Received: 6 November 2022 | Revised: 15 March 2023 | Accepted: 29 May 2023

DOI: 10.1002/evan.21988

REVIEW ARTICLE

The estimation and evolution of hominin body mass

Christopher B. Ruff1 | Bernard A. Wood2

1
Center for Functional Anatomy and
Evolution, Johns Hopkins University School of Abstract
Medicine, Baltimore, Maryland, USA
Body mass is a critical variable in many hominin evolutionary studies, with implications
2
Center for the Advanced Study of Human
Paleobiology, George Washington University,
for reconstructing relative brain size, diet, locomotion, subsistence strategy, and social
Washington, District of Columbia, USA organization. We review methods that have been proposed for estimating body mass
from true and trace fossils, consider their applicability in different contexts, and the
Correspondence
Christopher B. Ruff, Center for Functional appropriateness of different modern reference samples. Recently developed
Anatomy and Evolution, Johns Hopkins techniques based on a wider range of modern populations hold promise for providing
University School of Medicine, 725 N. Wolfe
St., Baltimore, MD 21205, USA. more accurate estimates in earlier hominins, although uncertainties remain,
Email: cbruff@jhmi.edu particularly in non‐Homo taxa. When these methods are applied to almost 300 Late

Funding information
Leakey Foundation; Wenner‐Gren
Foundation, Grant/Award Number: 6084;
National Science Foundation,
Grant/Award Numbers: SBR‐891955, SBR‐
8919749, BSC‐0642297
Miocene through Late Pleistocene specimens, the resulting body mass estimates fall
within a 25–60 kg range for early non‐Homo taxa, increase in early Homo to about
50–90 kg, then remain constant until the Terminal Pleistocene, when they decline.
KEYWORDS
Australopithecus, body size, Homo, human evolution, ontogeny, scaling, skeleton

1 | INTRODUCTION
The importance of body mass (or weight) as a fundamental biological
1–5
trait of organisms has been emphasized by many authors. Variation
in body mass affects almost all other aspects of an animal's morphology
and physiology, including the mechanical strength of its bones for
support and movement, neurological requirements for control of the
body, and food‐processing capabilities to supply metabolic needs. Thus,
analyses of characteristics as disparate as locomotor behavior, relative
brain size (or encephalization), and dietary adaptations must take body
mass into account. Furthermore, many critical ecological and life history
variables, such as home range size, life span, reproductive rate, and so
forth are highly correlated with body mass.2,6,7
Given its significance in so many biological and behavioral contexts,
body mass as a variable has taken on increasing importance in
8–27
paleoanthropological research over the past several decades.
Because body mass cannot be measured directly in past individuals, it
must be estimated indirectly from their preserved remains. A variety of
techniques have been proposed to do this, almost all of them based on
skeletal dimensions, but other approaches, such as use of footprint
size,25 are also possible. Here we review and critically examine these
techniques, noting their inherent assumptions, advantages, and limita-
tions, and provide a number of formulae for body mass estimation that
are applicable to human archaeological and paleontological specimens.
We end by presenting a summary of temporal trends in body mass in
the hominin lineage, from the late Miocene through living humans.
2 | HOW T O E STIMATE B ODY M ASS
2.1 | Morphometric versus mechanical
Methods of body size estimation from skeletal remains can be divided
into two basic approaches: morphometric, in which the body is actually
reconstructed from the available material, and mechanical, in which a
functional relationship between a particular skeletal feature and body

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2 |
mass is used to develop a prediction equation.28 An example of the first
approach is the so‐called “Fully” or “anatomical” technique for
estimating stature, in which the heights or lengths of skeletal elements
from the foot to the head are summed, along with an addition for soft
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RUFF and WOOD
tissue.29,30 For estimating body mass, an analogous method using
estimated stature and bi‐iliac (maximum pelvic) breadth has been
developed31,32 (see Figure 1). These two dimensions encapsulate much
of the variation in body size and shape present among living and earlier
hominins.31,33,34 When combined, stature and bi‐iliac breadth can
predict body mass with an average absolute prediction error of less than
10% in healthy, nonoverweight individuals.20 Because of differences in
body shape between males and females (shoulder to hip breadth
proportions), separate equations are used for the two sexes, or, if sex is
uncertain, an average of male and female estimates is used. A formula to
convert skeletal to living bi‐iliac breadth is available for application in
skeletal samples (see Table 1).20,21 Equations have been developed
based on a worldwide sample (see below), and have been applied in a
variety of archaeological and paleontological contexts.38–41
Another type of morphometric method for estimating body mass
that has not been widely used in human evolutionary studies is based on
so‐called “volumetric” reconstruction of the body performed by digitally
fitting a surface over an articulated skeleton.42 When this method was
applied to A.L. 288‐1 (“Lucy”), one of the most complete australopith
skeletons known, it resulted in a very low body mass estimate compared
to other methods.43 Possible factors contributing to this result included
problems with the cadaveric reference sample used to generate the
predictive equations and inaccuracies in reconstructing the full skeleton
of A.L. 288‐1. Body masses of a test sample of macaques and modern
humans were also generally underestimated, sometimes considerably.
More testing of the method on additional primate skeletons of known
body mass is needed to establish its reliability.44
Mechanical methods rely on predicted and empirically observed
mechanical relationships between skeletal dimensions and body mass.
The most reliable dimensions are those that reflect the size of structures
involved in supporting body mass (e.g., lower limb joint size). Two such
dimensions are the superoinferior breadth of the femoral head and
mediolateral breadth of the proximal articular surface of the tibia (see
Figure 1). These are relatively straightforward to measure and are
preserved in many early (as well as later) hominins.20 Regression
equations including these dimensions have been developed from a
number of modern reference samples,9,16,45–47 but the most recent and
comprehensive samples include many more individuals representing a
greater variety of body sizes and shapes,20,36 and are used here.
Because it has been shown that early, non‐Homo hominins (i.e.,
australopiths) have relatively small femoral heads,16,20 possibly due to
differences in gait biomechanics relative to modern humans,48,49 an
adjustment for this proportional difference is made first for such taxa
before applying the femoral head formula20 (see Table 1). The possibility
F I G U R E 1 Modern human skeleton showing dimensions used in
estimating body mass. Red arrows: stature and bi‐iliac breadth. Green
arrows: long bone articular breadths, including femoral head
superoinferior breadth, proximal tibial mediolateral breadth, humeral
head superoinferior breadth, and distal humeral articular mediolateral
breadth. Purple: femoral midshaft cortical area (which may be
combined with femoral maximum length, not shown). Blue: maximum
foot length and forefoot breadth, measured on footprints.
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RUFF and WOOD | 3

TABLE 1 Body mass estimation equations for adults.

