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«Item type text; Dissertation-Reproduction (electronic) Authors Munro, Natalie Dawn Publisher The University of Arizona. Rights Copyright © is held ...»

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animals represent less biomass per individual than ungulates, their living density per unit area is much greater than that of large game, increasing the likelihood that they may live or die naturally at archaeological sites. Many types of small animals are actually attracted to caves which provide shelter for nesting or burrowing. Several small animal species are also drawn to the niche created by human habitations due to the presence of food scraps, crevices and disturbed sediment and in the case of the Natufian, dry wall architecture. Finally, cave sites also attract a range of small predators including mammalian carnivores and predatory birds, that may feed on small animals and deposit their remains as pellets or scats in cave sites.

This analysis seeks to narrow the list of small animals in Natufian cave sites to those species potentially collected by humans. Very small species were excluded from consideration from the start, as they lack any evidence for human activity other than very low incidences of burning. These species include small Passeriforme birds, all rodents except squirrels, amphibians, and reptiles smaller than the lizard Agama stellio. The recovery of articulated skeletons of microfauna and concentrations of their bone elements beneath long-term roosts of bam owls within Hayonim Cave (E. Tchemov, personal communication 2000), suggest that these species are intrusive in the two sites as victims of natural death or deposition by predators, most likely bam owls {Tyto alba). Thus only species with an average body mass weighing more than one half kilogram, but less than two or three kilograms are considered as likely small prey of humans and systematically investigated for signs of human activity. At Hayonim Cave and Hilazon Tachtit, species

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Ophisaurus apodus), a variety of avian species (except small perching birds), hares {Lepus capensis), hedgehogs {Erinaceus europaeus) and squirrels (Sciurus anomaliis). A few avian species are larger than the 2-3 kg upper weight cutoff, including the large raptors such as eagles and vultures, as well as several species of waterfowl (e.g., Gms grus, Otis tetrax, O. tarda, Ardea cinerea and Anser albifrons), but are also incorporated within the small game group.

Potential Collectors of Small Prey Skeletons It is possible to construct a list of potential bone-collectors based partially on their dietary preferences. A predator's body size and feeding strategies limits the size of the prey that it can kill or consume, an important factor governing prey species choice. Many of the common predators in the southern Levant during the Natufian were small or medium-sized and focused primarily on small to medium-sized prey. The list of potential collectors of small game thus greatly exceeds that for ungulates or carnivores.

Birds of prey merit more thorough consideration here. The Levantine Falconiformes vary dramatically in size from the lesser kestrel {Falco naumanni, 95-130 grams) to the black vulture {Aegypiiis monachus), the latter can grow as large as 13 kilograms (Paz 1987). Upward of thirty varieties of falcons, hobbies, merlins, hawks, eagles and vultures nest and/or migrate annually through the southern Levant, the west Palearctic's most important Falconiforme spring and autumn migration route (Paz 1987).

Falconiformes that live in or regularly visit the Levant including three vultures (Aegypius monachus. Gyps fulvus. Neophron monachus) and two eagles {Aquila chrysaetos,

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vultures are scavengers and feed primarily on carrion but may transport bones with adhering flesh to young in the nest. Of the two eagles, the golden eagle (A. chrysaetos) is the only likely contributor to the two cave assemblages, since H. abicilla hunts and nests close to major water sources. Though the large raptors commonly nest and perch on rocky crags and cliff faces, which are available near Hayonim Cave and Hilazon Tachtit, they are unlikely to nest inside caves unless appropriate open ledges are available. Most of these species make enormous nests, averaging 1.5 meters in both width and height but reaching dimensions up to 5 meters wide and 3 meters high (Paz 1987), eliminating most cave interiors as suitable locations.

