Original Paper
Brain Behav Evol 2018;91:136–147
DOI: 10.1159/000488889
Human Paleoneurology and the
Evolution of the Parietal Cortex
Emiliano Bruner
Centro Nacional de Investigación sobre la Evolución Humana, Burgos, Spain
Keywords
Brain evolution · Hominids · Parietal lobes · Precuneus ·
Visuospatial integration
Abstract
Paleoneurology deals with the study of brain anatomy in fos-
sil species, as inferred from the morphology of their endocra-
nial features. When compared with other living and extinct
hominids, Homo sapiens is characterized by larger parietal
bones and, according to the paleoneurological evidence,
also by larger parietal lobes. The dorsal elements of the pos-
terior parietal cortex (superior parietal lobules, precuneus,
and intraparietal sulcus) may be involved in these morpho-
logical changes. This parietal expansion was also associated
with an increase in the corresponding vascular networks,
and possibly with increased heat loads. Only H. sapiens has a
specific early ontogenetic stage in which brain form achieves
such globular appearance. In adult modern humans, the pre-
cuneus displays remarkable variation, being largely respon-
sible for the longitudinal parietal size. The precuneus is also
much more expanded in modern humans than in chimpan-
zees. Parietal expansion is not influenced by brain size in fos-
sil hominids or living primates. Therefore, our larger parietal
© 2018 S. Karger AG, Basel
E-Mail karger@karger.com
www.karger.com/bbe
Published online: August 10, 2018
cortex must be interpreted as a derived feature. Spatial mod-
els suggest that the dorsal and anterior areas of the precu-
neus might be involved in these derived morphological vari-
ations. These areas are crucial for visuospatial integration,
and are sensitive to both genetic and environmental influ-
ences. This article reviews almost 20 years of my collabora-
tions on human parietal lobe evolution, integrating func-
tional craniology, paleoneurology, and evolutionary neuro-
anatomy. © 2018 S. Karger AG, Basel
Paleoneurology and Functional Craniology
Morphology is not only a study of material things
and of the forms of material things, but has its dynamical
aspect, under which we deal with the interpretation,
in terms of force, of the operations of energy.
D’Arcy Wentworth Thompson
(On Growth and Form, 1942)
Traditionally, paleoneurology has been defined as the
study of fossil endocasts, namely the casts of the cranial
cavity in extinct species [Falk, 1987; Holloway et al.,
2004]. The tight contact between brain and braincase, and
Emiliano Bruner
Centro Nacional de Investigación sobre la Evolución Humana
Paseo Sierra de Atapuerca 3
ES–09002 Burgos (Spain)
E-Mail emiliano.bruner @ cenieh.es
the fact that the vault bones are largely shaped by the
brain during growth, makes it possible to provide some
inferences from skull (endocranial) morphology on brain
anatomy [Bruner, 2017a]. This is particularly relevant
when dealing with species for which we only have bone
remains. Brain tissues do not fossilize, but the endocra-
nial cavity can supply information on brain size and pro-
portions, as well as on sulcal patterns, and in the past cen-
tury these have been the most investigated paleoneuro-
logical features. On the endocranial wall, there are also
the imprints of some vascular elements, such as the mid-
dle meningeal artery and the venous sinuses. The endo-
cranial cavity provides this information by itself, but
nonetheless paleoneurology (which should be more
properly termed paleoneurobiology) largely deals with its
positive mold, the endocranial cast. After all, we are pri-
mates, and we are used to relying on our eyes and hands
to interact with our environment. That is why we prefer
to have “something to see and to handle,” instead of an
empty cavity. Of course, much information is lost from a
real brain to its endocranial cast, which is a sort of geo-
metrical model able to supply only some scattered infor-
mation on cerebral gross cortical features and general
proportions. Furthermore, the meninges and the cere-
brospinal fluid are interposed between the cerebral cortex
and the internal table of the braincase, separating their
respective surfaces and smoothing the endocranial im-
prints, most of all in larger skulls [Kobayashi et al., 2014;
Van Minh and Hamada, 2017]. Yet, despite these limita-
tions, this is the only direct information we can have on
the brains of extinct taxa and, therefore, deserves atten-
tion.
However, not every endocranial variation is due to
brain anatomy. Even the word “brain” is a bit misleading
and very general when dealing with morphology. Its size
and shape depends on neurons but also upon glial cells,
cerebrospinal fluid, and blood vessels. Furthermore, its
form and geometry are not a real property of the organ
itself, but are largely due to internal blood pressure and
external meningeal tension [Moss and Young, 1960]. Ad-
ditionally, the intimate spatial relationship between skull
and brain makes influences between bone and cortex re-
ciprocal, and the final phenotype is hence shaped by the
effects of both soft and hard tissues. In terms of both on-
togeny an phylogeny, skull and brain must co-evolve, fol-
lowing a principle of structural and functional balance
[Richtsmeier et al., 2006; Bruner, 2015]. In the dorsal re-
gions (vault), the brain largely molds the bones because
its growth processes are compensated by bone deposition
and modeling. However, in the ventral regions (base), the
Evolution of the Parietal Lobes
skull exerts important constraints on the brain form at
the three endocranial fossae. The frontal lobes are con-
strained by the orbital structures, the temporal lobes are
constrained by the facial block and by mandibular ele-
ments, and the cerebellum is constrained by the cranial
base flexion. Therefore, when handling endocasts, paleo-
neurology must carefully consider not only brain anato-
my but also the general spatial and morphogenetic rela-
tionships between skull and brain, so as to determine
what changes of the brain form are due to brain evolution,
and what are secondary outcomes due to cranial influ-
ences. Both kinds of variations are relevant in evolution-
ary anthropology, but only the former might possibly be
associated with neural (and ultimately cognitive) varia-
tions.
Parietal Lobes and the Study of Brain Evolution
Parietal lobes have interested paleoanthropologists for
a long time. In the early days of the discipline, almost one
century ago, Raymond Dart and Franz Weidenreich al-
ready mentioned morphological differences of the pari-
etal region when studying australopithecines and Homo
erectus, respectively. Ralph Holloway [1981] was proba-
bly the first to perform an endocranial shape analysis on
living and fossil hominoids, evidencing a remarkable pa-
rietal variability among different species. In humans, dis-
tinctive parietal traits have been known since the first half
of the 20th century also in neurobiology [Catani et al.,
2017], and advances in the study of the parietal cortex
have been stimulating and promising [Mountcastle,
1995]. Nonetheless, the posterior parietal cortex was, un-
til recently, an “uncharted region” [Zilles and Palomero-
Gallagher, 2001], and parietal lobes have never been tak-
en into consideration when dealing with key changes in
human evolution.
