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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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detecting hunting intensity do not require elaborate aging systems. One only needs to divide the sample into adult and juvenile age groups. When the data allows, the adult component will also be separated into prime and senescent (old) groups. The intensity of hunting will be monitored by examining the proportion of juvenile gazelles and tortoises in archaeological assemblages. Because, reptilian and mammalian bone obey different patterns of growth, different methods must be applied to age them.

Age Determination for Gazelles Ungulate skeletal remains can be effectively aged using tooth eruption and wear sequences, as well as bone fusion. Tooth eruption and wear provide the most discrete and precise age stages for adults, but the robusticity of the technique is often hampered by small sample sizes, and, in the Natufian, heads of some prey apparently never reached the site. Because tooth samples are often small, bone fusion data is examined as an additional line of evidence.

Tooth Eruption and Wear Sequences Ungulate teeth erupt according to a genetically determined ontogenic plan and at species-specific rates. Through the study of individuals of known ages, the eruption sequence of the milk and adult dentition and their associated ages have been established for some ungulate species such as gazelles (Davis 1980a, 1983). The stage of development at the time of death can be determined most accurately if a complete tooth row is present, although individual teeth can also be evaluated using the degree of crown and root formation (Hillson 1986). The dentitions of ungulates are adapted for processing

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enamel that forai peaks and troughs and provide a rough grinding surface for food processing. Enamel is a much denser, tougher structure than dentine, thus the two substances wear at different rates. Ungulate teeth retain a sharp, uneven grinding surface throughout their functional life.

Food processing results in gradual attrition of a tooth's occlusal surface, exposing a distinctive sequence of patterns in the folds of dentine and enamel, known as wear stages. By combining eruption sequences with tooth wear, teeth can be assigned to a series of relative age categories with considerable precision especially in the younger age groups. Attrition occurs at a variable but predictable constant rate within a species (e.g., Gifford-Gonzales 1991; Severinghaus 1949). Animals of the same age are expected to exhibit similar stages of wear if they consume the same basic diet. Some researchers have attempted to assign absolute ages to wear stages using modem specimens of known age for comparison. Though provocative, and perhaps possible under certain tightly controlled conditions, rates of wear can vary in response to many factors, including but not limited to climate, sex and amounts of dietary grit. For this reason, as well as the sample size issue, wear stages are best collapsed into broader yet more reliable groups representing the more basic stages of life (following Stiner 1990, 1994).

In archaeology the use of tooth wear and eruption techniques was pioneered by Sebastian Payne (1973), who developed a series of relative wear stages for domestic sheep and goat in Anatolia. His notation is widely used and has been adapted by researchers to derive age wear stages for many other species (Grant 1982; Levine 1982;

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Appendix 3), based on comparative collections of mountain gazelle skulls of known ages housed in the mammalian collections at Tel Aviv University and the Hebrew University in Jerusalem, Israel. Stutz follows Payne's basic wear stages for sheep and goat but with attention to species-specific differences in attrition rate and the structural relationship between enamel and dentine for gazelle in particular. Due to small sample sizes and the simplicity of the data required to address questions of hunting intensity, Stutz's wear stages will be collapsed into three basic age categories following Stiner (1990: 312).

These stages include juvenile, prime, and old categories and correspond to the three major phases of an animal's full potential life span. The juvenile stage is devoted to growth and includes the period between birth and reproductive maturity in females. The prime stage of life is dedicated to reproductive activity and corresponds with an animal's peak reproductive years. Senescent animals are those past their reproductive prime, in a state of decline towards death.

No tooth element in the dental row is in continuous wear throughout a gazelle's lifetime. The eruption and wear of the lower deciduous fourth premolar (dP4) in combination with the fourth permanent premolar (P4) or the third permanent molar (M3), however, provide the closest approximation of a continuous sequence of wear spanning from an animal's birth to its death. The dP4 and M3 eruption and wear sequence is used here to construct gazelle age profiles. The M3 is chosen in lieu of the P^ since the division between prime and old animals is more difficult to assess using the wear stages for gazelle P4S. The Mj erupts just prior to the loss of the dP4, thus there is only a short

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wear stage and side of each tooth were cross-checked to ensure that no two teeth could have come from the same mandible. If an Mj and dP^ could have originated from the same individual, one was eliminated from the sample.

Stages of Bone Fusion The fusion of bone elements corresponds loosely to the transition from youth to adulthood. In the first phase of a mammal's life, energy is primarily devoted to growth.

As growth slows this energy becomes available for processes of maintenance and reproduction, the latter beginning in females around the time an animal reaches full size.

Unfused bones provide secure markers of juvenile animals, while fiised bones indicate that an animal is well on its way to adulthood, if not already there.

Fusion does not occur at the same time or in all long bone elements, but like teeth, it follows a predetermined species-specific sequence. Fusion sequences and ranges of absolute ages are provided for different species based on the study of modem individuals of known ages. Davis (1980a, 1983) estimated absolute ages for the fusion of gazelle long bones, obtained from a modem assemblage of wild mountain gazelle {Gazella gazella gazella) collected in Israel and curated in the Department of ESE at the Hebrew University of Jerusalem. Davis determined the ages of the skeletons using tooth wear and eruption stages that he developed from a second population of gazelles of known age in the Museum of Zoology at Tel Aviv University. This somewhat roundabout approach may involve minor errors, but Davis' data are of great value as a relative sequence of long

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Table 8.1: Age of fusion of gazelle long bone epiphyses, adapted from Davis (1980a: 132).

Despite overlap in the age of fusion among elements, bones fuse in a pre-determined order, recorded here from 1 to 13.

