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10.1146/annurev.anthro.31.040402.085407

Annu. Rev. Anthropol. 2002. 31:211–32

doi: 10.1146/annurev.anthro.31.040402.085407
Copyright ° c 2002 by Annual Reviews. All rights reserved
First published online as a Review in Advance on May 21, 2002

VARIATION IN HUMAN BODY SIZE AND SHAPE

Christopher Ruff
Center for Functional Anatomy and Evolution, Johns Hopkins University School of
Medicine, 1830 E. Monument St., Baltimore, Maryland 21205; email: [email protected]

Key Words hominin, climate, nutrition, growth, adaptation

■ Abstract Evolutionary trends in human body form provide important context for
interpreting variation among modern populations. Average body mass in living humans
is smaller than it was during most of the Pleistocene, possibly owing to technological
improvements during the past 50,000 years that no longer favored large body size.
Sexual dimorphism in body size reached modern levels at least 150,000 years ago and
probably earlier. Geographic variation in both body size and shape in earlier humans
paralleled latitudinal clines observed today. Climatic adaptation is the most likely
primary cause for these gradients, overlain in more recent populations by nutritional
effects on growth. Thus, to distinguish growth disturbances, it is necessary to partition
out the (presumably genetic) long-term differences in body form between populations
that have resulted from climatic selection. An example is given from a study of Inupiat
children, using a new index of body shape to assess relative body mass.

INTRODUCTION

Body size and shape vary considerably among living human populations. Mean
body mass (weight) varies by 50% or more, within sex, in a worldwide sampling
of populations (Ruff 1994), even if Pygmies are not considered. Mean body height
(stature) and breadth (bi-iliac or maximum pelvic breadth) also vary between the
same samples, although in different ways. Variation in height is smaller (about
10%) and does not follow any particular geographic trend. Variation in breadth is
larger (about 25%) and shows a clear latitudinal gradient. The explanation for this
difference, and some other systematic human body shape differences, may lie in
basic physiological adaptive mechanisms, as discussed below.
There is abundant evidence that both body size and shape were even more
variable among Plio-Pleistocene hominins (e.g., Jungers 1988, Ruff 1991, Aiello
1992, McHenry 1992) and, within geographically dispersed taxa, followed clines
that were similar to those found among modern humans (Trinkaus 1981; Ruff &
Walker 1993; Ruff 1994; Holliday 1997a,b). General temporal trends in body size
are also apparent in the fossil record (e.g., Ruff et al. 1997). An appreciation of such
variation among human ancestors is important for several reasons: (a) Body size
(and within-taxon variation in body size) is related to many other characteristics
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of a species, including life history parameters, ecology, and social organization

(Calder 1984, Schmidt-Nielson 1984); thus, it is often used to predict these traits
in fossil taxa [notwithstanding some significant problems in doing so (see Smith
1996)]. (b) Body size is the usual “denominator” for assessing key evolutionary
trends in the hominin lineage, including changes in relative brain size (encephaliza-
tion), tooth size (megadontia), bone strength (robusticity), and gut size (Pilbeam &
Gould 1974; McHenry 1976, 1984, 1988; Ruff et al. 1993, 1997; Aiello & Wheeler
1995; McHenry & Coffing 2000). (c) Differences in body shape have been used as
population or taxonomic markers among past hominins and to identify migrational
and possible interbreeding events (Trinkaus 1981; Holliday 1997a,b; Duarte et al.
1999; Ruff et al. 2002). (d) The body size and shape of earlier humans can serve
as effective baselines for assessing more recent temporal or geographic variation,
for example, the effects of changes in subsistence strategy and the significance of
recent secular trends (Frayer 1984, Tobias 1985, Ruff et al. 1997) (see below). Ap-
preciation of long-standing differences in body form between human populations
can help inform decisions regarding the most appropriate methods for assessing
the health and nutrition of living individuals. Both absolute measures of body
size (height, weight) and relative indices reflecting body shape [body mass index
(wt/ht2)] are often used as standards for evaluating growth and under- and over-
nutrition (WHO 1995). To the extent that these characteristics vary systematically
in response to factors other than health and nutrition, such standards may not be
universally applicable across populations.

EVOLUTIONARY TRENDS IN BODY SIZE

Methodological Considerations
Because body size reconstruction in the fossil record almost always depends on
extrapolation from fragmentary remains, it is important to clearly understand the
rationale behind different reconstruction methods. A purely statistical approach
considers the relationship between body mass (or stature) and skeletal/dental fea-
tures in a modern reference sample and applies resulting prediction equations to
the fossil fragments, perhaps preferring those equations with the smallest estima-
tion errors in the modern sample. The problem with such an approach is that it
implicitly assumes equality—proportional and/or functional—between the refer-
ence sample and the individual to which it is applied, which may or may not be
true. For example, equations for predicting stature from long bone lengths devel-
oped in European samples severely overestimate stature in East Africans because
of their relatively longer limb-to-trunk lengths (Allbrook 1961). This result is pre-
dictable given known systematic differences in limb proportions among modern
human populations (Roberts 1978). The same rationale applies to reconstruction
of stature in the Homo erectus KNM-WT 15000, who also had apparently very
long limbs (Ruff & Walker 1993). Another example would be the use of tooth
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HUMAN BODY SIZE AND SHAPE 213

