THE ANATOMICAL RECORD 294:1548–1556 (2011)

A Bivariate Approach to the Variation

of the Parietal Curvature in the
Genus Homo
EMILIANO BRUNER,1* JOSÉ MANUEL DE LA CUÉTARA,1
2
AND RALPH HOLLOWAY
1
Departamento de Paleobiologı́a, Centro Nacional de Investigación sobre la Evolución
Humana, Burgos, España
2
Department of Anthropology, Columbia University, New York

ABSTRACT
The parietal bones approximately cover the extension of the underly-
ing parietal lobes. Although the boundaries of these two anatomical ele-
ments do not coincide, during morphogenesis the growth of the parietal
bones is largely induced by the pressure exerted by the parietal lobes.
Modern humans display larger parietal chords and arcs compared with
non-modern human species. However, the variation of these variables
have not been analyzed before according to the covariation with the gen-
eral endocranial diameters. When the curvature of the parietal bones is
regressed onto the main neurocranial distances, modern humans show
larger relative values, suggesting not only an absolute enlargement but a
definite allometric change. Taking into account the morphogenetic rela-
tionships with the parietal lobes, these results further support previous
hypotheses suggesting a relative enlargement of these cortical areas in
Homo sapiens, by using simple and reliable homologous neurocranial
arcs. Anat Rec, 294:1548–1556, 2011. V C 2011 Wiley-Liss, Inc.

Key words: parietal bone; parietal lobes; paleoneurology;

human brain evolution

INTRODUCTION the principles of functional craniology (Moss and Young,

Neurocranial morphogenesis is the result of the inte-
gration between soft and hard tissue, most of all
between the bones of the braincase (in particular vault
and cranial base, with some minor but relevant influ-
ence of the facial complex) and brain cortical volumes
(Richtsmeier et al., 2006). Ever since paleontologists
have been aware of an intimate relationship between
brain and cranial evolution (Weidenreich, 1941), taking
into consideration that the same factors shaping the
neurocranium during growth and development may be
involved in generating changes and differences between
species at evolutionary level (Bruner, 2007; Bastir,
2008). Whereas the morphology of the endocranial base
is more sensitive to nonneural variations associated with
structural and biomechanical factors (McCarthy, 2001;
Bruner and Ripani, 2008; Bastir and Rosas, 2009;
Neubauer et al., 2009; Bastir et al., 2010), the morphol-
ogy of the upper endocranial bones is largely the result
of the size and shape changes induced by the underlying
brain cortical components (Enlow, 1990). According to
1960), neurocranial changes in size are supposed to be
mainly induced by the pressure exerted by the underly-
ing brain volumes, whereas, neurocranial changes in
shape are largely influenced by the redistribution of the
growth forces according to the biomechanical properties
of internal tensors. Hence, from one side the pressure of
the growing brain volume separates the vault bones
inducing bone modelling by tissue deposition at the
sutures and on the outer surface and resorption on the
endocranial wall. At the same time spatial changes
through the development are influenced by interhemi-
spheric connective layers of the dura mater (falx cerebri
and tentorium cerebelli) (Moss and Young, 1960), as well
*Correspondence to: Emiliano Bruner, Centro Nacional de
Investigación sobre la Evolución Humana, Paseo de Atapuerca
s/n, 09002 Burgos, España. E-mail: emiliano.bruner@cenieh.es
Received 1 October 2010; Accepted 10 June 2011
DOI 10.1002/ar.21450
Published online 1 August 2011 in Wiley Online Library
(wileyonlinelibrary.com).

