AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY 133:1047–1059 (2007)

Role of Wild Plant Foods Among Late Holocene Hunter-Gatherers From Central and North Patagonia (South America): An Approach From Dental Evidence Valeria Bernal,1* Paula Novellino,2 Paula N. Gonzalez,1 and S. Ivan Perez1 1

CONICET, Divisio´n Antropologı´a, Facultad de Ciencias Naturales y Museo, Universidad Nacional de La Plata, La Plata, Buenos Aires, Argentina 2 CONICET, Laboratorio de Bioarqueologı´a, Museo de Historia Natural de San Rafael, San Rafael, Mendoza, Argentina KEY WORDS

wild plants; gathering; caries ratio; dental wear

ABSTRACT This study evaluates the role of plant foods in the subsistence of hunter-gatherers that inhabited the Central East, Northwest, and Northeast Patagonia (Argentina) during the late Holocene (ca. 3,000–500 years BP). The goal of the present study is to assess the temporal variation of dental caries ratio and wear rate in skeletal samples to ascertain if the biological information supports the dietary shift toward greater consumption of wild plant foods around 1,500 years BP, suggested by other types of evidence. The authors registered caries, antemortem and postmortem tooth loss, and tooth wear from eight samples belonging to hunter-gatherers from Patagonia for which chronological sequences from early late Holocene (ca. 3,000–1,500 years BP) up to final late Holocene (ca. 1,500–

500 years BP) are available. The results indicate that caries percentages in Patagonian samples fall within the range established for hunter-gatherers but there are significant geographical differences. In addition, caries ratio does not change significantly through time, so the amount of carbohydrates consumed seems to have remained fairly constant since 3,000 years BP. In contrast, there is a marked temporal trend toward the reduction of wear rates in the three areas, suggesting a faster rate in early late Holocene than in final late Holocene. These results would reflect a change to less hard and/or abrasive diets in the final late Holocene, probably owing to differences in food processing methods employed. Am J Phys Anthropol 133:1047–1059, 2007. V 2007 Wiley-Liss, Inc.

The studies about subsistence of prehistoric huntergatherers have traditionally emphasized the role of hunting, and consequently these groups have been regarded as meat-based whereas the role of plant foods in their diets tended to be ignored (Kelly, 1995; Hather and Mason, 2002). Since the middle 1960s, a reassessment of the subsistence of these societies led to acknowledge the vast range and variability of the resources consumed by many ethnographic groups and the importance of plant foods (Murdock, 1967; Lee and Devore, 1968; Hayden, 1981; Kelly, 1995; Binford, 2001). Although there is a growing trend toward the recognition of the role of wild plants for prehistoric hunter-gatherers, their relative contribution to their diets is often underestimated and the view of hunter-gatherer subsistence as largely meat-based has persisted. From a methodological viewpoint, this is partly due to the greater occurrence of faunal remains and the lower probability of recovering direct evidence of vegetables in archaeological sites of hunter-gatherers, since plant remains are generally poorly preserved (Hather and Mason, 2002). Therefore, the integration of archaeofaunistic, archaeobotanical, bioarchaeological (e.g., stable isotopes and dental caries), and technological evidences is necessary to elucidate both the past roles of wild plants and the probably subtle dietary changes in hunter-gatherer populations. In this study we focus on the role of plant foods in the subsistence of hunter-gatherers that inhabited the East Central, Northwest, and Northeast Patagonia, Argentina (Fig. 1), during the late Holocene (ca. 3,000–500 years BP). On the basis of several lines of evidence, the sub-

sistence of these populations was established to have been based mainly on hunting of guanaco (Lama guanicoe), with lesser and variable inclusion of rheas (Rhea americana, Pterocnemia pennata), wild plant foods and tubers, seafood, and other marine resources in the coastal groups (Gomez Otero et al., 2000; Kelly et al., 2001; Hernandez, 2002; Neme and Gil, 2002; Martinez, 2004; Neme et al., 2005). Likewise, a shift in subsistence strategies toward increased consumption of vegetables was suggested to have taken place between 2,000 and 1,000 years BP (Neme, 2002; Barrientos and Perez, 2004; Martinez, 2004; Neme et al., 2005). The greater reliance on plants was inferred based on (a) indirect evidences such as the amount of ground-stone tools and ceramic found in archaeological sites from final late Holocene (ca. 1,500–500 years BP), which could have been used for processing, cooking, and storing such resources (Moreno, 1874; Outes, 1907; Torres, 1922; Vignati, 1931; Ları´a,

C 2007 V

WILEY-LISS, INC.

C

Grant sponsor: CONICET, Universidad Nacional de La Plata. *Correspondence to: Valeria Bernal, Divisio´n Antropologı´a, Facultad de Ciencias Naturales y Museo, Museo de La Plata, Paseo del Bosque s/n, La Plata (1900), Argentina. E-mail: [email protected] Received 27 November 2006; accepted 26 March 2007 DOI 10.1002/ajpa.20638 Published online 6 June 2007 in Wiley InterScience (www.interscience.wiley.com).

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V. BERNAL ET AL.

Fig. 1. Geographic localization of the samples analyzed.

1961; Bo´rmida, 1964; Sanguinetti de Bo´rmida, 1999; Go´mez Otero et al., 1999, 2000; Neme, 2002; Martinez, 2004; Prates, 2004); (b) the availability of many plant species belonging to Espinal and Monte phytogeographic provinces (Cabrera and Yepes, 1960; Cabrera, 1976) indicate that they could have provided food for people (L’Heureux, 2002; Barrientos and Perez, 2004; Martinez, 2004); (c) the findings of macro-remains of plant foods in archaeological sites from Northwest Patagonia dated as final late Holocene that support the use of wild vegetables such as fruits of several Prosopis species and tubers (Lagiglia, 1957; Fernandez, 1988; Hernandez, 2002); (d) the Spanish chronicles that refer to the production of a type of bread made with \algarrobo" (Prosopis sp.) flour and the consumption of tubers by hunter-gatherers, at least in postcontact times (Vignati, 1941; Bo´rmida and Casamiquela, 1958–1959; Nacuzzi and Perez de Micou, 1983–1985; Hernandez, 2002). The studies concerning prehistoric subsistence in Patagonia have been mainly focused on archaeological evidence whereas systematic research of skeletal remains has not been undertaken across this region, with the exception of stable isotope analyses. Numerous analyses of carbon and nitrogen isotopes have been performed on several samples of human skeletal remains from the region studied to obtain more accurate information about the diets of these populations (Novellino and Guicho´n, 1999; Gomez Otero et al., 2000; Kelly et al., 2001; Martinez, 2004; Gil, 2003; Gil et al., 2006a; Gomez Otero, 2006; Martinez et al., 2006). The use of stable isotope methods provides direct evidence about diet composition and has proved to be a very useful way of establishing the proportion of marine and terrestrial resour-

ces and discriminating between C3 and C4 plants (see revision in Katzenberg, 2000; Larsen, 2002). The positive carbon and nitrogen isotope ratios in human bone collagen and apatite indicate that the subsistence of the people living in coastal zones of Central East and Northeast Patagonia was mainly based on terrestrial resources but also contained a variable amount (less than 35%) of marine items (Gomez Otero et al., 2000; Kelly et al., 2001; Martinez, 2004; Gomez Otero, 2006; Martı´nez et al., 2006). The dC13 values obtained for Northwest Patagonia would result from a diet made up of C3 resources and herbivores that consumed C3 and C4, and the results also suggest that a small proportion of cultivable C4 plants were consumed during the final late Holocene (Novellino and Guicho´n, 1999; Gil, 2003; Novellino et al., 2004; Gil et al., 2006a). However, the relative contribution of plants to the diet of Patagonian populations is difficult to establish based on these data because most wild plants occurring in this region are C3, and therefore carbon isotopes do not allow the distinction between consumption of C3 plants and C3-eating herbivores (Barberena, 2002; Gil et al., 2006a,b). In consequence, other evidences are needed to evaluate possible temporal trends in the consumption of wild plants. From a bioarchaeological standpoint, dietary information based on the analysis of dental wear and caries is of considerable value since it offers an independent check against reconstructions of prehistoric subsistence based on the analysis of floral, faunal, and technological evidence (Walker, 1978). Caries proportion and dental wear rates are informative about subsistence practices, both in terms of type of food consumed and food preparation techniques (Scott and Turner, 1988). Dental caries is a disease process involving the demineralization of dental tissue due to the production of organic acids from bacterial fermentation (e.g., Streptococcus mutans) of dietary carbohydrates (Larsen et al., 1991). Although the cause of dental caries is controversial, a vast dental literature documents the close association between caries and the amount of carbohydrates, sugar, and sticky nutrients consumed in the diet (Turner, 1979; Larsen et al., 1991; Lukacs, 1992; Larsen, 1997). In consequence, caries ratio can provide an index of the proportion of carbohydrate and protein intake, and thus is a valuable source for the reconstruction of dietary habits in prehistoric populations. Dental wear is a continuous long-lasting process that results from the mutual contact of opposite crowns as well as from their contact with food or abrasive material incorporated into food (Smith, 1972; Walker, 1978; Smith et al., 1984). Hence, dental wear is informative about dietary texture (hardness vs. softness) resulting either from food properties or the processing methods employed (Molnar et al., 1983; Powell, 1985; Teaford and Oyen, 1989). Considering the relevance of dental evidence for the discussion of prehistoric diets, the goal of the present study is to assess the temporal variation in dental caries ratio and wear rate in samples of prehistoric huntergatherer populations from Patagonia. Particularly, if a temporal change toward greater dependence upon plant food among hunter-gatherers took place around 1,500 years BP, caries ratio is expected to be greater in samples from after that time.