size to predict body mass in the hominin lineage, a common practice in primate
paleontology (e.g., Gingerich et al. 1982, Conroy 1987) but one that is obviously
biased in this case because of temporal (and probably taxonomic) differences
in relative tooth size among hominins (Pilbeam & Gould 1974, McHenry 1984,
McHenry & Coffing 2000, Teaford et al. 2002). Another not-so-obvious example is
the use of long-bone diaphyseal breadth to predict body mass in hominins, a com-
monly employed procedure in the past (McHenry 1976, Oleksiak 1986, Rightmire
1986, Hartwig-Scherer 1994). It is evident that long-bone diaphyses change their
diameters in response to mechanical loading (Trinkaus et al. 1994) and that Plio-
Pleistocene hominins had relatively thicker diaphyses than modern humans (Ruff
et al. 1993, 1994; Ruff 1998). Thus, use of diaphyseal cross-sectional dimensions
and a modern reference sample will lead to systematic overestimates of body mass
in fossil hominins.
These examples illustrate the importance of considering the functional signif-
icance of skeletal/dental traits when using them for body size reconstruction. In
many cases this will involve their mechanical significance because mechanical fac-
tors have such a pervasive influence on skeletal form (see references above). The
mechanics of weight-bearing in bipeds, in fact, argues for the use of lower limb-
bone dimensions for body mass reconstruction in hominins. Articular dimensions
have been shown to be less sensitive to differences in activity level than are diaphy-
seal breadth dimensions (Ruff et al. 1994, Trinkaus et al. 1994, Lieberman et al.
2001); thus, they should more accurately reflect variations in body mass without
the potentially confounding effects of individual behavioral differences. For these
reasons, lower limb articular size, and particularly femoral head size (because the
femoral head is often preserved and is easily measurable), is an effective predictor
of body mass in hominins (Ruff et al. 1991, 1997; McHenry 1992; Grine et al.
1995).
Mechanically based methods for estimating body mass can be distinguished
from morphometric methods (Ruff 1994, 2000b) that rely on direct reconstruc-
tion of body dimensions from preserved bone dimensions. The latter usually in-
volve estimation of stature or body length, followed by estimation of body mass
from stature, assuming some specified relationship between the two (Mathers &
Henneberg 1995, Porter 1995). Using a cylindrical model of the human body (Ruff
1991), body mass should be predictable from stature and body (cylinder) breadth.
In fact, body mass estimates from stature and bi-iliac breadth in living humans
are remarkably good (Ruff 1994, 2000b). It is critical when estimating stature, or
bi-iliac breadth (if necessary), to use appropriate modern reference samples, i.e.,
modern populations with body proportions similar to the specimens being esti-
mated (Holliday & Ruff 1997). Fortunately, strong geographical trends in body
shape, as well as information gleaned from more complete specimens, can provide
guidance with the more usual fragmentary specimens (Ruff et al. 1993; Ruff &
Walker 1993; Trinkaus & Ruff 1999a,b). Body mass estimates from femoral head
size and a multiple regression using stature and bi-iliac breadth give similar results,
on average, when applied to Pleistocene hominin specimens (Ruff et al. 1997).
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General Temporal Trends

Figure 1 shows body mass estimates for individual specimens of Pleistocene Homo
species derived from regressions on femoral head size and/or bi-iliac breadth and
stature (Ruff et al. 1997), mean estimates for other Plio-Pleistocene hominin taxa
taken from the literature, mainly McHenry (1992), and modern human sex-specific
population means from a worldwide sampling, not including Pygmies (Ruff 1994).
All estimates for skeletal/fossil specimens were derived from postcranial elements.
The Pleistocene Homo specimens include 163 individuals used in a previous anal-
ysis (Ruff et al. 1997) plus two recently described Middle Pleistocene specimens
from Yinnuishan, China (Rosenberg et al. 1999) and Atapuerca, Spain (Arsuaga
et al. 1999). Three other specimens attributed to Homo habilis sensu stricto
(Olduvai 8 and 36 and KNM-ER 3735), whose body masses were estimated by
McHenry (1992), are shown enclosed in parentheses because their attribution to the
genus Homo has been questioned (Wood & Collard 1999). [Other early Homo spec-
imens plotted here, including KNM-ER 3228, 1481, and 1472, are not definitely
attributable to any particular species but appear to be most similar in morphology
to later H. ergaster/erectus rather than H. habilis sensu stricto (McHenry 1994b,
McHenry & Coffing 2000).] Among the Homo specimens, those from higher
(above 30◦ ) and lower latitudes are distinguished by different symbols. (All Aus-
tralopithecus specimens are from lower latitudes.) Two dotted lines representing
the mean body masses of the modern higher- and lower-latitude samples are plotted
for reference. Because of differences in data sampling density, the temporal axis
is given in three different scales with break points at 2 million and 100,000 years
ago.
Several temporal trends in hominin body mass are apparent from Figure 1.
First, there is a marked increase in body size with the appearance of early Homo
(except H. habilis sensu stricto) about 2 million years ago. This contrast between
Homo and earlier and contemporaneous australopithecines has been previously
noted (e.g., McHenry 1994a, Wood & Collard 1999, McHenry & Coffing 2000).
The explanation for the increase in body size in Homo is not clear but may be
related to a commitment to a fully terrestrial lifestyle, inhabitation of more open
environments, increased foraging distances, and/or dietary shifts (Foley 1987,
Wheeler 1992, McHenry 1994a, Leonard & Robertson 1997, Klein 1999). The
lack of such an increase in H. habilis sensu stricto, with its attendant behavioral
implications, is one argument against the inclusion of this taxon within the genus
Homo (Wood & Collard 1999).
Second, there is an increase in average body size at about 500,000 years ago,
corresponding to the first hominin postcranial remains recovered from higher lati-
tudes. It is well documented that among modern humans body mass is distributed
clinally, increasing in higher latitudes (Roberts 1978, Ruff 1994, Katzmarzyk &
Leonard 1998), as reflected in the living-human data points in Figure 1. This has
been interpreted as an example of Bergmann’s Rule (Mayr 1956), which is gen-
erally explained as an adaptation to decreased heat loss in colder environments.
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Figure 1 Estimated body mass of Plio-Pleistocene hominins. Letters represent species means of early taxa: A, Ardipithecus ramidus;
B, Australopithecus anamensis; C, Australopithecus afarensis; D, Australopithecus africanus; E, Paranthropus robustus; F, Paranthropus
boisei. (A) Based on similarity in size of humerus to middle of range for A. afarensis (White et al. 1994); (B) Leakey et al. 1995; (C–F ):
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McHenry 1992. Symbols represent individual Homo specimens ( Ruff et al. 1997, Arsuaga et al. 1999, Rosenberg et al. 1999). Symbols in
HUMAN BODY SIZE AND SHAPE