There could be at least 5 main reasons for this. First, a
large part of the parietal cortex is not superficial but po-
sitioned under the cerebral surface (precuneus and intra-
parietal sulcus). These folds are more difficult to discern
in terms of dissection and visualization, and they can pass
unnoticed in many observations and surveys. Second,
this deep position makes damage to these areas less com-
patible with survival after impairment. Functional dam-
age – important in detecting an association between
structure and function in the past centuries – is less easy
to observe in these deep cortical areas when compared
with lesions in other regions (e.g., language or personal-
ity alterations after frontal injury). Third, the parietal cor-
Brain Behav Evol 2018;91:136–147 137
DOI: 10.1159/000488889
tex is generally labeled with the elusive term “association
cortex,” which denotes multiple and integrative func-
tions. The parallel involvement of many distinct func-
tions makes experiments and inferences less straightfor-
ward. Fourth, many parts of the parietal lobes are associ-
ated with visuospatial integration which, in general, refers
to a set of functions traditionally interpreted as basic spa-
tial management, something that was not expected to be
involved in “higher” cognitive functions. Fifth, humans
and nonhuman primates display many anatomical differ-
ences which often hamper a direct comparative approach.
Patent inconsistencies between the monkey and human
parietal cortical maps were described since Brodmann’s
studies back in 1909 [Zilles and Palomero-Gallagher,
2001; Zilles and Amunts, 2010]. This lack of correspon-
dence, instead of stimulating the study of these areas, may
have paradoxically demoted interest in these cortical ele-
ments.
At the end of the 1990s, I began applying shape analy-
sis and multivariate statistics to paleoanthropology and
craniology, and it soon became apparent that parietal
bone shape was characteristic in our own species. Parietal
bone shape differences could have been only a matter of
cranial architecture, but then I found similar results when
working with endocasts, brains, or vessels. I was surprised
to find an unexpected paucity of information on many
cranial, cerebral, and vascular traits associated with pari-
etal regions, so I began to investigate not only fossils but
also living humans, not only hominoids but also other
primates, not only evolutionary aspects but also medical
ones, not only the isolated features but also their relation-
ships. This article is hence a review of almost 20 years of
my personal collaborations on the evolution of parietal
anatomy, a long trail that did not aim to look for answers
but instead search for proper questions.
The Fossil Evidence: Parietal Bones, Lobes,
and Vessels
The modern human skull has always been defined as
“globular” when compared with extinct human species,
which instead display a flatter braincase [Lieberman et al.,
2002]. The frontal bone can partially contribute to this
globular shape because of a more pronounced curvature
of the frontal squama. However, there is no evolutionary
evidence of macroscopic morphological changes in the
modern human frontal lobes or in the frontal endocra-
nial profile [Bookstein et al., 1999; Semendeferi et al.,
2002; Bruner, 2017b], and therefore such frontal curva-
138 Brain Behav Evol 2018;91:136–147
DOI: 10.1159/000488889
ture is probably simply a spatial rearrangement due to the
peculiar position of our face situated under the anterior
cranial fossa [Beaudet and Bruner, 2017; Pereira-Pedro et
al., 2017a]. Instead, shape analyses revealed that our glob-
ular vault is largely due to the size and proportions of the
parietal bones [Bruner et al., 2004]. In fact, Homo sapiens
has a parietal bone which is larger and more bulging than
in any other human species [Bruner et al., 2011a]. This
feature is not due to the large size of the braincase, and in
fact the Neanderthals, with a similar cranial capacity, dis-
played short and flat parietal bones [Bruner, 2014]. The
longitudinal size and curvature of the parietal bones do
apparently not influence the position of other endocra-
nial regions, but have an effect on the general orientation
of the skull, thus influencing the general head position
and its functional axis [Bruner et al., 2017a].
Moving from skull to brain, the boundaries of the
parietal lobes can be tentatively recognized on endocasts
[Pereira-Pedro and Bruner, 2018]. However, while mor-
phometrics of the bone are based on fixed and recogniz-
able craniometric landmarks (sutures), morphometrics
of the lobe must be necessarily performed on anatomical
inferences. Sulcal imprints are faint and smooth, and
their localization is based on the experience of the anato-
mists. Nonetheless, the localization of cortical references
on endocasts relies not only on the recognition of the spe-
cific grooves or bosses on the endocranial surface, but
also on the relative position of other surrounding cortical
elements. The use of sample or hemispheric averages can
further limit this anatomical uncertainty, providing a
rough but reasonable estimation of the position of the
central sulcus, postcentral sulcus, parieto-occipital sul-
cus, intraparietal sulcus, supramarginal gyrus, and angu-
lar gyrus. According to such estimations, modern hu-
mans are not only characterized by larger parietal bones
but also by larger parietal lobes (Fig.  1) [Bruner et al.,
2003, 2018a; Bruner, 2004]. Spatially, this difference is ap-
parently associated with the dorsal parts of the parietal
cortex, possibly the superior parietal lobules or the intra-
parietal sulcus [Bruner, 2010]. A similar situation can be
described when comparing the endocasts of living homi-
noids, with modern humans displaying a remarkable ex-
pansion of the superior parietal lobule (Fig. 2).
An increase in the size and proportions of the parietal
bone can be explained as an increase in the underlying
brain elements (larger parietal cortex) or, alternatively, as
a spatial readjustment of the cranial architecture (i.e.,
flexion and curvature of the skull) with no major changes
in brain proportions. However, in the case of an increase
in the parietal lobe, the latter explication does not stand.
Bruner
 a b
d e
Fig. 1. When compared with all the other fossil hominids, Homo
sapiens’ rounded skull (a), in terms of midsagittal geometry (b), is
largely due to absolute and relative enlargement of the parietal
bone (c – digital replica of skull and endocast of an australopith,
Sts5, and shape deformation associated with modern human cra-
nial form). Some anatomical references of the parietal lobe (cs,
central sulcus; pos, parieto-occipital sulcus; ips, intraparietal sul-
cus; smg, supramarginal gyrus; ag, angular gyrus) can be tenta-
tively inferred on endocasts (d). When compared with more ar-
chaic human species like Homo erectus (e), Neanderthals display a
lateral bulging of the dorsal parietal surface (LD: lateral dorsal ar-
A volumetric change of a specific cortical region cannot
be interpreted as a geometric inflation due to spatial cur-
vature and is more likely to be associated with the increase
in some cerebral components (neurons, connections,
glia, etc.) included within those boundaries.
Additional evidence comes from the vascular system:
only H. sapiens has a complex and reticulated endocra-
Evolution of the Parietal Lobes
c
f
eas), and modern humans are further characterized by a general
longitudinal expansion of the same region (SD, sagittal dorsal ar-
eas). The former variation spatially matches the position of the
intraparietal sulcus and superior parietal lobules, while the latter
corresponds to the position of the superior parietal lobules and
precuneus. Neanderthals and modern humans share a similar cra-
nial capacity, but the latter shows a general enlargement of the pa-
rietal lobes (f – digital replica of skull and endocasts of the Nean-
dertal Saccopastore 1, and shape deformation associated with
modern human endocranial anatomy). Images after Bruner [2004,
2010, 2014] and Bruner et al. [2004, 2014a, 2015c].
