Juvenile hominoid cranium from the late Miocene of
southern China and hominoid diversity in Asia
Jay Kelleya,1 and Feng Gaob
a
Institute of Human Origins and School of Human Evolution and Social Change, Arizona State University, Tempe, AZ 85287; and bDepartment of
Paleoanthropology, Yunnan Institute of Cultural Relics and Archaeology, Kunming, Yunnan 650118, China
Edited by Erik Trinkaus, Washington University, St. Louis, MO, and approved March 23, 2012 (received for review January 24, 2012)
The fossil ape Lufengpithecus is known from a number of late
Miocene sites in Yunnan Province in southern China. Along with
other fossil apes from South and Southeast Asia, it is widely con-
sidered to be a relative of the extant orangutan, Pongo pygmaeus.
It is best represented at the type site of Shihuiba (Lufeng) by
several partial to nearly complete but badly crushed adult crania.
There is, however, an additional, minimally distorted cranium of a
young juvenile from a nearly contemporaneous site in the Yuan-
mou Basin, which affords the opportunity to better assess the rela-
tionships between Lufengpithecus and Pongo. Comparison with
similarly aged juvenile skulls of extant great apes reveals no fea-
tures suggesting clear affinities to orangutans, and instead reveals
a morphological pattern largely consistent with a stem member of
the hominid (great ape and human) clade. The existence at this
time of other hominids in South Asia (Sivapithecus) and Southeast
Asia (Khoratpithecus) with clear craniofacial affinities to Pongo
suggests both more diversity among Asian Late Miocene apes
and more complex patterns of dispersal than previously supposed.
Major differences in the associated mammal faunas from the south-
ern China sites and those from South and Southeast Asia are con-
sistent with these findings and suggest more than one dispersal
route of apes into East Asia earlier in the Miocene.
T he hominoid Lufengpithecus from the late Miocene of
southern China is known mostly from hundreds of isolated
teeth, principally from the sites of Shihuiba (Lufengpithecus
lufengensis; ca. 6–7 Ma) (1, 2) and several somewhat older sites in
the Yuanmou Basin (Lufengpithecus hudienensis; ca. 7–8 Ma) (2–
4). The molar teeth are strikingly similar to the highly distinctive
teeth of the modern orangutan, with intricately wrinkled enamel
over relatively flattened occlusal basins. There are also several
badly crushed crania from Shihuiba (5–7). Restoration has been
attempted for one of these (8), but some of the morphology
remains ambiguous or is clearly still distorted. The condition of
these crania has allowed for different interpretations of their
morphology and, consequently, different opinions regarding the
phylogenetic position of Lufengpithecus (4, 8–11). Nevertheless,
most recent analyses propose a relationship to the Pongo clade,
as either its nearest relative or as a stem member of the clade
(12–17).
The cranium of L. hudienensis from Yuanmou, YV0999, pre-
serves nearly the entire facial skeleton, the palate with partial
dentition, and most of the frontal bone. In contrast to the Shihuiba
crania, it is remarkably well preserved and minimally distorted,
with the right side being almost completely free of distortion (Fig.
1, Fig. S1, and SI Text) (18, 19). Because of its preservation, this
cranium can be more informative about the phylogenetic rela-
tionships of Lufengpithecus than are the damaged adult crania
from Shihuiba. However, its juvenile status (first molar just coming
into occlusion) introduces an additional challenge for interpre-
tation because there are no other relatively complete juvenile
crania, and even few juvenile cranial fragments, known for other
Miocene anthropoids. Additionally, little has been done to identify
the developmental stages at which the derived features (apomor-
phies) characteristic of adult great ape crania first become evident.
www.pnas.org/cgi/doi/10.1073/pnas.1201330109
Although YV0999 was recovered in 1988 (18) and has been the
subject of a preliminary analysis (20), it has not been analyzed
within an adequate comparative framework of similarly aged
juveniles of extant great apes. Therefore, a database was assem-
bled of measurements and discrete, nonmetric characters from
extant great ape individuals, representing Gorilla gorilla, Pan
troglodytes and Pongo pygmaeus (each species sample comprising
a single subspecies), at approximately the same stage of dental
development as YV0999. The measurements reflect general shape
and proportional relationships of the facial skeleton and were
evaluated using both cluster analysis and discriminant analysis
(Materials and Methods and SI Text) to determine the overall
ANTHROPOLOGY
phenetic resemblance of YV0999 with respect to the extant great
ape crania. The nonmetric characters reflect consensus or near
consensus cranial apomorphies of each of the extant great apes
(21) that were found to be at least incipiently expressed at a stage
of development equivalent to that of YV0999. YV0999 was scored
for these characters, which were evaluated individually rather
than by formal, computational cladistic analysis.
Results
Results of the cluster analysis are shown in Fig. 2. YV0999 groups
as an outlier to the majority of the Pan crania. Interestingly, Pan
clusters with Pongo rather than with its sister taxon, Gorilla,
highlighting the distinctiveness of the proportions and overall
shape of the gorilla facial skeleton at this stage of development.
Although most of the great ape crania group with their con-
specifics, there is a fourth, taxonomically heterogeneous cluster
that includes crania of all three taxa, but primarily Pongo, and that
is linked to the Pan plus Pongo cluster (Fig. 2). This cluster reveals
that possession by some individuals of certain metric character-
istics that are more typical of one of the other genera was enough
to create a separate grouping of these somewhat anomalous
individuals. It also highlights the fact that, although the juvenile
crania of each genus largely express their genus-specific metric
characteristics, there is greater similarity in the shape of the facial
skeleton among the three great ape genera at the early juvenile
stage than as adults.
The discriminant analysis was more definitive. The results are
highly significant (Wilk’s lambda P value, <0.001), and all 30
extant great ape crania were correctly classified. YV0999 was
entered as an unknown and did not, therefore, contribute to the
computation. Among the three extant great ape group means,
YV0999 is closest to Pan (Mahalanobis D2 values to the group