V
C 2011 WILEY-LISS, INC.
 PARIETAL BONE AND HUMAN GENUS
Fig. 1. Extension of the parietal bones on the cortical brain surface
(left) and on the endocast (right). Bone landmarks (capital fonts): aste-
rion (ast), bregma (br), lambda (la), and pterion (pt). Approximations of
the main cerebral parietal areas underlying the parietal bones (small
as by the biomechanical effect of the neural fibers (Van
Essen, 1997; Hilgetag and Barbas, 2006). According to
this morphogenetic interaction, volumetric differential
growth and development of specific brain components
are supposed to be associated with evolutionary changes
in endocranial proportions, changing the structural
influence of the cortical surface on the vault form.
While the frontal and occipital bones are affected by
the endocranial base dynamics, the parietal bones are
supposed to be almost entirely shaped by the endocra-
nial brain changes, both in terms of ontogeny and phy-
logeny. The reciprocal endocranial influence between
bones and brain may be not strictly direct, being possi-
bly mediated by mechanical transduction and biochemi-
cal signalling through the vascular components or the
connective tissues of the dura mater (Henderson et al.,
2004). Thus, for the parietal bones we can assume that
the ontogenetic and phylogenetic variation of their mor-
phology are largely influenced by variation of the brain
cortical mass. This is particularly true for the upper pa-
rietal areas, while at the junction with the temporal
squama some biomechanical influences of the cranial
base cannot be ruled out. Also, although frontal and pa-
rietal bones both contribute to the architecture of the
upper braincase and are both formed by direct ossifica-
tion, a certain ontogenetic and evolutionary independ-
ence may be assumed by virtue of their different
embryological origin (neural-crests for the frontal bone,
mesoderm for the parietal bones—Jiang et al., 2002;
Morriss-Key and Wilkie, 2005). A further characteristic
which suggests relatively linear morphogenetic dynamics
of the parietal bone is its origin from one single centre
of ossification, and not from the interaction between
multiple bone components.
The upper part of the parietal bone covers the parietal
convolutions (Fig. 1). As a matter of fact in current bio-
medical practice parietal bones have more relevance in
terms of neurosurgical references (Ribas et al., 2006) than
1549
fonts): ag: angular gyrus; cs: central sulcus; ips: intraparietal sulcus;
ls: lateral sulcus; pos: postcentral gyrus; pre: precentral gyrus; smg:
supramarginal gyrus; spl: superior parietal lobule; stg: superior tempo-
ral gyrus. Digital skull and endocast on the right are from Mladeč 1.
as possible source of morphogenetic and pathological
defects (Gupta et al., 2008). At the same time, such a tight
relationship between parietal brain areas and parietal
bones make these latter a useful proxy to study evolution-
ary cortical changes from a paleoneurological perspective.
In the current paleoanthropological debate a clear-cut
separation between modern and nonmodern variation is
still disputed, also taking into account the many chrono-
logical and geographical gaps in the fossil record
(Stringer, 2002). Nonetheless, there is a certain agree-
ment in recognising a brachycephalic morphology and a
globular braincase as characters definitely associated
with anatomical modernity (Lieberman et al., 2002;
Pearson, 2008). The parietal areas are largely related to
such changes in the neurocranial proportions, being the
structural bridge between the other bone components.
The possible role of the parietal areas in hominid evo-
lution has been evidenced since the early studies in pale-
oneurology (Dart, 1925), revealing peculiar patterns of
variation throughout the entire hominoid clade (Hollo-
way, 1981a). Modern humans have been hypothesized to
be characterized by enlargement of the parietal cortical
areas, according to results obtained from the analysis of
the ectocranial profile (Bruner et al., 2004), by using
endocranial brain metrics (Bruner et al., 2003; Bruner,
2004), and considering both shape and form differences
(Bruner, 2008, 2010a; Gunz et al., 2010). When com-
pared with more archaic species of the genus Homo,
Neandertals display a lateral widening of the lower and
upper parietal districts, whereas, modern humans show
a general enlargement of the whole parietal area. Such
changes are not allometric consequences of general brain
size increase, but probably species-specific changes of
the neurocranial morphogenetic patterns. Recently,
areas around the intraparietal sulcus were hypothesized
to be involved in the differences characterizing the mod-
ern human brain geometry (Bruner, 2010b; Bruner
et al., 2010).
1550 BRUNER ET AL.