nial vascular network as far as we can see for the menin-
geal and diploic vessels [Bruner et al., 2005; Bruner and
Sherkat, 2008; Rangel de Lázaro et al., 2016]. Vascular
morphology can show important individual variation,
and the mechanisms and factors involved in such diver-
sity are still not known. However, all extinct human spe-
cies display simpler vascular networks when compared
Brain Behav Evol 2018;91:136–147 139
DOI: 10.1159/000488889
Fig. 2. A principal component analysis on endocranial 3D coordi-
nates from hominoid endocasts using parietal lobe landmarks
shows a PC1 (48% of the total variation) separating humans from
all apes (above) because of parietal lobe enlargement (middle; red
wireframe shows the shape change toward humans). If we con-
sider only the parietal landmarks (below), PC1 (67%) separates
again humans from apes because of the enlargement of the dorsal
and anterior regions of the parietal lobe. In this example, endocasts
from 1 representative individual of each species (adult males) were
used, averaging 10 replicas and the 2 hemispheres, so as to limit
with modern humans, with almost no anastomotic con-
nections between the main branches. Apparently, vascu-
lar complexity does not depend on brain size, and it par-
ticularly concerns the parietal surface. This is relevant be-
140 Brain Behav Evol 2018;91:136–147
DOI: 10.1159/000488889
uncertainty due to landmark localization (data after Pereira-Pedro
and Bruner [2018]). Parietal landmarks: ab, angular boss; cs, cen-
tral sulcus (midline estimation); ls, posterior end of the lateral sul-
cus; pos, parieto-occipital sulcus (midsagittal estimation); sb, su-
pramarginal boss. Species: BON, Pan paniscus; CHIM, Pan troglo-
dytes; GIB, Hylobates lar; GOR, Gorilla gorilla; HUM, Homo
sapiens; ORA, Pongo pygmaeus; SIA, Symphalangus syndactylus.
Morphometrics was computed with MorphoJ 1.06d [Klingenberg,
2011].
cause the endocranial vascular system may have a central
role in brain thermoregulation [Bruner et al., 2011b]. Nu-
merical simulations showed that the dorsal parietal sur-
face in more platycephalic species is important for endo-
Bruner
cranial heat exchange, and the bulging of the parietal
region characteristic of modern humans can hence con-
tribute to increase the heat load of the corresponding
deeper cerebral areas [Bruner et al., 2012, 2014a].
Ontogenetic analysis of the endocranial form suggest-
ed that modern human brain globularity is achieved ear-
ly postnatally [Neubauer et al., 2009] or even prenatally
[Ponce de León et al., 2016]. Either way, it is due to a mor-
phogenetic stage which is specific to H. sapiens and absent
in chimps or Neanderthals [Gunz et al., 2010; Neubauer
et al., 2010; Scott et al., 2014]. In sum, head and brain
globularity in our species is largely due to parietal lobes
and bones, and associated with a species-specific growth
period. As far as the paleoneurological evidence can show,
such morphological differences are definitely more ap-
parent than many subtle or individual variations com-
monly debated in paleoanthropology or evolutionary
neuroanatomy. They should deserve, at least, attention.
The Living Evidence: Parietal Lobe Morphology
Parietal lobes are located below the parietal bones, al-
though the respective size and positions may vary [Brun-
er et al., 2015a]. The parieto-occipital sulcus, separating
the parietal and occipital cortex, is more stable in its cra-
nial location, roughly close to the boundary between the
parietal and occipital bones (lambda). In contrast, the po-
sition of the anterior boundary of the lobe (central sulcus
and postcentral gyrus) is more variable, and it becomes
relatively closer to the boundary of the frontal and pari-
etal bones (bregma) in brains with larger parietal lobes.
Although these positions and proportions are influenced
by many factors and distinct ontogenetic processes, there
is still a significant correlation between parietal bone and
lobe length even among adult humans. In adults, the form
of the parietal bone and lobe is integrated with the form
of the occipital bone and lobe [Gunz and Harvati, 2007;
Bruner et al., 2018b], and bulging of one of the two re-
gions is associated with the flattening of the other. In con-
trast, the size of the parietal lobe is inversely correlated
with the size of the frontal and temporal lobes, and not
correlated with the size of the occipital cortex [Allen et al.,
2002]. This may suggest more structural constraints with
the occipital region and more functional constraints with
frontal and temporal regions – an entangled morphoge-
netic position, indeed.
A key element of the dorsal parietal cortex is the pre-
cuneus, which displays remarkable morphological vari-
ability (Fig. 3). When dealing with the whole midsagittal
Evolution of the Parietal Lobes
brain geometry in adult humans, the most variable fea-
ture regards the precuneus longitudinal extension [Brun-
er et al., 2014b]. Namely, adult individuals show a notice-
able anatomical diversity because of the length of the pre-
cuneus. Differences mainly deal with the dorsal regions,
and enlargement of the precuneus is associated with in-
crease in its cortical surface area [Bruner et al., 2015b].
Parietal lobes generally present larger size, more connec-
tions, and larger surface area in males than females, a dif-
ference associated with distinct visuospatial performance
[Koscik et al., 2009]. Nonetheless, the overall precuneus
morphology apparently does not depend on brain size,
sex, hemisphere, or geographic ancestry [Bruner et al.,
2017b]. The individual differences in sulcal variability
may also be noticeable in these areas because of a variety
of precuneal and subparietal sulci, and precuneus expan-
sion is partially associated with additional folding ele-
ments [Bruner et al., 2017b]. Precuneus differences al-
most entirely occur along its longitudinal and vertical size
and extensions, influencing the dorsal midsagittal mor-
phology of the brain but not particularly the lateral (para-
sagittal) brain dimensions [Pereira-Pedro and Bruner,
2016]. Actually, precuneus lateral development (mainly
due to the lateral extension of the subparietal sulcus) is
less variable and does not apparently influence the exter-
nal brain form.
Interestingly, if we compare the midsagittal brain
morphology in humans and chimpanzees, we also find
that their main difference concerns the extension of the
precuneus, which is much larger in our species [Bruner et
al., 2017c]. This difference seems, again, to be localized in
the dorsal and anterior part of the precuneus, a region
that matches the area 7a according to Scheperjians et al.
[2008]. This area includes the anterior portion of the me-
dial surface of the precuneus but also the external dorsal
surface of the superior parietal lobules.
In terms of spatial correspondence, the morphological
changes associated with the precuneus at intra- and inter-
specific levels match the parietal bulging described in the
endocranial evolution of modern humans [Bruner et al.,
2014a].
It can be argued that modern human parietal expan-
sion may be due to some intrinsic allometric trend of the
brain and proportional scaling due to encephalization.
However, the available evidence largely rejects this pos-
sibility. In fact, concerning the fossil record, Neander-
thals had a brain size comparable with ours but no pari-
etal longitudinal expansion [Bruner et al., 2003; Bruner,
2004; Gunz et al., 2010]. Concerning nonhuman pri-
mates, the precuneus is particularly variable within other
Brain Behav Evol 2018;91:136–147 141
DOI: 10.1159/000488889
 b

Fig. 3. The vertical extension of the precuneus (a) is responsible for the main coronal variation among adult hu-
mans, strictly influencing the height of the dorsal parietal cortex (b). The width of the precuneus (c) is, in contrast,
less variable. The length of the dorsal part of the precuneus is also responsible for the main midsagittal brain shape
differences among adult humans, because of variations in its cortical surface area (d). A similar shape change also
represents the main difference between humans and chimps (e, f). Data after Bruner et al. [2014b], Pereira-Pedro
and Bruner [2016], Bruner et al. [2017b]. Morphometrics was computed with PAST 2.17c [Hammer et al., 2001].

