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Craniofacial morphology in the Argentine center-west Consequences of the transition to food production.

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AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY 130:333–343 (2006)
Craniofacial Morphology in the Argentine Center-West:
Consequences of the Transition to Food Production
Marina L. Sardi,1,2* Paula S. Novellino,2,3 and Héctor M. Pucciarelli1,2
1
Departamento Cientı́fico de Antropologı́a, Facultad de Ciencias Naturales y Museo,
Universidad Nacional de La Plata, 1900 La Plata, Argentina
2
Consejo Nacional de Investigaciones Cientı́ficas y Técnicas, Buenos Aires, Argentina
3
Laboratorio de Bioarqueologı́a, Museo de Historia Natural de San Rafael, 5600 San Rafael, Mendoza, Argentina
KEY WORDS
subsistence economy; functional cranial components; masticatory stress
ABSTRACT
The Argentine Center-West was the
southernmost portion of the Andes where domestication
of plants and animals evolved. Populations located in the
southern portion of this area displayed a hunter-gatherer
subsistence economy up to historical times, and coexisted
with farmers located to the north. Archaeological and biological evidence suggests that the transition to food production was associated with the consumption of a softer
diet and a more sedentary way of life. This study tests
the hypothesis that diet-related factors influenced morphological differentiation, by comparing functional cranial
components of farmers and hunter-gatherers. Three-dimensional changes on eight minor functional components
(anteroneural, midneural, posteroneural, otic, optic, respiratory, masticatory, and alveolar) were measured on
skulls derived from both subareas. Volumetric and morphometric indices were calculated to estimate the absolute and relative size of components, respectively. Results
of a paired t-test indicated that farmers have a smaller
craniofacial size than hunter-gatherers. The components
that varied the most were masticatory and posteroneural,
showing smaller absolute and relative sizes in farmers.
Discriminant analyses indicated that lengths and widths
were the most affected dimensions of these and other
components. The pattern of differentiation, which involves
specific components, enabled us to exclude differential
gene flow and stochastic mechanisms as the main causes.
Instead, results support the hypothesis that diet-related
factors associated with both subsistence economies influenced craniofacial morphology. A proportion of the observed variation associated with size differences can be
explained by two systemic factors: the lesser quality of
nutrition due to a low protein content in the diet, and a
decrease of growth hormone circulation induced by a
lower mobility due to sedentism. However, differentiation
is better explained by a localized factor: the reduction in
the masticatory and posteroneural components in farmers
resulted from a decrease of masticatory stresses and
workload on the head and neck, linked to the consumption of a softer diet. Am J Phys Anthropol 130:333–343,
2006. V 2006 Wiley-Liss, Inc.
One of the most important changes in human evolution during the Holocene was the transition from hunting and gathering to food production. The increased
dependence on agriculture and pastoralism implied
several transformations of biological characteristics in
human populations worldwide (Pechenkina et al., 2002;
Richards, 2002; Larsen, 2003; Eshed et al., 2004). In
South America, the earliest evidence of agriculture
appeared in Peru and Ecuador, yielding dates of around
6,900 years BP (Pearsall, 1992) or even older (Smith,
1998). The Argentine Center-West was the southernmost
portion of the Andes where food production evolved (Gil,
2003). Populations located in the southern portion of this
area maintained a hunter-gatherer subsistence economy,
coexisting with farmers located in the north.
Because morphology arises from the translation of
genotype into phenotype through several epigenetic processes, phenotypes of a group of populations express their
genetic relationships as well as the influence of environmental factors acting across ontogeny. Nevertheless,
studies are commonly more concerned with distances
among populations from a taxonomic point of view than
with contextual factors and adaptation (Armelagos and
Van Gerven, 2003) that may affect differentiation. Moreover, which structures might be more plastic to these
factors is a subject frequently disregarded (GonzálezJosé et al., 2005).
BIOCULTURAL CONTEXT
C 2006
V
WILEY-LISS, INC.
C
The Argentine Center-West is an archaeological area
comprised of the provinces of San Juan and Mendoza
(Fig. 1), which are longitudinally variable in geomorphological and ecological terms (Capitanelli, 1972; Roig,
1972; Ruiz Leal, 1972). Three ecological zones can be distinguished: 1) the Andean mountains, with more than
2,000 m of altitude acting as a barrier to humid Pacific
winds and promoting arid conditions; 2) a piedmont plateau, between 1,000–2,000-m altitude, which is entirely
crossed by valleys, although most of them almost constantly lack water; and 3) plains, at less than 1,000-m
altitude.
Grant sponsor: Universidad Nacional de La Plata, Argentina;
Grant sponsor: Fondation Fyssen, France.
*Correspondence to: Marina L. Sardi, Departamento Cientı́fico de
Antropologı́a, Museo de La Plata, Paseo del Bosque S/N, 1900 La
Plata, Argentina. E-mail: msardi@fcnym.unlp.edu.ar
Received 3 March 2005; accepted 10 August 2005.
DOI 10.1002/ajpa.20379
Published online 18 January 2006 in Wiley InterScience
(www.interscience.wiley.com).
334
M.L. SARDI ET AL.