Fig. 2. Endocranial diameters investigated in the present paper: BAC: bregma–asterion chord; BLA:
bregma–lambda arc; BLC: bregma–lambda chord; HL: hemispheric length; MWA: maximum width arc
MWC: maximum width chord (as: endoasterion; br: endobregma; cs: trace of the coronal suture; eu:
endoeurion; lm: endolambda; ls: trace of the lambdoid suture).

Nonetheless, endocranial morphometric analyses using
anatomical or geometric cortical references have been of-
ten questioned because of the difficulties in localizing
brain landmarks in fossil specimens. The recognition of
cortical components on the endocast surface is often
based upon the personal experience of the researcher.
Also, many cortical references are formed by smooth
surfaces of continuous profiles, requiring a priori meth-
odological choices when they are represented by specific
diameters or coordinates. On the other hand, parietal
bone landmarks are commonly referred as to homologous
and reliable metric references, throughout the history of
anthropometrics. Although it is recognized that modern
humans have larger parietal bones, the relationship
between their dimensions and the other neurocranial
diameters have not been quantified in the human genus.
The present analysis is aimed at considering the length,
width, and curvature of the parietal bones taking into
account the covariation with the major neurocranial and
endocranial diameters, to analyze the allometric pat-
terns characterizing the variability of the genus Homo.
Apart from supplying a quantitative framework of varia-
tion for the parietal bone by using a large sample size,
this analysis provides a further test for the endocranial
parietal differences already described between modern
and nonmodern human taxa, by using cranial homolo-
gous variables instead of endocranial biological referen-
ces (sulci and gyri), whose localization may be more
subjective or less reliable. The null hypothesis states
that modern and nonmodern human species rely on the
same allometric trajectory for the major parietal bone
diameters, the differences between species being the
direct consequence of the allometric variation.
MATERIAL AND METHODS
The traces of the cranial suture are generally visible
on the endocasts in fossil and extant hominids, and bony
metrics can be directly measured on the endocranial
mould. Sutures are not perfectly perpendicular to the
bone surface, producing minor discrepancies between
outer and inner points, but there is a good correspon-
dence between ectocranial and endocranial bone land-
marks (Holloway et al., 2004). Hence, measuring bone
landmarks on the endocranial surface allows us to quan-
tify and to compare directly endocasts morphology
through reliable and homologous cranial references. In
this analysis, we considered the endobregma–endo-
lambda chord and endobregma–endolambda arc (BLC,
BLA), the endobregma–endoasterion chord and endo-
bregma–endoasterion arc (BAC, BAA), the hemispheric
length (as the interlandmark distance between the fron-
tal and occipital poles, averaging the two hemispheres;
HL), and the maximum endocranial width chord and
maximum endocranial arc (MWC, MWA) (Fig. 2; see Hol-
loway et al., 2004). Generally, endobregma lies anterior
to the central scissure, between the posterior areas of
the prefrontal convolutions and the precentral sulcus.
Endolambda is generally posterior (lower) to the perpen-
dicular scissure, covering the upper part of the occipital
lobe. Endoasterion is localized around the meeting area
between the occipital, temporal, and cerebellar lobes.
Accordingly, the extension of the parietal bone includes
all the parietal lobes, the posterior part of the frontal
lobes, and the anterior portion of the occipital lobes (Fig.
1) (Ribas et al., 2006).
The endocranial variation of the parietal bones has
been quantified in a sample of 79 adult modern human
specimens from RLH’s collection. Nineteen individuals
