Expert Review
org
Evolution of the human birth canal
Philipp Mitteroecker, PhD; Barbara Fischer, PhD
Introduction
Many anatomical, physiological, medi-
cal, and psychological factors contribute
to the success of labor. However,
compared with childbirth of other pri-
mate species, human childbirth is a long
and risky process because the large head
of the fetus has to pass and rotate
through a relatively small, rigid, and
twisted birth canal. Thus, human child-
birth is associated with a considerable
risk of morbidity and mortality, espe-
cially where access to obstetrical care is
limited. This raises the question as to
how and why this small human birth
canal has evolved.
Humans are not the only primate
species with a relatively small birth canal.
Macaques, for instance, also show a tight
fetopelvic fit (Figure 1), yet maternal
morbidity and mortality seem to be rare.
Several mammals, such as rodents and
bats, even give birth to fetuses that are
considerably larger than human fetuses
compared to the body mass of their
mothers. Which other evolutionary so-
lutions did these species evolve and why
have these evolutionary paths not been
accessible to humans?
The last years have witnessed growing
interest and a series of empirical and
theoretical studies on these topics. Here,
we reviewed this body of work and
Unit for Theoretical Biology, Department of
Evolutionary Biology, University of Vienna,
Vienna, Austria.
Received May 2, 2022; revised Sept. 7, 2022;
accepted Sept. 7, 2022.
The authors report no conflict of interest.
This study received funding from the Austrian
Science Fund, Elise Richter Project 826-B.
Corresponding author: Philipp Mitteroecker,
PhD, philipp.mitteroecker@univie.ac.at
0002-9378
ª 2022 The Author(s). Published by Elsevier Inc. This is
an open access article under the CC BY license (http://
creativecommons.org/licenses/by/4.0/).
https://doi.org/10.1016/j.ajog.2022.09.010
ajog.
It seems puzzling why humans have evolved such a small and rigid birth canal that entails
a relatively complex process of labor compared with the birth canal of our closest rel-
atives, the great apes. This study reviewed insights into the evolution of the human birth
canal from recent theoretical and empirical studies and discussed connections to ob-
stetrics, gynecology, and orthopedics. Originating from the evolution of bipedality and the
large human brain million years ago, the evolution of the human birth canal has been
characterized by complex trade-off dynamics among multiple biological, environmental,
and sociocultural factors. The long-held notion that a wider pelvis has not evolved
because it would be disadvantageous for bipedal locomotion has not yet been empirically
verified. However, recent clinical and biomechanical studies suggest that a larger birth
canal would compromise pelvic floor stability and increase the risk of incontinence and
pelvic organ prolapse. Several mammals have neonates that are equally large or even
larger than human neonates compared to the size of the maternal birth canal. In these
species, the pubic symphysis opens widely to allow successful delivery. Biomechanical
and developmental constraints imposed by bipedality have hindered this evolutionary
solution in humans and led to the comparatively rigid pelvic girdle in pregnant women.
Mathematical models have shown why the evolutionary compromise to these antago-
nistic selective factors inevitably involves a certain rate of fetopelvic disproportion. In
addition, these models predict that cesarean deliveries have disrupted the evolutionary
equilibrium and led to new and ongoing evolutionary changes. Different forms of assisted
birth have existed since the stone age and have become an integral part of human
reproduction. Paradoxically, by buffering selection, they may also have hindered the
evolution of a larger birth canal. Many of the biological, environmental, and sociocultural
factors that have influenced the evolution of the human birth canal vary globally and are
subject to ongoing transitions. These differences may have contributed to the global
variation in the form of the birth canal and the difficulty of labor, and they likely continue
to change human reproductive anatomy.
Key words: biocultural evolution, bipedality, cesarean delivery, childbirth, evolutionary
medicine, human evolution, pelvic floor disorders, pelvis, obstetrical dilemma, relaxin
summarized the current understanding skeletal adjustments. To balance the up-
of the evolutionary conundrum under- per body over the hip and legs, the pelvis
lying the human birth canal. We tried shortened and broadened, the sacrum
to provide a multidisciplinary perspec- lowered, and the iliac blades evolved
tive that interprets medical evidence stronger lateral flare. The overall width
within an evolutionary framework. of the pelvis, largely determined by bi-
In particular, we highlighted the inter- iliac breadth, had to suffice not only
action of cultural, medical, and biolog- the biomechanics of locomotion but also
ical processes, so-called gene-culture the thermoregulation of the body (see
coevolution. In the Glossary, we briefly below). Overall, the pelvis evolved into a
explained some key terms from evolu- bowllike structure able to support the
tionary biology that may be less familiar weight of the spine and upper body.2e7
to the medical readership. Only much later, approximately 2
million years ago, early Homo started to
Fossil evidence of pelvic evolution evolve substantially larger brains and
At least 4 to 5 million years ago, homi- bodies. Until approximately 600,000
nids evolved upright walking. This new years ago, increase in brain size was
mode of locomotion required major primarily linked to an increase in total
MARCH 2024 American Journal of Obstetrics & Gynecology S841
Expert Review
FIGURE 1
Schematic representations of the size of the birth canal
Human
anteroposterior diameter
head
transverse diameter
of pelvic inlet
Schematic representations of the size of the birth canal (pelvic inlet, black) relative to the size of the
fetal head (gray) at the time of birth for humans, chimpanzees, gorillas, and macaques (scaled to the
same mediolateral width of the pelvic inlet, redrawn after1).
Mitteroecker & Fischer. Evolution of the human birth canal. Am J Obstet Gynecol 2024.
body mass. Subsequently, brain volume
increased by approximately 30% inde-
pendently of changes in body mass8,9
(Figure 2), which ultimately enabled
the evolution of the wide cognitive and
cultural abilities that humans show
today. However, the increasingly large-
headed babies had to be delivered
through pelvises that had earlier been
adapted to bipedalism, making birth
increasingly difficult.5,6,10 Australopith-
ecines already had pelvises with flared
iliac blades that extended laterally
beyond the hip joint and femoral neck
(Figure 2). These ilia allowed the gluteal
muscles to work as effective balancers in
bipedal walking. Although apes possess a
birth canal where the largest dimension
is oriented anteroposteriorly, australo-
pithecines, similar to modern humans,
had a pelvic inlet where the largest
dimension is the mediolateral one.
Australopithecines had relatively small
heads, and researchers long thought that
their births must therefore have been
comparatively easy.3,11e15 However,
recent research shows that they might
have experienced birth difficulties
similar to humans.16 The pelvises of
early Homo were similar in overall form
to earlier hominins; however, they
possessed derived traits that distinguish
them from australopithecines.15,16 Many
of these are thought to be related to
changes in locomotor behavior.
Although the pelvis of Homo erectus was
S842 American Journal of Obstetrics & Gynecology MARCH 2024
Chimpanzee Gorilla Macaque
broad compared with that of
modern humans, it had less laterally
flared iliac blades than the australopith-
ecines and a rounder pelvic inlet17,18
(Figure 2). As Homo erectus presumably
had both a mediolaterally wide pelvic
inlet and outlet, it is possible that the
birth process did not require the fetus to
rotate.15,17 Neanderthals probably also
had neonates with large heads and rela-
tively difficult birth, but studies disagree
on whether their birth mechanic
was humanlike.19,20 Approximately
300,000 to 150,000 years ago, our an-
cestors had essentially evolved the
modern human morphology, including
modern brain size and pelvic
morphology. Within the past 30,000
years, body size and brain size both
declined slightly in Homo sapiens and
increased again about the same magni-
tude during the last century.8
Fetal size and gestation length
Our earliest mammalian ancestors pre-
sumably were altricial species, that is,
their newborns were relatively helpless
and immature.21 Primates, by contrast,
generally are precocial as they give birth
to relatively mature and mobile
offspring. Humans are an exception in
this regard: compared with the new-
borns of other primates, our newborns
are altricial—a condition termed “sec-
ondary altriciality”22,23. For instance,
human newborns possess only
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approximately 24% of the adult brain
mass, compared with 40% to 70% in the
other great apes. This human condition,
where most brain growth occurs post-
natally, is generally interpreted as an
evolutionary adaptation to ameliorate
childbirth (but see Dunsworth et al24 for
a different interpretation). However,
many other mammals show a much
more pronounced altriciality (eg, squir-
rels, bears, dogs, or cats).25,26 So why do
humans not give birth at an even earlier
developmental stage of the fetus, at a
time when the fetus could easily fit
through the birth canal?
Multiple lines of medical evidence
suggest that birth at an earlier develop-
mental stage would have many disad-
vantages for human neonates. For
instance, delivery before term entails
considerable health risks and an
increased likelihood of impaired cogni-
tive function in later life.27e31 Similarly,
too fast brain growth after birth is asso-
ciated with an increased risk of devel-
oping autism spectrum disorder.32
Pushing birth to an even earlier devel-
opmental stage and accelerating post-
natal brain growth do not seem to be a
viable evolutionary path. Clearly, these
were not the selective factors that have
led to the evolution of the large human
brain; however, a shorter gestation
length seems to be incompatible with the
modern human pattern of neuro-
cognitive development. Conversely, it
has been suggested that not only partu-
rition but also the capacity of the
maternal metabolism imposes an upper
bound on gestation length.24 In other
words, modern human gestation length
and fetal brain development presumably
evolved by “trading off ” these health
risks and cognitive disadvantages against
easier childbirth and metabolic de-
mands. In the presence of such antago-
nistic selection regimes, a population is
expected to evolve a “compromise
phenotype distribution” that maximizes
mean population fitness (ie, average
reproductive success across individuals),
even though this distribution may entail
several diseased individuals with reduced
or even zero fitness33e35 (Figure 3). It
seems likely that modern human gesta-
tion length and fetal development are
ajog.org Expert Review

