New high-resolution computed tomography data of the
Taung partial cranium and endocast and their bearing
on metopism and hominin brain evolution
Ralph L. Hollowaya,1, Douglas C. Broadfieldb, and Kristian J. Carlsonc,d,e
a
Department of Anthropology, Columbia University, New York, NY 10027; bDepartment of Anthropology, Florida Atlantic University, Boca Raton, FL 33431;
c
Evolutionary Studies Institute, Palaeosciences Centre, and dSchool of Geosciences, University of Witwatersrand, Johannesburg 2050, South Africa;
and eDepartment of Anthropology, Indiana University, Bloomington, IN 47405
Edited* by C. Owen Lovejoy, Kent State University, Kent, OH, and approved July 16, 2014 (received for review February 17, 2014)
Falk and colleagues [Falk D, Zollikofer CP, Morimoto N, Ponce de
León MS (2012) Proc Natl Acad Sci U S A 109(22):8467–8470] hy-
pothesized that selective pressures favored late persistence of
a metopic suture and open anterior fontanelle early in hominin
evolution, and they put an emphasis on the Taung Child (Austral-
opithecus africanus) as evidence for the antiquity of these adaptive
features. They suggested three mutually nonexclusive pressures: an
“obstetric dilemma,” high early postnatal brain growth rates, and
neural reorganization in the frontal cortex. To test this hypothesis,
we obtained the first high-resolution computed tomography (CT)
data from the Taung hominin. These high-resolution image data
and an examination of the hominin fossil record do not support
the metopic and fontanelle features proposed by Falk and col-
leagues. Although a possible remnant of the metopic suture is ob-
served in the nasion–glabella region of the Taung partial cranium
(but not along the frontal crest), this character state is incongruent
with the zipper model of metopic closure described by Falk and
colleagues. Nor do chimpanzee and bonobo endocast data support
the assertion that delayed metopic closure in Taung is necessary
because of widening (reorganization) of the prefrontal or frontal
cortex. These results call into question the adaptive value of delay-
ing metopic closure, and particularly its antiquity in hominin evolu-
tion. Further data from hominoids and hominins are required to
support the proposed adaptive arguments, particularly an obstetric
dilemma placing constraints on neural and cranial development
in Australopithecus.
prefrontal reorganization | cranial capacity | human evolution
T he Taung natural endocast, a cast of the endocranial surface
of the cranium, has always been problematic because of
preservation and preparation. The endocast is distinguished by
numerous calcite deposits on its surface. Indeed, the midline
region is marked by numerous calcite deposits, indicating that
the true surface of the endocranium lies below these deposits
(Fig. 1). In addition, there are remnants of the endocranial table
adhering to the surface of the natural endocast, obscuring many
of its morphological features, including portions of its left frontal
lobe (Fig. 1). Finally, Dart’s exuberant use of his wife’s knitting
needles to extricate the fossil from the surrounding breccia may
have scarred the surfaces, forever obliterating some of the
original features of the natural endocast.
The metopic suture is a synostosis between the two halves of
the frontal bone, and although it is present in all mammals
during development, it normally fuses in humans shortly after
birth. Biological anthropologists most often refer to its persis-
tence in adults as a nonmetric trait (Fig. S1), the frequency of
which varies among different human populations (1). Its pres-
ence in primates has no documented relationship to the size and
form of the underlying prefrontal cortex, nor to corresponding
areas on endocasts. Specifically, known correlations between the
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presence of metopism/open anterior fontanelle and prefrontal/
frontal lobe reorganization have not yet been demonstrated.
13022–13027 | PNAS | September 9, 2014 | vol. 111 | no. 36
Falk and colleagues (2) recently argued in support of such
relationships. They proposed that delayed fusion of the metopic
suture coupled with delayed closure of the anterior fontanelle
provided a selective advantage during hominin evolution. The
advantages, they reasoned, stemmed from mutually nonexclusive
evolutionary pressures resulting from an “obstetric dilemma,”
high early postnatal brain growth rates, and reorganization in the
frontal neocortex (2: 8469). Falk and colleagues placed substantial
emphasis on the Taung Child (Australopithecus africanus) in their
arguments, as of the 63 fossil hominin crania they examined for
a metopic suture (2, Table S3), Taung was the only australopith in
which they observed a partial or unfused metopic suture. Therefore,
their argument that a persistent metopic suture and open anterior
fontanelle constitute an adaptation that extends substantially
backward in time within the hominin lineage (i.e., into pre-Homo
lineages) rests entirely on the status of these features in the Taung
Child. Given that they play such a prominent role in these recent
claims (2), we suggest it is prudent to critically reexamine the state
of these features in the Taung Child. In doing so, we offer high-
resolution computed tomography (CT) image data of the specimen.