MATERIALS AND METHODS We collected data on caries, antemortem tooth loss (AMTL) and postmortem tooth loss (PMTL), and tooth

American Journal of Physical Anthropology—DOI 10.1002/ajpa

ROLE OF FOOD PLANTS IN PATAGONIAN HUNTER-GATHERERS wear from nine archaeological samples derived from Argentinean prehistoric populations (Fig. 1, Table 1). The samples are housed at Museo de La Plata (La Plata, Argentina), Museo Etnogra´fico \J. B. Ambrosetti" (Buenos Aires, Argentina) and Museo de Historia Natural (San Rafael, Argentina). The nine samples include a total number of 327 adult individuals; subadults were not included because they are underrepresented in the samples analyzed. Eight of the samples belong to hunter-gatherers from three areas of Patagonia (Fig. 1, Table 1) for which chronological sequences from early late Holocene (ca. 3,000–1,500 years BP) up to final late Holocene (ca. 1,500–500 years BP) are available. In addition, one sample of farmers from Central West Argentina (SJ-LLH) was included for comparative purposes (Gambier, 1993; Novellino and Guicho´n, 1997–1998). Sex and age determinations were done using standard osteological procedures described by Buikstra and Ubelaker (1994). Pelvic and cranial traits were used when both anatomical structures were available. However, as most of the samples belong to museum collections, the majority of individuals were represented by skulls only, and therefore the determinations were restricted to cranial traits. Sex estimation on the basis of pelvic structures was done using four morphological traits following Phenice (1969) and Buikstra and Ubelaker (1994): ventral arch on the ventral surface of the pubis; subpubic concavity; medial aspect of the ischio-pubic ramus, and greater sciatic notch. The traits scored in the skull were glabella, supraorbital margin, mastoid process, arcus superciliaris, crista supramastoidea, and overall aspect of mandible, which comprises the appearance of corpus and rami as well as size and strength (Buikstra and Ubelaker, 1994; Graw, 2001; Walrath et al., 2004). Age estimation from the pelvis was based on examination of the pubic symphysis (Brooks and Suchey, 1990) and auricular surface (Lovejoy et al., 1985). Age estimation from the skull was based on suture closure, using the lateral–anterior region because it is a more reliable predictor of chronological age than the vault region (Meindl and Lovejoy, 1985). Each individual was assigned to one of the following age categories: young adult (20–34 years, Y-A); middle-age adult (35–49 years, M-A); old adult (50+ years, O-A) according to Buikstra and Ubelaker (1994). Individuals were assigned as adults if they presented an obliterated spheno-basilar suture. The percentage of individuals assigned to each age class for all the samples is summarized in Table 2. There is a lower percentage of individuals in O-A class in most of the samples analyzed; for this reason, the comparisons of dental wear and caries ratio were restricted to the other two age categories. Dental caries and wear were evaluated macroscopically. A carious lesion was recorded only when demineralization had formed a distinct cavity in the tooth, excluding those pulp cavities without evidence of demineralization (Hillson, 2001). When comparing caries frequencies among samples, other factors than diet that might affect this variable must be considered, such as age groups, teeth classes, and different preservation of upper and lower teeth (Jacks and Lubell, 1996). The number of carious lesions present in a person is correlated with the number of years their teeth have been exposed in the oral cavity; thus differences in age distribution should be taken into account (Walker and Erlandson, 1996; Larsen, 1997; Hillson, 2001; Duyar and Erdal,

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2003). In addition, different tooth classes have different susceptibility to pathology, as the complex topography and grinding function of molars and premolars increases the likelihood of caries in these teeth over that in the simple, cutting anterior dentition—incisors and canines (Jacks and Lubell, 1996; Walker and Erlandson, 1996; Larsen, 1997; Hillson, 2001). Consequently, to use caries ratio as a diet indicator, these sources of variation should be controlled for, because differences among samples can result from their unequal age profiles and tooth class distribution, among other factors. In this study, caries frequency was calculated on the basis of surviving teeth instead of the number of individuals. This is the most common approach in skeletal series since it allows the inclusion of isolated teeth (Duyar and Erdal, 2003). However, the existence of numerous methods to measure caries frequency make it difficult to compare different studies (Hillson, 2001). Epidemiological studies of living people employ DMFT and DMFS (decayed, missing, and filled tooth or surface) indices for describing the dental caries experience of individuals or populations (Klein et al., 1938). The former is calculated by summing the number of permanent teeth that are decayed, missing or filled, whereas the second index is based on tooth surfaces. One of the greatest problems of these indices is the assumption that most missing teeth have been lost as a result of carious lesions, since periodontal disease and abrasion are also important causes of tooth loss among adults (Lukacs, 1995; Hillson, 2001). In addition, there is little evidence of dental work in archaeological samples; thus, no filled teeth are found (Hillson, 2001). Because of these reasons, DMF indices are not frequently used in archaeological surveys. In this study, caries ratio was calculated as the proportion of total number of caries recorded with respect to the number of observable teeth in each sample (Hillson, 2001). Because of dental caries is an age-progressive process, the comparisons in caries frequency among samples were made by age class, grouping the individuals according to the abovementioned categories. We evaluated possible influence of sex on caries ratio by means of a Chi square test. Dental wear is a complex process that involves three main types of dental substance loss, called attrition, abrasion and erosion or corrosion (Bell et al., 1998; Kaidonis et al., 1998; Grippo et al., 2004). Attrition may be defined as the wear produced by tooth-on-tooth contact between neighboring teeth or opposing teeth, and it produces wear facets on the occlusal surface or at contact points between teeth. Abrasion is the loss of tooth substance due to friction between a tooth and an exogenous agent. Corrosion refers to the chemical dissolution of enamel and dentine by endogenous or exogenous acids not produced by oral bacteria. These mechanisms occur frequently in combination during the dynamics of mastication, and therefore some attempts have been made to differentiate between mechanical and chemical wear (Bell et al., 1998; Kaidonis et al., 1998; Kieser et al., 2001a,b). In this study, dental wear was scored for all teeth using the ordinal scale proposed by Smith and Holly (1984) and Scott (1979a) for anterior and posterior dentition, respectively. These methods are widely employed in studies about oral health in prehistoric populations; hence the results obtained here will be comparable with those obtained by other authors. However, these qualitative methods record the result of the conjoint action of these mechanisms, and do not allow to

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American Journal of Physical Anthropology—DOI 10.1002/ajpa

Early late Holocene (3,000–1,500 BP) Final late Holocene (1,500–500 BP) Final late Holocene (1,500–500 BP)

Early late Holocene (2,500–1,500 BP) Final late Holocene (1,500–500 BP)

Middle/late Holocene (4,000–2,500 BP)

Final late Holocene (1,500–500 BP)

Early late Holocene (2,500–1,500 BP)

Chronology

F: female; M: male; I: indeterminate.

Total

Center west Argentina (Cuyo)

Central East Patagonia

Northeast Patagonia

Northwest Patagonia

Geographic area

SJ-FLH

CH-FLH

15—Valley of Chubut river 16—Calingasta 17—Angualasto 18—Pachimoco 19—Huaco

46 28

IGSB-FLH CH-BLH

344

29

74

30

RN-FLH

12—Valley of Negro river 13—Isla Gama 14—San Blas 15—Valley of Chubut river

10

32

20

75

n

RN-BLH

RN-M/LH

MZ-FLH

MZ-BLH

Abrev.

12—Valley of Negro river

1—Jaime Prats 2—Can˜ada Seca 3—Rinco´n del Atuel 4—Ojo de Agua 5—Cerro Mesa 6—Dique Villa 25 de Mayo 7—Puesto Aisol 8—Capiz Alto 9—Tierras Blancas 10—Las Ramadas 11—Laguna del Juncal 12—Valley of Negro river

Archaeological sites

176

13

43

26 15

19

8

14

11

27

M

134

16

29

19 13

11

2

18

9

17

Sex F

34

0

2

1 0

0

0

0

0

31

I

Domesticated plants (Zea mays, Phaseolus vulgaris and Cucurbita sp.)

