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Evolution of cranial capacity revisited: A view from the late Middle Pleistocene
cranium from Xujiayao, China

Article  in  Journal of Human Evolution · February 2022

DOI: 10.1016/j.jhevol.2021.103119

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 Journal of Human Evolution 163 (2022) 103119

Contents lists available at ScienceDirect

Journal of Human Evolution

journal homepage: www.elsevier.com/locate/jhevol

Evolution of cranial capacity revisited: A view from the late Middle

Pleistocene cranium from Xujiayao, China
Xiu-Jie Wu a, b, *, Christopher J. Bae c, *, Martin Friess d, Song Xing a, b, e, Sheela Athreya f,
Wu Liu a, b
a
Key Laboratory of Vertebrate Evolution and Human Origins, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences,
Beijing 100044, China
b
CAS Center for Excellence in Life and Paleoenvironment, Beijing 100044, China
c
Department of Anthropology, University of Hawai'i at Manoa, Honolulu 96822, USA
d 
Departement Homme et Environnement, UMR 7206 du CNRS, Mus eum National d’Histoire Naturelle, Universit
e de Paris, Paris 75016, France
e
Centro Nacional de Investigacion sobre la Evolucio
n Humana (CENIEH), Paseo de la Sierra de Atapuerca s/n, 09002 Burgos, Spain
f
Department of Anthropology, Texas A&M University, TX 77843, USA

a r t i c l e i n f o a b s t r a c t

Article history: The Late Middle Pleistocene hominin fossils from the Xujiayao site in northern China have been closely
Received 6 February 2021 studied in light of their morphological variability. However, all previous studies have focused on sepa-
Accepted 15 November 2021 rated cranial fragments. Here, we report the first reconstruction of a fairly complete posterior cranium,
Available online xxx
Xujiayao 6 (XJY 6), confidently dated to ~200e160 ka, which facilitated an assessment of its overall
cranial size. XJY 6 was reconstructed from three of the original fragmentsdthe PA1486 (No.7/XJY 6a)
Keywords:
occipital bone, PA1490 (No.10/XJY 6b) right parietal bone, and PA1498 (No.17/XJY 15) left temporal
Xujiayao
bonedwhich originated from the same young adult individual. The XJY 6 endocranial capacity, estimated
Cranium
Cranial capacity
by measuring endocranial volume, was estimated using multiple regression formulae derived from
Late Middle Pleistocene ectocranial and endocranial measurements on select samples of Pleistocene hominins and recent
China modern humans. The results indicate that the larger pooled sample of both Pleistocene and recent
modern humans was more robust for the endocranial capacity estimate. Based on the pooled sample
using the ectocranial and endocranial measurements, we conservatively estimate the XJY 6 endocranial
volume to be ~1700 cm3 with a 95% confidence interval of 1555e1781 cm3. This is close to Xuchang 1,
which dates to 125e105 ka and whose endocranial volume is ~1800 cm3. Thus, XJY 6 provides the
earliest evidence of a brain size that falls in the upper range of Neanderthals and modern Homo sapiens.
XJY 6, together with Xuchang 1, Homo floresiensis, Homo luzonensis, and Homo naledi, challenge the
general pattern that brain size gradually increases over geological time. This study also finds that
hominin brain size expansion occurred at different rates across time and space.
© 2021 Elsevier Ltd. All rights reserved.

1. Introduction dating of deposits northwest of Locality 74093 resulted in an age

The Xujiayao hominin fossils were found at Locality 74093 of the
Xujiayao (Houjiayao) site, in the Nihewan Basin, northern China,
during excavations conducted in the 1970s (Fig. 1). The site was
originally estimated to be 125e104 ka by uranium series on
mammal teeth that were found in association with the hominin
fossils (Chen et al., 1984; for review of ages, refer to the study by
Norton and Gao, 2008). More recently, electron spin resonance
* Corresponding authors.
E-mail addresses: wuxiujie@ivpp.ac.cn (X.-J. Wu), cjbae@hawaii.edu (C.J. Bae).
bracket of 370 to 270 ka (Ao et al., 2017). However, given the
complexity of the deposits in the Nihewan Basin, dates estimated
for the area around Loc. 74093 cannot, and probably should not, be
considered representative of the age of the cultural layer that
contains the Xujiayao hominin fossils (Wang and Li, 2020). A series
of optically stimulated luminescence ages taken from samples from
Loc. 74093 place the human remains between 200 and 160 ka (Li et
al., 2014, 2016). The latter dates fit well with the cold-adapted fauna
from the site (Chia et al., 1979; Norton and Gao, 2008).
The Xujiayao hominin fossil collection is comprised of 21 frag-
ments: one partial left maxilla, three isolated teeth, 13 parietal

https://doi.org/10.1016/j.jhevol.2021.103119
0047-2484/© 2021 Elsevier Ltd. All rights reserved.
X.-J. Wu, C.J. Bae, M. Friess et al.
Figure 1. Map showing the location of Xujiayao and other sites mentioned in the article.
fragments, two occipital bones, one partial mandible, and one left
temporal bone. These fragments represent at least 10 individuals.
Since the hominin remains were found, they have been the subject of
a variety of morphological studies (e.g., Chia et al., 1976, 1979; Wu,
1980, 1986; Wu et al., 2012, 2013, 2014; Wu and Trinkaus, 2014,
2015; Xing et al., 2015; Trinkaus and Wu, 2017). Unfortunately, all of
these previous studies focused on separated cranial fragments. Here,
we present a new reconstruction of the posterior cranium, Xujiayao
6 (XJY 6), using three separate fragments that were pieced together,
which allows for the estimation and analysis of the previously un-
known endocranial volume (ECV) of this hominin. The purpose of
this study is fourfold: 1) to provide a detailed description of the
reconstruction of XJY 6 and to determine the overall size of the
Xujiayao cranium; 2) to estimate ECV of XJY 6 using multiple re-
gressions derived from both ectocranial and endocranial measure-
ments; 3) to explore which methods are most suitable to estimate
ECV for partial fossils; and 4) to discuss the evolutionary position of
XJY 6 within the fossil record.
2. Materials and methods
2.1. The Xujiayao 6 fossil fragments
XJY 6 was reconstructed using one occipital (PA 1486/No.7/XJY
6a), a right parietal (PA 1490/No.10/XJY 6b), and the left temporal
(PA 1498/No.17/XJY 15), along with mirror images of the parietal
and temporal bones (Fig. 2) using computed tomography (CT) scans
of the originals. The three fossils are curated in the Institute of
Vertebrate Paleontology and Paleoanthropology (IVPP), Chinese
Academy of Sciences, Beijing.
PA1486 (Fig. 2A) is an occipital bone, 82.6 mm in height and
110.9 mm in width. It is represented by an almost complete
2
Journal of Human Evolution 163 (2022) 103119
occipital planum and most of the nuchal planum. The fossil was
found in 1979 and was briefly described in the original report (Chia
et al., 1979). PA1490 (Fig. 2B) is a largely complete right parietal
bone, 132 mm in length and 126 mm in width. The fossil was found
in 1977, in association with seven other hominin fragments (Wu,
1980). PA 1498 (Fig. 2C) is a largely complete left temporal bone
which was found in 1979 and was first published in 1986 (Wu,
1986). More recent research indicates that the morphological fea-
tures of the temporal labyrinths are similar to those of Neander-
thals, with a low and narrow anterior semicircular canal, a small
and circular posterior semicircular canal, and a high and wide
lateral semicircular canal (Wu et al., 2014). Although these three
Xujiayao fragments have been studied before, they had never been
joined together to reconstruct a more complete cranium. We
consider these three fairly intact bones to originate from the same
individual based on three key facts. First, the three fossils were
found in the same cultural layer and are all heavy and thick with a
deep yellow color. Second, the preserved sagittal, lambdoidal,
sphenoparietal, and squamosal sutures are all fully open, suggest-
ing an adolescent or (at most) a young adult age at the time of
death. Finally, there is clear anatomical continuity between the
thickness, internal meningeal grooves, and occlusal sutures of these
fossils (Fig. 2D, E).
2.2. XJY 6 reconstruction
The three posterior cranium fossils were scanned in coronal
orientation using an industrial high-precision CT scanner (450 kV-
ICT) housed at the IVPP, with the following parameters: X-ray tube
voltage 450 kV, X-ray tube current 1.5 mA, pixel matrix 1024
1024,
and 0.2 mm pixel size. The three-dimensional (3D) virtual re-
constructions of the isolated cranial fragments and the mirror
X.-J. Wu, C.J. Bae, M. Friess et al. Journal of Human Evolution 163 (2022) 103119

