The evolution of the human pelvis:
changing adaptations to bipedalism,
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Review
Cite this article: Gruss LT, Schmitt D. 2015
The evolution of the human pelvis: changing
adaptations to bipedalism, obstetrics and
thermoregulation. Phil. Trans. R. Soc. B 370:
20140063.
Downloaded from https://royalsocietypublishing.org/ on 20 April 2022
http://dx.doi.org/10.1098/rstb.2014.0063
One contribution of 13 to a discussion meeting
issue ‘Human evolution: brain, birthweight and
the immune system’.
Subject Areas:
biomechanics, evolution
Keywords:
human evolution, pelvis, bipedalism,
functional anatomy, birth
Author for correspondence:
Laura Tobias Gruss
e-mail: lgruss@radford.edu
obstetrics and thermoregulation
Laura Tobias Gruss1 and Daniel Schmitt2
1
Department of Biology, Radford University, Radford, VA, USA
2
Department of Evolutionary Anthropology, Duke University, Durham, NC, USA
The fossil record of the human pelvis reveals the selective priorities acting on
hominin anatomy at different points in our evolutionary history, during
which mechanical requirements for locomotion, childbirth and thermo-
regulation often conflicted. In our earliest upright ancestors, fundamental
alterations of the pelvis compared with non-human primates facilitated
bipedal walking. Further changes early in hominin evolution produced a
platypelloid birth canal in a pelvis that was wide overall, with flaring ilia.
This pelvic form was maintained over 3–4 Myr with only moderate changes
in response to greater habitat diversity, changes in locomotor behaviour and
increases in brain size. It was not until Homo sapiens evolved in Africa and
the Middle East 200 000 years ago that the narrow anatomically modern
pelvis with a more circular birth canal emerged. This major change appears
to reflect selective pressures for further increases in neonatal brain size and
for a narrow body shape associated with heat dissipation in warm environ-
ments. The advent of the modern birth canal, the shape and alignment of
which require fetal rotation during birth, allowed the earliest members of
our species to deal obstetrically with increases in encephalization while
maintaining a narrow body to meet thermoregulatory demands and
enhance locomotor performance.
1. Introduction
The human pelvis is a remarkable structure that plays a central role in many
critical biological processes, most notably bipedal locomotion, thermoregula-
tion and parturition (childbirth). Each of these processes is essential enough
to survival and reproductive success as to be under strong pressure from natur-
al selection [1– 4]. As a result, the fossil record of the evolution of the human
pelvis over the past 4.5 Myr reveals a profound story concerning selective pri-
orities during different phases of human evolution, and elucidates the essential
constraints that formed our modern anatomical condition.
Pelvic anatomy impacts human performance. To walk upright in an energet-
ically efficient manner with a minimal risk of injury, the pelvis must be robust
and have a shape that maximizes muscle lever arms and minimizes load [5– 11].
The ability to regulate body temperature is affected by the width and depth of
the pelvis, which plays a crucial role in determining overall body proportions
and the body’s surface area-to-mass ratio, thereby influencing heat loss through
the body’s surface [12,13]. Finally, and most critically from a selective stand-
point, pelvic shape must allow the delivery of a healthy infant without harm
to the mother [14]. These goals are not all achieved by the same pelvic
morphology, yet all three demands must be met. Therefore, selection has
favoured compromises between these often contradictory pressures [15– 17]1.
The results of these compromises can be seen in our hominin2 ancestors’
pelvic morphology at every stage of evolution of the lineage, with different
aspects of hominin ecology playing a more or less dominant role in driving
pelvic form at different times. The changes in pelvic anatomy over time in
our lineage appear to reflect changes in locomotion, habitat, climate, brain
& 2015 The Author(s) Published by the Royal Society. All rights reserved.
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2

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3
3

1 1

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H. sapiens A. afarensis Ar. ramidus P. trogodytes

Figure 1. The pelvis in (L to R) modern human (Homo sapiens), early hominins and chimpanzee (Pan troglodytes). In the anterior view (upper), note shortening of
the height of the body (1) and blade (3) of the ilium in Homo compared with Pan, which lowers the centre of mass in the former. The iliac blade is also more
laterally placed and the sacrum is wider in humans (2), which eliminates the entrapment of the lower lumbar vertebrae seen in the chimpanzee (2). In the lateral
view (lower), note especially the coronal orientation of the iliac blade in the chimpanzee compared with the more laterally rotated blade in the hominins, which
allows gluteal muscles that arise from the external surface of the ilium and attach to the greater trochanter of the femur to act as abductors of the thigh and, when
the lower limb is in stance phase, to counteract tipping of the pelvis. In addition note the length of the ischium in Homo compared with Pan. Like chimpanzees, the
ischium in Ar. ramidus is relatively long, indicating a potential mosaic evolution of the pelvis in early hominin evolution. The ischium is the site of attachment for
hamstrings and sacrotuberous ligament; the small triangles indicate the orientation of the ischial tuberosity, which is directed inferiorly in Ar. ramidus and the
chimpanzee but more posteriorly in A. afarensis and modern humans. (Adapted from references [20,21].)

