American J Phys Anthropol - 1992 - Rosenberg - The Evolution of Modern Human Childbirth

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YEARBOOK OF PHYSICAL ANTHROPOLOGY 3589-124 (1992)
The Evolution of Modern Human Childbirth
KAREN R. ROSENBERG
Department of Anthropology, University of Delaware,
Newark, Delaware 19716
KEY WORDS Obstetrics, Pelvis, Childbirth, Human evolution
ABSTRACT Human birth follows a pattern which is unique among

mammals. Distinctions include the orientation of the fetus as it passes
through the birth canal, the way the fetus emerges from the birth canal,
difficulty during labor, and behavior by the mother andlor other individuals
around the time of birth. Birth has important implications for the morphol-
ogy of the pelvis, for sex differences in the pelvis, for such aspects of human
biology as size (and maturity) a t birth, and for behavior (including cooper-
ative behavior). This paper reviews the fossil and comparative evidence for
when and how the modern pattern of birth evolved.
The modern human pattern of birth evolved in a mosaic manner with
some unique features appearing early in human evolution and others quite
late. A human-like entry of the fetal head into the birth canal was already
present among australopithecines as a result of their wide pelvic apertures.
Other aspects of modern human birth such as the rotation of the head and
body within the birth canal and the emergence of the fetal head in an
occiput anterior position probably evolved later, when encephalization had
placed increasing selection on both the form of the pelvis and the timing of
birth. Cooperative behavior during and after birth accompanied the origin
of the fully modern human mechanism of birth.
The unique phenomenon of modern human birth did not evolve in re-
sponse to a single “obstetrical dilemma” but as part of a complex interplay
between changes in a number of aspects of human biology.
0 1992 Wiley-Liss, Inc.
Birth in modern humans is a process unlike that seen in any other living species.
This fact results from the interaction of several features of our evolutionary his-
tory, such as bipedal locomotion and posture, encephalization, and secondary al-
triciality. In turn, the human pattern of birth has profound implications for many
aspects of morphology and behavior in such areas as locomotor efficiency, infant
dependency a t birth, and possibly even cooperative behavior during birth. This
paper surveys what is currently known about when and how the human pattern of
parturition arose.
The idea that encephalization and bipedalism have combined to make parturi-
tion a difficult and laborious process for human females was discussed by Krogman
(1951), who called human birth one of the “scars of human evolution,” and Wash-
burn (1960), who described an “obstetrical dilemma” resulting from the conflict
between selection t o increase the size of the brain (and, therefore, the size of the
birth canal) and selection to decrease the overall breadth of the pelvis for locomotor
efficiency. In the last few decades, fossils have been discovered from several time
periods which have tremendously expanded our knowledge of the evolution of
0 1992 Wiley-Liss, Inc.

10968644, 1992, S15, Downloaded from https://onlinelibrary.wiley.com/doi/10.1002/ajpa.1330350605 by CAPES, Wiley Online Library on [21/02/2023]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
90 YEhRBOOK OF PHYSIChL ANTHROPOLOGY [Vol. 35, 1992
hominid pelvic morphology, while allowing us to infer how this “dilemma” was
resolved and provoking some intriguing suggestions about the evolution of human
childbirth. In spite of the increase in information and understanding of the unique
nature of birth in humans, this subject generally receives little attention in con-
siderations and summaries of the human fossil record such as those in introductory
textbooks.
This paper focuses on those aspects of birth which are discernible from the fossil
record. Therefore, it will not deal with such important phenomena as the role of the
placenta in parturition and the hormonal determinants of labor. Information on
the evolution of these phenomena can be gained only from a comparative study of
extant humans and other primates. Because this paper deals with parturition-
related morphology which is recoverable from the fossil record, the concern is
primarily with the skeletal morphology of the maternal pelvis and to a lesser
extent with the neonatal crar,iun;, as the latter is rarely fossilized. In addition, the
discussion will include aspects of birth which might be inferred from the fossil
record, such as behavior during parturition. This paper has two parts. The first
section reviews birth in living human and nonhuman primates and describes the
unique nature of human birth; the second section examines the fossil record to
determine how and when the characteristic pattern of human birth evolved.
Although obstetrically significant anatomy of the pelvis is emphasized here,
obstetric factors are far from the only selective forces affecting pelvic morphology.
Such factors as locomotion, posture, visceral support, and climate also influence
the form of the pelvis (Abitbol, 1987b, 1988; Lovejoy, 1973,1975,1988; Lovejoy et
al., 1973; Rak, 1990,1991; Reynolds, 1931; Reynolds and Hooton, 1936; Ruff, 1991;
Steudel, 1984; Susman et al., 1984). Selection resulting from these factors must
interact with obstetrical selection to produce the pelvic form characteristic of hom-
inids.
BIRTH IN LIVING HUMAN AND NONHUMAN PRIMATES
Birth in all mammals is associated with considerable risk of mortality for both
infants and mothers. Physiological, morphological, and behavioral factors all de-
termine the outcome of this process. One of the most important of these (and the
only one which can be addressed directly by an examination of skeletal morphol-
ogy) is the relationship between the size and shape of the maternal birth canal and
the size of the fetal cranium. A relationship between these dimensions which does
not allow sufficient room for successful passage of the fetus can result in death or
damage to the infant and/or to the mother. Both situations affect reproductive
success.
How is modern human birth differentfrom birth in other living primates?
In general, birth in humans is distinguished from birth in nonhuman primates
as well as other mammals in three respects: 1)the mechanism of birth, specifically,
the course that the fetus must follow as it passes through the maternal birth canal
and its orientation during that passage; 2) the duration and degree of difficulty of
labor; and 3) behavior during and after labor (including such considerations as
whether the birth is attended by other individuals).
Bo (19711, Brandt and Mitchell (19711, and, more recently and most thoroughly,
“revathan (1987) have reviewed the available evidence on primate parturition.
Data on this subject are difficult to obtain because primates, like many other
mammals, usually give birth in isolation where and when (Jolly, 1972, 1973;
Napier and Napier, 1986; Rawlins, 1979) they will be least vulnerable to predators
and least visible to primatologists.1 Since the act of parturition is generally quick
‘In both zoo and field settings, diurnal s ecies generally give birth at night, while nocturnal species tend to
give birth during the day. This su gests tRat time of birth in primates is an adaptation to avoiding normal
predators and/or attention from otier members of one’s own species. Human birth shows evidence of being
subject to the same selective pressures (Jolly, 1972).
Rosenbergl EVOLUTION OF BIRTH 91
in nonhuman primates and most other mammals, the probability of an observer

witnessing the birth is small. In addition, because observations are often fortuitous
rather than systematic, they often occur under unusual or unnatural conditions so
that many aspects of birth which might be quantified (such as duration of labor)
can only be subjectively reported.
Mechanism of birth
“Mechanism of birth” refers to the “way the baby adapts itself to and passes
through the maternal pelvic” (Oxorn, 1986: 94). This includes the passage of the
fetus through the birth canal, the orientation of the fetal head as it enters the
pelvic inlet, changes in orientation of the fetal head and body during the passage,
and the orientation of the fetus as it emerges from the mother’s birth canal.
Passage through the birth canal. Nonhuman primates. The birth canal in non-
human primates consists of a relatively straight tube which is greater in anterior-
posterior diameter than in transverse diameter at all planes (Abitbol, 198713;
Broek, 1914; Hager, 1989; Leutenegger, 1974; Schultz, 1949, 1969; Tague, 1991,
199213; Washburn, 1942). The fetal skull in primates is always longer in the sag-
ittal than in the biparietal diameter (Abitbol, 1991; Leutenegger, 1974, 1982b;
Schultz, 1949).
Among most nonhuman primates, birth is comparatively simple. Passage
through the birth canal rarely involves significant changes in orientation of the
fetus (Trevathan, 1987). For example, in nonhuman primate species in which the
fetal cranium is close to the size of the female pelvic aperture (small cebids, and
probably many cercopithecids), the head enters the inlet in a dorsoflexed position
so that the sagittal diameter of the skull which must pass through the shortest
diameter of the pelvic inlet is facial height rather than the longer cranial length
(Leutenegger, 1974,1982b). In other species, such as chimpanzees, the birth canal
is quite spacious relative to the neonatal head (Schultz,1949).The head of the fetus
aligns itself so that its longest diameter (the sagittal diameter) is along the sagittal
diameter of the pelvic inlet and proceeds through the birth canal without changing
that orientation and with no bony resistance (though soft tissue resistance may
cause cranial flexion) (Tague and Lovejoy, 1986).
Living humans. In humans, the passage of the fetus through the birth canal is
more tortuous. The normal mechanism of labor involves a number of movements
and alterations of the position of the head, often called the “cardinal movements of
labor” (Cunningham et al., 1989; Oxorn, 1986; Percival, 1980; DB Stewart, 1984bi.
Labor in humans is traditionally divided into three stages by clinicians (Baird,
1962; Cunningham, et al., 1989; Oxorn, 1986; Percival, 1980; Pritchard and Mac-
Donald, 1980). The first stage of labor is the period of effacement (thinning) and
dilation (opening) of the cervix. The second stage, “expulsion of the fetus,” is the
period from complete dilation of the cervix to the delivery of the infant. Finally, the
third stage is “expulsion of the placenta.” Clearly, the second stage of labor is most
relevant to cephalo-pelvic (fetal cranial-maternal pelvic bone size) relationships
and is, therefore, of most importance to this study. This stage will be outlined here.
Clinical data on living humans indicate that when labor begins, 99.5% of fetuses
lie along a longitudinal axis (Oxorn, 1986). That is, the long axis of the fetus is
parallel to that of the mother’s body. The great majority (96%)of all fetuses are in
a vertex presentation (with the head positioned over the pelvic inlet). The descrip-
tion which follows is for the most common fetal presentation in humans: the “left
occiput anterior position,” that is, with the fetal occiput and vertebral column on
the anterior and left side of the mother’s body (Oxorn, 1986). Figure 1 shows a
human fetus passing through the mother’s birth canal.
Engagement of the fetal head consists of the passage of the fetal biparietal
diameter into the maternal pelvic inlet. Typically, the fetal head is oriented so that
its longest axis (anterior-posterior or sagittal diameter) is aligned either along the
92 YEARBOOK OF PHYSICAL ANTHROPOLOGY Wol. 35, 1992
Fig. 1. The stages of labor in humans. In each box, a sagittal section through the maternal body during
labor is shown. Anterior is at the top of each picture and inferior is to the right. The maternal pubic bone
and vertebral column are shown in black. In the upper right hand corner of each box is a “midwife’seye”
view of the fetus as it rotates within and emerges through the birth canal.
transverse diameter of the maternal pelvic inlet or obliquely (Cunningham et al.,

1989; Kadar, 1985). Figure 1B shows the head engaged slightly obliquely to the
transverse diameter of the pelvic inlet.
Descent of the fetal head is continuous throughout labor and is brought about by
the downward force of uterine contractions and by muscular bearing down by the
mother.
Flexion of the fetal head may occur before the onset of labor. During descent, the
head meets resistance from the walls or floor of the pelvis or from the cervix, and
flexion occurs. Because the occipito-frontal diameter of the fetal head is larger
than the suboccipito-bregmatic diameter by as much as 2.5 cm (Cunningham et al.,
1989), the head flexes, bringing the chin into contact with the thorax, so that the
smaller of these diameters becomes the sagittal diameter of the cranium (Fig. 1B).
Internal rotation of the fetal head occurs when the head passes from the pelvic
inlet into the midplane (Figs. 1C,D). Because the longer axis of the midpelvis is the
anterior-posterior diameter rather than the transverse diameter, the head must
turn internally to align itself to the longest axis of the pelvic midplane. The fetus
is usually facing dorsally in the birth canal at this stage with its occiput against
the dorsal surface of the maternal pubis. The shoulders do not rotate, but continue
to be oriented transversely or obliquely, so that the fetal neck is twisted by as much
as 90 degrees.
Extension of the fetal head takes place after the head has rotated, so that its
sagittal axis is aligned with the sagittal axis of the maternal pelvis (Fig. 1E). This
brings the base of the occipital bone into contact with the pubic symphysis and the
neonatal face emerges from the birth canal below the subpubic angle.
Restitution occurs when the fetal head reaches the pelvic floor and the shoulders
enter the pelvis. When the head has emerged from the birth canal, the head twists
back 45 degrees so that it is in a normal relationship to the shoulders (Fig. 1F).
External rotation of the neonatal head (Fig. 1G) is the result of internal rotation
of the shoulders. The shoulders rotate to be aligned in the sagittal plane of the
pelvic outlet so that they can emerge under the pubic symphysis and the head
rotates passively with them.
Expulsion (Fig. 1H) occurs when the first (anterior) shoulder emerges from un-
der the pubic symphysis followed by the second (posterior) shoulder and remaining
portion of the fetal body.
Following the birth of the head and shoulders, the postcranial portion of the
fetus (which is not as wide as the cranium or shoulders) is born quickly and easily.
Presentation of the fetus. A key aspect of mechanism of birth is the presentation of