Prefrontal widening, although perhaps indicating a volume
increase (assuming no proportional decreases in lobe/region
volumes elsewhere), is an insufficient indicator of cortical re-
organization. Rather, documenting cortical (prefrontal/frontal)
reorganization in brains of early hominin taxa requires the dem-
onstration of differences in endocast convolutional details that can
be traced to underlying cytoarchitectonic changes. Logically, it
Significance
The Taung Child (Australopithecus africanus) has historical and
scientific importance in the fossil record as the first and best
example of early hominin brain evolution. It was recently
proposed that Taung exhibits adaptive morphology (e.g., per-
sistent metopic suture and open anterior fontanelle), permit-
ting important postnatal brain growth late into infancy. As
Taung provides the only purported pre-Homo fossil evidence
for the suggested adaptive mechanism, we test the hypothesis
that it displays these features. Using new high-resolution
images and in silico exploration, we did not observe the hy-
pothesized form of these features. If delayed metopic suture
closure were adaptive, for example, permitting substantial
postnatal brain growth and alleviating an obstetric dilemma,
there is no evidence this mechanism evolved before Homo.
Author contributions: R.L.H., D.C.B., and K.J.C. designed research, performed research,
analyzed data, and wrote the paper.
The authors declare no conflict of interest.
*This Direct Submission article had a prearranged editor.
Freely available online through the PNAS open access option.
1
To whom correspondence should be addressed. Email: rlh2@columbia.edu.
This article contains supporting information online at www.pnas.org/lookup/suppl/doi:10.
1073/pnas.1402905111/-/DCSupplemental.
www.pnas.org/cgi/doi/10.1073/pnas.1402905111

Fig. 1. Superior view of the original Taung partial cranium and endocast.
The frontal portion faces toward the bottom of the image. The bregmatic
region of the natural endocast is roughly 3 cm posterior to the broken edge
of the frontal bone in this image. Calcite deposits (black arrows) filling su-
tural voids during the fossilization process can be seen on the surface of the
natural endocast, most visibly along the right coronal suture. The left hemi-
sphere of the natural endocast is covered by remnants of the endocranial table
to varying extents and thicknesses. It appears that remnant bone covers the
same type of calcite infill associated with the left coronal suture, whereas such
infill is not as visible at the medial-most margin of the remnant bone anterior
to bregma (i.e., location of the purported metopic suture).
must be remembered that internal prefrontal/frontal reorgani-
zation (e.g., cytoarchitectonic changes) may occur without any
external evidence manifested on the surface of a brain or on its
representation via the endocast surface. Nonetheless, the pre-
frontal region of the Taung endocast has not been previously ar-
gued to share any demonstrable reorganizational features in sulcal
morphology with Homo to the exclusion of chimpanzees or other
apes, particularly in the third inferior frontal convolution (3, 4).
Indeed, some have steadfastly claimed that the prefrontal region
of the Taung endocast surface was apelike (5). Neural re-
organization of the australopith brain toward a derived Homo
configuration remains an intriguing possibility, however, given
the recent claim (6) that the presumably later australopith
from Malapa (i.e., MH1), Australopithecus sediba, has an endo-
cast that may display some reorganization in surface features of
its third inferior convolution.