Region Predictor(s)a Sex Equationb r %SEEc %PEd Sourcee

Worldwide ST, BI Male BM = 0.422(ST) + 3.126(BI) − 92.9 0.913 6.2 4.7 (1)

Female BM = 0.504(ST) + 1.804(BI) − 72.6 0.819 7.4 5.5 (1)

Europe f
FHSI Male BM = 2.800(FHSI) − 66.7 0.838 10.1 8.0 (2)

Female BM = 2.180(FHSI) – 35.8 0.671 7.8 6.1 (2)

Combined BM = 2.295(FHSI) − 41.7 0.793 9.0 7.2 (2)

TPML Male BM = 1.838(TPML) − 70.4 0.566 11.1 9.0 (3)

Female BM = 1.406(TPML) − 37.8 0.548 9.2 7.2 (3)

Combined BM = 1.564(TPML) − 49.2 0.736 10.2 8.3 (3)

Worldwide FHSIg Combined BM = 2.262(FHSI) − 38.7 0.831 8.7 7.2 (4)

TPML Combined BM = 1.623(TPML) − 52.7 0.755 9.7 8.4 (4)

HHSIh Combined BM = 2.067(HHSI) − 28.5 0.789 10.3 7.8 (5)

HDMLi Combined BM = 2.124(HDML) − 27.6 0.760 10.5 8.4 (5)

Worldwidej FCA Combined BM = 0.0797(FCA) + 27.8 0.782 10.9 8.5 (6)

FCA, FL Combined BM = 0.0448(FCA) + 0.1136(FL) − 5.3 0.823 9.3 7.2 (6)

Worldwide k
FT Combined BM = −0.250(FT) + 0.00099(FT ) + 59.1
2
0.861 13.7 10.1 (7)

FT, BMI Combined BM = 0.155(FT) + 2.48(BMI) − 32.9 0.965 7.0 5.4 (7)
a
ST: stature (cm); BI: living bi‐iliac breadth (cm) (living bi‐iliac = skeletal bi‐iliac × 1.17–3, except when skeletal bi‐iliac <24 cm, where living = skeletal + 1.0);
FHSI: femoral head superoinferior breadth (mm); TPML: tibial plateau mediolateral breadth (mm); HHSI: humeral head superoinferior breadth (mm);
HDML: humeral distal articular mediolateral breadth (mm); FCA: femoral midshaft cortical area (mm2); FL: femoral maximum length (mm); FT: footprint
area (length × breadth) (cm2); BMI: body mass index (body mass in kg/(stature in m)2). Femoral midshaft defined relative to femoral length' dimension.99
b
BM: body mass in kg.
c
%SEE: percent standard error of estimate [(standard error of estimate of y/mean y) × 100)].
d
%PE: absolute value of percent prediction error [|(observed y – predicted y)/predicted y) × 100|].
e
1: Ruff et al.32; 2: Ruff et al.36; 3: calculated from data in Ruff et al.36; 4: Ruff et al.20; 5: Ruff et al.24; 6: Ruff et al.37; 7: Ruff et al.25 (normal walking
condition).
f
Equations applicable to anatomically modern Homo sapiens from Europe.
g
For non‐Homo specimens, adjust FHSI: FHSI = 1.252(FHSI) – 5.7.
h
Applicable to Homo and Australopithecus sediba. For archaic Middle and Late Pleistocene Homo, adjust HHSI: HHSI = HHSI × (HHAP/HHSI)/0.935 ,
where HHAP is anteroposterior breadth of humeral head.
i
Applicable to Homo and Australopithecus sediba. For Late Pleistocene archaic Homo, multiply body mass by 1.14; for Middle/Late Pleistocene anatomically
modern H. sapiens, multiply body mass by 1.08.
j
Applicable to non‐habilis (sensu stricto) Pleistocene Homo.
k
Applicable to non‐habilis (sensu stricto) Homo. A different equation, based on the proportions of the Australopithecus afarensis A.L. 288‐1 skeleton
(“Lucy”) is recommended for A. afarensis: body mass = 0.218 × FT.

that different early hominin taxa may have exhibited somewhat
different scaling relationships between articular size and body mass,
due in part to variation in locomotor behavior, must also be considered,
however. An example of this as applied to upper limb bone articulations
is given below and in Supporting Information: Text S1. Juveniles have
different articular‐to‐body mass proportions than adults, so different
50
formulae must be applied to them (also see Section 2.5).
Lower limb diaphyseal (shaft) cross‐sectional dimensions have also
9,14–16,19
been used to estimate body mass, which is theoretically
plausible because these dimensions are also involved in support of body
mass. A potential complication with this approach, however, is the
developmental plasticity of long bone diaphyses, which respond to
mechanical loadings (forces) due to activity level and muscularity more
readily than articular surfaces.46,51–53 There is evidence that such
factors varied systematically between modern humans and earlier
hominins,48,54,55 which could bias body mass estimates if a modern
human reference sample were used to develop equations. In an attempt
to minimize such biases, a method has recently been developed that
uses a Pleistocene Homo sample with body masses estimated from
articular size to derive new equations based on femoral diaphyseal
cross‐sectional area and bone length.37 The equations are applicable to
other Pleistocene Homo specimens, but not to non‐Homo specimens.
Modern human upper limbs normally do not support body mass,
yet some modern human upper limb bone dimensions are still
correlated with body mass, albeit to a lesser degree than the lower
limb.24,56,57 Body mass estimation equations based on proximal and