the fetus. This refers t o that portion of the fetus which enters the pelvic inlet and
emerges from the birth canal first.
Nonhuman primates. The most common presentation in nonhuman primates
is a vertex presentation (head first) with the occiput posterior (Bo, 1971; Bowden
et al., 1967; Trevathan, 1987). The occiput posterior position has been observed in
the slender loris (Kadam and Swayamprabha, 19801, common marmoset (Rothe,
1974, cited in Trevathan, 1987; Stevenson, 19761, squirrel monkey (Hopf, 1967),
colobus monkey (Wooldridge, 19711, Java macaque (Kemps and Timmermans,
1982), chacma baboon (Gillman and Gilbert, 1946, cited in Trevathan, 1987), and
orangutan (Graham-Jones and Hill, 1962, cited in Trevathan, 1987). In other
cases, observers note that the presentation was vertex, without being more specific
about the position of the occiput. This is the case for the squirrel monkey and mona
monkey (Takeshita, 1961-21, howler monkey (Sekulic, 19821, hamadryas baboon
(Abegglen and Abegglen, 19761,olive baboon (Love, 19781,rhesus macaque (Mitch-
ell and Brandt, 1975; Rawlins, 1979), langur (Brockman and Lippold, 1975;
Oppenheimer, 1976), pygmy chimpanzee (Bolser and Savage-Rumbaugh, 1989),
and gorilla (Beck, 1984; Fisher, 1972;Lotshaw, 1971; Nadler, 1974;Stewart, 1984).
A vertex presentation apppears to have an adaptive significance in primates and
94 YEARBOOK OF PHYSICAL ANTHROPOLOGY [Vol. 35, 1992
other mammals. Schultz (1969) noted that “monkey babies are normally born head
first, so that there is no danger of the umbilical cord becoming clamped off between
foetus and pelvis before breathing can begin” (Schultz, 1969:153).
Breech presentations, though mentioned in some of the descriptions of primate
birth, are rare. Mitchell and Brandt (1975) suggested that in macaques, breech
births usually result in infant mortality. Breech births have been observed in a
feral anubis baboon, resulting in the death of the infant (Nash, 1974); in a suc-
cessful birth in a mantled howler monkey (Moreno et al., 1991); and in several
cases in which the infant was stillborn, namely, squirrel monkey (Bowden et al.,
1967), Java macaque (Kemps and Timmermans, 19821, and rhesus macaque
(Brandt and Mitchell, 1971, 1973; Mitchell and Brandt, 1975). In addition, breech
deliveries have been noted in slender loris (Kadam and Swayamprabha, 19801,
common marmosets (Rothe, 1974, cited in Trevathan, 19871, and rhesus macaques
(Gibber, 1986). As well as being dangerous t o the fetus, breech deliveries present
risks to the delivering mother. Although quantitative data are not available,
breech deliveries in nonhuman primates are probably associated with increased
mortality for both mothers and their offspring (as they are in humans).
Living humans. In humans, as mentioned above, the vast majority of fetuses
are oriented with the head down (a vertex presentation) and engaged in the pelvic
inlet a t the time labor begins. Following internal rotation, the head of the fetus
usually emerges from the birth canal with the occiput anterior (see Fig. 1E).
Although an occiput anterior delivery is not a universal emergence pattern,
Trevathan (1988:674) argues that it is “optimal for distribution of forces of uterine
contraction along the spine and for cervical dilation.” She points out, however, that
this presentation is disadvantageous as it makes it difficult for the mother to “meet
the immediate needs of the infant after the head is born” (Trevathan, 1988: 674)
such as moving the umbilical cord if it is wound around the infant’s neck, lifting
the infant towards her safely, or clearing the infant’s breath passage soon after the
head emerges from the birth canal.
In summary, the unique aspects of the mechanism of birth in humans are: 1)the
transverse or oblique position of the fetal head as it enters the maternal pelvic
inlet; 2) the rotation of the fetal head as it passes from pelvic inlet to midplane; and
3) the emergence of the fetal head from the birth canal in an “occiput anterior”
position.
Species differences in degree of difficulty during labor
Morphology. Schultz (1949, 1969) and Leutenegger (1974, 1982b) examined the
relationship between the size of the maternal pelvis and the size of the fetal head
in a range of primate species. Schultz noted that ease of delivery is dependent on
this relationship but showed that there was no consistent relationship between
these variables among primate species (Schultz, 1949). In a now classic figure
(redrawn here as Figure 2) he showed that monkeys and humans have a close
relationship between the size of the fetal head and the size of the maternal pelvic
inlet, while in the great apes the pelvis is much more spacious relative to the fetal
head (see Table 1).This relationship is dependent on 1)the overall body size of the
infant relative to the size of the mother, and 2) the degree of encephalization of the
infant at birth (Leutenegger, 1974). In nonhuman primates, the relationship be-
tween maternal and fetal body weight is negatively allometric so smaller bodied
species tend to have relatively large infants a t birth. In addition, newborns of
smaller-bodied primate species tend to have larger brains relative to body weight
than larger-bodied primates (Leutenegger, 1973b; Fig. 9 in Martin, 1983).
Bowden et al. (19671, Leutenegger (1982a,b),and Lindburg (1982) also observed
that variation among primate species in size of the infant’s cranium relative to the
mother’s pelvic inlet is correlated with difficulty during parturition. In callitri-
chids, cebids (except the howler monkey), cercopithecids, hylobatids, and humans,
the maternal pelvis is only slightly larger than the infant cranium if at. all, and
TABLE 1. Auerage transverse diameter of female peluic inlet, newborn cranial breadth, and relative
size of newborn cranium for primate species
Adult females Newborns
Transverse
diameter Cranial Relative
of inlet breadth cranial
Species N (mm) N (mm) breadth'
Callithrix jacchus 15 17.2 4 18.0 104.5
Saimiri sciureus 7 23.1 3 28.0 121.1
Cebus capucinus 4 35.1 2 43.0 118.2
Alouatta uillosa 2 47.0 1 38.0 80.8
Lagothrix lagothricha 4 44.3 1 48.0 108.3
Ateles geoffroyi 7 54.4 8 52.2 95.9
Macaca mulatta 41 50.9 28 50.7 99.6
Nasalis laruatus 15 51.8 1 49.0 94.6
Hylobates lar 87 55.9 6 52.7 94.3
Pongo pygmaeus 26 102.5 4 74.9 73.1
Pan troglodytes 29 98.0 9 71.0 72.4
Gorilla gorilla 10 122.6 4 79.0 64.4
Homo sapiens 10 121.6 10 123.8' 101.8
'Relative cranial breadth is newborn cranial breadth as a percentage of the transverse diameter of the maternal pelvis.
Index is calculated for the means of each species.
'Cranial length rather than breadth is included here, since that is the dimension that passes through the transverse
diameter of the inlet in humans.
From Leutenegger, 1982b.
birth is difficult, often involving dilation of pelvic ligaments and deformation of the
infant cranium. Infant mortality in these species is reported to be high (Leuteneg-
ger, 198213). Marmosets and squirrel monkeys in particular show a close correla-
tion between the size of the mother's pelvis and the size of the neonatal head2
(Leutenegger, 1982b). In fact, as mentioned above, in these animals the length of
the neonatal cranium is often greater than the sagittal dimension of the maternal
pelvis (see Table 1)and they are born with a face presentation (Trevathan, 1987).
Great apes have neonatal cranial dimensions which are small relative to maternal
pelvic aperture size. These relative dimensions are probably a consequence of the
large body size of these animals (Leutenegger, 1973c, 1974) and permit a quick and
easy delivery (Leutenegger, 1982b). Within the great apes, gorillas have a wider
pelvic inlet relative to neonatal head breadth than chimpanzees and orangutans.
Galdikas (1982) suggested that this may account for the relative ease of gorilla
birth compared to the other great apes.
Table 1 shows average pelvic dimensions of adult females and cranial dimen-
sions of newborns for a number of anthropoid primates. Note that with the excep-
tion of howler monkeys (see below), fetal crania are relatively large in monkeys
and relatively small in apes. Humans are more similar to monkeys than to apes in
the relationship between these variables.
Because selection resulting from parturition acts directly only on females, many
researchers have suggested that sexual dimorphism in the pelvis is a reflection of
differential selection on the two sexes (Black, 1970; Gingerich, 1972; Hager, 1989;
Leutenegger, 1973d, 1974, 1982a; Lovejoy, 1973, 1975; Lovejoy et al., 1973;
Schultz, 1949; Tague, 1989,1991; Tague and Lovejoy, 1986; Wood and Chamber-
lain, 1986). Schultz (1949) and Leutenegger (1974) hypothesized that the degree of
sexual dimorphism in the pelvis of a species is an indication of the intensity of
obstetrical selective pressure. This follows from the fact that while the pelvis in
both sexes is under selection due to locomotion which would favor a narrow pelvis,
'Goss e t al. (1968)is often cited as evidence of a 50%infant mortality rate in squirrel monkeys. This is almost
certainly a n overestimate. That paper reports on a small sample 118 females) of captive squirrel monkeys which
were subjected to frequent abdominal surgery for removal of embryos for research. Even if this estimate of infant
mortality is too high, it appears to be the case that birth in squirrel monkeys is more difficult (based on
observations) and may be riskier than in most other primates.
Fig. 2. The relationship between the size of the maternal pelvic inlet and the size of the infant’s head
in a number of primate species. Maternal pelvic and infant cranial outlines are diagrammatic, but scaled
so that the transverse diameter of all of the maternal pelvic inlets are the same. The outlined rectangle
represents the maternal pelvic inlet and the black rectangle represents the infant’s cranium. Note that
in the monkeys and gibbon, the dimensions of the infant cranium are only slightly smaller than the
dimensions of the mother’s pelvis. In great apes, the pelvic inlet is relatively spacious. In humans, the
infant cranium is actually longer than the anterior-posterior dimension of the pelvic inlet, requiring the
head to rotate in order to pass through. (After Schultz, 1949).
the female pelvis is additionally under selection favoring a wide birth canal and
hence a wider pelvis.
In an attempt to measure the osteological effects of differential selection on
human males and females, Tague (1989) tested a hypothesis proposed by Meindl et
al. (19851, who suggested that because selection differs in the two sexes, female
pelvic variability should be reduced relative to male. Tague (1989) found no sig-
nificant (or even systematic) differences in variation of pelvic metrics between
males and females. Nevertheless, in humans, visual cues of sexual dimorphism
(such as those used in Phenice’s (1969) method of sexing the pubis) are more
variable in males than in females (Meindl et al., 1985). Tague (1989) suggested
that sex differences in pelvic morphology may be the result of sex steroid influ-
ences on pelvic growth rather than directly genetically based.
Table 2 contains average ischio-pubic index [(pubiclengthlischial length) x 1001
for male and female primates and a measure of sexual dimorphism in that index.
Leutenegger (1974,1982a) and Schultz (1949) suggested that the greater the ceph-
alo-pelvic ratio, the greater will be the degree of pelvic sexual dimorphism. The
ischio-pubic index expresses the elongation of the pubis (and presumably of the size
of the pelvic aperture) relative to body size (this assumes that ischial length is
highly correlated with body size). Sexual dimorphism is calculated as female mean/
male mean x 100. Note that an index of sexual dimorphism calculated from the
TABLE 2. Relative elongation of the pubic bone lischio-pubic index) and sexual dimorphism i n that
index in living primates
Female Male
ischio-pubic ischio-pubic Sexual
SDecies index’ index dimomhism2
Perodicticus potto 188.7 183.3 102.9
Arctocebus cabalarensis 156.8 146.6 106.9
Callithrix jacchus 93.9 84.7 110.9
Saimiri sciureus 92.9 73.0 127.3
Cebus capucinus 90.8 75.8 119.8
Alouatta villosa 115.3 111.3 103.6
Lagothrix lagothricha 112.9 98.7 114.5
Ateles geoffroyi 100.8 89.9 112.3
Macaca mulatta 95.7 82.9 115.4
Nasalis laruatus 96.9 86.4 112 1
1ij.lvbutro lur 1Ul.ti 92.7 109.6
Pongo pygmaeus 107.0 97.8 109.4
Pan troglodytes 87.0 82.6 105.3
Gorilla gorilla 98.3 93.6 105.0
Homosapiens 95.0 79.9 118.9
‘The ischio-pubic index is the length of the pubic bone divided by the length of the ischium x 100.
‘Sexual dimorphism is here calculated as the female mean divided by the male mean x 100.
From Leutenegger, 1973, 1974.
ischio-pubic index is a reflection of the differential elongation of the pubic bone in