species too, but there are no consistent differences in size
and proportions between different species, including be-
tween species with very distinct brain size [Pereira-Pedro
et al., 2017b]. Therefore, according to the converging ev-
idence we have on extinct and extant species, we can con-
clude that the derived parietal proportions in H. sapiens
are not due to general allometric effects associated with a
large brain. It is worth noting that, when using apes as an
allometric reference, humans display an expected parie-
to-occipital volume for their brain size [Semendeferi and
Damasio, 2000] but smaller occipital volume [De Sousa
et al., 2010] which, implicitly, should mean a larger pari-
etal cortex.
In general, as expected after a visual inspection, hu-
mans and nonhuman primates display further differenc-
es in their whole parietal organization. The pattern of
connections is distinct, notably at the inferior regions
[Catani et al., 2017]. Although fossils have provided
scanty evidence of gross morphological differences asso-
ciated with the inferior parietal lobules, these areas are
known to be specialized in humans as well [Bzdok et al.,
2016]. The intraparietal sulcus is also far more complex

142 Brain Behav Evol 2018;91:136–147 Bruner

DOI: 10.1159/000488889
in humans than in other primates, in terms of size and
organization [Grefkes and Fink, 2005; Choi et al., 2006],
and its evolutionary changes in our species are possibly
related to tool use [Kastner et al., 2017]. The homology
between humans and nonhuman primates for their pari-
etal cortex is not clear, although the complexity of the
intraparietal sulcus in H. sapiens may have displaced and
repositioned internal and external folding regions, gener-
ating a lack of correspondence between human and non-
human cortical topology [Zlatkina and Petrides, 2014]. It
must be taken into account that most inferences are based
on comparisons between humans and macaques, two lin-
eages that are not closely related in terms of phylogeny,
and that underwent separation and independent evolu-
tion for some 20 million years. Therefore, most results
deal with only one genus of primates (Macaca) and in
particular with one that is not related to human evolution.
Many areas seem evolutionarily conserved in humans
and macaques [Orban, 2016], but for those elements that
are evolutionarily derived – in humans as well as in ma-
caques – we still miss a comprehensive comparative sce-
nario. Both modern humans and Neanderthals display a
lateral bulging and widening of the dorsal parietal surface
[Bruner et al., 2003], which could be tentatively interpret-
ed as a cortical increase and outfold of the intraparietal
cortex [Pereira-Pedro and Bruner, 2016].
Comparative data on the superior parietal lobule are
still lacking. According to the parcellation proposed by
Scheperjans et al. [2008], what we call superior lobule in
humans is largely the outer extension of the anterior and
posterior areas of the precuneus. This dorsal external sur-
face has not been sufficiently investigated yet. At least
three areas have been identified in the human dorsal re-
gion, but comparative data for nonhuman primates are
lacking. These outer cortical folds are apparently involved
in our derived parietal morphology, and future surveys
should be specifically dedicated to investigate this issue.
Visuospatial Integration and Body Cognition
The precuneus is involved in a functional integration
of body and vision, bridging somatosensory and occipital
signals [Cavanna and Trimble, 2006; Margulies et al.,
2009; Zhang and Li, 2012; Freton et al., 2014]. Its anterior
area deals mostly with body cognition, the posterior area
with visual cognition, and the intermediate area inte-
grates both signals. This process, generally named visuo-
spatial integration, not only deals with body-environ-
ment physical coordination, but also with visual imaging
Evolution of the Parietal Lobes
[Land, 2014] and conscious self-centered memory recall
[Fletcher et al., 1995]. Visual imagery is the basis for sim-
ulation and mental experiments. The body, in this sense,
is used as a measure of physical space, but also in terms
of chronological and social relationships (i.e., generating
a chronological space and a social space according to an
egocentric perspective) [Hills et al., 2015; Maister et al.,
2015; Peer et al., 2015]. The intraparietal sulcus is also
specialized in visuospatial capacities, particularly in eye-
hand coordination [Grefkes and Fink, 2005; Tunik et al.,
2007; Martin et al., 2011; Verhagen et al., 2012]. These
visuospatial functions are relevant for all primates, but
they are far more crucial for our species, which is charac-
terized by complex tools and technology, visual symbols,
and extensive social structure. Interestingly, potential
early modern humans, dated at around 100–200,000
years ago, did not display larger parietal bones and lobes,
suggesting that modern human phylogenetic origins may
have preceded the evolution of a modern human brain
form [Bruner and Pearson, 2013; Bruner et al., 2018a]. An
inclusive survey suggested that, since the phylogenetic
origin of our species, these changes may have followed a
gradual pattern [Neubauer et al., 2018]. Paleoanthropolo-
gists find fossils with bulging parietal bones and lobes
roughly since the same dates (≈50–100,000 years ago) ar-
chaeologists find distinctive modern cultural traits, like
complex technology, larger social groups, and visual cul-
ture (paintings and ornamentation, for example). The ex-
act timing and dynamics of these changes are not known,
but, on an evolutionary and geological time scale, the two
changes are almost matching. We should consider wheth-
er or not this is due to chance.
The case of Neanderthals merits attention, because pa-
leoneurology, skeletal biology, ecology, and archaeology
are supplying converging evidence on some aspects of
their visuospatial behavior [see Bruner and Lozano, 2014,
2015 for a detailed discussion]. As we mentioned, al-
though they had a similar or even larger brain volume
than modern humans, they did not have large parietal
lobes. At the same time, there is no evidence of a robust
and complex visual culture, their social groups and ter-
ritories were smaller, and there is not even any evidence
of projectile technology (fine spears, spear throwers, or
arrows). Also, according to the traces found on their front
teeth, they largely relied on the mouth to handle their
tools, more than any modern human population. This
distinctive (and hazardous) degree of use of the mouth as
a “third hand” may suggest that hand management may
have been less efficient in dealing with their material cul-
ture [Bruner et al., 2016]. Hypotheses in cognitive archae-
Brain Behav Evol 2018;91:136–147 143
DOI: 10.1159/000488889
ology cannot be tested to the same extent we can do in
neontological fields, but all this evidence together points,
independently, to possible limits in visuospatial resources
and body cognition. This is not necessarily a verdict for
extinction, and it may even suggest that their large brains
may have had some distinct specializations that we did
not evolve [Pearce et al., 2013]. Nonetheless, the com-
parison with Neanderthals further supports the hypoth-
esis of derived – and probably enhanced – visuospatial
cognition in modern humans, associated with body-
environment management and body-tool functional spe-
cialization.
In the last 2 million years, our culture was not simply
“object assisted” but instead “object dependent” [Plum-
mer, 2004]. Our technology is actually part of our cogni-
tive processes, and there are several theories in current
neuroscience interpreting cognition as the result of inte-
gration between brain, body, and tools [Malafouris, 2010;
Iriki and Taoka, 2012; Byrge et al., 2014]. In this sense,
our parietal cortex and visuospatial functions can play a
major role in the management of such interactions be-
tween the nervous system, the body interface, and tech-
nological resources [Bruner and Iriki, 2016]. Interesting-
ly, a comprehensive analysis on neural circuits in pri-
mates evidenced major human changes in the genetic
expression associated with the striatum, an element of the
basal ganglia deeply involved in body management [Sou-
sa et al., 2017].
