Monchot and Mashkour M., 2010. Hyenas around the cities. The case of Kaftarkhoun (Kashan- Iran). Journal of Taphonomy : 8 (1) 17-32.

P R O ME T H E US P R E S S / P A L A E O N T O L O G I C A L N E T W O R K F O UN D A T I O N (TERUEL) 2010 Journal of Taphonomy Available online at www.journaltaphonomy.com Monchot & Mashkour VOLUME 8 (ISSUE 1) Hyenas Around The City (Kashan, Iran) Hervé Monchot UMR 7194/CNRS, Département de Préhistoire, Muséum national d'Histoire naturelle Institut de Paléontologie Humaine, 1 rue René Panhard, 75013 Paris, France Marjan Mashkour* UMR 7209/CNRS, Département Ecologie et Gestion de la Biodiversité, Muséum national d'Histoire naturelle, Bâtiment d'Anatomie comparée, 55, rue Buffon, 75005 Paris, France Journal of Taphonomy 8 (1) (2010), 17-32. Manuscript received 19 September 2009, revised manuscript accepted 23 December 2009. This paper presents a taphonomic study of faunal remains of domestic and wild mammals found in a striped hyaena (Hyaena hyaena) den at Kaftar Khoun in the Karkars Piedmont near the city gate of Kashan (Iran). The Kaftar Khoun faunal assemblage is characterized by a low degree of bone breakage with many of the long bones complete, an intermediate frequency of tooth marking and a moderate amount of weathering damage to the bones. The species list, and mortality profiles of the main taxa, suggests that the hyenas collected remains of domestic stock that died naturally or were hunted/ scavenged (e.g. mules, donkeys), while the canids represent prey killed during conflicts over carcasses or were scavenged from road kills. The Kaftar Khoun den offers insights into the behaviour of striped hyenas in peri-urban environments. It shows that their behavioral adaptations are directly connected to modifications in their environment such that it may be considered as a commensal animal associated with human activities. Keywords: TAPHONOMY, EQUIDS, CANIDS, STRIPED HYENA, KAFTAR KHOUN, IRAN Consequently, in this study we examine the dynamics of animal bone accumulation due to carnivore activities as a means of understanding patterns of prehistoric site formation. Hyenas, like humans, hunt prey and/or scavenge carcasses. Their dens, like prehistoric human sites, are characterized by large accumulations of animal bones (e.g. Kerbis-Peterhans & Horwitz, 1992; Lam, 1992; Becker & Reed, Introduction Using modern case studies as analogues for exploring the past, even if it may carry some risk due to the chronological distance between the present-day and the past eco-environmental conditions, has undeniably demonstrated to prehistorians and archaeologists alike the value of such an approach (Brain, 1981). Article JTa095. All rights reserved. *E-mail: [email protected] 17 Modern striped hyena consumption 1979; Mendelssohn, 1985; Boneh, 1987; Qumsiyeh, 1996; Roberts, 1997). In Iran, the striped hyena was quite common throughout the nineteenth century A.D. and was found near big cities such as Tehran, but apparently was not greatly feared, as it does not attack humans (Polak, 1865). Unfortunately, there is very little statistical information on the demography of these animals on the Iranian Plateau, and increasing industrialization and the destruction of natural habitats has reduced the hyena's chances of expanding its zoogeographic range. 1993; Fosse, 1995, 1997; Brugal et al., 1997; Fosse et al., 1998; Lacruz, 2005; Pokines & Kerbis Perterhans, 2007; Andrews, 2008; Egeland et al., 2008; Kuhn et al., in press). Our case study is a faunal assemblage recovered from a modern striped hyena den complex located near the city of Kashan, in the centre of the Iranian Plateau. The striped hyena in Iran The striped hyena (Hyaena hyaena) is one of four extant species of Hyaenidae, the others being the spotted hyena (Crocuta crocuta), the brown hyena (Parahyaena brunnea) and the aardwolf (Proteles cristatus). The only extant species in Iran is the striped hyena, although the spotted hyena was the most common large carnivore in southwest Asia since the Early Pleistocene (e.g. Rabinovich, 2002; Monchot, 2006, 2008; Mashkour et al., 2009). Crocuta underwent a significant decrease in numbers during the last glaciation and disappeared from southwestern Asia, including the Iranian Plateau around 13-11 ka BP (Stuart, 1991; Stiner, 2004). Diet and foraging behaviour Adult striped hyenas have a body length of 40-47 in. (103-119 cm) plus an additional 10-18 in. (26-47 cm) for the length of the tail. Shoulder height is 23-37 in. (60-94 cm) and body weight ranges from 55-121 lb. (25-55 kg). Primarily a scavenger, the striped hyena is attracted to remains of kills made by other large carnivores, such as lions and spotted hyenas, but in general they avoid these animals. Smaller carnivores are either ignored or treated as prey. Striped hyenas are not known to actively hunt large or medium sized animals, but prefer very small prey, such as insects, lizards, birds, rodents, and rabbits (Kruuk, 1976; Mills, 1977; Roberts, 1997). Striped hyenas will also readily eat fruit. They drink regularly at water sources, but can survive off the moisture in the prey's body. They have been known to live in areas that are relatively dry. Striped hyenas often cache their food, like bones, pieces of flesh, or even some meat, in shallow holes dug with their snouts. The striped hyena forages individually and is rarely seen in groups. It does, however, associate in small family groups at the den. Geographical range and habitat The striped hyena’s current range extends across a large area, from the Near East to Central and Southern Asia, although in some regions they are virtually extinct (Harrison, 1968; IUCN, 1998). Today, in southwestern Asia, the striped hyena inhabits arid mountainous areas and piedmonts, avoiding true deserts, such as the steppes and semi-arid deserts of the Arabian Peninsula, the Negev desert of Israel, southern Iraq, southern, western, and northern regions of Afghanistan and Baluchistan (Harrison, 1968; Skinner & Ilani, 18 Monchot & Mashkour Piedmonts is a mountain chain stretching for over 100 km from Kashan to Ardestan and is mainly composed of igneous (plutonic and volcanic), pyroclastic and sedimentary tertiary rocks. KKH is located 8 km to the south of the prehistoric mound of Sialk in the city of Kashan at the edge of Dasht-e-Kavir, the Iranian central desert (Figure 1). The travertine is no longer active and the hollows which measure 50 to 70 cm height are interpreted as dens. KKH dens are located on a steep slope (60 to 70 degrees). The surface of the travertine has been deteriorated by weathering and is covered by a 2 to 5 cm layer of calcium carbonate, resulting in sparse vegetation cover around the dens (Heydari, 2004). The ground outside the hollows was strewn with bones and the long bones were identified in the field as having been eaten by hyenas. Each small hollow had obviously served as a refuge, such as a sleeping area. However, it was not possible to assess how many hyenas lived in Kaftar Khoun. During the third field season of excavation the second author and Saman Heydari Guran surveyed and collected bones from the dens (Mashkour, 2004). The area of the dens and surrounding slopes were gridded into 10 m squares and all the visible bones on the surface were systematically collected. Two years after the first visit and the collecting of the bones, no new evidence of hyena activity was observed, indicating that the hyenas had left the area. This was probably due to disturbance caused by the construction of an equestrian centre and a chicken breeding farm on the summit of the travertine. Striped hyenas are almost exclusively nocturnal, although they are occasionally seen lying in the sun or walking towards their lair in the evening or early morning. Roberts (1997) reports that striped hyenas spend the greater part of their active hours searching for food or moving to established foraging sites. They follow paths or roads very occasionally, almost invariably preferring to walk cross-country, covering new ground and investigating new bushes and boulders. Striped hyenas do not appear to have a set routine in moving around their range, although they regularly return to a good source of food such as a refuse tip, in some cases every night. A different route is taken every night, even when heading for the same lair. Consequently, there is very little path-formation, even close to the den (Kruuk, 1976). The den complex: Kaftar Khoun (N 33° 54E 51° 22) The site of Sialk, excavated originally by the French archaeologist