belonging to the H. ergaster/erectus hypodigm and 11
Neandertals have been used to provide estimations of
the allometric variation in nonmodern human species
(Table 1). Four individuals from the Afro-European Mid-
dle Pleistocene (H. heidelbergensis) have been also
included in the sample but not used in the regressions.
The variation within adult modern humans has been
quantified using univariate and bivariate statistics.
Relationships between variables has been investigated
by using a model II regression through reduced major
axis on the raw data and on the log–log transformed
variables. Differences among modern and nonmodern
taxa have been quantified by using the least-square
approach and analysis of covariance. Principal compo-
nent analysis (PCA) was computed on the whole set of
variables to describe the overall patterns of relationships
involved in generating the morphological variation of
 PARIETAL BONE AND HUMAN GENUS 1551
the parietal form. PCA was computed by using the corre- RESULTS
lation matrix on modern humans alone and on the whole
Table 2 gives the descriptive statistics for the varia-
sample. Only specimens with all variables were used, to
bles investigated, to provide a metric reference for the
avoid missing data imputation. Statistics have been com-
modern human species. The variation is lower for the
puted with PAST 2.08b (Hammer et al., 2001).
bregma–asterion diameters (4%), and higher for the
maximum width arc (8%). Table 3 provides the correla-
tion and determination coefficients for the pairwise
TABLE 1. Fossil sample regressions, the slopes according to the reduced major
Archaic humans (N ¼ 19) axis, and the allometric coefficients with 95% confidence
KNM-ER3733 intervals for the log–log approach. The highest correla-
KNM-ER3883 tion is shown between the midsagittal chord and arc
OH9 (90%). For this pair of variables isometry cannot be
Salé rejected, and the arc is about 11% larger than the chord.
Sambungmacan 3 On the other hand, the midsagittal parietal arc shows
Sangiran 2
Sangiran 4
positive allometry relatively to the hemispheric length
Sangiran 10 and the maximum width arc shows positive allometry
Sangiran 12 relatively to the endocranial width. Both regressions
Sangiran 17 show an allometric coefficient of 1.27. Hence, as the
Solo XI brain gets longer or wider, the parietal curvature
Solo V enlarges, increasing the bulging of the parietal surface.
Solo VI The relationship between bregma–asterion chord and
Trinil 2 arc is quite high (67%), and the allometric coefficient
WT15000 suggests an isometric pattern of variation. Table 4 shows
Zhoukoudien 1L
Zhoukoudien 2D
the same parameters for nonmodern humans.
Zhoukoudien 3E The comparison between modern and nonmodern allo-
Zhoukoudien 3L metric patterns indicates relatively larger parietal arcs
Afro-European Middle Pleistocene in H. sapiens, even when the endocranial diameters are
Arago taken into account (Fig. 3). Modern humans have a
Kabwe slightly larger bregma–lambda arc relative to the
Reilingen bregma–lambda chord (Fig. 3a). The ratio between BLA
Sima de los Huesos 5 and BLC is proportional to the degree of sagittal curva-
Neandertals (N ¼ 11) ture of the parietal bones. Hence, at similar midsagittal
Amud
La Chapelle aux Saints
parietal length modern humans have a more curved pa-
La Ferrassie rietal profile. This difference is definitely more apparent
Feldhofer when plotting the bregma–lambda arc onto the hemi-
Gibraltar 1 spheric length, that is comparing the parietal total
Guattari 1 length with the main (antero-posterior) brain diameter
Krapina 3 (Fig. 3b). A similar result is obtained by considering the
Krapina 6 correlation between the maximum width arc and the
La Quina maximum endocranial width, which ratio is proportional
Spy 1 to the degree of coronal curvature (Fig. 3c). In this sec-
Spy 2
ond case there is some overlapping in the distribution of