Fig. 1. Argentine Center-West. Localities where skulls were
recovered. 1, Uspallata; 2, Mendoza; 3, Capiz, 4; Campo Las
Julias; 5, Arroyo Imperial; 6, Arroyo El Tigre; 7, Dique Villa 25
(Los Reyunos); 8, Dique Villa 25 (La Hedionda); 9, Dique Villa
25 de Mayo; 10, Los Coroneles; 11, Médano Puerto Dı́az; 12,
Arroyo Los Jilgueros; 13, Rincón del Atuel; 14, Loma del Eje;
15, Cañada Seca; 16, Jaime Prats; 17, Cerro Negro; 18, Agua
del Médano; 19, El Nihuil; 20, Agua del Zapallo; 21, Cerro
Meson; 22, Respolar; 23, La Herradura; 24, Puerto Aisol; 25, El
Sosneado; 26, Puerto Tierras Blancas; 27, Cerro Mesa; 28, El
Manzano; 29, La Matancilla; 30, El Chacay; 31, La Cañada; 32,
Sur de Malargüe; 33, Calingasta; 34, San Juan; 35, Caliningasta
Barrealito; 36, Angualasto; 37, Pachimoco; 38, Huaco.
In ethnographic and archaeological terms, this area is
divided by the Diamante River into northern and southern subareas. The northern subarea comprises the provinces of San Juan and Mendoza up to the Diamante
River; the southern subarea is located to the south of
Mendoza (Fig. 1). According to ethnohistorical data, aboriginal populations inhabiting the Argentine CenterWest at the arrival of the Spaniards were divided in two
different cultural complexes with different subsistence
economies. Farmer groups, named Huarpes, inhabited the
northern subarea, while the hunter-gatherer Puelches
inhabited the southern subarea (Cabrera, 1929; Latcham,
1929; Canals Frau, 1937, 1953; Michieli, 1978; Prieto,
1989; Durán, 1994).
Evidence of Late Pleistocene and Early Holocene occupation was dated to 11,000 and 10,500 years BP in the
northern and southern subareas, respectively (Gambier,
1976; Garcı́a, 1997; Lagiglia, 2002). Populations based
their subsistence on the hunting of guanaco (Lama guanicoe), ñandú (Rhea americana), and other small mammals, and on the gathering of wild plants by seasonal
movements between lowlands and highlands (Gambier,
1993; Lagiglia, 2002). The earliest evidence of agriculture in the northern subarea is dated to around 4,400
years BP (Bárcena, 1985), and is associated with pottery
and sedentism (Lagiglia, 2002). The transition to food
production in the Argentine Center-West was part of a
general process that occurred in the South-Central
Andean region where, across the archaic cultural tradition, some changes in the archaeological record suggest
a diversification of diet and a demographic increase
(Castro and Tarragó, 1992; Planella and Tagle, 2004). As
part of this diversification, local groups gradually
adopted agriculture and pastoralism, as complementary
resources to hunting, gathering, and fishing (Castro and
Tarragó, 1992; Gambier, 1993). Potato (Solanum tuberosum), manioc (Manihot esculenta), beans (Phaseolus vulgaris), and maize (Zea mays) were some of the species
included in the diet. It is accepted that plant and animal
domestication was not a uniform process, but in general
terms, a continuity from archaic to formative groups
seems to be supported (Castro and Tarragó, 1992; Planella and Tagle, 2004), as occurred in the valleys of
northern Chile where continuity was demonstrated by
craniometrics and mitochondrial DNA (mtDNA) analyses
(Moraga et al., 2005).
Some archaeological indicators suggest changes associated with food production in the northern subarea.
Grinding-stone artifacts were utilized, and their morphology evidenced food preparation because they were
deep and associated with discoid manos (Lagiglia, 1997).
Some villages presented collective mills made of large
rocks with many pits (Lagiglia, 1997). Pottery remains
were diverse (Lagiglia, 2002), and pottery’s use in cooking might enable the boiling of corn and other vegetables
as in other Andean populations (Bruhns, 1994). Around
1,500 years BP, agriculture was of primary importance
in the economy of the northern group (Gambier, 1993).
Settlement patterns changed, revealing the formation of
permanent and semipermanent villages. In the northern
Mendoza province, the concentration of archaeological
assemblages and their association with collective mills
suggest an increase in sedentism (Lagiglia, 2002). In
San Juan province, the remains of small villages close to
river valleys were found, which showed water-control
features and dwellings for animals (Gambier, 1993). It is
important to state that northern farmers maintained a
broad-spectrum diet, with some reliance on hunting and
gathering. Nevertheless, at around 1,500 years BP, these
activities provided complementary resources, mainly
during the winter season (Gambier, 1993). Pastoralism
was more important for transport than for food. The use
of wild vegetables was reported for the Huarpes, although confined to the preparation of breads and alcoholic drinks (Gambier, 1993).
In the southern subarea, groups displayed mobile and
seasonal hunting and gathering up to historical times.
Cultigens (Zea mays) dated between 1,900–2,200 years
BP were found (Hernández, 2002). However, they do not
335
ARGENTINE CRANIOFACIAL DIFFERENTIATION
indicate the development of agriculture (Gil, 2002);
instead, their caltigens were probably obtained by exchanges
with northern populations. Animal domestication was
never adopted. Pottery began to be incorporated around
2,000 years BP (Lagiglia, 2002). Flat and small mills
were found, but they lack the traces induced by the
grinding of grains, which led some authors (Castro and
Tarragó, 1992; Lagiglia, 1997) to think that mills were
only used for processing pigments. Scrapers (used for skin
preparation) are a diagnostic element of habitat occupation, and since they are broadly dispersed, they suggest high mobility and short-term occupations (Lagiglia,
2002).