such evolved compromises of a combi-

FIGURE 2
nation of selective factors. Although a
higher degree of altriciality would ease
Evolutionary change of brain volume and pelvis shape
childbirth and decrease maternal mor-
tality, it would have too many disad-
vantages for the fetus.

Pelvic canal size

Human childbirth could not have been
eased by evolving smaller body and brain
sizes at birth. However, why did the birth
canal not evolve to be more spacious?
Given the considerable individual vari-
ation and heritability of pelvic di-
mensions and their obvious relevance
for survival and reproductive success,
one would expect that human evolution
should have brought about a more
spacious birth canal. Hence, a larger
pelvic canal—despite its advantage for
childbirth—must also entail some dis-
advantages, or the pelvic canal is some-
how else evolutionarily constrained.
In his influential 1960 article, the an-
thropologist Sherwood L. Washburn
coined the term “obstetrical dilemma”
and combined several ideas previously
mentioned by Portman,23 Krogman,36
and others into one hypothesis, sug-
gesting that a pelvis with a larger distance
between the sacroiliac and hip joints Brain volume started to increase approximately 2 million years ago, when the pelvis had already been
would be selected against as it would adapted to bipedal locomotion, which emerged at least 4 to 5 million years ago. Compared with
make upright walking and running australopithecines (shown is the pelvis of Australopithecus afarensis “Lucy” in frontal and superior
energetically more costly.4,6,37 Despite views), Homo erectus (shown is the Gona pelvis) and modern humans have a rounder pelvic canal
being the most popular explanation for and more upright-oriented iliac blades (pelvis images are from Ruff,7 scaled to the same bi-iliac
decades, this “obstetrical dilemma hy- breath).
pothesis” has been tested only in recent Mitteroecker & Fischer. Evolution of the human birth canal. Am J Obstet Gynecol 2024.
years. Several empirical studies, specif-
ically focusing on the mediolateral width
of the pelvis, argued that wide pelvises
are just as energy efficient as narrow
pelvises in bipedal locomotion.38e42 addition to energy efficiency, other biologist M. Maurice Abitbol in 1988. He
There is not much evidence that mod- locomotion-related selective forces were suggested that in upright humans, a
ern variation in pelvic dimensions is suggested to have shaped pelvic small bony pelvic canal contributes to
associated with locomotion efficiency. morphology. Broader pelvises might be the structural stability of the pelvic floor
This finding could be a consequence of more suitable and efficient for child- in men and women. It has further been
the minor importance of pelvic width for carrying,44,45 and certain pelvic mor- argued that a small pelvic canal is better
locomotion efficiency in recent human phologies are better for avoiding injury for supporting the weight of the
evolution or the lack of variation in the than others.46 For example, even small abdominopelvic organs and a large fetus
relevant pelvic dimensions caused by variation in the position and orientation during pregnancy and withstanding
stabilizing selection in the past. Pelvic of the acetabulum can substantially intra-abdominal pressure associated
dimensions important for bipedal loco- affect bipedal walking and cause with physical activities (eg, coughing)
motion vary less with body size than injuries.47e50 while maintaining continence. Thus, a
other body dimensions37,43 and might Another important, yet originally less more spacious pelvic canal would
simply have been restricted by selection influential, hypothesis was proposed by compromise the stability of the pelvic
to a functionally viable range. In the gynecologist, obstetrician, and floor and increase the risk of pelvic floor

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is necessary for a more balanced view of