parentheses are individual H. habilis sensu stricto (McHenry 1992). Living humans are sex/population-specific means (Ruff 1994). Open
symbols, lower latitude; solid symbols, higher latitude; dotted and dashed lines through living human means for higher and lower latitudes,
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respectively. Myr, millions of years before present. (Adapted from Ruff 2001, with permission from Elsevier Science.)
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Thus, it would be expected that a range expansion into colder climates would in-
crease the average body mass (and total range in body mass) of the genus Homo.
Body masses for individual Pleistocene Homo specimens are generally above the
means for living humans from the same latitudinal zones, on the order of 10%
higher on average (Ruff et al. 1997). The difference is even greater if only archaic
Homo specimens (Neandertals and other specimens dated to before 100,000 years
ago) are considered: Higher-latitude archaic Homo specimens average almost 20%
greater in body mass than the means for living higher-latitude populations. This
figure may be somewhat inflated by a male preservational bias in the fossil record
(Ruff et al. 1997), but even so, it is evident that Middle Pleistocene and early
Late Pleistocene humans were significantly larger on average than living humans.
A large body size is also consistent with evidence from material culture from this
time period, such as very large Middle Pleistocene throwing spears (Thieme 1997).
Third, there is a decline in average body mass beginning about 50,000 years
ago. The decline is best demonstrated among higher-latitude samples and is equiv-
alent whether or not archaic humans (Neandertals) are included (Ruff et al. 1997).
Lower-latitude samples with body mass estimates are unfortunately rare in the Late
Pleistocene. However, similar declines in the Late Pleistocene and early Holocene
in other body dimensions, including estimated stature and cranial size, have been
demonstrated in many areas of the world, including Europe, the Mediterranean re-
gion, sub-Saharan Africa, South Asia, and Australia (Frayer 1980, 1984; Kennedy
1984b; Meiklejohn et al. 1984; Jacobs 1985; Henneberg 1988; Brown 1992;
Henneberg & Steyn 1993; Formicola & Giannecchini 1998). The decline con-
tinues through the Neolithic, after which it is reversed in European samples. No
reversal in this negative trend in size has been reported for lower-latitude sam-
ples, except possibly for some very recent increases in some areas over the past
few decades (Katzmarzyk & Leonard 1998). As a result, living higher-latitude
populations are about as large as terminal Pleistocene samples, whereas living
lower-latitude populations are smaller on average than they were 10,000 years ago
(Figure 1) (see Ruff et al. 1997).
Several reasons, not mutually exclusive, have been proposed for the Late
Pleistocene–Early Holocene decline in human body size (see references above).
These include technological improvements that decreased the selective advantage
of a larger body (which is also metabolically expensive to maintain), a decline in
nutritional quality, climatic factors (adaptation to a warming environment), and
reduced gene flow (inbreeding). A reduction in body size has been observed in
many, although not all, populations undergoing a transition from a foraging to a
food producing economy, especially an intensive agricultural economy (Cohen &
Armelagos 1984a). Explanations for this trend usually focus on increases in mal-
nutrition, overcrowding, and spread of infectious diseases. Institution of a class
system with resulting social inequities in resource allocation may also be a factor
in some populations (Bogin & Keep 1999). However, because body size reduc-
tion occurred in many areas of the world prior to the adoption of food production
(or where food production was never adopted, i.e., Australia), this factor may be
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HUMAN BODY SIZE AND SHAPE 217

contributory but does not fully explain the trend. Given the timescales involved
and the known effects of malnutrition on growth, it seems likely that changes as-
sociated with food production were developmental rather than genetic in nature.
Recent positive secular trends over the past few hundred years are also most likely
to be developmental and related to increased nutritional and overall health levels
(Van Wieringen 1986, Eveleth 1994, Stinson 2000). Such positive trends have not
been observed in many areas of the world, primarily in lower latitudes that have
not enjoyed the same improvements in environment (Malina et al. 1983, Tobias
1985, Pretty et al. 1998). At the same time, the positive secular trend in body size
in (mainly) higher-latitude populations may have leveled off over the past few
decades (Kimura 1984, Stinson 2000).
One possible explanation for these observations is that the Late Pleistocene
reduction in body size was due primarily to genetic factors, possibly reduced se-
lection for large body size in association with technological improvements (Frayer
1984), whereas the succeeding fluctuations (decrease, then, in higher latitudes, in-
crease) in body size in the Holocene were due to environmental effects on growth,
e.g., nutrition. Thus, many higher-latitude populations may have recently achieved
their maximum “genetic potential” in body size (stature), originally established
in the terminal Pleistocene, whereas many lower-latitude populations have not.
Additional Late Pleistocene body-mass data from lower-latitude samples would
be useful in further testing this hypothesis.

