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Body proportions in ancient Andeans from high and low altitudes.

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AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY 128:569 –585 (2005)
Body Proportions in Ancient Andeans From High and
Low Altitudes
Karen J. Weinstein*
Department of Anthropology, Dickinson College, Carlisle, Pennsylvania 17013
KEY WORDS
limb proportions; ecogeographic variation; high altitude; Andes
ABSTRACT
Living human populations from high altitudes in the Andes exhibit relatively short limbs compared with neighboring groups from lower elevations as
adaptations to cold climates characteristic of high-altitude
environments. This study compares relative limb lengths
and proportions in pre-Contact human skeletons from different altitudes to test whether ecogeographic variation
also existed in Andean prehistory. Maximum lengths of
the humerus, radius, femur, and tibia, and femoral head
breadth are measured in sex-specific groups of adult human skeletons (N ⫽ 346) from the central (n ⫽ 80) and the
south-central (n ⫽ 123) Andean coasts, the Atacama
Desert at 2,500 m (n ⫽ 102), and the southern Peruvian
highlands at 2,000 –3,800 m (n ⫽ 41). To test whether limb
lengths vary with altitude, comparisons are made of intralimb proportions, limb lengths against body mass estimates derived from published equations, limb lengths
against the geometric mean of all measurements, and
principal component analysis. Intralimb proportions do
not statistically differ between coastal groups and those
from the Atacama Desert, whereas intralimb proportions
are significantly shorter in the Peruvian highland sample.
Overall body size and limb lengths relative to body size
vary along an altitudinal gradient, with larger individuals
from coastal environments and smaller individuals with
relatively longer limbs for their size from higher elevations. Ecogeographic variation in relation to climate explains the variation in intralimb proportions, and dietary
variation may explain the altitudinal cline in body size
and limb lengths relative to body size. The potential effects of gene flow on variation in body proportions in
Andean prehistory are also explored. Am J Phys Anthropol 128:569 –585, 2005. © 2005 Wiley-Liss, Inc.
Evolutionary biologists have long shown that ecogeographic patterns, such as the rules of Bergmann (1847)
and Allen (1877), explain much of the variation in body
size and shape among geographically widespread homeothermic species (e.g., Mayr, 1963; Porter and Gates, 1969;
Searcy, 1980; Aldrich and James, 1991; Graves, 1991;
Ashton et al., 2000). Populations from cold climates and
high latitudes tend to be heavier with relatively short
appendages compared with their conspecifics from
warmer climates and lower latitudes. Heavier, shorterlimbed individuals have a smaller surface area to body
mass ratio, and thus are better equipped at conserving
heat under the stress of cold climatic conditions. Conversely, a larger surface area to body mass ratio enables
taller, thinner individuals to shed body heat under hot
climatic conditions (e.g., Calder, 1984; Schmidt-Nielson,
1984).
Variation in body size and proportions in past and
present human groups generally conforms to these ecogeographic patterns. Among living human populations worldwide, body mass and relative sitting height are negatively
correlated with mean annual temperature and absolute
latitude (Schreider, 1950, 1957, 1964, 1975; Roberts, 1953,
1978; Ruff, 1991, 1994; Katzmarzyk and Leonard, 1998).
Mean annual temperature also explains regional variation in body mass and proportions among living human
populations from Africa (e.g., Hiernaux and Froment,
1976), Europe and the Mediterranean (e.g., Crognier,
1981), and North and South America (e.g., Newman, 1953;
Newman and Munro, 1955; Stinson, 1990). Ecogeographic
patterns in body size and shape have also been central in
discerning climatic adaptations in human skeletal re-
mains. Relative limb proportions and bi-iliac breadth in
modern human skeletons recovered from diverse geographical regions and climatic zones are correlated with
mean annual temperature and latitude (Trinkaus, 1981;
Jacobs, 1985, 1993; Ruff, 1991, 1994; Holliday, 1997). The
association between body proportions and climate, furthermore, is used to infer climatic adaptations and phylogenetic relationships among Homo groups during the
Pleistocene (Trinkaus, 1981; Ruff, 1991, 1994, 2002b; Ruff
and Walker, 1993; Ruff et al., 1997; Holliday, 1997, 1999,
2000; Rosenberg et al., 1999; Pearson, 2000). In particular, the long, linear body shape of the earliest modern
humans in Europe and the cold-adapted body shape of
Neandertals suggest that these two contemporaneous
groups had separate origins, despite living in close prox-
©
2005 WILEY-LISS, INC.
Grant sponsor: The Leakey Foundation; Grant sponsor: Nutter
Dissertation Fellowship, College of Liberal Arts and Sciences, University of Florida; Grant sponsor: Tinker Field Research Grant, Center for Latin American Studies, University of Florida; Grant sponsor:
Summer Scholar Award, Dickinson College Research and Development Committee.
*Correspondence to: Karen J. Weinstein, Department of Anthropology, Dickinson College, PO Box 1773, Carlisle, PA 17013.
E-mail: weinstek@dickinson.edu
Received 14 November 2003; accepted 9 July 2004.
DOI 10.1002/ajpa.20137
Published online 13 May 2005 in Wiley InterScience
(www.interscience.wiley.com).
570
K.J. WEINSTEIN
imity in Western Europe during the late Pleistocene (e.g.,
Trinkaus, 1981; Holliday, 1997, 1999, 2000).
Discerning climatic adaptations in human evolution entails comparing individual Pleistocene Homo specimens
against large skeletal samples of recent H. sapiens from
broadly distinct climatic zones. Comparative samples of
recent human skeletons are often grouped into circumpolar, temperate, subtropical, and tropical regions, and are
argued to represent the full range of modern human variation in body size and shape (Trinkaus, 1981; Ruff, 1991,
1994; Ruff et al., 1997; Holliday, 1997, 1999; Pearson,
2000). With the exception of circumpolar groups from
North America, however, the majority of comparative
skeletal samples come from the Old World, a sampling
choice that reflects the assumption that climatic adaptations have a genetic basis and therefore evolved in specific
populations over lengthy periods of time. Thus, limb proportions in Neandertals, the earliest H. sapiens, and other
Homo specimens are compared to those of cold-adapted
circumpolar recent H. sapiens on one extreme against
heat-adapted, tropical groups on the other. The plethora of
studies that yield similar results regarding ecogeographic
variation in body proportions in recent H. sapiens and
Pleistocene Homo attests to the efficacy of this comparative method and to the persuasiveness of these results.
Yet there have been no studies that examine variations
in body proportions in closely related populations across
small geographic distances or of prehistoric modern human groups from diverse environments in the Americas.
Comparisons of geographically proximate and closely related populations that differ in climate might reveal important information regarding the length of time required
to develop morphological adaptations to climate. Comparisons of geographically proximate groups, moreover,
might elucidate fine-grained clinal variations in body size
and shape that are not possible to discern from comparisons of populations separated by vast geographic and temporal distances.
For a number of reasons, pre-Contact modern human
skeletons from different altitudes in the Andes are ideal
for identifying clinal variations in body proportions and
for testing hypotheses about the development of climatic
adaptations in human prehistory. First, the South American Andes encompass multiple elevational zones within
short geographic distances. Lowland regions along the
coast of the Pacific Ocean have mild tropical to subtropical
climates. In contrast, climatic conditions at high altitudes
(elevations of 2,500 m and greater) are typically much
colder. Among many environmental stressors, ambient
temperatures at high altitudes dramatically fluctuate on a
diurnal basis, with nighttime temperatures often falling
below freezing year-round (Baker, 1960; Pawson and Jest,
1978; Barry, 1979; Billings, 1979).
The Andean archaeological record reveals a cultural
and ecological separation between coastal and highland
populations. Human populations first settled coastal and
highland regions in the Andes at 11,000 years BP (Moseley, 2001). Coastal populations exploited the rich marine
resources of the Pacific Ocean, and this maritime adaptation persisted for millennia (Moseley, 2001). Highland
populations, in contrast, practiced agropastoralism that
required “verticality,” a land-use pattern that simultaneously exploits three elevational zones: the high puna
above 4,000 m for camelid herding, mountainous valleys
at 2,500 – 4,000 m for the cultivation of tubers and hardy
grains, and elevations between 2,000 –2,500 m for the
cultivation of maize and other less resilient crops (Murra,
1972; Moseley, 2001). The absence of marine resources in
highland sites and vice versa suggest further that coastal
and highland Andean groups maintained largely separate
breeding populations until at least after 3,500 years BP
(Moseley, 2001). Thus, the rich Andean archaeological
record documents dietary variation and the degree of population interactions between coastal and highland settlements through time.
Finally, living human populations indigenous to the
Andes exhibit anthropometric variation in body size and
proportions that is correlated, in part, with climatic conditions associated with different altitudes. Similar to
other human groups from cold climates, highland Aymara
and Quechua have short limbs relative to stature compared with lowland Andean populations of comparable
socioeconomic status (Stinson and Frisancho, 1978; Palomino et al., 1979) and with other South American populations (Stinson, 1990), i.e., patterns that conform to
Allen’s rule. Whether similar differences in body shape
existed in Andean prehistory has yet to be established.
This study compares limb proportions in pre-Contact
human skeletons from different altitudes in the Andes in
order to test whether ecogeographic variation in body proportions associated with altitude applies to human populations in Andean prehistory. Considering the longevity of
human settlement in the coastal and highland Andes and
that body proportions in living Andeans vary with climate
and elevation, pre-Contact Andean peoples from different
altitudes should exhibit variation in body proportions that
reflect climatic adaptations. If this is the case, then preContact lowland groups should have relatively long limbs
for their size, while highland groups should have relatively short limbs. If factors other than climate influence
variation in body proportions, then relative limb lengths
should not vary along an altitudinal gradient. These factors may include variations in diet and subsistence-related activities, exposure to various diseases that might
affect growth, or large-scale population movements across
vast geographic regions.
MATERIALS AND METHODS
Human skeletal samples
To examine the relationship between body proportions
and altitude, four sex-specific samples of adult human
skeletons from the Andes were compared: two samples
from the Pacific coast, and two from the highlands (Table 1). The choice of skeletal samples was restricted to
large, well-preserved collections of complete or nearly
complete individuals. Time, dietary variation, and gene
flow are potentially confounding factors that may influence variation in body size and proportions. Thus, the
archaeological record and history of each of these skeletal
samples are described.