FIGURE 3
these issues. In a biomechanical study,
Evolution in response to antagonistic selection Stansfield et al53 constructed finite
element models of differently sized pel-
A 2 1
C 3 1
vic floors and applied pressure from
above to these pelvic floor models. They
found that larger pelvic floors deform
more and show higher strains and
stresses in the pelvic floor tissues in
response to pressure, implying that pel-
trait trait vic floor stability tends to decrease with
B D the size of the pelvic canal. These results
× ×
suggest a higher risk of pelvic floor dis-
probability density

probability density
1 2 1 3

orders in women with a large pelvic canal

and thus lend support to the “pelvic floor
hypothesis.” This association between
pelvic canal size and pelvic floor stabil-
ity—even if of secondary clinical rele-
trait trait
vance—does not only impose natural
Many organismal traits, especially those related to human childbirth, have evolved in response to selection for a small pelvic canal (see also
antagonistic selective pressures, that is, by “trading off” different functions. A, A trait affects 2 Discussion) but also may underlie the
functions (f1 and f2), where individual fitness continually increases with increasing trait value for relatively high frequency of incontinence
function f1 but decreases for f2. For instance, increasing the thickness of the pelvic floor muscles and prolapse in human populations.
increases pelvic floor stability but complicates childbirth because passage of the fetal head through Based on their biomechanical models,
the muscular diaphragm of the pelvic floor requires higher uterine pressure (see main text). B, Total Stansfield et al53 also found that a thicker
individual fitness (ie, the average reproductive success of individuals with a certain trait value) is a pelvic floor deforms and stretches less
result of these 2 trait functions (here simply the product f1
f2) and shows a maximum at an in- than a thinner one, which is consistent
termediate trait value, which is a “compromise solution” to these 2 functions. For such a symmetric with clinical reports of reduced thickness
fitness function, evolutionary theory predicts that the mean of the phenotype distribution (blue bell of the levator ani (the muscle forming
curve) evolves to match the trait value with maximal individual fitness because this maximizes the the main part of the pelvic diaphragm)
mean population fitness (average reproductive success across all individuals). For example, the in individuals with urinary or fecal in-
average pelvic floor thickness in a population is expected to be close to the optimal compromise continence and prolapse.60e62 Why did
between pelvic floor stability and parturition. C, Total fitness is again affected by 2 functions, f1 and evolution not compensate for the
f3, but the performance of f3 shows a threshold behavior: individual fitness drops rapidly at a certain biomechanical disadvantage of a large
trait value. For instance, increased birthweight continually increases neonatal survival rate up to the pelvic canal and large pelvic floor by
point where the neonate would be too large to pass the birth canal and, thus, fitness drops to zero increasing pelvic floor thickness? First, a
(without cesarean delivery). D, As a consequence, total individual fitness (f1
f3) is a highly complete compensation of an enlarged
asymmetric function. The evolutionary stable state where population mean fitness is maximized is pelvic canal would require a dispropor-
achieved at a mean trait value smaller than the trait value that maximizes individual fitness. tionate increase in pelvic floor thickness,
Moreover, it includes a fraction of individuals with zero fitness (red shaded area). In other words, the as demonstrated by modeling results.53
high fitness of the individuals on the left side of the fitness threshold is “traded off” against the zero Second, and more importantly, thicker
fitness of the individuals on the right side of the threshold. Hence, the population distribution of muscle tissue requires higher pressure to
birthweight that maximizes average reproductive success in a population necessarily involves a undergo the same amount of deforma-
certain fraction of individuals too large to be born vaginally. tion as thinner tissue. During the second
Mitteroecker & Fischer. Evolution of the human birth canal. Am J Obstet Gynecol 2024.
stage of labor, when the fetal head de-
scends through the birth canal, the pelvic
floor muscles, especially the funnel-
disorders, such as incontinence and limited morphologic variation in pelvic shaped levator ani, stretch up to more
pelvic organ prolapse.25,51e53 Some canal form observable in modern pop- than 3 times their original lengths, and
clinical studies support this hypothesis ulations and the presence of other, clin- the intrauterine pressure that women
by showing that women with a medi- ically more relevant risk factors that are produce during this stage (about 19 kPa)
olaterally wider pelvic canal are more unrelated to pelvic form. Furthermore, presumably is near the upper limit of
prone to develop incontinence and most clinical studies were conducted in what is possible.53,63,64 Even though
prolapse,54e56 whereas other studies did the Global North (Europe, North thicker pelvic floor muscles would in-
not find such associations.57e59 This America, and other high-income coun- crease pelvic floor stability, they may
incongruence presumably owes to the tries); more research in the Global South therefore complicate parturition.42 In

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general, physical exercise and pelvic floor
muscle training in pregnant women have
been shown to strengthen the pelvic
floor muscles but do not lead to longer
labor or affect the mode of delivery.65
Mild exercise even has a protective ef-
fect against incontinence65 and instru-
mental delivery.66 However, female
athletes performing high-impact sports
training over a long period can have
longer and more difficult births, pre-
sumably because of their very strong
pelvic floor muscles.67,68 In a biome-
chanical simulation of the second stage
of vaginal delivery, Li et al69 found that
the relatively thick levator ani muscles of
an athlete required a 45% increase in
peak force to push the fetal head through
the pelvic floor compared with a
nonathlete with a thinner pelvic floor.
Another selective factor, at least on a
global scale, may be environmental
temperature. It is well known that the
size and shape of the body affect its
metabolism and the ability of effective
thermoregulation, in both animals and
humans. The mass of a body increases
proportionally to the third power of
body size, whereas the body surface in-
creases only with its second power. For
instance, doubling the size of an organ-
ism leads to an 8-fold increase in volume
but only to a 4-fold increase in the sur-
face area. As the heat loss of an organism
is proportional to its surface area, this
geometric relationship implies that a
larger body has a smaller surface-to-
volume ratio and hence less heat loss
per unit mass. Accordingly, populations
and species of large-bodied animals are
typically found in colder environments,
whereas those of smaller sizes tend to be
found in warmer regions. This associa-
tion is known as the “Bergmann rule,”
named after the 19th-century biologist
Carl Bergmann.70 Moreover, species or
populations adapted to cold climates
often have stockier body proportions
and thicker limbs than those adapted to
warm climates as this further reduces
the surface-to-volume ratio and conse-
quently heat loss (the “Allen rule,”
named after Joel Asaph Allen71). These
patterns also apply to modern humans.
Populations near the poles, such as
Inuit, Aleut, and Sami people, are on
average heavier and tend to have
shorter limbs and broader trunks than
populations from middle latitudes.72e75
As pelvic form affects the proportions
of the lower trunk, temperature may
also impose natural selection on the
pelvis, at least on its upper part. Indeed,
bi-iliac breadth and the width of the
pelvic canal vary with latitude among
modern humans, with those from
warmer climates having smaller pelvic
breadths than those from colder
climates.7,73,76,77
Cliff-edge model of obstetric
selection
The capacity of the human birth canal
evolved as a compromise to several,
partly antagonistic selective factors, but
these factors are not symmetric in the
way they influence fitness. Medical
literature documents that evolutionary
fitness increases approximately linearly
with increasing fetal size and decreasing
pelvic canal size, as pelvic width corre-
lates with the risk of pelvic floor disor-
ders.33 However, the selection imposed
by childbirth is of truncational nature; if
the fetus is too large to pass through the
birth canal, fitness drops sharply to zero
(here, we assumed that no cesarean de-
livery was performed). Thus, the fitness
function associated with the size of the
fetus relative to the size of the maternal
birth canal (the variable D shown on the
horizontal axis in Figure 4) is highly
asymmetric; it resembles a cliff edge with
a linear increase and a sudden drop (blue
curve in Figure 4). By contrast the dis-
tributions of fetal and pelvic dimensions
in human populations are approximately
symmetric (bell-shaped black curve).
The incongruence of these two differ-
ently shaped functions impedes an
evolutionary resolution of fetopelvic
disproportion. The evolutionary equi-
librium with maximal mean population
fitness necessarily involves a certain
fraction of cases with fetopelvic dispro-
portion. In other words, evolution trades
off the benefits of the many births of
neonates as large as possible but still able
to pass the birth canal against the few
cases of fetopelvic disproportion.
Clearly, neither pelvic canal form nor
fetal dimensions can be properly
MARCH 2024 American Journal of Obstetrics & Gynecology
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described by a single variable. Nonethe-
less, this idealized “cliff-edge model”
describes why childbirth-related selec-
tion could not further reduce cases of
fetopelvic disproportion. Given the
observed rates of disproportion, the
cliff-edge model also allows for a rough
estimation of the strength of selection
acting on the relationship between fetal
and pelvic dimensions. Mitteroecker
et al33 showed that the selection on D, the
size of the fetus relative to the size of the
maternal birth canal, is surprisingly low.
The model suggests that the persistent
rates of fetopelvic disproportion are not
the direct result of strong selection for a
large fetus or a small pelvic canal; it is
primarily the asymmetric, cliff-edged
shape of the fitness function that ac-
counts for the maintenance of these
rates. A weak evolutionary advantage of a
large fetus or a small birth canal could
have been sufficient to inhibit an evolu-
tionary resolution of fetopelvic dispro-
portion in humans.
Hormonal effects on pelvic flexibility
In humans, the pelvic articulations and
especially the pubic symphysis soften
and gain flexibility during late pregnancy
under hormonal influence and prepare
the pelvis for parturition.78 Relaxin is a
peptide hormone present in both sexes,
which plays an important role in this
process as it causes extracellular matrix
breakdown and suppresses the synthesis
of new collagen, leading to increased
joint flexibility. Women possess a higher
density of relaxin receptors in the hip
than men, and serum concentration of
relaxin is highly elevated during preg-
nancy. This leads to increased flexibility
in the female sacroiliac joint and pubic
symphysis during pregnancy, which is
essential for human birth.78,79
However, in many other mammals,
the flexibility of the pubic symphysis is
much greater than in humans, particu-
larly in species that have to accommo-
date relatively large neonates.25 In guinea
pigs, for instance, the mean diameter of
the fetal head at birth is almost twice as
large as the pelvic canal in early preg-
nancy. To accommodate the fetal head,
the pubic bones separate at the pubic
symphysis throughout pregnancy. This
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more flexible in humans. A serious

