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from the Early to the Late phase. The proportion of ungulate species captured by Natufian hunters was partially determined by local availability and thus influenced by environmental conditions. The only site in which gazelles had reduced importance is Ain Mallaha, located in a riparian habitat that likely supported different animal communities than the surrounding Mediterranean forest. Other ungulate species, in particular wild goat {Capra aegegrus), played more important roles in human diets in arid regions of the Jordanian and Negev deserts (Table 10.4).

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Table 10.4 Proportion of ungulate species represented in Natufian faunas.

Gg = Gazella gazella. Ov/cp = Ovids and Caprids, Bp = Bos primigenius, Cc = Capreolus capreolus. Dm = Dama niesopotamica, Ce = Cervus elaphus, Cer\' = Cervidae, Eq = Equus sp.. Ab = Alcephalits biicephalus.

Despite their general dominance in the Natufian period, the ratio of gazelles in

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1999, 2000). The routine capture of a diverse array of small reptile, bird and mammal species has long been recognized in Natufian assemblages (cf. Bar-Yosef and BelferCohen 1989; Byrd 1989b; Davis et al. 1994; Tchemov 1993a, 1993b) and is generally consistent with the prediction of Flannery's (1969) Broad Spectrum Revolution (BSR) hypothesis. Recently, several researchers have questioned the value of the BSR hypothesis, arguing that prey diversity changes little from the Middle Paleolithic to the Natufian period (Bar-Oz et al. 1999; Neely and Clark 1993; Edwards 1989).

Nonetheless, the high proportions of small game quantified at Hayonim Cave, Hayonim Terrace, el-Wad, and Hilazon Tachtit in this study far exceed proportions in all earlier Paleolithic assemblages (see Figures 7.1 and 7.13), and there is a demonstrable trend toward more even dependence on the three categories of small prey, an indication of expanding dietary breadth. Stiner et al. (1999, 2000) show that small game, particularly slow-moving or sessile t'pes, were important contributors to Paleolithic diets in the Mediterranean region from the Mousterian onward. Birds become important prey during the early Upper Paleolithic, but the proportion of quick small game in human diets -birds and hares ~ increased most dramatically in the Natufian period (see Figures 7.1 and 7.2 for the Wadi Meged sequence). The small size of the new prey precluded them from replacing ungulates as the primary meat source, but high relative abundance indicate an increasing human dependence on lower-ranked small resources in habitats directly surrounding Mediterranean Natufian sites (see Figure 7.1). Unfortunately, the abundance

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assemblages (but see Pichon 1984; 1991; Tchemov 1993a), but hopefully more ambitious inter-site comparisons will become possible in the future.

Although diversity measures based on species-based counts have failed to detect it, clearly something wis up with small game use towards the end of the Late Pleistocene.

Part of the problem appears to be the level of inquiry, that is the tools used to detect change. How we rank prey and define the "broad spectrum economy" strongly affects the patterns we identify in the archaeological record. Recently Stiner (2001; Stiner et al.

1999, 2000) has argued that small prey should be grouped and ranked according to their predator evasion strategies (slow-moving, high-ranked small game types such as tortoises and many shellfish; fast low-ranked birds; and fast low-ranked small mammals), and then measured for evenness among types. When this technique is applied to Paleolithic faunas from the circum-Mediterranean region, it is clear that diets began to expand fi-om the Upper Paleolithic onward, but most of all in the Natufian. Natuflan faunas generally stand apart from earlier assemblages in the remarkable evenness in small game types when ranked by cost of capture. Yet this pattern is not true of every Natufian assemblage: it is most apparent in the large sites of the Mediterranean zone of the Early Natufian phase. The balance of small game types declines after the transition to the Late Natufian, returning in scale to that observed for the Epipaleolithic Kebaran.