4 |
distal humeral articular breadth dimensions in a large modern human
sample have been developed (see Figure 1).24 As might be expected,
though, because of their less tight association with body mass, there
are more uncertainties in applying these equations to earlier hominins
than with lower limb bone articular equations. In particular, the
equations cannot be applied to taxa that show evidence for different
upper limb bone‐to‐body mass proportions. These include all
australopith taxa for which data are available (except Australopithecus
sediba),24 and probably also Homo habilis (sensu stricto).58 For some of
these taxa, formulae based on chimpanzees, or a mean of chimpan-
24
zees and modern humans, appear to be more appropriate. Here we
take a conservative approach and use a mean of modern human and
chimpanzee‐based humeral estimates for these taxa (see Section 3
and Supporting Information: Text S1). Even in later Homo, some
adjustments should be made to account for the differently shaped
humeral head in archaic Middle and Late Pleistocene Homo, and
slightly but systematically different distal humeral to body mass
proportions in Late Pleistocene archaic and anatomically modern
Homo compared to very recent humans.24 These adjustments are
shown in Table 1 and discussed in more detail in Ruff et al.24
Morphometric approaches to body mass estimation have the
advantage that actual body size and shape are estimated directly,
rather than inferred from functional relationships that may vary, as
illustrated by the complexities in applying upper limb bone mechani-
cal equations. However, they have the disadvantage that much more
of the skeleton must be preserved to apply them. As a result, only a
very few early hominin specimens have been analyzed in this way,
and even among more recent archaeological samples such specimens
are relatively uncommon (although see Ruff et al.20,36). Mechanical
techniques are much more widely applicable, and when used with the
appropriate reference samples, they can provide relatively accurate
estimates.
Finally, preserved footprints in hominin trackways offer a
completely different approach to body mass estimation. Hominin
footprint records, which are available worldwide and from time
periods from the Pliocene to the present,59,60 provide unique
opportunities to sample groups of individuals,61–63 and potentially
carry out demographic analyses. In upright bipedal hominins, feet are
the sole contact with the ground during standing, walking, and
running, and are thus critical in weight support. A recent study of 143
unshod or minimally shod individuals from a variety of modern
populations demonstrated a high correlation between foot “area”
25
(maximum foot length × foot breadth) and body mass. However, the
relationship varied by body shape, with relatively heavier populations
(evaluated using the body mass index, BMI = body mass in kg/(stature
in m)2) having relatively smaller feet. Thus, if body shape can be
assessed in the target individual or sample, body mass estimates from
footprint size can be improved. Juveniles have relatively large feet, so
special procedures must be used to estimate their body masses. Body
masses estimated from footprints are similar to those estimated from
skeletal remains in taxonomic/geographic groupings of Pleistocene
Homo.25 However, equations based on modern humans overestimate
body mass in Australopithecus afarensis, due to the latter's relatively
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RUFF and WOOD
larger feet. Thus, a different technique based on the ratio of body
mass to foot size in the most complete A. afarensis, A.L. 288‐1
(“Lucy”), is more appropriate for this taxon.25
2.2 | Which reference sample?
All body mass estimation techniques begin with identifying an
appropriate reference (or “training”) sample, from which the
methodology to calculate body mass from available dimensions is
developed. Because such dimensions are almost always skeletal (with
the exception of footprints), this involves finding reference samples
with associated data for both skeletal dimensions and body mass.
One option employed by a number of past researchers is to use
osteological collections derived from cadaveric material, where body
weights were taken before dissection.8,9,11,15,16,19,45 Unfortunately, it
is very difficult to control for factors such as wasting before death,
the different treatments of bodies after death, and differences in the
time elapsed between death and when the body is weighed in such
samples. A recent comparison indicated that in one sample true living
body masses were probably 21%–29% higher than cadaveric body
masses, with large random as well as directional prediction errors
when using skeletal dimensions to estimate body mass.20 Another
option is to use skeletal dimensions derived from medical imaging
(radiography, computed tomography), or anthropometric measure-
ments (e.g., stature, bi‐iliac breadth) in living subjects with known
body masses.46,47,64–67 A common issue in this case is not under-
estimation of body mass, as with cadaveric samples, but rather
overestimation, due to the generally overweight condition of most
recent Western industrialized populations.68 Thus, equations devel-
oped from these samples may produce upwardly biased estimates,
unless they are restricted to nonoverweight individuals.20,47
Both cadaveric and living reference samples used to develop
body mass estimation equations commonly suffer from another
important limitation: they often incorporate only a limited range of
variation in body size and shape. Most such samples have been
derived from populations in the United States or Europe, or if they
include other regions of the world, are based on only a few
individuals or sex/population means.20 There is abundant evidence
that both body size and body shape varied greatly among hominins
over the past 6 million years.20,34,69,70 Some of this variation may be
attributable to climatic adaptation26,34,37,40,71,72 and some to broader
temporal/phylogenetic trends12,20,22,23,70,73 (see Section 3), but in
any event it argues for inclusion of as broad a range of body sizes and
shapes as possible in any modern reference sample to avoid undue
extrapolation when methods are applied to earlier hominins.
One body mass estimation method that does include a wide range of
body forms in the reference sample is the morphometric stature/bi‐iliac
breadth technique described in the preceding section. This method is
based on 58 sex/population mean dimensions, measured anthropometri-
cally, with a worldwide distribution, ranging from small‐bodied “Pygmies,”
including African Mbuti, Aka, and Kivu Twa, to large‐bodied European and
arctic populations.32,34 Body mass ranges from 38.2 to 82.9 kg and