FIGURE 4
negative side effect of high relaxin pro-
Cliff-edge model duction during pregnancy is pelvic girdle
pain, which is associated with difficulty
A B symmetric in walking.87,88 It has been estimated to
“ ” phenotype dist.
affect approximately 20% to 25% of

prob. density of
pregnant women89 and emerges as a
consequence of the increased flexibility
and the resulting lack of stability in the
pelvis. If the human pubic symphysis
0 0 would open to a degree as seen in several
“optimal“ mean rate of fetopelvic
nonhuman mammals, upright walking
dimensions disproportion would be almost impossible. Empirical
evidence for a dose-response relation-
C ship between serum relaxin concentra-
tion during pregnancy and pelvic girdle

prob. density of
pain is inconsistent,90,91 but as relaxin
acts as an endocrine and paracrine factor,
measured serum levels do not necessarily
reflect the degree of hormone activity.79
0
High relaxin levels can also have other
increased rate
deleterious side effects. In nonpregnant
women, relaxin cycles with other men-
A, Medical studies show that neonatal size positively correlates with infant survival rate, whereas strual hormones and peaks during the
pelvic width correlates with the rate of pelvic floor disorders. Thus, individual female fitness depends female menstrual cycle days 21 to 24.79
on both factors: it increases approximately linearly with the difference, D, between neonatal size and These cyclical relaxin peaks in nonpreg-
maternal pelvic canal size. For D>0, fetal size exceeds the capacity of the birth canal; without nant women have been shown to corre-
cesarean delivery, fitness sharply drops to zero because of fetopelvic disproportion. B, In contrast to late with the risk of musculoskeletal
the “cliff-edged” fitness function, maternal and fetal dimensions—hence also the phenotype D—are injuries, especially anterior cruciate lig-
approximately symmetrically distributed (black curve). By evolution through natural selection, the ament rupture.79,92,93 Another potential
mean phenotype (dashed line) will approach the value that maximizes population mean fitness side effect of high relaxin levels concerns
(average reproductive success). Because of the contrast between the asymmetric fitness function the fetus in utero. Sex differences in the
and the symmetric phenotype distribution, the “optimal” phenotype distribution inevitably entails a number of relaxin receptors in the hip
fraction of individuals with D>0 (red area) and hence with zero fitness because of fetopelvic seem to be present already in the fetus.
disproportion. C, The regular and safe use of cesarean deliveries has removed selection acting Hip dysplasia is much more frequent in
against fetopelvic disproportion and thus has disrupted the previously evolved equilibrium. As se- female neonates than in male neonates,
lection for large neonates and small birth canals (large D values) partly persists, this has led to an and serum relaxin concentrations in the
evolutionary increase of fetopelvic disproportion, according to this model. Mitteroecker et al33 mother have been linked to the risk of
estimated this evolutionary increase of fetopelvic disproportion to be in the order of half a per- hip dysplasia in the neonate.94e97 Hence,
centage point since the mid-20th century (after Mitteroecker et al33). high concentrations of maternal relaxin,
Mitteroecker & Fischer. Evolution of the human birth canal. Am J Obstet Gynecol 2024. excreted to enable birth, not only
compromise pelvic stability and the
ability to walk in the mother but may also
process is accompanied by bone resorp- pocket gopher, a burrowing rodent, the negatively affect the development of the
tion at the symphyseal surfaces of the pelvis forms a complete ring in both fetal hip, especially in female fetuses. The
pubic bones and the growth of an sexes, but in females, the pubes begin to relatively stiff and inflexible human pel-
interpubic ligament, which spans the gap resorb in the first breeding season, with a vic girdle likely is another evolutionary
between the pubic bones.80e83 After complete “pubiolysis” by the time of compromise solution, where bipedality
birth, the guinea pig pelvis approaches copulation. Thereafter, the female keeps does not allow for the evolution of a
its state before pregnancy. The estrogen- her open pelvis for the rest of her more flexible birth canal for parturition.
and relaxin-mediated modifications of life.76,84,85 Concerning pelvic flexibility, bats are
the connective tissue in the pubic sym- In humans, pubic symphyseal flexi- an interesting group. They give birth to
physis that lead to the formation of an bility is more limited. The mean increase the largest neonates relative to maternal
interpubic ligament occur not only in in width of the interpubic gap is only size among all mammals and they are the
guinea pigs but, for example, also in approximately 3 mm.78,86 This raises the only mammals that have evolved pow-
mice, bats, and macaques.25 In the question as to why the pelvic girdle is not ered flight. Bat neonates reach between

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10% and 45% of maternal body mass.