R. Ghirshman (1939), is currently being restudied within the Sialk Reconsideration Project (SRP), directed by S. Malek Shahmirzadi (2003, 2004). In 2003, two members of the SRP team, F. Biglari and S. Heydari, surveyed the area within a radius of 30 km of the site in order to document early human settlements in the Iranian Central plateau, where little investigation had been done (Biglari, 2004; Heydari, 2004, Mashkour, 2004). During the prospecting, they found Kaftar Khoun (KKH), which literally means the “hyena house” in Persian1, which comprises a series of twenty small cavities hollowed out of a travertine formation in the Karkars Piedmonts (Heydari et al., in press). The Karkars In Persian, the hyena is called kaftar, in Baluchi aftar and in Pashto kog. Other vernacular names are taras in Sindhi, and targh or charkh as well as lagar bagar (bhagga) in Urdu, Hindi, and Punjabi (Frembgen, 1998). 19 Modern striped hyena consumption Figure 1. Location and general view of Kaftar Khoun (KKH) near the city of Kashan (Central Iran). category a minimum number of individuals (MNI) of 44 was calculated (based on the distal tibia). The good quality of bone preservation facilitated the measurement of the entire equid assemblage. We present here only the metacarpal data as a representative sample of the equid assemblage, as this bone is the most diagnostic bone for securing identifications. The method used for the taxonomic distinction is the log difference method (Simpson, 1941), which has been adapted for equid bones by Eisenmann (1979) and Eisenmann & Beckouche (1986). Among the 17 measurable metacarpals, eleven Species representation In April 2005, the authors had the occasion to reexamine the entire faunal assemblage collected from the dens, which is currently housed at the SRP archaeological base in Kashan. This assemblage comprises 1019 identified bones representing a minimum of 81 animals (Table 1). The KKH assemblage is largely made up of domestic species, and is dominated by the remains of equids (72.1%), followed by canids (12.2%). When all the equid specimens (n=735) were pooled into a combined equid 20 Monchot & Mashkour Domestic dog (Canis familiaris), golden jackal (Canis aureus) and wolf (Canis lupus) are positively identified. Four specimens of red fox (Vulpes vulpes) were also identified in the collection. Cattle (Bos taurus) is represented by 83 bones, with an MNI count of 8 individuals. Identified caprines (n=31) include domestic sheep (Ovis aries) and domestic goat (Capra hircus). Sheep and goat have been pooled into a combined category termed Caprini due to the problem of distinguishing between these two species in fragmented bone assemblages. Age profiles showed that of 13 caprine mandibles, representing a MNI of 8, 6 belong to adults while 2 are juvenile. Domestic camel (Camelus dromedarius) is represented by 14 bones, with a MNI count of two: one young adult and old individual. Alongside the equids, these are the largest species identified in the KKH assemblage. Tortoise is also represented in the collection. There are 9 specimens, which are bones could be identified as belonging to donkeys and 6 belonging to mules (Figure 2). However, because of the lack of a large and reliable comparative collection for mules and hinnies it was not possible to distinguish clearly between the two hybrids. The key finding is that in the den there are two sizes of equids: one smaller and definitely identifiable as donkey, and the other larger, compatible with hybrids and which is on average smaller than horse. On the basis of teeth, a MNI of 12 equids were identified, all adults. Of these, two represent young adults and six very old animals. Equids are often the main species contributing to the diet of hyenas (spotted and striped), both during the past and present, and in the Near/Middle East and Europe (e.g. Kerbis Peterhans & Horwitz, 1992; Villa et al., 2004; Diedrick & Zak, 2006; Marchal et al., 2009). The 124 canid bones represent a MNI of 13 individuals, making canids the second most common species in the dens. Figure 2. The logarithmic comparison of the equid metacarpals from Kaftar Khoun. The 0 line represents the standard animal (Equus hemionus onager) average measurements of 37 specimens (Eisenmann & Mashkour, 2000). 