RUFF and WOOD
stature from 137.3 to 182.1 cm. Both narrow and wide‐bodied
populations are represented, with bi‐iliac breadth ranging from 23.4 to
29.8 cm. These ranges encompass the great majority of specimens in the
archaeological and paleontological record, the primary exceptions being
some very small‐bodied australopiths (such as A.L. 288‐1).20,22,34
Furthermore, for the most part the reference data were collected before
the last 50 years, on younger adults, avoiding recent increases in relative
body weight in many parts of the world74 as well as increases occurring in
middle age.75 When tested, the equations work well on average in both
elite athletes as well as “normal” nonoverweight subjects.20,31 Thus, they
should be broadly applicable to younger, healthy adults in the past as well.
The sex‐specific equations are given in Table 1.
As noted previously, though, relatively few archaeological and
paleontological specimens are complete enough to apply the stature/bi‐
iliac method directly. Mechanical methods that rely only on measurement
of long bone dimensions are much more widely applicable, but they
require appropriate reference samples with recorded body masses in
which the skeletal dimensions can be measured, which have the
limitations discussed above. To avoid these limitations, a “hybrid”
approach that combines morphometric and mechanical methods can be
employed, where body mass in a skeletal sample is first estimated from
the stature/bi‐iliac method, and then used to derive body mass prediction
equations based on other skeletal dimensions in the same sample. Since
many recent skeletal samples are available, this allows inclusion of a much
wider range of populations in the reference sample.
Two reference samples have been used in such analyses: a
36
Holocene European sample (n = 1145) and a Holocene worldwide
20,24
sample (n = 522–682). In each case individual values, not sex/
population means, were used as data points. The worldwide sample is
smaller because more stringent requirements were applied for
inclusion of specimens.20 It incorporates indigenous populations
from Europe, eastern and southern Africa, the southwestern US, and
US Euroamericans. Body mass estimation equations were derived
based on long bone articular breadths (Figure 1). Both sets of
equations have been applied to a variety of target populations or
individuals.20,76–82 The European equations are more appropriate
specifically for recent (Holocene) European samples, while the
worldwide equations can be applied to any hominin sample or
individual, with a correction for femoral head size in non‐Homo
specimens, as described earlier. The formulae are listed in Table 1,
including a new equation for the European reference sample based
on proximal tibial mediolateral articular breadth.
Because long bone cross‐sectional diaphyseal dimensions are
highly developmentally plastic, responding to differences in activity
level and muscular forces, and there is evidence for systematic
variation in such factors between modern and earlier Homo, a special
reference sample restricted to Pleistocene Homo specimens was used
to generate body mass prediction equations from these dimen-
sions.37 Femoral head size was used to estimate body mass in 69
Pleistocene specimens, and these values were then used to generate
body mass estimation equations based on femoral midshaft cortical
area, and on cortical area combined with maximum bone length
(Table 1). The equations are applicable to other Pleistocene Homo
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| 5
specimens, but not to non‐Homo specimens, which have larger
diaphyseal cross sections relative to body size than in Homo.48,83
Other skeletal dimensions, including vertebral body
breadths,11,13,14,16 breadths of the articular surfaces of the tibiotalar
(ankle) joint,11,16,84 and craniofacial dimensions,85,86 have also been
used to estimate body in fossil hominins. However, in all cases, the
human reference samples were cadaveric and/or included only a very
limited sampling of modern human morphological variation. Some
also included only species or sex/species means in broader
comparative analyses, rather than individual data, precluding assess-
ment of true individual‐level prediction errors. The potential
problems involved in using cranial dimensions to estimate body
mass—changing cranial/postcranial and intracranial proportions dur-
ing hominin evolution—have been discussed previously.85
To develop body mass estimation equations based on footprint
size, a special reference sample of habitually unshod (barefoot) or
minimally shod modern humans was used, because most of the target
samples were also habitually unshod and shoe wearing affects foot
dimensions.25 A number of geographically dispersed populations
from North and South America, Africa, and the Pacific Islands
(Samoa), exhibiting a large range of body sizes and shapes, were
included in the reference sample. This contrasted with earlier
attempts to develop such equations,87 which were limited geograph-
ically and/or ethnically. The importance of including a wider range of
body forms became apparent when footprint size relative to body
mass was found to be dependent in part on body shape, with
relatively wider‐bodied populations having relatively smaller feet.25
Thus, earlier equations produced unrealistic body mass estimates for
hominins that did not match the body shapes of the particular
reference sample. Equations were developed from anthropometric
measurements of foot size in the living reference sample, with
conversion factors for converting footprint dimensions to foot
dimensions provided. (For foot/footprint area in “normal walking”
conditions, no conversion is necessary, i.e., footprint area = foot area.)
A quadratic equation and a linear multivariate equation that includes
a body shape (BMI) variable are given in Table 1. Use of the latter
equation is recommended whenever possible, with BMI estimated
from related populations or individuals (see Ruff et al.25 for
examples). An unshod modern juvenile reference sample was used
to develop correction factors for the footprints of likely juveniles
(also see Section 2.5). Other comparisons between habitually unshod
and shod individuals within a single population provided a correction
factor for individuals who likely used footwear.25 As noted earlier,
these equations are only appropriate for Homo footprints; a different
equation for A. afarensis footprints is given in the Table 1 footnote.
2.3 | Statistical issues and general
recommendations
The information provided in Table 1 also raises several general
statistical issues. The equations based on articular dimensions were
all generated using Reduced Major Axis (RMA) regression rather than

6 |
Least Squares (LS) regression. This is because, as noted earlier, some
of the target specimens (e.g., small‐bodied australopiths) fall outside
the range of the reference sample, and in this situation RMA provides
more accurate estimates than LS.50,88 Two different measures of
predictive precision—the percent standard error of estimate (%SEE)
and absolute percent prediction error (%PE)—are given for each
equation. These measure the degree of random error around
estimates in relative terms. For example, the average %PE for a
body mass estimate of 60 kg derived using the worldwide sample
femoral head equation in Table 1 is ±4.8 kg (8.0%). The %SEE and %
PE can be used to help choose between equations based on their
predictive precision. Note that the correlation coefficient is an
imperfect indicator in this regard: because the correlation coefficient
is also strongly affected by factors such as sample size and range of x
and y values. a higher correlation does not necessarily equate with a
lower %SEE and %PE.89 All of the equations in Table 1 have average
%PEs of 10% or less, which is better than many equations reported
for intraspecies and interspecies reference groups.90–92 Among
articular dimensions, the femoral head produces the lowest predic-
tion errors, followed by the proximal tibia; this is not surprising, given
that they are both weight‐bearing joints. Combining femoral length
with femoral midshaft cortical area, which is equivalent to combining
morphometric and mechanically‐based approaches, improves predic-
tion over cortical area alone. The very low apparent prediction errors
for the stature/bi‐iliac and footprint/BMI equations (%PEs of around
5%) are almost certainly underestimates of true errors in individual
estimates. The stature/bi‐iliac reference sample consists of sex/
populations means, and variation among individuals would be
expected to be greater. However, tests of these equations among
samples of individuals of known body mass have generally produced
average %PEs of less than 10%,20 so in this respect they are still
comparable to the other equations in Table 1. The footprint equation
that includes a measure of body shape, BMI, assumes that BMI is
actually known for the target individual, whereas in practice it must
be estimated itself from other information, thus increasing overall
error. However, this technique is still recommended over the
equation that includes just footprint area, for reasons that are
discussed in detail in the original publication.25 All of the equations in
Table 1 are only applicable to adults, since juveniles have different
body proportions (see Section 2.5).
Finally, it should be emphasized that the measures of predictive
power in Table 1 are purely statistical in nature. True accuracy of the
equations is dependent not only on the strength of the relationship
between a predictor variable and body mass in the reference sample
but also to what extent the same relationship applies in the target
sample. For this reason, several of the equations have limitations in
terms of appropriate target specimens, and in other cases, either the
skeletal predictor variable or body mass estimate should be adjusted
to reflect proportional differences from the reference sample, as
noted in the table footnotes.
In summary, the stature/bi‐iliac method for estimating body mass
makes the fewest assumptions about body shape and skeletal
proportions, and thus should be used if sufficient skeletal material
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RUFF and WOOD
is available. Among “mechanical” techniques, those based on femoral
head and proximal tibial breadths will provide the most accurate body
mass estimates, although femoral head breadth should be adjusted in
non‐Homo taxa. The “Europe” equations are most appropriate for
Holocene Europeans, while the “Worldwide” equations can be
applied to any area of the world. In specimens that do not preserve
the femoral head or proximal tibia, humeral articular equations can be
used, but come with more caveats and are generally not applicable to
non‐Homo specimens. In the absence of any of these articulations,
body mass in Pleistocene Homo specimens can be estimated from
femoral midshaft cortical area, combined with femoral length,
although results should be viewed with more caution because of
possible confounding behavioral factors. Footprints are effective as
body mass predictors, especially when it is possible to incorporate
information regarding body shape (BMI).
2.4 | The perils of proxies
An alternative to estimating body mass directly is to use skeletal
proxies such as long bone lengths and breadths to investigate
intrataxon or intertaxon variation in body size.8,93–96 This approach
has the advantage that no dimensions are estimated, and the various
issues in estimating body mass noted above are avoided. However,
use of proxies rather than body mass itself has some significant
disadvantages. Most early hominin skeletal remains are fragmentary,
preserving different regions of the skeleton; thus, the same
dimensions cannot always be compared across specimens assigned
to the same, or a different taxon. However, using appropriate
equations, body mass can be estimated from a number of different
dimensions (Table 1), and can thus serve as a “common denominator”
for comparisons across individuals represented by different combi-
nations of elements (for example, see Section 3). Body mass is also
the most common “size” variable measured in ecological and
physiological studies of living species (i.e., you can more conveniently
weigh a living animal than measure the size of its femoral head); thus,
body mass estimates provide access to a much richer comparative
context for interpreting variation among fossil taxa than is available
for individual skeletal dimensions.
But perhaps the most important limitation of skeletal proxies is
that they assume similar proportionality across, and within, samples
between the particular skeletal dimension(s) employed and body
mass, which is often not the case. There are two factors that should
be considered here: overall differences in skeletal/body mass
proportions between groups, and changes in proportions within
groups that are correlated with body size. Both of these phenomena
are part of a broader field of study referred to as allometry, or scaling,
which examines how body size affects morphology.1,5
An illustration of how these factors can affect interpretations of
body size variation is provided by comparisons of femoral head size
scaling with body mass in anthropoid primates. Femoral head breadth
(or acetabular breadth) is one of the most commonly employed
proxies for body mass in comparative studies.8,35,93–97 Figure 2 is a