FIGURE 5
The birth of these enormous fetuses is
only possible because the pubic sym-
Pelvic sex differences in humans and chimpanzees
physis in the maternal pelvis opens up
widely, and a ligament forms in between
the pubic bones. The female bat pelvis
then stays open after pregnancy.25 In
some bat species, the pubic ligament can
span up to or even beyond the maximum
transverse diameter of the pelvic ca-
nal.25,98 In contrast, in the male pelvis,
the left and right pubic bones are fused
by synostosis. Bats possess forelimbs that
have evolved into wings, whereas the
functionality of their hindlimbs for
terrestrial locomotion is strongly
reduced. Instead, their hindlimbs are
adapted to serve as hooks to attach to a
surface for roosting head down.99,100
Because bats spend a considerable
portion of their life roosting, they expe-
rience comparatively little and infre- The upper row shows the average pelvis shape in human females and males (middle panels) along
quent pressure on the viscera and the with 2-fold extrapolations to ease the interpretation of these sex differences (left and right panels).
fetus on their pelvic floor. In contrast to The typical pattern of human sex differences is visible: women have a wider subpubic angle, a wider
the human pelvis and the pelvis of pelvic inlet, and a relatively shorter sacrum. The middle row shows the corresponding mean shapes
quadrupedal mammals, the evolution of for chimpanzees; their sex differences resemble the human pattern. To compare the sex differences
the bat pelvis was therefore not limited in humans and chimpanzees independently of the species differences, the bottom row depicts the
by the stability that is required for pattern of sex differences in chimpanzees added to the human sex means after scaling it to the same
terrestrial upright locomotion. Presum- magnitude as the human differences. The visualizations in the bottom row are virtually identical to the
ably, this has opened an evolutionary human sex differences shown in the upper row, reflecting the surprisingly similar pattern of sex
path for bats that was not accessible to differences in the species despite the difference in magnitude (from Fischer et al111).
humans: the evolution of a wide pubic Mitteroecker & Fischer. Evolution of the human birth canal. Am J Obstet Gynecol 2024.
opening and great flexibility in the fe-
male pelvic girdle to enable the birth of
large neonates.25
differences requires differential selection A comparative study of human and
Evolution of sex differences in the in males vs females. If selection (eg, to- chimpanzee pelvises revealed that the
human pelvis ward a wider pelvis) would be acting in two species are identical in their
The pelvis is the skeletal element with the one sex only, the other sex is expected to “pattern” of pelvic sex differences, even
strongest sex differences in humans.43 show a correlated response because the though humans have evolved about
During bone growth in childhood and genetic alleles favored in one sex are also twice the “magnitude” of sex differences
adolescence, the pelvic canal expands expressed in the other.110 Therefore, the as chimpanzees111 (Figure 5). This im-
more in women than in men because of fact that the male pelvis is narrower than plies that pelvic sex differences did not
the higher female levels of estrogen and the female pelvis in many mammalian evolve de novo in modern humans; the
relaxin.101e104 However, pelvic sex dif- species implies that an antagonistic se- last common ancestor of humans and
ferences have been identified in many lective force favoring a narrow pelvis in chimpanzees as well as extinct hominid
other primates and mammals as well. males must exist. This selective pressure species, such as australopithecines and
Generally, primate species with large might either act equally in males and neanderthals, likely have already
neonatal heads relative to the size of the females or be male specific. However, to possessed this pattern. The pattern of sex
maternal pelvic canal exhibit more pro- date, the identity of this selective force is differences in the pelvis is likely even
nounced sex differences.105e109 These still unclear. It could primarily be the much older and at least of early
differences have commonly been inter- biomechanical efficiency during walking mammalian origin because difficult la-
preted as a consequence of obstetric se- and running affecting both sexes equally bor owing to large fetuses can be found
lection, that is, selection acting on or pelvic floor function, which may also in several other species. Pelvic sex dif-
females for an obstetrically sufficient be relevant in males for erectile ferences have been documented for all
pelvis. However, the evolution of sex function.25,37,52 major Eutherian groups (mammals with

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Rotational birth is necessary because of

FIGURE 6
the complex, nonuniform shape of the
Biomechanical analyses of differently shaped pelvic floors birth canal, with its largest dimensions
oriented in different directions in the
A B three pelvic planes. Although the largest
dimension of the female pelvic inlet is

displacement (mm)
anterior

6
typically oriented mediolaterally, the
largest dimension of the outlet is usually

5
found in the anteroposterior direction.
posterior
Between the inlet and outlet lies the

4
narrowest plane of the birth canal, the

3
-1.0 -0.5 0 0.5 1.0
midplane, which is laterally delimited by
posterior
compartment log(AP/ML)
the ischial spines. In physiological ce-
phalic presentation, the fetus aligns the
anterior
compartment
largest dimension of its head with the
largest dimension of each of the three
A, Finite element model of the pelvic floor shown in sagittal view and superimposed on the muscles of planes, leading to a rotational movement
the pelvic floor. B, Results of the loading experiments (total displacement of the pelvic floor model), through the birth canal. In addition, the
showing how the shape of the pelvic floor affects its total displacement, separately for the anterior funnel-shaped levator ani with its sagit-
and posterior compartments, with the total area kept constant. The horizontal axis represents the tally oriented hiatus contributes to the
natural logarithm of the ratio of the AP to ML diameter of the pelvic floor, log(AP/ML). Negative values rotation of the fetal head.16 Presumably,
correspond to mediolaterally elongated shapes, positive values correspond to anteroposteriorly birth would be easier if the birth canal
elongated shapes, and a value of 0 corresponds to a circular pelvic floor with AP¼ML (depicted by was uniformly shaped. Great apes and
the gray ellipses). Displacement peaks for a mediolaterally oval pelvic floor shape, whereas ante- also other monkeys all have uniformly
roposteriorly oval pelvic floors are better in resisting displacement (from Stansfield et al118). anteroposteriorly oval-shaped birth
AP, anteroposterior; ML, mediolateral. canals.115e117 Stansfield et al118 pro-
Mitteroecker & Fischer. Evolution of the human birth canal. Am J Obstet Gynecol 2024. posed that the “twisted” human birth
canal is a consequence of different se-
lective forces acting on different parts of
a placenta), and the pattern of pelvic sex explained by obstetric selection, some the birth canal. Based on biomechanical
differences seems to be similar between examples seem to elude this explanation. models, they argued that an ante-
these groups, with differences concen- Some species possess mild sex differ- roposteriorly oval outlet is advantageous
trated in the pubic region.25,111,112 ences in the pelvis that resemble the for pelvic floor stability because it re-
The pattern of pelvic sex differences is human pattern, even though they are duces stress and strain in the pelvic floor
largely determined by the spatial distri- obstetrically unconstrained (eg, the Vir- compared with a round or medi-
bution of estrogen, androgen, and relaxin ginia opossum, a marsupial where olaterally oval outlet of the same area
hormone receptors in the pelvis and the neonatal mass amounts to only 0.01% of (Figure 6). In contrast, an ante-
hormonally induced bone remodeling maternal mass112). These sex differences roposteriorly oval inlet would require
during development.113,114 As most as- cannot be explained as adaptations for increased spinal curvature to bring the
pects of the endocrine system are highly birth. Fischer et al111 suggested that these center of mass back over the hips
conserved among vertebrates, Fischer differences might be a “vestigial pattern” (Figure 7), which seems to be selected
et al111 proposed that the genetic and inherited from a mammalian or amniote against because of the deleterious effects
developmental machinery producing ancestor, which initially evolved for this would have on spine health and
pelvic sex differences during develop- birthing large neonates. Complete structural stability of upright posture.
ment has also been preserved throughout removal of these differences might be Large spinal curvature and a large
primate evolution. However, the “knob” difficult to achieve evolutionarily lordotic angle have been shown to be
that regulates this machinery is likely because their hormonal induction is associated with back pain, displacement
much more evolvable. By changing the likely tied to other parts of the same of the vertebrae, and disc herniation.
amount and duration of hormone secre- reproductive system. Consequently, the only feasible option to
tion or the number and overall reactivity increase the area of the inlet without
of the corresponding receptors in the Why is the human birth canal increasing the length of the pelvis in the
pelvis, the magnitude of pelvic sex dif- twisted? anteroposterior direction was to evolve a
ferences can evolve quite rapidly while the Human birth commonly involves a mediolaterally wide inlet. A limitation of
pattern remains constant.111 rotational motion of the fetal head the biomechanical models of the pelvic
Although pronounced mammalian through the maternal birth canal, fol- floor by Stansfield et al118 is that they
sex differences in pelvic width are usually lowed by the rotation of the shoulders. were based on isotropic material