TABLE 2. Descriptive statistics for modern humans (see text for labels)
N Mean sd CV (%) Min 25q M 75q Max
HL 78 166 9 5.3 144 162 166 171 187
BLC 75 105 7 6.6 89 100 106 110 122
BLA 75 117 8 6.9 97 111 117 122 132
MWC 78 130 8 6.5 113 124 131 136 150
MWA 78 236 19 7.9 180 225 241 249 265
BAC 69 125 5 4.0 113 121 126 129 135
BAA 69 156 7 4.5 138 152 157 161 168

TABLE 3. Correlations and regressions for modern humans (see text for labels)
Log–Log
R R 2
Slope
95%CI Slope þ95%CI
HL BLA 0.62 0.38 0.89 1.06 1.27 1.57
BLC BLA 0.95 0.90 1.14 0.97 1.04 1.12
MWC MWA 0.59 0.35 2.24 0.98 1.27 1.54
BAC BAA 0.82 0.67 1.32 0.92 1.06 1.19
All correlations: P < 0.0001.
1552 BRUNER ET AL.

TABLE 4. Correlations and regressions for non-modern humans (see text for labels)
Log–Log
R R 2
Slope
95%CI Slope þ95%CI
HL BLA 0.84 0.70 1.02 1.39 1.75 2.12
BLC BLA 0.99 0.97 1.08 0.96 1.02 1.10
MWC MWA 0.80 0.64 2.61 1.12 1.56 1.90
BAC BAA 0.97 0.95 1.41 1.06 1.16 1.26
All correlations: P < 0.0001.

Fig. 3. Bivariate regressions (least-square fits). Groups: ARC, Archaic humans; MPL, Afro-European
Middle Pleistocene; NDR, Neandertals; MOD, Modern humans. Variables: BAA, endobregma–endoaste-
rion chord; BAC, endobregma–endoasterion arc; BLA, endobregma–endolambda arc; BLC, endobregma–
endolambda chord; HL, hemispheric length (average); MWA: maximum endocranial width arc; MWC,
maximum endocranial width chord.