Arica, Chile (AR). This sample, recovered from two
distinct temporal periods in Arica, Chile and the nearby
Azapa Valley, represents a lowland, south-central Andean
coastal population (Table 1). The earlier specimens, which
date to the Archaic period (3210 –1720 BC; Allison et al.,
1984; Standen et al., 1984, 1997; Focacci and Chacón,
1989), are part of the Chinchorro cultural tradition and
are biologically linked with the earliest inhabitants who
settled along the south-central Andean coast at
9,000 years BP (Allison et al., 1984; Arriaza, 1995; Moseley, 2001). Faunal remains of fish, shellfish, and marine
birds and mammals (Bird, 1943; Llagostera, 1989; Villaxa
571
BODY PROPORTIONS IN ANDEAN PREHISTORY
TABLE 1. Human skeletal samples, dates, and geographic locations
Machu Picchu and Cuzco, Peru (MPC)
San Pedro de Atacama, Chile (SP)
Coyo-3, Solcor-3, and Quitor-6
Arica, Chile (AR)
Mo-1, Mo-1/5, and Mo-1/6
AZ-140
Ancón, Peru (AC)
Males
Females
22
42
19
60
54
69
52
28
Dates1
15th century
AD 250–1240
3210–1720 BC
AD 750–1250
AD 1000–1476
Lat.2
Alt.3
Location4
13.55
22.55
2,000–3,800
2,500
FMNH, YPM
MSPA
18.33
⬍100
SMA
11.78
⬍100
FMNH
1
See text for full discussion of published dates and sites.
Absolute latitude in degrees from equator (Abate, 1991).
Elevation in m above sea level (Earth Info, Inc., 1996).
4
SMA, Museo Arqueológico San Miguel de Azapa, Universidad de Tarapacá, Arica, Chile; MSPA, Museo R.P. Le Paige, San Pedro de
Atacama, Chile; FMNH, Field Museum, Chicago, IL; YPM, Peabody Museum of Natural History, Yale University, New Haven, CT.
2
3
and Corrales, 1992; Arriaza, 1995) and the high frequency
of auditory exostoses, an osteological condition associated
with repetitive immersion in cold water (Standen et al.,
1997), indicate that these Archaic period individuals subsisted primarily on the rich marine resources of the adjacent Pacific Ocean. The later specimens date to AD 750 –
1250 (Table 1; Arriaza et al., 1995; Langsjoen, 1996;
Standen et al., 1997). During this period, south-central
coastal populations practiced a mixed subsistence economy based on the cultivation of maize, beans, and tubers,
and the harvesting of marine resources (Berenguer and
Dauelsberg, 1989; Arriaza et al., 1995; Langsjoen, 1996;
Standen et al., 1997). The earlier Arica skeletons and the
later Azapa individuals are pooled into one sample because previous analysis revealed that limb lengths and
proportions do not vary between individuals from these
two time periods (Weinstein, 2001).
Ancón, Peru (AC). This skeletal sample represents a
lowland population from the central Andean coast dating
to AD 1000 –1476 (Moseley, 2001; Table 1). G.A. Dorsey
excavated these individuals from a cemetery at Ancón for
inclusion in the 1893 Columbia Exposition in Chicago,
Illinois (Dorsey, no date). These individuals were buried in
seated, flexed positions and wrapped in textiles. Based on
associated grave goods, these individuals most likely engaged in a mixed subsistence strategy based on marine
resources and agriculture (Dorsey, no date).
San Pedro de Atacama, Chile (SP). SP represents
highland individuals from 2,500 m recovered from three
cemeteries dated to AD 250 –1240 (Table 1; Costa, 1988;
Llagostera et al., 1988; Neves and Costa, 1998; Neves et
al., 1999). Although 2,500 m is at the lower limit of high
altitude, San Pedro de Atacama’s elevation is high enough
for ambient temperatures to drop below freezing, especially during the winter months (Earth Info, Inc., 1996).
Archaeological evidence also suggests that SP represents
a highland population. Studies of the paleopathological
conditions in these individuals (Costa, 1988; Neves and
Costa, 1998; Neves et al., 1999) indicate heavy reliance on
agropastoralism, a south-central Andean highland subsistence practice since approximately 4,000 years BP, which
requires verticality: the simultaneous use of elevations
from 2,000 – 4,500 m (Núñez, 1991; Moseley, 2001). In
addition, individuals in this sample were buried with
large numbers of grave goods that originated in the highlands of Bolivia and northwest Argentina, which suggests
continuous economic, cultural, and biological contact between San Pedro de Atacama populations and the high-
land regions to the east (Núñez, 1991; Neves and Costa,
1998).
Machu Picchu/Cuzco (MPC). The human skeletons
from Machu Picchu (2,000 –2,850 m) and Cuzco (3,500 –
3,800 m) comprise the second highland sample (Table 1).
These individuals date to the 15th century, and were recovered by Hiram Bingham during his 1911–1914 archaeological expeditions to southern Peru (Eaton, 1916; Bingham, 1930; Burger and Salazar, 2003). While MPC dates
to a somewhat later time than the other samples, this
temporal gap overlaps the time range for AC and is less
than 200 years younger than that of AR and SP.
The individuals from Machu Picchu were buried in caves
and rock shelters in seated, flexed positions, wrapped in
textiles and rope, and associated with llama and alpaca
remains that are presumed to have been sacrificed at the
time of the individual’s death (Miller, 2003). The burials
often contain more than one individual (Eaton, 1916; Verano, 2003), which can be explained through a variety of
taphonomic factors, including Inca rituals regarding ancestor veneration and treatment of the deceased, soil acidity
and cave humidity, and postmortem activities of carnivores
and rodents (Miller, 2003; Verano, 2003). Bone chemistry
analysis and dental pathological conditions indicate that
MPC individuals consumed a diet high in maize (Burger et
al., 2003; Verano, 2003).
Much has been speculated about the geographic origins
and ethnic identities of Inca subjects in general and the
inhabitants of Machu Picchu in particular (e.g., Eaton,
1916; Bingham, 1930; Moseley, 2001; Burger and Salazar,
2003). According to Andean archaeologists, the Inca implemented a number of social policies that functioned to
subdue their subjects and conquer new territories. For
example, the Inca would resettle entire communities into
newly colonized territories, and adult males from throughout the Andes were drafted as labor for large-scale construction projects and military service (Moseley, 2001).
Dietary variation based on the stable carbon and nitrogen
isotope content of human bone suggests that Machu Picchu individuals were not exclusively highland in origin
(Burger et al., 2003). Cranial deformation styles and the
analysis of biological distance based on craniometrics also
suggest that Machu Picchu inhabitants represented diverse populations from throughout the Andes (Verano,
2003). Multivariate craniometrics on artificially deformed
skulls, however, are not especially reliable methods for
assessing biological distance between populations, given
that cranial shape undergoes marked changes in response
572
K.J. WEINSTEIN
Fig. 1. Intralimb proportions in males: log RAD vs. log
HUM (A) and log TIB vs. log FEM (B). Here and in all subsequent figures, individual data points are labeled according to
following symbols: ●, MPC; X, SP; ⫹, AR; ‚, AC. Note common
scatter of SP, AR, and AC, while most of MPC falls below
regression line.
to artificial deformation (Antón, 1989). Nevertheless, caution is required when interpreting body proportions in this
sample. If MPC represents people from across the Andes,
then body proportions in this sample should exhibit the
largest range of variation and should overlap the ranges of
skeletal samples from other elevations. MPC individuals
who are highland in origin should cluster with SP, the
other highland sample, whereas MPC individuals who are
lowland should exhibit similar body proportions as those
from AR and AC.
Osteometric measurements
Maximum lengths of the humerus (HUM), radius
(RAD), femur (FEM) (bicondylar length), and tibia
Fig. 2. Intralimb proportions in females: log RAD vs. log
HUM (A) and log TIB vs. log FEM (B). See Figure 1 for explanation of symbols. Similar to patterns seen in males, female SP, AR,
and AC have virtually indistinguishable intralimb proportions,
whereas those of MPC tend to be shorter.
(TIB), as defined in Martin (1928), were measured to the
nearest 0.5 mm with an osteometric board, and femoral
head breadth (FHB) to the nearest 0.1 mm with Mitutoyo digital sliding calipers. FHB is defined as the average of the antero-posterior and supero-inferior diameters of the femoral head (Grine et al., 1995). Since
articular surfaces of weight-bearing joints are correlated with body weight (e.g., Ruff et al., 1991; Lieberman et al., 2001), FHB provides an appropriate variable
to estimate individual body mass and to compare
against relative limb lengths.
Statistical analyses
Intralimb proportions. Previous studies indicate that
intralimb proportions, which represent the relationship
between the proximal and distal limb segments of the
573
BODY PROPORTIONS IN ANDEAN PREHISTORY
TABLE 2. Mean body mass estimates in kilograms1
Sample2
Males
Females
MPC
SP
AR
AC
Scheffe’s
test3
58.60 ⫾ 1.53
61.34 ⫾ 0.76
62.27 ⫾ 0.75
66.36 ⫾ 0.83
MPC ⬍ AC; SP ⬍ AC,
AR ⬍ AC
49.96 ⫾ 1.13
52.74 ⫾ 0.59
55.13 ⫾ 0.60
56.83 ⫾ 1.00
MPC ⬍ AR; MPC ⬍ AC;
SP ⬍ AR; SP ⬍ AC
1
Mean and standard error of body mass estimates are based on
average of three regression equations for each individual (Ruff et
al., 1991; McHenry, 1992; Grine et al., 1995).
2
See Table 1 for sample abbreviations.
3
Scheffe’s test results compare mean BM between two samples.
Inequality indicates significant difference (P ⱕ 0.05) in direction
shown. All other comparisons are not significant (P ⬎ 0.05).
Fig. 3. Mean BM with two standard errors in sex-specific
groups by site. Note progressive decrease in estimated body mass
with increasing altitude in both males and especially females. In
addition to this general pattern, Scheffe’s test results indicate
that AC males are significantly larger in estimated BM compared
with males from MPC, SP, and AR. Among females, AR and AC
are significantly larger compared with both MPC and SP.
upper and lower limbs, vary with climate in human populations due to shortening of the distal segments in coldadapted populations (e.g., Trinkaus, 1981; Ruff, 1991,
1994, 2002b; Holliday, 1997, 1999; Holliday and Ruff,
2001). In order to test whether intralimb proportions also
vary according to altitude, RAD is compared with HUM
and TIB with FEM in sex-specific groups according to site
using bivariate scatter plots (Figs. 1, 2).