Because the Taung Child is the only australopith put forth in
the adaptationist argument for metopism in hominins (2), critical
evaluation of this specimen is necessary to verify the accuracy
and evolutionary significance of these claims. Here we offer di-
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rect evidence for verifying the purported metopic suture and
open anterior fontanelle (2, 7), evidence that is less limited than
Holloway et al.
the image data available to Falk and colleagues (2) and less
affected by the aforementioned well-known preservation
issues. Specifically, we use these data to assess whether the
sharp-edged remnant of cortical bone covering much of the
left half of the frontal region of the natural endocast reflects
an open metopic suture, as has been suggested (2, 7). If direct
evidence for such a metopic suture is to be found, then we
would predict that any remaining evidence of a metopic suture
in the existing frontal should be found in more caudal regions
and should be continuous with the extent of the metopic su-
ture suggested by Falk and colleagues on the natural endocast
(i.e., support a “zipper” model of anterior to posterior closure),
and that evidence of a protruding ridge along the midline of
the natural endocast surface extending rostrally from bregma (or
rostrally from the purported open anterior fontanelle) should be
observable to some unknown extent, indicating the presence of
an open metopic suture that became filled by intrusive matrix
during formation of the natural endocast.
Results
A metopic suture is visible neither on the external table of the
frontal bone (Fig. 1) nor on the rendering of the partial cranium,
with the possible exception of the region between nasion and
glabella (Fig. 2 A and B). A coronally oriented virtual section
passing approximately through the rostral-most region of the
cranium receiving the frontal poles of the endocast (Fig. 2C),
which are visualized as two asymmetric elliptical regions (darker
areas) surrounded by diploic bone (lighter areas), displays a thin
suture internally connecting the periosteal surface near nasion
with the endocranial surface near the left frontal pole. Note that
the frontal squama in this virtual section (Fig. 2C) is uninterrupted
ANTHROPOLOGY
by such a feature. This internal structure appears to be manifested
superior to nasion on the external table of the cranial rendering as
a remnant metopic suture (Fig. 2B). Additional internal evidence
of a metopic suture (Fig. S1) in the frontal of the Taung partial
cranium is absent in a virtual section through glabella and through
virtual sections between glabella and the posterior break (Fig. 2 C
and D).
Fig. 3A illustrates the location of representative coronally
oriented virtual sections through the Taung natural endocast.
The thin remnant of the endocranial table covering the left side
of the frontal region can be differentiated from the underlying
surface of the natural endocast in the high-resolution image data
(Fig. 3B). In each of the illustrated sections, the remnant layer
comes to an abrupt end near the midline (i.e., immediately left of
the leftmost of the white arrows). In other words, there is not a
separate superiorly projecting ridge of matrix at each terminus
rising from the natural endocast beneath the bone, as would be
expected if an open (or partially open) metopic suture was
present during the formation of the natural endocast. Rather,
uniform material contrast throughout the remnant layer until its
sudden cessation at the midline suggests a protruding ridge was
absent, unlike what is observable on the natural endocast at the
right coronal suture (Fig. 1).
High-magnification stereomicrographic pictures also were
taken of the midline surface of the Taung natural endocast at
multiple locations between its midfrontal region and bregma to
independently assess evidence for the hypothesized protruding
ridge-like structure (Fig. S2). These images also provide no con-
clusive evidence of a protruding ridge similar to that appearing at
the right coronal suture. In general, these images corroborate the
coronally oriented virtual sections (Fig. 3B) in showing no support
for a superiorly projecting ridge of calcite rising from the midline
of the natural endocast rostral to bregma or rostral from the
purported open anterior fontanelle.
It is important to appreciate that a large degree of variation
characterizes the small sample of additional A. africanus endocasts.
Comparing them with a sample of 40 bonobo (Pan paniscus)
PNAS | September 9, 2014 | vol. 111 | no. 36 | 13023
 Fig. 2. Rendering of the Taung partial cranium produced from high-resolution image data obtained in this study, as well coronally oriented virtual sections
through the frontal of the specimen. The rendering (A) and close-up of the nasion–glabella region (B) were generated from volume data consisting of
isotropic 55.5 μm voxels. Three black arrows (B) arranged vertically indicate the approximate extent of a potential metopic suture persisting in the frontal.
Coronally oriented virtual sections through the partial cranium obtained at glabella (g), nasion (n), and midfrontal (mf) locations are depicted in C. The black
arrow in the virtual section through nasion indicates internal evidence of the potential persisting metopic suture. A coronally oriented virtual section through
the posterior region of the frontal, just anterior to its broken edge, is depicted in D. Note the absence of internal evidence (e.g., see C and Fig. S1) for
a persistent metopic suture in the midfrontal (mf) and posterior frontal (pf) sections.