The bone elements, their estimated age at fusion, and the relative order of fusion are presented in Table 8.1. Davis (1980a: 132, 1983) provides age ranges for the fusion of most bones, as well as the sequence in which the elements fuse. For example, it appears that the distal calcaneum and distal femur fuse at the same time, since they are both assigned an age of 10-16 months. However, in gazelle the tuber calcis of the calcaneum always fuses before the distal femur. There is no question that all unfused gazelle elements in the collection belong to animals younger than two years of age, by which point gazelles have ceased growing and females have reached reproductive maturity.

The following skeletal portions were selected to examine the age structures of gazelle populations following Davis (1983); distal tibia, tuber calcis of the calcaneum, distal femur, and distal radius. These portions all fuse between 10 and 15 months of age

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assemblage, such as metapodials and phalanges, but the inclusion of these elements greatly increases the risk of double counting the same individual. For example, each gazelle skeleton contains eight of each of the three phalanges. It is possible that one individual could be represented by as many as eight times in the age profile of a single element. All of the elements selected for analysis here are represented by only two bones per skeleton, but data on metapodials are included to later check the consistency of the other results. To evaluate the reliability of the fusion results, the proportion of unfused specimens for each of the selected bone portions are determined. The results for each are then presented in the order that they fuse to check for consistency within the assemblage.

Elements identified as "gazelle" and "small ungulate" are included as gazelles in the age profile. It is assumed that the vast majority of small ungulate bones belong to gazelles, since they constitute 99% of all small ungulates identified to species in this study. It is possible that a few roe deer or wild goat bones were categorized as small ungulate and are erroneously included in the gazelle assemblage, but their rate of occurrence is so low (1%) that they are expected to have no discernible effect on the final outcome of the analysis.

Age Determination for Tortoises The growth of reptilian bone is sensitive to food availability and quality, thus growth does not occur at such predictable rates as those seen in mammals and birds.

Ossification occurs from only a single center, leavings no lines of fusion or good visual indications that the bone belonged to a juvenile or a reproductively mature adult.

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As explained earlier, hunting pressure not only increases the proportion of juveniles in a tortoise population, it also reduces the mean average age by decreasing the likelihood that an adult will complete it's potential life span. In continually growing animals, a decrease in the average age of the population should thus translate to a decrease in mean body-size.

The fact that female and male tortoises grow at different rates may bias this assumption slightly if the relative proportion of males and females varies dramatically from assemblage to assemblage. At this point there is no evidence to suggest that this is the case here.

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Figure 8.6: Tortoise {Testudo graeca) humerus and femur, with the narrowest point on the shaft indicated, where the medio-lateral breadth measurement is taken.

Sketch is slightly modified, and courtesy of M.


The humerus and the femur are the largest and most easily identified weightbearing elements in the tortoise skeleton. The size of the limb bones correlates with body mass (Wainwright et al. 1976; Gideon Hartman personal communication 2001). The humerus and femur shafts are particularly resistant to destruction by archaeological processes, whereas the proximal and distal ends tend to snap off and are often missing or

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viewed anteriorly or posteriorly, and the narrowest point is easy to locate. The mediallateral breadth is measured at the narrowest point along the shaft (see Figure 8.6; see also Klein 1994; Klein and Cruz-Uribe 2000; Stiner et al. 1999, 2000). The samples of humerus measurements are larger than those for femurs, probably because the femur shaft is narrower and breaks more often, close to the point where the measurement is taken.

When available, the results for both elements are presented.


Age data from gazelle and tortoise populations are presented below to monitor hunting pressure in the Levantine Paleolithic with special reference to the Natufian period. To explore Natufian hunting intensity, long-term trends in hominid hunting strategies from the Middle Paleolithic period onward are explored to set the Natufian within a broader evolutionary framework. We will then delve into a more detailed investigation of the faunas from Natufian sites, and at Hayonim Cave in particular.

Long-Term Change in Gazelle Hunting in the Mediterranean Levant To examine long-term change across the Paleolithic period in the Mediterranean Levant, published reports were consulted for data on gazelle age profiles and tortoise body size. Data on gazelle tooth eruption and wear is spotty for the Paleolithic period, most often due to small assemblage sizes and the lack of published material.

Gazelle Bone Fusion Results Much of the pre-Natufian data on gazelle bone fusion presented here are adapted

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gazelles across the Paleolithic sequence in the Levant. Davis monitored the proportion of juvenile gazelles in faunal assemblages by tabulating the proportion of unfiised epiphyses from five long bones that fuse between 10-15 months of age — the distal tibia, the tuber calcis of the calcaneum, the distal femur, the distal radius, and the distal metapodial (Davis 1983). Though this method has its problems, not least of which is the potential to count an individual multiple times, it allows the combination of small, otherwise unusable databases to construct large samples. Davis' (1983) original Natufian sample is bolstered here with data from the current study and from recent publications (see Table 8.2). To maintain geographic consistency, only sites from the Mediterranean zone and the Jordan Valley are included here.

Difficulties arise when attempting to compile and compare data sets collected by different researchers using variable reporting techniques. The most serious problems with fusion data are the failure of the author to define the age of a juvenile animal or disclose which elements were used to age the assemblage. Some researchers calculate the proportion of juveniles based on the total number of unfused elements in the sample, mixing elements that fuse at different ages. To overcome various pitfalls associated with data reporting, and to maintain consistency, two aging procedures are followed here.

Because small sample sizes are common in the comparative data, these procedures were chosen to favor sample robusticity over accuracy. First, the proportion of unfused metapodials are used to estimate the proportions of juvenile animals. Although metapodials may encourage double counting, they fuse within a tightly restricted time

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