Sexual Dimorphism
Figure 2 shows sexual dimorphism in estimated body mass for early hominin taxa
and regional/temporal groupings of H. sapiens. Mean male and female body masses
for Australopithecus, Paranthropus, and early Homo are from McHenry & Coffing
(2000), mean sex-specific body masses for Pleistocene H. sapiens are from the data
set used in Ruff et al. (1997), and values for individual living human populations
(n = 19) are from Ruff (1994). The mean sexual dimorphism (male/female) in body
mass in the living human populations is about 15%, similar to average figures
derived from other living population samples (Stini 1974, McHenry & Coffing
2000). Sexual dimorphism is similar in higher-latitude (mean 15.5%) and lower-
latitude (mean 14.7%) living samples (p > 0.75, t test). The same is true for the Late
Upper Paleolithic sample—the only Pleistocene sample with a sufficient number
of individuals from higher and lower latitudes to subdivide in this way—with
16.1% and 15.1% sexual dimorphism for higher and lower latitudes, respectively.
Thus, whereas latitude affects average body mass, it appears not to affect sexual
dimorphism in body mass in any systematic way. This is the pattern that would be
expected if sexual dimorphism in body mass in humans were more influenced by
environmental/behavioral factors other than climate per se.
All taxonomic or regional/temporal groupings of Homo specimens have levels
of body-mass sexual dimorphism comparable to those of modern humans (7%–
18%) (Figure 2). In contrast, most earlier and contemporary australopithecines,
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Figure 2 Sexual dimorphism in body mass in hominin taxa or regional/temporal

groupings. See text for origin of data. “Early archaic Homo sapiens” corresponds to
“early Late Pleistocene” in Ruff et al. 1997. Line indicates living human mean.

including A. afarensis and A. africanus, and Paranthropus boisei, show clearly el-
evated levels of sexual dimorphism (37%–55%). Paranthropus robustus is slightly
elevated (25%) although just within the range of modern human populations. Some
of the estimates for the earlier species are heavily dependent on taxonomic attri-
butions, which can be problematic for isolated postcranial elements; thus, such
estimates can change substantially if groups are redefined or additional specimens
are added to particular taxa (McHenry 1992, 1994a; McHenry & Coffing 2000).
For example, based on newly identified and some reclassified specimens from
Swartkrans, South Africa, Susman et al. (2001) estimated body-mass sexual di-
morphism to be about 40% in P. robustus, equivalent to that in P. boisei and A.
africanus (Figure 2), and even larger in Homo sp. from the site, about 80%. This
latter figure is similar to earlier estimates of sexual dimorphism in H. habilis more
broadly defined (sensu lato) (McHenry 1992). Another problem in such compar-
isons is that almost none of the earlier specimens have any associated indicators
of sex; thus, sex assignment is made on the basis of size alone (larger = male,
smaller = female), often across different sites and temporal ranges, which may ex-
aggerate the degree of estimated dimorphism. In contrast, all of the Late Pleistocene
sex assignments shown in Figure 2 for H. sapiens are based either on associated
pelvic or cranial material or on comparisons within site-specific samples.
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HUMAN BODY SIZE AND SHAPE 219

Perhaps the most conservative assessment of the data shown in Figure 2 is that
sexual dimorphism in body mass definitely reaches modern levels with archaic
Homo sapiens1, and very possibly earlier with H. ergaster/erectus, but was likely
larger in australopithecines and possibly some early species of Homo. The be-
havioral and ecological significance of increased sexual dimorphism in body size
in at least some early hominin taxa has been debated (see McHenry 1991, 1992,
1994a; Plavcan & Van Schaik 1997; Plavcan 2001 and references therein). The
lack of concurrent marked sexual dimorphism in canine tooth size may imply a
social/behavioral system unique among primates (Plavcan & Van Schaik 1997,
Plavcan 2001). To the extent that sexual dimorphism in body size reflects social
organization, the data for archaic H. sapiens indicate that by the Late Pleistocene,
at least, patterns of social organization may have been similar to those of modern
human foragers. This is concordant with evidence from lower limb bone structure,
which indicates a sexual division of labor in archaic H. sapiens similar to that of
modern hunter-gatherers (Ruff 1987, 2000a).
More recent variations in body-mass sexual dimorphism, not shown in Figure 2,
likely reflect more subtle differences in subsistence strategy, diet, and possibly
sex-related buffering against the environment (Stini 1974; Frayer 1980, 1984;
Meiklejohn et al. 1984; Jacobs 1985; Stinson 1985; Jantz & Jantz 1999). Secular
trends in stature over the past few hundred years have been in similar directions for
males and females of the same populations (e.g., Meredith 1976, Bogin & Keep
1999), although again with possible subtle differences in magnitude (Jantz & Jantz
1999).