size and subsistence behaviour (behaviours related to the
acquisition of food), and therefore tell the story of human
evolution with remarkable depth.
To consider the role of the pelvis in locomotion, obstetrics
and thermoregulation and the way in which these different
pressures affected pelvic anatomy, we follow the approach
of Lovejoy et al. [20] and divide the pelvis into upper and
lower halves (dashed white line in figure 1). In the upper
half, changes in the height and position of the iliac blades
(alae or wings of the ilia) influence the leverage of muscles
that abduct or extend the hip joint, the range of motion in
the lumbar vertebral column, and the height of the centre
of mass, all of which affect the ability to balance the body
over the legs during a striding bipedal gait. In addition, the
position of the iliac blades affects trunk width and impacts
thermoregulation. But the upper half of the pelvis has a lim-
ited influence on parturition; the proportions of the birth
canal are determined by structures in the lower pelvis.
2. Pelvic morphology in humans and non-
human primates
The overall form of the pelvis in hominins is dramatically
different from other primates in many key ways that reveal
human adaptations to bipedalism, thermoregulation and par-
turition (see [22,23] and summaries in the literature; [7,9,24]
for more details). The mechanical goals of modern bipedalism
appear to be to walk with long strides, with high mechanical
efficiency, while not falling over [5]. Thus, the ability to bal-
ance our upper body on long extended limbs is seen as a
central pressure for pelvic anatomy in humans. The curvature
in the lower back (lumbar lordosis) helps balance the upper
body over the pelvis. In the upper half of the pelvis, the
size, shape and orientation of the iliac blades differ between
apes and humans in a way that reflects our commitment to
striding bipedal locomotion (figure 1). Our iliac blades are
shorter than those of apes, lowering our centre of mass and
avoiding entrapment of the lumbar vertebrae by the iliac
blades, thus allowing lumbar lordosis [11,20,21].
Our iliac blades are also reoriented compared with non-
human apes (figure 1). The iliac blades in non-human primates
are tall, flat plates and oriented in the coronal plane, allowing
the gluteal muscles, which arise from the external surface of
the iliac blade and insert on the proximal femur, to run poster-
iorly over the hip and extend (retract) the thigh [6,7,25–27]. In
modern humans, in contrast, the iliac blades curve around the
side of the body (facing laterally) and flare outward, produ-
cing the characteristic bowl shape of the modern human
pelvis and allowing the lesser gluteals—especially gluteus
 (a) (b) 3
P. troglodytes A. afarensis H. sapiens

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S1
S2
S3
inlet S4 inlet
S5

midplane

outlet

false
midplane pelvis
inlet

Phil. Trans. R. Soc. B 370: 20140063

true midplane
pelvis
outlet

outlet
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Figure 2. (a) ‘Midwife’s’ (inferior or lithotomy position) view of the birth canal in a chimpanzee (P. troglodytes), A. afarensis (A.L. 288– 1, ‘Lucy’) and a modern
human female. The position of the infant’s head at the three major planes of the birth canal is shown. Note the non-rotational birth in apes in which the infant’s
head faces anteriorly throughout, the contrasting non-rotational pattern in the australopithecine in which the infant’s head faces laterally throughout, and the
rotational pattern in the human in which the infant’s head enters the birth canal facing laterally, but then rotates at the midplane and exits the outlet
facing posteriorly. Also note that in the two hominins, the infant’s head exits the pelvis more anteriorly, under the ischiopubic rami. (b) Midsagittal (above)
and coronal (below) sections through a modern human female pelvis, illustrating the three major planes of the birth canal. (a adapted from [24,33]; b from [1]).