Of course, the nature behind this association between
form and function remains unclear. The parietal cortex is
sensitive to genetic programming [Chen et al., 2012] but
also to environmental influences, and its components are
highly susceptible to training [Quallo et al., 2009]. At
present, we ignore to what extent parietal differences be-
tween and within species are due to genes, culture, or to
different kinds of feedback between both factors. Culture
can influence phenotypic expression, but also genetic se-
lection, through direct effects (autocatalysis) or indirect
channeling of phenotypic plasticity [the so-called Bald-
win effect; see for example Crispo, 2007]. Thus, although
parietal differences begin now to be acknowledged in evo-
lutionary neuroanatomy, their causes, factors and mech-
anisms still remain, at present, largely unexplained.
Perspectives
The parietal cortex has been largely neglected in evo-
lutionary neuroanatomy (a case of “parietal neglect”!).
The precuneus is a large cortical element but, despite an
144 Brain Behav Evol 2018;91:136–147
DOI: 10.1159/000488889
early and abandoned term quadrate lobule due to its
square shape, it does not even have its own name, being
defined in relation to something else (precuneus means
anterior to the cuneus). Its broad designation includes
some inferior areas which intermingle with the posterior
cingulate and retrosplenial cortex. All this block, that
bridges the parietal lobe with the cingulate cortex, is high-
ly interconnected, but formed by distinct parts with dis-
tinct functions, and should be referred to as to posterior
medial cortex [Bzdok et al., 2015]. The inferior areas are
part of the default mode network and are crucial to func-
tions that are integrated but distinct from those of the
dorsal areas [Fransson and Marrelec, 2008; Margulies et
al., 2009; Utevsky et al., 2014; Yang et al., 2014; Barks et
al., 2015]. Therefore, probably the term precuneus should
be strictly limited to the dorsal areas above the subparietal
sulcus. I wonder whether the term quadrate lobule should
also be rescued and used to indicate this region. It also
remains to be established whether (or to what extent) the
superior parietal lobule is part of the same element. Of
course, the features described in this article should be in-
tegrated within a more comprehensive scenario. For ex-
ample, we currently recognize the crucial importance of
the frontoparietal system, these two regions being deeply
interconnected by functional and connective networks
[Jung and Haier, 2007; Caminiti et al., 2015].
The noticeable degree of morphological variation of
the parietal lobes, and in particular of the precuneus, has
been evidenced in distinct samples and through distinct
methods. It is clear that there are differences between hu-
mans and other primates, and probably the same region
displays a pronounced enlargement in H. sapiens com-
pared with fossil human species. There are multiple and
independent sources of evidence suggesting that the deep
folds of the parietal cortex have undergone some special-
ization in modern humans. A mandatory future step is,
therefore, to study the histological changes associated
with these macroanatomical differences.
Geometrical models point to the dorsal and anterior
areas as a main source of variation. Nonetheless, also the
posterior (parieto-occipital) region may show human
specialization for body-tool coordination and allocentric
imaging [Hutchison et al., 2015; Sulpizio et al., 2016]. En-
hanced morphological models can help to localize more
specific areas involved in these volumetric variations in
human fossils as well as in living primates.
The dorsal parietal cortex is largely involved in visuo-
spatial integration, and future surveys should investigate
possible functional or cognitive factors associated with its
morphological diversity. Although differences in the pa-
Bruner
rietal cortex are associated with visuospatial performance
[Koscik et al., 2009], at present, there is no evidence cor-
relating the overall precuneus morphology with tradi-
tional psychometric scores [Bruner et al., 2015b]. In this
sense, and taking into account the cortical diversity of the
precuneus, future neurofunctional analyses should con-
sider more specific cortical areas and probably more tar-
geted behavioral tasks.
Together with this “to-do” list, there is also a “not-to-
do” list, which particularly refers to things that should be
avoided when handling the fossil record. First, hypothe-
ses should be tested through samples and not through
specimens. Many paleoneurological traits display large
intragroup variation and scarce intergroup differences.
Therefore, individual fossils can hardly supply consistent
support to promote or demote complex evolutionary
scenarios. Second, quantitative analyses are necessary to
provide a proper comparative framework. Descriptive
studies should be restricted to preliminary and introduc-
tory surveys only. Third, speculations should be used
with discretion and common sense, and quickly substi-
tuted by specific hypotheses that can be (at least partially)
tested. Such hypotheses should be proposed and corrobo-
rated through multiple and independent lines of evi-
dence, and not following specific or punctual results.
Fourth, endocasts are valuable sources of information,
but their general shape is the result of distinct factors (cra-
nial and cerebral) and distinct elements (cortical areas).
Therefore, overall shape analyses should be limited to
preliminary geometrical surveys. More detailed morpho-
metrics, based on anatomical boundaries, should be used
to test specific hypotheses in evolutionary neuroanatomy
[Bruner and Ogihara, 2018]. Fifth, more information is
needed about modern human variation. Many paleoneu-
rological traits are not even known, in terms of variability
and function, in our species. Therefore, proper knowl-
edge of large living samples is mandatory before we make
inferences on a few fragmented bony remains.
Taking into account the many disputes and disagree-
ments on subtle anatomical or statistical issues in paleo-
anthropology and evolutionary neuroanatomy, it is cu­
rious how the large parietal differences have passed un-
noticed, and they are not even mentioned in many
scientific reviews. Nonetheless, at this time, neglecting
such evidence seems to be unreasonable. Once the differ-
ence has been noticed, it should be investigated properly.
And, if visuospatial functions have a major role in extend-
ing our “prosthetic” capacity in terms of space, time, tech-
nology, body cognition, and social perception, we should
once more consider that brain evolution is basically a
matter of anatomy, while cognitive evolution must deal
with additional components (body and culture) that go
beyond the neuronal content of a braincase.
Acknowledgments
I would like to thank Ashley Morhardt and Georg Striedter for
giving me the opportunity to participate in the 29th Karger Work-
shop “From Fossils to Function: Integrative and Taxonomically
Inclusive Approaches to Vertebrate Evolutionary Neuroscience.”
Many and diverse studies on parietal evolution have been possible
thanks to the collaboration with many people, including Giorgio
Manzi, Ralph Holloway, Jim Rilling, Todd Preuss, Naomichi Ogi-
hara, Manuel Martín-Loeches, Roberto Colom, Heidi Jacobs, Marina
Lozano, Philipp Gunz, Simon Neubauer, Atsushi Iriki, Duilio Garo-
foli, Amélie Beaudet, Barbara Saracino, Fred Coolidge, Roberto Mac-
chiarelli, and Xiujie Wu. I am particularly grateful to all the students
who have supported and developed my laboratory over the years,
namely Sofia Pereira-Pedro, Gizéh Rangel de Lázaro, José Manuel de
la Cuétara, Annapaola Fedato, Alannah Pearson, María Silva Gago,
Hideki Amano, and Stana Eisová. These research lines were primar-
ily funded by the Proyecto Atapuerca (Spanish Government), coor-
dinated by José María Bermudez de Castro, as well as by the Italian
Institute of Anthropology and the Wenner-Gren Foundation.