Figure 2. The three fossil and virtual fragments used to reconstruct Xujiayao 6 cranium. A1e3: PA1486, the occipital bone in exterior, superior, and interior views; B1e3: PA1490,
the right parietal bone in exterior, interior, and superior views; C1e4: PA 1498, the left temporal bone in exterior, medial, lateral, and inferior views; D: Virtual parietal, occipital and
the mirror image of the parietal bones, showing the morphological continuities between the parietal and occipital bones; E: Virtual temporal and the mirror image of the parietal
bones, showing morphological continuities between the parietal and temporal bones. Gray, mirror-imaged portion.

images of the left parietal and right temporal bones were carried out
using Mimics v. 20.0 (Materialise Inc., Belgium). The pieces in the 3D
virtual reconstruction were repositioned and reoriented along the
sutures in combination with relying on anatomical continuity of the
morphological features in virtual space.
After reconstruction, XJY 6 (Figs. 2D, E and 3AeF) is represented
by an almost complete posterior skull cap, consisting of largely
complete parietal bones, almost complete temporal bones, and an
almost complete occipital planum and most of the nuchal planum.
The morphological continuities between the parietal, temporal, and
occipital bones and endocranial cast reveal excellent anatomical
fits. The right parietal bone joins with the occipital bone at the right
lambdoid suture and fits well with the mirror-imaged left temporal
bone at the temporoparietal suture, and its mirror-imaged portion
articulates well with the left temporal bone.
The endocranial cast (endocast) of the preserved XJY 6 neuro-
cranium was virtually reconstructed in 3D using the existing
external cranial contours (Fig. 3GeI). It consists of the complete
right parietal lobes, a large portion of the left temporal lobe, and
most of the occipital lobe which joins with two upper smaller
portions of the cerebellar hemispheres. The base is missing. The
superior sagittal sinus is visible beginning as a narrow and shallow
groove between the two cerebral lobes and bifurcates at the
confluence, turning to the left and right transverse sinuses sepa-
rately. Although there is a gap between the temporal and occipital
lobes, the right transverse sinus crosses the temporal bone across
the posterior parietomastoid suture and continues to the sigmoid
sinus, which indicates that the reconstruction is reliable.
2.3. Comparative samples
To explore suitable formulae to estimate partial fossil ECV, a total
of 93 crania (Table 1) and 87 endocasts (Table 2), representing four
separate samples (Early Pleistocene, Middle Pleistocene, Late
3
Pleistocene and recent modern humans), were included in the
comparative analysis. For the Pleistocene comparative specimens, all
were adult and preserved most of the parietal and occipital lobes. For
the recent modern humans, the specimens were adult, as defined by
the presence of fully erupted permanent teeth. The comparative
specimens were sampled at the IVPP, American Museum of Natural
History (New York, USA), and from the Ralph Holloway endocast
collection (Columbia University, New York, USA).
2.4. Linear measurements
Taking into account the measurements that were available on the
XJY 6 reconstruction, two sets of measurements were selected for
ECV estimation: one for the ectocranium (seven measurements) and
one for the endocranium (eight measurements). For the ectocra-
nium, the seven linear measurements were as follows: maximum
cranial width, maximum frontal breadth, auricular-to-vertex height,
bregmaeasterion chord, lambdaeasterion chord, biasterionic
breadth, and parietal chord length (Fig. 4). The eight endocranial
measurements were as follows: maximum endocast width,
maximum endocast frontal width, endo-biasterionic breadth, supe-
rior cerebral height, endo-bregma-to-occipital pole length, parietal
lobe chord length, endo-bregma-to-asterion length, and endo-
lambda-to-asterion length (Fig. 5).
Linear measurements were taken on XJY 6 and the comparative
specimens using both sliding calipers on physical specimens and
Rapidform XOR3 software (INUS Technology, Inc) on 3D virtual
reconstructions. All measurements were taken by the same
observer (X.W.), except for Harbin 1, whose measurements were
culled from the study by Ni et al. (2021). Each specimen was
measured three times, and the average was used as the final
measurement. The measurements were to the nearest 0.1 mm. The
raw data are presented in Supplementary Online Materials (SOM
Tables S1 and S2).
X.-J. Wu, C.J. Bae, M. Friess et al. Journal of Human Evolution 163 (2022) 103119

Figure 3. The Xujiayao 6 reconstruction. Virtual external cranium reassembly in A) superior, B) anterior, C) left lateral, D) right lateral, E) posterior, and F) inferior views. Virtual
endocranial cast reassembly in G) superior, H) posterior, and I) right lateral views. Gray: mirror-imaged portion.

Table 1
Comparative cranial groups and specimens used for XJY 6 ECV analyses.

Groups Cranial specimens

Early Pleistocene (n ¼ 5) KNM-ER 3733a; KNM-ER 3833a; Sangiran 2a, 10a, 17a
Middle Pleistocene (n ¼ 23)  1b; Broken Hilla; Jebel Irhoud 1, 2b; Omo-Kibish 2b; Laetoli hominid 18b;
Sale
Sambungmacan 1, 3b; Narmada 1a; Atapuerca Sima de los Huesos 4, 5b; Petralona 1b; Steinheim 1b;
Zhoukoudian 3, 5, 10, 11, 12a; Nanjing 1a; Hexian 1c; Dali 1a; Jinniushan 1a; Harbind
Late Pleistocene (n ¼ 35) Ngandong 6, 7, 12, 13, 14a; Shanidar 1b; Saccopastore 1b; Amud 1a; La Chapelle aux Saints 1a;
La Ferrassie 1b; Spy 1, 2b; Le Moustier 1a; Guattari 1a; Gibraltar 1a; Teshik Tash 1b; Xuchang 1a;
Qafzeh 6, 9; Skhul 5b; Liujiang 1c; Upper cave 101, 102, 103a; Minatogawa 1, 4; Brno 2, 3a; Cro-Magnon 1a;
Mladec 1, 5b; Prĕdmostí 3, 4a; Oberkassel 1, 2a.
Recent modern humans (n ¼ 30) Chinac
a
Casts curated at the Institute of Vertebrate Paleontology and Paleoanthropology (IVPP), Chinese Academy of Sciences.
b
Casts curated at the American Museum of Natural History, New York, US.
c
Original fossils or skulls curated at the IVPP.
d
Data taken from the study by Ni et al. (2021).