Ten species of owl inhabit the southern Levant today, but the only species large enough to have collected the prey of interest here are the eagle owl {Bubo bubo) and the bam owl {Tyto alba). The eagle owl is the largest owl in the southern Levant, weighing between 1 and 3 kilograms (Paz 1987). It feeds on small to medium-sized game, including partridges, doves, crows, waterfowl, fish, hedgehogs, bats, and rodents, and even several species of diurnal raptors. Bam owls are much smaller (250-310 g) and prefer correspondingly smaller prey, primarily rodents, songbirds, and insects, though they generally avoid reptiles and amphibians. What bam owls consider prey is determined more by prey body size than taxonomic affiliation (Tchemov 1993a). A rich database of pellet studies from Israel shows that modem bam owls repeatedly favor sparrows, voles and shrews (Paz 1987). Both bam and eagle owls are common in Mediterranean habitats and roost and nest in crevices in rocks and caves. They are

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Criteria for Separating Collectors Although only limited work on the composition and condition of bones deposited by avian predators has been undertaken in Southwest Asia, comparable data are available for similar and in some cases the same species in many other parts of the world (e.g., Andrews 1990; Cruz-Uribe and Klein 1998; Hockett 1996; Hockett and Bicho 2000;

Sampson 2000; Schmitt 1997). Though the owls inhabiting these diverse areas may not have identical diets, it is expected that they will target prey of similar body size, and generally leave the same types of damage on prey skeletons despite differences in the species consumed.

Large owls {Bubo bubo and Tyto alba) typically swallow their prey whole, digest the meat and then regurgitate the inedible bones and ftir in a dense pellet. As noted earlier the size of prey they consume is limited by their own body size, and specifically by the dimensions of their throat. The eagle owl consumes larger prey than the bam owl and may first prepare larger species by plucking or skinning before swallowing the remaining animal whole. Smaller, bam owls consume correspondingly smaller prey types. The owl's strategy of swallowing prey whole limits bone fragmentation, so that skeletal elements are regurgitated largely intact (Andrews 1990; Hockett 1996; Kusmer 1990; Saavedra and Simonetti 1998). Owls have relatively weak gastric acids, suited for the digestion of flesh but not bone, the latter of which is regurgitated (Andrews 1990;

Hockett 1996). Bones from owl pellets thus exhibit only weak if any corrosive damage and are at most marked by patches of polish and light pitting. In sum, the bones of

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largely complete, (c) display only light corrosion or polish if any, and (d) be represented by a high degree of skeletal completeness.

Falconiformes strip flesh from animal bones and consume carcasses in chunks.

The size of their prey is limited to species they are strong enough to kill and carry in flight, unless the prey is scavenged. In the Levant large eagles regularly kill and consume prey on the larger end of the small game spectrum (e.g., adult hares). Vultures consume carrion and therefore prey choice is not limited by prey body size. Small and mediumsized raptors target the same general types of small prey as bam owls. By stripping meat, raptors ingest fewer bones, particularly large ones (Hockett 1996), and thus fewer bones are exposed to digestive acids. Those that are swallowed are subject to intensive digestion, however the gastric acids of Falconiformes are very potent, especially in comparison to those of owls, and can heavily scar, pit, and thin the walls of prey bones (Andrews 1990; Brain 1981; Hockett 1996; Hockett and Bicho 2000; Schmitt 1997).

Raptors may also damage prey skeletons during capture, transport, or consumption by puncturing bones with their talons or beaks. Prey assemblages collected by raptors are expected to yield punctured crania, but fewer imprints on elements of the post-cranial skeleton (Cruz-Uribe and Klein 1998; Hockett 1991, 1996; Schmitt 1997). Golden eagles frequently behead their prey before transporting the carcass to the nest, and thus collect inflated proportions of postcranial body parts there. A few studies on the bone collecting behaviors of the golden eagle in North America show that the representation of prey body parts is significantly biased against heads and forelimbs, unlike those collected by owls

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raptors are thus expected to (a) be composed of small to medium sized prey, depending on the strength and size of the predator, (b) include high proportions of fragmented bones, (c) be frequently scarred by high incidences of corrosion, polish, and thirming from digestive acids, (d) be possibly biased toward the representation of prey hind limbs, and (e) exhibit punctures.