females relative to males. In this measure of sexual dimorphism, females exceed
males (the percentage sexual dimorphism exceeds 100) in all species.
Theoretically, the tighter the fit between the neonatal head and the maternal
pelvis, the greater the specialization of the female pelvis to parturition and the
greater the degree of sexual dimorphism in the ischio-pubic index. Leutenegger
(19738 interpreted the lack of sexual dimorphism in the ischio-pubic indices of
African lorises as evidence of the lack of overall body size dimorphism as well as
the “absence of selective pressure for expansion of the birth canal, the latter due to
the small size of the fetus a t term in relation to the dimensions of the female pelvic
inlet.” Leutenegger (1974) provided evidence that the degree of sexual dimorphism
in the ischio-pubic index is highly correlated with the relationship between the
dimensions of the maternal pelvis and those of the infant cranium. Figure 3 shows
the relationship between sexual dimorphism in the ischio-pubic index and the
relative breadth of the infant’s head. Note that, generally, species with relatively
spacious pelvic inlets (small relative cranial breadths) and low peluic sexual di-
morphism (at,the left in Fig. 3) are large in body size while those with relatively
small pelvic inlets and high pelvic sexual dimorphism (at the right in Fig. 3) are
small in body size. As stated above, the reason is that smaller primates have
relatively larger newborns (Leutenegger, 1973b, 1974, 1982a). Two exceptions to
the generalization about body size are Alouatta (howler monkeys) and humans.
Gould (1975) suggested that the relatively small brain size of howler monkeys is a
result of a recent increase in body size (phyletic giantism). Leutenegger (1982b)
suggested as an alternative that the highly specialized and enlarged vocal appa-
ratus of howler monkeys involves substantial modifications of their neurocrania
and limits the size of their brain.
The position of humans on Figure 3 is of most relevance to the current review.
Interestingly, based on a regression of neonatal brain weight on neonatal body
weight in anthropoid primates, humans have a brain size a t birth that matches
what is predicted from their body weight (Leutenegger, 198213). That is, humans
are not relatively encephalized a t birth compared to other living anthropoids.
What is different about humans is that their neonates are relatively large in body
size for a primate species of such considerable adult body size (Frost, 1987; Jor-
daan, 1976a,b; Leutenegger, 1973b; Lynch et al., 1983); i.e., human neonates de-
viate from a regression line of neonatal body weight on maternal body weight in
anthropoid primates by having large neonates relative to maternal body weight
130 r
100 ‘
60 70
I
60
I
90
1
100
I
110
1 1
120
I
130
Relative Cranial Breadth
Fig. 3. The relationship between pelvic sexual dimorphism on the vertical axis and relative size of the
neonatal cranium on the horizontal axis. Schultz predicted that species in which the neonatal cranium
was relatively larger would have more intense obstetrical selection on females and hence, greater sexual
dimorphism. After Leutenegger (1974), data in Tables 1 and 2.
(Fig. 1in Leutenegger, 1982b). The fact that humans have a position on Figure 3
close to primates of much smaller body size is a result of the relatively large size
of their newborns relative to maternal body weight (Leutenegger, 1982b).
Tague (1991) has taken another approach to addressing the hypothesized rela-
tionship between pelvic dimorphism and relative size of newborns. He examined
the patterns of pelvic dimorphism (that is, how different pelvic measures rank in
degree of dimorphism) in seven species of anthropoids. He ranked dimensions of
the pelvis in amount of sexual dimorphism and compared the ranking among
species. He found that species with relatively large newborns did not share a suite
of pelvic dimorphisms. Instead, shared patterns of dimorphism are associated with
taxonomic group (pongids and humans versus monkeys). On this basis he sug-
gested that “the hypothesis fails to be supported that interspecific commonalities
in pelvic dimorphism are closely related to obstetrics” (Tague, 1991:168).Note that
his test of the hypothesis involves shared patterns of dimorphism, whereas Leu-
tenegger’s (1974) test of the hypothesis discussed above involves degree of dimor-
phism. Two points make it unlikely that Tague’s approach might have supported
the hypothesis: 1)the fact that closely related groups share a recent evolutionary
history and locomotor pattern means that overall patterns of sexual dimorphism
will probably be shared; and 2) as Tague pointed out, pongids and humans are
dimorphic in the transverse plane of the pelvis, while monkeys are dimorphic in
the sagittal plane. This is almost certainly related to differences in trunk shape
and also to differences in the way these animals give birth. As discussed above,
humans and monkeys have different mechanisms of birth and different dimensions
of the pelvis which are most significant in assuring successful birth (Tague, 1991).
Therefore, one would not expect the pattern of their pelvic dimorphism to be
similar even if they share relative size of the newborn. One might expect, however,
that degree of pelvic dimorphism would be shared in these phylogenetically distant
species.
Duration and difficultyof labor. Duration of labor gives some indication of degree
of difficulty. The significance of difficulty during labor is that it reflects the danger
of birth and therefore of the extent to which natural selection may be acting.
Table 3, taken largely from Trevathan (19871, presents data on the average
TABLE 3. Length of labor in selected primate species’

First stage Second stage Third stage Total
Species (min) (min) (min) (min)
Sifaka2 ca.42 15
Marmoset3 30-300 10-30 10-24
Langur4 21 4
Colobus’ 240 20
Guenon‘ 30 30 32
Macaque7 23-210 1-2 1-7 11 14-270
Squirrel monkey’ 62
Mona monkey’ 54
Baboon9 42-67 4 4
Hamadryas baboon’’ 45
Orangutan 120 30
Gorilla” 18-155 1-18
n
2 ew
Chimpanzee” ca. 120 8
Humans’* 800 (450) 57 (18) 5 >862
‘From Trevathan, 1987, and other sources as noted.
‘Richard (1976) cited in Trevathan (1987).
3Stevenson (1976), cited in Trevathan (1987).
40ppenheimer (1976),cited in Trevathan (1987).
5Wooldridge (1970, cited in Trevathan (1987).
‘Brandt and Mitchell (1971).
7Tinklepaugh and Hartman (1930),cited in Trevathan (19871,Rawlins (1979),Adachi et al. (1982).Total estimate is from
Mitchell and Brandt (1975).
qakeshita (1961).
gLove (1978), Howell et al. (1978, cited in Trevathan (1987).
“Abegglen and Abegglen (1976).
”Galdikas (1982).
“Nadler (1974), Beck (1984), and Stewart (1984).
‘3Goodall and Athumani (1980).
I40xorn (1986). First figures is for primiparas, number in parentheses is for multiparas.
length of labor in primates. In spite of disparate data for primates, labor in humans
has the potential to be longer than in other species. Nevertheless, there seems to
be variation between different primate species in the length of time for delivery to
take place once labor has begun. These data do suggest that there is a n association
between the length of labor and the spaciousness of the maternal pelvis relative to
the neonatal head. Schultz (1969) noted that “the act of birth is a difficult and
prolonged experience for most monkeys” (p. 188)but that ape births “occur speed-
ily and rarely with noteworthy difficulty” (p. 154). Lindburg and Hazel1 (1972)
reported durations of labor in great apes ranging from 30 minutes to 8 hours, with
most times being less than 2 or 3 hours. This is in contrast to humans and mon-
keys, in which birth seems to be both longer and more difficult (Schultz, 1949,
1969). Labor in squirrel monkeys is apparently prolonged compared to other pri-
mates: “the similarity in squirrel monkeys and humans is not surprising since the
two species have among the largest neonatal dimensions, relative to maternal
pelvic dimensions in the order” (Trevathan, 1987:78). In terms of length of labor
and difficulty during labor, squirrel monkeys may be the living primate most
similar to modern humans.
Obviously, degree of difficulty during labor is problematic to quantify, or even to
describe in a way which will allow meaningful comparisons between members of
the same species, or across different species. Nevertheless, it is possible to extract
from the literature some subjective information about whether females appear to
find parturition a painful experience. Although vocalization during parturition
occurs in some primate species (Bo, 19711, in many cases females give birth quietly
and without any obvious facial expressions suggestive of discomfort (Trevathan,
1987). As parturition constitutes a vulnerable time for females and infants, one
might expect selection to favor female behavior at birth which does not call atten-
tion to the participants, hence the common nonhuman primate pattern of not
crying out during labor. Trevathan (1987) cited some evidence suggesting that
nonhuman primates experience what the observers interpret as pain during con-
tractions of labor. Collectively these observations suggest that while “difficulty”
100 YEARBOOK OF PHYSICAL ANTHROPOLOGY [Val. 35, 1992
during labor is by no means unique to humans (Trevathan, 1988) most primates