2.5. Endocranial volume estimation 3. Results

The ECV of XJY 6 was estimated by using multiple regressions
derived from the metric variables for the comparative crania and
endocasts. Because morphological features vary across hominin
taxa, to make comparisons and choose suitable formulae to esti-
mate the ECV of partial hominin fossils, five separate regressions
were run for the ectocranial measurements (Table 1) and five for
the endocranial measurements (Table 2). The groups used for the
regressions include: Early Pleistocene, Middle Pleistocene, Late
Pleistocene, recent modern humans, and Pleistocene þ recent
modern humans. Multiple regression parameters were then used to
estimate the XJY 6 ECV. Significance level for the multiple re-
gressions was set at a p-value of 0.05 and an F-value of 3.84. All
statistical analyses were performed using SPSS v. 20.0 (IBM com-
pany, Beijing).
3.1. Linear metric comparisons
The metric data for XJY 6 and the comparative crania and endo-
casts are presented in Table 3. In general, XJY 6 is large, low, and
wide. The broadest region is situated in the posterior area of the
temporal bone. The maximum cranial width (175.0 mm), maximum
frontal breadth (135.0 mm), parietal chord length (118.0 mm),
bregmaeasterion chord (155.0 mm), and lambdaeasterion chord
(99.0 mm) are all in the top part of the range of the comparative
Pleistocene and recent modern human samples. Biasterionic breadth
(122.0 mm) falls within the middle range of the Pleistocene hominin
sample. The auricular-to-vertex height of XJY 6 is 106.0 mm, which is
high for Early Pleistocene Homo and in the range of Middle and Late
Pleistocene Homo and recent modern humans. However, when

4
X.-J. Wu, C.J. Bae, M. Friess et al. Journal of Human Evolution 163 (2022) 103119

Table 2
Comparative endocast groups and specimens used for XJY 6 ECV analyses.

Groups Endocast specimens

Early Pleistocene (n ¼ 5) KNM-ER 3733a; KNM-ER 3833a; Trinil 2a; Sangiran 2, 17b
Middle Pleistocene (n ¼ 13) 1b; Broken Hillb; Zhoukoudian 2, 3, 5, 10, 11, 12a; Hexian 1a; Sambungmacan 3a; Arago 21b; Swanscombeb; Jebel Irhoud 1b
Sale
Late Pleistocene (n ¼ 19) La Chapelle 1b; La Ferrassie 1b; Neanderthal lb; Spy 1, 2b; Guattari 1b; Reilingen lb; Amud 1b;
Ngandong 1, 6, 7, 12, 14b; Xuchang 1a; Liujiang 1a; Prĕdmostí 3, 4, 9, 10a
Recent modern humans Worldwidea: China ¼ 20, Europe ¼ 15, Africa ¼ 15
(n ¼ 50)
a
Curated at the Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences.
b
Curated at the Ralph Holloway collection at Columbia University, New York.

scaled against its broad cranial base, the breadth-height index is very
low (60.8), where only a few H. erectus specimens (KNM-ER 3883,
D2700, Sangiran 2, 4, and 17) are lower.
Endocranially (Fig. 3), the endocast is very low and very wide as
seen in most Middle and a few Early Late Pleistocene fossils from
East Asia. The maximum endocast width (156.5 mm), maximum
endocast frontal width (124.0 mm), and the endo-biasterionic
breadth (122.0 mm) are wider than Early and Middle Pleistocene
hominins. The superior cerebral height (66.5 mm) is close to Late
Pleistocene and recent modern humans. The endo-bregma-to-
occipital pole length (150.5 mm), the parietal lobe chord length
(108.5 mm), the endo-bregma-to-asterion length (132.5 mm), and
endo-lambda-to-asterion length (87.5 mm) are longer than those of
the other comparative specimens.
3.2. Endocranial volume estimation using ectocranial and
endocranial measurements
The results of the multiple regressions using ectocranial mea-
surements and the derived ECVs for XJY 6 are provided in Table 4.
Using ectocranial measurements, four of the five regressions were
significant at p < 0.001, with adjusted r2 (r2adj) values ranging be-
tween 0.607 and 0.986 (Table 4). The ECV estimates and 95% con-
fidence intervals (CIs) are 1644 (1360e1929) cm3, 1784
(1529e2039) cm3, 1779 (1718e1840) cm3, and 1669 (1555e1778)
cm3. When including only the Early Pleistocene group (n ¼ 5) for
the final model with ECV as the dependent variable, standardized
residuals and the standard error (std. error) of the estimate could
not be computed because the correlation coefficient is 1.000 and F
¼ 0.
The results of the multiple regression equations using endo-
cranial measurements and the derived ECVs for XJY 6 are provided
in Table 5. When using endocranial measurements, four of the five
regression equations were significant at p < 0.05, with r2adj values
ranging between 0.903 and 0.957. The ECV estimates and 95% CIs
are 1578 (1361e1794) cm3, 1767 (1552e1983) cm3, 1648
(1556e1740) cm3, and 1719 (1657e1780) cm3. As with the ectoc-
ranial measurements, when including only the Early Pleistocene
group (n ¼ 5) for the final model with ECV as the dependent var-
iable, standardized residuals and the std. error of the estimate could
not be computed because the correlation coefficient is 1.000 and F
¼ 0.

Figure 4. The ectocranial measurements used for ECV estimation on the XJY 6 cranium. The cranium was placed on the Frankfort Horizontal using the similar length of Dali as
reference. AeC: posterior, superior, and right lateral views. XCB ¼ maximum cranial width; XFB ¼ maximum frontal breadth on the lateral side of the coronal suture;
AVH ¼ auricular-to-vertex height; BAC ¼ bregmaeasterion chord; LASC ¼ lambdaeasterion chord; ASB ¼ biasterionic breadth; PAC ¼ parietal chord length.

5
X.-J. Wu, C.J. Bae, M. Friess et al. Journal of Human Evolution 163 (2022) 103119

Figure 5. The linear measurements used for ECV estimation on the XJY 6 endocast. The endocast was placed on a flat surface with the occipital poles (left side) horizontal along the
axis of the frontal and occipital poles (left side) parallel to the flat surface. The frontal poles were taken using the similar size of Xuchang 1 as reference. The asterion is defined at the
intersection of endo-lambdoid and endo-parietotemporal sutures. AeC: posterior, superior, and right lateral views. EXCB ¼ maximum endocast width; EASB ¼ endo-biasterionic
breadth; SCH ¼ superior cerebral height; BOCL ¼ endo-bregma-to-occipital pole length; EPAC ¼ parietal lobe chord length; EBAC ¼ endo-bregma-to-asterion length; ELASC ¼ endo-
lambda-to-asterion length; EXFB ¼ maximum endocast frontal width on bilateral endo-coronal suture.

Table 3
Metric data (mm) on XJY 6, XUC 1, and comparative Pleistocene and recent humans for ECV estimation.

Measurements XJY 6 Early Pleistocene Middle Pleistocene Late Pleistocene Recent modern humans

Maximum cranial width 175.0 146.0 ± 8.3 150.8 ± 9.1 147.3 ± 9.1 136.6 ± 8.2
Maximum frontal breadth 135.0 105.1 ± 8.7 116.2 ± 9.7 121.7 ± 7.3 115.4 ± 8.7
Parietal chord length 118.0 92.4 ± 8.7 105.1 ± 10.5 112.7 ± 8.7 110.9 ± 7.4
Bregmaeasterion chord 155.0 123.7 ± 7.2 132.8 ± 8.5 138.7 ± 6.1 130.6 ± 6.5
Lambdaeasterion chord 99.0 77.0 ± 4.9 84.8 ± 7.7 89.5 ± 6.1 85.2 ± 5.0
Biasterionic breadth 122.0 117.5 ± 5.2 121.4 ± 9.6 119.1 ± 9.1 107.5 ± 6.3
Auricular-to-vertex height 106.0 90 ± 3.9 102.0 ± 7.8 111.2 ± 6.3 110.6 ± 4.6
Maximum endocast width 156.5 122.4 ± 3.3 128.1 ± 8.8 139.3 ± 8.1 133.3 ± 7.3
Maximum endocast frontal width 124.0 101.4 ± 4.5 106.4 ± 8.2 119.6 ± 7.6 115.4 ± 6.7
Endo-biasterionic breadth 118.0 94.7 ± 2.1 98.9 ± 6.4 108.1 ± 8.5 102.5 ± 6.2
Superior cerebral height 66.5 52.9 ± 3.3 59.7 ± 4.6 67.6 ± 7.7 74.8 ± 3.7
Endo-bregma-to-occipital pole length 150.5 111.5 ± 4.4 122.7 ± 9.8 134.5 ± 7.7 132.3 ± 6.7
Parietal lobe chord length 108.5 85.3 ± 8.4 94.7 ± 7.6 105.6 ± 9.7 105.5 ± 5.8
Endo-bregma-to-asterion length 132.5 105.1 ± 4.1 114.7 ± 8.7 125.6 ± 7.9 125.4 ± 5.1
Endo-lambda-to-asterion length 87.5 62.6 ± 2.4 69.2 ± 7.0 76.7 ± 6.0 79.2 ± 5.9