S848 American Journal of Obstetrics & Gynecology MARCH 2024

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properties and did not consider the de-
tails of muscle and fiber directions.
Shifting trade-offs
Modern human pelvic form and the
resulting rate of fetopelvic disproportion
are evolutionary compromise solutions
to the multiple, partly opposed selective
forces described above. However, the
actual magnitudes of these selective fac-
tors have changed over prehistoric and
historic time scales, and they also differ
among modern human populations.
Not only temperature varies globally,
also average birth weight varies consid-
erably from approximately 2.7 to 3.6 kg
across human populations,119e121
imposing variable magnitudes of ob-
stetric selection. Similarly, average
neonatal head circumference and gesta-
tion length vary globally.122e125
Maternal stature and pelvic dimensions
also vary across human
populations126e128 and thus alter the
risk of obstructed labor. For instance,
average adult female stature, a well-
known obstetrical risk factor,129e131
varies from 149 to 170 cm across coun-
tries.132 Maternal stature is correlated
not only with the capacity of the birth
canal but also with gestation length and
birthweight via both genetic and epige-
netic pathways.43,133e136
Cultural differences among pop-
ulations may be at least as important for
the local evolutionary dynamics as the
many biological differences. The actual
effect of birthweight on infant survival
and health (and thus the strength of se-
lection for high birthweight) depends on
food availability, hygiene conditions,
maternal care, and the availability of
medical treatment. All these factors have
changed considerably throughout hu-
man history and still differ globally.
Similarly, the effect of pelvic dimensions
on the energetics of locomotion and the
risk of pelvic floor disorders depend,
among other factors, on maternal and
fetal body size and weight, physical ac-
tivities, subsistence strategy, and diet,
which vary across countries as well as by
ethnicity and sociocultural background
and thus expose different magnitudes of
selection on the birth canal.137e139 In
addition, gestational length and meta-
bolic capacity during pregnancy are
affected by nutritional conditions.24,26
Moreover, transitions in environmental
and socioeconomic conditions can affect
the relationship between fetal and
maternal size, thus influencing the
difficulty of labor and the strength
of obstetric selection.30,140e142 For
instance, because of the important role
of maternal nutrition for fetal growth,
rapid improvement of living conditions
in a country can increase average fetal
size and thus inflate the rate of fetopelvic
disproportion.141
Because of all these varying biological,
environmental, and sociocultural fac-
tors, the evolutionary trade-off dy-
namics and the “optimal” compromise
solutions are likely to vary globally and
have led to different pelvic dimensions.
Mitteroecker et al76 referred to this as
“shifting trade-off dynamics.” Morpho-
logical divergence in pelvic form among
human populations has been found to
clearly exceed neutral genetic population
divergence, which is evidence of past
divergent natural selection in human
populations.76
The role of midwifery and obstetrics
Some forms of birth assistance (support
from other individuals during birth) or
midwifery have been present since the
stone age and exist in variable forms in
all human populations to date.1,143e145 It
has been suggested that assistance during
birth became obligatory in humans at
the time when bipedalism evolved,
particularly because of the rotational
birth mechanism in humans.6,10,146
Typically, the baby emerges with the
head extended, in the occiput anterior
position (ie, facing backward). The
mother cannot help the baby emerge, as
is the case in other primates, because this
would mean pulling the neonatal head
backward and potentially injuring the
spine. Therefore, midwifery is important
for human reproduction; it can be
considered an “extended phenotype”
sensu Dawkins.147 Assisted birth is
rather uncommon in other primate
MARCH 2024 American Journal of Obstetrics & Gynecology
Expert Review
FIGURE 7
Relationship between pelvic
depth and spinal curvature
A spine
B
sacrum
pelvis
hip joint
The pelvis and spine are shown schematically in
sagittal view. A, Normal spinopelvic relationship
where the center of mass (indicated by the
vertical dashed line through the last cervical
vertebra, the so-called C7 plumbline) is posi-
tioned sagittally above both the hip joints and the
superior endplate of the sacrum. B, In an
anteroposteriorly elongated pelvis (as indicated
by the red double arrow), the center of mass
must be brought back above the hip joints by a
forward tilt of the sacrum. This leads to an
overall increased curvature of the spine,
particularly an increased lumbar lordosis and a
deviation of the center of mass from the sacral
endplate, which is associated with multiple or-
thopedic disorders, such as spondylolisthesis
and disc herniation (from Stansfield et al118).
Mitteroecker & Fischer. Evolution of the human birth canal.
Am J Obstet Gynecol 2024.
species, although examples have been
documented in species where pair bonds
are formed (eg, in Callicebus, Callithrix,
and Saguinus10). Here, the male some-
times takes an interest in birth but is not
actively involved with helping in the
birth process. Actively assisted birth oc-
curs only in humans.
Because midwifery and obstetrical
care enable successful childbirth despite
a tight fetopelvic fit, these practices have
affected the evolutionary dynamics un-
derlying the human birth canal. This is
an excellent example of gene-culture
coevolution: cultural practices and
genetically determined traits mutually
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Expert Review
influence each other and thus
coevolve.148,149 At the same time, from
an evolutionary perspective, the impor-
tant contributions of midwifery and
obstetrics to human healthcare may have
hindered the evolution of a larger birth
canal. Without the development of
midwifery and obstetrical care, the se-
lective pressure for a wider birth canal
would have been stronger, which may
have led to the evolution of a wider birth
canal and eventually easier childbirth
(presumably at the cost of pelvic floor
stability). In evolutionary biology, this
phenomenon has been dubbed “behav-
ioral inertia” or the “Bogert effect,” after
Charles Bogert, who first popularized
it150e152: behavioral flexibility hinders
evolutionary adaptation by buffering
organisms against natural selection. A
typical example of this effect is thermo-
regulation in desert snakes and lizards,
which are not particularly adapted to
heat. Instead, by actively adjusting their
behavior to the local thermal environ-
ment, such as seeking shelter or shifting
activity to the night, their thermal
physiology has diverged surprisingly lit-
tle from squamate reptiles living in
colder environments. In other words,
behavioral plasticity has enabled survival
in a wide range of thermal environments
without requiring evolutionary adapta-