RUFF and WOOD
F I G U R E 2 Scaling of body mass on femoral head breadth in
modern humans, nonhuman great apes (Gorilla, Pan, Pongo), and
baboons (Papio). All data natural log‐transformed. Regression lines fit
through the three groups using Reduced Major Axis regression.
plot of body mass relative to femoral head breadth in 682 Holocene
modern humans, 38 nonhuman great apes (Pan, Gorilla, and Pongo,
hereafter “great apes”), and 20 baboons (Papio cynocephalus). All
individuals are adults, and all of the nonhuman specimens were from
wild populations. Body masses were directly measured in the
nonhuman taxa90,98 and were estimated from stature and bi‐iliac
breadth in the human sample.20 Data were logarithmically trans-
formed, allowing easier assessment of proportional changes using
linear regression. In this type of analysis, line elevations reflect overall
proportions of a group, while slopes indicate changes (or not) in
proportions over a size range within a group. All lines were fit with
69
RMA regression, appropriate for investigating scaling relationships.
Because body mass can be assumed to be proportional to body
volume, and body volume is proportional to linear dimensions to the
third power, isometry, or geometric similarity, is indicated here by a
slope of 3.0 in logarithmic space (i.e., body mass is proportional to
3
femoral head breadth ).
It is apparent from Figure 2 that: (a) the three great apes have the
same scaling relationship between femoral head breadth and body
mass and (b) modern humans, great apes, and baboons have different
scaling relationships between femoral head size and body mass.
Modern humans have the relatively largest femoral heads, adaptive
for their bipedal locomotion where all of body mass is supported on
the lower limbs. Both great apes and baboons are quadrupedal and
have relatively smaller femoral heads than modern humans, but
femoral head size is relatively larger in the great apes than in
baboons. The latter is likely related to great apes' style of vertical
climbing, involving increased hip mobility, which is facilitated by a
relatively larger femoral head.92,99,100 In addition to these general
proportional differences, the slopes of the lines through the three
groups are significantly different. Great apes are isometric, with a line
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| 7
slope very close to 3.0. In contrast, the slope of the line through
modern humans is highly nonisometric, with femoral head breadth
increasing much faster than body mass. Baboons show the opposite
pattern, with femoral head size increasing more slowly than body
mass. Thus, large and small modern humans, or large and small
baboons, will show different proportions of body mass to femoral
head size purely as a result of general scaling trends within these taxa
(the underlying functional explanations of these different scaling
trends have been discussed elsewhere).20 This also means that
differences in general proportions between taxa will be in part
dependent on the size range considered; for example, small modern
humans have proportions more similar to those of great apes than do
large modern humans.
Thus, it is clear that using femoral head size as a surrogate for
body mass in comparisons both between and within taxonomic
groups will lead to significant bias in many cases, due to different
average proportions between groups and nonisometric scaling within
some groups. (For further discussion of the dangers inherent in using
other proxies for body size, see Hill et al.101 and Smith102). This is true
for comparisons between nonhuman anthropoid primates as well as
between modern humans and nonhuman primates, and even within
taxonomically/functionally similar groups, such as large and small
modern humans. Furthermore, scaling trends differ from one part of
the skeleton to another (e.g., in modern humans knee joint breadth
scales differently than femoral head breadth),20 so trends in one
feature cannot be extrapolated to other skeletal regions. The results
shown in Figure 2 also argue against combining groups such as
modern humans and nonhuman hominoids to develop taxonomically
broader body mass estimation equations.11,15,16 Fitting a regression
line through the combined great ape‐modern human sample in
Figure 2 would result in an uninterpretable scaling relationship not
observed in any modern taxon. Rather, if a morphologically and
functionally intermediate proportion is hypothesized, as for example
in some early hominins that combined terrestrial bipedality with some
degree of arboreality,48,103 then equations for different groups can
be applied and an average or range of estimates provided. This is the
approach taken here for estimating body mass from upper limb bone
dimensions in non‐Homo taxa (see Section 3).
2.5 | Teenagers are all knees and elbows
Juvenile postcranial specimens can provide critical information on the
timing of developmental events and the relative importance of
genetic and environmental effects on adult morphology.50,104–109
Body size is a potentially important component of such analyses, and
yet until recently there were no equations available for estimating
body mass from juvenile skeletal remains.50,110,111 Consequently,
earlier studies used indirect methods for assessing body size in
juveniles, such as weight for height relationships or body (pelvic)
breadth dimensions.107,112 Even today, body mass estimation
equations for juvenile skeletal specimens are available from just
one reference sample—the Denver Growth Study sample,50,110,111