When comparing the ECV results using the ectocranial and
endocranial measurements, the larger pooled sample furnishes a
slightly smaller 95% CI for the XJY 6 estimate. When utilizing the
larger pooled sample of all the ectocranial specimens, the multiple
regression provides an ECV estimate for XJY 6 of 1669 cm3 with a 95%
CI of 1555e1781 cm3. When using the larger pooled sample of all the
endocranial specimens, the multiple regression provides an ECV
estimate for XJY 6 of 1719 cm3 and a 95% CI of 1657e1780 cm3.
Accordingly, we consider ~1700 cm3 with a 95% CI of 1555e1781 cm3
to be a reasonable ECV estimate for XJY 6.
4. Discussion
4.1. Endocranial volume estimations of incomplete hominin fossils
The evolution of brain size has been a focus of discussion in
paleoanthropology almost since its inception as a scientific field
(e.g., Dart, 1925; Weidenreich, 1936; Tobias, 1963; Jerison, 1970;
Holloway, 1973; Falk, 1991; Leigh, 1992; Aiello and Wood, 1994; De
Miguel and Henneberg, 2001; Lee and Wolpoff, 2003; Rightmire,
2004; Wu et al., 2010; Bruner, 2021). As such, brain size has long

6
X.-J. Wu, C.J. Bae, M. Friess et al. Journal of Human Evolution 163 (2022) 103119

Table 4
Multiple regression results for XJY 6 ECV (cm3) estimates using ectocranial measurements.

Groups r2adj F-value p- standard error XJY 6 ECV Multiple regression equations
value (95.0% CIs)

Early Pleistocene e e e e 992 e

(n ¼ 5)
Middle Pleistocene 0.789 12.740 <0.001 86.0 1644 (1360e1929) ECV ¼ 1438.388 þ 3.366 * XCB þ 5.345 * XFB þ 1.639
(n ¼ 23) * AVH þ 6.801 * BAC þ 3.650 * LASC e 5.295 * ASB þ7.024 * PAC
Late Pleistocene 0.607 8.515 <0.001 124.1 1784 (1529e2039) ECV ¼ 2961.186 þ 5.509 * XCB þ 5.452 * XFB þ 4.771
(n ¼ 35) * AVH - 0.461 * BAC þ6.683 * LASC þ3.610 * ASB þ12.785 * PAC
Recent modern 0.986 286.133 <0.001 19.4 1779 (1718e1840) ECV ¼ 2262.100 þ 6.196 * XCB þ 6.188 * XFB þ 5.044
humans * AVH þ1.280 * BAC þ 7.661 * LASC e 0.237 * ASB þ 5.585 * PAC
(n ¼ 30)
All specimens 0.813 58.224 <0.001 101.0 1669 (1555e1781) ECV ¼ e2217.770 þ 3.267 * XCB þ 6.827 * XFB þ 7.636
(n ¼ 93) * AVH þ 0.516 * BAC þ8.458 * LASC e 0.945* ASB þ6.624 * PAC

Abbreviations: XCB ¼ maximum cranial width, XFB ¼ maximum frontal breadth, AVH ¼ auricular-to-vertex height, BAC ¼ bregmaeasterion chord, LASC ¼ lambdaeasterion
chord, ASB ¼ biasterionic breadth, PAC ¼ parietal chord length.

Table 5
Multiple regression results for XJY 6 ECV (cm3) estimate using endocranial measurements.

Groups r2adj p- F-value Standard XJY 6 ECV Multiple regression equations

value error (95.0% CIs)

Early e e e e 2069 e
Pleistocene
(n ¼ 5)
Middle 0.919 <0.007 18.057 48.0 1578 (1361e1794) ECV ¼ 1470.622 þ 2.843 * EXCB þ 2.430 * EASB þ 7.241 * BOCL þ 2.207
Pleistocene * EPAC þ 0.352 * EBAC þ 1.674 * ELASC þ 8.222 * SCH þ 1.995 * EXFB
(n ¼ 13)
Late Pleistocene 0.957 <0.001 50.721 49.9 1767 (1552e1983) ECV ¼ 2439.914 þ 8.535 * EXCB þ 10.342 * EASB þ 4.247 * BOCL e 3.659
(n ¼ 19) * EPAC þ 6.310 * EBAC e 5.195 * ELASC þ 8.156 * SCH þ 3.913 * EXFB
Recent modern 0.903 <0.001 58.263 46.1 1648 (1556e1740) ECV ¼ 2055.597 þ 2.048 * EXCB e 1.117 * EASB þ 6.469 * BOCL þ 1.189
humans * EPAC þ 4.812 * EBAC þ 4.301 * ELASC þ 4.265 * SCH þ 8.988 * EXFB
(n ¼ 50)
All specimens 0.937 <0.001 159.998 60.6 1719 (1657e1780) ECV ¼ 2233.668 þ 4.304 * EXCB þ 4.124 * EASB þ 4.576 * BOCL þ 0.176
(n ¼ 87) * EPAC þ 6.589 * EBAC þ 2.123 * ELASC þ 4.140 * SCH þ 6.049 * EXFB

Abbreviations: EXCB ¼ maximum endocast width, EASB ¼ endo-biasterionic breadth, BOCL ¼ endo-bregma-to-occipital pole length, EPAC ¼ parietal lobe chord length,
EBAC ¼ endo-bregma-to-asterion length, ELASC ¼ endo-lambda-to-asterion length, SCH ¼ superior cerebral height, EXFB ¼ maximum endocast frontal width.