medius—to cross laterally over the hip, making them abduc-
tors rather than extensors and allowing them to play a role
as pelvic balancers in bipedal walking [9–12,22,27–29].
During bipedal walking when the body is supported by a
single leg, the pelvis has a tendency to tip towards the unsup-
ported side. When apes walk bipedally, they compensate by
leaning their trunk towards the supported side or stretching
out their arms [30,31], but such side-to-side weight shifts are
energetically costly [30]. In humans, the gluteal muscles on
the support side are able to balance the trunk more efficiently
by pulling up the unsupported side of the pelvis [22,23,32].
Changes in the lower half of the pelvis—sacrum, ischium
and pubis—affect the anteroposterior (A –P) and transverse
diameters of the birth canal as well as the attachment sites
for ligaments that connect the ischium and sacrum, and for
the hamstrings and adductor muscles. Most notably, the
ischium in apes is long, and the ischial tuberosity where
the hamstrings and the sacrotuberous ligament attach faces
downward (figure 1), whereas in humans, the ischium is
shortened [22,23]. Because the human thigh is relatively
long, this means that the insertion site for the hamstrings is
pulled further away from the origin [7,11,22]. In addition,
the ischial tuberosity is more robust than in apes, and
angled with part of it facing upwards [7,11,22]. The position
and form of the ischial tuberosity in humans not only
improves the leverage for the hamstrings, but also reflects
increased tension from the sacrotuberous ligament. The lum-
bosacral joint is located anterior to the sacroiliac joint in
humans, and so body weight tends to rotate the sacrum anter-
iorly, depressing the sacral promontory and elevating its apex
(lower end), along with the coccyx. The sacrotuberous liga-
ment, which runs from the sacrum and coccyx to the ischial
tuberosity, resists sacral rotation and is loaded under tension
by body weight during bipedal standing and walking
[7,11,22,31].
The shape of the lower half of the pelvis (along with soft
tissues, including the pelvic floor muscles) dictates the shape
of the birth canal and shows important differences between
apes and modern humans related to neonatal head size. In
modern humans, the sacrum is shorter, wider, more ventrally
concave and rotated anteriorly compared with apes ([23];
figure 1). In modern apes, the birth canal is large and poster-
iorly oriented, with the largest diameter anteroposteriorly,
owing to the superiorly placed sacrum and coronally oriented
iliac blades [33]. Combined with a relatively small neonatal
head size, this makes for a comparatively easy and non-
hazardous birth process in apes (although not necessarily
in smaller primates: [14,24,34]). In all primates, the greatest
diameter of the infant’s head—the A–P diameter—aligns
with the greatest diameter of the mother’s birth canal, so
ape infants are thought to pass through the entire birth
canal facing forward and without rotation (figure 2;
[1,24,33,35]; although further research, [34], is needed to
verify the process of ape parturition).
The situation is quite different in humans. Humans have a
relatively narrow birth canal in the A–P plane, possibly, in
part, because of the needs for efficient balancing in bipedal
locomotion. The human sacrum is unique in forming a bony
posterior/superior wall of the birth canal [23,33]. The result
is a pelvic inlet that is wider mediolaterally than anteroposter-
iorly (i.e. transverse diameter is long relative to the anatomical
conjugate, between the pubic symphysis and sacral promon-
tory; [1,33,35,36]). In compensation, the modern human birth
canal at its more inferior levels is enlarged in humans,
especially in women. This expansion is accomplished through
the anterior rotation and sagittal curvature of the sacrum,
 along with overall A–P expansion of the pelvis provided
mostly by lengthening of the pubic rami [33,36]. These
changes are accompanied by shortening and flaring of the
ischia, which result in a wider subpubic angle and a greater
distance between the ischial spines (figure 2). When looked
at as a canal with a superior inlet and inferior outlet, the chan-
ging shape of the human birth canal is clear: although there is
variation [37], in general, the human female pelvis is wide
mediolaterally at its inlet, whereas the midplane and outlet
are wider anteroposteriorly (figure 2; [1,35]).
The contrasts between the human pelvis and that of apes,
in combination with the relatively large body size and cranial
size of human neonates, have a profound effect on human
infant delivery: we experience a birth process that is unusual,
if not unique [1,24,34]. A human infant must perform a
series of twists and turns during parturition to enable the
widest dimensions of its body to pass through the changing
dimensions of the maternal birth canal, a process called
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rotational birth. A human infant’s head enters the inlet facing
laterally, and then rotates, usually to face backwards, as it
exits through the more anteroposteriorly oriented outlet
(figure 2). Although the precise orientation of the infant may
vary depending on variations in the shape of the maternal
birth canal, this rotation during birth is consistent among
humans [4]. In contrast with an ape baby, who passes through
the outlet posteriorly between the ischial tuberosities, a human
baby emerges more anteriorly, under the ischiopubic rami,
owing to the inferior position of the sacrum [33,35].
Finally, even if the delivery of its large head is successful,
a human infant’s broad, rigid shoulders pose another signifi-
cant obstacle to birth [1,35]. We share wide shoulders in
common with the great apes, but ape mothers are generally
able to deliver their infants’ shoulders without difficulty
because of their relatively large and comparatively straight
birth canals. In humans, however, shoulder dystocia is a
serious potential complication in childbirth, and further con-
tortions are required from the infant to squeeze its shoulders
through the birth canal. After its head emerges, the baby
must rotate twice more, so that its shoulders can align first
with the pelvic inlet and then with the outlet [24]. Trevathan
and Rosenberg have argued [24,35,38] that passage of the
infant’s shoulders may have influenced the size and shape
of the human pelvis as much as head size.
Skeletal anatomy, whether of the mother’s pelvis or the
infant’s head and shoulders, is not the only factor influencing
the pattern and degree of difficulty of birth in humans. The
soft tissues of the pelvic cavity and vagina are compressed or
stretched during delivery, facilitating passage of the infant.
The effects of the hormone relaxin are another important
factor promoting successful parturition. Relaxin production
by the ovaries and uterus increases towards the end of preg-
nancy, increasing the elasticity of the pelvic ligaments [39,40].
This allows some mobility in the maternal pelvis in response
to pressure from the fetal head and shoulders: the bones of
the pelvis can spread apart somewhat at the pubic symphysis
and sacroiliac joints, and the sacrum can rotate anteriorly to
facilitate exit of the fetal head through the pelvic outlet
[39,40]. In addition, the bones of the fetal skull are pliable
and are moulded during passage through the birth canal
[41]. However, endocrine effects on parturition in extinct ani-
mals are difficult if not impossible to ascertain, and neonatal
remains are vanishingly rare in the fossil record owing to
their great fragility [1]. The morphology of the maternal
pelvis, therefore, provides the best information about the pro- 4
cess of birth in extinct hominins, and this review focuses on
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inferences made from fossil pelvic anatomy.
3. Pelvic evolution in early (non-Homo)
hominins
An examination of pelvic anatomy in the fossil record reveals the
varying pressures on the pelvis during human evolution. The
evolution of the pelvis in the earliest hominins—Ardipithecus
ramidus, Australopithecus afarensis, Au. africanus and the more
Phil. Trans. R. Soc. B 370: 20140063
recent Au. sediba—shows derived features relative to apes, pat-
terns that make enormous logical biomechanical sense in
terms of the appearance and evolution of bipedalism in our lin-
eage. It is well known that there is considerable and often
rancorous debate about the nature of locomotion in early hom-
inins: was it like modern humans, like that of chimpanzees, or
something in between? It is not the goal of this paper to revisit
this debate; the details can be found in the original works
[7,26,27,42], in more recent summaries [10,11,21,43–47] and a
myriad of articles in between. With the recent description of
Ar. ramidus [20,48], there is no longer debate about whether
there was a phase of non-human-like bipedalism in our lineage.
The debate now is simply about how long that form of loco-
motion lasted, and how it is reflected in the anatomy of the
pelvis. Some argue that full human-like locomotion arose
with Au. afarensis (see reference [11] for a summary), whereas
others argue that it appeared only in Homo erectus (see [46] for
a summary).
Our earliest detailed knowledge of the hominin pelvis comes
from fossils of Ardipithecus ramidus, from Ethiopia 4.4 million
years ago (Ma)3 [20,52]. Although the Ar. ramidus pelvis
required extensive reconstruction, most agree that it displays
a ‘mosaic’ pattern: the upper half of the pelvis is substantially
changed relative to apes, whereas the lower half retains more
primitive features [20]. The iliac blades are repositioned and
flare laterally compared with apes, improving gluteal lever-
age and allowing control of side-to-side balance during
walking (figure 1). In addition, the change in iliac blade
shape and position appears to reduce lumbar entrapment
and to facilitate the development of lumbar lordosis. Reflect-
ing on the selective factors that drove changes in iliac height,
Lovejoy & McCollum ([21], p. 3297) recently argued that ‘the
favourable position of the anterior gluteal muscles in homin-
ids that allows them to control pelvic tilt during single
support can now be seen to have been largely a refinement
that followed the initial primary adoption of a lordotic
spine with an emancipated caudal-most lumbar vertebra’.
In contrast to changes in the ilium, the long ischium of
Ar. ramidus is reported to be more like that of living
apes, with a hamstrings tuberosity that points downward
(figure 1). Lovejoy et al. ([20], p. 71) argue that this suggests
that Ar. ramidus was ‘ . . . an effective upright walker’ and
that ‘it could also run, but probably with less speed and
efficiency than humans’.
The shape of the pelvis in the early African australopithecines
Australopithecus afarensis and Au. africanus, which post-date
Ar. ramidus by 1–2 Myr, has been described in detail
previously (see [6,7,9,11,26,27,53–56]), and the two species are
for the purpose of this paper considered similar (though see
reference [56] for a sense of the variation) and will be discussed
together. As in Ar. ramidus, the iliac blades of Au. afarensis and
 (a) (b) 5