Wild plants (Prosopis sp., Arjona tuberose)

Marine resources such as mollusks (Mytilus edulis, Aulacomya ater ater), mammals (Otaria flavescens) and fishes Lama guanicoe (guanaco), Rhea americana, Pterocnemia pennata

Domesticated plants (Zea mays)

Lama guanicoe (guanaco), Rhea americana, Pterocnemia pennata Wild plants (Prosopis sp., Arjona tuberose)

Lama guanicoe (guanaco), Rhea americana, Pterocnemia pennata Wild plants (Prosopis sp., Arjona tuberose).

Food resources available

TABLE 1. Geographic localization, chronology, and number of individuals by sex of the samples analyzed

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ROLE OF FOOD PLANTS IN PATAGONIAN HUNTER-GATHERERS TABLE 2. Total number y percentage of individuals assigned to each age class by sample

TABLE 3. Proportion of anterior and posterior teeth lost postmortem

Age class

PMTLa

Sample

Y-A

M-A

O-A

MZ-ELH

10 15% 5 25% 25 78% 19 63% 22 51% 12 42% 44 60% 4 14% 141

39 65% 14 70% 5 15% 10 33% 14 32% 14 50% 27 36% 19 69% 142

17 20% 1 5% 2 7% 1 4% 7 17% 2 8% 2 4% 5 17% 37

MZ-FLH RN-M/LH RN-FLH IGSB-FLH CH-ELH CH-FLH SJ-FLH TOTAL

discriminate the relative contribution of each factor to dental substance loss. Because dental wear is an age-related process, this source of variation must be controlled for to compare subsistence base among populations. This can be done easily by estimating rates of wear rather than mean degrees (Scott and Turner, 1988). Considering the natural eruption sequence of the permanent molars, in which there is an approximate 6-year interval between the eruption of first, second and third molars in all human populations, it is possible to derive age-independent rates of wear with acceptable confidence (Scott and Turner, 1988). Since wear rate is calculated based on molars, the anterior dentition was not included in this study. The principal axis analysis proposed by Sokal and Rohlf (1979) was applied to estimate wear rates (Scott, 1979b; Richards, 1984; Benfer and Edwards, 1991; LevTov Chattah and Smith, 2006). This method is more suitable than regression since it does not assume a causal relationship between variables (Sokal and Rohlf, 1979). The slope of the principal axis equation can be used as an indicator of the relationship between the adjacent molars, and therefore of the rate of wear. A high slope would indicate a rapid rate of wear and a low slope a slower rate of wear (Scott, 1979b). In this study, principal axis analysis was applied to wear rates between first and second upper molars whereas lower molars were excluded because mandibles were underrepresented in most analyzed samples. Third molars were excluded because their eruption time is more variable and they are frequently retained. In addition, first and second molars showing no exposed dentine or total loss of enamel were excluded. AMTL was recorded when a tooth socket displayed evidence of alveolar bone absorption, whereas PMTL was recorder when there was no evidence of bone absorption. Subsequently, the percentage of AMTL and PMTL in relation to the total number of sockets observed was calculated to establish possible biases in the number of caries observed. Both AMTL and PMTL percentages were estimated separately for anterior and posterior dentition. Finally, Chi square and Fisher tests were performed to estimate if the frequency of caries, AMTL, and PMTL differed significantly among samples grouped chronologi-

MZ-ELH MZ-FLH RN-M/LH RN-FLH IGSB-FLH CH-ELH CH-FLH SJ-FLH

293 101 168 152 308 152 378 166

(49.25) (49) (57.36) (49.94) (53.16) (54.87) (56.62) (57.44)

PMTLp 302 108 148 125 236 113 379 123

(50.75) (51) (42.64) (50.06) (46.84) (45.13) (43.38) (42.56)

Values in parentheses are in percentage.

Fig. 2. Percentage of antemortem anterior and posterior tooth lost corresponding to young adults (a) and middle-age adults (b) for each sample.

cally and geographically. The comparisons were made by age class for the first two variables, whereas for the last variable the comparisons were made by sample, grouping all individuals, since PMTL was not expected to be correlated with age. All statistical analyses were performed using R 1.9.1 software (Ihaka and Gentleman, 1996).

RESULTS The proportion of anterior and posterior teeth lost postmortem (PMTLa and PMTLp) is similar in all the

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V. BERNAL ET AL. TABLE 4. Comparison of antemortem tooth lost in posterior tooth group for young adults

MZ-ELH MZ-FLH RN-M/LH RN-FLH IGSB-FLH CH-ELH CH-FLH SJ-FLH

MZ-ELH

MZ-FLH

RN-M/LH

RN-FLH

IGSB-FLH

CH-ELH

CH-FLH

SJ-FLH

– 0.42 0.02* 0.30 0.02* 0.04* 0.02* 1

– 0.16 0.87 0.18 0.28 0.13 0.61

– 0.02* 0.86 0.57 0.9 0.11

– 0.04* 0.09 0.02 0.62

– 0.67 0.74 0.12

– 0.46 0.18

– 0.1

–

Probability associated to the Chi square test and Fisher test. * P < 0.05. TABLE 5. Comparison of antemortem tooth lost in posterior tooth group for middle-age adults

MZ-ELH MZ-FLH RN-M/LH RN-FLH IGSB-FLH CH-ELH CH-FLH SJ-FLH

MZ-ELH

MZ-FLH

RN-M/LH

RN-FLH

IGSB-FLH

CH-ELH

CH-FLH

– 0.08 0.00* 0.00* 0.10 0.00* 0.00* 0.00*

– 0.00* 0.00* 0.93* 0.00* 0.00* 0.00

– 0.11 0.00* 0.31 0.00* 0.14

– 0.00* 0.24 0.24 0.77

– 0.60 0.01* 0.21

– 0.00* 0.21

– 0.03

SJ-FLH

Probability associated to the Chi Square test and Fisher test. * P < 0.05.

samples, close to 50% for both tooth classes (Table 3). This indicates that the degree of preservation is similar in the skeletal samples analyzed. Conversely, in the case of AMTL there are some differences in the percentages of posterior and anterior teeth lost (AMTLp and AMTLa, respectively), with the percentage being lower for the latter in all the samples (Fig. 2). Likewise, the results show that the percentage of AMTLp increases with age, whereas AMTLa remains fairly constant at very low values (Fig. 2a,b). Because of this, the comparisons among samples by means of Chi square test and Fisher test were only made for the posterior teeth group. The results obtained for young adults indicate that MZ-BLH, MZ-FLH, SJ-FLH, and RN-FLH samples form a group characterized by few AMTLp, whereas the rest of the samples show higher number of posterior teeth lost antemortem (Fig. 2a). Table 4 shows that only some of these differences are significant. A slightly different pattern is observed for middle-age adults, the two samples from Northwest Patagonia (MZ-BLH, MZ-FLH) and IGSBFLH sample exhibit the lowest percentage of AMTLp and differ significantly from the other samples (Fig. 2b; Table 5). With regard to the frequency of caries, the results indicate that the sample from West Central Argentina (SJ-FLH) shows the highest values for both age classes analyzed (i.e., young adults and middle-age adults), followed by the samples from Central East and Northwest Patagonia (Fig. 3; Table 6). The samples from Northeast Patagonia present the lowest percentages of caries for both age classes (Fig. 3; Table 6). The results of Chi square and Fisher tests show that the group comprising MZ-BLH, CH-BLH, and SJ-FLH differs significantly from the rest of samples in the first age class (Table 7). For middle-age adults, only the samples from West Central Argentina and Northwest Patagonia differ significantly from the other samples (Table 8). There are no significant chronological differences between the two periods analyzed by area, either for Y-A or M-A (Tables 7

Fig. 3. Percentage of caries corresponding to young adults (a) and middle-age adults (b) for each sample.

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ROLE OF FOOD PLANTS IN PATAGONIAN HUNTER-GATHERERS TABLE 6. Total number of caries by age and sex for each sample Young adults Sample MZ-ELH MZ-FLH RN-M/LH RN-FLH IGSB-FLH CH-ELH CH-FLH SJ-FLH

F 5 0 2 1 1 6 17 5

M

(66) (73) (107) (80) (88) (155) (320) (53)

0 1 0 0 2 4 7 1

(33) (44) (81) (62) (130) (47) (279) (12)

Middle-age adults I

2 (29)

Total 7 1 2 1 3 10 24 6

(128) (117) (188) (142) (218) (202) (599) (65)

F 4 6 0 0 0 1 4 12

(65) (124) (15) (1) (62) (55) (133) (76)

M 4 5 0 0 1 3 7 6

(135) (122) (7) (80) (108) (144) (257) (95)

I

Total

8 (115)

16 11 0 1 1 4 11 18

(315) (246) (22) (81) (170) (199) (390) (171)

The number of teeth observed is indicated between the brackets. F, female; M, male; I, indeterminate. TABLE 7. Comparison of caries frequencies by sample for young adults