Limb lengths and overall size. In order to examine
individual variation in limb lengths relative to body size and
to test whether this variation is associated with altitude, it is
necessary to choose an appropriate estimate of body size.
Methods for determining overall size in skeletal remains
include estimating body mass from regression equations
based on femoral head breadth (e.g., Ruff et al., 1991, 1997;
McHenry, 1992; Grine et al., 1995), reconstructing overall
body shape based on regression equations of bi-iliac breadth/
stature (e.g., Ruff, 1994, 2002b; Ruff et al., 1997), and estimating overall size by calculating the geometric mean of all
variables measured on each individual (e.g., Darroch and
Mosimann, 1985; Falsetti et al., 1993; Jungers et al., 1995).
This study applies two of these methods: body mass (BM)
estimated from FHB, and overall size estimated from the
geometric mean (GM). Reconstructing body shape from biiliac breadth/stature is not possible in this study because a
large number of individuals lack pelvic remains. Thus calculating bi-iliac breadth/stature would require the omission
of too many individuals within each sample, rendering comparisons across samples meaningless.
Estimating BM is based on the following four regression
equations:
BM共kg兲 ⫽ 2.268 ⫻ FHB ⫺ 36.5 (Grine et al., 1995)
BM ⫽ 2.239 ⫻ FHB ⫺ 39.9 (McHenry, 1992)
BM ⫽ 2.741 ⫻ FHB ⫺ 54.9 for males (Ruff et al., 1991)
BM ⫽ 2.426 ⫻ FHB ⫺ 35.1 for females (Ruff et al., 1991)
Each of these equations was developed by regressing
FHB against body weight in particular reference samples. Ruff et al. (1997) found that while the equations of
Ruff et al. (1991) and Grine et al. (1995) tend to slightly
overestimate body mass compared with that of
McHenry (1992), the mean difference between these
three equations was only 4%. Using Ruff et al. (1997) as
an example, in which the authors apply these equations
to Pleistocene Homo, BM is estimated by averaging the
results of each equation for each individual grouped by
site (Table 2, Fig. 3). In order to test whether estimated
body mass differs by site, ANOVA and Scheffe’s pairwise comparisons of mean BM are performed on sexspecific groups where group means are considered significantly different at P ⱕ 0.05 (Table 2, Fig. 3). Limb
lengths (HUM, RAD, FEM, and TIB) are then plotted
against BM (Figs. 4, 5) in order to determine whether
limb lengths relative to BM vary in individuals from
different sites and altitude.
Relative limb lengths are also compared against GM, an
estimate of overall size that is independent from the methods described above. Although not equivalent to body
weight, GM estimates overall size based on all of the
variables measured for each individual (e.g., Mosimann
and James, 1979; Darroch and Mosimann, 1985; James
and McCulloch, 1990; Falsetti et al., 1993; Jungers et al.,
1995). GM is calculated based on the following five variables: HUM, RAD, FEM, TIB, and FHB, according to the
following equation (Sokal and Rohlf, 1995):
Gm y ⫽ antilog 1/n
冘
log Y
Similar to previous analyses, each limb length (dependent
variable) is plotted against GM (independent variable) for
each individual in sex-specific groups according to site
(Figs. 6, 7).
Tests for differences in proportions. All bivariate
scatterplots in this study are illustrated with the reduced
major axis (RMA) regression line based on individuals
grouped by sex. RMA regression models are appropriate
for these comparisons because the independent variables
are measured with error (Sokal and Rohlf, 1995). For each
scatterplot that compares intralimb proportions and limb
lengths against BM and GM, tests for elevational differences between samples are undertaken using the nonparametric “quick test” of Tsutakawa and Hewett (1977)
(Table 3). This test reveals whether or not body proportions in each sample significantly differ from other groups
by identifying the number of individuals from each sample
who fall above or below a common RMA regression line
through the pooled sample. Samples are considered to
574
K.J. WEINSTEIN
Fig. 4. Limb lengths vs. estimated BM in males: HUM (A), RAD (B), FEM (C), and TIB (D). See Figure 1 for explanation of
symbols. Note differences between SP, which tends to fall above regression line, compared with AR and especially AC, which tend to
fall below line. MPC males tend to scatter above and below regression line in all variables except for HUM (A).
significantly differ from each other with respect to their
position above or below the common regression line at P ⱕ
0.05.
As an independent method for comparing proportional
differences between groups in both intralimb proportions
and comparisons of limb lengths against BM and GM,
log-transformed indices are generated according to the
methods described in Ruff (2002a). Each index is calculated as log(Y/Xb), where Y is the dependent variable in
the regression model, X is the independent variable, and b
is the predicted slope if the independent and dependent
variables scale isometrically. In the case of intralimb proportions and limb lengths against GM, the predicted isometric slope is 1.0, whereas in the comparisons of limb
lengths against BM, the predicted isometric slope is 0.666.
As in Ruff (2002a), mean log(Y/Xb) values are compared
between samples, using ANOVA and Tukey multiple comparison tests (Tables 4 – 6). Group means are considered
significantly different at P ⱕ 0.05.
Principal component analysis (PCA). I also test further for differences in body proportions by site and altitude through PCA of an unrotated variance covariance
matrix of the five variables (HUM, RAD, FEM, TIB, and
FHB; Table 7). The first PCA axis generally captures
individual variation due to differences in size, while the
second PCA axis tends to explain individual variation
resulting from differences in shape while controlling for
size (e.g., Darroch and Mosimann, 1985; Falsetti et al.,
1993; Jungers et al., 1995). Thus, the second PCA axis
should separate individuals from different altitudes if
body proportions vary according to altitude. Alternatively,
BODY PROPORTIONS IN ANDEAN PREHISTORY
575
Fig. 5. Limb lengths vs. estimated BM in females: HUM (A), RAD (B), FEM (C), and TIB (D). See Figure 1 for explanation of
symbols. Note highland/lowland contrast based on tendency of highland groups (SP and MPC) to fall above regression line, and
lowland groups (AR and especially AC) to fall below.
if altitudinal variation does not explain differences in
overall size or relative limb proportions, then each group
should be indistinguishable along this axis. This variation
is illustrated by plotting individual factor scores (Fig. 8).
All variables were transformed into their natural logarithms for all statistical analyses. All statistical tests were
completed using Systat version 8.0 for Windows (SPSS,
Inc., 1998).
RESULTS
Intralimb proportions
Comparisons of intralimb proportions reveal two general
patterns. First, SP males and females have intralimb proportions that are generally similar to the two lowland samples, which are generally similar to each other. Scatterplots
illustrate that intralimb proportions overlap between SP,
AR, and AC (Figs. 1, 2), and this pattern is also evident in the
Tsutakawa and Hewett nonparametric quick test results
(Table 3). Among males in these three groups, all pairwise
comparisons in individuals above and below the RMA regression lines are not significant, with the exception of larger
SP compared with AR in TIB relative to FEM. Among females of these three groups, all pairwise comparisons are not
significant except for AC females, which are larger in RAD
relative to HUM compared with AR, and larger in TIB relative to FEM compared with SP. Tukey multiple comparison
tests of log-transformed indices (Table 4) also indicate that
SP males and females do not significantly differ from their
lowland counterparts in upper and lower intralimb proportions.
576
K.J. WEINSTEIN
Fig. 6. Limb lengths vs. GM in males: HUM (A), RAD (B), FEM (C), and TIB (D). See Figure 1 for explanation of symbols. Note
that despite considerable overlap between groups, MPC and SP tend to separate slightly from AC.
Second, MPC has markedly shorter RAD and TIB relative to HUM and FEM, respectively, compared with the
other three samples. Although their range of variation
overlaps the other groups, MPC consistently tends to fall
below the regression line in comparisons of the distal
segment against the proximal segment for both the upper
and lower limbs (Figs. 1, 2). Quick test comparisons also
indicate that, with some exceptions, MPC males and females exhibit significantly shorter RAD and TIB relative
to HUM and FEM compared with the other groups (Table
3). Male MPCs, for example, have shorter intralimb proportions compared with all other groups except for TIB
relative to FEM in AR. Female MPCs have significantly
shorter TIB relative to FEM compared with AR and AC,
but do not differ from the lowland groups in RAD relative
to HUM or in any intralimb proportions compared with SP
(Table 3). The Tukey multiple comparison tests of intralimb proportion indices reveal even more pronounced differences between MPC and the other three samples: MPC
males and females are consistently significantly shorter in
distal to proximal limb length proportions for both the
upper and lower limbs compared with all other groups
(Table 4).
Limb lengths relative to body size
Reconstruction of body mass using the equations of
McHenry (1992), Grines et al. (1995), and Ruff et al. (1997)
indicate that the two highland groups are lighter than the
lowlanders in terms of mean BM (Table 2, Fig. 3). Among
males, MPC has the smallest estimated body masses, followed by SP, whose range of BM overlaps considerably
BODY PROPORTIONS IN ANDEAN PREHISTORY
577
Fig. 7. Limb lengths vs. GM in females: HUM (A), RAD (B), FEM (C), and TIB (D). See Figure 1 for explanation of symbols. As
in male comparisons, there is considerable overlap across samples, except for HUM (A), in which MPC and SP are slightly separated
from AC in particular.
with that of AR. Male ACs are the heaviest in BM and do
not overlap the others. One-way ANOVA and Scheffe’s
tests that compare mean BM among males reveal that
despite the progressive decrease in BM with increasing
altitude, the only group to significantly differ from the
others is AC (Table 2, Fig. 3). Among females, MPC exhibits the smallest BM, which overlaps that of SP, while
AR and AC exhibit overlapping ranges which, based on
one-way ANOVA and Scheffe’s tests, are significantly
heavier than those of the two highland groups (Table 2,
Fig. 3).