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A. afarensis H. erectus
(c) (d)
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H. erectus A. afarensis H. sapiens

H. heidelbergensis H. sapiens
Figure 3. Anterior and superior views of the pelvis in (a) female A. afarensis (‘Lucy’), (b) female H. erectus (Gona), (c) male H. heidelbergensis (Atapuerca Pelvis 1)
and (d ) female modern H. sapiens. The bones are shown to approximately the same scale. (e) Comparison of pelvic shape and body proportions in H. erectus,
A. afarensis and modern H. sapiens. Note the relatively broad pelvis in A. afarensis, and the great degree of iliac flare in both fossil hominins. As iliac flare and pelvic
width decrease the body shape becomes relatively narrow, promoting effective heat loss. (a,b and d from [57]; c from [37]; e by J. Gurche from [58]).

Au. africanus are vertically shortened and have moved laterally
compared with living apes ([27]; but see reference [7] for a
contrary view), and also exhibit great lateral flare, meaning
they extend laterally out over the hip joint and femoral neck
(figure 3). The pelvis overall is extremely wide, wider than in
any modern human of their small size [55], with a wide inter-
acetabular distance, making it broader mediolaterally than
anteroposteriorly (i.e. platypelloid in shape).
The extreme width of the australopithecine pelvis has pre-
sented enormous challenges for interpretation. Lovejoy and
co-workers [6,11,20,21,25,59] see it as part of a larger complex
of lateral iliac flare and long femoral necks that produce
human-like hip kinetics. Ruff [10,60] has also argued that
this morphological complex is related to hip mechanics,
suggesting that the iliac flare provides better leverage for
the gluteal muscles to balance the pelvis (but see [61,62]),
and moderates forces applied to the lower limb joints. How-
ever, in contrast to Lovejoy and co-workers, Ruff contends
that this arrangement would also increase mediolateral
(M –L) loading of the femur and require a non-human-like
gait in the australopithecines [10]. Other researchers argue
that the australopithecine pelvic morphology only makes
sense for a biped who may still spend some time in the
trees or whose bipedalism differs in style or frequency from
that of modern humans [26,44,46].
Rak [8] argued that as the wide australopithecine pelvis
rotated in the transverse plane during walking, its great
bi-acetabular diameter allowed more forward movement of
the lower limb for each degree of rotation. This increased
stride length in these short-legged bipeds (much shorter rela-
tive to body size than in modern humans ([63– 65], but see
[66]; figure 3) while controlling vertical oscillation of the
body’s centre of mass. Longer strides, in turn, decrease stride
frequency and the cost of locomotion. We investigated Rak’s
idea with an experimental study [67] that tested for relation-
ships between lower limb length and locomotor variables
(stride length, speed and pelvic rotation) in modern humans.
Our work supports Rak’s hypothesis: when people with
short legs (as in australopithecines) walk faster, they show
pronounced increases in pelvic rotation, which increases
their stride length. Whitcome et al. [68] similarly found that
women take longer strides relative to leg length when walking
than men do, probably thanks to their broader pelvises. How-
ever, it is unclear how much the energetic cost of locomotion
may actually have been elevated by the broad pelvis in aust-
ralopithecines [32,62], and whether any increase in cost
would have been offset by the energetic advantages of a
longer stride. Stern [46] raised concerns that we share about
the costs of vertical oscillations and stride length, suggesting
that specific stride lengths and levels of centre of mass
oscillation may provide energetic efficiency. Work by Wall-
Scheffler and co-workers [69,70] suggests that a broad pelvis
may provide the additional important benefit of increas-
ing speed flexibility relative to locomotor cost, especially
in short-legged individuals. Furthermore, Wall-Scheffler and
co-workers have found that a wider pelvis reduces the cost
of carrying an infant, which would have been an important
energy drain for early female bipeds [69,71,72].
 The functional significance of anatomical changes in the
lower half of the pelvis is even more ambiguous. Australo-
pithecus afarensis and Au. africanus have anatomy that is
intermediate between Ar. ramidus and modern humans.
The sacrum is placed relatively posteriorly compared with
apes, and like the pelvic inlet, the lower planes of the birth
canal in early hominins are platypelloid in shape (figures 2
and 3; [11]). The position and shape of the ischial tuberosity
and attachment sites for the hamstrings (figure 1) have been a
subject of significant debate [6,7,11,20,26,27,53,73]. Some
researchers [6] argued that australopithecines had poor
leverage for their hamstrings, whereas others [7] argued
that the leverage was strong; these differing results could
be explained, in part, by differing methodologies and indices
against which leverage is measured. In addition, the relative
flatness of the sacrum and the orientation of the ischial tuber-
osity in Australopithecus compared with modern humans
suggest that the sacrotuberous ligament was less robust,
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and vertical loading during locomotion was reduced [7].
The presence of a platypelloid birth canal from the inlet
through to the outlet in Au. afarensis and Au. africanus
suggests to most authors that Australopithecus had a unique
type of non-rotational birth (figure 2). The neonate’s head
would have entered the inlet facing laterally, and remained
in that position until it exited the outlet [24,33,74]. Import-
antly, Trevathan & Rosenberg [38] have pointed out that
the birth could not have been truly non-rotational, because
as in modern humans, the infant would have to have rotated
after emergence of the head to allow passage of its wide, rigid
shoulders. In notable contrast, Berge & Goularas [56] recon-
structed Sts 14 as less platypelloid than previously thought,
and said that Au. africanus had a ‘human-like movement of
rotation and flexion of the fetal skull’ during birth ([56],
p. 262; see also [36,75]). Neonatal head size relative to the
size of the birth canal in australopithecines was probably
closer to apes than humans ([33,75 –77]; but see [1,36]), and
the M –L wide and A–P narrow pelvis is seen by most
authors as a barrier to birth of infants with relatively large
brains (see [16] for a review of encephalization and birth in
fossil hominins).
The relative importance of obstetric versus locomotor or
thermoregulatory demands (or even neutral evolutionary
processes) in determining pelvic morphology in extinct hom-