Stable-isotope analyses of human bone (Gil, 2003; Novellino et al., 2004) indicate a small proportion of cultivable plants’ consumption in the southern subarea.
Hunter-gatherer skeletons show biological markers of
diet (dental wear and presence of caries) and health
(porotic hyperosthosis and dental hypoplasia) with a similar pattern to those observed in other hunter-gatherers
(Novellino et al., 1996; Novellino, 2002). In contrast,
northern farmers show significant differences from
southern hunter-gatherers due to a higher frequency of
caries (Novellino and Guichón, 1997–1998), which may
indicate the increased role of carbohydrates in their diet
(Larsen, 1995). Hunter-gatherers also show a higher
degree of dental wear than farmers, which may indicate
the incidence of greater masticatory loads and/or abrasion. These differences in dental wear are most likely
due to differences in masticatory stresses, because farmers were also as affected by dental abrasion as huntergatherers due to the inclusion of particles of sand in the
food, which was promoted by the use of mills (Lagiglia,
2002). Once the farmers of the Argentine Center-West
acquired new techniques of cooking, such as grinding
and boiling with the use of mills and pottery, which was
a common practice among farmers of the Andean region
(Bruhns, 1994), they consumed a softer diet than
hunter-gatherers.
THE FUNCTIONAL PARADIGM IN
CRANIOFACIAL STUDIES
In the 1960s, the ‘‘functional paradigm’’ appeared in
the study of craniofacial growth, in opposition to the
‘‘genomic paradigm’’ that pointed out the preeminence of
genes in the expression of growth patterns (Carlson,
1999). The paradigm was developed by Melvin Moss,
who postulated the functional matrix hypothesis (Moss,
1973, 1997a–c), which states that cranial shape reflects
its primary functions of support and protection of the
related functional tissues and spaces (Moss and Young,
1960). Each function of the head, such as digestion and
vision, is performed by a functional cranial component
comprised of the functional matrix and the skeletal unit
(Moss, 1973; Carlson, 1999). All soft tissues, organs, and
cavities necessary to carry out a function comprise the
functional matrix, classified as: 1) periosteal matrix
(muscles and neurovascular structures of the direct functional environment), and 2) capsular matrix (cavities
and larger organs such as the brain and eyes). The
group of hard tissues (bone and cartilage) and others
(tendon and ligaments) that give biomechanical support
to the functional matrix comprises the skeletal unit,
classified as: 1) the microskeletal unit, which expresses
the constraints made by the periosteal matrix, such as
tuberosities or ridges for muscle attachment; and 2) the
TABLE 1. Samples derived from Argentine Center-West
Females,
n (%)
Males,
n (%)
Farmers not deformed
Farmers deformed
13 (10.5)
20 (16.1)
7 (5.6)
18 (14.5)
Hunter-gatherers
not deformed
Hunter-gatherers
deformed
Total
18 (14.5)
31 (25.0)
5 (4.0)
12 (9.6)
56 (45.1)
68 (54.8)
Total,
n (%)
20
38
58
49
(16.1)
(30.6)
(46.7)
(39.5)
17 (13.7)
66 (53.2)
124 (100)
macroskeletal unit, which expresses the constraints
made by the capsular matrix and associated skeletal
structures.
The functional matrix hypothesis proposes that bone
does not regulate the rate and direction of its growth by
means of its own genetic control. Instead, bone is epigenetically modified by the growth of the functional matrix
associated with it (Moss, 1973, 1997c). Thus, each component is relatively independent in form (size and shape)
and spatial position (Moss and Simon, 1968). Due to the
particular association of functional matrices and skeletal
units, components can be classified as: 1) contiguous (a
single skeletal structure and different functional matrices, such as the mandible), and 2) adjacent (different
skeletal structures associated with the same functional
matrix, such as the zygomatic arch and the mandibular
angle) (Moss and Simon, 1968).
Hunter-gatherers and farmers of the Argentine Center-West might differ in craniofacial morphology. Differences could evolve due to environmental factors, stochastic mechanisms, and differential gene flow with populations from outside the Argentine Center-West. Among
environmental factors, the subsistence economy is the
most important, since geography and ecology throughout
the northern and southern subareas are quite similar.
Biological consequences of the transition to food production are usually attributed to a decline in the quality of
nutrition and reduction in workload (Cohen and Armelagos, 1984; Ruff et al., 1984; Ruff, 1987; Larsen, 1995).
The effect of diet-related factors on craniofacial structures
was demonstrated by comparative (e.g., Carlson and Van
Gerven, 1979; Hinton, 1983; Varrela, 1992; Kaifu, 1997;
Sardi et al., 2004) and experimental (e.g., Beecher et al.,
1983; Lieberman et al., 2004) analyses. A general hypothesis to be tested in this study is that diet-related factors
intervene in the differentiation of farmers and huntergatherers of the Argentine Center-West. If this is true,
differentiation should not be stochastically distributed
throughout the skull. Instead, it should involve specific
functional components, mainly those that participate in
mastication and reflect differences in workload on the
skull due to differences in food consistency.
MATERIALS AND METHODS
Samples
Adult skulls of both sexes, derived from northern and
southern subareas of the Argentine Center-West, were
analyzed (Table 1, Fig. 1). The material is housed at the
Museo de Historia Natural (San Rafael), Museo Etnográfico (Buenos Aires), and Museo de La Plata (La Plata) in
Argentina. All individuals present closure of the sphenooccipital synchondrosis (Buikstra and Ubelaker, 1994).