MZ-ELH MZ-FLH RN-M/LH RN-FLH IGSB-FLH CH-ELH CH-FLH SJ-FLH

MZ-ELH

MZ-FLH

RN-M/LH

RN-FLH

IGSB-FLH

CH-ELH

CH-FLH

SJ-FLH

– 0.04* 0.00* 0.02* 0.03* 0.50 0.45 0.24

– 0.62 0.70 0.57 0.04* 0.08 0.00*

– 1 0.37 0.01* 0.02* 0.00*

– 0.48 0.02* 0.05 0.00*

– 0.03* 0.06 0.00*

– 0.5 0.23

– 0.05

–

Probability associated to the Chi Square test and Fisher test. * P < 0.05. TABLE 8. Comparison of caries frequencies by sample for middle-age adults

MZ-ELH MZ-FLH RN-M/LH RN-FLH IGSB-FLH CH-ELH CH-FLH SJ-FLH

MZ-ELH

MZ-FLH

RN-M/LH

RN-FLH

IGSB-FLH

CH-ELH

CH-FLH

SJ-FLH

– 0.7 0.3 0.02* 0.00* 0.07 0.06 0.08

– 0.36 0.04* 0.01* 0.12 0.16 0.01*

– 1 0.89 0.64 0.56 0.09

– 0.69 0.25 0.14 0.00*

– 0.2 0.09 0.00*

– 0.69 0.00*

– 0.00*

–

Probability associated to the Chi Square test and Fisher test. * P < 0.05.

and 8; with the exception of MZ-BLH and MZ-FLH samples for Y-A). Finally, it is important to note that caries ratio decreases with age in the samples from Central East and Northeast Patagonian, whereas it remains constant in the other samples (Fig. 3). Table 9 shows the results of principal axes analysis. A temporal reduction of wear rate is observed within each region analyzed. The samples from early late Holocene have greater wear rates than the final late Holocene ones. The rates range between 20 and 30 in the first period, except for RN- M/LH with a wear rate value of 9.99. In contrast, the rates calculated for final late Holocene range from 1.34 to 7.37. The lowest value observed corresponds to the farmer sample (SJ-FLH).

DISCUSSION Dental caries is closely associated with the amount of carbohydrates, sugar, and sticky nutrients consumed in the diet; therefore, the proportion of caries may provide information about carbohydrate and protein intake ratios (Turner, 1979; Larsen et al., 1991; Lukacs, 1992; Larsen, 1997). Numerous studies show that the proportion of caries is relatively low among hunter-gatherers

whose diets contain substantial quantities of animal protein, but increase markedly when populations adopt carbohydrate-rich diets based on cultivated plants (Turner, 1979; Milner, 1984; Larsen et al., 1991; Hillson, 2000). According to Turner (1979), hunter-gatherers show a proportion of caries ranking from 0.0 to 5.3%, mixed-diet consumers show between 0.4 and 10.3%, and individuals whose economy is based on agriculture present from 2.3% to 29%. Our results show that the sample of agriculturalists (i.e., SJ-FLH) displays the highest percentage of caries (9.5% in young adults and 10.53% in middle adults). In hunter-gatherer from Northwest and Central East Patagonia the percentage of caries in young and middle-age adults ranges between 2.75% and 5.25%, close to the upper limit proposed by Turner (1979) for these societies. Conversely, the samples from Northeast Patagonia display lower percentages of caries (between 0.5 and 1.25%). Within some samples analyzed, the percentage of carious lesions is lower in middle-age individuals than in young ones (Fig. 3). Similar findings, which apparently contradict the generally accepted statement that dental caries is an age progressive process, were also found by other authors (Delgado-Darias et al., 2006). This can be partially explained by the fact that a part of

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V. BERNAL ET AL. TABLE 9. Results of principal axis analysis

Chronology Early Late Holocene (3,000–1,500 BP)

Final Late Holocene (1,500-500 BP)

Sample

Slope

n

RN-M/LH RN-ELH MZ-ELH CH-ELH CH-FLH RN-FLH IGSB-FLH MZ-FLH SJ-FLH

9.99 23.54 20.73 29.5 2.93 7.37 2.86 2.25 1.34

36 8 22 20 60 32 44 26 34

n, number of teeth employed to calculate the principal axis equation.

older individuals have lost more teeth, as documented by the greater percentage of AMTL in middle-age than in young adults (Fig. 2). However, it cannot be assumed that all AMTL is caused by caries, since multiple causes, such as wear, periodontal diseases, pulp chamber infection, and trauma, may also result in AMTL (Powell, 1985; Lukacs, 1992; Duyar and Erdal, 2003). Nevertheless, the low frequency of caries found in young adults, combined with the high wear rates, seems to indicate that the exposition of pulp cavity due to occusal wear was the main cause of AMTL among the hunter-gatherers analyzed. In contrast, the higher frequency of caries and lower wear in the farmers sample suggest that a large proportion of AMTL in middle-age adults would have been caused by carious lesions. There is an important variation in the percentage of caries among hunter-gatherers from different regions of Patagonia, suggesting probable differences in the amount of carbohydrates consumed. For comparative proposes we considered two cases from Patagonia that represent two very dissimilar situations: samples from populations from Tierra del Fuego whose diet incorporated a small proportion of vegetables and was primarily meat-based (Barberena, 2002; Yesner et al., 2003), and samples from Neuque´n province (North Patagonia) where it is known that Holocene hunter-gatherers included a large proportion of seeds of Araucaria sp. in their diets (Fernandez and Panarello, 2001; Kozameh and Barbosa, 1992). Previous studies of skeletal indicators in several samples from Tierra del Fuego show a percentage of carious teeth ranging between 0% and 1.2% (Perez-Perez and Lalueza Fox, 1992; Kozameh, 1993; Guicho´n, 1995). Such low frequency of caries is consistent with the fact that the available resources in Tierra del Fuego are marine (e.g., sea lion, seafood) and terrestrial (Lama guanicoe) animals whereas the amount of wild plants is negligible (Yesner et al., 2003). In addition, ethnographic and archaeological data indicate that Fueguian populations included a very small proportion of vegetables in their diets (Gusinde, 1982; MansurFranchomme et al., 1997–1988; Yesner et al., 2003). Conversely, a high percentage of carious teeth (30%) was found in a skeletal sample from Neuque´n, which is consistent with the consumption of Araucaria seeds that have a large percentage of carbohydrates (Fernandez, 1988; Kozameh and Barbosa, 1992). The comparison of these results suggests that the populations from Northeast Patagonia could have consumed a diet with a proportion of carbohydrates as low as that of the populations from Tierra del Fuego, whereas in the Central East and Northwest areas there was a higher consumption of carbohydrates.

As noted above, a temporal trend toward greater dependence upon plant foods among hunter-gatherers from North and Central Patagonian was suggested based on archaeological evidences. Under the assumption that the plant parts consumed are cariogenic, it would be expected that such processes result in a greater proportion of carious teeth in final late Holocene populations with respect to older ones. Although the marked increase in the prevalence of dental caries is generally used to infer the adoption and intensification of domesticated plants, it is important to consider the cariogenic potential for both wild and domesticated plants. Therefore, we considered the chemical composition of vegetables available for consumption in North and Central Patagonia, mainly for the species of genus Prosopis whose exploitation is well known from ethnohistorical evidence (Bo´rmida and Casamiquela, 1958–1959; Hernandez, 2002). Although the chemical composition of the fruits of different species might differ slightly, the information corresponding to Prosopis alba and Prosopis flexuosa shows that they have a large percentage of carbohydrates (more than 50%) with a variable proportion of fibers and proteins (FAO, 1993; Roig, 1993). Another factor to consider in the case of late Holocene populations from Northwest Patagonia is that isotopic evidence indicates that a small proportion of C4 plants, i.e., maize, was consumed. Maize grains are a highly cariogenic food, since approximately 70% of its weight corresponds to starch (FAO, 1993). This area is the South American frontier of Pre-Hispanic maize expansion, but isotopic information demonstrates that C4 resources were never a quantitatively significant part of human diets (Gil et al., 2006a). In base to ethnographic descriptions and archaeological evidence it was suggested that maize was not locally cultivated, instead hunter-gatherers obtained it by exchange with their northern farming neighbors (Gil, 2003; Gil et al., 2006a). Considering that the composition of the wild plants available in the region studied includes a large amount of carbohydrates, increased consumption of these items is expected to involve higher caries frequency. However, the percentages of caries did not change significantly through time within the three areas of Patagonia (Fig. 3; Tables 7 and 8). Conversely, our results suggest that the proportion of plant foods consumed would have been constant at least during the last ca. 3,000 years BP. The trend toward increasing reliance on plants among Holocene hunter-gatherers from North and Central Patagonia was proposed based on the presence of ground stone tools (e.g., mortars, \manos"—grinding slabs, \metates"—handstones) that could be used for processing the gathered plants (Go´mez Otero et al., 2000; Neme, 2002; Barrientos and Perez, 2004; Martinez, 2004). Ground stone tools are usually employed to study subsistence strategies and to infer shifts in prehistoric strategies under the assumption that such artifacts were designed for food-processing, although groundstone tools may have been used in non-food-related activities (Wright, 1994; Adams, 1999). Notwithstanding this fact, little work has been done to establish their functionality in Patagonian prehistoric populations. Thus, the ethnographic evidence available indicates that such artifacts were related to the preparation of plants as well as salt, fish flour and mineral pigments and the processing of a type of dry meat named \charqui" (Moreno, 1874; Bo´rmida and Casamiquela, 1958–1959). The few analyses about ground stone tools functionality were performed