inin species could be illuminated by an assessment of sexual
dimorphism, because obstetric constraints affect females,
whereas other selective pressures would presumably affect
both sexes equally. In modern humans, for example, pelvic
breadth is related to climate in the same manner in males
and females [13], but a number of features of the female
bony pelvis increase the size of the birth canal relative to
males [78]. Unfortunately, the fossil record of the hominin
pelvis is so sparse that it is usually impossible to make
comparisons between male and female anatomy for any par-
ticular species [17,36,79]. In fact, because evolution has
produced in fossil hominin pelvises ‘a combination of traits
in an extinct animal that is duplicated by no living creature’
[80, p. 110], it is often very difficult even to assign a sex to
any given fossil [2,33,36,81 –83].
A rare glimpse of pelvic sexual dimorphism in an early
hominin is provided by the recently described 1.9 Myr old
South African species Australopithecus sediba. These fossils
have garnered attention because of their age (close to some
of the oldest specimens of Homo erectus) and because of
arguments that they may be ancestral to Homo [84]. Pelvic fos- 6
sils from a male and a female have been recovered, and
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although conclusions must be tempered by the male’s juvenile
status, Kibii et al. ([84], p. 1408) report that Au. sediba ‘may not
have expressed the marked sexual dimorphism characteristic
of modern humans’; neither the male nor the female exhibits
the suite of traits that characterizes the respective sex in
H. sapiens. Furthermore, despite the fact that Au. sediba’s
pelvic inlet (in the male and the female specimen) is somewhat
rounder compared with earlier australopithecines, it also had a
small adult brain size, and therefore Kibii et al. argue that any
Homo-like aspects of the pelvis are not related to increased
Phil. Trans. R. Soc. B 370: 20140063
neonatal cranial size and are not explicable by obstetric
requirements. They conclude that ‘birthing of larger-brained
babies was not driving the evolution of the pelvis at this
time’ ([84], p. 1407), and that if these changes in Au. sediba
are functionally relevant rather than part of a phyletic signal,
they represent locomotor rather than obstetric demands.
The overall pattern of pelvic morphology in Au. sediba is
complex and differs from both earlier australopithecines and
early Homo. Kibii et al. [84] report that these specimens not
only retain the wide interacetabular distance, long pubic rami
and sacral shape of prior australopithecines, but also have
Homo-like features including shorter, more vertical ilia and a
shortened ischium with an everted ischial tuberosity. Their
explanation for these features is complex, noting at some
points that it is possible that the shortened ischium is due to
pleiotropy associated with other pelvic changes, and at others
arguing for a phyletic affinity with Homo, rather than novel
locomotor or obstetric requirements. Kibii et al. also point out
that Au. sediba’s relatively M–L broad birth canal (and long
femoral neck), but smaller degree of iliac flare, may present a
challenge for Ruff’s model of australopithecine hip biomechan-
ics [10,84]. A conclusive interpretation of the complex and
potentially mosaic features of the pelvis of Au. sediba will require
more material from adult members of this species and a more
extensive biomechanical analysis like those applied to earlier
fossil hominins.
What is there to make of the overall picture of the aust-
ralopithecine pelvis? Changes in the iliac blade in our early
ancestors compared with other primates are a response to
fundamental needs for pelvic balance in bipedal walking,
whereas changes in ischial and pubic shape are more com-
plex. Some researchers are convinced that the nature of
locomotion in early hominins was close to that of modern
humans, and differences between australopithecines and
Homo reflect obstetric changes [11,20,21]. Other researchers
take a different view and argue that there were major loco-
motor differences between australopithecines and Homo (see
[7,46,84]), and counter that pelvic differences reflect both
locomotor and obstetric needs. In this formulation, there
were multiple phases of locomotor evolution in australopithe-
cines, and obstetric concerns played a relatively minor role
until the evolution of Homo species and possibly not until
H. sapiens specifically.
In our estimation, early stages of hominin pelvic evolu-
tion reflect clear pressures for locomotion without an
obvious nod towards either obstetric or thermoregulatory
pressures. Although such pressures were certainly present
prior to 2 Ma, they were neither dominant nor even equal to
pressures of bipedal locomotion on pelvic form. Whatever
the adaptive or functional significance of australopithecine
pelvic morphology may be, it is intriguing that virtually the
 same suite of anatomical features is seen in the pelvis of
another hominin from the opposite side of the globe and
millions of years later. The small-bodied, short-legged, small-
brained hominin Homo floresiensis (the ‘Hobbit’)—from Java
as recently as 17 000 years ago [85,86]—has a pelvis that is
strikingly similar to that of Au. afarensis [87,88], suggesting
similar adaptive pressures despite great geographical and
chronological distance.
4. Pelvic evolution in fossil Homo
The genus Homo emerged in the early Pleistocene, just after
2 Ma, and the first representatives of H. sapiens appear in the
fossil record around 200 thousand years ago (kya). The scarcity
of relevant fossils in the intervening period makes interpretation
difficult, but some evolutionary patterns over time are evident.
The pelvises of early Homo, although similar in overall form to
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earlier hominins, do have derived traits that distinguish them
from australopithecines. Many of these are probably related to
changes in locomotor behaviour. Once established in the Early
Pleistocene, pelvic morphology in archaic Homo seems to be
largely characterized by morphological stasis over time. How-
ever, some trends towards modern human anatomy can be
seen later in the Pleistocene, most likely adaptations to birthing
larger-brained babies. A new additional source of variation in
genus Homo is thermoregulatory adaptation to increasingly
diverse environments.
(a) Early Homo
Patterns of pelvic evolution in earliest Homo are obscured by
continuing uncertainty about taxonomy [19,60,83,89–92] and
the almost total absence of pelvic fossils that are associated
with diagnostic craniodental remains (the one notable exception
being the Nariokotome H. erectus skeleton from Kenya [93]).
Furthermore, although hominins spread out of Africa during
