with comparable values (5.1%e8.8% in B-I; 3.8%e8.5% in B-II). Since the distinction between Bos and Bison was not possible for all specimens, the two genera have been grouped together (when the

Table 1 Number of large mammal remains from 1998 to 2006 excavations at San Teodoro Cave.

Identifiable bones Teeth Bone splinters Coprolites Total

Unit B

Reworked

Total

845 277 3467 9960 14549

45 28 202 126 401

890 305 3669 10086 14950

distinction was possible, remains of Bos absolutely predominate over Bison). The herbivore to carnivore ratio by NISP and MNI is illustrated in Fig. 5 and Fig. 6 respectively. The MNI ratio shows a higher proportion of hyena, with a frequency of 26.9% in B-I and 13.3% in B-II, probably biased by the lower degree of fragmentation of hyena remains; the MNI frequency of hyena from B-I is very close to that of Cervus elaphus siciliae, which is the most abundant prey but has the highest degree of fragmentation of remains, as indicated by the NISP to MNI ratio (see Tables. 4, 5). The other carnivores, C. lupus and Vulpes vulpes, are extremely scarce. At present, skeletal remains of Crocuta crocuta spelaea are lacking in B-III, where there is, however, evidence of hyena occupation, such as coprolites and bone damage. B-I and B-II show similar faunal associations, referred to the “San Teodoro Cave-Pianetti” Faunal Complex of the late Late Pleistocene of Sicily; this faunal complex, for which only the radiometric dating from San Teodoro Cave is available, is characterized by a low degree of endemism (Bonfiglio et al., 2001, 2008). The assemblages are characterized by significant frequencies of the Pleistocene European cave hyena, which is genetically closely related to the extant spotted hyena (Rohland et al., 2005). The frequency of cave hyena from various European fossil dens shows a wide range of variation, probably related to climate and environmental conditions: the highest frequencies of hyena are observed in the assemblages of temperate climates indicating an open grassland-like landscape; low frequencies characterize assemblages indicating forest landscape; frequencies from cold climate assemblages are strongly variable (Fosse, 1994, 1996, 1997). At San Teodoro Cave the NISP representations of hyena from B-I (8%) and B-II (12.3%) are not too different; they fall at the lower end of the range of values indicated in several European fossil hyena dens (Table 6). The MNI representations of hyena from B-I (26.9%) and B-II (13.3%) have higher values, of which the former is comparable with the recognized values from various fossil hyena dens of Southern Africa (hyena MNI frequency > 20%)

Table 2 Distribution of large mammal remains from the three excavated levels of San Teodoro Cave. B-I

Identifiable bones Teeth Bone splinters Coprolites Total

B-II

B-III

NR

%NR

NR

%NR

NR

%NR

744 239 2846 8249 12078

6.2 2 23.5 68.2

92 38 608 1670 2408

3.8 1.6 25.2 69.3

9

14.3

13 41 63

20.6 65

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Table 3 Approximate extension and volumetry of the three excavated levels and distribution of remains per cubic metre at San Teodoro Cave.

Area (m2) Volume (m3) Identifiable bones þ teeth per m3 Bone splinters per m3 Coprolites per m3 Total objects per m3

B-I

B-II

B-III

22 17 57.8

14 6 21.6

4 2 4.5

167.4 485.2 710.4

101.3 278.3 401.2

6.5 20.5 31.5

(Cruz-Uribe, 1991; Klein and Cruz-Uribe, 1984). However, in modern spotted hyena dens MNI values are much lower (<10%, according to Kerbis Peterhans, 1990; <13%, according to Kuhn et al., 2010). 6.2. Body-part representation The body-part representation based on the NISP is given in Table 7 and shown, separately, in Fig. 7 and Fig. 8. The medium-size ungulates (Cervus elaphus siciliae, Sus scrofa, E. hydruntinus) are prevalently represented by cranial portions, among which isolated teeth and antlers predominate, while mandibles are less abundant; the postcranial portions are well represented, with a predominance of metapodials and phalanges. Due to the intense fragmentation, deer antler fragments are particularly abundant (17% of the identifiable remains in B-I and 35% in B-II) and mainly represented by basal portions belonging to both shed and unshed antlers. Hyena is mainly represented by cranial portions (teeth, mandibles, skulls), while postcranial parts are very few. Bovids (Bos primigenius siciliae and Bison priscus siciliae) are mostly represented by postcranial bones and vertebrae in B-I and by cranial remains and vertebrae in B-II. The endemic elephant P. mnaidriensis is prevalently represented by teeth and vertebrae. Excluding only one hyena vertebra, at San Teodoro Cave all preserved axial portions belong to the large herbivores (bovids and elephant). High percentages of antlers have been observed in fossil hyenacollected assemblages from Italy, where cervids are generally well represented (Piperno and Giacobini, 1990e91; Pitti and Tozzi, 1971; Stiner, 1991, 2004); antler fragments from Pleistocene hyena dens of western and eastern Europe show variable percentages (Diedrich  and Zák, 2006; Fosse, 1994; Guadelli, 1989; Marra et al., 2004; Turner, 1981). In modern spotted hyena dens the proportions of horn fragments, mainly produced by skull fragmentation, are generally low (Kuhn et al., 2010; Pickering, 2002; Pokines and Kerbis Peterhans, 2007). According to Cruz-Uribe (1991) and Klein and Cruz-Uribe (1984), one important criterion for the recognition of fossil