8 |
which is limited both ethnically and environmentally.113 Given that
body mass estimation equations are available for adults from a much
wider range of populations, as discussed above, it is reasonable to ask
whether such equations might be applied to juveniles as well, thus
broadening the distribution of body sizes and shapes represented.
This question was addressed in an investigation of scaling relation-
ships between skeletal dimensions and body mass in the Denver
Growth Study sample.50 It was found that during ontogeny both
femoral length and femoral head breadth “grow ahead” of body mass in
late childhood and early adolescence, that is, they approach young adult
values earlier, and are thus larger relative to body mass than in adults. As
a consequence, body mass estimation equations based on adults
overestimate body mass in juveniles, particularly in this age range. This is
illustrated in Figure 3, which plots the percent difference between body
mass predicted from femoral head breadth using an adult equation (the
worldwide equation in Table 1) and true body mass for 20 individuals in
the Denver Growth Study sample followed longitudinally from 6 to 20
years of age (there are fewer points in the oldest age groups because of
subject dropout from the study). The maximum deviation between
values occurs between about 9 and 13 years of age, averaging 20% or
more, but by late adolescence (18–20 years) there is no average bias in
estimates, as individuals reach adult proportions.
The effects of using an inappropriate adult reference sample for
a fossil juvenile can be illustrated by applying adult and age‐matched
formulae to KNM‐WT 15000, an early African Homo erectus male
whose skeletal maturity was equivalent to that of a modern 11‐13‐
F I G U R E 3 Percent difference between body mass predicted
from femoral head breadth using an adult equation and true body
mass in a modern human juvenile longitudinal sample (extracted from
the Denver Growth Study). Line fit through data using LOWESS
nonparametric regression.114 [Correction added on 19 July 2023,
after first online publication: Minor corrections were made to the y
axis label of Figure 3 and in the Figure 3 caption for clarification.]
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RUFF and WOOD
year‐old boy.50 If age‐appropriate femoral head equations for 11–13
year‐olds in the Denver sample are applied, body mass estimates
between about 50 and 53 kg are obtained.50 If the worldwide adult
femoral head formula is used instead, his estimated body mass is
about 63 kg, 19%–26% higher, and larger than would be expected
given his reconstructed stature of less than 157 cm.20,50 Thus, it is
clear that adult body mass estimation equations based on femoral
head breadth—and likely any long bone articular breadth—should not
be used for juveniles. Furthermore, the amount of adjustment needed
for adult estimates is age‐specific, so no constant correction factor
can be applied. Obtaining data similar to those available for the
Denver Growth Study sample113 for other populations would be
useful for further exploring possible effects of body proportions on
juvenile body mass estimation, although fortunately such effects
appear smaller (in adults) than on stature estimation.28,107,115
Foot size relative to body mass also varies significantly between
juveniles and adults, with juveniles having relatively larger feet.25
Thus, adult equations for estimating body mass from footprint size
cannot be used directly on juvenile footprints. Again, like femoral
head size, the deviation in proportions between juveniles and adults
is not constant, although in this case, the greatest deviation occurs in
early childhood, with a progressive trend towards adult proportions,
which are reached at about 16 years of age. The developmental
difference in proportions is likely due to a combination of earlier
maturation of the foot than most of the rest of the skeleton, and
changes in body shape (increases in BMI) during growth.25 Because
age is very difficult if not impossible to estimate based on footprint
size alone, a technique was developed whereby the size of a juvenile
footprint relative to an average adult from the same population
(footprint sample) is used to calculate a correction factor for body
mass estimated using the adult formulae.25 The difference in foot/
body mass proportions between juveniles and adults appears to be
relatively constant between populations of different body shape, so
the same correction equations should be universally applicable.
When applied to a juvenile Neanderthal sample,61 they produced
reasonable body mass estimates.25
3 | TEM PO RAL TREN DS IN HO MININ
BOD Y MAS S
The methods described in the preceding sections and summarized in
Table 1 were applied to 299 late Miocene (6.0 Ma [million years ago])
to terminal Pleistocene (12,000 BP [years before present]) hominin
postcranial specimens (individuals) from Africa, Europe, and Asia.
When more than one equation was applicable, the worldwide femoral
head and proximal tibial breadth equations were favored because of
their low prediction errors and more direct functional association
with body mass (stature/bi‐iliac equations were very rarely applicable
and were not used here). Femoral head breadths were adjusted for all
non‐Homo taxa before entry into the equation, as described in
Table 1. Equations based on humeral articular dimensions were used
for Homo, with some adjustments for particular groups as described
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RUFF and WOOD | 9

in Table 1. For non‐Homo taxa estimated using humeral articular
dimensions, a mean of body mass estimates derived from modern
human and chimpanzee reference groups was used, with chimpanzee
equations derived from reference.90 Supporting Information: Text S1
gives more details. In addition to these skeletal samples, 102 sex/
population means for living humans distributed among the same
three geographical regions were included for comparison. All
individual data are given in Supporting Information: Table S1. Subsets
of these data have been published previously.20,24,37
A plot of individual body mass estimates against time in shown in
Figure 4; a dual time scale is used to better differentiate data points
after 200,000 BP. Non‐Homo are plotted in red and Homo specimens
in blue, with specimens of indeterminate or controversial taxonomic
20,24,37
status in purple. Taxonomy follows previous studies, supple-
mented by recent analyses of humeral morphology116,117 and general
postcranial variation among early hominins.118 However, it must be
recognized that most early hominin taxa are identified and diagnosed
on the basis of cranial and not postcranial morphology.118 There is
also the possibility that taxa that are cranially and dentally distinctive
may not necessarily be distinguishable on the basis of postcranial
evidence. Several researchers have recently provided helpful reviews
of the latter,117,118 and, where possible, we have used these to
allocate specimens to taxa. Early (>1.7 Ma) Homo not assigned to H.
habilis (sensu stricto) are simply designated “early Homo,” and
nonanatomically modern Middle and Late Pleistocene Homo, except
for Homo naledi and Homo floresiensis, are assigned to “archaic Homo.”
Individuals assigned to non‐Homo taxa (Orrorin, Ardipithecus,
Australopithecus, Paranthropus) vary in body mass from about
25–60 kg, and show no evidence of any systematic directional
change in body mass between 6.0 and 1.0 Ma. The average body
mass of these taxa over this time period is 43.8 kg (±10.3 SD). Three
H. habilis (s.s.) specimens, dating to 1.9–2.0 Ma, have a similar mean
body mass of 42.2 kg (range 41–45 kg). Thus, as argued previ-
ously,119,120 in terms of body size, there is no distinction between H.
habilis (s.s.) and non‐Homo taxa.
In contrast, non‐habilis early Homo (1.8–2.0 Ma) is somewhat
larger‐bodied, with four specimens averaging 58.6 kg (range
52–67 kg). By 1.4–1.6 Ma there is a substantial increase in body
mass in H. erectus/ergaster, with six specimens averaging 75.0 kg
(range 65–90 kg). Following this, average body mass in Homo remains
relatively constant, with individual specimens ranging between about
50 and 90 kg, until the Terminal Pleistocene (post‐25,000 BP), when it
declines. There are three exceptions to this general trend, however.
The first is the specimen from Gona, Ethiopia (BSN49/P27), assigned
by its discoverers to H. erectus.121 The taxonomic status of this
specimen has been questioned, however, based largely on its very