Limb lengths relative to BM also tend to vary across an
altitudinal gradient, although in a pattern that does not
conform to the predictions of Allen’s rule. The smallerbodied highland samples tend to have relatively long
limbs for their size, whereas the larger-bodied lowland
samples tend to have relatively shorter limbs. This highland/lowland contrast in limb lengths against BM is perhaps most apparent in comparisons of SP with AC, in
which SP males and females fall predominantly above the
regression line for all limb lengths against BM, while AC
males and females are positioned below the regression
line for these same variables (Figs. 4, 5). Tsutakawa and
Hewett quick test results (Table 3) and Tukey multiple
comparisons of mean log-transformed indices of limb
lengths against BM (Table 5) further support the distinction between SP and AC, in which SP males and females
consistently exhibit long limbs for their relatively small
body masses compared with larger-bodied ACs. Tukey
multiple comparisons of mean log-transformed indices indicate that, with the exception of HUM and RAD in males,
SP and AR significantly differ in all limb lengths relative
578
K.J. WEINSTEIN
TABLE 3. Bivariate positional comparisons of samples using “quick test”1
2
Measurement
Males
RAD v HUM
TIB v FEM
HUM v BM
RAD v BM
FEM v BM
TIB v BM
HUM v GM
RAD v GM
FEM v GM
TIB v GM
Females
RAD v HUM
TIB v FEM
HUM v BM
RAD v BM
FEM v BM
TIB v BM
HUM v GM
RAD v GM
FEM v GM
TIB v GM
MPC/SP
MPC/AR
MPC/AC
MPC ⬍ SP
MPC ⬍ SP
ns
ns
ns
ns
ns
ns
ns
ns
MPC ⬍ AR
ns
ns
ns
ns
ns
ns
ns
MPC ⬎ AR
ns
MPC ⬍ AC
MPC ⬍ AC
ns
ns
ns
ns
ns
ns
MPC ⬎ AC
ns
ns
SP
ns
ns
SP
SP
ns
ns
SP
SP
ns
ns
ns
ns
ns
ns
MPC ⬎ SP
ns
ns
ns
ns
MPC ⬍ AR
ns
ns
ns
ns
ns
ns
MPC ⬎ AR
ns
ns
MPC
MPC
ns
ns
MPC
MPC
MPC
ns
MPC
ns
ns
SP
ns
SP
SP
ns
ns
SP
ns
⬍ AC
⬎ AC
⬎ AC
⬎ AC
⬎ AC
⬎ AC
SP/AR
⬎ AR
⬎ AR
⬎ AR
⬎ AR
⬎ AR
⬎ AR
⬎ AR
⬎ AR
⬎ AR
SP/AC
ns
ns
SP
SP
SP
SP
SP
SP
SP
ns
⬎
⬎
⬎
⬎
⬎
⬎
⬎
AC
AC
AC
AC
AC
AC
AC
ns
SP
SP
SP
SP
SP
SP
SP
SP
ns
⬍
⬎
⬎
⬎
⬎
⬎
⬎
⬎
AC
AC
AC
AC
AC
AC
AC
AC
AR/AC
ns
ns
AR
AR
ns
ns
AR
AR
ns
ns
⬎ AC
⬎ AC
⬎ AC
⬎ AC
AR ⬍ AC
ns
AR ⬎ AC
ns
ns
ns
AR ⬎ AC
ns
ns
ns
1
ns, equality of position about RMA line in two samples. Inequality indicates significant difference (P ⬍ 0.05) about RMA line in
direction shown.
2
See text for measurement definitions and abbreviations. See Table 1 for sample abbreviations.
TABLE 4. Log-transformed indices and Tukey multiple comparisons of intralimb proportions
Ratio
Sample
n
Mean
SD
Tukey test results
Males log (RAD/HUM)
MPC
SP
AR
AC
MPC
SP
AR
AC
MPC
SP
AR
AC
MPC
SP
AR
AC
13
41
38
52
18
38
48
51
10
56
56
28
12
54
66
28
⫺0.282
⫺0.244
⫺0.244
⫺0.251
⫺0.181
⫺0.154
⫺0.153
⫺0.155
⫺0.293
⫺0.263
⫺0.262
⫺0.256
⫺0.182
⫺0.161
⫺0.155
⫺0.145
0.045
0.035
0.039
0.028
0.026
0.019
0.029
0.026
0.046
0.030
0.029
0.027
0.025
0.023
0.023
0.023
MPC ⬍ SP, P ⫽ 0.003
MPC ⬍ AR, P ⫽ 0.001
MPC ⬍ AC, P ⫽ 0.02
Males log (TIB/FEM)
Females log (RAD/HUM)
Females log (TIB/FEM)
to BM (Table 5), although the Tsutakawa and Hewett
quick test results reveal that these differences are significant only in the comparisons of lower limbs in males and
in all limbs except RAD in females (Table 3). In general,
MPC males and especially females have small body
masses, and in terms of limb lengths relative to BM, MPC
females particularly tend to cluster near the smallest SP
individuals (Figs. 4, 5). Tukey multiple comparison tests,
however, indicate that indices of limb lengths with BM in
MPC significantly differ with AC only (Table 5), although
quick test results show that these differences are only
significant between MPC and AC females for comparisons
of HUM and TIB against BM (Table 3).
All four limb lengths scale isometrically with GM in
both males and females. Confidence intervals of the regression slopes for the scatterplots of HUM, RAD, FEM,
and TIB include 1.0, and all groups, including MPC, exhibit considerable overlap (Figs. 6, 7). A progressive decrease from highland to lowland in limb lengths relative to
overall size, however, is apparent when comparing mean
log-transformed indices of limbs and GM (Table 6). Male
and female MPCs have the largest mean ratios of
MPC ⬍ SP, P ⫽ 0.001
MPC ⬍ AR, P ⬍ 0.001
MPC ⬍ AC, P ⫽ 0.001
MPC ⬍ SP, P ⫽ 0.02
MPC ⬍ AR, P ⫽ 0.02
MPC ⬍ AC, P ⫽ 0.004
MPC ⬍ SP, P ⫽ 0.02
MPC ⬍ AR, P ⫽ 0.002
MPC ⬍ AC, P ⬍ 0.001
log(HUM/GM) and log(FEM/GM), followed by SP, AR, and
then AC, and Tukey multiple comparison tests show that
many of these mean values, especially between SP and
AC, significantly differ. This highland/lowland progressive decrease in limb length relative to GM is less apparent in comparisons of RAD and TIB: mean values are more
randomly distributed between groups, and most of the
group mean differences, as revealed through Tukey multiple comparison tests, separate SP from AR and AC, and
AC from MPC. Quick test results that examine pairwise
comparisons of individuals across RMA regression lines of
limb lengths against GM also indicate varied results (Table 3). Among males, MPC significantly differs from AR
and AC in FEM; SP significantly differs from AR in FEM
and TIB, and from AC in HUM, RAD, and FEM; and AR
and AC significantly differ from each other in HUM and
RAD. Among females, MPC significantly differs from SP
in HUM, from AR in FEM, and from AC in all variables
except for FEM (Table 3). Female SPs also significantly
differ in their RMA line position from ARs in terms of
FEM and from ACs in all variables except TIB. Finally,
AR and AC females differ in their positions of HUM. Thus,
579
BODY PROPORTIONS IN ANDEAN PREHISTORY
TABLE 5. Log-transformed indices and Tukey multiple comparisons of limb lengths against BM
Ratio
Sample
n
Mean
SD
Males log (HUM/BM)
MPC
SP
AR
AC
MPC
SP
AR
AC
MPC
SP
AR
AC
MPC
SP
AR
AC
MPC
SP
AR
AC
MPC
SP
AR
AC
MPC
SP
AR
AC
MPC
SP
AR
AC
10
40
42
51
10
41
39
51
16
39
49
51
15
39
47
51
9
53
61
28
8
55
58
28
12
54
68
28
10
52
66
28
2.970
2.968
2.953
2.909
2.715
2.725
2.710
2.659
3.290
3.295
3.258
3.235
3.103
3.138
3.105
3.080
2.992
2.990
2.954
2.904
2.717
2.726
2.692
2.649
3.318
3.321
3.281
3.240
3.151
3.160
3.124
3.094
0.021
0.051
0.054
0.060
0.077
0.053
0.060
0.062
0.055
0.050
0.050
0.051
0.054
0.054
0.059
0.058
0.051
0.048
0.054
0.058
0.043
0.050
0.056
0.062
0.047
0.049
0.049
0.064
0.047
0.053
0.052
0.060
Males log (RAD/BM)
Males log (FEM/BM)
Males log (TIB/BM)
Females log (HUM/BM)
Females log (RAD/BM)
Females log (FEM/BM)
Females log (TIB/BM)
Tukey test results
AC ⬍ MPC, P ⫽ 0.006
AC ⬍ SP, P ⬍ 0.001
AC ⬍ AR, P ⫽ 0.001
AC ⬍ MPC, P ⫽ 0.035
AC ⬍ SP, P ⬍ 0.001
AC ⬍ AR, P ⬍ 0.001
AC ⬍ MPC, P ⫽ 0.001
AR ⬍ SP, P ⫽ 0.005
AC ⬍ SP, P ⬍ 0.001
AR ⬍ SP, P ⫽ 0.038
AC ⬍ SP, P ⬍ 0.001
AC
AR
AC
AC
AC
AR
AC
AC
AC
AR
AC
AC
AC
AR
AC
AC
⬍
⬍
⬍
⬍
⬍
⬍
⬍
⬍
⬍
⬍
⬍
⬍
⬍
⬍
⬍
⬍
MPC, P ⫽ 0.001
SP, P ⫽ 0.001
SP, P ⬍ 0.001
AR, P ⬍ 0.001
MPC, P ⫽ 0.01
SP, P ⫽ 0.005
SP, P ⬍ 0.001
AR, P ⬍ 0.003
MPC, P ⬍ 0.001
SP, P ⬍ 0.001
SP, P ⬍ 0.001
AR, P ⫽ 0.002
MPC, P ⫽ 0.022
SP, P ⫽ 0.002
SP, P ⬍ 0.001
AR, P ⬍ 0.074
TABLE 6. Log-transformed indices and Tukey multiple comparisons of limb lengths against GM
Ratio
Sample
n
Mean
SD
Males log (HUM/GM)
MPC
SP
AR
AC
MPC
SP
AR
AC
MPC
SP
AR
AC
MPC
SP
AR
AC
MPC
SP
AR
AC
MPC
SP
AR
AC
MPC
SP
AR
AC
MPC
SP
AR
AC
9
38
35
51
9
38
35
51
9
38
35
51
9
38
35
51
6
50
54
28
6
50
54
28
6
50
54
28
6
50
54
28
0.352
0.338
0.338
0.328
0.079
0.093
0.101
0.077
0.665
0.663
0.647
0.653
0.491
0.509
0.493
0.498
0.347
0.346
0.340
0.324
0.070
0.083
0.079
0.068
0.684
0.678
0.667
0.659
0.512
0.517
0.511
0.514
0.019
0.019
0.024
0.021
0.033
0.021
0.025
0.024
0.010
0.015
0.020
0.013
0.025
0.018
0.023
0.023
0.016
0.019
0.021
0.016
0.023
0.020
0.023
0.021
0.013
0.016
0.018
0.020
0.016
0.021
0.018
0.016
Males log (RAD/GM)
Males log (FEM/GM)
Males log (TIB/GM)
Females log (HUM/GM)
Females log (RAD/GM)
Females log (FEM/GM)
Females log (TIB/GM)
Tukey test results
AC ⬍ MPC, P ⫽ 0.008
MPC ⬍ AR, P ⫽ 0.075
AC ⬍ SP, P ⫽ 0.015
AC ⬍ AR, P ⬍ 0.001
AR ⬍ MPC, P ⫽ 0.01
AR ⬍ SP, P ⬍ 0.001
AC ⬍ SP, P ⫽ 0.025
AR ⬍ SP, P ⫽ 0.013
AC ⬍ SP, P ⫽ 0.11
AC ⬍ MPC, P ⫽ 0.03
AC ⬍ SP, P ⬍ 0.001
AC ⬍ AR, P ⫽ 0.001
AC ⬍ SP, P ⫽ 0.016
AC ⬍ MPC, P ⫽ 0.01
AC ⬍ SP, P ⬍ 0.001
AR ⬍ SP, P ⫽ 0.007
No significant differences
580
K.J. WEINSTEIN
TABLE 7. PCA in males and females
Eigenvector coefficients
Males
RAD
FHB
FEM
TIB
HUM
Eigenvalue
% of total variance
Females
Axis I
Axis II
Axis I
Axis II
0.505
0.350
0.446
0.485
0.443
0.009
72.27
0.259
⫺0.931
0.080
0.199
0.145
0.002
16.59
0.505
0.287
0.430
0.500
0.477
0.008
71.26
0.131
⫺0.955
0.181
0.101
0.167
0.002
19.19
while the scatterplots illustrate considerable overlap between sites in limb lengths relative to GM, statistical tests
reveal a subtle altitudinal cline from smaller-bodied highland groups to larger-bodied lowlanders similar to that
seen in comparisons of limb lengths relative to BM. From
smallest to largest, MPC and SP tend to have relatively
long limbs, and the larger AR and AC tend to have relatively shorter limbs.