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ROLE OF FOOD PLANTS IN PATAGONIAN HUNTER-GATHERERS on artifacts recovered in archaeological sites from Patagonia and a neighboring area (i.e., South Pampa), which was inhabited by hunter-gatherer populations biological and culturally related with those from the region under study (Barrientos and Perez, 2004; Perez, 2006). These studies indicate the presence of red mineral pigment (ochre) and microremains (i.e. silica phytoliths) of vegetables in \manos" and mortars (Gomez Otero et al., 1999; Babot et al., 2005; Bonomo, 2005; Tassara and Osterrieth, 2005; Zucol and Bonomo, 2005). Because ground stones could have multiple functions, their frequency of occurrence cannot be used as direct evidence to infer the amount of plant foods consumed (L’Heureux, 2002). On the other hand, the radiocarbon data available for mortars found in stratigraphic positions, at one archaeological site in South Pampa (Nutria Mansa I; Zucol and Bonomo, 2005) and one site in North Patagonia (Aquihueco´; Della Negra and Novellino, 2005; Della Negra, personal communication) indicate an antiquity of 3,500–3,000 years BP. However, chronology is not easy to estimate for most of these artifacts because they appear on the surface rather than in stratigraphic position, and consequently the moment when their use began to be widespread in the area is difficult to establish. The prevalence of caries found among the hunter-gatherer samples analyzed in this study could be related to the effect of occlusal wear, which could theoretically obliterate teeth fissures and might therefore remove fissures of the earlier erupting molars before they became carious as well as the carious tissue (Hillson, 2000). This suggests a negative relationship between dental wear and caries (Maat and van der Velde, 1987), assuming that wear rate happens very rapidly to outstrip the type of caries development that occurs deep in the fissures. In contrast, some authors have proposed that both variables are positively correlated, so that the rapid wear was an important factor intimately involved in the development of carious lesions (Moody, 1960; van Reenen, 1966). Finally, other studies suggest that dental caries and wear would be independent variables and that samples with different diets will show different relationships between these variables (Meiklejohn et al., 1992; Larsen, 1997). According to our results, occlusal wear rates changed considerably through time, whereas caries ratio remained constant. The results obtained show temporal variation in wear rates in the three Patagonian areas studied, suggesting a faster rate in early late Holocene compared to final late Holocene. Therefore, the low prevalence of caries in the latter period cannot be attributed to effect of occlusal wear but to low carbohydrate consumption. Wear rate is highly correlated with food abrasiveness and texture (hardness vs. softness) and the amount of abrasives incorporated into the diet, which would include abrasive particles from the air, grit from food processing through ground stone tools, cooking directly over a hearth or in ashes, sand, and grit associated with shellfish and other food obtained from litoral zones, among others (Walker, 1978; Scott and Turner, 1988; Lalueza Fox et al., 1996; Lev-Tov Chattah and Smith, 2006). Thus, variation in wear rates can be used as an independent line of evidence to evaluate shifts either in the type of food consumed or in the food processing methods employed (Smith, 1972; Walker, 1978; Smith et al., 1984; Gold, 2000). This is supported by experimental studies and by the observation of ethnographic

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groups whose diet is known. For instance, it was demonstrated by means of experimental designs that hard diets result in faster tooth wear than soft ones (Teaford and Oyen, 1989) and a diet incorporating significant amounts of stone-ground maize can indeed lead to accelerated rates of tooth microwear (Teaford and Lytle, 1996). The reduction of mechanical and abrasive stresses on dental structures that occurred in recent aboriginal populations due to the adoption of softer foods and changes in cooking styles is well documented (Molnar et al., 1983). The comparisons of populations undergoing subsistence type changes show a consistent pattern of decline in severity of wear from hunter-gatherers to agriculturalists (Molnar, 1971; Gold, 2000; see revision in Larsen, 1997). However, as Scott and Turner (1988) have noted, both hunter-gatherers and agriculturalists can show pronounced degrees and fast rates of wear. Furthermore, it was suggested that prehistoric agriculturalists may exhibit more severe dental wear because plant materials contain more abrasives than meat does (Larsen, 1995; Hillson, 1996). Some studies have attributed dental microwear to abrasion by phytoliths (Baker et al., 1959; Walker et al., 1978), which are considerably harder than tooth enamel. However, a recent study tested the hardness of silica phytoliths and mammalian teeth enamel and found that the former are softer than enamel (Sanson et al., 2007). These authors suggested that exogenous particles, such as dust and grit, are the major source of enamel wear since plant organs are exposed to, and accumulate, such sort of materials. In addition, when seeds and nuts are processed by means of grinding stones, wear rates could be greatly accelerated by the fine abrasives added to the food. Therefore, food preparation techniques should be examined, since the use of grinding stones to process the same food items results in increased rates of dental wear, whereas dental wear rates decline markedly with the development of ceramic cooking vessels (Molleson et al., 1993; Sciulli, 1997). The relevance of food preparation and consumption patterns is stressed by Walker et al. (1998) in their analysis of ethnographic data of Yanomamo, Yora, and Shiwiar groups from Amazons. All of these groups obtain their food through hunting, gathering, and slash-and-burn agriculture, so their diets are very similar. However, these authors found large differences in caries ratio and dental wear rate among groups, which are explained by small differences in their patterns of food consumption and processing. Because of the aridity of the geographic areas under study, airborne dust and sand particles would had been a common source of abrasive particles incorporated into the food. The methods for preparation of dried meat used by the human groups that inhabited this region during the 19th century, which included exposure of strips of lean meat to the sun and a combination of salting followed by air drying, favored the incorporation of abrasive particles into food (Dobrovsky, 1946; Musters, 2005). Ethnohistoric sources also mention that cooking meat directly over a hearth or ashes, which results in the addition of hard particles to the food, was a common food preparation technique (Musters, 2005). Although these customs were recorded only after European contact, it is reasonable to infer that similar practices existed in earlier times. These sources of abrasive particles were probably common for all Holocene populations in Patagonia; nevertheless, some differences could have existed between coastal and inland groups. Since

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particles associated with marine resources such as sand and grit represent an important source of enamel wear, it would be expected that coastal populations exhibit greater wear rates than inland populations. However, the results of principal axis analysis do not support this hypothesis, because the slope values for samples from Central East and Northeast Patagonia, which correspond to populations situated near the coast, are very similar to the values for samples from Northwest Patagonia. The temporal trend toward reduced wear rates observed in the three Patagonian areas studied (Table 9) suggests that a shift occurred after ca. 1,500 years BP involving either a change in the type of food consumed or in the processing methods employed. The lack of temporal changes in caries ratio between the two periods analyzed in this study does not support the hypothesis of a change of items consumed, at least not in the proportion of resources derived from plants and animals. With respect to the preparation techniques, the use of groundstone tools could have increased the amount of hard abrasives entering the diet because these tools are constructed from materials such as quartz, which contain particles of grit harder than enamel (Mahoney, 2006). However, temporal trends in the use of such artifacts cannot be accurately evaluated since, as mentioned above, there are not radiocarbon dates for most of them. In this sense, the presence of ceramic in the archaeological sites from the final late Holocene is noteworthy. Pottery with remains of vegetables and evidences of cooking were found in Central Patagonia, in sites dated ca. 880 years 14C BP (La Azucena 1 and 2, Gomez Otero et al., 1999). Likewise, the available information for Northeast Patagonia indicates the presence of ceramic in archaeological sites dated later than ca. 1,500 years BP (La Petrona, ca. 500–300 years 14C BP, Martinez, 2004; La Eloisa 1 ca. 1,340 years 14C BP, Las Olas 5 ca. 570 years 14C BP, Sanguinetti de Bo´rmida, 1999). Finally, radiocarbon dating indicates the presence of ceramic in Northwest Patagonia around ca. 1,700–1,500 years BP (Lagiglia, 2002). These evidences suggest that food processing would become more intensive in the latter period through the practice of boiling in ceramic vessels. Therefore, the introduction of pottery in these areas could have led to softer diets, which resulted in a slower wear rate. Previous studies of dental occlusal microwear have noted that the introduction of pottery has profound effects on dietary hardness (Molleson et al., 1993; Schmidt, 2001). However, the lack of analyses of microwear in the areas under study does not allow differentiation of the effects of food hardness and abrasiveness.