336
M.L. SARDI ET AL.
TABLE 2. Functional matrix of components and interlandmark measurements1
Component
Functional matrix
Anteroneural
Neural structures related to anterior
cranial fossa (mainly anterior lobes)
Midneural
Neural structures related to middle and
part of posterior cranial fossae and most
parts of brain hemispheres
Cerebellum
Posteroneural
Otic
Bones and organs for hearing
and equilibrium
Optic
Ocular globe
Respiratory
Cavity for respiration and smell
Masticatory
Temporal and part of masseter muscles
Alveolar
Teeth and tissues of oral cavity
Interlandmarks measurements
L: glabella-bregma2
W: pterion-pterion
H: bregma-hormion
L: bregma-lambda2
W: eurion-eurion
H: basion-bregma
L: opisthion-opisthocranium2
W: asterion-asterion
H: lambda-opisthion2
L: posterior-inferior limit of tympanic bone,
to midpoint of inner extreme of petrous bone
W: external auditive meatus width2
H: external auditive meatus height2
L: dacryon-optic foramen
W: dacryon-ectoconchion
H: midpoint of supraorbitary border to midpoint
of infraorbitary border
L: subspinale-posterior nasal spine
W: widest extension of anterior nasal aperture
H: nasion-subspinale
L: zygomaxillare-posterior border of glenoid cavity2
W: anterior sulcus of sphenotemporal crest-lower point
of zygotemporal synchondrosis2
H: lower border of zygotemporal synchondrosis-upper
temporal line at coronal intersection2
L: external prosthion-posterior alveolar border2
W: from left to right alveolar borders, at unions
between second and third molars
H: intermaxillary synchondrosis-alveolar border,
at unions between second and third molars2
1
L, length; W, width; H, height.
Projected measurements. They must be done in relation to auricular-infraorbitary equalization (Frankfurt plane). In contrast,
direct measurements may be made out of Frankfurt orientation.
2
Sex determinations were done on the pelvis and skull,
following the standards described by Buikstra and Ubelaker (1994): ventral arch, subpubic concavity, ischiopubic
ramus ridge, sciatic notch, preauricular sulcus (on the
pelvis), nuchal crest, mastoid process, supraorbital margin, and glabellar and mental eminences (on the skull).
The sample from the southern subarea is represented
by 66 adult skulls. Seventeen individuals present occipital artificial deformation (Table 1). Individuals were
found in three ecological zones: mountains (n ¼ 3), piedmont (n ¼ 4), and plains (n ¼ 59) (Fig. 1). Thirty-five
percent of the sample was provided by the ‘‘Jaime Prats’’
cemetery, with two dates at 2,040 6 120 and 1,755 6 80
years BP (Novellino and Guichón, 1999). Archaeological
assemblages of the whole sample express that individuals displayed a mobile hunter-gatherer economy. For at
least half of the sample, assemblages do not include pottery (Novellino et al., 1996).
The sample derived from the northern subarea is represented by 58 skulls. Among them, 38 individuals present
occipital and front-occipital deformation (Table 1). They
were recovered from localities in mountains (n ¼ 43),
piedmont (n ¼ 8), and plains (n ¼ 7) (Fig. 1). There are
no radiocarbon dates associated with this material; thus,
chronology was inferred through archaeological indicators, such as pottery, dwellings, and the presence of irrigation systems. They can most likely be assigned to the
Late Agriculturalist period (500–800 BP) (Gambier,
1993) and to the ethnographic Huarpes (Lehmann Nitsche, 1910).
A third sample was added as a reference to measure
internal variation of samples of the Argentine Center-
West. These individuals of known age and sex derive
from a cemetery in the city of Coimbra (Portugal), born
during the last part of the 19th century and studied by
Rosas and Bastir (2002), Albanese (2003), and Sardi and
Ramı́rez Rozzi (2005), among others. The sample comprises 102 females and 100 males of Portuguese origin,
between 18–39 years of age, with closure of the sphenooccipital synchondrosis. They are housed at the Museu
Antropologico of Coimbra (Coimbra, Portugal).
Craniometric method
The morphological assessment was based on the functional matrix hypothesis, as applied in previous studies
(Pucciarelli et al., 1990; Sardi et al., 2004; González-José
et al., 2005). The neurocranium and face are major components of the skull. The neurocranium was divided into
four functional components: anteroneural, midneural,
posteroneural, and otic. Another four comprise the face:
optic, respiratory, masticatory, and alveolar (Table 2).
Length, width, and height were measured for each
component with spreading, sliding, coordinate, and Poech
calipers.
In this method, no component is over- or underrepresented because each has the same quantity of measurements, expressing three-dimensional changes which render the method sensitive to more aspects of cranial variation (O’Higgins, 1989). Another property of the method
is that interlandmark measurements do not overlap
among them. Thus, the correlation matrix of measurements represents ‘‘organic’’ correlations, because it measures associations between different cranial regions,
ARGENTINE CRANIOFACIAL DIFFERENTIATION
337
TABLE 3. Methods for estimating absolute and relative size of components
Indices
Volumetric (VI)
Neural morphometric
Facial morphometric
Difference between means
Equation
p
3ffi
(length 3 breadth 3 height)
Neural VI/SVI anteroneural, midneural, posteroneural, and otic
Facial VI/SVI optic, respiratory, masticatory, and alveolar
(Mean of hunter-gatherers’ VI mean of farmers’ VI)/[(mean of hunter-gatherers’ VI þ
mean of farmers’ VI)/2]
rather than ‘‘spurious’’ correlations which reflect redundant measurements of the same structure (Armelagos
and Van Gerven, 2003). Moreover, since different functional matrices have relatively independent embryological origins and growth patterns, specific coordinates
should measure them. This method is more suitable than
the macromeasurements commonly used in biological
anthropology to identify which region shows the greatest
amount of variation.