PCA
The PCA reveals the same contrasts between MPC and
the other groups in terms of relative limb lengths, and
between the two highland groups and the lowlanders in
terms of differences in overall size. The first PC axis
explains approximately 71–72% of the total sample variance for males and females (Table 7). This axis represents
differences in overall size: all eigenvector coefficients load
strongly and positively along this axis. Individual factor
scores, moreover, are highly correlated with GM (r2 ⫽
0.999, P ⬍ 0.001). Similar to the bivariate regression
analyses, MPC tends to plot negatively along this axis: all
but three males have slightly negative factor scores, and
all but one female have markedly negative scores (Fig. 8).
Male and female SP are virtually indistinguishable from
the lowland groups along the first axis. While the separation between MPC and the other three groups is not especially marked, this axis shows that MPC falls at the
small end of the size range for these variables.
The second PCA axis accounts for approximately 16 –
19% of the total sample variance (Table 7). This axis
contrasts FHB from limb lengths. FHB loads strongly and
negatively along this axis, while all four limbs, and especially TIB and RAD, score positively. This second axis
separates highland groups from the lowland samples. In
particular, SP males and females tend to plot positively
along this axis and are separate from those of AC, who
score negatively, especially AC females. Male and female
ARs appear to be intermediate between these two extremes (Fig. 8). All but two males and one female from
MPC exhibit positive factor scores along this axis, and
tend to cluster within the SP range of variation. Thus, the
second axis illustrates a subtle altitudinal cline in femoral
head size and limb lengths with the lighter individuals
from highland regions and the heavier individuals from
the lowlands.
DISCUSSION
In these samples, body size and proportions vary across
an altitudinal gradient in two patterns. First, individuals
from the highest altitudes (MPC) have the shortest relative intralimb length proportions, whereas those from
slightly lower elevations at 2,500 m (SP) have intralimb
length proportions that are indistinguishable from the
lowland groups, a pattern that is perhaps best explained
as a response to climatic stress. Second, overall size and
limb lengths relative to overall size tend to vary across an
elevational cline, with the largest individuals with relatively shorter limbs from the central (AC) and southcentral (AR) coasts, and smaller individuals with relatively longer limbs for their size from 2,500 m (SP) and
2,000 –3,800 m (MPC). Unlike intralimb length proportions, the highland/lowland contrasts in overall size and
limb lengths relative to overall size do not follow a pattern
indicative of climatic stress. In this sample, the largest
individuals with relatively short limbs for their size are
from coastal regions with mild climates rather than from
higher elevations with harsher ambient temperatures as
predicted by Bergmann’s and Allen’s rules. While a variety of biological and ecological factors may be at work,
nutritional and dietary variation may best explain this
subtle contrast in body size and limb lengths relative to
overall size between highland and lowland groups. Considering that gene flow may be a confounding factor, archaeological evidence for widespread movements among
the populations from which these samples were derived is
also explored.
Intralimb proportions, climatic adaptations, and
biological diversity
Morphological adaptations to climate perhaps best explain the short intralimb proportions in MPC. Given that
MPC individuals were recovered from approximately
2,000 –3,800 m in elevation where ambient temperatures
frequently fall below 0°C, and that living Andeans from
high altitudes are presumed to be cold-adapted (Stinson
and Frisancho, 1978), the short relative intralimb proportions in this highland skeletal sample most likely reflect a
cold adaptation, as predicted by Allen’s rule. Furthermore,
comparisons of pre-Contact human skeletons from the
highland Andes (data from MacCurdy, 1923) with those
from North American populations from Arctic climates
that are known to be cold-adapted (data from Trinkaus,
1981) illustrate that highland Andeans have intralimb
proportions that are equally as short as Eskimo/Inuit
groups, demonstrating further the development of a coldadapted body shape among prehistoric highland Andean
populations (Weinstein, 1998, 2001). In contrast, relative
intralimb proportions in SP are virtually indistinguishable from individuals from lowland environments. This
contrast between MPC and SP suggests that climatic conditions in the southern Peruvian highlands are severe
enough to favor the selection of a cold-adapted body shape,
whereas temperatures in the Atacama Desert of northern
Chile are less severe.
Climatic adaptations in Andean prehistory, as inferred
from intralimb proportions, are useful to debates about
the evolution of modern human biological diversity and
the worldwide dispersal of modern humans during the
Pleistocene. As discussed earlier, numerous paleoanthropological studies compare intralimb proportions of recent
Homo sapiens from different climates in order to discern
climatic adaptations, migrations, and phylogenetic relationships in human groups during the Pleistocene (e.g.,
Trinkaus, 1981; Ruff, 1991, 1994, 2002b; Ruff and Walker,
1993; Holliday 1997, 1999, 2000; Pearson, 2000). With the
exception of skeletal samples from Arctic regions in North
America, however, recent human skeletons from prehistoric North and South America are rarely included in
comparative studies of climatic adaptation in human evo-
BODY PROPORTIONS IN ANDEAN PREHISTORY
581
proportions within MPC and significantly greater proportions in groups from lower elevations, while admittedly
less pronounced than the contrasts between Old World
populations that are separated by vast geographic and
temporal distances, indeed suggest that climatic adaptations are one of many selective factors that shape the
biology of prehistoric Andeans in particular: results that
are similar to Newman (1953, 1960) and Beals et al. (1984)
regarding climatic adaptations in the Americas in general.
Archaeological evidence suggests that humans have inhabited South America since at least 12,500 years BP and
potentially earlier (e.g., Dillehay, 2000; Roosevelt et al.,
2002). Moreover, by 11,000 years BP, regional populations
developed diverse cultural adaptations to their unique
environments, including the Andean highlands and coast,
temperate forests, and tropical lowlands (Dillehay, 1997;
Moseley, 2001; Roosevelt et al., 2002). Dillehay (2000)
emphatically argue that the peopling of the Americas
should be interpreted as part of the worldwide dispersal of
Homo sapiens that began ca. 40,000 years ago, that there
is great time depth of Amerindian groups in widely varied
environments, and that prehistoric populations within the
Americas developed a large degree of cultural diversity.
The results of this study suggest that prehistoric Amerindian populations should exhibit a similar degree of biological diversity as well. Given the diverse environments that
the earliest Americans colonized, including high altitudes,
arid deserts, and tropical and temperate regions, it seems
plausible that prehistoric populations of the Americas
would develop biological adaptations to suit these environmental extremes. Future research should continue to examine body proportions across a number of populations
from the Americas to more readily capture the degree of
biological diversity and clinal variation in body size and
proportions that most surely existed within the pre-Contact period.
Body size, relative limb lengths, and
dietary variation
Fig. 8. Individual factor scores for first and second axes of
five-variable PCA in males (A) and females (B). See Figure 1 for
explanation of symbols. Axis I represents size differences. Note
that with exception of three males and one female, MPC plots
negatively along this axis, especially in females. Axis II contrasts
FHB with limb lengths. Although there is some overlap in range
of variation, SP tends to plot positively along this axis, with MPC
falling within same range. AC tends to plot negatively, and AR
appears intermediate in relation to others.
lution. Moreover, Amerindian groups which are included
as comparative samples in studies of intralimb proportions in both living populations (e.g., Eveleth and Tanner,
1976, 1990) and in skeletal samples (e.g., Trinkaus, 1981;
Ruff, 1991, 1994; Holliday, 1997, 1999, 2000; Pearson,
2000) are generally categorized as “Northern Asian” or
“Asiatic.” While never explicitly stated, “Northern Asian”
and other similar labels imply that North and South
Americans are biologically unchanged from the earliest
inhabitants of the New World who are presumed to be
cold-adapted. Yet reduced distal/proximal limb-length
The highland groups (MPC and SP) tend to be smaller
in overall size with relatively longer limbs, while the lowland groups (AR and AC) tend to be larger with relatively
shorter limbs: a pattern that runs counter to Bergmann’s
and Allen’s rules. While a number of factors could be at
work, studies of living populations at high altitudes suggest that dietary variation and its differential effects on
growth may offer the most robust explanation for the
highland/lowland contrasts in body size and limb lengths
relative to body size in these samples.