the Early Pleistocene and into Europe and Asia [94,95], pelvic
fossils of early Homo have been recovered exclusively from
Africa [2]. The available evidence, however, suggests that the
morphology of the pelvis in these early members of our genus
(specimens typically referred to H. erectus, but sometimes
to H. ergaster, H. habilis or H. rudolfensis [60]) remained largely
consistent for at least a million years (see [60,96–99]). The
pelvis retains the primitive overall form established in early
hominins—mediolaterally broad and anteroposteriorly
narrow, with a platypelloid birth canal and laterally flaring
iliac blades (figure 3; [2,60,66,74,100])—but there are changes
compared with earlier australopithecines that appear to reflect
ecological and behavioural changes occurring in our genus.
Ruff [10,60] has proposed that many features of pelvic
morphology in early Homo are part of a single biomechanical
complex that increases the leverage of the gluteal abductors,
as in the australopithecines. However, changes in locomotor
behaviour also appear to be reflected in some aspects of
pelvic morphology [60,96–99]. Open, grassy savannahs first
became a major component of the African landscape in the
Early Pleistocene [101–104], and many authors have suggested
that greater mobility in early Homo, along with increased
exposure to heat from the sun, required improvements in loco-
motor efficiency [10,105–109]. The pelvis of H. erectus, while
broad compared with modern humans, was narrower relative
to body height than in the australopithecines (figure 3; [2,60]).
This change in proportions, combined with longer lower limbs
and higher crural indices [110], would have allowed H. erectus 7
to walk farther at a lower energetic cost (although some
rstb.royalsocietypublishing.org
studies suggest that a broad pelvis may improve some aspects
of walking economy [70,111]).
Endurance running may have been a component of the
hominin locomotor repertoire for the first time in H. erectus
[108,112]. Novel adaptations to running may include the
overall narrower pelvis, large and deep acetabulae, more
sagittally curved iliac blades, increased ligamentous stabiliz-
ation of the pelvic joints and more powerful muscles
[82,108,111]. With these morphological changes, movements
of the trunk and hip were decoupled, allowing more move-
Phil. Trans. R. Soc. B 370: 20140063
ment between the two, the trunk was better stabilized, and
joint stress was reduced, making running a viable subsistence
behaviour for H. erectus.
Other features of the pelvis in early Homo are probably
related to parturition. The primitive platypelloid birth canal
suggests that these hominins would have also retained the
presumed primitive non-rotational birth mechanism of the aust-
ralopithecines [2,60,74]. However, some expansion in relative
A–P dimensions of the birth canal did occur, primarily through
upward rotation of the pubic rami (figure 3; [60,82]). This enlar-
gement is probably related to an increase in neonatal brain size.
There was an initial jump in encephalization (relative brain
size) over australopithecines with the origins of the genus
[60,113,114], and non-rotational birthing of a relatively larger-
brained and larger-shouldered infant would explain the
continued need for the M–L broad true pelvis [2,38].
Subsequently, levels of encephalization in H. erectus remained
largely unchanged, because further brain size expansion
accompanied proportional increases in body size [60,113,114].
This stasis in relative brain size may account for the long-term
consistency in pelvic morphology in the species.
Finally, thermoregulation, while probably not a major
selective force acting on the australopithecines, became increas-
ingly important for Homo owing to climatic change and
geographical radiations into more varied habitats [2,13]. As dis-
cussed in §3, a wider body conserves heat, whereas a narrower
body increases heat loss [8,9]. In a series of exhaustive studies,
Ruff has demonstrated that pelvic bi-iliac breadth (intercristal
diameter) varies with latitude among modern humans
[2,12,13,115], and fossil hominins show the same pattern, with
those from warmer climates, including African earliest Homo,
having smaller pelvic breadths than those from colder climates
[2,12,13,116] (although see [18]).
An exception to the prevailing pattern of pelvic morphology
in early Homo is the Ethiopian Gona pelvis, dating to 1.4–0.9 Ma
and identified by its discoverers as an adult female H. erectus
(figure 3; [82]). Although Gona is similar in some ways to
other H. erectus specimens, it differs in key features. Most nota-
bly, the Gona pelvis is even broader overall, but has a more
gynecoid (rounder) birth canal with a particularly large pelvic
outlet. Simpson et al. [82,83] argue based on this specimen
that adaptations to birthing larger-brained infants, including
perhaps rotational birth, arose earlier than previously sus-
pected. Another surprise, especially considering the great
breadth of the Gona pelvis, is the very diminutive body size
implied by its small acetabulae, which are more similar in size
to those of australopithecines than to other H. erectus [2,82]. In
contrast to the majority of the fossil evidence, which suggests
that body sizes in H. erectus were in the modern human range
(summarized in reference [2]; although see [90,117,118]), Gona
indicates either very high levels of sexual dimorphism or
 great intraspecific variation in body size [82,83]. Ruff [2] has
questioned these unexpected conclusions about H. erectus,
countering that Gona may in fact represent Australopithecus
(see [83] for Simpson et al.’s response).
In sum, early Homo provides another illustration of power-
ful, conflicting selective pressures acting on the pelvis: while a
greater overall width was beneficial for non-rotational partur-
ition, a narrower pelvis was more advantageous in terms of
both locomotor efficiency and thermoregulation in hot cli-
mates. However, it is important to keep in mind the lessons
of the Gona fossil, which, like Au. sediba, shows that there
was significant postcranial diversity among Pleistocene homin-
ins, and details of mosaic evolution in specific populations
may contrast with overall long-term trends [92].
(b) Middle Pleistocene Homo
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Africa, Europe, the Middle East and East Asia have all pro-
duced fossil human pelvises from the Middle Pleistocene
(about 800–100 kya). As in earlier Homo, the available evi-
dence again suggests that the overall primitive pelvic form
was retained in archaic (pre-modern) humans without major
changes throughout most of this time period [37]. The biomech-
anics of bipedalism had reached an essentially modern state in
H. erectus in the Early Pleistocene [108], so any changes in