EVOLUTIONARY TRENDS IN BODY SHAPE

There is abundant evidence that many early hominin taxa (australopithecines and
H. habilis sensu stricto) had basic body proportions that were significantly different
from those of modern humans. Upper limbs were large relative to lower limbs, and
body breadth (and mass) was large relative to stature (McHenry 1978, Johanson
et al. 1987, Leakey et al. 1989, Hartwig-Scherer & Martin 1991, Ruff 1991, Aiello
1992, McHenry & Berger 1998, Asfaw et al. 1999). These differences are probably
best accounted for by retained arboreal capabilities combined with the allometric
effects of small body size, which itself may be related to locomotor patterns (see
above). By 1.5 million years ago body proportions, together with body size, in
H. ergaster/erectus were well within the range of modern human variation (Ruff
& Walker 1993). From this point on, within the Homo lineage, a fully terrestrial

1
The Sima de los Huesos skeletal sample from Atapuerca, Spain, dated to about 200,000
years ago, preserves the remains of many individuals (Arsuaga et al. 1999). Most lower
limb remains from the site have not yet been formally described, but a number of acetabular
breadths are given in the above reference, some of which have sexually diagnostic regions of
the pelvis preserved. The males average 60 mm and the females 52 mm, a 15% difference.
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lifestyle can be inferred, with variations in body shape viewed as adaptations to

environmental variables other than basic locomotor behavior.
One environmental variable that has long been postulated to influence modern
human body shape is climate (see references in Ruff 1994). Relative limb length
(Allen’s Rule) and body breadth vary systematically with temperature in modern
human populations, with those from colder environments having relatively shorter
limbs (and shorter distal segments within limbs) and wider bodies than those from
warmer environments (Roberts 1978, Trinkaus 1981, Ruff 1994). These clinal vari-
ations, and the observed cline in body mass discussed earlier, can all be viewed as
part of the same adaptive strategy to reduce body surface area/body mass in colder
climates and increase it in warmer climates (Schreider 1964; Ruff 1991, 1994).
Of course, body proportions can be affected by other environmental variables as
well. For example, relative limb length may increase with improved nutrition and
health status (Tanner et al. 1982). However, this factor cannot explain the rela-
tively longer limbs observed in modern tropical populations, most of whom are
more nutritionally deprived than higher-latitude populations. Thus, genetic fac-
tors (climatic selection) are likely involved (Stinson 2000). Interestingly, there
is evidence that body (bi-iliac) breadth is not as affected by nutritional or other
developmental influences as limb length proportions, perhaps in part because of sta-
bilizing selection owing to obstetric requirements (Ruff 1994). In terms of climatic
adaptation, changes in absolute body breadth will always affect surface area–to–
body mass ratios, whereas changes in stature will not (Ruff 1991). Thus, (skeletal)
body breadth shows very strong latitudinal trends, whereas stature does not
(Ruff 1994).
Where possible to evaluate, body proportions in earlier hominins have been
found to follow the same geographic clines as modern humans. Thus, for example,
Neandertals have relatively short distal limb segments (Trinkaus 1981) and broad
bodies (Ruff 1994), whereas the opposite is true for KNM-WT 15000, a juvenile
H. ergaster/erectus from East Africa (Ruff & Walker 1993). Furthermore, with the
discovery of new fossil specimens over the past several years, it is now apparent
that the Neandertal body type is actually part of a broader pattern shared with
other higher-latitude archaic Homo specimens. These new specimens include the
Yinnuishan female, dated to about 280,000 years ago (Rosenberg et al. 1999), and
the Atapuerca “Pelvis 1” male, dated to at least 200,000 years ago (Arsuaga et al.
1999). Both specimens preserve enough of the pelvis to allow measurement or
accurate estimation of bi-iliac breadth. Yinnuishan also preserves an ulna from
which forearm length can be assessed. The Atapuerca Sima de los Huesos remains
are all unassociated, but an approximate ulnar length can be calculated from a com-
plete humerus recovered at the same site (“Humerus II”) (Carretero et al. 1997).
The pairing of the pelvis and humerus is reasonable, because both were at the
upper end of the size range represented at the site and so very likely represented
large males. Ulna length was calculated from humeral length using a regression
formula based on a combined sample of modern East Africans and Pecos Pueblo
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HUMAN BODY SIZE AND SHAPE 221

Amerindians (Ruff 1995): ulna = 0.873· humerus − 12.4 (r = 0.910, standard er-
ror of estimate = 8.4 mm). This is a conservative estimate in terms of testing
whether the Atapuerca specimen had relatively short limbs because East Africans
have typically tropical proportions (long forearms); using only the Amerindians
as a reference sample would produce an ulna length estimate several millimeters
shorter.
Figure 3 plots ulna length against bi-iliac breadth for the Yinnuishan and
Atapuerca specimens, together with similar data for several other Pleistocene spec-
imens and some modern comparative samples. The modern samples include East
African (Ruff 1995) and combined Alaskan Inupiat (Eskimo) and Aleut skele-
tal samples. Because modern males have relatively longer forearms than females
(Trinkaus 1981), sexes are distinguished in the plot.
As expected, the modern tropical sample has much longer forearms relative to
body breadth than the high-latitude sample: There is no overlap between same-
sex individuals from the two groups. Interestingly, a line indicating equivalent