Living highland Andeans exhibit delayed growth and
short adult stature compared with lowland Andean populations and with European migrants to high altitudes
(e.g., Frisancho, 1978; Greksa et al., 1984, 1985; Greksa,
1986). High-altitude hypoxia is one potential environmental factor that can adversely affect growth during the
prenatal period, infancy, and childhood, leading to small
body sizes and short stature in adulthood (Frisancho and
Baker, 1970; Beall et al., 1977; Mueller et al., 1978, 1980;
Stinson, 1980, 1982; Greksa, 1986; Bogin, 1999) Among
living highland populations in the Andes, however, limited nutrition and low energy availability (two biocultural
stressors that are also characteristic of high-altitude environments) may more profoundly influence the small
body sizes in the highland groups. In highland southern
Peru and in Ecuador, individuals with low socioeconomic
status experience annual and seasonal shortages of ade-
582
K.J. WEINSTEIN
quate food resources, leading to delayed growth during
childhood and adolescence, which are interpreted as the
primary cause of short adult stature in these populations
(e.g., Leonard, 1989; Leonard et al., 1990, 1995, 2000).
Nutritional stress, exposure to infectious diseases, and
their adverse affects on growth also explain highland/
lowland contrasts in body sizes in living populations from
Ethiopia (e.g., Harrison et al., 1969; Clegg et al., 1972) and
the Himalayas (e.g., Bharadwaj et al., 1973; Gupta and
Basu, 1981). Furthermore, fluctuations in dietary quality,
disease load, and health status led to secular changes in
adult stature and body mass in Central and South American populations over the last 8,000 years (Bogin and
Keep, 1999), in the United States over the last 200 years
(Meadows Jantz and Jantz, 1999), and in populations
from developing nations over the last 40 years (Katzmarzyk and Leonard, 1998). Limb lengths relative to overall
size may also be affected by diet and nutritional status.
Among living populations suffering from mild to moderate
chronic undernutrition, limb lengths tend to be short relative to stature (Bogin, 1999). When undernourished
groups experience improved diets, however, lower limb
lengths tend to increase relative to stature, effectively
increasing lower limb proportions (Bogin, 1999; Stinson,
2000). Thus, living human populations experiencing both
chronic mild undernutrition and improved diets demonstrate the plasticity of body size and of limb lengths relative to body size in response to nutritional conditions.
The altitudinal cline in overall body size in human skeletons from the pre-Contact Andes in this study follows a
pattern similar to that of living human populations with
different diets and energy expenditure. Archaeological evidence infers that the highland and lowland groups differed in their diets and subsistence base. The larger-bodied lowlanders relied heavily on marine resources as a
dietary staple, with an increase in terrestrial resources
through time (e.g., Arriaza, 1995; Moseley, 2001). Individuals from AC exhibited rather high frequencies of cribra
orbitalia and porotic hyperostosis (personal observation),
osteological conditions that are indicative of iron-deficiency anemia (Stuart-Macadam, 1992) and that are common in pre-Contact coastal groups exposed to parasites
from marine-based diets (Walker, 1986). The other coastal
group from AR, however, does not exhibit these pathological conditions in high frequency (personal observation).
Highland MPC and SP, in contrast, consumed maize, tubers, and camelid meat (e.g., Costa, 1988; Núñez, 1991;
Costa Junquiera and Llagostera, 1994; Miller, 2003;
Burger et al., 2003). Both highland groups exhibit relatively low frequencies of pathological conditions associated with nutritional deficits (Costa, 1988; Neves and
Costa, 1998; Neves et al., 1999; Verano, 2003). While
dietary reconstruction based on faunal remains, stable
isotopic analysis of human bone, and paleopathological
conditions of human skeletal remains do not identify any
obvious nutritional deficits in the highland groups, the
agropastoral highland diet most likely encompassed a
smaller degree of food diversity compared with the diets of
coastal groups. Low food diversity can lead to insufficient
intake of specific nutrients, and as is the case in living
Andean highland populations, delayed growth and short
adult stature (Leonard et al., 1990; Bogin, 1999). Thus
small body sizes in the highlanders in this study seem
likely to reflect a less varied diet compared to that of the
lowlanders. Although ecogeographic principles predict
that relative limb lengths should be shorter in coldadapted populations, variation in limb lengths relative to
overall size in these samples (the relatively longer limbs in
the smaller highlanders, and the relatively shorter limbs
in the larger lowlanders) may also reflect morphological
plasticity in response to dietary conditions, as documented
in living populations. Future studies may reveal whether
nutritional deficits, high-altitude hypoxia, ambient temperature, or a combination of numerous environmental
stressors had an impact on relative limb proportions and
body size in pre-Contact Andean populations.
Coastal-highland gene flow and
morphological variation
Archaeologists and biological anthropologists have long
debated the nature of population interactions between
coastal and highland sites in Andean prehistory (e.g.,
Murra, 1972; Moseley, 2001; Burger and Salazar, 2003).
Therefore, any examination of morphological variation in
pre-Contact Andean populations requires a discussion of
the possible effects of gene flow between highland and
coastal groups. While the exact cultural and historical
processes involved in highland-coastal interactions in Andean prehistory are beyond the scope of this paper, the
results of this study suggest that, with the exception of
some MPC individuals, gene flow had minimal effect on
the biological diversity observed in these skeletal samples.
Analysis of biological distance based on epigenetic (nonmetric) cranial and dental traits indicates genetic continuity in Arica populations from the Archaic period (3000 –
1000 BC) through the Late Intermediate period (AD
1000 –1476) (Sutter, 2000; Sutter and Mertz, 2004). Identical body proportions in human skeletons from the Archaic and Late Intermediate periods further suggest that
Arica populations experienced little biological change
through time (Weinstein, 2001). Similarly, archaeological
evidence not only indicates human settlement along the
central Andean coast starting at 9,000 years BP, but also
a geopolitical separation between the central coast and the
south-central highlands that continued through the Late
Intermediate period (Moseley, 2001). Thus rather than an
influx of highland biological and cultural influences
through time, coastal populations appear to be biologically
distinct and relatively isolated from their highland neighbors, at least in the coastal groups represented in this
study. Likewise, archaeological evidence from San Pedro
de Atacama suggests that this highland population had
strong biological and cultural ties to the highlands to the
east rather than to the coastal settlements to the west.
Núñez (1991), for example, argued that San Pedro de
Atacama was at the center of prehistoric trade routes
linking highland regions to the Northeast and Southwest.
A paucity of coastal artifacts in San Pedro de Atacama
archaeological sites suggests further that biological and
cultural contacts with coastal populations were minimal
(Moseley, 2001).
The issue of coastal-highland population movements is
more problematic in MPC. Archaeologists have long speculated about the ethnic diversity of the Andes during the
Inca period in general, and of the populations at Machu
Picchu and Cuzco during this time period in particular
(e.g., Bingham, 1930; Moseley, 2001; Burger and Salazar,
2003). Burger and Salazar (2003) argued that the individuals recovered from Machu Picchu were nonelite servants
selected from throughout the Andes to serve at this Inca
royal estate. Dietary variation in the Machu Picchu skeletal sample, as inferred from stable carbon and nitrogen
isotopic signatures in human bone, support the hypothesis
BODY PROPORTIONS IN ANDEAN PREHISTORY
that these remains represent diverse coastal and highland
populations (Burger et al., 2003). Verano (2003) argued
that the Machu Picchu skeletal sample is equally divided
into individuals from coastal and highland regions, based
on his analysis of cranial deformation styles and craniometrics. Given that artificial cranial deformation modifies
craniofacial shape (e.g., Antón, 1989) and is a rather ubiquitous practice in Andean prehistory, craniometrics is not
a reliable method to assess biological distance and geographic origin in human skeletal remains. While comparisons of body size and shape do not directly address questions about genetic affinities, MPC does exhibit the
greatest range of variation in body size and proportions of
all samples represented in this study, results which suggest the possibility that a few MPC individuals may be
coastal migrants. Despite this possibility, the majority of
MPC exhibited the smallest intralimb proportions, a pattern in accordance with ecogeographic rules. Climatic conditions at high altitudes seem to influence, in part, the
short intralimb proportions of highland individuals. This
cold-adapted body shape that is characteristic of the majority of MPC, moreover, is contrasted with those from
lower elevations, further illustrating the presence of ecogeographic variation in Andean prehistory.
CONCLUSIONS
Pre-Contact human skeletal remains exhibit body proportions that vary along an altitudinal cline due to both
climatic conditions and dietary factors. Despite the possibility of gene flow from coastal regions, highland individuals from MPC exhibit markedly shorter intralimb length
proportions compared with individuals from lower elevations, features that are indicative of a morphological adaptation to cold climatic conditions. Highland individuals
from both MPC and SP tend to be smaller in estimated
body mass and have relatively long limbs for their small
body sizes compared with their lowland counterparts, an
altitudinal contrast that may reflect differences in diet
and subsistence between highland and lowland populations. Thus both climatic conditions and dietary variation
appear to influence variation in body size and proportions
in Andean prehistory. The results of this study reveal
subtle variation in postcranial morphology across relatively short geographic distances and temporal periods.
ACKNOWLEDGMENTS
I thank the following individuals for their guidance
during the course of this research: Susan Antón, Sue Boinski, Michael Moseley, Dave Steadman, and William Leonard. I am grateful to the following individuals and institutions for providing access to skeletal collections: M.A.
Costa and all of the staff from the Museo Arqueológico
R.P. Gustavo Le Paige, San Pedro de Atacama, Chile;
Juan Chacama, Vivien Standen, and the staff from the
Museo San Miguel de Azapa, Universidad de Tarapacá,
Arica, Chile; P. Ryan Williams and Will Pestle from the
Department of Anthropology, Field Museum, Chicago, IL;
and Richard Burger and Roger Colten from the Division of
Anthropology, Peabody Museum of Natural History, Yale
University, New Haven, CT. I also thank Clark Spencer
Larsen and two anonymous reviewers for their helpful
comments.