the specimens, but the difference in the regression fits is
clear. No differences in these allometric trajectories can
be seen between H. ergaster/erectus and Neandertals.
Considering the relationship between bregma–asterion
chord and arc the differences between modern human
and Neandertals are less marked, but visible neverthe-
less (Fig. 3d). It is worth noting that in this case there is
a definite gap between archaic humans and Neandertals.
The small sample size for both groups hampers robust
statistical approaches, but the latter display relative
larger values for the arc. The ratio between these two
diameters is proportional to the degree of diagonal cur-
vature of the parietal bone, including both the antero-
posterior parasagittal component and the latero-lateral
component. This may be relevant considering that this
arc crosses both the upper and lower parietal lobules,
including the supramarginal area. In all comparisons
the African and European Middle Pleistocene specimens
always fit within the nonmodern trajectories, but in this
last comparison they are included in the Neandertal
lower range.
Analysis of covariance of these allometric trajectories
confirms the graphic results: homogeneity of slopes can-
not be rejected for any bivariate analysis (P > 0.05), and
modern humans show larger values for the parietal arcs
(both sagittal and coronal curvatures) when corrected
for the endocranial main diameters (P < 0.0001). Also
for the regression between bregma–asterion chord and
arc, the differences between modern humans and the
nonmodern taxa are significant, including the difference
with Neandertals (P ¼ 0.05). Hence, the null hypothesis
of no differences in the allometric relationships between
modern and nonmodern humans can be rejected for the
endocranial parietal metrics analyzed in the present
study: among the human taxa the within-species allo-
metric trends (coefficients) are similar, but in modern
humans the relative parietal dimensions exceed the fig-
ures expected for nonmodern groups, most notably for
the sagittal and coronal arcs.
PCA on the modern human sample and on the whole
sample are shown in Fig. 4a and b, respectively. Accord-
ing to a Jolliffe cut-off criterion, in both cases only the
 PARIETAL BONE AND HUMAN GENUS
Fig. 4. Biplots of the PCA for the modern humans (a) and for the
whole sample (b). In both cases only the first two principal compo-
nents are significant. Group variation is shown by 95% confidence
ellipse for modern humans, and by convex hulls for the inter-specific
comparison. The arrows show the variable vectors projected onto the
multivariate subspace (see text for labels).
first two principal components are significant. In the
analysis of the modern variation, the first component
(62% of variance) is a size vector. The second component
(19%) is associated with decrease of the endocranial
lengths (hemispheric and parietal) and increase of the
other variables (mostly the maximum width arc). Also in
the analysis of the whole sample the first component
(72%) is a size vector, separating at the two extremes H.
ergaster/erectus and Neandertals. Modern humans show
intermediate values, largely overlapping with the Nean-
dertal variation. The second component (12%) is associ-
ated with decrease of the parietal lengths and increase
in the maximum width chord. This second component
further separates modern humans and Neandertals,
with the latter group at higher values. It is worth noting
that in this subspace, accounting for 84% of the var-
iance, H. ergaster/erectus is completely separated from
the other groups, whereas, Neandertals and modern
humans show a certain degree of overlapping. Afro-Eu-
ropean Middle Pleistocene specimens approach the lower
range of Neandertal variation, totally included within
the modern human variability. However, in this case the
sample size is too small to allow any robust
interpretation.
DISCUSSION
Changes at the parietal lobes have been often
hypothesized to be relevant in hominid brain evolution.
Very early in the history of paleoneurology, Franz Wei-
denreich (1936) suggested that ‘‘it is the parietal and not
the frontal regions which undergo the greatest degree of
1553
differential enlargement’’, remarking that this ‘‘appears
to be conclusions from which there is no escape.’’ The pa-
rietal cortical surface shows a large variation within
humans and within hominids (Holloway, 1981a), and
cortical reorganization at the parietal lobes have been
hypothesized to be associated with the evolution of the
earliest Pliocene hominids (Dart, 1925; Holloway, 1995).
A lateral enlargement of the upper parietal lobules has
been described for Neandertals (Bruner et al., 2003),
leading to the classic en bombe profile of the vault in
posterior view. However, at least in terms of gross mor-
phology of the endocranium, a general enlargement of
the whole parietal cortical volumes is the most outstand-
ing trait associated with modern human endocranial
morphology (Bruner et al., 2003; Bruner, 2004; Gunz
et al., 2010). Neurocranial globularization is regarded as
a derived modern human character (Lieberman et al.,
2002), and bulging of the fronto-parietal cranial profile
separates modern humans from the extinct human taxa
through a discrete morphological gap (Bruner et al.,
2004). Such bulging of the parietal lobes separates H.
sapiens from the allometric trajectory characterizing
brain shape changes in nonmodern humans associated
with the encephalization processes (Bruner et al., 2003;
Bruner, 2008). Structurally, these variations may be
interpreted taking into account allometric constraints
associated with the connective tensors of the brain (Bru-