endocasts from the collection of R.L.H. is instructive. Maximum
width measurements of australopith prefrontal regions across the
breadth of third inferior convolutions are: Taung, 68 mm;
Sterkfontein (Sts) 60, 72 mm; Sts 71, 72 mm; Sterkfontein type 2,
78 mm; Sts 5, 80 mm; and Sterkfontein (StW) 505, 80 mm.
Where only a single side is available, width was doubled to
obtain these full breadths. Two endocasts [endocranial volumes
(EVs) of 359 and 372 mL] randomly chosen from among the 40
bonobos have maximum prefrontal widths of 73 and 74 mm,
respectively. A third bonobo endocast (EV, 403 mL) with
roughly the same EV as Taung has a prefrontal width of 77 mm.
None of the 40 bonobo endocasts in the sample display third
inferior convolutional reorganization beyond what is usually
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argument of Falk and colleagues regarding prefrontal widening
indicating reorganization (2), one would have to conclude that
the bonobo prefrontal was reorganized and more advanced than
the prefrontal of several australopiths. Moreover, when aug-
menting the bonobo sample with an additional 35 chimpanzee
(Pan troglodytes) endocasts from the collection of R.L.H., the
combined mean EV becomes 364 mL (SD, 43 mL), with a com-
bined mean maximum prefrontal breadth between Broca’s caps
(third inferior convolutions) of 68.5 mm (SD, 4.3 mm). The
Spearman’s rho correlation (rs) between EV and prefrontal
breadth in this sample of 75 Pan endocasts is 0.678 (P < 0.001).
Clearly, convolutional patterns provide a more conclusive means

observed in anthropoids. If one compares these bonobo endocasts of comparing prefrontal/frontal organization than widening
with those of A. africanus mentioned earlier and applies the or EV.

13024 | www.pnas.org/cgi/doi/10.1073/pnas.1402905111 Holloway et al.


Fig. 3. Rendering of the Taung natural endocast produced from high-res-
olution image data obtained in this study, as well as coronally oriented
sections through the specimen. The rendering (A) was generated from vol-
ume data consisting of isotropic 65 μm voxels. Coronally oriented virtual
sections through the natural endocast (B) were obtained at two frontal
locations, one anterior (fa) and one posterior (fp). White arrows in each of
the two virtual sections point to the remnant endocranial table adhering to
the left hemispheric surface of the natural endocast. Note that the remnant
layer varies in thickness and appears to abruptly end immediately to the left
of the leftmost white arrows in each section. There is no discernible ridge of
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material protruding superiorly from the endocast surface in either location,
as occurs along the coronal suture.
Holloway et al.
Discussion
State of Metopic Suture in Taung. The form of metopic suture in-
dicated in the high-resolution image data set of the Taung partial
cranium is inconsistent with the form of metopic suture and open
anterior fontanelle described by Falk and colleagues (2). A
persistent metopic suture in the region between nasion and
glabella would contradict an anterior to posterior zipper model
of closure suggested by Falk and colleagues. Moreover, using the
high-resolution image data set of the Taung natural endocast, we
find no evidence of a distinct superiorly protruding ridge on the
Taung natural endocast indicating a possible open (or partially
fused) metopic suture rostral to bregma. Thus, neither of the
predicted features necessary to support the form of metopic
suture in the Taung Child suggested by Falk and colleagues (2)
are observed in the present study. Standard medical CT images do
not provide spatial resolution equivalent to that of the high-res-
olution image data set generated in the present study (Fig. S3),
and thus it would appear that Falk and colleagues (2) may have
been unknowingly constrained by this deficiency. According to
surface renderings produced from medical CT data (e.g., see
their figure 1), it would appear possible that they may have
interpreted the midline border of the remnant endocranial
table as a superiorly protruding ridge indicative of an open (or
partially fused) metopic suture.
In modern humans, the metopic suture normally closes 3–9 mo
postpartum (8, 9), whereas in Pan, all of the sutures (including the
metopic suture) normally close before birth (10, 11), notably earlier
than in Homo. This is likely because suture fusion is more strongly
related to dental development than it is to brain development (11, 12).
With an estimated age between 3.73 and 3.93 y at death, Taung is
well beyond the age for normal closure of the metopic suture exem-
ANTHROPOLOGY
plified by modern humans or chimpanzees and other primates
(11, 13), unless the suture were to persist as a nonmetric trait (1).