Data analyses
Assessment of within-samples variation. Since some
factors like sex and deformation can inflate variance
within samples, contingency tables were constructed to
evaluate if dependency existed among these factors.
Another factor that inflates variances is artificial deformation, which also affects morphology (Anton, 1989), even
in those structures that do not undergo deformation
directly (Anton and Weinstein, 1999). To test whether
deformation had an influence or not on within-samples
variation, linear measurements of each component were
compared after size-correction through the Q-standardization proposed by Darroch and Mossiman (1985). Shape
changes of functional components between deformed and
undeformed skulls were evaluated on pooled samples and
on each sample by means of discriminant analysis. Components that were not affected by deformation were able
to be included in further comparisons.
Another factor of variation among individuals within a
subarea may be their ecological origin (mountains, piedmont, and plains) or localities, and even the distribution
of sex with respect to localities. Since some localities provided very few individuals, it is not possible to perform
statistical tests on hypotheses about differentiation
among individuals according to their ecological distribution or the distribution of sex across localities. Nevertheless, the internal variation of samples of the Argentine
Center-West was assessed, taking a third sample (Coimbra) as a reference. The hypothesis about the equality of
variances of linear measurements was evaluated with
Bartlett’s test. In this comparison, all individuals were
included after a z-standardization within sexes, which is
a common method to remove sex-related size variation
(Williams-Blangero and Blangero, 1989; Relethford,
2001). The Coimbra sample can be considered homogenous with respect to ethnicity, geography, and temporal
range. If samples of the Argentine Center-West do not
differ in variances with respect to the Coimbra sample,
then it is possible to deduce that such factors that may
inflate internal variance are not important enough to
affect results, and it would be justifiable to pool skulls
together for comparisons between both subareas.
Assessment of between-samples variation. Two types
of indices were calculated with measurements (Pucciarelli et al., 1999; Sardi, 2002; González-José et al., 2005)
(Table 3): 1) volumetric indices, represented by the geo-
metric mean of three dimensions, estimate the absolute
size of functional components; and 2) morphometric indices, represented by the proportion between a given volumetric index and the sum of volumetric indices that
belong to the neurocranium or face, estimate the relative
size of components. A paired t-test was carried out with
these indices to evaluate differences between means of
farmers and hunter-gatherers, and F-ratios were calculated to evaluate the equality of variances. In order to
assess relative size between samples, differences
between means were calculated with volumetric indices
(Table 3). In order to know which dimensions intervened
in the components’ differentiation, discriminant analyses
with a complete estimation (i.e., including the three variables of each component) were performed on sex-pooled
samples after a z-standardization within sexes. In those
components that showed significant F-values (P < 0.05),
the discriminant analyses were remade with backward
stepwise estimation to establish which measurements
were useful for discrimination between both samples.
The alpha of F-values to enter or remove variables from
functions was set at 0.05.
RESULTS
Within-samples variation
Dependence between area, sex, and deformation.
Table 4 indicates that cranial deformation is a factor
dependent on subarea, showing higher frequencies
among farmers (Table 1), but it is uniformly distributed
across the sexes. The sex factor shows a significant
dependence on subareas (Table 4), because females are
more frequent among farmers (Table 1).
Cranial deformation effect. Results shown in Table 5
indicate that artificial cranial deformation did not influence the shape of facial components. Skulls underwent
deformation on the frontal and occipital bones. Such an
effect would not affect the cranial base, which directly
influences facial growth (Hilloowalla et al., 1998; Lieberman et al., 2000), not modifying facial components, but
modifying the midneural and otic components (Table 5).
Since deformed crania showed differences in the shape
of neural components, they were excluded from the following analyses. However, they were included in analyses of facial components, because the face was not
affected by deformation.
Homogeneity of variances. Most variances (19/24) of
samples of the Argentine Center-West and Coimbra were
homeocedastic (Table 6). Heterocedastic measurements
included midneural width (P < 0.05), posteroneural
length (P < 0.05), posteroneural width (P < 0.01), respiratory length (P < 0.05), and masticatory width (P <
0.01). Note that Coimbra showed the greatest variances
(Table 7) for most of the heterocedastic measurements.
These results indicate that there was no specific factor
338
M.L. SARDI ET AL.
TABLE 4. Tests of dependence between factors (subarea, sex, and deformation)1
Subarea vs. presence/absence of deformation
Subarea vs. sex
Sex vs. presence/absence of deformation in farmers
Sex vs. presence/absence of deformation in hunter-gatherers
Sex vs. presence/absence of deformation across subareas
Subarea vs. sex of nondeformed skulls
Subarea vs. sex of deformed skulls
1
n
df
v2
P-value
124
124
58
66
124
69
55
1
1
1
1
3
1
1
18.19
5.20
0.39
0.06
7.14
3.51
1.70
0.000
0.023
0.532
0.802
0.068
0.061
0.192
For comparisons with df ¼ 1, Yates-corrected v2 were calculated.