21 Modern striped hyena consumption We do not know how long hyenas had been gathering bones in KKH dens. In the western United States, Miller (1975) studied abandoned and broken bones on the plateau of southern Idaho, and concluded that they would mostly decompose in four years. However, the Kashan region is warmer and drier; abandoned bone would more quickly dehydrate, thus slowing bacterial action. We think bones would disintegrate more slowly, so that they would remain present and identifiable for more than four years and perhaps for considerably longer. On the whole, the KHH bone elements are relatively complete and readily identifiable. The distribution of bone weathering stages at the den is shown in Table 3. The majority are well-preserved bones (71.5%), with only 5.4% falling in weathering stages 4-5. This suggests that the den had been used for at least 10-15 years, assuming similar climatic conditions to those observed by Behrensmeyer (1978) in Amboseli National Park in Kenya. We can thus conclude that most of the bones were recently brought to the site: within 10-15 years of their being collected (this does not equate to the duration of functioning of the site) (Prendergast & Domínguez-Rodrigo, 2008). probably of the the spur-thighed tortoise, Testudo graeca, which lives in the area. Striped hyena (Hyaena hyaena) is represented by six cranial remains - two skulls and four mandibles. They testify to the presence of at least 4 individuals and indicate the practise, referred to by several authors, of hyenas sometimes eating their own kind, although it is unclear whether such individuals first died natural deaths or were killed by other hyenas (Becker & Reed, 1993). Humans: a fragment of a skull (parietal bone) attests to the presence of Homo sapiens in this assemblage. This is not a surprising find, as striped hyenas are well known as desecrators of graves (Kruuk, 1976, Horwitz & Smith, 1988, 1997). Weathering stage Weathering is defined by Behrensmeyer (1978) as the process by which the original microscopic organic and inorganic components of bone are separated from each other and destroyed by physical and chemical agents operating on the bone in situ, either on the surface or within the soil zone. Consequently, each weathering stage (WS) represents a discontinuous point in time along the continuous process of bone deterioration. According to direct observations on bones and descriptions based on easily observable criteria (macroscopic and mechanical, rather than chemical), we defined five WS (WS1 through WS5, increasing in the degree of weathering) in relation to the characteristics of the bones studied here (Table 2). These descriptions emphasize the fact that WS are not mutually exclusive categories, but rather arbitrary divisions operating within a continuous spectrum (Behrensmeyer, 1978; Lotan, 2000; Andrews & Whybrow, 2005; Todisco & Monchot, 2008). Body part representation Anatomical studies of the striped hyena (Spoor & Badoux, 1986) have shown that the musculature of the forelimb, neck and head are more strongly developed than in canidae or felidae. This is interpreted as an adaptation for lifting and carrying heavy prey. Consequently, the presence of whole equids and camel limb elements in the bones assemblage is not surprising. For the smaller taxa, like caprini and canidae, maxillae and mandibles were by the 22 Monchot & Mashkour Table 1. Census of Bones removed from the Kaftar Khoun area den (by species, number of identified specimens [NISP], minimum numbers of individuals [MNI] and percentage of species representation). NISP % NISP MNI Species Equids: Mule, Donkey, Hemione (Equus hemionus/E. asinus) Canids: domestic dog/Golden Jackal/Wolf (Canis familiaris/C. aureus/C. lupus) Domestic cattle (Bos taurus) Sheep/Goat (Capra hircus/Ovis aries) Domestic camel (Camelus dromedarius) Striped hyena (Hyaena hyaena) Red fox (Vulpes vulpes) Human (Homo sapiens) Tortoise Unidentified large herbivores 735 72.1 44 124 12.2 13 83 31 14 6 4 1 9 12 8.1 3.0 1.4 0.6 0.4 0.1 0.9 1.2 8 8 1 4 1 1 1 -- Table 2. Definition of weathering stages (Todisco & Monchot, 2008 modified from Behrensmeyer, 1978). WS 1 2 3 4 5 Definition of weathering stage Fresh bone aspect with no cracking or flaking. Surface smooth