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Huanglong Cave: A Late Pleistocene human fossil site in Hubei Province, China
Wu Liu
a
,
*
, Xianzhu Wu
b
,
c
, Shuwen Pei
a
, Xiujie Wu
a
, Christopher J. Norton
d
a
Key Laboratory of Evolutionary Systematics of Vertebrates, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, Beijing 100044, China
b
Institute of Archeology and Cultural Relics of Hubei Province, Wuhan 430077, China
c
Chongqing Normal University, Chongqing 400047, China
d
Department of Anthropology, University of Hawaii, Manoa, USA
article info
Article history:
Available online 30 June 2009
abstract
This contribution discusses recent paleoanthropological findings from Huanglong Cave, a Late Pleisto-
cene human fossil site from Yunxi County, Hubei Province, China. Three excavations in the Huanglong
Cave from 2004 to 2006 yielded seven human teeth, some stone and bone tools, possible burnt sediment
and other evidence possibly related to hominin activities. Based on the presence of extinct faunas (20% of
total taxa identified), the deposits dated to the Late Pleistocene. Electron spin resonance (ESR) and
uranium-series (U-series) dating analyses on associated teeth and speleothems have resulted in diver-
gent chronometric ages (ESR: 44–34 ka; U-series: 103–79 ka). Analysis indicates: (1) most of the
morphological and metric features of the human teeth from Huanglong Cave fall within the range of
variation of modern Chinese, but a few characters may still link them to more archaic hominins; (2) some
activity-induced abrasion and other tooth use-marks were identified, including pronounced tooth
chipping and interproximal grooves on the anterior teeth; (3) the sample of blackened deposit has a high
carbon content (over 70%), experienced high temperatures, and likely was of cultural origin and not
natural; (4) the mammal fossils represent the ‘‘Ailuropoda-Stegodon’’ faunal unit which lived in southern
China throughout the Pleistocene. Synthesizing all of these findings, especially the human teeth that
display modern human characteristics, Huanglong Cave will offer some new insights into various issues
currently being debated in Late Pleistocene human evolutionary research.
Ó2009 Elsevier Ltd and INQUA. All rights reserved.
1. Introduction
Based on a series of new fossil discoveries and an improved
dating framework, the earliest anatomically modern humans or
early modern humans currently appear in East Africa (White et al.,
2003; McDougall et al., 2005; Grine et al., 2007). In contrast, to the
African record, the nature of early modern humans in East Asia is
more debated (Shen, 2004; Shang et al., 2007). Currently, Late
Pleistocene hominin fossils have been found in more than 40 sites
in China (Wu and Poirier,1995; Wu and Wu,1999)(Fig.1). However,
most of the fossils are very fragmentary, hindering detailed
morphological studies. In addition, most of the Late Pleistocene
hominin fossils found in China date to within 30 ka (Wu and Poirier,
1995)(Table 1). A recent discovery and related research show that
the mandible and postcrania from Tianyuan Cave near Zhou-
koudian, China, which have direct AMS
14
C dates of w35 ka may
represent the oldest modern human fossils from mainland East Asia
(Tong et al., 2004; Shang et al., 2007). Nonetheless, there have
been claims that the earliest modern humans in China can be
traced back to 50 ka or even earlier (e.g., Shen et al., 2002, 2005;
Shen, 2004). However, either because of stratigraphic uncer-
tainties or lack of reliable dating, all of the evidence which support
such claims is weak (Wu and Poirier, 1995). Two key barriers in
China have been the lack of accurately dated hominin fossils from
between 100 ka and 50 ka (Shang et al., 2007) and the paucity of
more detailed studies of the morphological variation in Chinese
Late Pleistocene hominins. With this background in mind, more
fossil discoveries of early modern humans from China are required
to explore various issues related to the emergence and spread of
modern humans in East Asia. This paper reports a newly discov-
ered Late Pleistocene human fossil site from central China:
Huanglong Cave.
2. Background
Huanglong Cave (Huanglongdong) is located in Yunxi County,
Hubei Province, China (33
07
0
62.8
00
N; 110
13
0
04.3
00
E) (Fig. 2). The
huge cave has a northeast facing opening and is 598 m above sea
*Corresponding author. Tel.: þ86 10 88369120; fax: þ86 10 68337001.
E-mail address: liuwu@ivpp.ac.cn (W. Liu).