LITERATURE CITED
Abate FR. 1991. Omni gazetteer of the United States of America,
volume 9: Pacific. Detroit: Omnigraphics, Inc.
583
Aldrich JW, James FC. 1991. Ecogeographical variation in the
American robin (Turdus migratorius). Auk 108:230 –249.
Allen JA. 1877. The influence of physical conditions in the genesis
of species. Rad Rev 1:108 –140.
Allison M, Focacci G, Arriaza B, Standen V, Rivera M, Lowenstein J. 1984. Chinchorro momias de preparación complicada:
métodos de momificación. Chungara 13:155–173.
Antón SC. 1989. Intentional cranial vault deformation and induced changes of the cranial base and face. Am J Phys Anthropol 79:253–267.
Arriaza BT. 1995. Beyond death: the Chinchorro mummies of
ancient Chile. Washington, DC: Smithsonian Institution Press.
Arriaza BT, Salo W, Aufderheide AC, Holcomb TA. 1995. PreColumbian tuberculosis in northern Chile: molecular and skeletal evidence. Am J Phys Anthropol 98:37– 45.
Ashton KG, Tracy MC, de Queiroz A. 2000. Is Bergmann’s rule
valid for mammals? Am Nat 156:390 – 415.
Baker PT. 1960. Climate, culture, and evolution. Hum Biol 32:
3–16.
Barry RG. 1979. High altitude climates. In: Webber PJ, editor.
High altitude geoecology. Boulder, CO: Westview. p 55–74.
Beall CM, Baker PT, Baker TS, Haas JD. 1977. The effects of high
altitude on adolescent growth in southern Peruvian Amerindians. Hum Biol 49:109 –124.
Beals KL, Smith CL, Dodd SM. 1984. Brain size, cranial morphology, climate, and time machines. Curr Anthropol 25:301–330.
Berenguer J, Dauelsberg P. 1989. El norte grande en la orbita de
Tiwanaku. In: Iván S, editor. Culturas de Chile prehistoria:
desde sus orı́genes hasta los albores de la conquista. Santiago:
Andrés Bello. p 129 –180.
Bergmann C. 1847. Ueber die Verhaltnisse der Warmeokonomie
der Thiere zu ihrer Grosse. Gottinger Stud 3:595–708.
Bharadwaj H, Singh AP, Malhotra MS. 1973. Body composition of
the high-altitude natives of Ladakh: a comparison with sealevel residents. Hum Biol 45:423– 434.
Billings WD. 1979. High mountain ecosystems: evolution, structure, operation and maintenance. In: Webber PJ, editor. High
altitude geoecology. Boulder, CO: Westview. p 97–125.
Bingham H. 1930. Machu Picchu: a citadel of the Incas. New
Haven: Yale University Press.
Bird J. 1943. Excavations in northern Chile. Anthropol Pap Am
Mus Nat Hist 38:173–318.
Bogin B. 1999. Patterns of human growth, 2nd ed. Cambridge:
Cambridge University Press.
Bogin B, Keep R. 1999. Eight thousand years of economic and
political history in Latin America revealed by anthropometry.
Ann Hum Biol 26:333–351.
Burger RL, Salazar LC. 2003. Preface. In: Burger RL, Salazar LC,
editors. The 1912 Yale Peruvian scientific expedition collections from Machu Picchu: human and animal remains. Yale
University publications in anthropology 85. New Haven: Department of Anthropology, Yale University, and Division of
Anthropology, Peabody Museum of Natural History. p xiii–xv.
Burger RL, Lee-Thorp JA, van der Merwe NJ. 2003. Rite and crop
in the Inca state revisited: an isotopic perspective from Machu
Picchu and beyond. In: Burger RL, Salazar LC, editors. The
1912 Yale Peruvian scientific expedition collections from Machu Picchu: human and animal remains. Yale University publications in anthropology 85. New Haven: Department of Anthropology, Yale University, and Division of Anthropology,
Peabody Museum of Natural History. p 119 –137.
Calder WA. 1984. Size, function, and life history. Cambridge, MA:
Harvard University Press.
Clegg EG, Ashton EH, Flinn RM. 1972. The growth of children at
different altitudes in Ethiopia. Philos Trans R Soc Lond [Biol]
272: 403– 437.
Costa MA. 1988. Reconstitución fı́sica y cultural de la población
tardı́a del cementerio de Quitor-6 (San Pedro de Atacama).
Estud Atacamenos 9:99 –126.
Costa Junquiera MA, Llagostera A. 1994. Coyo-3: momentos finales del perı́odo medio en San Pedro de Atacama. Estud Atacamenos 11:73–107.
Crognier E. 1981. Climate and anthropometric variations in Europe and the Mediterranean area. Ann Hum Biol 8:99 –107.
584
K.J. WEINSTEIN
Darroch JN, Mosimann JE. 1985. Canonical and principal components of shape. Biometrika 72:241–252.
Dillehay TD. 1997. Monte Verde: a late Pleistocene settlement in
Chile. Volume 2. The archaeological context and interpretation.
Washington, DC: Smithsonian Institution Press.
Dillehay TD. 2000. The settlement of the Americas: a new prehistory. New York: Basic Books.
Dorsey GA. No date. Unpublished field notes on file, Department
of Anthropology, Field Museum of Natural History, Chicago,
Illinois.
Earth Info, Inc. 1996. Global climate categories CD-ROM. Boulder, CO: Earth Info, Inc.
Eaton GF. 1916. The collection of osteological materials from
Machu Picchu. Mem Conn Acad Arts Sci 5:1–96.
Eveleth PB, Tanner JM. 1976. Worldwide variation in human
growth. Cambridge: Cambridge University Press.
Eveleth PB, Tanner JM. 1990. Worldwide variation in human
growth, 2nd ed. Cambridge: Cambridge University Press.
Falsetti AB, Jungers WL, Cole TM III. 1993. Morphometrics of
the callitrichid forelimb: a case study in size and shape. Int J
Primatol 14:551–572.
Focacci G, Chacón S. 1989. Excavaciones arqueológicas en los
faldeos del Morro de Arica, sitios Morro 1/6 y 2/2. Chungara
22:15– 62.
Frisancho AR. 1978. Human growth and development among
high-altitude populations. In: Baker PR, editor. The biology of
high-altitude peoples. Cambridge: Cambridge University
Press. p 117–171.
Frisancho AR, Baker PT. 1970. Altitude and growth: a study of
the patterns of physical growth of a high altitude Peruvian
Quechua population. Am J Phys Anthropol 32:279 –292.
Graves GR. 1991. Bergmann’s rule near the equator: latitudinal
climates in body size of an Andean passerine bird. Proc Natl
Acad Sci USA 88:2322–2325.
Greksa LP. 1986. Growth patterns of European and Amerindian
high altitude natives. Curr Anthropol 27:72–74.
Greksa LP, Spielvogel H, Paredes-Fernandez L, Paz-Zamora M,
Caceres E. 1984. The physical growth of urban children at high
altitude. Am J Phys Anthropol 65:315–322.
Greksa LP, Spielvogel H, Caceres E. 1985. Effect of altitude on
the physical growth of upper-class children of European ancestry. Ann Hum Biol 12:225–232.
Grine FE, Jungers WL, Tobias PV, Pearson OM. 1995. Fossil
Homo femur from Berg Aukas, Northern Namibia. Am J Phys
Anthropol 97:151–185.
Gupta R, Basu A. 1981. Variations in body dimensions in relation
to altitude among the Sherpas of the eastern Himalayas. Ann
Hum Biol 8:145–151.
Harrison GA, Küchemann CF, Moore MAS, Boyce AJ, Baju T,
Mourant AE, Godber MJ, Glasgow BG, Kopec AC, Tills D,
Clegg EJ. 1969. The effects of altitudinal variation in Ethiopian
populations. Philos Trans R Soc Lond [Biol] 256:147–182.
Hiernaux J, Froment A. 1976. The correlations between anthrobiological and climatic variables in sub-Saharan Africa: revised
estimates. Hum Biol 48:757–767.
Holliday TW. 1997. Postcranial evidence of cold adaptation in
European Neandertals. Am J Phys Anthropol 104:245–258.
Holliday TW. 1999. Brachial and crural indices of European late
Upper Paleolithic and Mesolithic humans. J Hum Evol 36:549 –
566.
Holliday TW. 2000. Evolution at the crossroads: modern human
emergence in western Asia. Am Anthropol 102:54 – 68.
Holliday TW, Ruff CB. 2001. Relative variation in human proximal and distal limb segments. Am J Phys Anthropol 116:26 –
33.
Jacobs K. 1985. Climate and the hominid postcranial skeleton in
Würm and early Holocene Europe. Curr Anthropol 26:512–514.
Jacobs K. 1993. Human postcranial variation in the Ukranian
Mesolithic-Neolithic. Curr Anthropol 34:311–324.
James FC, McCulloch CE. 1990. Multivariate statistical methods
in ecology and systematics: panacea or Pandora’s box? Annu
Rev Ecol Syst 211:129 –166.
Jungers WL, Falsetti AB, Wall CE. 1995. Shape, relative size,
and size-adjustments in morphometrics. Yrbk Phys Anthropol
38:137–161.
Katzmarzyk PT, Leonard WR. 1998. Climatic influences on human body size and proportions: ecological adaptations and secular trends. Am J Phys Anthropol 106:483–503.
Langsjoen OM. 1996. Dental effects of diet and coca-leaf chewing
on two prehistoric cultures of northern Chile. Am J Phys Anthropol 101:475– 489.
Leonard WR. 1989. Nutritional determinants of high altitude
growth in Nuñoa, Peru. Am J Phys Anthropol 80:341–352.
Leonard WR, Leatherman TL, Carey JW, Thomas RB. 1990.
Contributions of nutrition vs. hypoxia to growth in rural Andean populations. Am J Hum Biol 2:613– 626.
Leonard WR, Dewalt KM, Stansbury JP, McCaston MK. 1995.
Growth differences between children of highland and coastal
Ecuador. Am J Phys Anthropol 98:47–57.
Leonard WR, Dewalt KM, Stansbury JP, McCaston MK. 2000.