Gazelle Mortality Profiles As first argued by Davis (1983), and confirmed in many Natufian assemblages

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Mediterranean Natufian sites show consistently high proportions of juveniles, particularly in comparison to earlier Paleolithic assemblages. The strong juvenile bias in Natufian assemblages crosscuts many other archaeological variables, including site size and time period. In Chapter 8 it was shown that inflated proportions of juveniles are expected in prey populations subjected to persistent, heavy hunting pressure. The greater the harvesting pressure, the more pronounced the expected juvenile bias. This is not the only way juvenile bias can come about (i.e., Nunamiut skin hunting in spring, Binford 1978, Stiner 1990), but in the Natufian case it occurs everywhere and all of the time, indicating sweeping change in predator-prey relationships for the entire period. This exploitation pattern was not sufficient to drive gazelles to extinction — they shifted instead to a remarkable variety of other resources — some with great intensity such as hares and cereals.

Average Body Size of Hunted Tortoise Populations Like the age profiles of gazelles, the average body size of hunted tortoise provides an indirect indication of human harvesting pressure, since large specimens are both more visible and expected to be preferred when encountered. Because individual tortoises and their populations grow very slowly, increased harvesting pressure is expected to cause a reduction in the average body size of tortoise populations (see Chapter 8). Trends in tortoise body size were investigated earlier using the minimum medio-lateral breadth of the humeral shaft, which preserves well in the archaeological record. Measurements of

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sample includes the Early and Late phases from Hayonim Cave and the Late phase at Hilazon Tachtit. Comparative measurements are also available for the entire Paleolithic sequence of occupation at Hayonim Cave, and the Kebaran layers from Meged Rock Shelter (Stiner et al. 1999, 2000). The results show that the average breadth of tortoise humeri from Natufian assemblages are similar regardless of their origin in the Early or Late phase, Natufian tortoises as a group are also somewhat smaller than those from all preceding Paleolithic assemblages (see Figure 8.11). This means that the minimum pressure required to keep them small was continuous throughout the Natufian period.

Implications of Resource Pressure During the Natufian Period The three lines of faunal evidence reviewed above point to two major conclusions.

First, Natufian hunters in the Mediterranean zone exerted sufficient pressure on highranked prey (tortoises and gazelle) to affect, though not destroy, their populations on a regional scale. Second, human hunting pressure on high-ranked prey populations was stable throughout the Natufian period. It was not reduced in response to increases in population mobility during the Late Natufian phase. Evidence for greater mobility among small game types in the Late Natufian instead comes from a rising emphasis on tortoise, among small game types, in contrast to the situation in the Early Natufian, but much like that of the Kebaran.

The depression of gazelle populations ~ a decrease in the average age of hunted gazelles in the Mediterranean area in comparison to preceding Paleolithic periods ~ co

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Natufian archaeofaunas. Increased proportions of small game indicate that the Natufians were meeting more of their meat requirements with animals available in the local environments immediately surrounding their habitation sites. This suggests that during the Natufian the cost/benefit ratio of acquiring ungulate species increased in relation to that of capturing small local prey. Search costs for gazelles rose owing to decreased availability and/or constraints imposed by territorial restrictions created by human social or ecological pressures. The high percentages of juvenile gazelles at many core Natufian sites indicate depression of gazelle populations on a regional scale. The proportions of juveniles indicate that gazelle populations were pushed into a state of continuous growth, an extreme condition, suggesting that average annual mortality was higher than in preceding Paleolithic periods. Other possible explanations were discounted in Chapter 8.

Finally, the depression of tortoise populations is also indicated by a small decrease in the average body size of Natufian tortoises from Hayonim Cave and nearby sites in comparison to earlier Paleolithic periods (see Figure 8.11). The stability in average tortoise body size across the climatic extremes of the Younger Dryas indicates that climatic change is not responsible for sustained diminution. The reduction in body size is instead interpreted as a signal of sustained predator pressure on tortoise populations throughout the Natufian period. The implications for tortoise size reduction are more geographically localized than the age data for gazelles, however, since tortoises live out their lives in smaller areas. People continued to take tortoises whenever possible.