Contents lists available at ScienceDirect
Quaternary International
journal homepage: www.elsevier.com/locate/quaint
1040-6182/$ – see front matter Ó2009 Elsevier Ltd and INQUA. All rights reserved.
doi:10.1016/j.quaint.2009.06.017
Quaternary International 211 (2010) 29–41
level. In May 2004, during a field survey conducted as part of local
highway construction, some mammal fossils were discovered in the
cave. Three archaeological excavations were conducted in 2004,
2005 and 2006 by the Hubei Province Institute of Archaeology and
the Institute of Vertebrate Paleontology and Paleoanthropology.
The three excavations resulted in the discovery of seven human
teeth, some stone and bone artifacts, and evidence of fire use (Wu
et al., 2006, 2007a; Pei et al., 2008; Wang et al., 2008; Liu et al.,
2009a,b).
The main axis of the cave is in a northeast–southwest direction
with its entrance width of 27.8 m and height of 11 m. About 50 m in
front of the site is the Dashui River, a branch of the Hanjiang River.
The entrance of the cave is about 7.8 m above the current river level.
Huanglong Cave extends more than 400 m (the end has not been
reached). Within the cave, five regions were mapped, and four of
the regions were excavated (see Fig 3). The excavations in Region 1,
Region 2, Region 4, and Entrance were of varying sizes. Excavations
in the four regions indicate that the stratigraphy is similar
Fig. 1. The late Pleistocene hominin fossil sites in China.
Table 1
The main late Pleistocene hominin fossil and their chronological dates in China.
Sites Province Main specimens Chronological Dates or
Geological Epoch
References
Upper Cave Beijing 3 crania and bone fragments 34–29 ka (AMS) Wu and Poirier, 1995
Liujiang Gaungxi 1 cranium and 4 bone fragments 67 ka? (U) Wu and Poirier, 1995
Ziyang Sichuan 1 cranium 30 ka (
14
C) Wu and Poirier, 1995
Tianyuan Cave Beijing 1 mandible, several limb bones, both scapulae and
some hand and foot bones
42–39 ka (
14
C) Tong et al., 2004; Shang et al., 2007
Chuandong Guizhou 2 skulls Late Pleistocene Wu and Poirier, 1995
Jianping Liaoning 1 humerus Late Pleistocene Wu and Poirier, 1995
Lijiang Yunnan A cranium Late Pleistocene Wu and Poirier, 1995
Miaohoushan Liaoning 2 parietal fragments and 1 radius 28 ka (
14
C) Wu and Poirier, 1995
Salawusu Inner
Mongolia
1 parietal and 2 frontal fragments, 1 child mandible, 2 femurs,
1 tibia and 1 incisor
50–37 ka (U) 35.3–19 ka
(
14
C)
Wu and Poirier, 1995
Huanglong Gansu 1 skull cap preserving most frontal and anterior part
of parietal bones
Late Pleistocene Wu and Poirier, 1995
Jingchuan Gansu A cranium without facial and basal portion 48–15 ka (OSL) Wu and Poirier, 1995; Liu et al., in press
Shiyu Shanxi A occipital fragment 28.9 (
14
C) Wu and Poirier, 1995
Wushan(Yuanyang) Gansu A skull cap including a complete frontal bone,
two parietals and parts of nasal and sphenoid bones
38.4 (
14
C) Wu and Poirier, 1995
Laibin Guangxi A cranial base Late Pleistocene Wu and Poirier, 1995
Mengzi Yunnan 4 fragmentary crania Late Pleistocene? Wu and Poirier, 1995
Huanglong Cave Hubei 7 teeth 103–44 ka (U and ESR) Wu et al., 2006
W. Liu et al. / Quaternary International 211 (2010) 29–4130
throughout the surveyed areas with five stratigraphic layers iden-
tified (Fig. 4):
(1) Capping flowstone layer (2–28 cm): Milk-white or brown-
yellow layer widely developed in the cave and most of the layer
has a pure and dense structure.
(2) Silt clay–sandy gravel layer (1–201 cm): Silty clay and sandy
gravel are in the same layer but in different levels the deposit of
the layer near the entrance comprises mainly gravels bedded in
a clay matrix. The gravels decrease in both density and size
gradually when moving toward the interior of the cave. In the
deeper areas of the cave the deposit becomes silty clay.
(3) Red silt clay layer (58–101 cm): Two thin flowstone layers
developed with a break. Below the flowstone layers there are
isolated stalagmites which have no direct connection with the
flowstone layer.