Given the results of the study and the accepted age of Taung, the
latter scenario would seem more likely than would a metopic suture
persisting as a selectively advantageous trait implicated in brain
development. Indeed, reorganization and development of the pre-
frontal and frontal cortex in humans exhibits little influence on fusion
of the metopic suture, as development of these areas continues
well beyond the age at which the metopic suture normally fuses.
The high-resolution image data generated in the present study
are no more definitive than the medical CT image data used by
Falk and colleagues (2) in determining whether there was an
open anterior fontanelle in the Taung Child, except that the
higher spatial resolution permits more precise visualization of
borders of the purported structure. The borders are not as sharp
as one would expect if the anterior fontanelle remained open
(Fig. 3A), but the aforementioned damage during extraction of
the fossil or decades of subsequent wear could be contributing
factors to the observed lack of sharpness. It is also worth noting
that an open anterior fontanelle would be extraordinarily rare if
Taung was indeed 3.5–4 y of age (13, 14), particularly if Taung
followed a growth pattern assumed to be more similar to chim-
panzees than humans (15, 16).
In addition to refuting the presence of an open metopic suture
and anterior fontanelle in the Taung Child, this evidence calls
into question the proposed adaptive significance of delayed
metopic suture and anterior fontanelle closure in hominin evo-
lution, particularly in australopiths. We suggest that a more
reasonable and parsimonious explanation for the observed form
of metopic suture in Taung (i.e., possibly retained between
nasion and glabella) would be its manifestation as an epigenetic
trait, as in modern humans (1) and many other primates.
Frontal Widening and Reorganization. Prefrontal widening in iso-
lation, although perhaps indicating a volume increase (assuming
no proportional decreases in lobe/region volumes elsewhere),
PNAS | September 9, 2014 | vol. 111 | no. 36 | 13025
 should not be considered indicative of cortical reorganization. As
demonstrated here, some great ape endocasts (e.g., those of
bonobos and chimpanzees) display prefrontal widening that
exceeds prefrontal breadths of Taung, Sts 60, and Sts 71 (Fig. S4
and Fig. S5) while simultaneously exhibiting EVs smaller than
that of Taung. These same Pan endocasts exhibit convolutional
details that are reflective of ape patterns and that are not more
derived toward Homo patterns than are the patterns exhibited by
the australopith endocasts.
Even considering an exclusively hominin sample of endocasts,
the relationship between prefrontal breadth and cortical orga-
nization is uncertain. Although illustrations of prefrontal outlines,
as shown in ref. 17, clearly indicate a greater degree of “point-
edness” to prefrontal regions of “robust” australopith endocasts
compared with prefrontal regions of “gracile” australopith endo-
casts (e.g., those of A. africanus), these outlines cannot be directly
translated into neurological/functional differences in cortical re-
organization, given the considerable differences in cranial and facial
development and architecture of these groups (18, 19). Convolu-
tional details of both robust and gracile forms also are simply mute
and do not allow conclusions regarding potential or actual differ-
ences in their neural organization.
Obstetric Dilemma and Metopism. As suggested by Falk and col-
leagues (2), it might seem reasonable to implicate persistent
metopism as an adaptive trait when facing an obstetric dilemma.
Tague and Lovejoy (20) and subsequent others (e.g., see ref. 21),
however, suggest that although hominin parturition may have
become more difficult compared with that of extant anthropoids
by the time of A. afarensis, the crucial dilemma would not have
occurred until encephalization began to increase dramatically in
Homo. Reconstructions of hominin pelves (e.g., Sts 14 and MH2)
support the absence of a crucial obstetric dilemma in additional,
later species of Australopithecus (20, 22). The absence of an adap-
tive metopic suture in Taung also supports the notion that this di-
lemma was not present in australopiths.