F I G U R E 4 Body mass trends in hominins from the late Miocene to living humans. Note change in temporal scale at 0.2 Ma (200,000 BP). Red
symbols: non‐Homo; blue symbols: Homo; purple symbols: contested or indeterminate taxonomy. Other subdivisions of the sample are given in
the symbol key (A.: Australopithecus; P: Paranthropus; H.: Homo). “Other non‐Homo” includes Orrorin tugenensis, Ardipithecus ramidus,
Australopithecus anamensis (all labeled), Paranthropus aethiopicus, Australopithecus prometheus, and Australopithecus sp. (unlabeled). Early African
H. erectus includes specimens sometimes attributed to H. ergaster. Unlabeled “archaic Homo” includes H. antecessor, H. heidelbergensis, and H.
neanderthalensis (or H. sapiens neanderthalensis). Living humans are sex/population means.

10 | RUFF and WOOD
small estimated body mass (35.1 kg), which is much lower than any
other Early‐Middle Pleistocene Homo specimen known (Figure 4),
122
including all other specimens assigned to H. erectus/ergaster. This
specimen is left taxonomically unassigned in Figure 4, as are four
partial tibiae and one partial humerus without taxonomically
identifiable features. The second exception is the H. naledi sample,
with a mean body mass of 46.2 kg (range 41–67 kg), which is
considerably lower than other broadly contemporary archaic Homo.
The final exception is the tiny H. floresiensis individual, with an
estimated body mass of only 31.4 kg. The H. naledi and H. floresiensis
specimens demonstrate the existence of temporally and geograph-
ically restricted groups of smaller‐bodied Homo in the Middle and
Late Pleistocene. In the case of H. floresiensis its small size has been
73
attributed to island endemism, that is, insular dwarfism.
The decline in average body mass at the end of the Pleistocene is
statistically significant whether or not one includes Late Pleistocene
archaic Homo (i.e., Neanderthals). In fact, Late Pleistocene archaic and
anatomically modern (AM) H. sapiens, before 25,000 BP, are not
significantly different in body mass.20 The average decline within AM
H. sapiens between 40,000 and 12,000 BP is about 6 kg, or 9%. Living
humans are smaller on average than Terminal Pleistocene
(25,000–12,000 BP) AM humans (p < .001, t test), but this is in part
confounded by geographic differences in body size. If comparisons
are carried out within regions (Africa, Europe, Asia), there are no
statistically significant differences in body mass between Terminal
Pleistocene and living humans. The significance of regional differ-
ences in body size within Homo throughout the Pleistocene has been
discussed elsewhere.37
Sexual dimorphism in body mass is another factor influencing the
distribution of values within both taxa and temporal periods. Sexual
dimorphism in body mass is greater within non‐Homo taxa than
within Homo taxa or temporally/geographically defined groups,
20
adding to the dispersion of values among the former. The lowest
levels of sexual dimorphism are found among populations of
anatomically modern H. sapiens, although their greater geographic
distribution adds to overall variability in body mass in H. sapiens, due
34
in part to climatic adaptation. These factors may affect apparent
temporal trends in body mass, particularly where sample sizes within
taxa or temporal periods are relatively small, for example, in early
Homo and some non‐Homo taxa.
Analyses of hominin footprints support the body mass trends
shown in Figure 4.25 Presumed Australopithecus afarensis footprints
63
from Laetoli produce estimated body masses between 31 and
59 kg, similar to the range for this taxon determined from skeletal
remains. Footprints likely made by Homo erectus/ergaster at Ileret,62
dated to about 1.52 Ma, produce body mass estimates that average
about 65 kg, within the range of early Pleistocene archaic Homo
(except H. habilis s.s.). Three adult male presumed Neanderthal
footprints from Le Rozel61 (about 80,000 BP) give average estimated
body masses of 78–90 kg, similar to larger Late Pleistocene archaic
Homo skeletal estimates. Estimates of sexual dimorphism in body
mass in the Laetoli and Ileret footprint samples are also similar to
those calculated from skeletal remains matched taxonomically,
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geographically, and temporally.20,25 This confirmation is significant
because skeletal samples, unlike footprints, are rarely limited to single
populations.
These results are broadly consistent with a number of previous
studies, but there are some important differences as well. Our
conclusion that body mass increased in the earliest members of the
genus Homo that were not H. habilis (s.s.), relative to previous and
contemporary non‐Homo, is in agreement with several stud-
ies,18,22,23,120 but disagrees with others.9,12 A major source of this
discrepancy in results is the use of different body mass estimation
equations based on different modern reference samples, with
equations based on cadaveric samples providing systematically
smaller estimates and less differentiation in body mass between
non‐Homo and early Homo samples.9,12 All previous studies agree,
however, that body mass increased significantly in Homo after
1.7 Ma, although there are some differences in interpretation of
subsequent trends within archaic Homo. One study concluded that
consistently high body masses within Homo were not achieved until
about 0.4–0.5 Ma, in part as a function of climatic adaptation to
higher latitudes.22 In contrast, our results indicate that average body
mass was essentially stable within archaic Homo (with the exception
of H. naledi and H. floresiensis) from 1.6 Ma to near the end of the
Pleistocene. Again, part of the divergence in results is due to
methodological differences, including the use in Will et al.22 of lower
estimates for early Middle Pleistocene (0.77–0.9 Ma) specimens
based on cadaveric‐derived equations.
Fine‐tuning body mass estimations can have important ramifica-
tions regarding body size variation and its consequences throughout
hominin evolution. For example, an increase in body size within Homo
relative to other hominin taxa may have affected important variables
such as home range size, age at maturity, lifespan, subsistence
strategy, and social organization,18,120 so the timing of this transition
is significant. While earliest non‐habilis Homo can be differentiated
from non‐Homo in body size, it is clear that the major increase in body
mass within Homo did not occur until after the initial expansion of
hominins out of Africa, which was before 1.7 Ma.123,124 This has
implications for interpreting both the biological and behavioral
requirements for geographical expansion in Homo as well as patterns
of body size variation throughout the Old World during the
Pleistocene.37,125 The discovery of much smaller‐bodied forms of
Homo later in the Pleistocene (H. naledi and H. floresiensis) raises
interesting questions regarding localized adaptations within the
genus,70,126 including divergent locomotor behavior.127,128 The decline
in body mass at the end of the Pleistocene among anatomically
modern H. sapiens has been attributed to a number of possible causes,
including changes in diet, demographics, and technology.129,130
4 | CONCLUSIONS
Body mass has become an increasingly important component of
many hominin adaptive scenarios. The accuracy of body mass
estimates will be affected by several factors, including the type of