Disclosure Statement
The author declares no conflict of interest.

References
Allen JS, Damasio H, Grabowski TJ (2002): Nor-
mal neuroanatomical variation in the human
brain: an MRI-volumetric study. Am J Phys
Anthropol 118:341–358.
Barks SK, Parr LA, Rilling JL (2015): The default
mode network in chimpanzees (Pan troglo-
dytes) is similar to that of humans. Cereb Cor-
tex 25:538–544.
Beaudet A, Bruner E (2017): A frontal lobe surface
analysis in three archaic African human fos-
sils: OH 9, Buia, and Bodo. Comptes Rendus
Palevol 16:499–507.
Bookstein F, Schafer K, Prossinger H, Seidler H,
Fieder M, Stringer C, Weber GW, Arsuaga JL,
Slice DE, Rohlf FJ, Recheis W, Mariam AJ,
Marcus LF (1999): Comparing frontal cranial
profiles in archaic and modern Homo by mor-
phometric analysis. Anat Rec 257:217–224.
Bruner E (2004): Geometric morphometrics and
paleoneurology: brain shape evolution in the
genus Homo. J Hum Evol 47:279–303.
Bruner E (2010): Morphological differences in the
parietal lobes within the human genus: a neu-
rofunctional perspective. Curr Anthropol
51:S77–S88.

Evolution of the Parietal Lobes Brain Behav Evol 2018;91:136–147 145

DOI: 10.1159/000488889
Bruner E (2014): Functional craniology, human
evolution, and anatomical constraints in the
Neanderthal braincase; in Akazawa T, Ogi-
hara N, Tanabe HC, Terashima H (eds): Dy-
namics of Learning in Neanderthals and
Modern Humans. Tokyo, Springer, vol 2, pp
121–129.
Bruner E (2015): Functional craniology and brain
evolution; in Bruner E (ed): Human Paleo-
neurology. Basel, Springer, pp 57–94.
Bruner E (2017a): The fossil evidence of human
brain evolution; in Kaas J (ed): Evolution of
Nervous Systems, ed 2. Oxford, Elsevier, vol
4, pp 63–92.
Bruner E (2017b): Language, paleoneurology, and
the fronto-parietal system. Front Hum Neu-
rosci 11:349.
Bruner E, Amano H, de la Cuétara JM, Ogihara N
(2015a): The brain and the braincase: a spatial
analysis on the midsagittal profile in adult hu-
mans. J Anat 227:268–276.
Bruner E, Amano H, Pereira-Pedro AS, Ogihara
N (2018a): The evolution of the parietal lobes
in the genus Homo; in Bruner E, Ogihara N,
Tanabe H (eds): Digital Endocasts. Tokyo,
Springer, pp 219–258.
Bruner E, de la Cuétara JM, Holloway RL (2011a):
A bivariate approach to the variation of the
parietal curvature in the genus Homo. Anat
Rec 294:1548–1556.
Bruner E, de la Cuétara JM, Masters M, Amano
H, Ogihara N (2014a): Functional craniology
and brain evolution: from paleontology to
biomedicine. Front Neuroanat 8:19.
Bruner E, de La Cuétara JM, Musso F (2012):
Quantifying patterns of endocranial heat dis-
tribution: brain geometry and thermoregula-
tion. Am J Hum Biol 24:753–762.
Bruner E, Esteve-Altava B, Rasskin-Gutman D
(2018b): Networking brains: modeling spatial
relationships of the cerebral cortex; in Bruner
E, Ogihara N, Tanabe H (eds): Digital Endo-
casts. Tokyo, Springer, pp 191–204.
Bruner E, Grimaud-Hervé D, Wu X, de la Cué-
tara JM, Holloway R (2015c): A paleoneuro-
logical survey of Homo erectus endocranial
metrics. Quat Int 368:80–87.
Bruner E, Iriki A (2016): Extending mind, visuo-
spatial integration, and the evolution of the
parietal lobes in the human genus. Quat Int
405:98–110.
Bruner E, Lozano M (2014): Extended mind and
visuo-spatial integration: three hands for the
Neandertal lineage. J Anthropol Sci 92: 273–
280.
Bruner E, Lozano M (2015): Three hands: one
year later. J Anthropol Sci 93:191–195.
Bruner E, Lozano M, Lorenzo C (2016): Visuo-
spatial integration and human evolution: the
fossil evidence. J Anthropol Sci 94:81–97.
Bruner E, Mantini S, Musso F, de la Cuétara JM,
Ripani M, Sherkat S (2011b): The evolution of
the meningeal vascular system in the human
genus: from brain shape to thermoregulation.
Am J Hum Biol 23:35–43.
Bruner E, Mantini S, Perna A, Maffei C, Manzi G
(2005): Fractal dimension of the middle men-
ingeal vessels: variation and evolution in
Homo erectus, Neanderthals, and modern hu-
mans. Eur J Morphol 42:217–224.
Bruner E, Manzi G, Arsuaga JL (2003): Encepha-
lization and allometric trajectories in the ge-
nus Homo: evidence from the Neandertal and
modern lineages. Proc Natl Acad Sci USA
100:15335–15340.
Bruner E, Ogihara N (2018): Surfin’ endocasts:
the good and the bad on brain form. Palaentol
Electr 21.1.1A:1–10.
Bruner E, Pearson O (2013): Neurocranial evolu-
tion in modern humans: the case of Jebel
Irhoud 1. Anthropol Sci 121:31–41.
Bruner E, Pereira-Pedro AS, Bastir M (2017a):
Patterns of morphological integration be-
tween parietal and temporal areas in the hu-
man skull. J Morphol 278:1312–1320.
Bruner E, Pereira-Pedro AS, Chen X, Rilling JK
(2017b): Precuneus proportions and cortical
folding: a morphometric evaluation on a ra-
cially diverse human sample. Ann Anat 211:
120–128.
Bruner E, Preuss T, Chen X, Rilling J (2017c): Evi-
dence for expansion of the precuneus in human
evolution. Brain Struct Funct 222:1053–1060.
Bruner E, Rangel de Lázaro G, de la Cuétara JM,
Martín-Loeches M, Colóm R, Jacobs HIL
(2014b): Midsagittal brain variation and MRI
shape analysis of the precuneus in adult indi-
viduals. J Anat 224:367–376.
Bruner E, Román FJ, de la Cuétara JM, Martín-
Loeches M, Colóm R (2015b): Cortical surface
area and cortical thickness in the precuneus of
adult humans. Neuroscience 286:345–352.
Bruner E, Saracino B, Ricci F, Tafuri M, Passarello
P, Manzi G (2004): Midsagittal cranial shape
variation in the genus Homo by geometric
morphometrics. Coll Antropol 28:99–112.
Bruner E, Sherkat S (2008): The middle menin-
geal artery: from clinics to fossils. Childs Nerv
Syst 24:1289–1298.