been considered to be an important indicator of the evolutionary
position of a hominin fossil.
Brain size is the actual size of a brain and is usually expressed by
a measure of weight and volume. For fossils, the actual brain is not
preserved. In this case, brain size, usually referred to as endocranial
capacity, is estimated for fossils by measuring the ECV
(Weidenreich, 1936). Numerous methods have been developed to
calculate ECV, but the vast majority of these approaches have
focused on complete modern human crania (Manjunath, 2002).
Because the shape of the modern human cranium is more rounded/
encephalized with different proportions than in other Homo taxa,
measurements based on modern human crania are dependent on
variables such as bone thickness and neurocranium morphology.
Thus, formulae derived using external measurements of recent
modern human skulls are usually considered unsuitable for direct
calculations of hominin fossil ECVs (Wu and Zhang, 2019). There are
three main methods to calculate the cranial capacity of fossil
hominins: for well-preserved or nearly complete crania, ECV is
usually estimated by 1) water displacement, 2) segmentation of CT
data from the endocasts that are virtually reconstructed manually
or (semi) automatically in 3D (e.g., Weidenreich, 1936; Conroy et al.,
1984; Weber et al., 1998; Falk et al., 2000; Wu et al., 2008; Wu et al.,
2011; Ni et al., 2021), and 3) by linear regression models, the
formulae for which are derived from ectocranial and/or endocranial
measurements (Lestrel, 1975; Wu and Bruner, 2016; Li et al., 2017;
Zhang et al., 2017).
A number of changes in cranial size, cranial thickness, and
general cranial morphological features have appeared in early
hominins from the Pliocene to the Pleistocene. In this regard, ECV
estimations need to consider variables such as taxonomic assign-
ment and chronospatial distribution when selecting appropriate
comparative samples. When taking into account the morphological
differences between Pleistocene and recent modern human crania,
the most reliable way to estimate ECV of a hominin fossil skull is
through internal cranial vault reconstructions using either physical
or 3D virtual endocranial casts (e.g., Weidenreich, 1936; Jerison,
1970; Holloway, 1973, 1981; Begun and Walker, 1993; Zhang et
al., 2017; Wu and Zhang, 2019; Ni et al., 2021). Due to the rarity
of fully intact fossil skulls, paleoanthropologists have continually
sought to develop more reliable methods to reconstruct ancient
cranial size values. Regression equations derived from Pleistocene
and recent human skull measurements have become the most
popular methods for calculating ECVs of ancient hominin fossils.
For example, H. erectus usually has a small, low, and thick skull,
which differs markedly from recent humans. Thus, formulae
derived using recent human skull measurements are unsuitable for
calculating the ECV of H. erectus. Given that Pleistocene early
modern humans and recent modern humans are more similar in
anatomical features, a regression equation derived using recent
modern human skull measurements can be utilized to estimate the
ECVs of Pleistocene early modern humans (Wu and Zhang, 2019).
XJY 6 was present in the late Middle Pleistocene, dating to be-
tween 200 and 160 ka (Li et al., 2014, 2016). As noted earlier, the
cranium of XJY 6 is very wide and low with a notably enlarged
neurocranium. In fact, XJY 6 is quite similar to Xuchang 1 (125e105
ka), in that they share very large, very low, and very wide cranial