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small game. The proportion of tortoises to quick small game increases in the Late Natufian sites, meaning less intensive site occupation and associated increases in human population mobility. Ultimately, it is a shift in the relative frequencies of tortoises among small game that provides the key to understanding major population shifts in the Late Natufian.

No evidence for the absolute intensity of plant exploitation is available. Increases in the proportion and diversity of groundstone, the introduction of sickle blades, and a marked increase in the attrition of human teeth, at least in comparison to Mousterian populations (Smith 1991), indicate that the Natufians increased their dependence on small, labor-intensive plant resources such as cereals and nuts. Though the intensive use of local plant resources ultimately allows increased energy yields per unit area, a significant decrease in foraging efficiency should be experienced (Wright 1994), since more energy must be invested for each unit of energy returned. Similarities between Early and Late Natufian groundstone and sickle blade assemblages suggest that the intensity of plant exploitation was roughly the same in the Early and Late phases. The difference between the Early and Late Natufian is principally one of declining human population density.

It has been suggested that fairly intensive occupation continued in the Mediterranean zone during the Late Natufian (Bar-Yosef and Belfer-Cohen 1989, 1991b;

Henry 1989, but compare Belfer-Cohen 1991b; Belfer-Cohen and Bar-Yosef 2000; Byrd

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intensively occupied than those in surrounding areas, the results presented here suggest a clear reduction in the intensity of site use during the Late Natufian at least at in the Mediterranean zone. There are no examples of Late Natufian sites in the core area or elsewhere in the southern Levant that match the intensity of Early Natufian occupations at Mediterranean base camps (e.g., Ain Mallaha, Hayonim Cave, el-Wad).


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This research has generated a series of hypotheses on human demography, site use intensity, hunting pressure, and settlement and subsistence strategies during the Natufian period. Most importantly, it provides an empirical test based foremost on the relative proportions of small game animals that can distinguish variation in human site occupation intensity, diachronic and synchronic shifts in human demography within the Natufian period, and its implications for many other facets of cultural change.

The preceding archaeological evidence and published interpretations encourage a synthesis of current hypotheses on cultural change within the Natufian period. Together, the evidence presented above demonstrates three general trends. First, there is a decrease in site use intensity fi-om the Early to the Late Natufian phase, particularly within the core Mediterranean area. This interpretation is supported by the return to a nearly exclusive focus on high-ranked game (tortoises and gazelles), changes in internal site organization,

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camps. A second, related implication is the shift toward increased mobility during the Late Natufian phase in the Mediterranean area. This interpretation is based primarily on changes in settlement pattern, including the founding of new settlements along the border of the Mediterranean zone (Bar-Yosef and Meadow 1995; Goring-Morris 1987), and on evidence for reduced site use intensity. These two conclusions have also recently been proposed by researchers studying other Natufian data classes such as graves, and settlement patterns (Bar-Yosef and Belfer-Cohen n.d.; Belfer-Cohen 1988, 1995; BelferCohen et al. 1991; Byrd and Monahan 1995; Valla 1998). Third, the data on resource extraction indicates that the Natufians intensively harvested wild plants and animals.

Pressure was exerted to the point of causing resource depression in high-ranked, prey species (gazelles and tortoises). Although little direct evidence for plant exploitation exists, secondary evidence provided by plant harvesting equipment suggests that the Natufians invested significant amounts of energy to collect and prepare small, energy rich foods such as grains and nuts for consumption. Of particular interest in the case of both fioral and faunal resources is the tenacity of the Natufian subsistence system in the face of strong climatic fluctuations and settlement change from beginning to end. These observations have important implications for understanding the nature of the Natufian adaptation and the transition to agriculture.

Was there Population Growth in the Early Natufian?

In the Early Natufian, the Mediterranean forest was at its maximum extent, and

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