Byrge L, Sporns O, Smith LB (2014): Develop-
mental process emerges from extended brain-
body-behavior networks. Trends Cogn Sci 18:
395–403.
Bzdok D, Hartwigsen G, Reid A, Laird AR, Fox
PT, Eickhoff SB (2016): Left inferior parietal
lobe engagement in social cognition and lan-
guage. Neurosci Biobehav Rev 68:319–334.
Bzdok D, Heeger A, Langner R, Laird AR, Fox PT,
Palomero-Gallagher N, Vogt BA, Zilles K,
Eickhoff SB (2015): Subspecialization in the
human posterior medial cortex. NeuroImage
106:55–71.
Caminiti R, Innocenti GM, Battaglia-Mayer A
(2015): Organization and evolution of parie-
to-frontal processing streams in macaque
monkeys and humans. Neurosci Biobehav
Rev 56:73–96.
Catani M, Robertsson N, Beyh A, Huynh V, de
Santiago Requejo F, Howells H, Barrett RLC,
Aiello M, Cavaliere C, Dyrby TB, Krug K, Pti-
to M, D’Arceuil H, Forkel SJ, Dell’Acqua F
(2017): Short parietal lobe connections of the
human and monkey brain. Cortex 97: 339–
357.
Cavanna AE, Trimble MR (2006): The precuneus:
a review of its functional anatomy and behav-
ioural correlates. Brain 129:564–583.
Chen CH, Gutierrez ED, Thompson W, Panizzon
MS, Jernigan TL, Eyler LT, Fennema-Notes-
tine C, Jak AJ, Neale MC, Franz CE, Lyons MJ,
Grant MD, Fischl B, Seidman LJ, Tsuang MT,
Kremen WS, Dale AM (2012): Hierarchical
genetic organization of human cortical sur-
face area. Science 335:1634–1636.
Choi H-J, Zilles K, Mohlberg H, Schleicher A,
Fink GR, Armstrong E, Amunts K (2006): Cy-
toarchitectonic identification and probabilis-
tic mapping of two distinct areas within the
anterior ventral bank of the human intrapari-
etal sulcus. J Comp Neurol 495:53–69.
Crispo E (2007): The Baldwin effect and genetic
assimilation: revisiting two mechanisms of
evolutionary change mediated by phenotypic
plasticity. Evolution 61:2469–2479.
De Sousa AA, Sherwood CC, Mohlberg H,
Amunts K, Schleicher A, MacLeod CE, Hof
PR, Frahm H, Zilles K (2010): Hominoid vi-
sual brain structure volumes and the position
of the lunate sulcus. J Hum Evol 58:281–292.
Falk D (1987): Hominid paleoneurology. Annu
Rev Anthropol 16:13–30.
Fletcher PC, Frith CD, Baker SC, Shallice T,
Frackowiak RSJ, Dolan RJ (1995): The mind’s
eye – precuneus activation in memory-related
imagery. NeuroImage 2:195–200.
Fransson P, Marrelec G (2008): The precuneus/
posterior cingulate cortex plays a pivotal role
in the default mode network: evidence from a
partial correlation network analysis. Neuro-
Image 42:1178–1184.
Freton M, Lemogne C, Bergouignan L, Delaveau
P, Lehéricy S, Fossati P (2014): The eye of the
self: precuneus volume and visual perspective
during autobiographical memory retrieval.
Brain Struct Funct 219:959–968.
Grefkes C, Fink GR (2005): The functional orga-
nization of the intraparietal sulcus in humans
and monkeys. J Anat 207:3–17.
Gunz P, Harvati K (2007): The Neanderthal “chi-
gnon”: variation, integration, and homology.
J Hum Evol 52:262–274.
Gunz P, Neubauer S, Maureille B, Hublin J-J
(2010): Brain development after birth differs
between Neanderthals and modern humans.
Curr Biol 20:R921–R922.
Hammer Ø, Harper D, Ryan P (2001): PAST: pa-
leontological statistics software package for
education and data analysis. Palaeontol Electr
4:9.
Hills TT, Todd PM, Lazer D, Redish AD, Couzin
ID (2015): Exploration versus exploitation in
space, mind, and society. Trends Cogn Sci 19:
46–54.
Holloway RL (1981): Exploring the dorsal surface
of hominoid brain endocasts by stereoplotter
and discriminant analysis. Philos Trans R Soc
Lond B 292:155–166.
Holloway RL, Broadfield DC, Yuan MS (2004):
The Human Fossil Record, vol III: Brain En-
docasts: The Paleoneurological Evidence.
Hoboken, Wiley-Liss.

146 Brain Behav Evol 2018;91:136–147 Bruner

DOI: 10.1159/000488889
Hutchison RM, Culham JC, Flanagan JR, Everling
S, Gallivan JP (2015): Functional subdivisions
of medial parieto-occipital cortex in humans
and nonhuman primates using resting-state
fMRI. NeuroImage 116:10–29.
Iriki A, Taoka M (2012): Triadic (ecological, neu-
ral, cognitive) niche construction: a scenario
of human brain evolution extrapolating tool
use and language from the control of reaching
actions. Philos Trans R Soc Lond B Biol Sci
367:10–23.
Jung RE, Haier RJ (2007): The Parieto-Frontal In-
tegration Theory (P-FIT) of intelligence: con-
verging neuroimaging evidence. Behav Brain
Sci 30:135–154.
Kastner S, Chen Q, Jeong SK, Mruczek REB
(2017): A brief comparative review of primate
posterior parietal cortex: a novel hypothesis
on the human toolmaker. Neuropsychologia
105:123–134.
Klingenberg CP (2011): MorphoJ: an integrated
software package for geometric morphomet-
rics. Mol Ecol Resour 11:353–357.
Kobayashi Y, Matsui T, Haizuka Y, Ogihara N,
Hirai N, Matsumura G (2014): Cerebral sulci
and gyri observed on macaque endocasts:
in Akazawa T, Ogihara N, Tanabe HC, Tera­
shima H (eds): Dynamics of Learning in
Neanderthals and Modern Humans. Tokyo,
Springer, vol 2, pp 131–137.
Koscik T, O’Leary D, Moser DJ, Andreasen NC,
Nopoulos P (2009): Sex differences in parietal
lobe morphology: relationship to mental rota-
tion performance. Brain Cogn 69:451–459.
Land MF (2014): Do we have an internal model of
the outside world? Philos Trans R Soc Lond B
Biol Sci 369:20130045–20130045.
Lieberman DE, McBratney BM, Krovitz G (2002):
The evolution and development of cranial
form in Homo sapiens. Proc Natl Acad Sci
USA 99:1134–1139.
Maister L, Slater M, Sanchez-Vives MV, Tsakiris
M (2015): Changing bodies changes minds:
owning another body affects social cognition.
Trends Cogn Sci 19:6–12.
Malafouris L (2010): The brain-artefact interface
(BAI): a challenge for archaeology and cultur-
al neuroscience. Soc Cogn Affect Neurosci 5:
264–273.
Margulies DS, Vincent JL, Kelly C, Lohmann G,
Uddin LQ, Biswal BB, Villringer A, Castella-
nos FX, Milham MP, Petrides M (2009): Pre-
cuneus shares intrinsic functional architec-
ture in humans and monkeys. Proc Natl Acad
Sci USA 106:20069–20074.