means of, respectively, Pan, Gorilla, and Pongo: 73.26, 119.13,
and 155.64), and in a forced classification, it was grouped with
Pan (P > 0.999).
Author contributions: J.K. designed research; J.K. performed research; J.K. and F.G. ana-
lyzed data; and J.K. wrote the paper.
The authors declare no conflict of interest.
This article is a PNAS Direct Submission.
1
To whom correspondence should be addressed. E-mail: jkelley.iho@asu.edu.
This article contains supporting information online at www.pnas.org/lookup/suppl/doi:10.
1073/pnas.1201330109/-/DCSupplemental.
PNAS Early Edition | 1 of 4
 S2), minimal offset from the nasoalveolar clivus to the palate in-
tranasally with a highly constricted incisive canal, a continuously
concave midfacial profile, and lack of a frontal sinus (21, 22). In
these, YV0999 is decidedly unlike Pongo and more closely resem-
bles Pan and Gorilla, especially the former, which both express
variants of the ancestral hominid condition for these features (21).
In most features of facial morphology, YV0999 most closely
resembles Pan, arguably the least derived of the extant great apes
in terms of cranial morphology. The principal differences are
in the relatively large orbits of YV0999 and their influence on
widening the upper face and the absence of the most clearly de-
rived features of Pan, a supraorbital torus and sulcus and a sharply
posteriorly inflected malar region. YV0999 also has a shorter,
flatter, and more steeply inclined premaxilla, a premaxillary su-
ture that is completely fused over its entire extent (in none of the
extant ape crania is this suture completely fused at this stage of
development, and it is mostly fused in only five of the 30 indi-
viduals in the sample), and two distinct, large incisive foramina
palatally, indicating a fully partitioned incisive canal. The incisive
Fig. 1. Oblique frontal view (Left) and lateral view (Right) of YV0999, canal is at most only partly partitioned at this stage of de-
Lufengpithecus hudienensis. Note the missing bone at the inferior nasal velopment in the African ape crania, whereas it is unpartitioned in
margin in the frontal view. In the lateral view, the actual facial contour is
Pongo and remains so in adulthood.
somewhat obscured by the broken and everted left nasal margin, but the
slight anterior projection of the inferior nasal bones, resulting in a biconcave
Discussion
facial profile, is nonetheless evident. (Scale bar: 1 cm.)
Assuming that YV0999 is representative of cranial morphology
in other Lufengpithecus species (SI Text), we consider it most
Despite the small, taxonomically heterogeneous group in the probable that Lufengpithecus represents a stem hominid lineage
cluster analysis, the results from the cluster and discriminant that also expresses a small suite of uniquely derived features (4,
analyses together affirm the distinctiveness of facial form among 9, 23). It is also possible given the general resemblance to Pan,
the three extant great apes at this early stage of development and exclusive of the derived features of extant African apes noted in
demonstrate the resemblance of YV0999 to Pan rather than Table 1, that Lufengpithecus represents a stem member of the
to Pongo. African ape and human clade (20) (discussed further below).
Results of the discrete character analysis are shown in Table 1. Although the absence of Pongo apomorphies in YV0999 does
Many of the most distinctive apomorphic features of extant great not absolutely preclude Lufengpithecus being a member of the
ape crania are clearly expressed at the developmental stage rep- Pongo clade, these absences take on added significance in light of
resented by incipient first molar occlusion. For none of these does the fact that most of the key derived facial skeletal features of
YV0999 show the presumed derived condition, regardless of the Pongo are expressed in the nearly contemporaneous Sivapithecus
extant ape that expresses the particular apomorphy. It instead sivalensis from the late Miocene Siwaliks of South Asia (13, 22),
expresses a generally plesiomorphic morphology that would be whereas at least one is expressed in Khoratpithecus piriyai from
expected in the common ancestor of the hominid (great ape and the late Miocene site of Khorat in Thailand (24, 25) and K.
human) clade. Most important, it shares none of the consensus ayeyarwadyensis from the Irrawaddy Formation in Myanmar (26).
derived features of Pongo. Six of these in particular are highly Sivapithecus shares all of the derived features of Pongo listed in
distinctive among apes, both fossil and extant: superiorly-inferiorly Table 1. Mandibles of K. piriyai and K. ayeyarwadyensis show no
elongate orbits, a relatively very narrow interorbital region (Fig. evidence of attachment of anterior digastric muscles, an absence
otherwise recorded only in Pongo among both fossil and extant
apes. In addition, the pattern of mandibular postcanine root
morphology in both Khoratpithecus species is most similar to that
Gorilla
Gorilla of Pongo among extant great apes (27). The presence of key de-
Gorilla
Gorilla rived features of Pongo in these other, contemporary South Asian
Gorilla
Gorilla genera reinforces the argument that the absence of these features
Gorilla
Gorilla in Lufengpithecus indicates exclusion from the Pongo clade.
Gorilla
Pan A phylogenetic link between Sivapithecus and Khoratpithecus
Pongo
Pongo to the exclusion of Lufengpithecus is also supported by the
Pongo
Pongo
mammalian faunas from the different fossil sites. There are
Pan
Pan
major differences at the family level between the large mammal
Pan
Pan
faunas from southern China on the one hand and those from
Pan
Pan
South (Siwaliks) and Southeast Asia (Khorat, Irrawaddy For-
Pan
Pan
mation) on the other (Table 2). Certain families that are com-
Pan
Yuanmou
mon or dominant elements in the southern China faunas, some
Pongo shared with faunas from northern China, are absent from the
Pongo
Pongo faunas of South and Southeast Asia. Likewise, certain families
Pongo
Pongo that are common in the South and Southeast Asian faunas are
Pongo
Pongo absent from those of southern China. There are further differ-
ences in the composition of these faunas. Bovids, for example,
0.0 0.1 0.2 0.3 0.4 0.5
Distances
are both common and diverse in the South/Southeast Asian
faunas but are either rare/absent (Yuanmou) or uncommon and
Fig. 2. Cluster tree of extant great ape juvenile crania and YV0999 based on with low diversity (Shihuiba) in the faunas from southern China.
craniofacial measurements (Materials and Methods and SI Text). The faunas from South and Southeast Asia also have in common