TABLE 5. Effect of cranial deformation by discriminant analyses with Q-standardized variables
Pooled
Component
Anteroneural
Midneural
Posteroneural
Otic
Optic
Respiratory
Masticatory
Alveolar
Hunter-gatherers
P-value
F
P-value
F
P-value
5.82
29.08
16.85
0.55
2.21
0.55
2.46
1.36
0.001
0.000
0.000
0.646
0.089
0.644
0.065
0.259
1.47
15.69
11.36
4.71
0.18
0.49
1.46
0.49
0.230
0.000
0.000
0.005
0.910
0.688
0.232
0.685
4.23
12.01
5.99
0.61
1.25
0.67
1.40
2.65
0.009
0.000
0.001
0.608
0.298
0.570
0.253
0.057
TABLE 6. Barttlet’s test to evaluate equality of variances in
samples of Argentine Center-West and Coimbra
Anteroneural length
Anteroneural width
Anteroneural height
Midneural length
Midneural width
Midneural height
Posteroneural length
Posteroneural width
Posteroneural height
Otic length
Otic width
Otic height
Optic length
Optic width
Optic height
Respiratory length
Respiratory width
Respiratory height
Masticatory length
Masticatory width
Masticatory height
Alveolar length
Alveolar width
Alveolar height
Farmers
F
Bartlett’s v2
P-value
0.41
1.84
2.50
5.76
6.94
0.51
8.96
17.91
1.74
1.11
1.37
2.90
0.54
3.55
1.17
7.21
0.56
0.79
5.58
17.83
4.73
2.57
2.70
0.73
0.814
0.397
0.286
0.056
0.031
0.773
0.011
0.000
0.419
0.574
0.503
0.234
0.764
0.169
0.556
0.027
0.755
0.673
0.061
0.000
0.094
0.276
0.259
0.693
that inflated variances in samples of the Argentine Center-West, such as locality or ecological zone, enabling the
samples to be compared.
TABLE 7. Variances of measurements that differed between
groups with Bartlett’s test
Midneural width
Posteroneural length
Posteroneural width
Respiratory length
Masticatory width
Hunter-gatherers
Farmers
Coimbra
0.69
0.53
0.57
1.07
1.07
0.57
0.25
0.25
0.57
0.64
1.09
0.74
1.08
1.03
0.48
farmers (Table 8). With respect to facial components,
hunter-gatherers showed highly significant smaller respiratory and bigger masticatory indices than farmers.
The pattern of differentiation in both sexes was quite
similar: females differed by the midneural, optic, and
masticatory volumetric indices, and the respiratory and
masticatory morphometric ones; males differed by the
otic and masticatory volumetric indices, and by the anteroneural, posteroneural, and masticatory morphometric ones (Table 8). Figure 2 presents differences between
means of volumetric indices, where the masticatory and
posteroneural components showed the highest values.
Significant discriminant functions were calculated for
the midneural, posteroneural, otic, optic, and masticatory components (Table 9). Measurements retained in
the discriminant functions after backward stepwise estimations were lengths and widths, which showed higher
values in hunter-gatherers than in farmers (Table 9).
The otic and optic components differentiated farmers
and hunter-gatherers, whereas their respective volumetric indices did not change (Table 8).
Between-samples variation
DISCUSSION
Volumetric indices showed that hunter-gatherers’
skulls were bigger for most of the components (Table 8).
When the sexes were pooled, the midneural, posteroneural, and masticatory volumetric indices were larger in
hunter-gatherers. Morphometric indices varied, because
hunter-gatherers presented a significantly smaller anteroneural index, and a bigger posteroneural one than
Results indicate that farmers present a smaller size
for most components. The masticatory and posteroneural
components show the greatest differences, with farmers
presenting an absolutely and relatively smaller size than
hunter-gatherers (Table 8, Fig. 2), mainly due to a
reduction in length and width (Table 9). Differential
gene flow with populations from outside the Argentine
339
ARGENTINE CRANIOFACIAL DIFFERENTIATION
TABLE 8. Paired t-test and F-ratios for comparing means and variances, respectively, of volumetric and
morphometric indices (hunter-gatherers farmers)
Volumetric indices
Pooled sexes
Anteroneural
Midneural
Posteroneural
Otic
Optic
Respiratory
Masticatory
Alveolar
Females
Anteroneural
Midneural
Posteroneural
Otic
Optic
Respiratory
Masticatory
Alveolar
Males
Anteroneural
Midneural
Posteroneural
Otic
Optic
Respiratory
Masticatory
Alveolar
Morphometric indices
t
P-value
F
P-value
t
P-value
F
P-value
0.25
3.08
3.95
0.54
1.60
0.55
4.99
0.71
0.804
0.003
0.000
0.585
0.111
0.578
0.000
0.480
1.00
1.39
2.47
1.01
1.04
1.39
1.45
1.24
1.000
0.437
0.034
1.000
0.861
0.204
0.149
0.407
2.53
0.46
2.78
1.87
1.91
3.05
4.51
0.88
0.013
0.647
0.007
0.065
0.057
0.002
0.000
0.380
1.58
1.61
1.55
1.18
1.78
1.42
1.07
1.21
0.275
0.256
0.297
0.712
0.027
0.173
0.777
0.442
0.80
2.76
1.90
0.71
2.30
0.84
4.56
0.89
0.430
0.009
0.067
0.480
0.025
0.405
0.000
0.374
1.46
2.07
4.28
1.03
1.04
1.65
1.99
1.04
0.510
0.203
0.014
0.977
0.944
0.189
0.074
0.890
1.13
0.74
0.89
1.84
1.63
2.74
3.58
0.57
0.266
0.462
0.381
0.076
0.107
0.008
0.000
0.567
2.54
2.28
2.61
1.33
2.32
1.17
1.23
1.31
0.106
0.149
0.096
0.617
0.029
0.674
0.576
0.511
0.53
1.72
4.24
0.03
0.15
0.00
2.87
0.15
0.600
0.093
0.000
0.973
0.877
0.999
0.005
0.877
1.58
1.27
6.79
1.10
1.05
1.20
1.02
1.47
0.371
0.595
0.023
0.767
0.923
0.632
0.975
0.311
2.73
0.13
3.59
0.84
1.15
1.73
2.98
0.68
0.009
0.891
0.000
0.406
0.255
0.087
0.004
0.494
1.25
1.02
1.98
1.12
1.42
1.66
1.10
1.17
0.844
1.000
0.398
0.742
0.363
0.186
0.767
0.638
Fig. 2. Percent differences between means (DM) of volumetric indices between hunter-gatherers and farmers. Positive values indicate greater mean value in hunter-gatherers.