RUFF and WOOD
material available for estimation and the appropriateness of the
reference sample used for developing the estimation equations. In
more complete specimens, a morphometric approach may be applied,
where body mass is estimated from direct reconstruction of the body
using estimated stature and bi‐iliac (maximum pelvic) breadth. In less
complete specimens a mechanical approach based on the relationship
of functionally relevant skeletal dimensions to body mass may be
applied. The most reliable skeletal dimensions are lower limb articular
breadths, especially hip (femoral head) and knee (proximal tibial)
breadths, with an adjustment for the relatively small size of the
femoral head in non‐Homo specimens. Equations using shoulder
(humeral head) and elbow (distal humeral) articular breadths have also
been developed, but they require more adjustments in either
dimensions or resulting body mass estimates depending on the
target sample, and are not applicable to non‐Homo taxa. A technique
based on femoral midshaft cortical area and femoral length can be
applied to specimens that do not preserve articular surfaces, but it is
subject to caveats stemming from behavioral effects on long bone
diaphyses, and again is not applicable to non‐Homo specimens.
Footprint size can also be used to estimate body mass, with better
estimates obtained when body shape is factored in.
In choosing reference samples, a general objective should be to
include individuals as representative as possible of those of the target
sample. For example, in forensic applications, a modern, more
overweight sample may be most appropriate. However, for earlier
hominins, who it can be assumed were relatively physically fit, such a
sample will likely produce overestimates of body mass. Cadaveric
samples, which are subject to wasting and loss of soft tissue, may
have the opposite problem, producing underestimates of body mass
in normal, healthy individuals. Both types of samples generally have a
limited range of variation in body size and shape, while earlier
hominins varied greatly in these characteristics. To address these
issues, a worldwide anthropometric sample of non‐overweight
populations with a wide range of body morphologies was used to
develop body mass estimation equations based on stature and bi‐iliac
(pelvic) breadth. In addition to direct application to relatively
complete target specimens, these body mass estimates were also
used to develop new articular equations based on a wide range of
modern humans. Because cross‐sectional dimensions of long bone
diaphyses are very developmentally sensitive to differences in
behavior, which likely varied between modern and Pleistocene Homo,
a reference sample of Pleistocene Homo with articular‐based body
mass estimates was used to develop estimation equations for other
Pleistocene Homo specimens using femoral cortical area and bone
length. Ontogenetic effects are another example of the importance
of matching reference and target samples: juveniles have different
proportions than adults in both relative articular size and foot size,
necessitating separate formulae based on age‐matched samples.
One general observation that can be drawn from these analyses
is that applicability of equations based on modern humans will
progressively decline when moving further away from modern
humans taxonomically and/or functionally. This is particularly true
for non‐Homo specimens, who are the most distant in both respects.
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| 11
Due to probable use of the upper limb in locomotion (i.e., tree
climbing) in many non‐Homo taxa, increasing upper limb bone
dimensions relative to body size, equations based on upper limb
bone dimensions are not applicable to these taxa.24 Chimpanzee‐
based formulae appear to work better for at least some non‐Homo
taxa24; here, an average between chimpanzee and modern human‐
based estimates was used. However, such an approach obviously
depends on the extent to which locomotion (and body proportions) in
various non‐Homo taxa differed from modern humans, something
that is difficult to determine precisely. Long bone diaphyses (of both
limbs) are also stronger relative to body size in non‐Homo taxa48,83;
so methods based on femoral cortical area, even in Pleistocene Homo,
cannot be applied to these taxa. Australopiths have relatively small
femoral heads,48 so an adjustment to femoral size must be
incorporated into equations based on this dimension. Australopithe-
cus afarensis has relatively large feet,25 necessitating a different
equation for estimating body mass from footprint size. All of these
considerations reflect increased uncertainty of body mass estimates
among non‐Homo taxa. Discovery of additional associated skeletons
with preservation of multiple skeletal regions, such as A.L. 288‐1
(“Lucy”), will help to further address such uncertainties by allowing
comparisons between different kinds of estimates.
The use of skeletal proxies as substitutes for body mass in
comparative analyses is discouraged. Specific skeletal features show
different scaling relationships to body mass in different taxa, and
even within taxa in different body size ranges. Thus, skeletal proxies
have a high probability of introducing errors into comparisons, unless
it can be shown that all units of comparison have the same
proportions. A safer alternative is to actually estimate body mass
using appropriate reference samples, and use that as the body size
parameter.
When recently‐developed body mass estimation equations are
applied to Miocene to recent hominins, some broad patterns emerge.
Body mass in the earliest hominins varies between about 25 and
60 kg, with no consistent temporal trend until the appearance of
Homo, at 1.8–2.0 Ma, which is somewhat larger. This does not apply
to H. habilis (s.s.), however, which remains small‐bodied. Body mass
increases greatly in Homo after 1.7 Ma and remains in the general
range of 50–90 kg until the end of the Pleistocene (12,000–25,000
BP), when it declines slightly to reach modern values. The small‐
bodied Middle Pleistocene H. naledi (41–51 kg) and very small‐bodied
Late Pleistocene H. floresiensis (31 kg) are exceptions. Variation in
hominin body mass has important implications for reconstructing and
interpreting changes in such fundamental characteristics as locomo-
tion, diet, brain size, subsistence strategy, social organization, and
many life history variables. Careful consideration of the assumptions
and limitations of any body mass estimation technique is mandatory
before its application.
ACKNOWLEDGME NT S
Many people over the years have contributed to the collection of
data used in this study. For assistance in collating hominin skeletal
data for Middle and Late Pleistocene specimens, we thank in
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12 | RUFF and WOOD

particular Erik Trinkaus. Please see the original publica- 16. McHenry HM. 1992. Body size and proportions in early hominids.
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RUFF and WOOD | 15

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Bernard A. Wood is University Professor of Human Origins at
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A U T H O R B I O G R A P H IE S SUPP ORTING INFO RM ATION

Additional supporting information can be found online in the
Christopher B. Ruff is Professor Emeritus and former Supporting Information section at the end of this article.
director of the Center for Functional Anatomy and Evolution,
Johns Hopkins University School of Medicine. His research
interests focus on evolution and environmental adaptation of How to cite this article: Ruff CB, Wood BA. The estimation
the hominoid postcranium, skeletal growth and development, and evolution of hominin body mass. Evolutionary
and behavioral reconstruction through biomechanical Anthropology. 2023;1‐15. doi:10.1002/evan.21988
analyses.