Martin K, Jacobs S, Frey SH (2011): Handedness-
dependent and -independent cerebral asym-
metries in the anterior intraparietal sulcus
and ventral premotor cortex during grasp
planning. NeuroImage 57:502–512.
Moss ML, Young RW (1960): A functional ap-
proach to craniology. Am J Phys Anthropol
18:281–292.
Mountcastle VB (1995): The parietal system and
some higher brain functions. Cereb Cortex 5:
377–390.
Evolution of the Parietal Lobes
Neubauer S, Gunz P, Hublin J-J (2009): The pat-
tern of endocranial ontogenetic shape chang-
es in humans. J Anat 215:240–255.
Neubauer S, Gunz P, Hublin J-J (2010): Endocra-
nial shape changes during growth in chim-
panzees and humans: a morphometric analy-
sis of unique and shared aspects. J Hum Evol
59:555–566.
Neubauer S, Hublin J-J, Gunz P (2018): The evo-
lution of modern human brain shape. Sci Adv
4:eaao5961.
Orban GA (2016): Functional definitions of pari-
etal areas in human and non-human pri-
mates. Proc Biol Sci 283:20160118.
Pearce E, Stringer C, Dunbar RIM (2013): New
insights into differences in brain organization
between Neanderthals and anatomically
modern humans. Proc Biol Sci 280:1758.
Peer M, Salomon R, Goldberg I, Blanke O, Arzy S
(2015): Brain system for mental orientation in
space, time, and person. Proc Natl Acad Sci
USA 112:11072–11077.
Pereira-Pedro AS, Bruner E (2016): Sulcal pat-
tern, extension, and morphology of the precu-
neus in adult humans. Ann Anat 208:85–93.
Pereira-Pedro AS, Bruner E (2018): Landmarking
endocasts; in Bruner E, Ogihara N, Tanabe H
(eds): Digital Endocasts. Tokyo, Springer, pp
127–142.
Pereira-Pedro AS, Masters M, Bruner E (2017a):
Shape analysis of spatial relationships be-
tween orbito-ocular and endocranial struc-
tures in modern humans and fossil hominids.
J Anat 231:947–960.
Pereira-Pedro AS, Rilling JL, Chen X, Preuss TM,
Bruner E (2017b): Midsagittal brain variation
among non-human primates: insights into
evolutionary expansion of the human precu-
neus. Brain Behav Evol 90:255–263.
Plummer T (2004): Flaked stones and old bones:
biological and cultural evolution at the dawn
of technology. Yrb Phys Anthropol 47: 118–
164.
Ponce de León MS, Bienvenu T, Akazawa T, Zol-
likofer CPE (2016): Brain development is sim-
ilar in Neanderthals and modern humans.
Curr Biol 26:R665–R666.
Quallo MM, Price CJ, Ueno K, Asamizuya T,
Cheng K, Lemon RN, Iriki A (2009): Gray and
white matter changes associated with tool-use
learning in macaque monkeys. Proc Natl
Acad Sci USA 106:18379–18384.
Rangel de Lázaro G, de la Cuétara JM, Píšová H,
Lorenzo C, Bruner E (2016): Diploic vessels
and computed tomography: segmentation
and comparison in modern humans and fossil
hominids. Am J Phys Anthropol 159: 313–
324.
Richtsmeier JT, Aldridge K, de Leon VB, Panchal
J, Kane AA, Marsh JL, Yan P, Cole TM (2006):
Phenotypic integration of neurocranium and
brain. J Exp Zool 306B:360–378.
Brain Behav Evol 2018;91:136–147
DOI: 10.1159/000488889
Scheperjans F, Hermann K, Eickhoff SB, Amunts
K, Schleicher A, Zilles K (2008): Observer-in-
dependent cytoarchitectonic mapping of the
human superior parietal cortex. Cereb Cortex
18:846–867.
Scott N, Neubauer S, Hublin JJ, Gunz P (2014): A
shared pattern of postnatal endocranial devel-
opment in extant hominoids. Evol Biol 41:
572–594.
Semendeferi K, Damasio H (2000): The brain and
its main anatomical subdivisions in living
hominoids using magnetic resonance imag-
ing. J Hum Evol 38:317–332.
Semendeferi K, Lu A, Schenker N, Damasio H
(2002): Humans and great apes share a large
frontal cortex, Nat Neurosci 5:272–276.
Sousa AMM, Zhu Y, Raghanti MA, Kitchen RR,
Onorati M, Tebbenkamp ATN, Stutz B, Mey-
er KA, Li M, Kawasawa YI, Liu F, Perez RG,
Mele M, Carvalho T, Skarica M, Gulden FO,
Pletikos M, Shibata A, Stephenson AR, Edler
MK, Ely JJ, Elsworth JD, Horvath TL, Hof
PR, Hyde TM, Kleinman JE, Weinberger DR,
Reimers M, Lifton RP, Mane SM, Noonan JP,
State MW, Lein ES, Knowles JA, Marques-
Bonet T, Sherwood CC, Gerstein MB, Sestan
N (2017): Molecular and cellular reorganiza-
tion of neural circuits in the human lineage.
Science 358:1027–1032.
Sulpizio V, Committeri G, Lambrey S, Berthoz A,
Galati G (2016): Role of the human retro­
splenial cortex/parieto-occipital sulcus in per-
spective priming. NeuroImage 125:108–119.
Tunik E, Rice NJ, Hamilton A, Grafton ST (2007):
Beyond grasping: representation of action in
human anterior intraparietal sulcus. Neuro-
Image 36:T77–T86.
Utevsky AV, Smith DV, Huettel SA (2014): Pre-
cuneus is a functional core of the default-
mode network. J Neurosci 34:932–940.
Van Minh N, Hamada Y (2017): Age-related
changes of sulcal imprints on the endocrani-
um in the Japanese macaque (Macaca fusca-
ta). Am J Phys Anthropol 163:285–294.
Verhagen L, Dijkerman HC, Medendorp WP,
Toni I (2012): Cortical dynamics of senso­
rimotor integration during grasp planning. J
Neurosci 32:4508–4519.
Yang Z, Chang C, Xu T, Jiang L, Handwerker DA,
Castellanos FX, Milham MP, Bandettini PA,
Zuo XN (2014): Connectivity trajectory
across lifespan differentiates the precuneus
from the default network. NeuroImage 89:
45–56.
Zhang S, Li CR (2012): Functional connectivity
mapping of the human precuneus by resting
state fMRI. NeuroImage 59:3548–3562.
Zilles K, Amunts K (2010): Centenary of Brod-
mann’s map – conception and fate. Nat Rev
Neurosci 11:139–145.
Zilles K, Palomero-Gallagher N (2001): Cyto-,
myelo-, and receptor architectonics of the hu-
man parietal cortex. NeuroImage 14:S8–S20.
Zlatkina V, Petrides M (2014): Morphological
patterns of the intraparietal sulcus and the an-
terior intermediate parietal sulcus of Jensen in
the human brain. Proc Biol Sci 281:20141493.
147