pelvic morphology during the Middle Pleistocene are unlikely
to reflect major locomotor adaptations. Ecogeographic patterns
in pelvic breadth indicate that thermoregulatory constraints
continued to play an important role [2] throughout the Pleisto-
cene. However, the demands of parturition became ever more
critical: trends towards modern human morphology during
this time become more clear and are probably related to
increases in encephalization and concomitant requirements
for birthing larger-brained babies. Populations from the later
Middle Pleistocene exhibit mosaic evolution and diversity in
anatomical details, and the modern human pelvic form (with
narrower overall breadth and a birth canal requiring rotational
birth) seems to appear between 200 and 100 kya, with the
origin of H. sapiens.
Pelvic anatomy in H. heidelbergensis (sometimes referred to as
‘Archaic H. sapiens’4) is known primarily from high-latitude
individuals from Europe and Asia (figure 3; [37,97,116,
119,120]). These pelvises are very large overall and remark-
ably wide: the specimens from Atapuerca (Spain) and
Jinniushan (China), for example, have the largest bi-iliac
breadths in the human fossil record, and are wider than the
average for any modern human population studied by Ruff
[2]. This great overall breadth reflects both a very wide trans-
verse diameter of the birth canal and a high degree of iliac
flare in H. heidelbergensis—although generally less than in
earlier Homo [2,37,100,116,119,120]. Again, these two aspects
of pelvic breadth are probably linked biomechanically [2,60].
The birth canal is broad both at the inlet, partly because of
long superior pubic rami, and at the outlet; the shape of the
midplane is not clear owing to poor preservation of the
iliac spines in the relevant fossils [2,37,60,74,116,119].
The pelvis in Neanderthals is typically very similar to
H. heidelbergensis in its morphology and large size [1,74,100,
120–126] (but see [127]). Except for a few uniquely derived
characteristics, particularly in the cross-sectional shape of the
superior pubic ramus (see [100,120] for reviews), recent work
has shown that most of the features of the Neanderthal
pelvis that were once thought to be distinctive are in fact 8
retained primitive traits [37,100,119–121,123,128].
rstb.royalsocietypublishing.org
The shape of the birth canal in most Middle Pleistocene
Homo specimens was primitive relative to modern humans
(although some may have indices within the modern human
range [16,37,116]). This implies a primitive (non-rotational)
birth mechanism in both H. heidelbergensis and Neanderthals,
although this is debated partly because of the paucity and
incompleteness of pelvic fossils [1,2,16,37,96,124,129]. How-
ever, birth canal size in Middle Pleistocene Homo is clearly
larger than in earlier hominins, generally within the modern
human range [16,74,79,119,121,124,127]. Presumably this enlar-
Phil. Trans. R. Soc. B 370: 20140063
gement is related to increases in infant size. Adult body size in
H. heidelbergensis and the Neanderthals, which was within or
above the modern human range, changed little during the
Middle Pleistocene, but adult brain size did increase
[24,60,74,113,114,116]. Fossil evidence suggests that Nean-
derthal infants were born with a body and head size similar
to modern newborns, implying a similar level of obstetric dif-
ficulty in Neanderthals and modern humans [1,16,74,77,124]
and probably ‘obligate midwifery’ (cf. [35,130]). Finally, it is
also of interest that these are the first humans in which
sexual dimorphism in the pelvis can be clearly observed, and
the pattern differs from modern humans. While the birth
canal is larger in females than males, this is primarily owing
to rotation of the pubis, rather than lengthening as in
modern humans. In archaic Middle Pleistocene Homo, the
pubic rami are actually longer in males than in females
[74,79,116,120,121,124].
5. The evolution of the modern human pelvis
European Neanderthals maintained this archaic complex of
pelvic anatomy (and possibly birth mechanism) well into the
Late Pleistocene ([125,131]; but see [127]). However, some
human populations during the late Middle Pleistocene, particu-
larly in Africa and the Middle East, where H. sapiens evolved,
were characterized by a mix of primitive traits and ones that
are derived in the direction of modern humans. The Late
Middle Pleistocene fossil pelvis from Broken Hill (Kabwe),
Zambia, for example, is largely modern in appearance but is
associated with a clearly archaic skull [132] and does have
some primitive traits (see [2,133,134]). Arensburg & Belfer-
Cohen [123] identify a similar mosaic in populations of late
Middle Pleistocene hominins from Israel. They point to
modern traits in pelvic fossils generally classified as Nean-
derthals (such as Kebara 2 and Tabun C) and retained
primitive features in early ‘anatomically modern H. sapiens’
such as Skhul IV and IX [123] (but see [122]). This mix of
traits is what might be expected in closely related groups
with a rapidly evolving new morphology. An alternative explan-
ation could be interbreeding between groups that display
different morphologies, but this hypothesis would be difficult
to test based on the small samples available.
When exactly fully modern human pelvic anatomy first
appeared is unclear at this time, but it must be near the end
of the Middle Pleistocene, and certainly by 100 kya. The earl-
iest likely examples are found in individuals from East
Africa and the Middle East, although the late Middle Pleisto-
cene pelvic fossil record from Africa is particularly sparse
[2]. The Omo I partial skeleton from Ethiopia, described as
‘the oldest well-dated anatomically modern human fossil’
 ([135], p. 421), has been dated to approximately 195 kya
[136]—interestingly, twice the age of the intermediate or
mosaic populations from Israel. However, there are traits
throughout the skeleton that differentiate it from modern
humans, some of which appear to be primitive [135]. Pearson
et al. [135] described the Omo I partial hip bone, but primarily
with respect to sex and age inferences; no archaic features are
mentioned, and no assessments of functional anatomy or phy-
logenetic implications have been made thus far (and may not
be feasible given the fragmentary nature of the specimen).
Whether the modern human pelvis was present at 200 kya
will have to wait for more complete fossils or at least a more
detailed analysis of the Omo I specimen. The 100–95 000
year old Qafzeh 9 H. sapiens skeleton from Israel provides
the next example of likely anatomically modern pelvic anat-
omy. Although it is badly crushed, its morphology appears
to match that of H. sapiens [122,123]. Qafzeh 9 is contempor-
aneous with the Skhul individuals [137], providing further