without cracks/fissures. Unweathered stage. Cracking parallel to fiber structure (longitudinal) in long bone. Incipient cracks on no long bone. Exfoliation started. Very limited surface weathering with a slight crazed appearance. Flaking of outer surface usually associated with cracks; flakes are long and thin with one edge attached to bone. Fissures growth in size, length and breadth. Advanced exfoliation. Bone surface coarse, rough and fibrous; large and small splinters loosely attached; weathering penetrates to inner cavities; cracks largely open. Spongy bone visible. Bone mechanically failing apart into pieces, very fragile, dusty. Bone generally unidentified, without cortical bone (only trabeculated, spongy bone). Bone severely deteriorated. Table 3. Summary of Weathering Stages (WS) for 975 KKH bone specimens. WS LOW WS MEDIUM WS HIGH WS WS 1 WS 2 WS 3 WS 4 WS 5 NISP 172 526 224 52 1 % 17.6 53.9 23.0 5.3 0.1 23 Table 4. Body part representation for different species given as number of identified specimens (NISP), minimum number of element (MNE) and minimum number of individuals (MNI) from KKH. The number of tooth marked specimens is given in the NISP column in parenthesis. Modern striped hyena consumption 24 *: calculations exclude values for isolated teeth. Monchot & Mashkour the Arad den in Israel, which shows no clear gradient by species size for skeletal elements and all species are over-represented by skull elements. For the equids at this site, for example, the skull elements represented 27.3% of the skeletal elements, with mandibles (n=22) and maxilla (n=25) for a MNI of 26 (Kerbis Peterhans & Horwitz, 1992). This body part distribution seen at the site suggests a selective transportation of skulls from smaller species, while those of large species were left behind at the kill/scavenge site. Another explanation for this pattern has been suggested (Horwitz, 1992) as being due to differential destruction of body parts in the different sized species: the resistance of bone to destruction is mediated by several factors which include its shape, internal composition, and size. An adult hyena could and would carry a whole sheep, goat or dog, but larger animals would necessarily have been dismembered by the hyena where the dying or dead equid was found and only a part carried off. The question is, do the ratios of skeletal elements found in the dens reflect what was found and transported by the hyenas, or does it reflect what they consumed? Considering the power of the jaws of hyenas in breaking off parts of bones, particularly the ends of long bones, the intact or near intact conditions of most of the dog skulls was immediately noticeable, and we concluded that a hyena’s jaw, even though equipped with large sharp carnassial teeth, did not have sufficient gape to allow it to break or crush the skull of a medium-sized dog, although they sometimes broke or chewed off the occipital condyles (Becker & Reed, 1993). far the most common elements represented, whereas the larger taxa (equids, cattle and camel) were better represented by post-cranial elements, especially limb bones (Table 4). For the equids, there are only 5 individuals represented by maxillae and 12 by mandibles; for the canids there are 15 by maxillae and 21 represented by mandibles; for the cattle there are 3 represented by maxillae and 8 by mandibles; and the caprine material includes one individual represented by maxillae and 8 by mandibles. Summing the post-cranial representation for the dominant taxa gives: for the equids, 33 individuals are represented by forelimbs and 44 by hindlimbs; for the canids, 9 individuals are represented by forelimbs and 9 by hindlimbs; for the cattle 3 individuals are represented by forelimbs and 5 by hindlimbs; and for the caprines only 1 individual is represented by both forelimbs and hindlimbs. Hindlimb representation is derived from the highest number of individuals represented by pelves, femora, tibiae or metatarsals. Forelimb totals are derived from the highest number of scapulae, humeri, radii and metacarpals. We can note an under-representation of trunks elements (vertebrae and costae) for all the species. Following Monchot & Horwitz (2002), we have summarized the skeletal representation data into body part classes (Table 5). This shows that, within hyena prey species, there is a further gradient which is size-related. Smaller animals are disproportionately represented by cranial remains compared with limbs, while larger ones are represented by greater numbers of limb elements. Similar findings were reported at the Wadi el Stbû site in Nubian Egypt (Becker & Reed, 1993) and by Cruz-Uribe (1991) or Brugal et al. (1997) for other hyena species, like brown hyenas (Parahyaena brunnea) and spotted hyena (Crocuta crocuta). As exception is Pattern of damage Despite the hyena bone crushing apparatus, no bones at KHH had been reduced to fragmentary 25 Modern striped hyena consumption Table 5. Summary in NISP (number of identified region. Skeletal group Equids Head 56 (7.6%) Axial 17 (2.3%) Forequarter 187 (25.4%) Hindquarter 72 (9.8%) Forefoot 171 (23.3%) Hindfoot 130 (17.7%) Foot 102 (13.9%) Total 735 (100%) specimens) for the main species of KKH by anatomical Dogs 56 (45.9%) 16 (13.1%) 19 (15.6%) 0 29 (23.8%) 1 (0.8%) 1 (0.8%) 122 (100%) Cattle 30(36,6%) 5 (6.1%) 7 (8.5%) 4 (4.9%) 15 (18.3%) 18 (22%) 3 (3.7%) 82 (100%) Caprini 17 (54.8%) 8 (25,8%) 2 (6.5%) 0 1 (3.2%) 1 (3.2%) 2 (6.5%) 31 (100%) Table 6. Equids long bone fragmentation summary (from Richardson, 1980; Todd & Rapson, 1988). Long bone Complete % Complete Proximal Diaphysis Distal Max. % Difference Humerus Radio-ulna Metacarpal Femur Tibia Metatarsal 13 29 49 0 24 61 17.8 35.8 76.6 0 24.5 69.3 2 22 6 0 6 7 9 8 0 12 10 0 49 22 9 35 58 20 62 51 58 35 82 81 61.03 0 0.03 100 46.42 8.72 %Difference = ([Col.1 + Col.2] − [Col.1 + Col.3])x100 (Col.1 + Col. 2) + (Col.1 + Col. 3) Table 7. Carnivore damages, gnawing trace (G) and tooth mark-puncture (TM) summary observed on equids limb bone (these marks were identified by naked-eye only). Proximal limb Humerus Radius + Ulna Metacarpal Femur Tibia Metatarsal G: 5 G: 8 G: 6 G: 2 G: 1 TM: 1 TM: 3 TM: 2 TM: 4 TM: 7 TM: 1 Shaft G: 1 G: 1 G: 1 Distal limb Total G: 3 G: 1 TM: 1 TM: 3 G: 4 TM: 1 G: 8 G: 9 G: 1 G: 7 G: 7 G: 1 TM: 1 TM: 1 26 TM: 2 TM: 6 TM: 3 TM: 5 TM: 8 TM: 1 Monchot & Mashkour & Kerbis Peterhans, 2007). Clearly, hyenas favour protuberances due to their association with areas of muscle and tendon attachment. The result is preferential damage of the soft, spongy trabecular bone. In contrast, the compact bone of the midshaft region suffers less due to its robusticity. As described for the Arad den (Kerbis Peterhans & Horwitz, 1992), the humerus and tibia are good examples to illustrate the damage pattern in the larger species: the proximal tibia (mainly comprised of trabecular bone) is destroyed whereas the distal part is relatively untouched (hard, compact bone). In contrast to the Arad den assemblage, no characteristic bone cylinders, which remain after both destroyed epiphyses, (Haynes, 1980; Hill, 1981; Brain, 1981; Binford, 1981; Bunn, 1983), were seen in the KKH assemblage. The absence of identifiable bones of young animals like those of the smaller sized sheep and goat, was probably largely due to the fact that such bones were more easily crushed and swallowed by the hyenas and had thus been preferentially destroyed. There is abundant literature from the past three decades showing the importance of traces left by hyenas in the formation process of osteological assemblages (e.g. Brain, 1981; Binford et al., 1988; CruzUribe, 1991; Fosse, 1995, 1997; Pickering, 2002; Blumenschine & Pobiner, 2007; Kuhn et al., in press). The low rate of observed traces at KKH is however not a unique case. Wezmeh, a Pleistocene cave in Iranian Kurdistan, shows very similar evidence of low bone modification (Mashkour et al., 2009), like the modern striped and spotted hyena sites of Djibouti (Ara Ide, Doumali, Dataganou, Fosse, unpublished data). Several factors can be put forward to explain this phenomenon: the quality and type of collection (surface collection vs. excavation), scraps. A measure of long bone fragmentation, developed by Richardson (1980) in a study of bone modification by African predator/ scavengers, examines percentage difference between representation of proximal and distal articular ends and can be useful in recognizing patterns of differential destruction of articular ends. The percentage difference value provides a technique