2 of 4 | www.pnas.org/cgi/doi/10.1073/pnas.1201330109 Kelley and Gao

Table 1. Discrete craniofacial features in extant great apes and YV0999
Morphology Derived condition and taxon in which it is expressed YV0999 resembles

Frontal squama orientation More horizontally angled back from supraorbital area (Pan/Gorilla) Pongo
Supraorbital topography Torus and sulcus (Pan/Gorilla); thin supraorbital costae (Pongo) Neither
Orbital shape Superiorly-inferiorly elongate/ovoid (Pongo) Pan/Gorilla
Interorbital septum Relatively narrow in relation to biorbital breadth (Pongo) Pan/Gorilla
Infraorbital foramina Positioned well medial to zygomatico-maxillary suture at orbit (Pongo) Pan/Gorilla
Malar orientation Gradually posteriorly convex (Pan/Gorilla) Pongo
Nasoalveolar clivus Strongly convex antero-posteriorly and highly prognathic (Pongo) Pan
Subnasal topography Minimal offset from nasoalveolar clivus to nasal floor (Pongo) Pan/Gorilla
Incisive canal Highly restricted (Pongo) Pan/Gorilla
Midfacial profile Continuously concave from glabella to nasospinale (Pongo) Pan
Frontal sinus Absent (Pongo) Pan/Gorilla

several species of deinotheres, anthracotheres, rhinoceroses, and
suids, whereas there are no ungulate species in common between
the South/Southeast Asian faunas and those from southern China
(25, 26, 28–30). Although these faunal patterns do not bear di-
rectly upon the question of whether Lufengpithecus is a member
of the Pongo clade, they do underscore that the fossil apes
showing clear links to the Pongo clade occur in faunal contexts
and East Asia, there was greater suprageneric taxonomic di-
versity among apes than recognized in most recent taxonomies.
Materials and Methods
A database was assembled of metric and nonmetric characters for extant
great apes of approximately the same stage of dental development as
YV0999. The database included 10 specimens of Pan troglodytes troglodytes,