(4) Mantle limestone layer (41–66 cm): Loose layer mainly
composed of mantled limestone breccia and clay matrix
residue.
(5) Brown-red silt clay layer (18–26 cm): It has a dense structure
without embodied materials.
All of the human fossils and most of the animal fossils were
excavated from Region 1. A few animal fossils were found in Region
2. No fossils were found in Region 4 and the Entrance region. In
Region 1, about 50 m
2
were excavated. The 2004 and 2005 exca-
vations are represented by the white area in Fig. 4, and the dark
area indicates the 2006 excavation. Among the seven human teeth
Fig. 2. The location of Huanglong Cave and its horizontal profile.
Fig. 3. The plan view of the Huanglong Cave and the excavation area (modified after Wu et al., 2007a).
W. Liu et al. / Quaternary International 211 (2010) 29–41 31
unearthed from Huanglong Cave, five of them were found during
the 2004–2005 excavations in square zones N1W2, N1W3, N3E1,
N2W1 and N1W3. The other two human teeth were found during
the 2006 excavation in square zones N2E6 and N1E7. All the seven
hominin teeth, all the stone artifacts and most of the animal fossils
are derived from Layer 3. The hominin teeth and stone artifacts
were found at the bottom of the layer, in association with the
stalagmites.
During the excavations, the fossil yielding layer was carefully
examined to ascertainif there is any possible evidence indicating the
fossils and artefacts were transported, sorted or redeposited. For the
following reasons, the materials are considered to have been buried
rapidly. First, Layer 3 is comprised of sharp-edged breccia. Little
evidence of fluvially transported sediment is present. No horizon-
tally or cross-bedded sediment was observed in the deposit. Second,
the faunal assemblage contains different species of varying sizes
with no evidence of apparent sorting. Both intact complete small
mammal bones and limb bones of large animals are well preserved.
Third, evidence of human fire use, which will be discussed below, is
distributed in multiple spots. If there had been a flow of water, then
no evidence of fire would be anticipated. However, the influence of
rodents cannot be completely ruled out (see below).
3. Dating
Chronometric dating analysis for the layer yielding the human
teeth (Layer 3) was carried out in three different laboratories. The
first Uranium-series (U-series) dating on two rhinoceros teeth was
analyzed at the Laboratory of the Institute of Karst Geology, Chinese
Academy of Geological Sciences. An age range of 94.712.5
79.4 6.3 ka (Table 2) was derived (Wu et al., 2006, 2007a).
The second dating attempt, again using U-series, was made on
a pure and dense stalagmite sample collected from the same hori-
zontal level that yielded the human fossils. The speleothem sample
was analyzed at the TIMS Laboratory in the Department of Geology
and Geophysics, University of Minnesota. Two small pieces from the
stalagmite sample were taken out and tested separately. The results
indicated an age range of 103,739 103,119 1616 1348 years
(Table 3).
To further assess the age of the hominin tooth layer, electron spin
resonance (ESR) dating on a rhinoceros tooth was conducted. The
rhinoceros tooth was analyzed at the State Key Laboratory of Earth-
quake Dynamics, Institute of Geology, China Earthquake Adminis-
tration, resulting in an age of 44,180 34,780 3280 4540 years
(Tabl e 4), which is much younger than the dates from the U-series
studies.
Among the three dating procedures for Layer 3, the two U-series
dates give an age range between 79,400 and 103,739 years. The ESR
date is much younger, between 44,180 and 34,780 years. Because the
exact provenience of the speleothem sample and its relationship
with the human fossils need further verification, more detailed study
of the stratigraphy and context from which the samples were
derived are necessary. In particular, if the speleothem overlies the
human fossil layer, the age of the human fossils could be older than
103 ka. However, if the stalagmite underlies the human fossil layer,
then the hominin deposition could be much younger. In order to
attempt to narrow the age range of the deposit, AMS
14
C analysis will
be conducted on associated bones. If the AMS dates fall outside the
accepted range (>50 ka), it would lend support for the older U-series
dates. If the AMS dates are younger than or around 40 ka, it would
support the ESR date. At this point, it is believed that the age of the
Huanglong Cave hominin teeth can be conservatively bracketed
between 100 ka and 34 ka.