tion of thermal physiology. Similarly,
behavioral and cultural flexibilities in
humans, including medical treatment
and assisted birth, may have reduced
anatomic and physiological adaptations
in our species.
In many countries of the Global
North, modern obstetrical care and
especially the availability of safe cesarean
deliveries and antibiotic treatment have
sharply reduced maternal and fetal
mortality. Cesarean deliveries have been
performed widely since the 1950s and
1960s, which has effectively removed
childbirth-related selection for a wider
birth canal in these countries. Therefore,
the previously established evolutionary
equilibrium has been disrupted, and new
evolutionary change has been triggered
(Figure 4, C). According to the cliff-edge
model of obstetric selection in humans,33
the remaining selection for large neo-
nates and small birth canals has
S850 American Journal of Obstetrics & Gynecology MARCH 2024
evolutionarily increased the rate of feto-
pelvic disproportion by up to 10% to
20% since the widespread use of cesarean
deliveries in the mid-20th century, which
equals roughly half a percentage point.
Compared to the actual cesarean section
rates in high-income countries to date,
this evolutionary effect is negligibly
small. However, it is remarkable that
medical treatment can induce evolu-
tionary change in human anatomy that is
noticeable in a time range of only de-
cades. Evolution is often considered a
process requiring thousands or millions
of years, but rapid evolutionary changes
within a century or less because of
changing selective pressures are increas-
ingly recognized in different animal
species, also in response to ongoing
climate change.153,154 In the Global
North, the evolutionary increase of feto-
pelvic disproportion likely has slowed
down again because of advancements in
neonatology and in the treatment of pel-
vic floor disorders, which has reduced the
selection for large neonates and small
birth canals.
It is difficult to identify these small
changes in fetal and maternal dimensions
empirically because variance within and
between human populations is substan-
tial and because the expected effect sizes
of this human-induced evolutionary
change are small compared with the ef-
fects of reduced gestational age and more
frequent preterm births. However, the
underlying model has been validated by
successfully predicting the heritability of
fetopelvic disproportion.155 Note that
in this evolutionary context, elective
cesarean deliveries are irrelevant as only
lifesaving interventions matter for evolu-
tionary dynamics. Cesarean deliveries
performed for other reasons do not alter
survival rates.
Discussion
From an individual perspective, several
anatomical, physiological, psychological,
and medical factors contribute to suc-
cessful human labor. Most of these fac-
tors vary to some degree geographically
and among sociocultural groups. It has
recently been argued that anatomic dif-
ferences are of minor importance for the
success of labor and that focusing on
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anatomical features, especially on feto-
pelvic disproportion, distracts from fac-
tors that are much more relevant from a
modern public health perspective.156
Therefore, some researchers have
claimed that there is no “obstetrical
dilemma” in humans.157 It has also been
argued that studying the evolution of
childbirth, especially in modern soci-
eties, is impossible without taking into
account the complex medical, sociocul-
tural, and psychological circum-
stances.158 We agree that anatomical
factors are, statistically and clinically,
only a minor risk factor and that vaginal
birth is a physiological process that
should not be pathologized. Most
women are able to give birth vaginally
when receiving adequate midwifery care;
fetopelvic disproportion affects only a
small percentage of all birthing women.
However, that other factors are more
variable among individuals and, clini-
cally, thus have a larger effect on the
success of labor does not render anat-
omy irrelevant. Most importantly, it
does not imply that human childbirth
and the involved anatomy cannot be
studied from an evolutionary point of
view. Compared with the birth canal of
other primates, the human birth canal
is relatively small and rigid. For many
women, parturition is a relatively long
and difficult process that can lead to
maternal and neonatal morbidity, even
though the birth canal is usually large
enough to encompass the fetus. From a
research perspective, the important
question is whether anatomical aspects
can be studied independently of other
biological and nonbiological aspects to
understand their separate influences.
This depends on the correlation among
these factors in the studied populations.
As a hypothetical example, if women
with a small birth canal would tend to
give birth in private hospitals or on
Friday afternoons (when cesarean de-
liveries are more common for nonbio-
logical reasons), these biological and
cultural factors could not be studied in
isolation because of their cooccurrence.
But, as such correlations are highly
unlikely, it is feasible to study these
traits and their evolutionary dynamics
independently despite the fact that
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Glossary
Amniote: A group of 4-limbed vertebrates that include reptiles, birds, mammals, and the extinct ancestors of these groups. They lay
eggs on land or retain the embryo within the mother. Unlike fishes or amphibians, they can reproduce independently of water because
they possess a membrane (amnion) that protects the embryo. Amniotes have a long, continuous embryonal and fetal development
without a larval stage.

Australopithecus (Latin: “southern ape”) is an extinct genus (taxonomic unit) of early hominins that existed in Africa from approxi-
mately 4.2 to 2.0 million years ago. Australopithecines were able to walk upright, but their brain volume was only slightly larger than that
of chimpanzees (approximately one-third of the modern human brain size). Their body height ranged from approximately 1.1 to 1.5 m,
with males being significantly taller than females.

Bipedalism: Bipedal animals move by walking on 2 legs. Bipedalism evolved independently several times, for example, in ostriches,
kangaroos, and in certain dinosaurs. Humans are obligatory bipeds, but also several other primates, for example chimpanzees, show
occasional bipedal locomotion. The evolution of bipedalism in the human lineage occurred approximately 4 to 6 million years ago and
was accompanied by anatomical adaptations in the entire body, especially in the legs, pelvis, spine, and cranium.

Hominid and hominin: Hominids are a taxonomic group consisting of all modern and extinct great apes (humans, chimpanzees,
bonobos, gorillas, and orangutans and all their immediate ancestors). By contrast, hominins are all species in the human lineage that
emerged after the evolutionary split from the chimpanzee lineage. This group consists of modern humans and extinct human ancestors,
including the genera Homo, Australopithecus, Paranthropus, and Ardipithecus.