Though several carnivores may have played roles in the collection of the Hayonim Cave and Hilazon Tachtit faunas, and much work has been done to separate their signatures from those of humans, only general criteria that characterize carnivores as a group (Andrews and Nesbit-Evans 1983; Mondini 1995; Stallibrass 1984) and separate them from Falconiformes, Strigiformes and humans are relevant here. Much variation in the behavior of carnivores exists, especially in the range of prey body parts, taxa and age groups that they collect, but the skeletal damage caused by carnivores is more easily summarized than that of raptors or owls. Carnivore tooth marks include punctures, scoring (drag marks), crenelation, and percussion marks. Some carnivores like hyenas intentionally consume bone, but most of the carnivores of interest here ingest only small to medium-sized fragments and small compact elements as riders with the flesh they eat.

The bones of small prey are often consumed whole, even by small carnivores such as foxes and jackals. The digestive acids of most carnivores are strong enough to "polish", corrode, pit, and thin the walls of bone fragments, but only the gastric acids of hyenas can entirely dissolve bone (Horwitz 1990; Stiner 1994; Sutcliffe 1970). Bones collected by smaller carnivores (e.g., foxes, jackals) are therefore expected to show intermediate

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small tooth marks including punctures, scoring and crenelation. Distinct patterns in the representation of prey body parts and fragment size are also expected in assemblages generated by carnivores, but these vary by species and often overlap, and thus will not be discussed here.

Humans are obvious contenders for the role of bone collector, particularly when assemblages are recovered from sites rich in cultural remains, as is the case here. Still in general, the collection of small prey species by humans tends to be more contentious than for bones of large game, since small taxa are less likely to preserve obvious signatures of human processing such as cut marks and percussion fractures. Humans also tend to be more opportunistic and omnivorous than their carnivorous counterparts, employing a broader definition of what constitutes prey, particularly with respect to body size.

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Despite their generalist ways, humans do leave telltale signs on small animal remains. Cut marks, the classic indicators of human butchery, are sometimes left on small animal bones, but usually in such low frequencies that their absence is not sufficient to eliminate humans as potential collecting and modifying agents. Small animal bones rarely preserve evidence of percussion damage, due to the small size and gracility of bone structure. One exception is the tortoise, whose thick carapace and plastron segments fracture according to the same principles as mammalian cortical bone when struck with a hammer (see Figure 4.3). Though not infallible, burning is one of the most common human signatures on small animal remains, particularly localized burning on specific elements. Though burning may also result fi^om indirect associations between bones and fire after deposition, variation in the frequencies of burning among small game taxa may enable the separation of taxa burned post-depositionally from those cooked or disposed of in hearths.

Because humans are less likely to adhere to a single hunting and carcass processing routine than other predators, it is difficult to construct generalized expectations about their impact on small game skeletons. According to ethnographic research, human hunters tend to transport complete small game carcasses regardless of distance, since they represent easily transportable packages (e.g., Hudson 1991; Yellen 1991b). Humans may also consume small animal carcasses during excursions away from a home base. In these cases they may also selectively return certain skeletal parts as raw materials valued for secondary use as tools or ornaments, a point that will be taken up

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Damage on Small Game Assemblages Many general statements can be made about the impact of predators on small animal prey, yet clearly there is much overlap in the patterns created by different predator species. As palimpsests, archaeological records are notorious for mixing the signatures of different collectors, leaving the archaeologist with the burden of separation. The most effective way to address this problem is to cross-reference multiple lines of evidence (Brain 1980; Cruz-Uribe 1991; Haynes 1983; Lyman 1994; Noe-Nygaard 1989; Stiner 1994). Prey body part representation, age structures, and bone fragmentation are summarized in Table 4.2 and provide general guidelines for the identification of the relative contributions of different collectors to archaeological assemblages. The observed damage and skeletal representation of several categories of small fauna from Hayonim Cave and Hilazon Tachtit are presented in Tables 4.3 and 4.4.

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Table 4.2: Damage bone collectors are expected to leave on small game prey elements.

The small prey taxa evaluated in Tables 4.3 and 4.4 can be divided into three groups, based on profiles of bone damage and skeletal representation. Hares, partridges,

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