Table 5 MNI frequency of the large mammal taxa from San Teodoro Cave. B-I

Palaeoloxodon mnaidriensis Bos primigenius siciliae/Bison priscus siciliae Equus hydruntinus Sus scrofa Cervus elaphus siciliae Crocuta crocuta spelaea Vulpes vulpes Canis lupus Total MNI

B-II

B-III

MNI

%MNI

MNI

%MNI

MNI

%MNI

4 5

7.7 9.6

1 2

6.6 13.3

1 1

20 20

6 5 16 14 1 1 52

11.5 9.6 30.8 26.9 1.9 1.9

2 3 4 2 1

13.3 20 26.6 13.3 6.6

2 1

40 20

15

5

hyena-collected assemblages is that the ungulate cranial/postcranial ratio, based on MNI, tends to decrease with increases in ungulate size, as hyenas are considered able to transport adult skulls of small ungulates but tend to abandon the skulls of larger ungulates. The decreasing of the cranial/postcranial ratio for ungulates has been recognized in modern and fossil hyena dens (Cruz-Uribe, 1991; Fosse, 1994; Guadelli, 1989; Klein and CruzUribe, 1984; Pokines and Kerbis Peterhans, 2007; Turner, 1981), but several fossil hyena sites from Europe show the opposite trend (Fosse, 1997). Even though this criterion may be biased by differential preservation of cranial remains, especially teeth, and some refinements have been proposed (Kerbis Peterhans, 1990; Pickering, 2002), at San Teodoro Cave the MNI cranial/postcranial sample size is too low to be significant. Cruz-Uribe (1991) and Klein and Cruz-Uribe (1984) also stated that the small compact bones of prey species, such as carpals and tarsals, should be absent or very rare in hyena accumulations. At San Teodoro Cave the percentages of carpals, tarsals, and phalanges of the smaller herbivore taxa (Cervus elaphus siciliae, S. scrofa) are 15.5% from B-I and 16.9% from B-II (see Table 7), in agreement with the observed data from modern dens (Kuhn et al., 2010; Pickering, 2002; Pokines and Kerbis Peterhans, 2007). According to Stiner (2004), balanced representation of cranial (horn/antler þ skull þ mandible) and postcranial (excluding phalanges) portions of the medium-size ungulates should reflect a hyena foraging system mainly based on hunting; on the contrary, horn/antler-dominated faunas could suggest scavenging. At San Teodoro Cave the three medium-size ungulate taxa from B-I and BII are well represented by both cranial and postcranial elements, and antler fragments are very abundant. Recognizing scavenging or hunting from fossil remains is very difficult. Since Crocuta crocuta

Table 4 NISP frequency of the large mammal taxa from San Teodoro Cave. B-I

Palaeoloxodon mnaidriensis Bos primigenius siciliae/Bison priscus siciliae Equus hydruntinus Sus scrofa Cervus elaphus siciliae Crocuta crocuta spelaea Vulpes vulpes Canis lupus Total NISP

B-II

B-III

NISP

%NISP

NISP

%NISP

NISP

%NISP

68 50

7 5.1

5 7

3.8 25.3

1 2

11.1 22.2

62 87 625 79 8 4 983

6.3 8.8 63.5 8 0.8 0.4

5 11 84 16 2

3.8 8.5 64.7 12.3 1.5

3 3

33.3 33.3

130

9

Fig. 5. NISP representation of the herbivore to carnivore ratio from the B-I and B-II levels of San Teodoro Cave.