to identify elements that have had one end preferentially destroyed or removed (Table 6). For the equid bone assemblage, the percentage of complete bones of forelimbs increases from the humerus down to the metacarpal, and we observe the same pattern for the rear limb from the femur to the metatarsal. The percentage difference for the articular ends of the forelimb bones decreases from proximal to distal ends, with large difference for the articular ends of the humeri, and almost no difference in the articular ends values of the radii and the metacarpals (Figure 3). The same tendency was observed for the rear limb. This pattern is not unusual and was found in many carnivore site e.g. wolf kills (Binford, 1981) but also an anthropogenic site (Todd & Rapson, 1988). At this point, it must be kept in mind that this approach merely documents patterns of differential destruction and does not directly indicate the processes responsible for them (Todd & Rapson, 1988:313). The damage patterns on a camel or equid from modern dens is comparable to those of an eland from prehistoric site. At KHH, there was evidence of tooth marks, and pitting on epiphyses, and a minority of the limb bones were gnawed at the epiphyseal ends (Table 7; Figure 3). The jagged and sharp edges remaining on these long bones are similar to descriptions of post-cranial remains from striped/spotted hyena dens in other regions (e.g. Sutcliffe, 1970; Henschel et al., 1979; Binford, 1981; Brain, 1981; Hill, 1981: Lupo, 1995; Pokines 27 Modern striped hyena consumption The striped hyena has shown their adaptability to occupy a very different habitat in modern times from their occurrence in archaeological sites. However, the limit of this adaptation and tolerance seems to have been reached today by the total destruction of their habitat by systematic intrusions. The effect of this is either migration to similar habitats, if they are not killed in the course of this migration, or by the investigation of new habitat perhaps higher up in the mountain beyond the reach of humans. Being situated only a few kilometres out of town, KKH was already threatened by various intrusions, which culminated at the site by the construction of an equestrian center and a poultry farm immediately in their habitat. The change in hyena behaviour is directly connected to environmental modifications, and the striped hyena appears now to be a commensal animal of human activities. The term commensalism derives from the Latin com mensa, meaning sharing a table and the definition is an interaction between two living organisms, where one organism benefits and the other is neither harmed nor helped. Originally it was used to describe the use of waste food from one species by a second species, like scavengers who follow hunting animals but wait until they have finished their meal. Intense human pressure (hunting) involving a quasi-disappearance of the wild fauna has meant that hyenas have had to change their natural subsistence behaviour, thus forcing them to prowl around cities, villages in search of abandoned carcasses, garbage heaps or weak animals in farmed herds (young, old or injured individuals). This phenomenon is not only found in Iran, but elsewhere in the Near and Middle East, and East Africa. Only the African natural the nature and size of the hunted/scavenged species, the type of environment (climate, landscape, interaction with other carnivores, etc.), the degree of human pressure, and finally the variable behaviour between different types of hyena (Hyena versus Crocuta). Clearly the nature of the site and the length of time carcasses were accessible to hyenas will affect the amount of surviving traces. Indeed KKH could be categorised more as a butchery and skinning site rather than as a den sensu largo. All these observations question the use of such traces as a straight forward criteria for identifying hyena activity on bones, and stress the importance of using modern examples to develop a better understanding of fossil european, asiatic and african den sites. Conclusions The present study consists of an analysis of a collection of bones, mostly of domestic mammals, collected by striped hyenas (Hyaena hyaena) via hunting or scavenging near the city of Kashan. Weathering data