experience parturition as a simpler, shorter, and very likely less painful process.
Behavior during and after parturition
Trevathan (1987) suggested that in species in which maternal pe1vic:neonatal
cranial dimensions are close, selection should favor behaviors or postures during
labor which would increase the pelvic opening. For example, colobus and macaque
mothers have been observed to shift their posture during labor in ways which may
serve to increase the size of the pelvic aperture (Brandt and Mitchell, 1973). Many
species crouch forward or arch their backs, behaviors interpreted as increasing the
pelvic aperture (Trevathan, 1987).
Although a sitting, or semi-reclining position is most common, humans assume
a wide range of other postures during parturition (Gelis, 1991; Oxorn, 1986). These
include “squatting, kneeling, standing and !eft lateral or Sirn’s position” ( n e -
vathan, 1987:103), which increase the diameter of the pelvic opening (Gordon,
1985; Oxorn, 1986), and the “lithotomy position” (in which the woman lies on her
back, with her legs in stirrups and her buttocks at the edge of the table) which is
commonly used in the United States and is most convenient for obstetricians (Ar-
ney, 1982).
Primate mothers (and other nonhuman mammals) frequently manually assist
the process of parturition by probing the perineal region during labor and breaking
the amniotic sac or by pulling on the fetus as it emerges from the birth canal
(Schultz, 1969). Trevathan’s review (1987) of nonhuman primate birth gives abun-
dant anecdotal evidence for this phenomenon. In addition, once the fetal head
appears, mothers commonly reach for their infants with their hands and lift them.
“Primate mothers have been observed licking or wiping the mucus and fetal mem-
branes from the infant’s noses and mouths which clears a breathing pathway even
before the body has emerged’’ (Trevathan, 1988:678). It is difficult for human
females to reach their own perineal region during birth in order to provide manual
assistance to the emerging fetus or in order to pick it up. Not surprisingly, a female
assisting her own birth is much less common in humans than in other primates.
A final extremely significant difference between birth in nonhuman primates
and humans is the presence of birth attendants (Trevathan, 1987, 1988). Among
the former there is often curiosity and interest in the process of delivery especially
by females and juveniles, who watch and may touch the infant or mother, but
actual interference or assistance in the birth process is rare. When such assistance
does occur it most often takes the form of protection rather than interference in the
delivery process. For example, Abegglen and Abegglen (1976) reported a case of a
wild hamadryas baboon in which a female literally delivered an infant over the
edge of a cliff. The infant fell over the cliff, dangling by the umbilical cord. The
mother retrieved her offspring, but in the meantime, an adult male who had been
watching the process approached ready to help. The significance of birth atten-
dants in humans will be discussed below.
Human obstetric morphology
Having reviewed the differences between human and nonhuman primate birth
in three areas (mechanism of birth, degree of difficulty of birth, and behavior
during and after childbirth) we can now discuss the morphology associated with
those differences. Since information about the evolution of human birth will ulti-
mately come from the fossil record, an understanding of the skeletal morphology
associated with obstetrics is critical.
Clinicians most often divide the pelvis into the true pelvis and the false pelvis
(see Fig. 4). The false pelvis is that part of the pelvis that is (anatomically) ante-
rior-superior to the birth canal or true pelvis (i.e., the birth canal). The birth canal
in humans forms a tunnel of changing cross-sectional size and shape. Usually,
three planes are considered of clinical or obstetrical importance: the inlet (at the
iliopectineal line or linea terminalis), the midplane (at the level of the ischial
FALSE
PELVIS
TRUE
PELVIS
Fig. 4. The human pelvis in sagittal and coronal section, indicating the positions of the three pelvic
planes: the pelvic inlet, midplane, and outlet. After Oxorn (1986).
spines), and the outlet (at the level of the ischial tuberosities and the apex of the
sacrum) (see Fig. 4). In modern human females, the inlet, midplane, and outlet
have different cross-sectional shapes. The inlet is transversely elongated, the mid-
plane is sagittally or anterior-posteriorly elongated, and the outlet is approxi-
mately circular. This changing shape determines the mechanism of birth in hu-
mans just as the constant shape in nonhuman primates allows the passage of the
infant without any reorientation along the way (Tague and Lovejoy, 1986). Of the
three planes, the midplane is smallest in circumference and the inlet the largest in
modern human females (Tague, 1992a).
Figure 5 [after Tague and Lovejoy (1986)l summarizes the obstetrically signifi-
cant features of hominid pelvic anatomy compared to nonhuman primates (in this
case, Pan).Briefly, in hominids:
1. The ilium is short and broad in hominids compared to most other primates, so
that the distance between the sacroiliac and hip joints is reduced (Berge et al.,
1984; Jordaan, 1976c; Lovejoy, 1973, 1975, 1978, 1988; Lovejoy et al., 1973; Sig-
mon, 1986; Straus, 1929; Tague and Lovejoy, 1986). Without other changes, this
would reduce the sagittal (anterior-posterior) diameter a t the pelvic inlet and
outlet.
2. The ilium in hominids is “extended dorsally” compared to other primates,
which separates the sacroiliac and hip joints sagittally “thereby maintaining an
obstetrically adequate anterior-posterior diameter of the pelvic inlet” (Tague and
Lovejoy, 1986:238).
3. In addition, hominid ilia are more sagittally oriented than those of other
102 YEARBOOK OF PHYSICAL ANTHROPOLOGY IVol. 35, 1992
Fig. 5. A chimpanzee pelvis (top), the reconstructed pelvis of A.L. 288-1 (middle), and the pelvis of a
modern human female (bottom)in approximately anterior and superior view. After Tague and Lovejoy
(1986).
primates, with the iliac blades flaring laterally, increasing the transverse diam-
eter of the pelvic inlet (Tague and Lovejoy, 1986).
4. The sacrum is wider in hominids than in other primates (Abitbol, 1987a;
Leutenegger, 1977; Lovejoy, 1988; Tague, 1991; T a p e and Lovejoy, 1986) sepa-
rating the ilia posteriorly.
5 . An important aspect of hominid pelvic morphology from the point of view of
obstetrics is the position of the sacrum. In monkeys, the apex of the sacrum does
not extend far into the birth canal. This means that the fetal head does not have
to pass the sacrum and pubic symphysis simultaneously, making passage through
the lower portion of the pelvic aperture fairly easy (Abitbol, 1987a; Schultz, 1949).
In apes, although the fetal head must pass these structures simultaneously, the
pelvic aperture is sufficiently spacious that this presents no barrier to successful
delivery (Trevathan, 1988). In hominids, the sacrum has become the dorsal wall of
the entire pelvic aperture and substantially reduces the anterior-posterior diam-
eters of the birth canal a t the midplane and outlet (Tague and Lovejoy, 1986).
6 . Finally, in humans the pelvic outlet faces more ventrally than in most non-
human primates so that the fetus must bend in order to emerge from the pelvic
outlet, rather than passing in a straight line behind the ischial tuberosities as in
nonhuman primates (Berge et al., 1984).
In humans and nonhuman primates, there are several important mechanisms
which increase the space available in the birth canal for passage of the fetal head.
As mentioned previously, posture assumed by the mother during labor can in-
crease the diameter of the birth canal (particularly the outlet) (Oxorn, 1986).
Russell and Richards (1971) reported that a squatting posture can produce an
outlet 25% to 30% larger in area than a supine posture in the same woman. This
occurs because of movement at the sacroiliac joint and a t the pubic symphysis and
has the effect of tilting back the sacrum and separating the pubic bones increasing
both the transverse and anterior-posterior diameters of the birth canal (Gordon,
1985).
During pregnancy, increased levels of the hormone relaxin cause a relaxation of
the ligaments a t the sacroiliac articulations and the pubic symphysis (Baird, 1962;
Borell and Fernstrom, 1957; Oxorn, 1986; Percival, 1980; Pritchard and MacDon-
ald, 1980; Putschar, 1976; DB Stewart, 1984a). This increases joint mobility and
the dimensions of the pelvic inlet by a considerable amount (Borell and Fernstrom,
1957),.
Finally, during labor, the fetal head is subjected to compressive forces arising
from maternal uterine contractions when it is pressed against the maternal pelvic
bones. Because the sutures of the fetal cranial bones are not fused and the fetal
cranium is quite plastic, movement and deformation can occur between the indi-
vidual bones. Generally, force is exerted along the sub-occipital-bregmatic or sub-
occipital-frontal diameter, so either of these dimensions reduces with compensa-
tory elongation in the mento-vertical diameter (Guha, 1990). Molding may account
for a reduction in fetal cranial diameter of 5-10 mm (Kriewall and McPherson,
1981; Pritchard and MacDonald, 1980). The resulting “cone-head” shape o f the
fetal skull (visible in Fig. 1) IS characteristic of newborn humans born vaginally
especially after prolonged labor. Stewart and Philpott (1980) showed that extent of
fetal head molding is positively correlated with the degree of cephalo-pelvic dis-
proportion. Although fetal cranial molding is a n important adaptive response to a
tight fit in the birth canal, excessive molding is disadvantageous as i t is linked to
a number of disorders in infants, including cerebral trauma, mental retardation,
and cerebral palsy (McPherson and Kriewall, 1980a,b). Schultz (1969) reported
that the fontanelles, which facilitate molding, have nearly or entirely disappeared
by the time of birth in monkeys, are small at birth in apes, but are still large in
human neonates. This raises the possibility that the human fetal cranium may be
more malleable than that of other primates, presumably in response to selection
resulting from obstetrical constraints.
None of these mechanisms of alleviating feto-pelvic disproportion is necessarily
unique to living humans: all are seen in varying degrees in other primates. There-
fore, we can assume that they most likely also occurred in extinct hominids.
Selection during birth in living humans
Before discussing the evolution of birth in hominids, the issue must be addressed
as to whether there is evidence for selection acting on the pelvis in living humans.
There are, of course, many environmental factors such as nutritional deficiencies
which may affect the size of the pelvic aperture (Abitbol, 1987b; Angel, 1978;
Stuart-Macadam, 1989). However, there is at least some genetic component to
variation in pelvic morphology. Evidence for selection acting on the pelvis consists
principally3 of clinical data which shows the relationship between the size and
shape of the female pelvis and outcome of birth. Many classifications of anatomical
variations in the female pelvis have been created to aid obstetricians or midwives
in making accurate predictions about a woman’s success in childbirth from a vag-
inal or radiographic examination of her pelvis (Caldwell and Moloy, 1933; Cald-
well et al., 1939; DeSouza, 1913; Greulich and Thoms, 1938, 1939; Heyns, 1946).
One approximate indication of whether a particular woman’s birth canal was suf-
ficiently spacious to allow passage of her neonate is whether her attendants de-
termined that operative intervention, (cesarean section or forceps delivery) was
necessary.4 For example, Greulich and Thoms (1938) showed t h a t 30% of all de-
3Although it is difficult to observe or infer selection in a prehistoric population, Tague (1986, 1992c) showed
a significant difference between younger (18-24 years of age) and older (25 years of age and older) adult females
in three prehistoric populations in linea terminalis length. One explanation was that this was the result of
selection in the form of maternal mortality due to cephalo-pelvic disproportion and that women with larger
pelvic inlets survived longer (and presumably left more offspring) than women with smaller pelvic inlets. An
alternative explanation is that this correlation is the result of aging (Tague, 1986, 1 9 9 2 ~ ) .
4Note that I do not defend the seriously inflated cesarean section rate in the United States today (approxi-
mately 24% of all births [Public Citizen Health Research Group, 1989; Taffel et al., 19921, many of which are
performed for reasons other than cephalopelvic disproportion or for that matter the welfare of patients). How-
ever, it is certainly the case that more women who undergo cesarean section or other operative intervention have
pelvic constriction than women who have successful normal vaginal deliveries. In addition, there is a potential
for circularity in that physicians base their decisions about whether to perform cesareans at least in part on
pelvic size.
liveries on women with platypellic (anterior-posteriorly flattened) pelves resulted

in operative intervention while only about 18%of deliveries in women with other
types of pelvic shape resulted in such intervention. Following this, a number of
studies in the late 1940s through the early 1960s attempted to predict the likeli-
hood of dystocia (abnormal labor) using radiographic pelvimetry (Allbrook, 1962;
Allbrook and Sibthorpe, 1961; Allen, 1947a,b; Baird, 1952; Bore11 and Fernstrom,
1960). Dimensions of the maternal pelvis and sometimes of the fetal cranium were
taken and compared to standards. Although this practice was ultimately aban-
doned because of dangers associated with radiation, there is abundant evidence for
a causal connection between maternal pelvic and infant cranial dimensions on the
one hand and the risk of dystocia or cesarean section (advised by the attending
physician because of their judgement that dystocia was inevitable) on the other (for
example, Amirikia et ai., 1981;Jagani et al., 1Y81; Joyce et al., 1975; Kolawole et
al., 1978; Mandry et al., 1983; Morgan et al., 1986; OBrien and Cefalo, 1982;
Thurnau et al., 1991; Yamazaki and Uchida, 1982). With the caveat that many
cesarean sections performed today are probably not necessary from the point of
view of alleviating cephalo-pelvic disproportion, these data nevertheless suggest
that in the absence of medical intervention, selection would act in modern times to
reduce the prevalence of pelvic morphology that interferes with successful passage
of the neonate (Harrison, 1985).
Data from Shorter (19821, discussed in Rosenberg (19891, show that in mid-19th
century maternity hospitals in Paris, there was a positive correlation between
degree of pelvic constriction and percentage of mothers dying in childbirth. Be-
cause many of these women had constricted pelves due to rickets, these data are
not evidence of selection against genetically determined pelvic contraction. Nev-
ertheless, they provide support for the important statement that pelvic contraction
is associated with increased maternal mortality.
There is additional evidence that selection operates against individuals who
frequently have breech presentations. Breech births are associated in living hu-
mans with increased infant mortality (approximately three times the rate for
cephalic presentations when delivered vaginally in modern clinical settings) and
increased risk of injury to the infant (damage to the infant occurs in between 3.5%
and 20% of cases, depending on the type of breech) (Oxorn, 1986).
These data as a whole suggest that in the absence of medical intervention, there
is selection acting in humans as a result of obstetrical constraints. This selection
acts in the realms of maternal pelvic diameters, mechanism of birth, and fetal size.
Human birth helpers
Trevathan (1987, 1988) argued that because human infants are born in an oc-
ciput anterior position, it is difficult, if not impossible, for human mothers to assist
their fetuses manually as their heads emerge from the birth canal. Because the
infant emerges facing away from the mother, any attempt to lift the infant towards
the mother would pull the infant against the normal flexion of its body, running
the “risk of injury or even paralysis especially in the neck region” (Trevathan,
1988:678).Human mothers are also in an awkward position during labor to reach
the umbilical cord, which may be wound around the infant’s neck, or to clear the
infant’s air passages after the emergence of the head. All of these difficulties can
be easily alleviated by an attending person. In cases of difficult delivery, “mor-
tality has been close to 100% for unassisted breech deliveries” (Trevathan, 1987:
94). Women who choose to labor in the presence of other individuals, preferably
ones at least somewhat experienced in the birth process, are clearly a t an advan-
tage in these contexts. Trevathan suggested that “the human adaptation to this
challenge has been the behavioral characteristic of seeking assistance a t b i r t h
(Trevathan, 1988:678)or “obligate midwifery.” In almost all human societies, birth
is an event which is attended by other adults, usually females, rather than under-
gone alone (Newton and Newton, 1972). One possible exception to this generali-
zation is the !Kung, who have a cultural ideal of women giving birth alone. In
practice, however, !Kung women are virtually always attended by female relatives
a t their first birth. By the fourth or fifth birth, about half of all women do labor
unattended, but always within earshot of potential helpers (Konner and Shostak,
1987).
The suggestion that attendants are important in human birth was also made by
Masters (1978) and Barash (1979), who argued that humans have a uniquely high
rate of maternal mortality during childbirth and that this would set a maximum
limit on possible brain size in infants. Because death in childbirth would be higher
for mothers who gave birth alone than for those who had assistance, Masters
(1978) and Barash (1979) argued that altruistic behavior in childbirth may have
allowed brain size to increase since relatives assisting each other in childbirth
would help to increase the probability of a successful birth outcome. They suggest
that such cooperative behavior has been an important factor in allowing human
brain size t o increase. “I rdihar like the idea that midwitery and obstetrics may
literally be our oldest professions, or a t least the ones that ‘gave birth’ to our
humanity” (Barash, 1979:167).
Because unattended delivery is unusual in humans, there are few accounts of
mortality rates. It is likely that it is associated with increased mortality [Ohio
State Medical Association (1975) and Edwards (19731, cited in Trevathan (1988)l.
Clinical evidence provides a different kind of support (than that suggested by
Trevathan) for the assertion that birth attendants can play an important role in
increasing the probability of a successful birth. In a study of women giving birth in
hospitals, continuous emotional support provided by a female companion or doula
significantly reduced the need for surgical intervention and significantly short-
ened labors (Duignan, 1985; Kennel1 et al., 1991; Kitzinger, 1985; Sosa et al.,
1980). Although these studies compared women giving birth in a hospital with
medical intervention but no emotional support with those giving birth in the same
environment but with emotional support, rather than looking at women giving
birth totally unassisted, they are relevant in suggesting that one should not un-
derestimate the importance of birth attendants in providing assistance beyond
actual physical intervention. Although data are not available to address this ques-
tion directly, “moral support” seems to be a significant factor in cases of prolonged
labor and “assisted” birth could be adaptive for this reason as well.
In addition, other types of assistance during pregnancy andlor childbirth can
improve the likelihood of successful birth. This could include many types of inter-
vention by a person experienced in childbirth. For example, fetuses which are in a
breech position can sometimes be turned using a procedure referred to by mediral
professionals today as “external version.” ‘Phis requires no modern medical tech-
nology but does depend on knowledge and experience of the attendant who must be
able to determine the position of the fetus by external palpation of the mother’s
abdomen and to turn it using externally applied manual pressure. Although a
fetus often returns to a breech position following external version, there is evi-
dence that this procedure reduces the number of breeches a t the time of delivery
(Cunningham et al., 1989). Such behavior in early humans could certainly have
reduced the rate of birth complications as well.
In summary, the major differences between living humans and nonhuman pri-
mates in the birth process and associated pelvic morphology are: 1)The position of
the sacrum relative to the birth canal. In humans, the sacrum forms the dorsal
wall of the bony birth canal and must be passed by the fetus a t the same time as
the pubic symphysis; in nonhumans the sacrum is passed in advance of the pubis.
2) Position of the outlet. In humans the pelvic outlet is in front of the ischial
tuberosities so that the fetus emerges in front of them, bending as it does so; in
nonhumans the fetus emerges behind the ischial tuberosities with no bending. 3)
Shape of the birth canal. In humans the birth canal is transversely elongated a t
the inlet, sagittally elongated a t the midplane, and approximately circular a t the
outlet. In nonhuman primates the birth canal is sagittally elongated a t all planes.
4) The orientation of the fetal head as it enters the pelvic inlet. In humans the fetal
head is transversely or obliquely oriented a t the pelvic inlet; in nonhumans it is