CONCLUSION In this article, we showed that caries frequencies remained constant from early to final late Holocene samples from the same areas. The ratio of carbohydrates consumed seems to have remained fairly constant since 3,000 years BP, and therefore these results do not support a dietary shift to a greater reliance on plant foods in final late Holocene populations. Although all the Patagonian samples fall within the range established for hunter-gatherers, there are some geographical differences. According to our results, the populations from Northeast Patagonia consumed a diet with lower proportion of carbohydrates than that of the populations from Central East and Northwest Patagonia. Finally, there is a marked temporal trend toward the reduction of wear

rates in the three areas, which would reflect a change to less hard and/or abrasive diets in the final late Holocene, probably due to differences in the processing methods employed. The results obtained stress the variety of subsistence strategies among Patagonian hunter-gatherers. However, there are some aspects that need a grater development in the area under study. Analyses of dental microwear are needed to compare the hardness of food items consumed in different geographical regions, with availability of different kind of resources and environmental setting, through time during the late Holocene. The application of techniques aimed to recover and identify micro and macro remains of plant foods in archaeological sites of hunter-gatherers can help to establish which species were consumed by prehistoric populations and whether they were the same than those documented by the chronicles in historical times. Functional analysis of ground stone tools and pottery and radiocarbon dates are needed for assessing their prehistoric use and obtaining a chronological control about their incorporation in these populations. The use of these data can contribute to a better understanding about the role of plant foods among Patagonian hunter-gatherers, as well as identify the food processing and consumption techniques employed.

ACKNOWLEDGMENTS We are grateful to Hector M. Pucciarelli [Divisio´n Antropologı´a. Facultad de Ciencias Naturales y Museo of La Plata (Argentina)], Ine´s Baffi, and Leandro Luna [Museo Etnogra´fico \J. B. Ambrosetti" of Buenos Aires (Argentina)] for granting access to the human skeletal collections under their care. Clark Spencer Larsen and two anonymous reviewers offered insightful comments during the review process.

LITERATURE CITED Adams JL. 1999. Refocusing the role of food-grinding tools as correlates for subsistence strategies in the U.S. Southwest. Am Antiq 64:475–498. Babot MP, Mazzia N, Beyon C. 2005. Potencialidad de los ana´lisis sobre artefactos de molienda. IV Congreso de Arqueologı´a de la Regio´n Pampeana Argentina. p 105–106 Baker G, Jones LHP, Wardrop ID. 1959. Cause of wear in sheep’s teeth. Nature 184:1583–1584. Barberena R. 2002. Los limites del mar. Iso´topos estables en Patagonia meridional. Buenos Aires: Sociedad Argentina de Antropologı´a. Barrientos G, Perez SI. 2004. La expansio´n y dispersio´n de poblaciones del norte de Patagonia durante el Holoceno tardı´o: evidencia arqueolo´gica y modelo explicativo. In: Civalero MT, Fernandez PM Guraieb AG, editors. Contra viento y marea. Arqueologı´a de Patagonia. Buenos Aires: Instituto Nacional de Antropologı´a y Pensamiento Latinoamericano. p 179–195. Bell EJ, Kaidonis J, Townsend G, Richards L. 1998. Comparison of exposed dentinal surfaces resulting from abrasion and erosion. Aust Dent J 43:362–366. Benfer RA, Edwards DS. 1991. The principal axis method for measuring rate and amount of dental attrition: estimating juvenile or adult tooth wear from unaged adult teeth. In: Kelley MA, Larsen CS, editors. Advances in dental anthropology. New York: Wiley-Liss. p 325–340. Binford RL. 2001. Constructing frames of reference: An analytical method for archaeological theory building using ethnographic and environmental data sets. Berkeley-Los Angeles: University of California Press.

American Journal of Physical Anthropology—DOI 10.1002/ajpa

ROLE OF FOOD PLANTS IN PATAGONIAN HUNTER-GATHERERS Bonomo M. 2005. Costeando las llanuras: arqueologı´a del litoral marı´timo pampeano. Buenos Aires: Sociedad Argentina de Antropologı´a. Bo´rmida M. 1964. Arqueologı´a de la costa Norpatago´nica. Trabajos de Prehistoria 16:7–108. Bo´rmida M, Casamiquela R. 1958–59. Etnografı´a Gu¨nu¨na-ke¨na. Testimonio del u´ltimo de los tehuelches Septentrionales. Runa 9:153–193. Brooks S, Suchey J. 1990. Skeletal age determination base on the os Pubis: A Comparison of the Acsa´di-Nemeske´ri and Suchey-Brooks Methods. Hum Evol 5:227–238. Buikstra J, Ubelaker D. 1994. Standards for data collection from human skeletal remains Arkansas Archaeological Survey Research Series No. 44. Fayetteville: Arkansas Archaeological Survey. Cabrera AL. 1976. Regiones fitogeogra´ficas argentinas. Buenos Aires: Acme. Cabrera A, Yepes J. 1960. Mamı´feros Sudamericanos. Tomo I. Buenos Aires: EDIAR. Delgado-Darias T, Velasco-Va´zquez J, Arnay-de-la-Rosa M, Martı´n-Rodrı´guez E, Gonza´lez-Reimers E. 2006. Dental caries among the prehispanic population from Gran Canaria. Am J Phys Anthropol 128:560–568. Della Negra CE, Novellino PS. 2005. ‘‘Aquihueco´’’: Un Cementerio Arqueolo´gico, en el Norte de la Patagonia, Valle del Curi Leuvu´ – Neuque´n, Argentina. Magallania 33:165–172. Dobrovsky M. 1946. Abrasiones dentarias en cra´neos de Indios Patagones. Revista del Museo de La Plata. T II:301–347. Duyar I, Erdal YS. 2003. A new approach for calibrating dental caries frequency of skeletal remains Homo 54:57–70. FAO. 1993. El maı´z en la nutricio´n humana. In: Alimentacio´n y nutricio´n, N825. Organizacio´n de las Naciones Unidas para la Agricultura y la Alimentacio´n. Roma. Fernandez J. 1988. Roedores, guanacos, huevos, semillas de araucaria y almeja fluvial. Estacionalidad, subsistencia y estrategia locacional en Haichol, Cordillera andina del Neuque´n. Precirculados de las Ponencias Cientı´ficas Presentadas a los Simposios del IX Congreso Nacional de Arqueologı´a Argentina. Buenos Aires: Universidad de Buenos Aires. p 130–139. Fernandez J, Panarello HO. 2001. Cazadores recolectores del Holoceno Medio y Superior de la Cueva Haichol, regio´n cordillerana central del Neuque´n, Republica Argentina. Relaciones 26:9–30. Gambier M. 1993. Prehistoria de San Juan, Argentina. San Juan: Editorial Universidad Nacional de San Juan. Gil A. 2003. The Zea mays in the archaeological record on the frontier American: chronology and their role in the diet. Curr Anthropol 44:295–300. Gil A, Shelnut N, Neme G, Tykot R, Michieli CT. 2006b. Iso´topos estables y dieta humana en el centro oeste: datos de muestras de San Juan. Cazadores Recolectores del Cono Sur. Revista de Arqueologı´a 1:149–162. Gil A, Tykot R, Neme G, Shelnut N. 2006a. Maize on the frontier. Isotopic and macrobotanical data from Central-Western Argentina. In: Staller J, Tykot R, Benz B, editors. Histories of maize multidisciplinary approaches to the prehistory, biogeography, domestication, and evolution of maize. p 199– 214. Gold DL. 2000. \Utmost confusion" reconsidered: bioarchaeology and secondary burial in later prehistoric interior Virginia. In: Lambert PM, editor. Bioarchaeological studies of life in the age of agriculture. Tuscaloosa: University of Alabama Press. p 195–218. Go´mez Otero J. 2006. Iso´topos estables y dieta en Patagonia Centro-Septentrional. Programa y resu´menes del Taller de Arqueologı´a e iso´topos estables en el sur de Sudame´rica. Discusio´n e Integracio´n de resultados. Notas del Museo de Historia Natural de San Rafael 60:19. Gomez Otero J, Belardi JB, Su´nico A, Taylor R. 1999. Arqueologı´a de cazadores recolectores en Penı´nsula Valdez (Costa Central de Patagonia): primeros resultados. In: Soplando en el Viento. Actas de las Terceras Jornadas de Arqueologı´a de la Patagonia. INAPL/Universidad Nacional del Comahue. p 393– 417.