Although questions still exist about the chronometric age of the
human fossils, based on the biostratigraphy, it can be minimally
concluded that the deposits are Late Pleistocene and clearly not
Holocene. The animal fossils from Huanglong Cave were classified
into 91 taxa (Wu et al., 2006, 2007a), representing a mixed
assemblage comprising extinct (e.g., Macaca robustus,Cricetinus
varians,Crocuta ultima,Ailuropoda melanoleuca baconi,Ursus thi-
betanus kokeni,Stegodon orientalis,Rhinoceros sinensis,Megatapirus
augustus), and extant species (e. g., Neofelis nebulosa,Cuon alpinus,
Canis lupus,Arctonyx collaris,Paguma larvata,Sus scrofa,Rusa uni-
color, and Cervus nippon). Among the 91 identified taxa found in the
Huanglong Cave, 20% are extinct species (see Table 5 for details).
The Huanglong Cave faunal assemblage is typical of the Oriental
biogeographic region (Norton et al., in press). The faunal composition
indicates both tropical and subtropical forest environments. For
example, Neofelis nebulosa is the representative taxa of tree living
animals in subtropical forests (Sheng, 1994), and Belomys pearsoni
lives in the evergreen broadleaved forest (Corbet and Hill, 1980).
There are also many fossils of Rhinolophus, an animal normally living
in broad forest environments. The Huanglong Cave fauna also
includes animals that typically exist in the Qinling Mountains. These
animals include Capricornis sumatraensis,Naemorhedus goral,and
Caryomys ineg. The faunal composition reflects its geographic loca-
tion in the southern part of the Qinling Mountains, and its border
position separating the Palearctic and Oriental biogeographic zones.
4. Hominin paleontology
The seven human teeth found in the Huanglong Cave were
identified as upper central and lateral incisors, upper canine, upper
third molar, lower lateral incisor, lower second and third molars
(see Fig. 5 and Table 6).
4.1. Age and individuals
According to the occlusal wear and root formation, the ages of
the individuals were estimated. Six teeth were estimated to be from
adults between 20 and 45 years of age, and only one tooth may
Fig. 4. Stratigraphic context of the Huanglong Cave deposits (from Wu et al., 2006).
Table 2
U-series analysis and age result for the Huanglong Cave rhinoceros teeth.
Sample No. Uranium
content (Ug/g)
234
U/
238
U
230
TH/
234
U Age (Ka) Corrected
age (ka)
020 1.445 1.553 0.576 85.3 6.9 79.4 6.3
021 1.583 1.469 0.595 92.4 12.1 94.7 12.5
W. Liu et al. / Quaternary International 211 (2010) 29–4132
represent a sub-adult less than 20 years old (see Table 6 for details).
From the occlusal views of these teeth (see Fig. 5), very different
wear patterns can be observed. Thus, the seven teeth came from
multiple individuals. Currently, with just these isolated teeth, the
sex of the individuals cannot be determined.
4.2. Research method
The morphology of the human teeth was assessed by referring
to the methods described by several authors (e.g., Weidenreich,
1937; Turner et al., 1991; Irish, 1998; Irish and Guatelli-Steinberg,
2003; Bailey et al., 2008). Other studies (e.g., Wolpoff, 1979; Ber-
mudez de Castro, 1988, 1993) were also consulted. The Arizona
State University Dental Anthropology System (ASUDAS) (Turner
et al., 1991) was used to grade some morphological features. Dental
metric analysis followed the descriptions by Bermudez de Castro
(1986, 1993), the mesiodistal (MD) and buccolingual (BL) dimen-
sions of the Huanglong Cave teeth were recorded to the nearest
0.1 mm. The crown base area (CBA ¼MD BL) and crown shape
index (CSI ¼BL 100/MD) were calculated from these tooth
dimensions. For those teeth which have light interproximal wear,
original borders were estimated by referring to overall crown shape
and the buccolingual extent of the wear facet (Bailey, 2004).
4.3. Comparative samples
The comparative samples include Chinese (Homo erectus,
archaic Homo sapiens, Late Pleistocene humans and recent Chinese
populations), Neandertals, European Middle Pleistocene humans
(Krapina and Atapuerca SH), European Late Pleistocene humans,
modern Europeans and the early modern humans of Skhul and
Qafzeh (Weidenreich, 1937; Wolpoff, 1971, 1979; Brace, 1976;
Frayer, 1977; Brace et al., 1984; Bermudez de Castro, 1993; Liu and
Yang, 1999; He, 2000).