ner, 2004). Functionally, changes at the intraparietal
area may be responsible for such differences, with cogni-
tive implications ranging from self-perception (as well as
perception of others and objects) to simulation and
abstract representation (Bruner, 2010b).
Most of these results have been achieved analyzing
the geometry of the endocasts by using anatomical refer-
ences associated with the cortical brain morphology.
Such references are very useful in terms of homologous
and functional brain components, but need caution
because of the difficulties in inferring brain anatomy
from endocasts morphology. Most of the traits do not cor-
respond to specific landmarks but to ‘‘endocranial areas’’
(bosses or depressions on the endocast surface), needing
experience and always entailing a certain degree of sub-
jectivity. Resampling and averaging (specimens or taxa)
are generally used to minimize possible uncertainties,
and the reliability of the results may be evaluated con-
sidering the stability of the statistical outputs.
The present analysis is aimed at supplying further
evidence of the parietal variations in modern humans
through homologous type I cranial landmarks (for those
metrics involving bregma, lambda, and asterion) and
type II cranial reference largely used in anthropometric
literature (for those metrics using euryon) instead of
type II and type III brain cortical references like those
landmarks used to localize brain bosses and sulci, and
assuming the parietal bone curvature as proxy for the
parietal lobe curvature. In terms of anatomy, the parie-
tal bones covers entirely the parietal lobes, plus a small
portion of the frontal lobes anteriorly, of the occipital
lobes posteriorly, and of the temporal lobes inferiorly.
Such correspondence is intimately associated with the
spatial organization between vault bones and brain cort-
ical organization, and as a matter of fact the anatomical
references used in this study have importance in neuro-
surgical context (Ribas et al., 2006). In terms of geome-
try, the vault morphology is largely shaped by the
1554 BRUNER ET AL.
pressure exerted by the underlying cortical masses
(Moss and Young, 1960). Hence, the curvature of the pa-
rietal squama may be largely interpreted as the result of
the morphogenetic intrinsic or extrinsic influence of the
underlying parietal lobes. This is quite relevant also
when considering the parietal bone within the general
framework of the cranial architecture. On the other
hand, the parietal bone morphology is highly integrated
with the occipital and temporal bone morphology in
human ontogeny and evolution (Bookstein et al., 2003;
Gunz and Harvati, 2007). At the same time, the form of
the cranial vault has an important genetic and phyloge-
netic signal, being less influenced by external factors
such as climate when compared with other anatomical
districts (Harvati and Weaver, 2006). In this article, we
quantify the degree of variation and allometric patterns
of the main parietal bone diameters and arcs in modern
and fossil humans, testing to what extent the endocra-
nial size may influence the parietal bone proportions
and curvature.
The current analysis showed that in adult modern
humans the variation of the latero-lateral curvature of
the parietal bones (8%) is only slightly larger than the
variation of the sagittal curvature (7%). Although the
length and curvature of the parietal bones showed simi-
lar proportions at different size, the parietal bulging
(both latero-lateral and midsagittal) increases as the
brain gets larger. Hence, in adult modern humans there
is a correlation between brain size and degree of parietal
bulging, with larger endocasts showing more curved pa-
rietal squamas. Interestingly, the allometric coefficient is
similar in both sagittal and lateral directions (1.27).
Apart from a general size vector, within modern
humans and within the human genus a second pattern
of variation contrasts the parietal length and the parie-
tal width. However, within modern humans the lateral
parietal curvature (the parietal arc) has a major role in
generating the sample variation. In contrast, in the total
sample more weight is gained by the parietal width
(maximum width chord), because of the Neandertal mor-
phology polarizing this component. This suggests that it
is the overall width more than the curvature character-
izing the Neandertal vault form.
The allometric patterns are similar in all human spe-
cies, showing the three humans groups investigated (ar-
chaic humans, Neandertals, and modern humans) have
comparable slopes. However, while differences between
Neandertals and archaic humans seem mostly related to
size variation, H. sapiens shows larger relative values
for the parietal curvature. This is definitely apparent
when considering the curvature along the midsagittal
and coronal planes, which are strictly related to the ge-
ometry of the underlying parietal lobes. The same pat-
tern can be confirmed also for the bregma–asterion
metrics, even if in this case the difference between mod-
ern humans and Neandertals is less noticeable. Asterion
and the lower parts of the parietal bone cover the poste-
rior tract of the temporal lobes and the anterior occipital
area, whereas, the upper tract of the bregma–asterion
lengths covers the posterior areas of the frontal lobes, as
well as the supramarginal gyrus (Ribas et al., 2006). For
this chord Neandertals seem to have larger values than
less encephalized human species, even if figures are rel-
atively smaller than modern humans. Hence, this may
be related to discrete changes of this taxon associated
with a lateral enlargement of the supramarginal boss, as