Influence of dietary quality on the growth of highland and
coastal Ecuadorian children. Am J Hum Biol 12:825– 837.
Lieberman DE, Devlin MJ, Pearson OM. 2001. Articular area
responses to mechanical loading: effects of exercise, age, and
skeletal location. Am J Phys Anthropol 116:266 –277.
Llagostera A. 1989. Caza y pesca marı́tima (9000 a 1000 AC). In:
Iván S, editor. Culturas de Chile prehistoria: desde sus orı́genes hasta los albores de la conquista. Santiago: Andrés Bello.
p 57–79.
Llagostera A, Torres CM, Costa MA. 1988. El complejo psicotrópico en Solcor-3 (San Pedro de Atacama). Estud Atacamenos
9:61–98.
MacCurdy GG. 1923. Human skeletal remains from the highlands of Peru. Am J Phys Anthropol 6:217–329.
Martin R. 1928. Lehrbuch der Anthropologie. Jena: Gustav Fischer.
Mayr E. 1963. Animal species and evolution. Cambridge, MA:
Harvard University Press.
McHenry HM. 1992. Body size and proportions in early hominids.
Am J Phys Anthropol 87:407– 431.
Meadows Jantz L, Jantz RL. 1999. Secular change in long bone
length and proportion in the United States, 1800 –1970. Am J
Phys Anthropol 110:57– 67.
Miller GR. 2003. Food for the dead, tools for the afterlife: zooarchaeology at Machu Picchu. In: Burger RL, Salazar LC, editors.
The 1912 Yale Peruvian scientific expedition collections from
Machu Picchu: human and animal remains. Yale University
publications in anthropology 85. New Haven: Department of
Anthropology, Yale University, and Division of Anthropology,
Peabody Museum of Natural History. p 1– 63.
Moseley ME 2001. The Incas and their ancestors, 2nd ed. London:
Thames and Hudson.
Mosimann JE, James FC. 1979. New statistical methods for allometry with application to Florida red-winged blackbirds. Evolution 33:444 – 459.
Mueller WH, Schull VN, Schull WJ. 1978. A multinational Andean genetic and health program: growth and development in
an hypoxic environment. Ann Hum Biol 5:329 –352.
Mueller WH, Murillo F, Palamino H, Badzioch M, Chakraborty R,
Fuerst P, Schull WJ. 1980. The Aymara of western Bolivia: V.
growth and development in an hypoxic environment. Hum Biol
52:529 –546.
Murra J. 1972. El “control vertical” de un maximo de pisos ecológicos en la economia de las sociedades Andinas. In: Murra J,
editor. Visita de la provincia de Leon de Huánuco (1562) Iñigo
Ortiz de Zúñigam Visitador, volume II, Huánuco. Peru: Universidad Hermillo Valdizen. p 429 – 476.
Neves WA, Costa MA. 1998. Adult stature and standard of living
in the prehistoric Atacama desert. Curr Anthropol 39:278 –281.
Neves WA, Barros AM, Costa MA. 1999. Incidence and distribution of postcranial fractures in the prehistoric population of San
Pedro de Atacama, northern Chile. Am J Phys Anthropol 109:
253–258.
Newman MT. 1953. The application of ecological rules to the
racial anthropology of the New World. Am Anthropol 55:311–
327.
BODY PROPORTIONS IN ANDEAN PREHISTORY
Newman MT. 1960. Adaptations to the physique of American
aborigines to nutritional factors. Hum Biol 32:288 –313.
Newman RW, Munro EH. 1955. The relation of climate and body
size in U.S. males. Am J Phys Anthropol 13:1–17.
Núñez AL. 1991. Cultura y conflicto en los oasis de San Pedro de
Atacama. Santiago de Chile: Editorial Universitaria.
Palomino H, Mueller WH, Schull WJ. 1979. Altitude, heredity,
and body proportions in northern Chile. Am J Phys Anthropol
50:39 –50.
Pawson IG, Jest C. 1978. The high-altitude areas of the world and
their cultures. In: Baker PT, editor. The biology of high-altitude
peoples. Cambridge: Cambridge University Press. p 17– 45.
Pearson OM. 2000. Activity, climate, and postcranial robusticity.
Curr Anthropol 41:569 – 607.
Porter WP, Gates DM. 1969. Thermodynamic equilibria of animals with environment. Ecol Monogr 39:227–244.
Roberts DF. 1953. Body weight, race and climate. Am J Phys
Anthropol 11:533–558.
Roberts DF. 1978. Climate and human variability, 2nd ed. Menlo
Park: Cummings.
Roosevelt AC, Douglas J, Brown L. 2002. The migrations and
adaptations of the first Americans: Clovis and pre-Clovis
viewed from South America. In: Jablonski NG, editor. The first
Americans: the Pleistocene colonization of the New World.
Memoirs of the California Academy of Sciences, no. 27. San
Francisco: California Academy of Sciences. p 159 –235.
Rosenberg KR, Ruff CB, Lu Z. 1999. Body size, body proportions
and encephalization in the Jinniushan specimen. Am J Phys
Anthropol 28:235 [abstract].
Ruff CB. 1991. Climate and body shape in hominid evolution. J
Hum Evol 21:81–105.
Ruff CB. 1994. Morphological adaptation to climate in modern
and fossil hominids. Yrbk Phys Anthropol 37:65–107.
Ruff CB. 2002a. Long bone articular and diaphyseal structure in
Old World monkeys and apes, I: locomotor effects. Am J Phys
Anthropol 119:305–342.
Ruff CB. 2002b. Variation in human body size and shape. Annu
Rev Anthropol 31:211–232.
Ruff CB, Walker A. 1993. Body size and body shape. In: Walker A,
Leakey R, editors. The Nariokotome Homo erectus skeleton.
Cambridge, MA: Harvard University Press. p 234 –263.
Ruff CB, Scott WW, Liu AYC. 1991. Articular and diaphyseal
remodeling of the proximal femur with changes in body mass in
adults. Am J Phys Anthropol 86:397– 413.
Ruff CB, Trinkaus E, Holliday TW. 1997. Body mass and encephalization in Pleistocene Homo. Nature 387:173–176.
Schmidt-Nielsen K. 1984. Scaling: why is animal size so important? Cambridge: Cambridge University Press.
Schreider E. 1950. Geographical distribution of the body-weight/
body-surface ratio. Nature 165:286.
Schreider E. 1957. Ecological rules and body-heat regulation in
man. Nature 179:915–916.
Schreider E. 1964. Ecological rules, body-heat regulation, and
human evolution. Evol 18:1–9.
Schreider E. 1975. Morphological variations and climatic differences. J Hum Evol 4:529 –539.
Searcy WA. 1980. Optimum body sizes at different ambient temperatures: an energetics explanation of Bergmann’s rule. J
Theor Biol 83:579 –593.
585
Sokal RR, Rohlf FJ. 1995. Biometry, 3rd ed. New York: W.H.
Freeman.
SPSS, Inc. 1998. Systat version 8.0 for windows. Chicago, Illinois.
Standen V, Allison M, Arriaza B. 1984. Patologı́as óseas de la
población Morro-1 asociada al complejo Chinchorro: norte de
Chile. Chungara 13:175–185.
Standen V, Arriaza BT, Santoro CM. 1997. External auditory
exostosis in prehistoric Chilean populations: a test of the cold
water hypothesis. Am J Phys Anthropol 103:119 –129.
Stinson S. 1980. Physical growth of high altitude Bolivian Aymara children. Am J Phys Anthropol 52:377–385.
Stinson S. 1982. The effect of high altitude on the growth of
children of high socioeconomic status in Bolivia. Am J Phys
Anthropol 59:61–71.
Stinson S. 1990. Variation in body size and shape among South
American Indians. Am J Hum Biol 2:37–51.
Stinson S. 2000. Growth variation: biological and cultural factors.
In: Stinson S, Bogin B, Huss-Ashmore R, O’Rourke D, editors.
Human biology: an evolutionary and biocultural perspective.
New York: Wiley-Liss. p 425– 463.
Stinson S, Frisansho AR. 1978. Body proportions of highland and
lowland Peruvian Quechua children. Hum Biol 50:57– 68.
Stuart-Macadam P. 1992. Anemia in past human populations. In:
Stuart-Macadam P, Kent S, editors. Diet, demography, and
disease: changing perspectives on anemia. New York: Aldine de
Gruyter. p 151–169.
Sutter RC. 2000. Prehistoric genetic and culture change: a bioarchaeological search for pre-Inka Altiplano colonies in the
coastal valleys of Moquegua, Peru, and Azapa, Chile. Lat Am
Antiq 11:43–70.
Sutter RC, Mertz L. 2004. Nonmetric cranial trait variation and
prehistoric biocultural change in the Azapa Valley, Chile. Am J
Phys Anthropol 123:130 –145.
Trinkaus E. 1981. Neanderthal limb proportions and cold adaptation. In: Stringer CB, editor. Aspects of human evolution.
London: Taylor and Francis. p 187–224.
Tsutakawa RK, Hewett JE. 1977. Quick test for comparing two
populations with bivariate data. Biometrics 33:215–219.
Verano JW. 2003. Human skeletal remains from Machu Picchu: a
reexamination of the Yale Peabody Museum’s collections. In:
Burger RL, Salazar LC, editors. The 1912 Yale Peruvian scientific expedition collections from Machu Picchu: human and
animal remains. Yale University publications in anthropology
85. New Haven: Department of Anthropology, Yale University,
and Division of Anthropology, Peabody Museum of Natural
History. p 65–117.
Villaxa A, Corrales J. 1993. Descripción y comentario de la fauna
malacológica del sitio Acha-2. In: Muñoz I, Arriaza B, Aufderheide A, editors. Acha-2 y los origenes del poblamiento humano
en Arica. Arica: Universidad de Tarapacá. p 81–90.
Walker PL. 1986. Porotic hyperosteosis in a marine-dependent
California Indian population. Am J Phys Anthropol 69:345–
354.
Weinstein KJ. 1998. Morphological evidence for cold adaptation
in pre-contact Andean skeletons from high and low altitudes.
Am J Phys Anthropol [Suppl] 26:229 [abstract].
Weinstein KJ. 2001. Comparative skeletal morphology of modern
humans and macaques from high and low altitudes. Ph.D.
dissertation, Department of Anthropology, University of Florida. Ann Arbor: University Microfilms International.
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