sagittally oriented. 5) Mechanism of birth. In humans, the fetus rotates internally
to emerge from the birth canal in a different orientation from the one in which it
entered the inlet; in nonhumans there is no such change in orientation. 6) Emer-
gence pattern. Human infants usually emerge with the occiput anterior; in non-
humans the occiput is more often posterior. 7) Presence of helpers. Human birth is
almost invariably attended and assisted; nonhuman birth is rarely if ever assisted.
8 ) Degree of difficulty. Though not unique in being difficult, human birth is gen-
erally more difficult, more prolonged, and presumably more dangerous than birth
in nonhuman primates or other mammals that have low cephalo-pelvic ratios.
THE FOSSIL RECORD
There are three important issues about birth that have been addressed using the
fossil record: 1)When and how did the unique mechanism of birth seen in humans
arise? 2) When and how did secondary altriciality [in which infants are born before
rapid “fetal” growth is completed and remain in that “exterogestate” state (Mon-
tagu, 1961) for some time following birth] arise? and 3) When did the patterns of
pelvic dimorphism that are seen in adult humans arise?
The size and shape of the birth canal directs the passage of the infant and
mechanism of birth (Tague and Lovejoy, 1986). Information about the first of these
questions, the origin of the human mechanism of birth, can be deduced directly
from an understanding of pelvic morphology. Second, with regard to the origin of
secondary altriciality in humans, Brown et al. (19851, Gould (19851, and Trinkaus
(1984) have argued that information about secondary altriciality can also be ob-
tained by studying the size of the pelvic aperture. In chimpanzees there is a decline
in the rate of brain growth from the fetal rate to the post-fetal rate after the time
of birth (Martin, 1983). In humans, this decline in rate of growth takes place about
1 year after birth (Martin, 1989). This means that humans are born before their
“fetal” brain growth is completed, that is, they are more altricial than chimpan-
zees. Although the issue of hominid maturation rates has been investigated ex-
tensively from an analysis of dental development, discussion of that controversial
literature is beyond the scope of this paper. However, some authors have suggested
that an analysis of pelvic aperture size in extinct hominids can indicate how big
the brain of a neonate was and that this information may be useful in determining
maturation rates. Third, an investigation of pelvic morphology in fossil hominids
can indicate when modern patterns and levels of pelvic sexual dimorphism arose.
How did the modern human pattern of childbirth arise?
When and how did the unique pattern of birth seen in humans arise? As Tague
and Lovejoy (1986:246)have emphasized, “the shape of the maternal pelvis deter-
mines the pattern of birth.” A corollary of this statement is that the mechanism of
birth can be inferred from the pelvic morphology of extinct species.
Primitive hominoid pelvic morphology
Primitive hominoid pelvic morphology has been assumed to be similar to that of
living great apes (Leutenegger, 198213; Lovejoy, 1973, 1975, 1988; Lovejoy et al.,
1973; Tague and Lovejoy, 1986). Great apes have pelvic inlets that are elongated
cranio-caudally and expanded mediolaterally relative to living monkeys as a result
of their torso morphology and locomotion (Abitbol, 1991; Schultz, 1949; Leuteneg-
ger, 1974). Unfortunately, no late Miocene proto-hominid pelvis is known in the
fossil record. Ward (1991) has argued that Proconsul nyanzae (KNM-MW 13142),
an early Miocene hominoid, may be representative of the primitive condition for
catarrhines. The relevant features of this specimen are a narrow pelvis, typical of
most pronograde quadrupeds, a relatively short distance between the sacro-iliac
and hip joints (unlike great apes), short pubic bones, and a narrow sacrum. Al-
though it did not have a tail, Proconsul pelvic morphology was not particularly
hominoid-like (Ward et al., 1991). “Although KNM-MW 13142 was a male, it
certainly seems as if this species would have been characterized by a relatively

narrow birth canal, and a pattern of obstetrics like that of extant monkeys in
which the infant is close in size to the pelvic inlet” (Ward, 1991:322). If Proconsul
does represent the primitive hominoid condition, i t is possible t h a t hominids may
never have passed through a great ape-like condition with respect to pelvic mor-
phology (Ward, 1991). Ward (1991)offers the hypothesis that the ape-human com-
mon ancestor may have been less derived in torso morphology and the short iliac
blade in hominids may reflect the primitive rather than the derived condition. A
further implication is that orangutans are convergent on African apes in torso
morphology. Finally, these data suggest that ancestors of the earliest hominids
may not necessarily have had relatively spacious pelvic inlets comparable to living
great apes. Instead they may have had a n obstetric configuration with more con-
striction between the maternal pelvis and the neonatal head, as in living monkeys
(Ward, i99i). This intriguing hypothesis can only be tested by further discoveries
of middle and late Miocene fossils.
Australopithecines
Our knowledge of obstetrically significant morphology in australopithecines
comes almost entirely from two specimens: Sts 14 from Sterkfontein, South Africa
(Broom et al., 1950; Robinson, 1972), and A.L. 288-1 (Lucy) from Hadar, Ethiopia
(Johanson et al., 1982). Other australopithecine pelvic material is either much
more fragmentary, severely distorted, or represents juvenile individuals. Although
the anterior-posterior and transverse diameters of the inlet can be measured in Sts
14, distortion on that specimen indicates (Berge et al., 1984; Day, 1973; Robinson,
1972; Tague and Lovejoy, 1986) that A.L. 288-1 is a “more anatomically accurate
specimen upon which to assess the early hominid obstetric pelvis” (Tague and
Lovejoy, 1986:252). Sts 14 preserves the right and most of the left hip bone as well
as a fragmentary sacrum. A.L. 288-1 is represented by a complete sacrum and left
hip bone on which transverse measurements of the pelvis were made to the midline
and doubled (Tague and Lovejoy, 1986).
Discussions of the birth process in australopithecines have reached a wide range
of conclusions, namely, that australopithecines had a birth process which was 1)
“quick and easy” (Leutenegger, l972,1982b, 1987; Wolpoff, 1978,1980);2) similar
to modern humans either in degree of difficulty (McHenry, 1986) or in “mechanism
of birth” (Abitbol, 198713; Berge et al., 1984; Fisher, 1982; Jordaan, 1976c; DB
Stewart, 1984a); and 3) unlike any seen in living humans or nonhuman primates
(Tague and Lovejoy, 1986; Lovejoy, 1988).
These analyses of obstetric relationships in australopithecines have concen-
trated on two critical aspects of the pelvic aperture: size and shape. Size and shape
are analyzed to determine degree of difficulty during labor and to infer the birth
mechanism. Issues addressed include whether australopithecine fetuses entered
the birth canal with the head oriented as in modern humans and whether the fetus
experienced rotation of the head and/or body during passage. The discussion below
will first consider the size of the pelvic inlet and then the shape of the birth canal
a s a whole.
Size. Even before the discovery of A.L. 288-1, Lovejoy (1973, 1975; Lovejoy et al.,
1973) argued that australopithecine pelvic morphology was singularly adapted to
bipedal locomotion, and that as encephalization took place during the Pleistocene,
selection led to a n increase in the size of the birth canal, which also increased
interacetabular breadth. He argued that differences in pelvic morphology between
Australopithecus and Homo (such as reduction of iliac flare which increased torque
about the hip and a less advantageous position for the abductors muscles) could be
explained by increase in the size of the birth canal a s a result of encephalization.
These changes led to a n increase in femoral head pressure and involved commen-
surate modifications in the anterior superior iliac spine, the iliac pillar, femoral
TABLE 4.The relationship between neonatal cranial site and maternal pelvic inlet size i n Homo,
Australopithecus, and Pan
Anterior-posterior' Transverse2
Pun (females, N = 25)
Pelvic inlet (cm) 14.2 10.5
Neonatal cranium (cm) 8.3 7.1
Relative cranial size3 58.4% 67.7%
Homo (females, N = 106)
Pelvic inlet (cm) 10.4 13.4
Neonatal cranium (cm) 12.4 9.8
Relative cranial size assuming no rotation 119.2% 73.1%
Relative cranial size assuming rotation 94.7% 92.5%
A.L. 288-1
Pelvic inlet (cmf 7.6 13.2
Pan neonatal cranium (cm) 5.3 '7.1
Relative cranial size assuming no rotation 109.2% 53.8%
STS 14
Pelvic inlet (cmf 8.5 9.9
Pun neonatal cranium (cm) 8.3 7.1
Relative cranial size assuming no rotation 97.6% 7i:7%
'Anterior-posterior diameter of the pelvic inlet is from the ventral surface of the sacral promontory to the dorsomedial
aspect of the pubic symphysis. Anterior-posterior diameter of the neonatal cranium is the fronto-occipitallength,
*Transverse diameter of the pelvic inlet is the maximum distance between the iliopectineal lines. Transverse diameter
of the neonatal cranium is biparietal diameter.
3Relative cranial size is the diameter of the cranium divided by the relevant diameter of the inlet. Where rotation is
assumed, the biparietal diameter of the cranium is divided by the anterior-posterior diameter of the pelvic inlet, and the
fronto-occipitaldiameter of the cranium is divided by the transverse diameter of the pelvic inlet.
From Tague and Lovejoy, 1986, and, Leutenegger, 1987.
neck length, and femoral head size (Lovejoy, 1973,1975,1978;Lovejoy et al., 1973;
McHenry, 1986). Although he did not explicitly discuss australopithecine birth, it
was implicit in Lovejoy's early statements that obstetrics had not become an im-
portant cause of selection on the hominid pelvis until the beginning of rapid brain
size increase.
Data presented by Leutenegger (1972) and Tague and Lovejoy (1986) are
relevant to the question of how large the pelvic inlet was in australopithecines
relative to estimated newborn cranial size. These authors, as well as most others
in the past, assumed that Pan (with a relatively spacious pelvic aperture) provides
the best living model for the ancestral pre-hominid pelvis. Table 4 gives data on
the relationship between maternal pelvic inlet size and neonatal cranial size in
Pan, A.L. 288-1, and Homo. In the case of Pan, the dimensions of the neonatal
cranium are much smaller than the corresponding dimensions of the pelvic inlet.
Consequently, the cranium can pass into the pelvic inlet in essentially any
orientation, though it usually does so with the sagittal axis of the neonatal skull
aligned with the sagittal axis of the maternal pelvis. Relative size of the neonatal
cranium is low (58.4% for the anterior-posterior diameter and 67.7% for the
transverse diameter) reflecting the spaciousness of the pelvis. In Homo the long
axis of the fetal cranium is actually greater than the anterior-posterior diameter
of the pelvic inlet. Therefore, i f the fetal cranium entered the pelvic inlet in the
same orientation as in Pan there would be constriction in the anterior-posterior
diameter (where relative cranial size would be 119.2% and successful passage
could not occur). However, as mentioned above, the human neonatal skull enters
the pelvic inlet aligned either along the transverse diameter or obliquely, so that
there is sufficient space (relative cranial size decreases to 94.7% in the ante-
rior-posterior diameter and 92.5% in the transverse diameter). Note that even
taking this orientation into account, the size of the infant cranium relative to the
pelvic inlet is considerably greater in Homo (in both the anterior-posterior and
transverse diameters) than it is in Pan. That is, humans have a less spacious
pelvis than chimpanzees.
In order to determine how spacious the pelvic inlet was in australopithecines,