Figure 3 Ulna maximum length against bi-iliac (maximum pelvic) breadth in Pleis-
tocene and modern Homo. Open symbols, modern East Africans; filled symbols, mod-
ern Eskimos/Aleuts; squares, males; circles, females. Open stars, KNM-WT 15000
and Sahaba 16; filled stars, La Chapelle 1, Kebara 2, Atapuerca “Pelvis 1”, Yinnuishan
(see text); diagonal line, equivalent ulnar and bi-iliac measures.
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ulnar length and bi-iliac breadth, included in the plot, almost perfectly divides
the two groups. Multiple analyses of variance (ANOVA) indicate that both sex
and latitude have highly significant effects on proportions in the modern sam-
ples (p < 0.0001). The Yinnuishan and Atapuerca specimens are “hyper-arctic,”
with very wide bodies and short forearms. Two Neandertals with pelvises suffi-
ciently preserved to measure or estimate bi-iliac breadth [Kebara 2 (Rak & Arens-
burg 1987) and La Chapelle 1 (Boule 1911, Ruff et al. 1993)] also fall in the
high latitude range. (Ulna length for La Chapelle was estimated from its radius
length using a formula derived from modern East Africans and Pecos Amerindi-
ans: ulna = 1.008 · radius + 16.2, r = 0.988, standard error of estimate = 3.2 mm.)
The Atapuerca and La Chapelle specimens are quite similar in proportions, with
Yinniushan falling below these two. The extreme proportions of Yinnuishan may
be attributable in part to sexual dimorphism: The difference between this female
specimen and the Atapuerca and La Chapelle 1 males is about equal to the aver-
age difference between modern females and males from the same latitudinal zone
(Figure 3). Kebara 2 (also a male) is somewhat less hyper-arctic, reflecting the
longer forearms (and possibly somewhat more linear body form) characteristic
of Near Eastern as opposed to European Neandertals (Trinkaus 1981, Ruff 1994,
Ruff et al. 2002).
Two Pleistocene specimens from tropical regions are also included in Figure 3:
KNM-WT 15000, a juvenile male H. ergaster/erectus (Walker & Leakey 1993),
and Jebel Sahaba 26, a female from a terminal Pleistocene site in Nubia (Wendorf
1968). The juvenile status of KNM-WT 15000 should have relatively little effect
on comparisons (Ruff & Walker 1993). Both specimens fall close to the average
proportions for sex-matched modern individuals from East Africa.
Thus, the available evidence supports latitudinal clines in body shape in earlier
Homo similar to those found in modern humans. This in turn suggests a similar
physiological mechanism as an explanation, i.e., climatic adaptation. As noted
earlier, there are very few available Middle and Early Late Pleistocene specimens
from lower latitudes, and none with enough postcranial elements to directly assess
body proportions. These would provide very useful comparisons with archaic
Homo specimens from higher latitudes because the latter show the most extreme
examples of apparent adaptation to colder climates. Without this direct contrast
it can be argued, for example, that the broad pelvises of higher-latitude archaic
Homo have more phylogenetic rather than physiological implications (Arsuaga
et al. 1999). However, other less direct evidence also supports a latitudinal cline in
body shape within the Middle Pleistocene. The complete tibia from Broken Hill,
Zambia (E691), is very long, in the Pleistocene equaled in length only by specimens
with high crural indices (long tibias relative to femora) (Kennedy 1984a). On the
basis of shaft external breadth or cross-sectional properties to length proportions,
the Broken Hill tibia and other lower-latitude tibias from the Middle Pleistocene
are clearly distinct from the broadly contemporaneous higher-latitude Boxgrove 1
tibia (Stringer et al. 1998, Trinkaus et al. 1999). The most parsimonious explanation
is that the Boxgrove specimen had a relatively wide body and short limbs (like
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HUMAN BODY SIZE AND SHAPE 223

Neandertals, Atapuerca, and Yinnuishan), whereas the lower latitude specimens

had relatively narrow bodies and longer limbs (like KNM-WT 15000).
The transition in Europe from Neandertals to “early anatomically modern”
(Late Paleolithic) humans 40,000 to 25,000 years ago and subsequent changes in
morphology within the latter group, are especially interesting in that they may
provide evidence of adaptation following migration to a new climatic zone if these
populations were derived from farther south, as suggested by the preponderance
of current evidence (Klein 1999). The lack of change between European Early and
Late Paleolithic samples in distal-to-proximal limb length proportions (crural and
brachial indices) was initially puzzling in this regard because a reduction would
have been predicted if climatic adaptation were taking place (Trinkaus 1981).
However, more recent work has shown that relative to measures of trunk (vertebral
column) height, limb length did decrease significantly within the Upper Paleolithic
in Europe, beginning at proportions similar to those of sub-Saharan Africans and
ending at proportions similar to those of modern Europeans (Holliday 1997a).
Comparisons of long bone lengths to bi-iliac breadths in available European Upper
Paleolithic specimens (n = 15–19, about a third from the Early Upper Paleolithic)
also indicate significant reductions in limb length to body breadth between the Early
and Late Upper Paleolithic (unpublished results based on data given in Ruff et al.
1997, supplementary information). Thus, body shape did change significantly in
Upper Paleolithic Europeans after exposure to colder climatic conditions, although
the change was mosaic in nature, beginning with a general reduction in limb
lengths followed by a reduction in distal-to-proximal limb element proportions.
The possible significance of this difference in timing has been further explored
elsewhere (Ruff et al. 2002).