described for many Neandertal specimens (Holloway,
1981b). This difference may be smaller in the metrics
involving maximum endocranial width, because this di-
ameter relies on geometrical landmarks which are not
homologous and are generally localized in a lower posi-
tion in archaic human species (Holloway, 1980). How-
ever, these lower cranial areas are in structural
interaction with the cranial base, whose multifactorial
structural organization (Lieberman et al., 2000; Bruner
and Ripani, 2008; Bastir et al., 2010) hampers a straight
evolutionary and morphogenetic relationship between
neural and cranial components. At last, the temporo-pa-
rietal boundaries are characterized by the squamosal
suture, which is highly variable in hominids, partially
influenced by cranial shape factors, and associated with
discontinuous changes through human phylogeny (Ter-
hune and Deane, 2008). Hence, robust paleoneurological
inferences on the relationship between hard and soft tis-
sues in this area are less easy to provide. The involve-
ments of the occipital, temporal, and frontal lobes (for
which no marked differences in their posterior morphol-
ogy are known between modern humans and Neander-
tals) and the implications of the cranial base
multifactorial dynamics decrease the differences in the
bregma–asterion curvature, which nonetheless can still
be detected.
These results demonstrate that at the same brain size
modern humans have larger parietal arc than nonmo-
dern species. This is confirmed relatively to the parietal
chord, but it is even more apparent when considering
the hemispheric length or endocranial width. As in pre-
vious studies based onto anatomical brain references,
also in the present analysis the good fit of the regres-
sions and the good correlation levels give strength to the
reliability of the results. These data once more support
the differences between modern and nonmodern human
species concerning the parietal bone/brain morphology,
suggesting a definite role of this trait in characterizing
the neurocranial model of our own species. While allo-
metric differences among the fossil groups cannot be dis-
carded, the available sample size may not be sufficient
to provide a robust statistical power to support this con-
clusion. Nevertheless, such differences would not be as
marked as the differences between modern and nonmo-
dern humans, and we can state that the extinct human
species show at least very similar scaling relationships
and proportions.
The outer and inner tables of the parietal bone show
different patterns of distribution of physical parameters
like mineral density, stiffness orientation, and bone
thickness, and many details of the functional and struc-
tural organization of this vault component are still
scarcely understood (Peterson and Dechow, 2002). It is
worth noting that the parietal lobes show a certain his-
tological continuity with the occipital lobes (Eidelberg
and Galaburda, 1984), and the parietal bones show a
morphological integration with the occipital squama
(Gunz and Harvati, 2007). Such a structural and func-
tional relationship has often hampered an independent
consideration of the parietal volumes alone in compara-
tive studies on the brain components, although eviden-
ces of H. sapiens specific neuroanatomical features are
now emerging (Vanduffel et al., 2002; Orban et al.,
2006), as well as important parietal changes during
 PARIETAL BONE AND HUMAN GENUS
endocranial morphogenesis (Neubauer et al., 2009, 2010;
Gunz et al., 2010) and relevant neurofunctional factors
mostly related to the intra-parietal area (Bruner, 2010b).
Interestingly, areas in the depth of the parietal lobes
may be also associated with mental speed (Bruner et al.,
2011).
Unfortunately comparative neuroanatomical data on
living apes are scarce. Semendeferi and Damasio (2000)
demonstrated no relative differences between humans
and apes for the parieto-occipital volumes, but data on
the parietal areas alone are not available. Moreover, a
reduction of the visual (occipital) components in modern
humans has been recently suggested (De Sousa et al.,
2010), further supporting the hypothesis of a relative
expansion of the parietal districts. Although comparative
data on hominoids are definitely necessary, we must
however take into consideration that living apes are ter-
minal lines of evolution, and not species ancestral to the
human lineage. The assumption that living apes display
the ancestral human character states is hence not neces-
sarily true. In this sense, the paleontological evidence
can be useful when considering the polarity of the evolu-
tionary changes.
Considering the fossil record associated with the mod-
ern human lineage, it is worth noting that there are
specimens without bulging parietal bones like Jebel
Ihroud 1 (Hublin, 2002) and specimens presenting a
modern parietal morphology like Herto (White et al.,
2003). Taking into account the phenetic similarity
between these fossils, the comparable chronology, the
different geography (North and East Africa, respec-
tively), the differences in parietal morphology, and the
endocranial evolutionary process associated with modern
humans, this may suggest that the origin of the modern
human lineage may have not coincided with the evolu-
tion of the modern human brain.
ACKNOWLEDGEMENTS
The authors are grateful to Tom Schoenemann and
Markus Bastir for their comments and suggestions. This
article is supported by the Program GR. 249, Junta de
Castilla y León (Spain), by the Project CGL2009-12703-
C03-01 Ministero de Ciencia e Innovación (Spain), and
by the Italian Institute of Anthropology.
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