Center-West, stochastic mechanisms, and factors associated with the subsistence economy might explain this
pattern.
The fact that the Argentine Center-West samples
showed low variances (Tables 6 and 7) with regard to
Coimbra enables us to discard differential gene flow with
populations from outside the Argentine Center-West as a
main factor in the increase of differentiation between
hunter-gatherers and farmers. On the other hand, other
factors could increase variation in Coimbra, such as less
canalization, evoked by the low biomechanical demands
associated with industrialization. Gene flow with morphologically different populations of adjacent areas
might exist in the Argentine Center-West. However, it
was expected that if gene flow as well as stochastic
mechanisms affected morphology, then the pattern of differentiation should involve many components, without
any dynamic relationship among them. In this study, the
main between-samples differentiation occurred in some
localized components, and it is better explained by systemic and/or localized factors associated with both subsistence economies.
Greater skeletal size can be attained by the volumetric
growth of functional matrices and/or by deposition of
skeletal tissue (Moss et al., 1987). Among systemic factors, differences in the composition of diet and hormonal
circulation would account for differentiation. Cordain
(2000) estimated that most worldwide hunter-gatherer
groups obtain more than 50% of their diet from animal
foods, which are associated with a high percentage of
energy derived from proteins, at about 19–35%. With a
greater dependence on agriculture, the accessibility of
animal proteins was reduced, as reported for many parts
of the Andean region (Castro and Tarragó, 1992). Likewise, the higher frequency of caries in farmers than
hunter-gatherers of the Argentine Center-West (Novellino and Guichón, 1997–1998) suggests an increased role
of carbohydrates in the diet. Thus, a diet more deficient
in proteins could lead to the smaller size in farmers, as
occurred in some other worldwide regions (Cohen and
Armelagos, 1984). Experimental models offered insights
into the effects of low-protein diets acting upon phenotypes, suggesting that these diets were associated with
size reduction in many cranial and postcranial variables
(Pucciarelli, 1980, 1981; Pucciarelli and Goya, 1983; Pucciarelli et al., 1990; Dressino and Pucciarelli, 1999;
Miller and German, 1999; Reichling and German, 2000).
However, although the protein content was lower in the
farmer diet than in the hunter-gatherer diet, it is not
possible to state that the farmers’ diet was so poor in
proteins as were diets of those experimental models,
which were under 10% of protein content.
340
M.L. SARDI ET AL.
TABLE 9. Discriminant analyses performed with measurements of components
Backward stepwise estimation
Complete estimation
F
P
Anteroneural
Midneural
Component
0.27
5.37
0.844
0.002
Posteroneural
Otic
7.43
3.39
0.000
0.023
11.08
2.15
12.04
0.000
0.097
0.000
1.01
0.388
Optic
Respiratory
Masticatory
Alveolar
Variables retained with
alpha ¼ 0.05
Length, F ¼ 8.07
Width, F ¼ 7.74
Width, F ¼ 16.26
Length, F ¼ 5.24
Width, F ¼ 6.33
Length, F ¼ 29.14
Length, F ¼ 12.66
Width, F ¼ 6.64
Hormonal action may be linked to differences in mobility. Weltman et al. (2001) found a direct association
between growth hormone circulation and physical activity.
In humans, the circulation of growth hormones was
recorded from birth (Geary et al., 2003). Growth hormone
promotes the increment in skeletal (Vogl et al., 1993; Barr
and McKay, 1998; Banu et al., 2001; Forwood et al., 2001)
and muscle mass (Vogl et al., 1993), and it influences craniofacial growth (Oyhenart and Pucciarelli, 1992; Cónsole
et al., 2001). The settlement pattern in the northern subarea, the greater concentration of archaeological material,
and the use of animals for transport revealed that farmers were less mobile than hunter-gatherers. But if the
reduced circulation of growth hormone led to a size reduction in farmers, then farmers should present greater
reduction in the face than in the neural skull, because
facial structures are under greater hormonal influence.
Moreover, the neural skull grows at high rates at early
infancy, and differences in mobility between hunter-gatherers and farmers during this ontogenetic period seem
unlikely. However, hormonal influence in a certain proportion of facial variation, at least, cannot be completely
excluded. Hormonal factors might interact with nutritional factors, since the mechanisms that regulate growthhormone secretion are sensitive to nutritional status. In
this sense, Cónsole et al. (2001) found that a low-protein
diet in monkeys induced a decrease in growth hormone
and prolactin cell populations, resulting in changes of craniofacial morphology.