7
X.-J. Wu, C.J. Bae, M. Friess et al.
Figure 6. Comparison of some representative hominin fossil endocranial volumes (cm3) by region (Africa, China, Indonesia, Europe, and Israel) and time (ka). The Xujiayao 6, La
Ferrassie, Amud 1, and some Late Pleistocene exceptions are highlighted in bold. Endocranial volumes are from the studies by Holloway et al. (2004), Li et al. (2017), Wu et al. (2019),
Wu and Zhang (2019), and Ni et al. (2021).
vaults. The endocranial proportions of XJY 6 and Xuchang 1 closely
approximate those of Late Middle and Early Late Pleistocene
hominins and contrast with the higher and more rounded skulls of
the recent human samples. Therefore, the more complete Xuchang
1 skull may provide reliable ECV estimates for XJY 6 given that they
appear morphologically similar.
When comparing the multiple regression results from ectocra-
nial and endocranial measurements using the five comparative
groups (Early Pleistocene, Middle Pleistocene, Late Pleistocene,
recent modern humans, and all specimens), it appears that the
regression equations calculated from the small sample sizes are not
reliable for the XJY 6 ECV estimate. For instance, utilizing the Early
Pleistocene dataset (n ¼ 5) yields an estimated ECV based on
ectocranial measurements of 992 cm3, whereas endocranial mea-
surements result in a twofold increase in estimated ECV of 2069
cm3. When comparing the results using the groups of Early Pleis-
tocene, Middle Pleistocene, Late Pleistocene, recent modern
humans, and Pleistocene þ recent modern humans, the larger
pooled sample utilizing the endocranial measurements has a
smaller 95% CI and std. error. The XJY 6 ECV 95% CI of 1555e1781
cm3 overlaps with Xuchang 1, whose 95% CI is 1687e1916 cm3 (Li et
al., 2017).
4.2. Endocranial volume variation by region
It is generally understood that hominin ECVs increase over time
and that Late Pleistocene fossil hominins have the largest ECVs for
the genus, although clear exceptions exist (e.g., H. floresiensis).
8
Journal of Human Evolution 163 (2022) 103119
Among the hominin fossils discovered in China, the Early Pleisto-
cene H. erectus specimen from Lantian has a small ECV <900 cm3
(Woo, 1966; Wu and Zhang, 2019). Nanjing, Zhoukoudian, and
Hexian H. erectus were present in China during the Middle Pleis-
tocene and have ECVs in the range of 780e1225 cm3 (Wu et al.,
2011). The Maba, Dali, Jinniushan, Hualongdong 6, and Harbin
skulls appeared in China during the Late Middle and Early Late
Pleistocene, and their ECVs range between 1150 and 1420 cm3 (Wu
and Poirier, 1995; Wu and Bruner, 2016; Wu et al., 2019; Ni et al.,
2021). These latter specimens are typically classified as mid-
Pleistocene Homo or archaic H. sapiens (and Homo longi in the
case of Harbin) and are considered generally larger in brain size
than Chinese H. erectus (Wu and Poirier, 1995; Bae, 2010; Liu et al.,
2014; Wu et al., 2019). The ECV of Harbin is 1420 cm3, which is close
to the Jinniushan ECV of 1390 cm3.
In Europe, the ECVs of mid-Pleistocene Homo generally
increased, whereas Neanderthals suddenly increased in the Early
Late Pleistocene. With the recent announcement of Homo bodoensis
sp. nov. (Roksandic et al., 2021), it is likely that at least some of
these European fossils will be assigned to this new taxon, while
others will be reassigned to early Neanderthals. The ECV estimates
of mid-Pleistocene Homo from China (Wu and Poirier, 1995; Liu et
al., 2014; Wu and Bruner, 2016; Li et al., 2017; Wu et al., 2019; Ni
et al., 2021) are similar to mid-Pleistocene Homo from Europe:
Hualondong 6 (300 ka) is 1150 cm3; Jinniushan (280 ka) is 1390
cm3; Dali (230 ka) is 1120 cm3; Maba (300e130 ka) is 1300 cm3;
Harbin (300e145 ka) 1420 cm3; with the exception being Xuchang
1 (125e105 ka) at 1800 cm3 and as presented here, XJY 6 (200e160
X.-J. Wu, C.J. Bae, M. Friess et al.
Figure 7. Natural logarithm of endocranial volume (Ln ECV) vs. natural logarithm of geological age (Ln chronology) for XJY 6 and the comparative samples. ECVs are from the studies
by Holloway et al. (2004), Li et al. (2017), and Ni et al. (2021).
ka) at 1700 cm3. Given the large size of XJY 6 and Xuchang 1, these
fossils likely represent large male individuals. However, within any
given population in the Middle and Early Late Pleistocene, the large
ECVs of 1700 and 1800 cm3 should be considered quite unusual
(Leigh, 1992; Lee and Wolpoff, 2003; Holloway et al., 2004). Thus,
XJY 6 from the Late Middle Pleistocene (200e160 ka) currently
provides the earliest evidence of a brain size that falls within the
upper range of Neanderthals and recent humans. Xuchang 1,
discovered at the Lingjing site in Henan Province, has the
comparatively largest ECV of 1800 cm3 in the Early Late Pleistocene,
according to the manually reconstructed endocast (Li et al., 2017).
Xuchang 1 dates to the early Late Pleistocene, a period during
which ECV was at its peak in some Homo species, such as Nean-
derthals and modern humans. The ECVs of Xuchang 1, some Ne-
anderthals (e.g., Amud 1), and as presented here, XJY 6, are at the
high end of the range for fossil hominins across time and space
(Holloway et al., 2004; Li et al., 2017).
Comparisons across time and space indicate that ECV generally
increases over time but that regional variation (Africa, China,
Indonesia, Europe, and Israel) is present (Fig. 6). In Africa, the
general increase was interrupted by the appearance of Homo naledi,
which has a very small ECV of 560 cm3 (Berger et al., 2015).
Hominin fossils have been found in Indonesia that date to the Early,
Middle, and Late Pleistocene. However, all of the Indonesian fossils
have small ECVs: the H. erectus fossils from Sangiran, Trinil, Sam-
bungmacan, Ngawi, and Ngandong have ECVs ranging between 918
and 1251 cm3 (Holloway et al., 2004), which is similar to Chinese H.
erectus. Homo floresiensis (LB 1), known only from the Late Pleis-
tocene, had an ECV of 400 cm3 (Falk et al., 2005), which is much
smaller than H. erectus.
One of the oldest early modern human fossils found in Israel is
from Nesher Ramla, dating to between 140 and 120 ka (Hershkovitz
et al., 2021). Although the skull is not complete, based on the small
size of the parietal bone, it should not have a large ECV. However, by
the advent of the Late Pleistocene, fossil hominins in Israel already
had large ECVs: ECVs for the Skhul hominins range between 1518
9
Journal of Human Evolution 163 (2022) 103119
and 1587 cm3, whereas ECVs for the Qafzeh hominins range be-
tween 1531 and 1569 cm3 (Holloway et al., 2004). The Amud 1 fossil
from the Levant has an ECV of approximately 1740 cm3, which is the
largest among the Neanderthals (Holloway et al., 2004).
4.3. Endocranial volume variation by time period
The expansion of the hominin brain into the modern human
range begins to appear by the advent of the Late Middle Pleistocene
(Holloway et al., 2004; Rightmire, 2004). The estimated cranial
capacities of XJY 6 and Xuchang 1 are well outside the range of H.
erectus (600e1251 cm3) and mid-Pleistocene Homo (1070e1450
cm3), and fall in the upper range of Neanderthals (1065e1740 cm3),
Early modern humans (1090e1880 cm3), and recent humans
(1077e1693 cm3; Holloway et al., 2004; Li et al., 2017). The largest
cranial capacities for mid-Pleistocene Homo are Ehringsdorf H
(1450 cm3) and Reilingen (1440 cm3; Holloway et al., 2004). The
cranial capacity of Neanderthals varies from 1065 cm3 to 1740 cm3,
with the larger cranial capacities including Amud 1 (1740 cm3), La
Ferrassie 1 (1689 cm3), Shanidar 1 (1600 cm3), Shanidar 5 (1550
cm3), and Spy 2 (1553 cm3; Holloway et al., 2004). However, these
Neanderthals (130e40 ka) are much younger than the Xujiayao
hominins.
In general, brain size tends to increase throughout geological
time (Rightmire, 1988; Leigh, 1992; Lee and Wolpoff, 2003). How-
ever, when comparing hominin fossil cranial capacities by region
and time (Fig. 6), it is clear that H. floresiensis, Homo luzonensis, H.
naledi, Xuchang 1, and XJY 6 challenge this trend (Falk et al., 2005;
Spoor et al., 2007; Berger et al., 2015; Detroit et al., 2019).
Furthermore, the change in brain size is different between Africa,
Europe, and Asia as it is clear that a high degree of variation in
Homo exists across time and space. In Africa and western Eurasia,
brain size tends to gradually increase from the Middle to Late