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Table 6 Frequency of hyena remains from some Pleistocene European sites (after Fosse, 1996, 1997; Guadelli, 1989; Marra et al., 2004; Piperno and Giacobini, 1990e1991; Pitti and Tozzi, 1971; Stiner, 1994; Turner, 1981). % NISP Grotta Guattari - Italy Grotta dei Moscerini - Italy Buca della Iena - Italy Cueva de Las Hienas - Spain Bois Roche - France Lunel Viel lower levels - France Morancourt - France Camiac - France Conives - France KirkdaleeUnited Kingdom Pin Hole - United Kingdom San Teodoro Cave (B-I) - Italy

Fig. 6. MNI representation of the herbivore to carnivore ratio from the B-I and B-II levels of San Teodoro Cave.

spelaea was the only large predator existing in the area of San Teodoro Cave, where other competitors for food were almost totally absent (the wolf is very scarcely represented in our accumulation and humans were absent), it is not unlikely that hunting was the predominant feeding strategy. This is consistent with modern observations on spotted hyena feeding behaviour (Holekamp et al., 1997; Kruuk, 1972; Lansing et al., 2009), even if the postcranial morphology of extant spotted hyena indicates more cursorial capability and, probably, a greater aptitude for active hunting than Pleistocene cave hyena (Lewis and Werdelin, 2000).

6.3. Age distribution The MNI age determination of the main large mammal taxa from B-I and B-II, basically calculated by the teeth eruption/abrasion and/or the epiphyseal fusion method, is presented in Table 8. The medium-size ungulates (Cervus elaphus siciliae, S. scrofa, E. hydruntinus) are distributed across all considered age classes, while the elephant and bovids remains surely attest the presence of young and adult individuals, but the probability that some adult postcranial remains belong to old individuals should not be excluded. In B-I, the hyena is represented by all age classes and adult individuals predominate. The hyena age distribution from B-I and B-II attests the contemporary presence of both juvenile and adult individuals, indicating the use of the cave as a communal den (see Section 6.6). In B-I, the mortality profile of the best represented prey, Cervus elaphus siciliae, based on DP4 and M2 crown heights (according to Klein et al., 1981), shows an attritional pattern (Fig. 9). Few data are available for comparison, but a variable pattern has been observed

% MNI

10 5.6 22 78.8 7.4 12 11.2 7.3 17 62 17 8

68.2 10.5 26 12.7 11.1

26.9

from Pleistocene European sites, where adult and/or young herbivores are generally predominant (Fosse, 1996). The U-shaped (attritional) prey mortality profile has been considered a distinctive feature of fossil hyena accumulations (Cruz-Uribe, 1991; Klein and Cruz-Uribe, 1984; Stiner, 1994). Hunting and/or scavenging by modern spotted hyenas can produce both attritional (Kruuk, 1972; Mills, 1984) and non-attritional prey mortality profiles (Pickering, 2002). 6.4. Bone modification The total number of recovered bone remains, excluding teeth, is 3590 from B-I and 700 from B-II, mainly represented by unidentifiable bone splinters (see Table 2). Surface preservation of the bones is very good. Most of the bone splinters have dimensions < 3 cm (Fig. 10); these specimens have also been observed by SEM and optical microscopy in order to identify the digestion traces. Distinctive damages by hyena activity have been recognized. Digestion traces (Binford, 1981; Brain, 1981; Lyman, 1994; Sutcliffe, 1970; Villa and Bartram, 1996) and edge polish (according to Pokines and Kerbis Peterhans, 2007; “ragged edges” in Maguire et al., 1980; Lyman, 1994; Villa and Bartram, 1996; “crenulated edges” in Binford, 1981) are the most common kinds of damages. Tooth grooves and tooth pits are less frequent and are sometimes associated with the edge polish (in this case they were not counted); only a few remains from B-I show the scooping out of cancellous bone (Fig. 11). The proportions of edge polish and digested bones from B-I and B-II are very different. In particular, the frequency of digested bones from B-II is much higher with respect to the edge polish. This can be explained considering that most of the bone remains from B-II are

Table 7 Anatomical part distribution of the large mammal taxa from San Teodoro Cave.