RELATIVE BODY MASS IN LIVING POPULATIONS

What can the patterns of variation in body size and shape observed in pre-Holocene
humans tell us about more recent human variation? If the body size of terminal
Late Pleistocene humans does indeed represent a genetic target (or limit) for body
size in living humans, then data on such populations, prior to the major changes
in diet, social organization, etc., characteristic of the Holocene, could serve as
effective baselines for evaluating current body size. As noted earlier, there is evi-
dence that many living populations from developing countries may not be as
large as their Pleistocene ancestors. Of course, this does not address the issue of
whether maximum potential body size is desirable or not (for a review see Stinson
1992).
Here I concentrate on another issue: how differences in relative body mass, i.e.,
body shape, between modern populations should be used in health assessment.
The above review indicates that body shape varies systematically along latitudi-
nal clines in both modern humans and Pleistocene Homo. Nutrition and health
status do not seem to explain these trends, which are, however, consistent with
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224 RUFF

climatic adaptation. The temporal depth of such clines and their persistence despite
other confounding factors argues that they represent long-standing mechanisms
of adaptation within our lineage and are probably at least in part genetically based.2
The body mass index (BMI), weight/height2 (also known as Quetelet’s Index),
is widely employed in assessing relative under- or over weight status of living
individuals and populations (WHO 1995). Standard percentiles for BMI at various
ages during childhood and in adults have been developed from U.S. population
surveys (WHO 1995). However, questions have been raised concerning the ap-
plicability of such standards to other populations, owing to the effects of varying
body proportions and lean body mass fraction on the index (Garn et al. 1986) (some
of these limitations were also noted in the WHO recommended guidelines cited
above). The effects of differences in relative lower limb length (or relative sitting
height) on body mass indices have been known for many years (Bardeen 1923).
Arctic populations represent an extreme in modern human body shape variation,
with relatively short limbs and wide bodies (see Figure 3). This contributes to their
high weight-for-height indices, despite the fact that they do not have increased
skinfold thicknesses compared with U.S. standards (Schaefer 1977, Johnston et al.
1982, Jamison 1986). This pattern is apparent among Eskimo (Inupiat and Inuit)
children and adults and has led to recommendations that different standards be
developed for identifying obesity in these populations (Schaefer 1977, Jamison
1986). One approach would be to develop population-specific weight-for-height
standards for such groups. The problems with this approach are that (a) it would
necessarily rely on a relatively small data set and would not take advantage of
the very extensive samples available in large population surveys that allow more
secure statistical inferences (e.g., Hamill et al. 1979), and (b) results could be
confounded by very recent and marked secular changes occurring in some of these
groups in diet, lifestyle, and anthropometrics (Schaefer 1970, Jamison et al. 1978).
An alternate approach would be to use a different index of relative weight that
is less affected by, or incorporates, variations in body build including relative limb
length and relative (skeletal) body breadth. Sitting height is a commonly taken
anthropometric measurement that reflects trunk height, i.e., a body length measure
that does not include the limbs. Sitting heights of Eskimo children were found to be
similar to those of age-matched U.S. whites and blacks, despite large differences
in stature (Johnston et al. 1982). Bi-iliac breadth is the best available trunk breadth
measurement (see Ruff 1991). As noted above, it also shows strong systematic
differences between human populations. The product of trunk height and bi-iliac
breadth provides an index of “trunk frame size” (Ruff & Jamison 2002). The ratio
of body mass to trunk frame size can then be considered a “trunk frame index”
(TFI). Both sitting height and bi-iliac breadth are expressed in cm and body mass
in kg; the resulting ratio is multiplied by 1000.

2
Note that this does not imply any close genetic ties between earlier Homo species and
modern humans from the same geographic regions, but rather the operation of the same
adaptational principles to both earlier and more recent populations.
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HUMAN BODY SIZE AND SHAPE 225

Figure 4 Two body mass indices applied to 9-year-old Inupiat and U.S. white chil-
dren (see text for derivation of samples). Horizontal lines indicate U.S. white medians.
(A) Traditional body mass index (BMI); (B) trunk frame index (TFI).

Figure 4 compares use of the TFI and BMI indices as applied to Inupiat and
U.S. white children. The Inupiat data are from northern Alaska and were collected
around 1970 by Jamison and colleagues (Jamison 1978, 1986, 1990). Individual
raw anthropometric data were obtained courtesy of P.L. Jamison (personal com-
munication). The U.S. white data are from the Denver Growth Study, collected
over several decades (McCammon 1970), although all of the measurements in-
cluded here were taken in the 1950s. Individual data for 20 subjects (10 males
and 10 females), followed longitudinally from near birth to late adolescence,
were obtained from the study database (R.M. Siervogel, personal communica-
tion). An age of 9 years was chosen for comparisons, because it avoids compli-
cations owing to differences in timing of the adolescent growth spurt and is an
age with little sexual dimorphism in the BMI, allowing pooling of males and fe-
males (WHO 1995). (The same similarity between the sexes at this age, within
group, was confirmed here for both BMI and TFI.) Anthropometric data were
available at this age for 19 Inupiat children (13 females and 6 males, “nonhybrids”
only).
BMI is significantly higher in Inupiats than in U.S. whites (p < 0.01, t test). In
fact, all of the Inupiat children fall above the median BMI for the Denver sample
(Figure 4A). In contrast, there is no significant difference between Inupiat and U.S.
white children for TFI (p > 0.50), and the distribution of individual data points,
with the exception of one high Inupiat outlier, is very similar in the two groups
(Figure 4B).
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226 RUFF