Pleistocene, but in East Asia, a noticeable increase in ECV is present
in XJY 6 and then Xuchang 1 during the Late Middle and Early Late
Pleistocene. Moving further into the Late Pleistocene in China, early
X.-J. Wu, C.J. Bae, M. Friess et al.
human ECVs decrease, as is evident from the Liujiang and Zhou-
koudian Upper Cave fossils (Fig. 6). To what extent this is due to
gaps in the hominin fossil record remains unclear.
Numerous hypotheses have been proposed to explain the
pattern in hominin brain expansion over geological time and why
gaps may exist. These hypotheses include walking upright, tool use,
hunting and meat consumption, high metabolic costs, climate and
environmental change, increasing social complexity, fire control
and cooking, and punctuated equilibrium (Robert, 1961; Holloway,
1966; Jerison, 1973; Gould and Eldgridge, 1977; Leakey and Hay,
1979; Fialkowski, 1986, 1988; Calvin, 1990; Aiello and Wheeler,
1995; Milton, 1999; Leonard et al., 2003; Wrangham and
Carmody, 2010; Gonza lez-Forero and Gardner, 2018). In all likeli-
hood, none of these hypotheses singularly fully explain variation in
brain size, particularly when a late-appearing hominin with a small
brain suddenly appears (e.g., H. floresiensis, H. luzonensis, and H.
naledi; Falk et al., 2005; Spoor et al., 2007; Berger et al., 2015;
Detroit et al., 2019) or in this case, when not-so-late-appearing
hominins with large brains (i.e., XJY 6 and Xuchang 1) are
discovered.
Alternative taxonomic assignments as well as uncertainties with
respect to absolute chronology and endocranial volume (Fig. 7) will
impact this analysis (r2adj here ¼ 0.75, which is moderate accuracy),
but the position of the ECV of XJY 6 together with that of Xuchang 1,
relative to the overall allometric trend within Homo, clearly un-
derscores its large ECV.
5. Conclusions
As Pleistocene hominin crania are rarely found completely
intact, reliability of estimates of their ECV depends primarily on the
overall preservation of the fossils and the methods used to calculate
the estimations. As many earlier hominins exhibit numerous
morphological differences throughout their evolutionary history,
comparative samples for Pleistocene hominin ECV estimations
need to consider, among other things, the geological age of the
specimen. Here, combining the results from ectocranial and
endocranial measurements using Pleistocene and recent modern
human samples, the ECV of XJY 6 is conservatively estimated to
~1700 cm3. XJY 6 is currently the earliest evidence of a hominin
brain size that falls in the upper range of later-appearing Nean-
derthals and modern H. sapiens. This very large cranial capacity, in
combination with a Neanderthal-like temporal labyrinthine
pattern (Wu et al., 2014; Li et al., 2017b), suggests that between
300,000 and 100,000 years ago it may be possible that more than
one type of ancient hominin existed in China. XJY 6 probably
belonged to a new type of previously unreported archaic hominin
group contemporary with other hominin populations. Considering
the current debates surrounding the possible presence of Deniso-
vans in the Chinese fossil record (Stringer and Barnes, 2015; Xing et
al., 2015; Bae et al., 2017; Buzhilova et al., 2017; Gokhman et al.,
2019), it seems likely that the Denisovans had a large ECV as well.
Declaration of competing interest
The authors declare that they have no known competing
financial interests or personal relationships that could have
appeared to influence the work reported in this article.
Acknowledgments
This work was supported by the Strategic Priority Research
Program of Chinese Academy of Sciences (grant number:
10
Journal of Human Evolution 163 (2022) 103119
XDB26000000) and the National Natural Science Foundation of
China (grant number: 41630102, 41872030), and M.F. received
financial support from the De partement Homme et Environne-
ment, MNHN. We are grateful to E. Trinkaus, R. L. Holloway, W.
Zhang, Y.M. Hou, and American Museum of Natural History for their
help in sharing comparative data and/or manuscript preparation.
We would also like to thank the editors and reviewers for their
earlier comments that substantially improved the final version of
this manuscript. We alone are responsible for any errors that may
appear.
Supplementary Online Material
Supplementary online material related to this article can be
found at https://doi.org/10.1016/j.jhevol.2021.103119.
References
Aiello, L., Wood, B., 1994. Cranial variables as predictors of hominine body mass.
Am. J. Phys. Anthropol. 95, 409e426.
Aiello, L.C., Wheeler, P., 1995. The expensive-tissue hypothsis: The brain and the
digestive system in human and primate evolution. Curr. Anthropol. 36,
199e221.
Ao, H., Liu, C.R., Roberts, A.P., Zhang, P., Xu, X., 2017. An updated age for the Xujiayao
hominin from the Nihewan Basin, North China: Implications for middle Pleis-
tocene human evolution in East Asia. J. Hum. Evol. 106, 54e65.
Bae, C.J., 2010. The late Middle Pleistocene hominin fossil record of eastern Asia:
Synthesis and review. Yearbk. Phys. Anthropol. 143, 75e93.
Bae, C.J., Douka, K., Petraglia, M.D., 2017. Human colonization of Asia in the Late
Pleistocene: An introduction to Supplement 17. Curr. Anthropol. 58, 373e382.
Begun, D., Walker, A., 1993. The endocast. In: Walker, A., Leakey, R. (Eds.), The
Nariokotome Homo erectus Skeleton. Harvard University Press, Cambridge,
pp. 326e358.
Berger, L.R., Hawks, J., de Ruiter, D.J., Churchill, S.E., Schmid, P., Delezene, L.K.,
Kivell, T.L., Garvin, H.M., Williams, S.A., DeSilva, J.M., Skinner, M.M.,
Musiba, C.M., Cameron, N., Holliday, T.W., Harcourt-Smith, W., Ackermann, R.R.,
Bastir, M., Bogin, B., Bolter, D., Brophy, J., Cofran, Z.D., Congdon, K.A., Deane, A.S.,
Dembo, M., Drapeau, M., Elliott, M.C., Feuerriegel, E.M., Garcia-Martinez, D.,
Green, D.J., Gurtov, A., Irish, J.D., Kruger, A., Laird, M.F., Marchi, D., Meyer, M.R.,
Nalla, S., Negash, E.W., Orr, C.M., Radovcic, D., Schroeder, L., Scott, J.E.,
Throckmorton, Z., Tocheri, M.W., VanSickle, C., Walker, C.S., Wei, P.P., Zipfel, B.,
2015. Homo naledi, a new species of the genus Homo from the Dinaledi
chamber, South Africa. eLife 4, e24232.
Bruner, E., 2021. Evolving human brains: Paleoneurology and the fate of middle
Pleistocene. J. Archaeol. Method Theor 28, 76e94.
Buzhilova, A., Derevianko, A., Shunkov, M., 2017. The northern dispersal route:
Bioarchaeological data from the late Pleistocene of Altai, Siberia. Curr.
Anthropol. 58, S491eS503.
Calvin, W.H., 1990. The Ascent of Mind: Ice Age Climates and the Evolution of In-
telligence. Bantam, New York.
Chen, T.M., Yuan, S.X., Gao, S.J., 1984. The study on uranium-series dating of fossil
bones as an absolute age sequence for the main Paleolithic sites of North China
(in Chinese with English abstract). Acta Archaeol. Sin. 3, 259e269.
Chia, L.P., Wei, Q., 1976. A Palaeolithic site at Hsü-Chia-Yao in Yangkao county,
Shansi Province (in Chinese with English abstract). Acta Archaeol. Sin. 2,
97e114.
Chia, L.P., Wei, Q., Li, C.R., 1979. Report on the excavation of Hsuchiayao man site in
1976 (in Chinese with English abstract). Vertebr. PalAsiat. 17, 277e293.
Conroy, G.C., Vannier, M.W., 1984. Noninvasive three-dimensional computer im-
aging of matrix-filled fossil skulls by high-resolution computed tomography.
Science 26, 456e458.
Dart, R., 1925. Australopithecus africanus: The man-ape of South Africa. Nature 115,
195e199.
De Miguel, C., Henneberg, M., 2001. Variation in hominid brain size: How much is
due to method? Homo 52, 3e58.
Detroit, F., Mijares, A.S., Corny, J., Daver, G., Zanolli, C., Dizon, E., Robles, E., Grün, R.,
Piper, P.J., 2019. A new species of Homo from the Late Pleistocene of the
Philippines. Nature 568, 181e186.
Falk, D., 1991. 3.5 million years of hominid brain evolution. Semin. Neurosci. 3,
409e416.
Falk, D., Radmond, J.C., Guyer, J., Conroy, C., Recheis, W., Weber, G.W., Seidler, H.,
2000. Early hominid brain evolution: A new look at old endocast. J. Hum. Evol.
38, 695e717.
Falk, D., Hildebolt, C., Smith, K., Morwood, M.J., Sutikna, T., Brown, P., Jatmiko, E.,
Saptomo, W., Brunsden, B., Prio, F., 2005. The brain of LB1, Homo floresiensis.
Science 242, 242e245.
Fialkowski, K.R., 1986. A mechanism for the origin of the human brain: A hypoth-
esis. Curr. Anthropol. 27, 288e290.