Antlers Skull Mandibles Teeth Axis Girdles Fore limb Hind limb Metapodials Podials Phalanges Total

Palaeoloxodon

Bos/Bison

B-I

B-I

4 18 27 4 4 5 1 5 68

B-II

3 2

5

1 1 5 8 1 4 10 8 10 2 50

Equus B-II

2 2 2

B-I 2 4 47

Sus B-II

4

1

1 1 3

1

7

4 62

5

B-I 3 2 22 2 1 12 8 15 7 15 87

Cervus B-II 1 2

2 2 3 1 11

Crocuta

B-I

B-II

198 19 27 111 8 8 40 21 62 47 84 625

26 3 4 21 1 2 1 5 4 17 84

B-I

B-II

13 17 30 1 2 7 6 1 2

3 2 5

79

16

1 3 1 1

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Fig. 8. Body-part representation by NISP from the B-II level of San Teodoro Cave. Fig. 7. Body-part representation by NISP from the B-I level of San Teodoro Cave.

represented by small flakes having dimensions < 3 cm (276 out of a total of 538 in B-II; 408 out of a total of 2049 in B-I). At Bois Roche, where complete coprolites are very scarce and small coprolite fragments are abundant, the very high number of small digested pieces has been related to the dissolution in the sediments of the droppings containing small digested bone fragments (Marra et al., 2004). As the B-II level of San Teodoro Cave is indicative of a more humid phase, this interpretation could explain the difference in frequency between the two levels. The edge polish has been recognized on both identifiable and unidentifiable remains, while digested bones are mainly represented by unidentifiable shaft splinters and by less frequent antler fragments and short bones; tooth groves and pits show variable width and depth (Fig. 12). At San Teodoro Cave the intense fragmentation of the remains has prevented the preservation of complete long bones, and the shaft portions are absolutely predominant. Table 9 presents data on the long bone representation of herbivore taxa from B-I. The proportions of long bones turned into “cylinders”, which is considered a typical feature of hyena activity (Binford, 1981; Bunn, 1983; Cruz-Uribe, 1991; Kerbis Peterhans, 1990; Klein and Cruz-Uribe, 1984; Pokines and Kerbis Peterhans, 2007), range from 5.7% (metapodials) to 23% (humerus). Most cylinders came from medium-size ungulates (Cervus elaphus siciliae, S. scrofa, E. hydruntinus), but

cylinders from the endemic reduced-size P. mnaidriensis (two juvenile humeri, one adult femur) and Bos primigenius siciliae/Bison priscus siciliae (one adult humerus) have also been recovered. The humerus and the tibia are dominated by the survival of the distal relative to the proximal end, probably due to the greater toughness and lesser nutritive value of their distal ends. The radius is not dominated by survival of either end. The femur appears mainly represented by distal portions, even if the data could be biased by the low value of the NISP (14). Metapodials have the highest proportion (24.6%) of complete elements, probably due to their lower nutritive value. Two complete and symmetrical tibiae of P. mnaidriensis, belonging to the same individual and showing very few hyena marks, have also been recovered (see Section 6.7). Gnaw marks by hyenas have been also recognized on materials from B-III. All recognized bone modifications at San Teodoro Cave are typical of hyena activity and attest Crocuta crocuta spelaea occupation during the sedimentation phases which created B-I, BII, and B-III. In particular, the high frequency of bone damages and the long bone modification pattern are consistent with modern spotted hyena assemblages (Bunn, 1983; Cruz-Uribe, 1991; Faith, 2007; Kuhn et al., 2010; Lam, 1992; Maguire et al., 1980; Pokines and Kerbis Peterhans, 2007).

Table 8 MNI age distribution of the large mammal taxa from San Teodoro Cave, based on the teeth eruption/abrasion and/or epiphyseal fusion method.a