Although it is true that pelvic dimensions changed from Aus-
tralopithecus to Homo, in part to accommodate the larger-brained
babies of the latter, there is little reason to suggest that fetal and
postpartum development was under any more selective pressure in
australopiths than in anthropoids and modern humans. Also, given
the average size of the birth canal in Australopithecus (20), the
estimated size of the neonate brain, and the growth trajectory in
australopiths, based on dental and osteological data (23), there is
no evidence that Australopithecus experienced selective pressures
necessitating delayed brain or skull development greater than
those observed in modern humans. Rather, the recently described
A. sediba pelvis (22), despite an EV of 420 mL (6), suggests that
the physical constraints of even the late Australopithecus birth
canal probably had less effect on parturition and the timing of
suture closure than in modern humans. It is also worth noting that
an additional contributing factor to an obstetric constraint on fetal
brain size has been suggested by Dunsworth and colleagues (24),
who posited that a metabolic limit exists that restricts the ability of
a female to keep a fetal brain growing.
Metopism in Homo. Although the present study focused on the
issue of metopism in australopiths, and specifically in the Taung
Child, our experience with this specimen leaves us skeptical of
claims that many early Homo specimens (e.g., OH 24, KNM-ER
1805, KNM-KNM-ER 1813, KNM-ER 3883, 3733, and so on, as
listed in table S3 of ref. 2) clearly and without ambiguity also
demonstrate persistent (or partially fused) metopic sutures or
open anterior fontanelles. Falk and colleagues (2) do not de-
scriptively substantiate or illustrate these examples and identify
support as coming from low-resolution CT data, which in the
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case of Taung, proved to be insufficient (Fig. S3). Although they
(2) claim to have “confirmed” features described by Prat (25),
13026 | www.pnas.org/cgi/doi/10.1073/pnas.1402905111
Prat neither demonstrates nor illustrates the metopic suture
claimed to be present in KNM-ER 1805 and other fossils. We
also note the absence of references to a metopic suture or open
anterior fontanelle in several original descriptive works, such as
those of Day and colleagues (26), Walker and Leakey (27), or
Schwartz and Tattersall (28), for KNM-ER 3883 or KNM-ER
3733. Although the latter two specimens are described by Leakey
and Walker (29) as retaining traces of a metopic suture, these
features are described as occurring in the glabellar region, which
contradicts the explicit locations (i.e., near bregma) suggested by
Falk and colleagues (2). In contrast to Falk and colleagues (2),
R.L.H. and D.C.B. find no evidence for an open metopic suture,
even ectocranially, in the Sambungmacan 3 Homo erectus cranium,
having studied the original fossil and made endocast ourselves (30).
Given the relevance of location to the zipper closure model en-
dorsed by Falk and colleagues (2), as it was with the Taung Child, it
would be worthwhile to study these (and other) specimens, using
more appropriate means (e.g., high-resolution CT) of determining
the state of metopic suture closure. Such analyses should precede
relying on these specimens to support any adaptive significance of
delayed metopic suture closure in hominin evolution. Ultimately,
even if future in silico techniques are successful in corroborating the
suggested elevated metopic suture frequencies in original Homo
fossils, a connection between metopic suture frequency and the
proposed adaptive benefits of delayed closure must be demonstrated.
For example, whether delayed closure facilitates reorganization
through convolutional changes that are distinctly non-ape must still
be corroborated in many early Homo endocasts (31).
In sum, we believe the claim of high frequencies of metopic
sutures in early hominins (2) is premature, and thus the propo-
sition that delayed metopic suture closure may have conferred
a selective advantage in early hominin evolution is equally pre-
mature. Rigorous analysis of a lynchpin specimen in this argu-
ment (i.e., the Taung Child) provides no support for the notion
that australopiths may have delayed metopic suture fusion, pos-
sibly for adaptive reasons. We suggest that the scenario hypothe-
sized by Falk and colleagues (2) requires substantially more
evidence to support the anatomic, neurologic, and adaptive asser-
tions promulgated therein. Where instances of direct disagreement
exist with original descriptions, the state of metopic sutures sug-
gested by Falk and colleagues should be precisely illustrated to
support the new characterizations. To this end, we suggest that
high-resolution image data sets, such as obtained here from the
Taung specimens, could be extremely helpful, if not necessary, to
substantiate claims of delayed metopic suture closure. Relatively
low-resolution images (e.g., medical CT) likely do not offer
enough spatial resolution to provide definitive evidence, as was
the case with the Taung Child. Also, location discrepancies ob-
served in Taung, and seemingly in other endocasts such as KNM-ER
1813, KNM-ER 3733, and KNM-ER 3883, must be reconciled with
the zipper suture closure pattern suggested by Falk and colleagues
(2) (i.e., a closure process concluding with fusion at bregma, rather
than nasion/glabella). Not all purported instances of partially fused
or open metopic sutures in hominin fossils support this closure
model, but some may instead reflect the underlying mechanisms
manifested in epigenetic or nonmetric traits.