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evidence that modern morphology evolved in a mosaic
fashion in the Middle East.
Early anatomically modern fossil pelvises, like those of
later H. sapiens, were narrower overall compared with archaic
humans, with a low degree of iliac flare. The birth canal was
equivalent or larger in size and had changed in shape,
becoming less platypelloid throughout, with a shortening of
the pubic rami, especially in males [100,120]. The birth
canal now varied in proportions along its length: the inlet
retained its M–L orientation (although less pronounced),
especially in females, whereas the midplane and outlet
became larger in the A–P dimension. The fossil evidence
suggests, therefore, that the modern human pattern of
rotational birth evolved only with the origin of H. sapiens
[2,74] (although there is some disagreement [124,129]).
What adaptive scenario can explain this dramatic change
in morphology? The answer may involve conflicts between
thermoregulation and obstetrics. In cold-adapted hominins
such as Jinniushan or the Neanderthals, there was no thermo-
regulatory constraint on widening the pelvis to increase birth
canal size (and therefore infant brain size [2,74]). However,
among tropical humans, obstetric and thermoregulatory
pressures on the pelvis are in direct conflict. Weaver &
Hublin [74] and Ruff [2] have proposed that during the
Middle Pleistocene, further increases in pelvic breadth
among low-latitude hominins were impossible because of
heat dissipation requirements. Instead, pressure for narrow-
bodied women to birth larger-brained babies eventually
resulted in yet another morphological compromise in the
pelvis: the capacity of the birth canal was enlarged by
increasing its A–P diameter, particularly at the midplane
and outlet, whereas its M– L dimensions simultaneously nar-
rowed. The resulting twist in orientation allowed the true
pelvis to accommodate a larger-brained infant without also
widening the body [138].
The relationship between body size and shape, overall pelvic
size and shape, and birth canal size and shape in modern
humans is complex [3,15,17,18,79,138–141]. With the evolution
of the modern pelvis, the size of the birth canal was essentially
decoupled from the size of the body: birth canal proportions
in modern humans do not necessarily correlate with body
proportions [138,141,142]. A recent study by Kurki [142] deter-
mined that among modern humans, birth canal transverse
breadth may be constrained by climate, with lower-latitude
populations having relatively narrower dimensions, but the
anteroposterior breadth of the birth canal appears more free to 9
vary in order to maintain obstetric capacity.
rstb.royalsocietypublishing.org
While it has been shown that thermoregulation is an
important factor influencing variation in pelvic dimensions
between geographically widespread human populations
[2,3,12,13], the way in which obstetrics influences variation
among populations, or even between males and females, is
unclear. Evolutionary theory would suggest that because
‘selection intensity and phenotypic variability are inversely
related’ ([143], p. 59), there should be sexual dimorphism in
the variability of pelvic morphology, with variation in the
size and shape of the female birth canal limited by obstetric
Phil. Trans. R. Soc. B 370: 20140063
demands. However, the nature and extent of sexual dimorph-
ism in the modern pelvis are particularly complex, and a large
number of studies have produced no clear indication that
obstetric constraints produced between-sex differences in
variability [3,18,138,139,141–143]. To further complicate mat-
ters, work by Betti and co-workers [3,18,141] has suggested
that among modern humans, neutral evolutionary processes
reflecting demographic history have also been important in
determining variation in pelvic form (see also [17]).
In any case, whatever the patterns of intraspecific variation,
recent and fossil H. sapiens populations from both warm and
cold climates show reduced pelvic breadth compared with
archaic humans from similar latitudes [2,13,138,144]. As anatom-
ically modern humans spread out from their origins in Africa
into higher latitudes [3], they adapted, as earlier hominins
had, by evolving wider bodies to improve heat retention
[2,12,13,138,145], but no modern H. sapiens populations re-
evolved the primitive platypelloid birth canal and great iliac
flare of archaic humans. In addition to its thermoregulatory
and obstetric benefits, the narrower modern pelvis and
accompanying decrease in body mass may have conferred bio-
mechanical and energetic advantages in walking [146] (but see
[15,32,62]), or perhaps improved the adaptive advantages of
high mobility, or of running as a subsistence behaviour
[111,147] (see also [148]). Through the evolution of the uniquely
shaped birth canal, modern humans have been able to birth
larger-brained infants, while maintaining the flexibility to
adapt body breadth to varying habitats and at the same time
improving locomotor performance.
Acknowledgements. We gratefully acknowledge The Royal Society; Pro-
fessors Eric Barrington, Graham Burton and Ashley Moffett for
inviting us to participate in the June 2014 Royal Society discussion
meeting, ‘Human Evolution: Brain, Birthweight and the Immune
System’; and other discussion meeting participants for thought-pro-
voking and helpful comments. We also thank Matt Cartmill, Yoel
Rak, Chris Ruff, Jack Stern, Steve Churchill, Michael Black and
Wenda Trevathan for stimulating conversations about pelvic mech-
anics and selective pressures on pelvic shape. Finally, we are
grateful to two anonymous reviewers for suggestions that improved
the quality of this paper. L.G. and D.S. researched, wrote and edited
this manuscript.
Funding statement. We are supported by Radford University Depart-
ment of Biology and College of Science and Technology and Duke
University Department of Evolutionary Anthropology.
Endnotes
1
Betti and co-workers [3,18] have shown that in addition to selection
based on thermoregulatory and locomotor constraints, neutral
evolutionary processes have also had a significant effect on pelvic
shape variation among modern humans (see §3). However, the
fossil record is insufficient to allow investigation of the extent of
neutral evolution of the pelvis in extinct humans.
 2 4
Hominins are defined those members of our lineage, both living
humans and our extinct relatives, that are more closely related to our-
selves than to apes. All known hominins are identified as being
bipedal [19].
3
Although other hominin species are known from as early as 6 –7 Ma
[49– 51], nothing is known about their pelvic morphology, because
no fossil pelvic material has been recovered.
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