Young Adult Old Total a

Palaeoloxodon

Bos/Bison

B-I

B-II

B-I

Equus

2 2

1

2 3

2

4

1

5

2

B-II

Sus

Cervus

Crocuta

B-I

B-II

B-I

B-II

B-I

B-II

B-I

B-II

3 2 1 6

1 1

3 1 1 5

2

5 9 2 16

1

3 9 2 14

1 1

2

1 3

3 4

2

The age determination from B-I is based on the following bones (the inventory numbers are also reported): two juvenile mandibles with deciduous molars (PL 2484, PL 2967) and two fused left fibulae (PL 16, PL 2964) of Palaeoloxodon mnaidriensis; one unfused left humerus (PL 2922), one not comparable unfused right femur (PL 191), two fused left tibiae (PL 4813, PL 2485), and one not comparable fused right tibia (PL 6188) of Bos/Bison; two lower deciduous teeth with different degrees of abrasion (PL 495, PL 753), one juvenile mandible with deciduous teeth (PL 2428), two adult left hemimandibles with little worn permanent molars (PL 102, PL 754), and one very worn upper molar (PL 739) of Equus hydruntinus; one maxillar bone with deciduous teeth (PL 1594), two left hemimandibles with not completely erupted permanent teeth (PL 4817, PL 4892), one maxillar bone with little worn permanent teeth (PL 1359), and one isolated very worn M3 (PL 320) of Sus scrofa; five right hemimandibles with deciduous teeth (PL 368, PL 1981, PL 2581, PL 2714, PL 3967), three left hemimandibles with not erupted molars (PL 612, PL 2939, PL 3787), six right hemimandibles with little worn molars (PL 766, PL 880, PL 1979, PL 4212, PL 4502, PL 6210), and two very worn right M3 (PL 2534, PL 4503) of Cervus elaphus siciliae; two left hemimandibles with deciduous teeth (PL 758, PL 4311), one isolated lower deciduous premolar (PL 3551), nine left mandibles with little worn permanent teeth (PL 2429, PL 2699, PL 2980, PL 3027, PL 3054, PL 3470, PL 4522, PL 4819, PL 4858), and two isolated very worn M1 (PL 3025, PL 4193) of Crocuta crocuta spelaea. The age determination from B-II is based on the following bones: one large atlas (PL 2948) of Palaeoloxodon mnaidriensis; two left hemimandibles with little worn M3 (PL 473, PL 2947) of Bos/Bison; one unworn (PL 115) and one worn (PL 67) upper molar of Equus hydruntinus; two very young long bones (tibia: PL 4784; metapodial: PL 4789) probably belonging to a newborn or a foetus, one unfused radius (PL 1398), and one very worn M3 (PL 49) of Sus scrofa; one maxillar bone with deciduous teeth (PL 78), one right hemimandible with very worn M3 (PL 1333), and two isolated very worn right M3 (PL 1559, PL 2262) of Cervus elaphus siciliae; two lower deciduous teeth (PL 116, PL 1416) and one left hemimandible having little worn teeth (PL 2974) of Crocuta crocuta spelaea.

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Fig. 9. Mortality profile of Cervus elaphus siciliae from the B-I level of San Teodoro Cave (according to Klein et al., 1981).

Fig. 11. Frequency of damage by hyena activity on fossil bones from the B-I and B-II levels of San Teodoro Cave.

6.5. Coprolites

6.6. Hyena cub remains

Hyena coprolites from the San Teodoro Cave assemblage are extremely abundant and very well preserved (see Table 2). Coprolites are homogeneously distributed in all investigated sectors. Numerous complete specimens come from B-I while fragmentary coprolites are prevalent in B-II (Fig. 13). Complete specimens show the typical sub-spherical shape, with a concave end and another, convex or pointed end, which has been observed in modern spotted hyena droppings (Horwitz and Goldberg, 1989; Larkin et al., 2000) (Fig. 14). Very small digested bone fragments have been observed inside the coprolites. The widest diameters of complete coprolites from San Teodoro Cave generally range from 2 to 5 cm (Fig. 15). Coprolite fragments from B-II show smaller dimensions with respect to the fragments from B-I and more than 50% of them have lengths < 2 cm (Fig. 16). High percentages of fragmentary coprolites indicate destruction in situ, probably due to humidity and/or  intensive trampling by hyenas (Diedrich and Zák, 2006; Marra et al., 2004). In our case, the high frequency of complete coprolites recovered from both B-I and B-II seems to exclude intensive trampling by hyenas. The greater frequency of fragments from B-II is probably related to more humid conditions, as attested by the lithology and molluscs fauna (Bonfiglio et al., 2008; Esu et al., 2007). There is no comparison between the impressive number of coprolites from B-I of San Teodoro Cave, whose preservation has probably been favoured by a low degree of humidity and by the absence of trampling, and other Crocuta fossil dens recognized in Europe.

A total of 20 remains of hyena cubs from B-I and six remains from B-II have been recovered, mainly coming from the b trench (22 remains). Remains from B-I are represented by four cranial remains, one maxillary bone with deciduous teeth, two mandibles with deciduous teeth, one isolated lower deciduous premolar, eleven germs of permanent teeth and one fragment of juvenile radius (Fig. 17); they represent a minimum number of three individuals. Remains from B-II are represented by two lower deciduous premolars, two germs of permanent teeth, one skull fragment, and one juvenile tibia, belonging at least to one individual. Only one isolated deciduous tooth from B-I, having resorbed roots, could have been naturally shed in the cave; all the other isolated

Fig. 10. Length distribution of the unidentifiable bone splinters from the B-I and B-II levels of San Teodoro Cave.