Leutenegger (1972) estimated cranial capacity for neonatal A. africanus. Based on
a birth weight of 1,660 g (estimated from maternal weight, which was in turn
estimated from adult female skeletal measures) and a neonatal cranial capacity of
9.8% of birth weight (since chimpanzees, 9.7%, and modern humans, 9.9%, share
essentially identical values) Austral0 ithecus africanus neonates were estimated
to have a cranial capacity of 163 cm! close to the size of a chimpanzee neonate
(Leutenegger, 1972) (see Table 4). Leutenegger concluded that A. africanus neo-
nates were small relative to the size of the female pelvic inlet. This suggested to
him that 1)birth was “quick and easy” (Leutenegger, 1972:569), and 2) selective
pressures for enlarging the birth canal were not yet present (Leutenegger, 1972).
If a Pan type of cranial orientation occurred in australopithecines, constriction in
the anterior-posterior dimensions would have been even more extreme than in
modern humaria, biit if a human-type head orientation occurred, constriction in
both diameters would have been below the level seen in modern humans but still
above the level seen in Pan.
Shape. The excellent preservation of A.L. 288-1 allowed Tague and Lovejoy (1986)
to evaluate not only the pelvic inlet, but the shape of the entire birth canal. The
shape of the pelvic inlet of A.L. 288-1 means that the pelvis could accommodate a
fetal cranium the size of newborn Pan only if the fetus entered in a Homo-like
orientation (Tague and Lovejoy, 1986) but the inlet and midplane were “amply
spacious relative to fetal size.” They suggest, however, (see below) that there was
contraction in the lower pelvis.
Using slightly different estimates of neonatal brain size in A. africanus and A.
afarensis than he had used in 1972, Leutenegger (198715argued that the austral-
ipithecine fetal cranium would have entered the pelvic inlet with an alignment
like that seen in modern humans. He disagreed with Tague and Lovejoy (1986),
however, that there was cephalo-pelvic constriction in the sagittal dimensions of
the lower pelvis. The difference in interpretations between these two studies may
be due to the slightly different estimates of neonatal brain size used. Tague and
Lovejoy (1986) used Leutenegger’s (1972) original estimate that A. africanus was
similar in neonatal brain size to living Pan. Leutenegger’s later estimates of neo-
natal brain size in A . afarensis and A. africanus were based on maternal body
weight (estimated from maternal femoral shaft dimensions and using a regression
based on hominoids) resulting in slightly lower estimates of neonatal brain size for
these extinct hominids. He reported estimates for brain sizes of 120.7 g for A .
afarensis and 115.3 g for A. africanus. If the lower estimates are used, A.L. 288-1
has a pelvic inlet exactly the size of the estimated dimensions of the A. afarensis
infant skull and probably would have required the cranium to enter the inlet in a
rotated position as in modern humans. In A. africanus, if the lower estimates of
neonatal brain size are used, the relative size of the cranium is below Homo and
approaches Pan. Given the estimated nature of these neonatal brain sizes, the
reasonable conclusion is that A.L. 288-1 shows evidence that the fetus entered the
pelvic inlet with a Homo type orientation of the head (regardless of which model
one uses for neonatal brain size) and that Sts 14 shows evidence of a pelvic inlet
that was slightly more spacious in the anterior-posterior diameter, but slightly less
spacious in the transverse diameter.
Tague (1991) analyzed variability in the anthropoid pelvis and determined that
pelvic shape [(anterior-posterior diameterhansverse diameter) x 1001 is highly
variable in modern anthropoids. Assuming that this variability was also true of
Australopithecus, he concluded that “A. afarensis’ pelvis was potentially more
spacious than that in humans at each pelvic plane” (Tague, 1991:173).
’The difference between Leutenegger’s original (1972) estimate and his later one (1987) is a t most 6 mm in
biparietal diameter and 7 mm in occipitofrontal diameter. Given the amount of soft tissue between the maternal
pelvis and the fetal cranium, the effect of such small differences in fetal skull size is uncertain.
110 YEARBOOK O F PHYSICAL ANTHROPOLOGY [Vol. 35, 1992
TABLE 5. Pelvic aperture indices’ of modern humans and A.L. 288-1’

Inlet Midplane Outlet
Males Females Males Females Males Females
Indian Knoll 85.2 83.6 124.4 108.5 109.3 90.5
7.0 (42) 5.8 (291 12.1 (10) 10.4 (4) 12.5 (24) 8.0 (22)
Pecos Pueblo 75.1 79.9 134.6 122.3
7.0 (37) 6.2 (46) 15.1 (10) 9.9 (6) 113.1
13.2 (32) 100.6
10.5 (25)
Libben 80.8 75.0 127.5 114.6 96.4
8.3 (24) 5.4 (11) 12.0 (4) 10.4 (17) 7.7 (5)
Haida 81.2 86.0 144.3 124.6
7.2 (22) 8.2 (19) 13.6 (14) 11.5 (8) 116.5
11.1 (16) 107.8
13.3 (10)
H a m a n n Todd 83.0 86.4 133.4 128.1
Whites 8.2 (47) 9.7 (49) 6.9 (11) 8.6 (3) 14.1 (44)
111.1 99.8
11.0 (46)
H a m a n n Todd 86.6 93 4 145 2 133.0 120.9 199.5.
_
Blacks 8.0 (49) 11.0 (49) 15.9 (23) 14.2 (21) -16.5 (48) 15.5 146)
“Modern 85 88 115 110 102 105
human^"^ 11(49) 8 (64) 13 (49) 11(64) 9 (48) 11(64)
A.L. 288-1* 57.6 71.3 74.0
‘All indices are calculated as (anterior-posterior diameteritransverse diameter) x 100.
’A11 data were from Tague (1989)except where indicated. Inlet dimensions were supplied by Tague.
3“Modern humans” are a mixed sample from Abitbol (1987b).
4Data from Tague and Lovejoy (1986).
Mechanism of birth in australopithecines. Berge et al. (1984) and Berge and

Kazmierczak (1986) argued that A.L. 288-1 and Sts 14 showed most of the obstet-
rical morphologies seen in modern humans. In particular, these authors suggested
that these specimens show a pelvic outlet which opens ventrally to the ischial
tuberosities, leading to an “ante-ischiatic” path of parturition in which the fetus
emerges from the birth canal anteriorly as in humans and (again, assuming a
neonatal size similar to Pan) an orientation of the neonatal skull in the inlet in a n
oblique or transverse position as in modern humans. Following this position of
engagement in the inlet, Berge and colleagues assumed that the fetus in Australo-
pithecus would have had to rotate as in modern humans and that constriction
existed, so that the fetal head did not “pass with any more ease than in Homo” and
that “an obstetrical traumatism already existed in the first hominids” (Berge et al.,
1984585). They suggested contra Lovejoy (1973, 1975) that the human mode of
parturition appeared early in human evolution and that the “human obstetrical
constraints thus seem not to be consequences of the cerebral development which
marked the transition from genus A ustralopithecus to genus Homo” (Berge et al.,
1984:585).
The pelvis of A.L. 288-1 is extremely wide, both absolutely and relative to stat-
ure. Tague and Lovejoy (1986) showed that the pelvis of A.L. 288-1 is transversely
elongated (platypelloid) a t all planes (see also Abitbol, 1991). This pelvic shape
differs from that seen in living nonhuman primates (in whom the inlet, midplane,
and outlet are all elongated in the anterior-posterior plane) and living humans (in
whom the shape of the pelvic aperture changes from transversely elongated at the
inlet, to elongated in the anterior-posterior plane in the midplane, to approxi-
mately equal in anterior-posterior and transverse diameters a t the outlet). Table 5
shows data on the index of pelvic shape [(anterior-posterior diameteritransverse
diameter) x 1001 for a number of human populations. At the pelvic inlet, A.L.
288-1 is almost two standard deviations below the average of the most platypelloid
population (Pecos Pueblol.6 At the midplane, A.L. 288-1 is also transversely elon-
gated, while none of the modern human populations shows this shape at the mid-
plane. At the outlet, A.L. 288-1 preserves a transversely flattened shape. The
sample means of modern females cluster around 100, indicating that at the outlet,
6Note that measurements made on the somewhat distorted Sts 14 specimen show it had a pelvic inlet which
was also transversely elongated [index at, the inlet is 85 6, within the range of the modern human samples
(Tague and Lovejoy, 198611.
transverse diameter (breadth between the ischial tuberosities) is approximately

equal to anterior-posterior diameter (from the dorsomedial aspect of the inferior
margin of the pubic symphysis to the apex of the fifth sacral vertebra). The shape
of the pelvic outlet in A.L. 288-1 is outside the range of modern human variation
represented in Table 5.
Tague and Lovejoy (1986) inferred the mechanism of birth from the shape of the
birth canal. They agreed with Berge et al. (1984) that the neonatal cranium would
have entered the inlet oriented as in modern humans. However, they suggested
that at the midplane there “would not have been bony resistance to fetal descent”
(Tague and Lovejoy 1986:248) and that the fetal head would have maintained its
original orientation as it passed through. A.L. 288-1 has a pelvis with a low index
of funnelling [(transverse diameter of outlet/transverse diameter of inlet) x loo)]
so the outlet is narrow relative to the inlet and there would have been “bony
resistance sufficient to promote cranial flexion” (Tague and Lovejoy, 1986:248).
Not only would fetal rotation within the birth canal have been unnecessary, such
rotation would have been impossible. This is because unlike modern humans, A.L.
288-1 has anterior-posterior diameters of the midplane and outlet which are even
narrower than the inlet. Because of the narrow anterior-posterior diameter of the
lower portion of the birth canal, Tague and Lovejoy (1986:249) argued that the
fetus could have passed through only if i t assumed “an asynclitic cephalic position
[in which the head is tilted so that the parietals do not enter the inlet simulta-
neously] and if the sacroiliac joints . . . increased in mobility during pregnancy.”
Tague and Lovejoy (1986:249) concluded that this would have reduced the trans-
verse diameter of the fetal head by presenting “the shorter subsuperparietal rather
than the biparietal” diameter. They suggested that in australopithecines the
transverse orientation of the head was maintained throughout the passage of the
fetus through the birth canal7.
For these reasons, Tague and Lovejoy (1986) concluded that australopithecine
birth involved a mechanism unlike that known for any other species. The mech-
anism of birth was similar to modern humans in the way in which the fetal
cranium entered the pelvic inlet, but simpler than modern humans in the path that
the fetus followed as it passed through the birth canal. They contended “that A.L.,
288-1’s pelvis is adequate to give birth to a newborn the size of Pan” (Tague and
Lovejoy, 1986:252) with no more constriction than many living anthropoids. The
implication is that secondary altriciality was not present in australopithecines
(contra Jordaan, 1976b, and Fisher, 1982). In arguing that pelvic morphology in
australopithecines shows adaptations to locomotor and posture related functions,
including visceral support. Tague and Lovejoy (1986)did not argue that birth was
necessarily “easy” in these hominids or that no modifications due to obstetrical
constraints had been made. The wide pelvis in Australopithecus was not exclu-
sively (or even largely) a n obstetric adaptation, but it had obstetrical implications.
The unique birth mechanism of A.L. 288-1 and other australopithecines was not a
product of encephalization or secondary altriciality (since these later human ad-
aptations had not yet evolved) but resulted from obstetrical constraints in a small-
brained hominid which had undergone massive alterations in pelvic morphology a s
a result of the shift to habitual bipedalism.
What is the advantage of the extreme platypelloidy seen in A.L. 288-1 relative
to modern humans? Lovejoy (1988) and Tague and Lovejoy (1986) argued that a
transversely oval pelvis functioned to accommodate and support the abdominal
contents and viscera in a n upright hominid. They suggested that morphological
7Trevathan (1988) argued that although Tague and Lovejoy may be correct that rotation of the fetal cranium
probably did not occur in australopithecines, the fetal head would have had to rotate after it passed through the
outlet so that the shoulders could pass through the inlet. She suggested that this would have prevented shoulder
dystocia (obstruction of the shoulders) which is associated with increased mortality of both mothers and infants
(Harris, 1984; Smeltzer, 1986). She maintained that delivery of the shoulders may have been a source of
difficulty (and hence a source of selection) in hominids like australopithecines and Neadertals, in whom shoul-
ders may have been broad.
modifications to bipedalism such as rotation of the ilia, narrowing the distance