1057

Go´mez Otero J, Belardi JB, Tykot R, Grammer S. 2000. Dieta y poblaciones humanas en la costa norte del Chubut (Patagonia argentina). In: Belardi JB, Carballo Marina F, Espinosa S, editors. Desde el Paı´s de los Gigantes. Perspectivas arqueolo´gicas en Patagonia. IV Jornadas de Arqueologı´a de la Patagonia. Rı´o Gallegos: Universidad Nacional de la Patagonia Austral. p 109–122. Graw M. 2001. Significance of the classical morphological criteria for identifying gender using recent skulls. Forensic Sci Commun [online] 3: 1. [15 March 2006] Available . Grippo JO, Simring M, Schreiner S. 2004. Attrition, abrasion, corrosion and abfraction revisited: a new perspective on tooth surface lesions. J Am Dent Assoc 135:1109–1118. Guicho´n RA. 1995. Vı´as de ana´lisis, problemas y discusiones en la antropologı´a biolo´gica de Tierra del Fuego. Relaciones 20:239–256. Guicho´n RA, Borrero LA, Prieto AI, Ca´rdenas P, Tykot R. 2001. Nuevas determinaciones de iso´topos estables para Tierra del Fuego. Rev Argent Antropol Biol 3:113–126. Gusinde M. 1982. Los indios de Tierra del fuego, Vol 1. Los Selknam. Buenos Aires: Centro Argentino de Etnologı´a. Hather JG, Mason SLR. 2002. Introduction: some issues in the archaeobotany in hunter-gatherers. In: Mason SLR, Hather JG, editors. Hunter-gatherer archaeobotany: perspectives from the northern temperate zone. London: Institute of Archaeology, University College London. p 1–14. Hayden B. 1981. Subsistence and ecological adaptations of modern hunter/gatherers. In: Harding R, Teleki G, editors. Omnivorous primates. New York: Columbia University Press. p 344–421. Hernandez AM. 2002. Paleoetnobotanica en el sur de Mendoza. In: Gil A, Neme G, editors. Entre montan˜as y desiertos: arqueologı´a del sur de Mendoza. Buenos Aires: Sociedad Argentina de Antropologı´a. p 157–180. Hillson S. 1996. Dental anthropology. Cambridge: Cambridge University Press. Hillson S. 2000. Dental pathology. In: Katzenberg MA, Saunders SR, editors. Biological anthropology of the human skeleton. New York: Wiley-Liss. p 249–285. Hillson S. 2001. Recording dental caries in archaeological human remains. Int J Osteoarchaeol 11:249–289. Ihaka R, Gentleman R. 1996. R: A language for data analysis and graphics. J Comput Graph Stat 5:299–314. Jackes M, Lubell D. 1996. Dental pathology and diet: second thoughts. In: Otte M, editor. Nature et culture: Actes du Colloque International de Lie`ge . Etudes et Recherches Arche´ologiques de L’Universite´ de Lie`ge. Vol. 68, p 457–480. Kaidonis J, Richards LC, Townsend G, Tansley GD. 1998. Wear of human enamel: a quantitative in vitro assessment. J Dent Res 77:1983–1990. Katzenberg MA. 2000. Stable isotope analysis: a tool for studying past diet, demography, and life history. In: Katzenberg MA, Saunders SR, editors. Biological anthropology of the human skeleton. New York: Wiley-Liss. p 305–327. Kelly J, Grammer S, Tykot RH, Belardi J, Borrero L, Gomez Otero J, Guicho´n R. 2001. Integrating archaeological, ethnographic and analytical subsistence data: a case study from Patagonia, South America. In: 66th Annual Meeting of the Society for American Archaeology, New Orleans, Louisiana. Kelly RL. 1995. The foraging spectrum: Diversity in huntergatherer lifeways. Washington: Smithsonian Institution Press. Kieser JA, Dennison KJ, Kaidonis JA, Huang D, Herbison PGP, Tayles NG. 2001a. Patterns of dental wear in the early Maori dentition. Int J Osteoarch 11:206–217. Kieser JA, Kelsen A, Love R, Herbison PGP, Dennison KJ. 2001b. Periapical lesions and dental wear in the early Maori. Int J Osteoarch 11:290–297. Klein H, Palmer CE, Knutson JW. 1938. Studies on dental caries: I Dental status and dental needs of elementary school children. Public Health Rep 53:751–765. Kozameh L. 1993. Patrones de abrasio´n dentaria en dos poblaciones prehisto´ricas argentinas. Bol Soc Esp Antrop Biol 14:81–104.

American Journal of Physical Anthropology—DOI 10.1002/ajpa

1058

V. BERNAL ET AL.

Kozameh L, Barbosa E. 1992. Patrones de abrasio´n dentaria en restos esqueletales. In: Fernandez J, editor. La cueva Haichol. Arqueologı´a de los Pinares Cordilleranos del Neuque´n. Universidad Nacional de Cuyo. An Arqueol Etnol , Vol III, p 613– 630. Lagiglia H. 1957. La presencia del patay en una tumba indı´gena de San Rafael (Mendoza). Notas del Museo 1. San Rafael. Museo de Historia Natural. Lagiglia H. 2002. Arqueologı´a prehisto´rica del sur mendocino y sus relaciones con el centro oeste Argentino. In: Gil A, Neme G, editors. Entre montan˜as y desiertos: arqueologı´a del sur de Mendoza. Buenos Aires: Sociedad Argentina de Antropologı´a. p 43–64. Lalueza Fox C, Jordi J, Rosa MA. 1996. Phytolith analysis on dental calculus, enamel surface, and burial soil: information about diet and paleoenviroment. Am J Phys Anthropol 101:101–113. Ları´a SC. 1961. Contribucio´n al estudio de la arqueologı´a de la regio´n este de Rı´o Negro. An Arqueol Etnol 16:247–257. Larsen CS. 1995. Biological changes in human populations with agriculture. Annu Rev Anthropol 24:185–213. Larsen CL. 1997. Bioarchaeology: interpreting behavior from the human skeleton. New York: Cambridge University Press. Larsen CS. 2002. Bioarchaeology: the lives and lifestyles of past people. J Archaeol Res 10:119–166. Larsen CS, Shavit R, Griffin MC. 1991. Dental caries evidence for dietary change: an archeological context. In: Kelley MA, Larsen CS, editors. Advances in dental anthropology. New York: Wiley-Liss. p 179–202. Lee R, Devore R. 1968. Man the hunter. New York: Aldine Publishing Company. L’Heureux GL. 2002. Inferencias paleodietarias a partir del ana´lisis de los patrones de desgaste dental y frecuencia de caries en muestras de restos humanos del Holoceno del Sudeste de la Regio´n Pampeana. In: Mazzanti D, Bero´n MA, Oliva FW, editors. Del Mar a los Salitrales. Diez mil An˜os de Historia Pampeana en el Umbral del Tercer Milenio. Mar del Plata: Universidad Nacional de Mar del Plata. Sociedad Argentina de Antropologı´a. p 127–140. Lev-Tov Chattah N, Smith P. 2006. Variation in occlusal dental wear of two Chalcolithic populations in the southern Levant. Am J Phys Anthropol 130:471–479. Lovejoy CO, Meindl RS, Pryzbeck TR, Mensforth RP. 1985. Chronological metamorphosis of the auricular surface of the ilium: a new method for the determination of adult skeletal age at death. Am J Phys Anthropol 68:15–28. Lukacs JR. 1992. Dental pathology and agricultural intensification in South Asia new evidence from Bronce Age Harappa. Am J Phys Anthropol 87:133–150. Lukacs JR. 1995. The ‘caries correction factor’: a new method of calibrating dental caries rates to compensate for ante-mortem loss of teeth. Int J Osteoarch 5:151–156. Maat GJR, van der Velde EA. 1987. The caries -attrition competition. Int J Anthropol 2:281–292. Mahoney P. 2006. Dental microwear from Natufian hunter-gatherers and early Neolithic farmers: comparisons within and between samples. Am J Phys Anthropol 130:308–319. Mansur-Franchomme ME, Orquera LA, Piana EL. 1987–1988. El alisamiento de la piedra entre cazadores-recolectores: el caso de Tierra del Fuego. Runa 17–18:111–206 Martinez G. 2004. Resultados preliminares de las investigaciones arqueolo´gicas realizadas en el curso inferior del rı´o Colorado (partidos de Villarino y Patagones provincia de Bs. As. In: Martinez G, Gutierrez M, Curtoni R, Beron M, Madrid P, editors. Aproximaciones contempora´neas a la arqueologı´a pampeana. Perspectivas teo´ricas, metodolo´gicas, analı´ticas y casos de estudio. Olavarria: UNPCBA. Martı´nez G, Zangrando AF, Prates L. 2006. Primeros resultados de d13C y d15N sobre restos o´seos humanos para el curso Inferior del Rı´o Colorado (Pcia. De Buenos Aires). Programa y Resu´menes del Taller de Arqueologı´a e iso´topos estables en el sur de Sudame´rica. Discusio´n e Integracio´n de resultados. Notas del Museo de Historia Natural de san Rafael 60:17.