X.-J. Wu, C.J. Bae, M. Friess et al.
Fialkowski, K.R., 1988. Origin of human brain as a preadaptation to enhanced
cognitive powers. Anthropol. Anzeiger 46, 317e325.
Gokhman, D., Mishol, N., De Manuel, M., De Juan, D., Shuqrun, J., Meshorer, E.,
Marques-Bonet, T., Rak, Y., Carmel, L., 2019. Reconstructing Denisovan anatomy
using DNA methylation maps. Cell 179, 180e192.
Gonz alez-Forero, M., Gardner, A., 2018. Inference of ecological and social drivers of
human brain-size evolution. Nature 557, 554e557.
Gould, S.J., Eldgridge, N., 1977. Punctuated equilibria: The tempo and mode of
evolution reconsidered. Paleobiology 3, 115e151.
Hershkovitz, I., May, H., Sarig, R., Pokhojaev, A., Grimaud-Herve , D., Bruner, E.,
Fornai, C., Quam, R., Arsuaga, J.L., Krenn, V.A., Martino  n-Torres, M., Bermúdez de
Castro, J.M., Martín-France s, L., Slon, V., Albessard-Ball, L., Vialet, A., Schüler, T.,
Manzi, G., Profico, A., Vincenzo, F.D., Weber, G.W., Zaidner, Y., 2021. A middle
Pleistocene Homo from Nesher Ramla, Israel. Science 372, 1424e1428.
Holloway, R.L., 1966. Cranial capacity and neuron number: A critique and proposal.
Am. J. Phys. Anthropol. 25, 305e314.
Holloway, R.L., 1973. New endocranial values for the East African early hominids.
Nature 243, 97e99.
Holloway, R.L., 1981. Volumetric and asymmetry determination on recent hominid
endocasts: Spy I and Spy II, Djebel Ihroud I, and the Sale  Homo erectus speci-
mens with some notes on Neandertal brain size. Am. J. Phys. Anthropol. 55,
385e393.
Holloway, R.L., Broadfiled, D.C., Yuan, M.S., 2004. The Human Fossil Record. Vol. 3,
Brain Endocasts e The Paleoneurological Evidence. Wiley-Liss, Hoboken.
Jerison, H.J., 1970. Gross brain indices and the analysis of fossil endocasts. In:
Noback, G.R., Montagna, W. (Eds.), The Primate Brain. Meredith Corporation,
New York, pp. 225e244.
Jerison, H.J., 1973. Evolution of the Brain and Intelligence. Academic Press, New
York.
Leakey, M.D., Hay, R.L., 1979. Pliocene footprints in the Laetolil beds at Laetoli,
northern Tanzania. Nature 278, 317e323.
Lee, S.H., Wolpoff, M.H., 2003. The pattern of evolution in Pleistocene human brain
size. Paleobiology 29, 186e196.
Leigh, S.R., 1992. Cranial capacity evolution in Homo erectus and Homo sapiens. Am.
J. Phys. Anthropol. 87, 1e13.
Lestrel, P.E., 1975. Hominid brain size versus time: Revised regression estimates.
J. Hum. Evol. 5, 207e212.
Leonard, W.R., Robertson, M.L., Snodgrass, J.J., Kuzawa, C.W., 2003. Metabolic cor-
relates of hominid brain evolution. Comp. Biochem. Physiol., A 136, 5e15.
Li, M.Y., Zhang, S.R., Xu, Q.H., Xie, F., Wang, F.G., 2016. New recognition of Houjiayao
relic site: Stratum age and environment (in Chinese with English abstract). Acta
Palaeontol. Sin. 55, 122e135.
Li, Z., Xu, Q., Zhang, S., Hun, L., Li, M., Xie, F., Liu, L., 2014. Study on stratigraphic age,
climate changes and environment background of Houjiayao Site in Nihewan
Basin. Quat. Int. 349, 42e48.
Li, Z.Y., Wu, X.J., Zhou, L.P., Liu, W., Gao, X., Trinkaus, E., 2017. Late Pleistocene
archaic human crania from Xuchang, China. Science 355, 969e972.
Liu, W., Wu, X.J., Xing, S., Zhang, Y.Y., 2014. Human Fossils in China (in Chinese).
Science Press, Beijing.
Manjunath, K.Y., 2002. Estimation of cranial volumeean overview of methodolo-
gies. J. Anat. Soc. India 51, 85e91.
Milton, K., 1999. A hypothesis to explain the role of meat-eating in human evolu-
tion. Evol. Anthropol. 8, 11e21.
Ni, X.J., Ji, Q., Wu, W.S., Shao, Q.F., Ji, Y., Zhang, C., Liang, L., Ge, J.Y., Guo, Z., Li, J.H.,
Li, Q., Grün, R., Stringer, C., 2021. Massive cranium from Harbin establishes a
new Middle Pleistocene human lineage in China. Innovation 2, 100130.
Norton, C.J., Gao, X., 2008. Hominin-carnivore interactions during the Chinese early
Paleolithic: Taphonomic perspectives from Xujiayao. J. Hum. Evol. 55, 164e178.
Robert, A., 1961. African Genesis. Dell Publishing Co., New York.
Rightmire, G.P., 1988. Homo erectus and later middle Pleistocene humans. Annu. Rev.
Anthropol. 17, 239e259.
Rightmire, G., 2004. Brain size and encephalization in early to middle Pleistocene
Homo. Am. J. Phys. Anthropol. 124, 109e123.
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Journal of Human Evolution 163 (2022) 103119
Roksandic, M., Radovic, P., Wu, X.J., Bae, C.J., 2021. Resolving the “muddle in the
middle”: The case for Homo bodoensis sp. nov. Evol. Anthropol. https://doi.org/
10.1002/evan.21929.
Spoor, F., Leakey, M.G., Gathogo, P.N., Brown, F.H., Anto n, S., McDougall, C.I.,
Kiarie, C., Manthi, F.K., Leakey, L.N., 2007. Implications of new early Homo fossils
from Ileret, east of Lake Turkana, Kenya. Nature 488, 688e691.
Stringer, C.B., Barnes, I., 2015. Deciphering the Denisovans. Proc. Natl. Acad. Sci. USA
112, 15542e15543.
Tobias, P.V., 1963. Cranial capacity of Zinjanthropus and other australopithecines.
Nature 23, 743e746.
Trinkaus, E., Wu, X.J., 2017. External auditory exostoses in the Xuchang and Xujiayao
human remains: Patterns and implications among eastern Eurasian Middle and
Late Pleistocene crania. PLoS One 12, e0189390.
Wang, F.G., Li, F., 2020. Discussions on stratigraphy and age of the Xujiayao hominin.
Acta Anthropol. Sin. 39, 16e172.
Weber, G., Recheis, W., Seidler, H., Scholze, T., 1998. ‘Virtual anthro-
pology’dmethodological aspects of measurements. Am. J. Phys. Anthropol.
Suppl. 26, 228e229.
Weidenreich, D.F., 1936. Observations on the form and proportions of the endo-
cranial casts of Sinanthropus pekinensis, other hominids and the great apes: A
comparative study of brain size. Palaeontol. Sin. 7, 1e50.
Woo, J.K., 1966. The skull of Lantian man. Curr. Anthropol. 7, 83e86.
Wrangham, R., Carmody, R., 2010. Human adaptation to the control of fire. Evol.
Anthropol. 19, 187e199.
Wu, M., 1980. Human fossils discovered at Xujiayao site in 1977 (in Chinese with
English abstract). Vertebr. PalAsiat. 18, 227e238.
Wu, M., 1986. Study of temporal bone of Xujiayao man (in Chinese with English
abstract). Acta Anthropol. Sin. 5, 220e226.
Wu, X.J., Trinkaus, E., 2014. The Xujiayao 14 mandibular ramus and Pleistocene
Homo mandibular variation. C. R. Palevol 13, 333e341.
Wu, X.J., Trinkaus, E., 2015. Neurocranial trauma in the late archaic human remains
from Xujiayao, Northern China. Int. J. Osteoarchaeol. 25, 245e252.
Wu, X.J., Bruner, E., 2016. The endocranial anatomy of Maba 1. Am. J. Phys.
Anthropol. 160, 633e643.
Wu, X.J., Zhang, W., 2019. Methods for estimating cranial capacity from Chinese
human fossils. Acta Anthropol. Sin. 38, 767e778.
Wu, X.J., Liu, W., Dong, W., Qu, J., Wang, Y., 2008. The brain morphology of Homo
Liujiang cranium fossil by three-dimensional computed tomography. Chin. Sci.
Bull. 53, 2513e2519.
Wu, X.J., Schepartz, L., Norton, C.J., 2010. Morphological and morphometric analysis of
variation in the Zhoukoudian Homo erectus brain endocasts. Quant. Int. 211, 4e13.
Wu, X.J., Holloway, R.L., Schepartz, L., Xing, S., 2011. The brain endocast of Nanjing 1,
Homo erectus. Am. J. Phys. Anthropol. 45, 452e460.
Wu, X.J., Maddux, S.D., Pan, L., Trinkaus, E., 2012. Nasal floor variation among
eastern Eurasian Pleistocene Homo. Anthropol. Sci. 120, 217e226.
Wu, X.J., Xing, S., Trinkaus, E., 2013. An enlarged parietal foramen in the late archaic
XJY 11 neurocranium from Northern China. PLoS One 8, e59587.
Wu, X.J., Crevecoeur, I., Liu, W., Trinkaus, E., 2014. The temporal labyrinths
of eastern Eurasian Pleistocene humans. Proc. Natl. Acad. Sci. USA 111,
10509e10513.
Wu, X.J., Pei, S.W., Cai, Y.J., Tong, H.W., Li, Q., Dong, Z., Sheng, J.C., Jin, Z.T., Ma, D.D.,
Xing, S., Li, X.L., Cheng, X., Cheng, H., De la Torre, I., Edwards, R.L., Gong, X.C.,
An, Z.S., Trinkaus, E., Liu, W., 2019. Archaic human remains from Hualongdong,
China, and Middle Pleistocene human continuity and variation. Proc. Natl. Acad.
Sci. USA 116, 9820e9824.
Wu, X.Z., Poirier, F.E., 1995. Human Evolution in China. Oxford University Press,
Oxford.
Xing, S., Martino  n-Torres, M., Bermúdez de Castro, J.M., Wu, X.J., Liu, W., 2015.
Hominin teeth from the early Late Pleistocene site of Xujiayao, northern China.
Am. J. Phys. Anthropol. 156, 224e240.
Zhang, Y.M., Wu, X.J., Schepartz, L., 2017. Comparing methods for estimating cranial
capacity in incomplete human fossils using the Jingchuan 1 partial cranium as
an example. Quat. Int. 434, 57e64.