indicate that bones accumulated at KKH over at least a decade or more and that the bone assemblage is therefore an accumulation of many individual episodes of carcass transport. The equids, including mules and donkeys, that dominate the assemblage, were brought to the den from the vicinity of Kashan. Since they are mostly old animals at the end of their lives, it is likely that they were abandoned by their owners and became easy prey for hyenas or large canids. The canid remains may represent prey killed by the hyenas during conflicts over carcasses or scavenged after road kills (KKH is situated not too far from the highway Tehran-Kerman, Figure 1). 28 Monchot & Mashkour 29 Modern striped hyena consumption Becker, B. & Reed, C.A. (1993). Studies of the bone detritus of the striped hyena (Hyaena hyaena) at a site in Egyptian Nubia, and the interpretation of the bone breakage by striped hyenas. In (Clason, A., Payne, S. & Uerpman, H.P., eds.) Skeletons in her Cupboard. Oxbow Monograph, pp. 157-182. Behrensmeyer, A.K. (1978). Taphonomic and ecologic information from bone weathering. Paleobiology, 4: 150-162. Biglari, F. (2004). The Travertin formations of the northern borders of the Karkas Mountains: Geoarchaeological surveys. In (Malek Shahmirzadi, S., ed.). The potters of Sialk. Report N°3, Archaeological Report Monograph Series 5, Iranian Cultural Heritage Organisation, pp. 123-128 (in Persian). Binford, L.R. (1981). Bones, Ancient Men and Modern Myths. New York: Academic Press. Binford, L.R., Mills, M. & Stone, N. (1988). Hyena scavenging behavior and its implications for the interpretation of faunal assemblage from FLK 22 (the Zinjfloor) at Olduvai Gorge. Journal of Anthropological Archaeology, 7: 99-135. Blumenschine, R.J. & Pobiner, B.L. (2007). Zooarchaeology and the ecology of the Oldowayan Hominin carnivory. In (Ungar, P.S., ed.) Evolution of the human diet. The known, the Unknown, and the Unknowable. Oxford: Oxford University Press, pp. 167-190. Boneh, D. (1987). Mystical powers of hyenas: interpreting a Bedouin belief. Folklore, 98: 57-64. Brain, C.K. (1981). The hunters or the hunted? An introduction to African cave taphonomy. Chicago and London: The University of Chicago Press. Brugal, J.P., Fosse, P. & Guadelli, J.L. (1997). Comparative study of bone assemblages made by recent and Pleistocene hyenids. In (Hannus, L.A., Rossum, L. & Winham, R.P., eds), Proceedings of the 1993 bone modification conference, Hot Springs, South Dakota. Occasional Publication 1 Archaeology Laboratory, Augustana College, pp. 58-187. Bunn, H.T. (1983). Comparative analysis of modern bone assemblages from a San hunter-gatherer camp in the Kalahari Desert, Bostwana, and from a spotted hyena den near Nairobi. In (Clutton-Brock, J. & Grigson, C., eds.) Animals and Archaeology I. Hunters and their Prey. BAR International Series 163, Oxford, pp. 143-149. Cruz-Uribe, K. (1991). Distinguishing hyena from hominid accumulations. Journal of Field Archaeology, 18: 467-486. Diedrich, C.G., & Zàk, K. (2006). Prey deposits and den sites of the Upper Pleistocene hyena Crocuta crocuta spelaea (Goldfuss, 1823) in horizontal and vertical caves of the Bohemian Karst (Czech Republic). Bulletin of Geosciences, 81: 237–276. reserves seem to have escaped this situation. Nevertheless, it shows how plastic the hyena behaviour is and how well they adapt to changing times. Acknowledgments We particularly thank Dr. Malek Shahmirzadi for having encouraged the study of this collection within the Sialk Reconsideration Project. Mr Saman Heydari Guran and Fereydoun Biglari are particularly thanked for their interest in undertaking the collection and study of this material. Thanks to Hamid Fahimi who was, at the time, head of the Sialk Reconsideration Project Base - ICHTO in Kashan for his help in organising our stay in Kashan. Also, we thank Mr Abbass Etemad for his help in providing information about the old city of Kashan. Thanks are due to Dr Véra Eisenmann who has advised on the specific identification of the Equid bones. Thanks to Michel Coutureau for the drawing and L.K. Horwitz for the helpful comments. Dr. Robin Bendrey and Dr. Mark Beech are thanked for providing comments and corrections on the text. 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