Fig. 12. Bones from San Teodoro Cave damaged by hyena activity. 1), 2): Edge polish on unidentifiable bone splinters. 3), 4), 5): Acid dissolution by partial digestion of unidentifiable bone splinters (3, 5) and astragalus of Cervus elaphus siciliae (4); 5a) detail by SEM analysis (58). 6), 7): Tooth pits on phalanges of Cervus elaphus siciliae. 8): Tooth groves on a fragmentary antler of Cervus elaphus siciliae.

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Table 9 Long bone representation by NISP of the herbivore taxa from the B-I level of San Teodoro Cave. Element

Portion

Humerus

Complete Cylinder Distal þ shaft Distal epiphysis (unfused) Shaft fragment Complete Cylinder Proximal þ shaft Proximal epiphysis (unfused) Distal þ shaft Distal epiphysis (unfused) Shaft fragment Complete Cylinder Proximal þ shaft Proximal epiphysis (unfused) Distal þ shaft Distal epiphysis (unfused) Shaft fragment Complete Proximal þ shaft Distal þ shaft Distal epiphysis (unfused) Shaft fragment Complete Cylinder Proximal þ shaft Distal þ shaft Distal epiphysis (unfused) Shaft fragment

Radius

Femur

Tibia

Metapodials

Total

M-size Ung. 3 3 10 4 1 2 2 8 2

Bovids

1 1

2

1

8 1 3 1

Elephant

1 1 1 1 1

1 2 3

1

2 1 3 12 3

1 1

2 18 5 14 18 10

3 5

15 157

2

2

2

21

Total

%

3 6 11 4

11.5 23 42.3 15.3

2 2 2 8 2

7.7 7.1 7.1 28.6 7.1

8 2

28.6 7.1

4 2 1 1 1

14.3 14.3 7.1 7.1 7.1

3 3

21.4 21.4

3 4 3 14 3

21.4 13.8 10.3 48.3 10.3

5 23 5 16 18 10

17.2 26.4 5.7 18.4 20.7 11.5

15

17.2

6

deciduous teeth, which have intact roots, and all the germs of permanent teeth, probably belong to cubs that were dead in situ. Gnaw marks are present on all cranial and postcranial remains. Most cubs of extant spotted hyenas are born into isolated natal dens, used by only one mother, and subsequently transferred by their mothers to the clan’s communal den, where cubs up to 9e15

Fig. 13. Distribution of complete and fragmentary coprolites from the B-I and B-II levels of San Teodoro Cave.

Fig. 14. Complete coprolites from San Teodoro Cave.

months old are present together with the adults (Boydston et al., 2006; East et al., 1989; Kruuk, 1972; Lansing et al., 2009; White, 2007). At the communal den young cubs begin to learn their social rank in the dominance hierarchy (Engh et al., 2000; Holekamp and Smale, 1993, 1998; Smale et al., 1993). Currently, communal dens are represented by burrows often having multiple entrances and chambers, leading to a network of tunnels (Kruuk, 1972; Mills, 1990; Pokines and Kerbis Peterhans, 2007), while cave dens are relatively uncommon and frequently formed by one or more outer chambers large enough to house adults and inner chambers or narrow cracks or tunnels accessible only to cubs (Henschel et al., 1979; Pokines and Kerbis Peterhans, 2007; Skinner et al., 1986). In modern spotted hyena dens, the presence of deciduous teeth and juvenile remains of hyena is an indicator of use as a communal den (Lansing et al., 2009). In communal dens, cub mortality can be due to either direct aggression (infanticide) or disruption of nursing activities by visitors or adult hyenas or sibling aggression (siblicide), probably correlated with a reduction in maternal resources (milk) (Golla et al., 1999; Kruuk, 1972; Holekamp and Smale, 1998; Smale et al., 1999; White, 2002, 2005, 2007).

Fig. 15. Size classes of complete coprolites from the B-I and B-II levels of San Teodoro Cave.

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Fig. 16. Length distribution of fragmentary coprolites from the B-I and B-II levels of San Teodoro Cave.

In extant spotted hyenas all deciduous teeth are replaced by permanent teeth at about 13e18 months (Van Horn, 2003); the San Teodoro Cave cubs’ remains probably belong to individuals that were dead by 13e18 months of age. Spatial distribution of cub remains in the innermost part of the San Teodoro Cave (b trench), very close to a narrow conduit along the eastern wall, partially filled with large fallen carbonate blocks, could suggest a partition of the den with inner and less accessible areas reserved for the cubs, similarly to the modern communal cave dens.