Methods
The Taung (UW-1) partial cranium and natural endocast were separately
scanned by K.J.C. in the Paleosciences Centre Microfocus X-ray CT facility of
the University of the Witwatersrand. The system is a Nikon Metrology XTH
225/320 LC dual-source industrial CT system (see www.wits.ac.za/microCT for
additional information on the system). Each specimen was securely wrapped
in standard bubble wrap to prevent movement during scanning and to
protect it from being damaged by the walls of the container. The container
was an acrylic tube with an external diameter of 110 mm and a wall thick-
ness of 5 mm. It was closed off on the bottom by a horizontal acrylic piece
glued to the bottom of the tube. Once a specimen was secured inside the
Holloway et al.
 acrylic container, the container was attached to the rotating sample holder
platform of the scanner by beeswax.
The partial cranium was scanned using the following parameters: 135 kV;
400 μA; 6,000 projections, where each projection represented the average of
two frames; an acquisition time of two frames per second; and copper (1.2
mm) and aluminum (3.6 mm) filtration. Data were acquired with an isotropic
voxel size of 55.5 μm. Projections were reconstructed into volume data using
proprietary Nikon metrology software (i.e., CT Pro; Nikon Metrology, Inc).
Automated protocols in CT Pro were applied for correcting beam hardening
and reducing noise. Thirty-two-bit float volume data produced from the
scanning were imported into VG Studio Max 2.1 (Volume Graphics GmbH)
and transformed into 16-bit unsigned volume data. While importing volume
data, the gray value histogram was calibrated to background and material
values of the specimen scanned in air.
The natural endocast was scanned using the following parameters: 190 kV,
330 uA, 5,000 projections, an acquisition time of four frames per second, and
copper (1.2 mm) and aluminum (3.6 mm) filtration. Data were acquired with
an isotropic voxel size of 65 μm. The ring artifact correction option in Nikon
proprietary software (i.e., Inspect-X; Nikon Metrology, Inc) was used, which
essentially averages three frames to reduce noise accumulation in the center
of the field of view. Projections were reconstructed into volume data, using
proprietary Nikon metrology software (i.e., CT Pro). Automated protocols in
CT Pro were applied for correcting beam hardening and reducing noise.
Thirty-two-bit float volume data produced from the scanning were imported
into VG Studio Max 2.1 and transformed into 16-bit unsigned volume data.
While importing volume data, the gray value histogram was calibrated to
background and material values of the specimen scanned in air.
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Holloway et al.
After importing volume data, the same procedure was used for pro-
ducing renderings and virtual sections of both specimens in the Virtual
Imaging in Paleontology laboratory of the University of the Witwatersrand.
Using the region growing tool, the object of interest was separated from
other voxels in the volume containing background and noise. Once the
region of interest was identified, a volume rendering of that region was
generated. The volume rendering was positioned in Frankfurt horizontal,
after which coronal sections were obtained by scrolling through the
repositioned volume. Each section was saved as a separate .tif file, along
with an image of the volume rendering indicating the location of the
virtual section.
ACKNOWLEDGMENTS. We are grateful to Dr. Bernhard Zipfel and Brendon
Billings for facilitating access to and assisting with transport of the Taung
specimens and modern human crania from the Dart Collection. We also are
grateful to the Department of Science and Technology of South Africa and
the National Research Foundation of South Africa for multiple funding
awards that permitted the creation of the Palaeosciences Centre Microfocus
X-ray CT facility and the Virtual Imaging in Palaeontology laboratory of the
University of the Witwatersrand. We thank Prof. Alan Morris for providing
information on examples of adult modern humans with metopic sutures in
the Dart collection. We also thank Dr. Tea Jashashvili for assistance with
scanning the modern human cranium (A 3163) and Kimberley Chapelle for
assistance with segmenting Taung images. We are grateful to Dr. Owen
Lovejoy for useful critical comments and, in particular, to Dr. Tim White
for critical comments and useful suggestions regarding future research on
the Taung specimen. We also thank two anonymous reviewers for their
constructive suggestions.
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