These Inupiat children, like other Eskimo children, do not have large skinfolds.
Mean skinfold breadths for 9-year-old boys and girls, respectively, are 8.1 mm
and 10.0 mm for triceps and 5.5 mm and 7.1 mm for subscapular [unlike BMI,
there is sexual dimorphism for skinfolds at this age (WHO 1995)]. The triceps
skinfolds are slightly below, and the subscapular skinfolds slightly above the 50th
percentiles for 9-year-olds based on U.S. standards (WHO 1995) or the Denver
Growth Study sample (Hansman 1970).
Thus, for the Inupiat sample as a whole, the TFI appears to more faithfully rep-
resent relative fatness than does the BMI. The same may be true on an individual
basis. The one high outlier for both BMI and TFI is also a high outlier for skinfold
values (above the 90th percentile for her age according to the same reference sam-
ples), i.e., both indices successfully identified her as overweight relative to direct
subcutaneous fat measurements. The next highest BMIs among the Inupiats were
from two boys, both above the 85th percentile for their age, the recommended
cutoff for “at risk for overweight” (WHO 1995). Interestingly, despite identical
BMIs in these boys (18.9), skinfold breadths were high in one (13.0 mm and 9.0 mm
for triceps and subscapular, respectively) but not in the other (6.0 mm and
5.0 mm). The individual with the large skinfolds had a relatively high TFI of
21.7—the second highest index in the group—whereas the individual with the
moderate skinfolds had a TFI of 20.1, closer to the median of the group (see
Figure 4). The identical BMIs but differing TFIs can be attributed to body build
differences: The individual with the smaller skinfolds was relatively larger in both
bi-iliac breadth and sitting height (>1 sample standard deviation above the other
boy for bi-iliac breadth/stature and sitting height/stature). Thus, the BMI tended to
overestimate his fatness level, whereas the TFI, which incorporates trunk shape,
did not. A noncorrespondence between secular changes in skinfold measurements
and BMI has also been noted for Eskimo (Inuit) adults (Leonard 2000), which
could be due to similar body shape differences (see Schaefer 1970).
Although this discussion has focused on identification of overweight status in In-
upiats, an equally valid question could be posed regarding underweight. As Jamison
(1990) has noted, we cannot tell from current weight-for-height data based on U.S.
standards where a borderline malnourished Inupiat child would fall. The same rea-
soning applies, of course, to populations in other areas of the world where malnutri-
tion is severe and that also exhibit apparently intrinsic (genetic) differences in body
proportions from U.S. standards, e.g., relatively longer limbs and narrower bod-
ies in lower-latitude populations. Incorporation of such proportional differences
into body mass indices should provide more sensitive and less population-biased
measures of under- and overweight. The apparent developmental lability of some
body proportions, e.g., lower limb length relative to stature (Tanner et al. 1982),
does not negate this argument. It is important to recognize such secular trends in
skeletal proportions if soft tissue differences are to be distinguished from general
body shape changes. For example, Takahashi (1986) found that among Japanese
older adolescent and younger adult women, stature had increased more than body
mass in the 1970s and early 1980s, which the author attributed primarily to dietary
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HUMAN BODY SIZE AND SHAPE 227

and lifestyle changes resulting from a desire “to be slimmer” and “to keep from
getting fat.” However, no clear secular trend in skinfold thicknesses could be dis-
cerned. The implication is that (skeletal) body breadth did not increase as fast as
stature during this time period. In fact, just such a secular trend was found among
Japanese-Americans living in Hawaii (Froehlich 1970) as well as in other popu-
lations with secular increases in stature (see Ruff 1994 for a discussion). Thus,
the inclusion of a body breadth measurement in the Japanese analysis could have
provided better discrimination of actual soft tissue changes.

CONCLUSIONS

Both body size and body shape vary systematically in modern humans, and did so in
earlier humans as well. Subsequent to the development of full bipedality and prior
to the adoption of food production, the most important environmental influences on
size and shape were likely climate and technological sophistication. Nutritional
and overall health levels may account for an increasing proportion of variation
during the Holocene, including very recent secular trends. Differences in body
form among modern humans are thus likely to result from an interaction between
long-term genetic factors, including climatic adaptation, as well as developmental
environment, including diet and disease. In assessing anthropometric variation
in living populations, it is important to consider the influence of both kinds of
factors in order to distinguish one from the other. Methodologies that incorporate
critical dimensions of body form can shed new light on human adaptation in both
paleontological and neontological contexts.

ACKNOWLEDGMENTS
I thank Paul Jamison and Roger Siervogel for making available individual data
for the Inupiat and Denver Growth Study samples, respectively. Erik Trinkaus
and Trent Holliday collaborated heavily in the original compilation of Pleistocene
Homo body mass data. Thanks also to the various museums and curators who made
available specimens and data records in their care, and to Ben Auerbach for his
Aleut skeletal data. Original data collection was supported in part by the National
Science Foundation and Wenner-Gren Foundation.

The Annual Review of Anthropology is online at http://anthro.annualreviews.org

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