6.7. Elephant bones All the elephant remains from B-I, B-II, and B-III of San Teodoro Cave belong to P. mnaidriensis, an endemic reduced-sized elephant which was about 2 m tall. Remains are generally fragmentary and show the same scattered distribution and the same damages produced by hyena activity that have been detected on the other large mammal remains. During the 2003 and 2004 excavations, a group of complete and/or partially articulated elephant bones were collected from B-I in the b trench; they are represented by two, symmetrical pairs of tibiae and fibulae, a semi-complete mandible, a tusk fragment, a large scapula fragment, some vertebral fragments, a large coxal fragment, a patella, a femur shaft, and a metatarsal. On the basis of anatomical representation, development stage, and

Fig. 17. Crocuta crocuta spelaea cubs’ remains from the B-I level of San Teodoro Cave. 1): Right maxillar bone with deciduous premolars D3eD4 and germ of P4. 2): Left germ of P4. 3): Fragment of left mandible with deciduous premolars D3eD4. 4): Left germ of lower permanent canine. 5): Right deciduous premolar D4.

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spatial distribution, these remains can be related to a single, adult individual elephant, which probably entered the cave and may then have been killed or scavenged by hyenas after its natural death (Mangano and Bonfiglio, 2010). As cyclic occupation of the den is observed in extant spotted hyenas (Lansing et al., 2009), it is possible that elephants inhabited the cave during a phase of abandonment by hyenas. Cave utilization by elephants is attested in the modern and fossil record (Bowell et al., 1996; Hadjisterkotis and Reese, 2008; Redmond, 1982). According to Haynes (1987), extant spotted hyenas would be able to prey upon large elephants, but remains of this herbivore are very rare in modern dens and are mainly scavenged (Lansing et al., 2009; and references therein). On the contrary, Mammuthus remains have often been recovered in the Crocuta fossil  dens from Europe (Diedrich and Zák, 2006; Fosse, 1997; Guadelli, 1989). Crocuta crocuta spelaea had larger dimensions than modern spotted hyena (Klein and Scott, 1989) and was probably able to hunt the reduced-sized P. mnaidriensis.

7. Discussion and conclusion Taphonomic analysis of the large mammal assemblage from 1998 to 2006 excavations at San Teodoro Cave took into account the stratigraphic context of the unit B deposits. Many similarities, but also some differences, have been detected between the three recognized fossiliferous levels. All the levels, which are not strictly contemporary, are characterized by evidence of Crocuta crocuta spelaea occupation, represented by skeletal remains of this carnivore (which was recovered in both B-I and B-II but not B-III), coprolites, and distinctive damages on the bones. These taphonomic features, together with the patterns of anatomical representation and age distribution, are consistent with those recognized from fossil hyena dens of Europe and Africa and attest to the use of the cave by Crocuta crocuta spelaea clans. Data from B-I and B-II, where adult and juvenile remains of hyena, including deciduous teeth, have been collected, attest to the use of the cave as a communal den. The excavated area is undoubtedly small with respect to the whole extension of the unit B fossiliferous deposit, which is present all over the cave area, as attested by several test-pits, and further investigations are needed to demonstrate possible den partitioning; however, the recovery of hyena cub remains from a less accessible area located in the innermost portion of the cave is significant. The very high density of coprolites from the two main fossiliferous levels could suggest the presence of an extended “latrine area” in this sector of the cave, similar to those of extant spotted hyenas, which can defecate inside the dens or in the immediate vicinity and tend to have specific “latrine areas” where they indulge in social defecation (Brain, 1981). Differences in the density of fossil remains from B-I, B-II, and B-III must be considered relative to the different lithological features of the three levels. Based on lithology, the high concentration of remains from B-I could be the result of a facilitated and more ready accumulation process in a sector of the cave without boulders, while the lower density from B-II could be due to a more difficult accumulation of remains in the sediments because of the presence of very large boulders. The sporadic presence of remains from B-III could indicate an area that was less frequented by hyenas, due to the possible presence of standing water in the cave. The lithological difference between B-I, B-II, and B-III may be due to their different topographic locations within the cave or to different environmental and/or climate conditions depending on the humidity. The differential distribution of complete coprolites and small digested pieces from B-I and B-II is very likely due to more humid conditions in B-II, which have biased the preservation of