between the sacroiliac and hip joints, tilting the pelvis to align these joints in a
vertical plane, and displacing the false pelvis sagittally (due to lumbar lordosis) all
reduce the volume of the false pelvis. One “anatomical solution to maintain the
capacity of the false pelvis is simple: flare the ilia laterally and broaden the
sacrum” (Tague and Lovejoy, 1986:250).
Several other suggestions have been made to explain the width of the pelvic inlet
and overall pelvis in A.L. 288-1 and probably other australopithecines. Abitbol
(1987b)maintained that locomotor selection was more important in producing the
morphology of the early hominid pelvis th an obstetrics. Rak (1991) suggested th a t
the wide pelvic inlet is an adaptation to reducing vertical displacement of the body
mass during bipedal gait in a short-legged hominid. Ruff (1991) argued that the
great relative width of the entire pelvis in australopithecines is consistent with
thermoregulatory constraints on body form observed in modern humans. Specifi-
cally, Rull suggested that the decrease In biiliac breadth relative to stature which
followed in the evolution of Homo erectus and modern humans resulted from the
necessity to maintain “physiological equivalence” (i.e., surface-to-volume ratios) in
hominids of different body sizes who inhabited the same climate. Clearly, the
ultimate cause of great pelvic diameter in australopithecines remains subject to
debate. Nevertheless, the consensus is that the cause of this aspect of pelvic mor-
phology was not obstetric in nature. However, such a pelvic form had profound
obstetrical consequences for australopithecines, resulting either in a different
mechanism of birth from either later humans or nonhuman primates (Tague and
Lovejoy, 1986; Lovejoy, 1988) or in degree of difficulty during labor (Berge e t al.,
1984).
Leutenegger (1977) argued that since the newborn in A. africanus was small in
relation to the size of the inlet in the “early stages of hominid evolution selection
pressure for enlarging the birth canal may have been minor or absent” (p. 310).
However, to say that encephalization following australopithecines placed obstet-
rical selection on hominids is not the same as saying obstetric selection was absent
in australopithecines. The morphological changes associated with the evolution of
bipedalism required changes in the pelvis for obstetric reasons even though en-
cephalization had not yet occurreds.
Homo erectus
One reasonably complete pelvis (KNM-WT 15000) is known for Homo erectus
(Brown et al., 1985). As also noted by Trinkaus and Tompkins (1990, footnote 7), its
usefulness for understanding obstetrics in extinct hominids is limited a s the pelvis
belongs to the skeleton of a male who died at approximately 12 years of age. The
estimated diameter of this individual’s adult pelvic aperture led Brown et al.
(1985:791) to suggest that in “early Homo erectus birth canal diameters would
have been significantly smaller than in H . sapiens and that passage of a modern-
sized, full-term fetus would have been impossible.” Brown e t al. (1985), Gould
(1985), and Shipman and Walker (1989) suggested that KNM-WT 15000 provides
evidence for the existence in Homo erectus of a human pattern of fetal and post-
natal brain growth, a gestation length that was “effectively shortened relative to
brain size” (Shipman and Walker, 1989:389), and secondary altriciality.
Following the description of KNM-WT 15000 (Walker and Leakey, 1993) and
their own pelvic reconstruction (Walker and Ruff, 19931, Walker and colleagues
have developed this argument. Assuming that the specimen is correctly sexed, that
growth in adolescent Homo erectus would have followed a modern human pattern,
that pelvic sexual dimorphism was approximately the same in Homo erectus as in
‘A very diff‘erent sort of hypothesis about the relationship between bipedalism and human childbirth was
offered by Quieroz do Amaral(1989) who suggested that in addition to putting constraints on the pelvic aperture,
bipedalism makes the human attern of pregnancy and childbirth difficult by increasing the rate of spontaneous
abortion and presenting “probyems of locomotion in the last months of pregnancy” (Quieroz do Amaral, 1989:341.
modern humans, and, finally, that the reconstruction of the pelvic inlet is correct
despite the fragmentary nature of the pubic bones (Walker and Leakey, 1993).
Walker and Ruff (1992) estimated the transverse diameter of the pelvic inlet in
adult female Homo erectus. Using data from modern populations to estimate the
thickness of soft tissue, the authors calculated that a fetus with a cranial size
similar to a modern 32-33-week human fetus would correspond to the inlet di-
mension. They concluded that in order to achieve adult Homo erectus cranial ca-
pacity, this species must have been characterized by a modern human-like pattern
of growth (in which rapid brain growth continues after birth) rather than an
ape-like pattern of growth (in which the timing of birth corresponds to a decrease
in brain growth rate).
Walker and Ruff (1993)proposed this as a test of Martin’s (1983) hypothesis that
the pelvic limitation of increased fetal brain growth was reached approximately
1.5 millien years ago. Martin had argued that until that time train s i ~ einciease
could have taken place simply by increasing the fetal rate of growth rather than by
altering the postnatal rate. This would mean that “secondary altriciality” evolved
after 1.5 million years ago. Martin concluded from this that “increasingly elaborate
parental care was required in Homo erectus and then still more in Homo sapiens to
cater for the increasingly helpless condition of the infant during the first months
of life” (Martin, 1983:49). However, Walker and Ruff (1993) maintained that in
Homo erectus 1.6 million years ago, the birth canal enlargement and related
changes in the hip complex which are seen in later hominids, had not begun, but
secondary altriciality and the modern pattern of fetal growth was “almost if not
fully as derived as that in modern humans” (Walker and Ruff, 1993). This would
mean that in Homo erectus, fetal rates of brain growth continued after birth as in
modern humans.
Ruff (1991) suggested that the KNM-WT 15000 pelvis represents a compromise
between obstetric, thermoregulatory, and biomechanical constraints on trunk
width, birth canal width, and hip mechanics. He proposed that in Homo erectus a
nonrotational mechanism of birth would have been impossible and hypothesized
that the modern human pattern of rotation of the fetus within the birth canal had
probably already evolved.
Archaic Homo sapiens
Given the number of publications devoted to the subject of birth in Neandertals
(Anderson, 1989; Rosenberg, 1986, 1988, 1989; Tague, 1992a; Trinkaus, 1976,
1984) we know surprisingly little about the shape of the birth canal and the
mechanism of labor in these homiiiids.
Although characteristic aspects of pelvic morphology in Neandertals have been
known for some time (Endo and Kimura, 1970; Heim, 1972, 1982,1983; McCown
and Keith, 1939; Smith, 1976; Stewart, 1960, 1963, 1977; Trinkaus, 1976;
Trinkaus and Howells, 1979), it was not until the 1980s that the first suggestions
were made that it might be possible to make inferences from this morphology to an
understanding of Neandertal obstetric constraints. Several hypotheses depended
on the assumption that the elongated superior pubic ramus in Neandertals was
associated with an enlarged birth canal. This followed from the facts that the
superior pubic ramus forms the front (ventral) wall of the bony birth canal in
humans and that it is longer in human females than in human males (Hanna and
Washburn, 1953; Thieme and Schull, 1957; Washburn, 1948, 1949).
The hypotheses differed in their explanations of why Neandertals would have
had larger birth canals than anatomically modern or more recent humans. Wolpoff
(1980) proposed that Neandertals had birth canals which were large relative to
their stature (compared to anatomically modern humans) because their crania
(adult and, by inference, fetal) were large relative to their stature. The gestation
length hypothesis (Trinkaus, 1983a,b, 1984, 1985, 1986, 1987) stated that Nean-
dertal neonates were large compared to modern neonates because they were born
at a later stage in their development than modern humans, namely after a gesta-
114 Y E A R B O O K OF P H Y S I C A L A N T H R O P O L O G Y [Vol. 35, 1992
tion of 11-12 months rather than 9 months.9 The accelerated fetal growth hypoth-
esis (Dean et al., 1986) suggested that Neandertal neonates were larger than those
of anatomically modern humans because Neandertals “emphasised a greater pro-
portion of total brain growth in utero but had gestation periods similar to modern
humans (Dean et al., 1986:307).The maternal body weight hypothesis (Rosenberg,
1986, 1988) stated that because Neandertal mothers were muscular and hence
heavy but short, they had large infants relative to their stature. Rosenberg argued
that the shift in body proportions in the transition to anatomically modern humans
was accompanied by a decrease in the size of the newborn relative t o maternal
stature and a resulting decrease in birth canal dimensions.
These hypotheses10 all accepted the assumption that the elongated pubis in
Neandertals was accompanied by a proportionately increased birth canal, al-
though no fossils which preserved all the relevant dimensions of the pelvic aper-
ture existed. Further, the hypotheses all accepted the assumption that Neandertal
neonates were larger than anatomically modern neonates, but they differed in
their explanations of such a difference in birth size (and in the implications for
Neandertal behavior, especially with regard to growth and development).
The various explanations of large neonatal size in Neandertals generated con-
siderable controversy in the literature (Anderson, 1989; Arensburg, 1989; Bask-
erville, 1989; Frayer, 1985; Greene and Sibley, 1986; Ivanhoe, 1985; Martin and
MacLarnon, 1991; Rak, 1990; Rak and Arensburg, 1987; comments to Rosenberg,
1988; Rosenberg, 1989; Stringer et al., 1990; Tompkins and Trinkaus, 1987;
Trinkaus, 1985, 1987). However, the validity of all of these obstetrically based
functional hypotheses was questioned by the analysis of a Neandertal discovered
in 1983 at Kebara in Israel (Bar-Yosef et al., 1986). Although it was a male,
Kebara 2 was the first Neandertal specimen which preserved a reasonably com-
plete pelvic aperture. The specimen exhibited the pelvic features previously de-
scribed as characteristically Neandertal (elongated and thinned superior pubic
ramus) based on more fragmentary specimens. Arensburg (19891, Rak and Arens-
burg (19871, and Rak (1990) argued that although Kebara 2 has a superior pubic
ramus which is elongated compared to modern humans, it did not have a propor-
tionately enlarged pelvic inlet. This suggested to them that the pelvic features
characteristic of Neandertals were not related to an increased pelvic capacity and
therefore were not evidence of larger neonates in Neandertals than in anatomi-
cally modern humans. Rak (1990)proposed that Kebara 2 differed from the Qafzeh
9 specimen as well as from more recent humans in that the inlet was more ante-
gMuch speculation about the evolution of gestation length in hominids has accompanied discussions of the
origin of secondary altriciality, and the need for intensive parental care in humans. Published estimates of
gestation lengths for living hominoid species are as follows: Hylobates klossi, 210 days (Harvey and Clutton-
Brock, 19851, Hylobates lar, 205 days (Harvey and Clutton-Brock, 1985), Symphalangus syndactylus, 231 days
(Harvey and Clutton-Brock, 19851,Pongopygmaeus, 250 days (Martin and MacLarnon, 1990), 260 days (Harvey
and Clutton-Brock 1985) and 275 days (Leutenegger, 1973a1, Pan troglodytes, 225 days (Leutene ger, 1973a),
228 days (Harvey ’and Clutton-Brock, 1985), and 235 days (Martin and MacLarnon, 19901, Gorifh, 256 days
(Harvey and Clutton-Brock, 1985),260 days (Martin and MacLarnon, 1990), and 268 days (Leutenegger, 1973a1,
and Homo sapiens 267 days (Harvey and Clutton-Brock, 1985; Martin and MacLarnon, 1990) and 280 days
(Leutenegger, 1973a). Humans therefore have gestation lengths quite close to that of living pongids. Something
close to this figure is very likely the primitive condition for hominids as well. Therefore, human gestation length
is not “short” in absolute terms, but is only short relative to brain size. This has surely been accomplished more
by an increase in brain size (which is well documented in the fossil record) than by a decrease in estation length.
There may be no reason then to suppose that human gestation length was ever longer (in absofute terms) than
it is today. If anything, human gestation length has likely increased slightly since the divergence from other
hominoids (Martin and MacLarnon, 1990). Although some authors (Gould, 1975; Shipman and Walker, 1989;
Walker and Leakey, 1992; Walker and Ruff, 1992) have been careful to speak of humans as having a “relatively
short gestation length,” it has often been assumed that human gestation length has shortened in absolute terms
and that a 12 month or longer gestation length was probable in our evolutionary past. There is no evidence
to suggest that this was the case.
’OIn addition, two hypotheses accepted the assumption that the elongated pubis in Neadertals was evidence
of an increased birth canal, but not the assum tion that this was related to larger neonatal size. Minugh-Purvis
(1988) suggested that thicker cranial vaults ofNeandertals could have made molding of the skull less likely and
parturition more difficult. She hypothesized that this could “explain the need for a larger pelvic aperture in
Neandertals” (Minugh-Purvis, 1988:260).Friedlander and Jordan (unpublished)have suggested that increased
bone density and robusticity in Neadertal mothers and infants might have made both the birth canal and the
fetal head less malleable, requiring a larger pelvic aperture in Neandertals than in modern humans.
riorly situated in relation to the acetabula and that the acetabula were more
laterally facing. He suggested that it was these locomotor related differences that
were related to the pubic elongation rather than a n enlarged pelvic inlet.11 He
concluded that Kebara 2 (and by extension, Neandertals as a group) differed from
later humans in “locomotion and posture-related biomechanics” (Rak and Arens-
burg, 1987).
How small is the pelvic aperture of Kebara 2? Tague (1992) systematically
compared Kebara 2 to a range of human populations not only at the pelvic inlet but
at all of the planes of the pelvic aperture. He showed that compared to modern
males, Kebara 2 is spacious in transverse and anterior-posterior diameters and
circumference12 of the pelvic inlet (greater than 94.1%, 95.8%, and 91.5% of the
males in his sample respectively). However, a t the midplane and outlet, Kebara 2
is actually constricted compared to modern males. For the index of pelvic funnel-
ling Lcoutlet circumferenceiinlet circumference) X lUUl, Kebara Z (74.9) is consid-
erably more funnelled than modern males (87.6 -t 4.8).13The index of outlet shape
[(anterior-posterior diameterkransverse diameter) x 1001 of Kebara 2 is 77.2,
whereas for modern males it is 114.6 2 14.4 (N = 181) and for modern females
102.2 2 13.8 (N = 154) (Tague, 1992). If Neandertal females were more elliptical
in this dimension than Neandertal males (which would have occurred if they
exhibited the same type of dimorphism as is present in modern humans), it is
unlikely t h a t they would have been capable of giving birth to a large brained
infant.
Figure 6 summarizes the shape of the pelvic aperture in Pan, Australopithecus
(A.L. 288-l), a Neandertal (Kebara 2), and modern Homo sapiens. Note that Ke-
bara 2 is shown as having a transversely oval outlet. Whether this outlet shape
would also occur in female Neandertals is obviously unknown.
Because the Kebara 2 specimen is male, like KNM-WT 15000, it is not ideally
suited for drawing conclusions about birth in a n extinct human population (Rosen-
berg, 1988). Nevertheless, Kebara 2 is by far the most complete Neandertal spec-
imen we have to date and provides a n opportunity to formulate hypotheses about
Neandertal female pelvic morphology. Tague (1992) suggested that while Nean-
dertal males and females probably had spacious pelvic inlets, it is likely t h a t at the
outlet, both sexes were at the lower end of the modern range of variation. If the
discovery of pelvic remains of Neandertal females confirms that they did have
relatively constricted pelvic outlets, then explanations of Neandertal pubic mor-
phology which predict larger newborns in Neandertals would have the implication
of less favorable cephalo-pelvic relations in Neandertals than in modern humans,
suggesting more difficult births (Tague, 1992). Interestingly, this is the opposite
of what Trinkaus (1984) suggested when he hypothesized that the transition from
Neandertals to anatomically modern humans might have been accompanied by a n
increase in cephalo-pelvic constriction and a compensatory improvement in obstet-
rical and neonatal care. Given the high risks associated with birth difficulties
today, a n explanation invoking even great difficulties in the past appears unlikely.
It is useful to remember, however, that we do not know what pelvic sexual
dimorphism was like in Neandertals. Assuming that our sample is representative,
Neandertals differed from living humans in that (at least for the small sample of
individuals who preserve a pubic bone), females actually had shorter pubic bones
“A very preliminary test of this hypothesis by Rosenberg (1990) failed to find a correlation within a single
population of modern humans (Central Europeans) between pubic length and several measures of pelvic aper-
ture position relative to the acetabulum. This hypothesis awaits further testing.
“Note that circumference of the pelvic inlet is calculated by taking the curved distance of linea terminalis,
doubling it and adding sacral breadth. Pubic length is the anterior component of that dimension.
I3Because pelvic circumferences and pelvic funnelling are measures of low variability in modern humans and
living primates, Tague (1992) suggested that they were probably of low variability in Neandertals. This in-
creases the confidence with which we treat a measurement from a single fossil specimen as representative for
its species.
Pan Ausrrubpirhecus Nadertal Homo