Homo is the genus that includes modern humans (Homo sapiens) and several extinct species classified as either ancestral to or closely
related to humans. The genus emerged with the appearance of Homo habilis approximately 2.3 million years ago. Homo erectus (which
lived approximately 2 million to 100,000 years ago) already had relatively modern gait and body proportions and was the first human
ancestor to spread throughout Eurasia. Homo erectus is an ancestor of both Homo sapiens and Homo neanderthalensis (neanderthals).
The oldest evidence of anatomically modern humans in Africa dates back to approximately 300,000 years.

Fitness: In most evolutionary contexts, the fitness of a certain genotype or phenotype is defined as the average number of offspring
(average reproductive success) of individuals with this genotype or phenotype. It also equals the average contribution of individuals with
this genotype or phenotype to the gene pool of the next generation. Through natural selection, genotypes or heritable phenotypes with
relatively high fitness tend to become more frequent in a population, which increases the “mean fitness” in the population. Evolutionary
theory predicts that a heritable trait evolves in a population until it reaches maximum mean fitness; then, evolution comes to a halt
(“evolutionary equilibrium”) unless it is constrained or disturbed by other influences (eg, drift or environmental changes).

Fitness function: For a continuous trait, such as body height or pelvic width, the fitness function describes the effect of the trait value
on the fitness value (Figures 3 and 4). Fitness functions are commonly used tools in quantitative evolutionary models.

Selection occurs if phenotypes are associated with different fitness values (different average reproductive success). For a continuous
trait, this implies that the fitness function is not flat but instead is sloped, and the strength of selection corresponds to the steepness of
that slope. Strong selection (large differences in fitness values) leads to a rapid phenotypic change in a population because of a fast
increase in the frequency of phenotypes with high relative fitness.
Total individual fitness is affected by many components, such as fecundity, age-specific survival rates, and life span. Thus, a trait can
influence fitness in several ways simultaneously. A trait is subject to “antagonistic selection” if it has a positive effect on one fitness
component but a negative effect on another component. For example, a large pelvic canal eases childbirth but, at the same time,
increases the risk of pelvic floor disorders and may have biomechanical disadvantages. Therefore, it is subject to antagonistic selection
(Figure 3).

childbirth is influenced by so many biocultural systems or processes as long For evolution by natural selection to
different biological and nonbiological as we know how to dissect them into occur, it is required that different trait
factors at the same time.159 In other largely independent or “near-decom- values are associated with different
words, it is possible to study complex posable”160,161 parts. fitness values (ie, the average

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Expert Review
reproductive success associated with a
trait value) and that the trait is heritable.
Although biologists are accustomed to
the fact that the statistical association
between a heritable trait and evolu-
tionary fitness is difficult to demonstrate
for many species, the situation is
different for humans: clinical and
epidemiologic studies offer a wealth of
information that allows one to infer how
traits are linked to fitness. Nevertheless,
it can be challenging to infer the effect of
a disorder on fitness under past living
conditions based on modern health im-
pairments. For instance, incontinence
and pelvic organ prolapse often
compromise social and sexual well-
being, but they are not lethal in the
Global North. By contrast, in sub-
Saharan Africa chronic incontinence,
often resulting from obstetrical fistulas,
is still a serious health threat to
women and can lead to social
ostracization.162e164 Moreover, inconti-
nence and prolapse predominantly affect
women after their reproductive period,
but both conditions also occur in
younger women.137,165e168 As the risk of
pelvic floor disorders is correlated with
the dimensions of the pelvic canal, these
syndromes likely have imposed natural
selection on the human pelvic form in
the past and still do so in countries where
no or insufficient medical care is avail-
able. For modern clinical practice, and
for the treatment of individual patients,
pelvic form may be a risk factor of minor
importance, but on the population level,
even a weak correlation with evolu-
tionary fitness is sufficient to impose
natural selection.
However, the presence of natural se-
lection in modern populations, as re-
flected by epidemiologic data, does not
necessarily imply that the same selective
force existed in the past or that it was
responsible for shaping the same traits in
early human evolution. For instance, it
likely was the efficiency of bipedal loco-
motion, not pelvic floor stability, that
drove the restructuring of the pelvis in
early hominin evolution. However, in
late human evolution, the selection
imposed by pelvic floor functionality
presumably hindered the pelvic canal
from evolving to be larger. The
S852 American Journal of Obstetrics & Gynecology MARCH 2024
disadvantages of a large pelvic canal
could, in turn, be compensated by the
evolution of stronger pelvic floor mus-
cles, but this would have consequences
for both parturition and continence.
Hence, functional and evolutionary in-
terpretations of isolated traits can be
misleading: many maternal, neonatal,
and cultural traits show tight mutual
dependencies and thus coevolve. Medi-
cal disorders often reflect “uncompen-
sated” trait variants that disrupt the
evolved integration of traits, but they do
not necessarily reflect the absolute limits
of evolution.
We showed that the human birth ca-
nal has evolved in response to several,
partly antagonistic selective forces.
Because of the asymmetric fitness func-
tion related to the relative size of the
birth canal, evolutionary theory predicts
that the evolved compromise necessarily
involves a certain rate of fetopelvic
disproportion in humans. This antago-
nistic selective regime may have started
to emerge with upright walking and was
aggravated with the evolutionary in-
crease of brain size in the Homo lineage.
Therefore, not only modern humans but
also earlier hominins likely experienced
a tight fetopelvic fit and difficult child-
birth, which is supported by recent
modeling approaches.16 Sociocultural
factors, including lifestyle, nutrition,
midwifery, and modern obstetrical
practices, are important components
contributing to these complex evolu-
tionary dynamics. Over many thousands
of years, midwifery practices have
become an integral part of the human
phenotype: they have enabled human
childbirth but, at the same time, may
have counteracted the evolution of a
wider birth canal. More recently, the
regular use of lifesaving cesarean de-
liveries likely has led to the disruption of
the previously established evolutionary
equilibrium underlying the dimensions
of the human birth canal and thereby
affects the future evolutionary trajectory
of modern human anatomy.
Environmental change and sociocul-
tural transitions in human history have
repeatedly affected the evolutionary dy-
namics underlying human childbirth in
our past. As environmental and
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sociocultural conditions still vary on a
global scale, the tightness of the feto-
pelvic fit and the difficulty of labor are
also likely to vary between human pop-
ulations today. In part, these population
differences contribute to the large global
variation in cesarean delivery
rates.30,140e142,169 Understanding the
evolutionary and biocultural dynamics
of the human birth canal helps us to
predict how medical advancements and
changing living conditions affect human
childbirth in the future, which, in turn,
may help to inform public health and
research strategies. -
ACKNOWLEDGMENTS
We thank our colleagues Nicole Grunstra,
Mihaela Pavlicev, and Katya Stansfield from the
University of Vienna and the Pelvis and Evolution
group, headed by Engelbert Hanzal from the
Medical University of Vienna, for many years of
fruitful discussion and collaboration. Further-
more, we thank the 6 reviewers and Roberto
Romero for many helpful comments.
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