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coprolites and the frequency of isolated small digested pieces coming from the dissolution of droppings. Based on the stratigraphic data, the three sedimentation phases represented by B-I, B-II, and B-III, may be not contemporary and reflect an intermittent occupation of the cave by hyenas, similar to modern spotted hyena dens. Extant spotted hyenas have the lowest potential to accumulate bone remains with respect to the brown and the striped hyenas. In modern spotted hyena dens the rate of bone accumulation is extremely low and affected by den type (burrow dens show the lowest rate of bone accumulation), clan size, seasonal prey availability, and duration of den occupation (Lansing et al., 2009; Pokines and Kerbis Peterhans, 2007). A sporadic utilization of the cave by elephants also could be attested by the recovery of the partially articulated elephant skeleton. The presented taphonomic data from the three fossiliferous levels of the San Teodoro Cave deposits point to a long-term, probably cyclic, occupation by hyenas of the cave, which is confirmed as one of the largest Pleistocene hyena dens in Europe. Acknowledgements I wish to thank the anonymous reviewers who helped me in improving the quality of the paper. Thanks are due to the Superintendence to Archaeological and Cultural Heritage of Messina for permission to excavate and to conduct this research, and to Dr. G. Sabatino for the photomicrographs. Excavations were directed by L. Bonfiglio, and G. Mangano collaborated. Works have been funded by the European Community, the Italian National Research Program, the Messina University, the Messina Provincial Administration, the Acquedolci City Council, and the Rotary Club of S. Agata di Militello. References Anca, F., 1860. Note sur deux nouvelles grottes ossifères découvertes en Sicile en 1859. Bulletin de la Société Géologique de France 17, 684e695. Arribas, A., Palmqvist, P., 1998. Taphonomy and palaeoecology of an assemblage of large mammals: hyaenid activity in the Lower Pleistocene site at Venta Micena (Orce, Guadix-Baza basin, Granada, Spain). Geobios 31, 3e47. Binford, L.R., 1981. Bones. Ancient Men and Modern Myths. Academic Press, New York. Bonfiglio, L., Esu, D., Mangano, G., Masini, F., Petruso, D., Soligo, M., Tuccimei, P., 2008. Late Pleistocene vertebrate bearing deposits at S. Teodoro cave (northeastern Sicily): preliminary data on faunal diversification and chronology. Quaternary International 190, 26e37. Bonfiglio, L., Locatelli, E., Mangano, G., Masini, F., Pavia, M., Petruso, D., Sala, B., Surdi, G., 2009. Vertebrate biological events and Pleistocene biogeographic history of Sicily. International Conference on Vertebrate Palaeobiogeography, 28e29 settembre 2009, Bologna, Italy. Abstracts, 19e22. Bonfiglio, L., Mangano, G., Marra, A.C., 1999. Late Pleistocene hyaena den from a large cave deposits of Sicily (Italy). INQUA XV International Congress, 3e11 August 1999, Durban, South-Africa. Abstracts, 27e28. Bonfiglio, L., Mangano, G., Marra, A.C., Masini, F., 2001. A new Late Pleistocene vertebrate faunal complex from Sicily (S. Teodoro Cave, North-Eastern Sicily, Italy). Bollettino della Società Paleontologica Italiana 40, 149e158. Bonfiglio, L., Mangano, G., Marra, A.C., Masini, F., Pavia, M., Petruso, D., 2002. Pleistocene Calabrian and Sicilian bioprovinces. Geobios 24, 29e39. Bonfiglio, L., Mangano, G., Masini, F., Pavia, M., Petruso, D., Spigo, U., 2004. In: Teodoro cave, S., Bonfiglio, L. (Eds.), Quaternary eustatic fluctuations and biochronology of vertebrate-bearing deposits correlated with marine terraces in Sicily. 32th International Geological Congress, Firenze 20e28 agosto 2004, Field Trip Guide Book, pp. 17e21. Bowell, R.J., Warren, A., Redmond, I., 1996. Formation of cave salts and utilization by elephants in the Mount Elgon region, Kenia. In: Appleton, J.D., Fuge, R., McCall, G.J.H. (Eds.), Environmental Geochemistry and Health. Geological Society Special Publication London, vol. 113, pp. 63e79. Boydston, E.E., Kapheim, K.M., Holekamp, K.E., 2006. Patterns of den occupation by the spotted hyaena (Crocuta crocuta). African Journal of Ecology 44, 77e86. Brain, C.K., 1981. The Hunters or the Hunted? Chicago University Press, Chicago. Brugal, J.P., Fosse, P., Guadelli, J.L., 1997. Comparative study of bone assemblages made by recent and Plio-Pleistocene Hyaenids. In: Hannus, L.A., Rossum, L., Winham, R.P. (Eds.), Proceedings of the 1993 Bone Modification Conference. Hot Springs, South Dakota, pp. 157e187.

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