troglodytes (AL 288-1) (Kebara) Sapiens
cr"
? ? ?
Met
E
Midplane
6
Outlet
~ E
Fig. 6. The shape of the pelvic aperture in Pun, Austrulopithecus (A.L. 288-1), a Neandertal (Kebara 21,
and modern Homo supiens. Data are taken from Tague and Lovejoy (19861, and Tague (1992). Scale is
approximate. Data for Pun and Homo are averages for females based on sample sizes of 25 and 119
respectively. Arrows indicate the direction of the longest axis of the pelvic aperture at each plane. The
pelvic outlet in Pun (shown in lighter outline) is accurate in shape but not homologous to the pelvic outlet
in bipeds because of the position of the sacrum. Note that in Pun the pelvic aperture is long in the
anterior-posterior direction at all planes. In Austrulopithecus, the pelvic aperture is elongated trans-
versely at all planes. The Neandertal and modern human are more complicated. Data from a single mule
Neandertal suggests that the pelvic inlet was long transversely as in other bipeds, but the outlet was
possibly also transversely long. Finally, in modern human females, the inlet is long transversely, the
midplane is long in the anterior-posterior dimension, and the outlet is approximately the same length in
both directions.
than males (Rosenberg, 1986,1988). Obviously, hypotheses about birth and about
obstetrically related anatomy in Neandertals must await the discovery of more
complete female remains.
Clearly, obstetric explanations are not adequate to account for all characteristic
aspects of Neandertal pelvic anatomy. For example, the extreme thinning of the
superior pubic ramus has never been satisfactorily addressed by any workers on
this morphology. However, it would be a mistake to reject ubsletric or reproductive
demands as one of a number of selective forces acting on the pelvis.
The large-brained Neandertals almost certainly experienced the same mecha-
nism of birth as exists in modern humans. This is likely since evidence suggests
that the origin of the human-like orientation of the fetal cranium in the pelvic inlet
was present in australopithecines (Tague and Lovejoy, 1986) and the rotational
aspect of the birth mechanism may have been present by the time of Homo erectus
(Ruff, 1991). The aspect of Neandertal birth that we know least about is the rel-
ative spaciousness of the Neandertal birth canal compared to modern humans. In
Neandertals, as in all species, pelvic morphology is a compromiseto selective forces
arising from locomotion and obstetrics as well as other factors. When we under-
stand more about the implication of possible locomotor differences between Nean-
dertals and modern humans we will be in a better position to evaluate the obstetric
implications of the small differences between Neandertals and modern humans in
pelvic morphology.
When did modern human pelvic dimorphism arise?
Sexual dimorphism in the human pelvis has been studied in a wide range of
recent populations (Davivongs, 1963; Day and Pitcher-Wilmott, 1975; Derry, 1923;
Genoves, 1954; Hager, 1989; Hanna and Washburn, 1953; Hauser and Jahn, 1984;
Holland et al., 1982; Ince and Young, 1940; Krukierek, 1951; Moerman, 1981;
Novotny, 1982,1983,1986; Orban, 1982; Orban-Segebarth, 1984; Schulter-Ellis et

al., 1983, 1985; Rosenberg, 1986; Segebarth-Orban, 1980; Seidler, 1980; Tague,
1986, 1991, 1992a; Thieme and Schull, 1957; Thoms and Greulich, 1940; Wash-
burn, 1948, 1949; Wu, 1982; Young and Ince, 1940). If pelvic sexual dimorphism is
in fact a response to differential selection on males and females, as suggested by
Schultz (1949) and supported by Leutenegger 1974,1982a), and if t h a t differential
selection is partly the result of encephalization which took place only recently in
human evolution, then a relevant question arises: When did the modern pattern of
pelvic sexual dimorphism first appear?
This question was addressed directly by Hager (1989) who argued that if modern
patterns of pelvic dimorphism are the result of differential selection on males and
females caused by obstetrics, then one would not expect modern patterns of dimor-
phism to be present in extinct hominids, especially those with small cranial ca-
pacity. She suggested that australopithecines may have had pelvic sexual dinmi--
phism more similar to that seen in living great apes. Hager is correct in pointing
out that we do not know (and should not make assumptions about) what pattern of
pelvic dimorphism characterized early hominids. It is important to reiterate, how-
ever, t h a t encephalization is not the only factor which might cause obstetric se-
lection. Modifications of the pelvis which occurred with the evolution of bipedalism
also placed obstetric constraints on the female pelvis and may have resulted in
pelvic dimorphism in australopithecines. Clearly the discovery of more complete
male australopithecine pelvic materials will be necessary before we can address
this issue.
The presence or absence of the ventral arc on the pubic is used in modern
humans as a reliable sex indicator (Phenice, 1969; Sutherland and Suchey, 1991).
Both Sts 14 and A.L. 288-1 have a ventral arc (Tague, personal communication)
which could indicate the existence of some modern sex differences among austral-
opithecines. Alternatively, since this feature is correlated with pubic length (Bu-
dinoff and Tague, 1990), i t may be a result of the relatively long pubic bones in
those specimens. Since no male pubic bones exist for australopithecines and since
elongated pubic bones appear to be the primitive condition for hominids (Frayer,
1985; Hager, 1989), it is impossible to say whether the presence of a ventral arc on
Sts 14 and A.L. 288-1 is related to the fact that they are females, whether i t is part
of the primitive condition, or whether i t would have been as accurate a n indicator
of sex among australopithecines as i t is in modern humans.
Tague and Lovejoy (1986) examined a series of pelvic measures which differ
significantly between males and females in modern human populations: the an-
gulation of the sacrum, the index of pelvic funnelling, the subpubic angle, and the
length of the iliopectineal line. All of these have the effect of increasing the size of
the pelvic aperture in females compared to males. Tague and Lovejoy (1986) found
that A.L. 288-1 which is sexed a s a female on nonpelvic grounds (Johanson and
White, 1979) is similar to modern males in all of these features except length of the
iliopectineal line. They concluded from this that the pattern of pelvic dimorphism
seen in modern humans was not yet present in australopithecines. Since our
knowledge of male australopithecine pelvic morphology is limited, we are not able
to argue that australopithecines had no pelvic dimorphism, just that they did not
exhibit the modern human pattern.
It is likely that modern patterns of pelvic dimorphisms began to appear by the
origin of Homo (Hager, 1989), when brain size increase had begun to place greater
differential selection on males and females than had been present among smaller
brained hominids. Like the modern human pattern of birth, pelvic sexual dimor-
phism evolved in a complex mosaic manner.
SUMMARY
An examination of the fossil and comparative evidence for the evolution of hu-
man birth provides support for the mosaic nature of this evolutionary process.
Different aspects of human birth did not evolve simultaneously. There was no
118 YEARBOOK OF PHYSIGAL ANTHROPOLOGY [Vol. 35, 1992
single obstetrical dilemma (Washburn, 1960)in human evolution. Rather the mod-
ern pattern of human childbirth evolved in a complex way (summarized below),
with particular aspects of that pattern evolving as obstetric responses to other
evolutionary changes in the skeleton.
1. The modern human-like position of engagement of the fetal head in the pelvic
inlet evolved early in hominid evolution after the morphological modifications of
the pelvis necessary for bipedalism had taken place.
2. The modern human-like rotation of the fetus within the birth canal probably
did not evolve until later, after encephalization had increased. This rotation was
probably also associated with the occiput anterior position, and with behaviors
such as birth assistance. Similarly, increase in degree of difficulty during birth
also evolved along with increasing brain size.
3. Modern patterns of sex differences in the metrics of the pelvis (but not nec-
essarily sex differences in pelvic morphology per se) probably also evolved in a
complex manner following the beginning of encephalization.
4. Assistance during childbirth probably became more important as difficulties
increased and as such assistance would have improved the chances of a successful
birth outccme. In addition, the birth of increasingly altricial infants has meant the
presence in hominid groups of family members requiring intensive parental care.
Assistance in childbirth and care of infants are aspects of cooperative behavior
which may have profound effects on human social organization.
ACKNOWLEDGMENTS
For critical comments on this paper, I thank Pat Bridges, Jonathan Friedlander,
Celeste Gagnon, Marquisa LaVelle, William Leonard, Thomas Rocek, Lynne
Schepartz, Carol Ward, and especially Robert Tague. Three anonymous reviewers
improved the final draft of the manuscript. I am grateful to Nancy Friedlander,
Robert Tague, and Alan Walker for permission to cite unpublished work and to
William Leonard and Mary Williams for assistance in obtaining references. Fig-
ures were drawn by Teryl Lynn and Sue Quinn.
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10968644, 1992, S15, Downloaded from https://onlinelibrary.wiley.com/doi/10.1002/ajpa.1330350605 by CAPES, Wiley Online Library on [21/02/2023].

The Evolution of Modern Human Childbirth

KEY WORDS Obstetrics, Pelvis, Childbirth, Human evolution

ABSTRACT Human birth follows a pattern which is unique among

0 1992 Wiley-Liss, Inc.

in nonhuman primates and most other mammals, the probability of an observer

transverse diameter of the maternal pelvic inlet or obliquely (Cunningham et al.,

Presentation of the fetus. A key aspect of mechanism of birth is the presentation of

ischio-pubic index is a reflection of the differential elongation of the pubic bone in

TABLE 3. Length of labor in selected primate species’

during labor is by no means unique to humans (Trevathan, 1988) most primates

liveries on women with platypellic (anterior-posteriorly flattened) pelves resulted

head is transversely or obliquely oriented a t the pelvic inlet; in nonhumans it is

certainly seems as if this species would have been characterized by a relatively

In order to determine how spacious the pelvic inlet was in australopithecines,

TABLE 5. Pelvic aperture indices’ of modern humans and A.L. 288-1’

Mechanism of birth in australopithecines. Berge et al. (1984) and Berge and

transverse diameter (breadth between the ischial tuberosities) is approximately

modifications to bipedalism such as rotation of the ilia, narrowing the distance

Pan Ausrrubpirhecus Nadertal Homo

Novotny, 1982,1983,1986; Orban, 1982; Orban-Segebarth, 1984; Schulter-Ellis et

infant development in captivity. Anim. Behav. approach to problems of cephalopeivic dispropor-

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