Meiklejohn C, Wyman JW, Schentag CT. 1992. Caries and attrition: dependent or independent variables? Int J Anthropol 7:17–22. Meindl RS, Lovejoy CO. 1985. Ectocranial suture closure: a revised method for the determination of skeletal age at death based on the lateral-anterior sutures. Am J Phys Anthropol 68:57–66. Milner GR. 1984. Dental caries in the permanent dentition of a Mississippian period population from de American Midwest. Coll Antropol 8:77–91. Molleson T, Jones K, Jones S. 1993. Dietary change and the effects of food preparation on microwear patterns in the late Neolithic of Abu Hureyra, northern Syria. J Hum Evol 24:455–468. Molnar S. 1971. Human tooth wear, tooth function and cultural variability. Am J Phys Anthropol 34:175–190. Molnar S, McKee JK, Molnar I. 1983. Measurements of tooth wear among Australian aborigines. I. Serial loss of the enamel crown. Am J Phys Anthropol 61:51–65. Moody JEH. 1960. The dental and periodontal conditions of aborigines at settlements in Arnhem land and adjacent areas. In: Mountford CR, editor. Records of the American-Australian scientific expedition to Arnhem land: anthropology and nutrition. Melbourne: Melbourne University Press. Vol. 2, p 60–71. Moreno F. 1874. Cementerios y paraderos prehisto´ricos de la Patagonia. Anales Cientı´ficos Argentinos 1:2–13. Murdock G. 1967. Ethnographic atlas. Pittsburgh: University of Pittsburgh Press. Musters GC. 2005. Vida entre los Patagones. Buenos Aires: El Elefante Blanco. Nacuzzi LR, Perez de Micou C. 1983–1985. Los recursos vegetales de los cazadores de la cuenca del rı´o Chubut. Cuadernos del Instituto Nacional de Antropologı´a.10:407–424. Neme G. 2002. Arqueologı´a del Alto valle del rı´o Atuel: Modelos problemas y perspectivas en el estudio de las regiones de altura del sur de Mendoza. In: Gil A, Neme G, editors. Entre Montan˜as y Desiertos. Arqueologı´a del sur de Mendoza. Buenos Aires: Sociedad Argentina de Antropologı´a. p 65–83. Neme G, Gil A. 2002. La explotacio´n faunı´stica y la frecuencia de partes esqueletarias en el registro arqueolo´gico del sur mendocino. In: Gil, A, Neme G, editors. Entre montan˜as y desiertos. Arqueologı´a del sur de Mendoza. Buenos Aires: Sociedad Argentina de Antropologı´a. p 141–155. Neme G, Gil A, Duran V. 2005. Late Holocene in southern Mendoza (northwestern Patagonia): radiocarbon pattern and human occupation. Before Farming 2: article 5. Novellino PS, Guicho´n R. 1999. Formas de subsistencias e iso´topos estables en el sur de Mendoza. Rev Argent Antropol Biol 2:323–334. Novellino P, Guicho´n R. 1997–1998. Comparacio´n de indicadores dieta y salud entre el sur de San Juan-Norte de Mendoza. Relaciones XXII–XXIII:125–138. Novellino PS, Gil A, Neme G, Duran V. 2004. El consumo de maı´z en el Holoceno tardı´o del Oeste Argentino: iso´topos estables y caries. Rev Esp Arqueol Am 34:85–110. Outes F. 1907. Arqueologı´a de San Blas (provincia de Buenos Aires). Anales del Museo Nacional de Buenos Aires 16:249– 273. Perez-Perez A, Lalueza Fox C. 1992. Indicadores de presio´n ambiental en aborı´genes de Fuego Patagonia. Un reflejo de la adaptacio´n a un ambiente adverso. An Inst Patagon Ser Cienc Hum 21:99–108. Perez SI. 2006. El poblamiento holoce´nico del Sudeste de la Regio´n Pampeana: un estudio de morfometrı´a geome´trica craneofacial. Ph.D. dissertation. Universidad Nacional de La Plata, La Plata, Argentina. Phenice TW. 1969. A newly developed visual method of sexing the os pubis. Am J Phys Anthropol 30:297–301. Prates LR. 2004. Arqueologı´a de la cuenca media del rı´o Negro (provincia de Rı´o Negro). Una primera aproximacio´n. Intersecciones Antropol 5:55–69. Powell ML. 1985. The analysis of dental wear and caries for dietary reconstruction. In: Gilbert RI, Mielke JH, editors. The analysis of prehistoric diets. Orlando: Academic Press. p 307–338.

American Journal of Physical Anthropology—DOI 10.1002/ajpa

ROLE OF FOOD PLANTS IN PATAGONIAN HUNTER-GATHERERS Richards LC. 1984. Principal axis analysis of dental attrition data from two Australian aboriginal populations. Am J Phys Anthropol 65:5–13. Roig FA. 1993. Informe nacional para la seleccio´n de germoplasma en especies de Prosopis de la Repu´blica Argentina. In: Unidades de Bota´nica y Fisiologı´a Vegetal, IADIZA, editors. Contribuciones Mendocinas a la Quinta Reunio´n Regional para Ame´rica Latina y El Caribe de la Red de Forestacio´n del CIID. p 1–37. Sanguinetti de Bo´rmida AC. 1999. Proyecto nordpatagonia. Arqueologı´a de la costa septentrional. Separata de Anales de la Academia Nacional de Ciencias de Buenos Aires. p 1–35. Sanson GD, Kerr SA, Gross KA. 2007. Do silica phytoliths really wear mammalian teeth? J Archaeol Sci 4:526–531. Schmidt CW. 2001. Dental microwear evidence for a dietary shift between two nonmaize-reliant prehistoric human populations from Indiana. Am J Phys Anthropol 114:139–145. Sciulli PW. 1997. Dental evolution in prehistoric native Americans of the Ohio Valley Area. I. Wear and pathology. Int J Osteoarchaeol 7:507–524. Scott EC. 1979a. Dental wear scoring technique. Am J Phys Anthropol 51:213–218. Scott EC. 1979b. Principal axis analysis of dental attrition data. Am J Phys Anthropol 51:203–212. Scott GR, Turner CG II. 1988. Dental anthropology. Annu Rev Anthropol 17:99–126. Smith, B Holly. 1984. Patterns of molar wear in hunter—gatherers and agriculturalists. Am J Phys Anthropol 63:39–56. Smith P. 1972. Diet and attrition in the Natufian. Am J Phys Anthropol 37:233–238. Smith P, Bar-Yosef O, Sillen A. 1984. Archaeological and skeletal evidence for dietary change during the late Pleistocene/ early Holocene in the Levant. In: Cohen MN, Armelagos GJ, editors. Paleopathology at the origins of agriculture. New York: Academic Press. p 101–136. Sokal RR, Rohlf FJ. 1979. Biometrı´a. Principios y me´todos estadı´sticos en la investigacio´n biolo´gica. Madrid: H. Blume Ediciones. Tassara G, Osterrieth M. 2005. Silicofitolitos en artefactos de ´ rea Interserrana, Buemolienda de sitios Arqueolo´gicos del A nos Aires. 3er Encuentro de Investigaciones Fitolı´ticas del Cono Sur Tafı´ del Valle, Tucuma´n. p 19–21. Teaford MF, Lytle JD. 1996. Brief communication: diet-induced changes in rates of human tooth microwear: a case study involving stone-ground maize. Am J Phys Anthropol 100:143– 147.

1059

Teaford MF, Oyen OJ. 1989. Differences in the rate of molar wear between monkeys raised on different diets. J Dent Res 68:1513–1518. Torres LM. 1922. Arqueologı´a de la Penı´nsula San Blas (Provincia de Buenos Aires). Revista del Museo de La Plata 26:473–532. Turner CG II. 1979. Dental anthropological indications of agriculture among the Jomon people of central Japan. Am J Phys Anthropol 51:619–636. van Reenen JF. 1966. Dental features of a low-caries primitive population. J Dent Res 45:703–713. Vignati MA. 1931. Investigaciones antropolo´gicas en el litoral marı´timo subatla´ntico bonaerense. Notas Preliminares del Museo de La Plata 1:19–31. Vignati MA. 1941. El ‘pan’ de los patagones protohistoricos. Notas del Museo de La Plata 6:321–336. Walker PL. 1978. A quantitative analysis of dental attrition rates in the Santa Barbara Channel Area. Am J Phys Anthropol 48:101–106. Walker PL, Erlandson JM. 1996. Dental evidence for prehistoric dietary change on the northern Channel Islands, California. Am Antiq 51:376–383. Walker AC, Hoeck HN, Perez L. 1978. Microwear of mammalian teeth as an indicator of diet. Science 201:908–910. Walker PL, Sugiyama L, Chacon R. 1998. Diet, dental health, and cultural change among recently contacted South American Indian hunter-horticulturalists. In: Lukacs JR, editor. Human dental development, morphology, and pathology: a tribute to Albert A. Dahlberg. Eugene: University of Oregon Anthropological Papers, Vol. 54. p 355–386. Walrath DE, Turner P Bruzek J. 2004. Reliability test of the visual assessment of cranial traits for sex determination. Am J Phys Anthropol 125:132–137. Wright KI. 1994. Ground-stone tools and hunter-gatherer subsistence in Southwest Asia: implications for the transition to farming. Am Antiq 59:238–263. Yesner DR, Figuerero Torres MJ, Guicho´n RA, Borrero LA. 2003. Stable isotope analysis of human bone and ethnohistoric subsistence patterns in Tierra del Fuego. J Anthropol Archaeol 22:279–291. Zucol AF, Bonomo M. 2005. Estudios etnobota´nicos del sitio arqueolo´gico Nutria Mansa 1 (partido de General Alvarado, provincia de Buenos Aires). II. Ana´lisis fitolı´ticos comparativos de artefactos de molienda. 3er Encuentro de Investigaciones Fitolı´ticas del Cono Sur Tafı´ del Valle, Tucuma´n. p 19– 21.

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