that are strikingly different from those at Yuanmou or Shihuiba.
The pattern of faunal differences between southern China
and South/Southeast Asia further suggests the possibility of dif-
ferent dispersal routes of apes into East Asia earlier in the Mio-
cene: one route through the South Asian subcontinent and thence
into Southeast Asia (members of the Pongo lineage) and a more
northern route between Europe and southern China through west
central China before uplift of the northern Tibetan Plateau.
Various lines of evidence indicate that major uplift of the northern
regions of the plateau occurred substantially later than in the
south, which would have led to a more prolonged period of
equable climatic conditions conducive to faunal dispersal through
central Asia, at least into the early late Miocene (31–35).
In light of these possibilities, comparisons between Lufengpi-
thecus and European Miocene apes, particularly Rudapithecus
from eastern Europe, a possible member of the African ape/
human clade (16, 17), and between their respective faunas, might
prove informative. They could help resolve the question of
whether Lufengpithecus is a stem hominid or a member of the
African ape/human clade. Preliminary dental comparisons be-
tween Lufengpithecus and Rudapithecus have revealed a number
of similarities (36). For example, Lufengpithecus shares with
Rudapithecus, and with most other western European Miocene
apes, very slender lower canines and a distinctive upper central
incisor morphology with a relatively narrow crown and a prom-
inent lingual pillar (1, 15, 17, 21, 36, 37). Although the phylo-
genetic affinities of Lufengpithecus are not yet certain, its
morphology demonstrates that during the late Miocene in South
ANTHROPOLOGY
9 specimens of Gorilla gorilla gorilla (both samples from the Hamman–Todd
collection at the Cleveland Museum of Natural History), and 11 specimens of
Pongo pygmaeus pygmaeus (2 from the Hamman–Todd collection and 9
from the Zoologische Staatssammlung, München, Germany).
Measurements were selected based on the preservation of YV0999. For
YV0999, unilateral measurements were taken on the right side or in the
midline. Some bilateral measurements were calculated by doubling the right-
side measurement. Raw measurements from the great ape crania and YV0999
were converted to Mossiman shape variables (38) to control for size differ-
ences among the different taxa. This involves calculating the geometric
mean (GM) for each specimen and dividing the raw measurements by the
geometric mean. Variables were first log-transformed and the shape varia-
bles calculated as log(y) − log(GM).
Eighteen measurements were used in both the cluster and discriminant
analyses of the crania to explore overall resemblance in cranial shape irre-
spective of cladistic criteria (SI Text). Both were carried out using SYSTAT
10.0. The cluster analysis incorporated the Join function. The distance metric
in this procedure is Euclidean distance. A single bivariate analysis of orbital
shape and relative interorbital distance was also carried out (Fig. S2) because
of the clear apomorphic condition of these features in extant Pongo. Rela-
tive interorbital distance equals the minimum interorbital width divided by
the maximum distance in the orbital plane from the midnasal suture to the
lateral margin of the zygomatico-frontal orbital pillar unilaterally. Orbital
shape equals maximum orbital height divided by maximum orbital width.
Relative interorbital distance and orbital shape were both calculated using
unscaled raw measurements.
ACKNOWLEDGMENTS. We thank Dr. Richard Kraft, Director of the Zoologi-
sche Staatssammlung, München; Dr. Yohannes Haile-Selassie, Curator of
Physical Anthropology; and Lyman Jellema, collections manager at the
Cleveland Museum of Natural History for allowing us to examine specimens
housed at their institutions. We also thank Liu Xu, Director, and Yang

Table 2. Faunal comparison between late Miocene hominoid sites in southern China (Yuanmou, Shihuiba), South
Asia (Siwaliks), and Southeast Asia (Khorat, Irrawaddy Formation)
Yuanmou Shihuiba Siwaliks Khorat Irrawaddy Formation

Ursidae Present Present Absent Absent Absent

Deinotheriidae Absent Absent Present Present Absent
Tapiridae Present Present Absent Absent Absent
Suidae Present Present Same species Same species Same species
Anthracotheriidae Absent Absent Present Present Present
Tragulidae Present Present Abundant, diverse ? Present
Giraffidae Absent Absent Present Present Present
Bovidae Absent or rare Low diversity Abundant, diverse Abundant Diverse
Cervidae Abundant, diverse Abundant, diverse Absent Absent Absent

Data are from Refs. 25, 26, 28–30.

Kelley and Gao PNAS Early Edition | 3 of 4

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