Among localized factors, mechanical loading acting
upon the masticatory complex and neck due to differences
in food consistency between hunter-gatherers and farmers
could promote differentiation. The bigger masticatory size
of hunter-gatherers may be the result of greater masticatory stress, which was reduced in farmers.
Masticatory forces regulate an important proportion of
craniofacial growth, affecting several structures (e.g.,
Hannam and Wood, 1989; van Spronsen et al., 1991;
Kiliaridis, 1995). According to Raadsheer et al. (1999),
the magnitude of masticatory forces is positively correlated with muscular thickness and facial measurements.
Kiliaridis (1995) proposed that masticatory hyperfunction
led to facial size enlargement through sutural growth
and bone apposition. Experimental studies suggest that
the mastication of soft diets contributes to many
changes, such as a reduction in cortical bone thickness
(Bresin et al., 1999), shortening of maxillary arches and
deformation of the palate (Beecher et al., 1983), reduction of muscular size (Ciochon et al., 1997), modifications
Mean of scores obtained with
discriminant function
Farmers
Hunter-gatherers
0.73
0.30
0.76
0.61
0.31
0.25
0.51
0.45
0.57
0.50
of the mandibular condile (Giesen et al., 2003), and
lesser growth, particularly in transverse dimensions and
in posterior portions of the skull (Lieberman et al.,
2004). Significant modifications located in the masticatory component were reported. Analyses of minipigs
showed that the masseter pulls on the zygomatic bone
during mastication, and great mechanical loadings are
produced in the zygomatico-squamosal suture and bone
surfaces (Rafferty et al., 2000; Herring et al., 2001). In
hyraxes, the highest strains generated by processing
hard food are located mainly in the zygomatic arch (Lieberman et al., 2004).
Comparative studies in temporal scales (Carlson and
Van Gerven, 1979; Varrela, 1992; Kaifu, 1997; Sardi
et al., 2004) are quite consistent with experimental models and with the findings of this study. Carlson and Van
Gerven (1979) compared Mesolithic and later groups of
Lower Nubia, and found that later groups presented a
reduction in size of the masticatory muscles and teeth, a
reduction in facial growth, changes in facial position, and
a cranial vault with a relatively shorter and higher shape.
Sardi et al. (2004) compared samples of Europe and North
Africa, and observed that post-Mesolithic groups were characterized by smaller size, narrower faces, and more reduced
masticatory volume than Paleolithic and Mesolithic huntergatherers. It is possible that the shift observed in both studies (Carlson and Van Gerven, 1979; Sardi et al., 2004) was
due to farmer demic diffusion and population replacement
(Ammerman and Cavalli-Sforza, 1984; Turner and Markowitz, 1990). However, the fact that differentiation is located
in specific structures supports the proposal that diet-related
factors account for at least part of the craniofacial variation.
Other studies (Smith, 1979; Lahr and Arensburg, 1995) proposed that a decrease of workload on the face during the
Neolithic transition was associated with a tendency toward
brachycephalization.
Masticatory forces can also explain the posteroneural
differentiation. Herring and Teng (2000) and Herring
et al. (2001) analyzed minipigs, and demonstrated that
the contraction of the masseter and temporalis during
natural mastication caused strains in some sutures of
the braincase. Since minipigs move their heads during
mastication, the neck muscles were also affected (Herring and Teng, 2000). In humans, a similar high degree
of coordination between concomitant mandibular and
head-neck movements during natural jaw activities was
reported (Zafar et al., 2000). This functional relationship
apparently relies on common neural connections that
control activities in both systems (Igarashi et al., 2000;
ARGENTINE CRANIOFACIAL DIFFERENTIATION
Zafar et al., 2000). Thus, the bigger postneural size in
hunter-gatherers may reflect the greater masticatory
activity on neck muscles, which are attached to the occipital bone.
CONCLUSIONS
The above results support the hypothesis that craniofacial differentiation between farmers and hunter-gatherers of the Argentine Center-West was affected by dietrelated factors. Farmers presented a smaller craniofacial
size, and showed absolute and relative reductions of the
masticatory and posteroneural components with respect
to hunter-gatherers. Two systemic factors, 1) the lower
quality of nutrition due to low protein content in the
diet, and 2) the diminution of growth hormone circulation because of lesser mobility due to sedentism, and a
localized factor, the reduction of masticatory stress and
workload on the head and neck, could generate this pattern. Although nutritional and hormonal factors can
account for a certain proportion of craniofacial size variation, the pattern of differentiation seems to suggest
localized rather than systemic factors. The size reduction
of the masticatory and posteroneural components in
farmers can be attributed to lesser muscular loadings
due to the softer consistency of the diet consumed after
the transition to food production.
ACKNOWLEDGMENTS
We thank our reviewers for their contributions to the
improvement of the manuscript; Rolando González-José
and Fernando Ramı́rez Rozzi, for useful commentaries
that helped clarify some aspects of our argument; Vı́ctor
Durán, Humberto Lagiglia, Gabriela Raviña, and Marta
Roa, for assistance in archaeological aspects; Pablo
Sardi, Rodrigo Lacruz, and Marı́a C. Muñe, for technical
assistance; and the curators of the collections for access
to specimens in their care. This work was made possible
by grants from the Universidad Nacional de La Plata
(Argentina) and the Fondation Fyssen (France) to M.L.S.
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