The following paragraphs describe the morphology and metric
features of the Huanglong Cave human teeth (see also Liu et al.,
2009a; Liu et al., 2009c).
4.4. Upper central incisor (I
1
)
The crown lingual surface is shovel-shaped. There is a weakly
developed basal prominence, but no finger-shaped projection can
be identified. The whole crown labial surface bulges slightly. Both
the mesial border and distal border at the labial side display
marked projections, which makes the whole crown labial surface
depressed and flat. Because both the lingual and buccal surfaces of
the crown are shovel-shaped, this upper central incisor has the
double shovel-shaped feature. According to ASUDAS, the degrees of
shoveling and double-shoveling are 4 and 3 respectively. The root is
awl-shaped. From the cervical region to the root tip, the root
gradually thins.
Previous studies of upper central incisors of Pleistocene homi-
nins found in China indicate that there are some morphological
features shared by the Chinese archaic hominins (Weidenreich,
1937; Wu et al., 1989). These features are pronounced labial
bulging, pronounced basal tubercle with finger-shaped projections
extending toward the incisal edge, the crown and root are not in the
same long axis, and the root is robust and awl-shaped. These
morphological features cannot be identified in the Huanglong Cave
upper central incisor. The crown lingual surface of the Huanglong
Cave upper central incisor is very flat and smooth. Its crown labial
surface is not only less bulging but shows obvious depressions. Both
the MD and BL diameter are within the modern human ranges, and
smaller than those of Chinese Homo erectus and archaic Homo
sapiens, and also smaller than Middle and Late Pleistocene homi-
nins in other regions of the world (see Table 7 for details). The long
axis of the crown and root of the Huanglong Cave upper central
incisor is not in the same line and the crown is inclined toward the
direction making a slight angle between the crown and root. These
morphological features indicate the upper central incisor is more
like modern humans. However, the robust root of the Huanglong
Cave specimen differs from that of modern humans, and resembles
that of earlier fossil hominins. The CSI is 76.5, which is closer to the
average of the same index of Homo erectus (75.7), and smaller than
those of archaic Homo sapiens and other Late Pleistocene humans
and modern humans (see Table 7).
4.5. Upper lateral incisor (I
2
)
The crown lingual surface is shovel-shaped (grade 3 of ASUDAS).
The crown lingual surface is flat without a basal tubercle and
a finger-shaped projection. The root is slender, flat and awl-shaped.
The crown mesiodistal and labiolingual dimensions fall within
range of modern humans, although the labiolingual dimension is
relatively large (see Table 7). In general, the labiolingual dimension
of modern humans’ upper lateral incisors is smaller than Homo
erectus and other Pleistocene humans. With the latter groups, CSI
Table 3
The U-series analysis and age results for the Huanglong cave stalagmite.
Sample
Number
238
U (ppb)
232
Th (ppt)
230
Th /
232
Th
(atomic 10
6
)
d
234
U
a
(measured)
230
Th /
238
U
(activity)
230
Th Age (yr)
(uncorrected)
230
Th Age (yr)
(corrected)
d
234
U
Initialb
(corrected)
HL-1 146.00.2 18922 58 113 1 379 2 0.8903 0.0051 106267 1009 103739 1616 508 4
HL-2 141.80.2 14132 38 147 1 384 2 0.8880 0.0048 105049 949 103119 1348 514 4
l
230
¼9.1577 x 10
6
y
1
,
l
234
¼2.8263 10
6
y
1
,
l
238
¼1.55125 10
10
y
1
.
a
d
234
U¼([
234
U/
238
U]
activity
1) 1000.
b
d
234
U
initial
was calculated based on
230
Th age (T), i.e.,
d
234
U
initial
¼
d
234
U
measured
e
l
234T
. Corrected
230
Th ages assume the initial
230
Th/
232
Th atomic ratio of
4.4 2.2 10
6
. Those are the values for a material at secular equilibrium, with the bulk earth
232
Th/
238
U value of 3.8. The errors are arbitrarily assumed to be 50%.
Table 4
The ESR analysis and age result for the Huanglong Cave rhinoceros tooth.
Lab Number 5103
Sample Number N1W2:59
Enamel thickness (mm) 2.1
Thickness of removed enamel (mm) 0.1
Dose (Gy) 56.16 2.38
U-enamel (ppm) 2.81
U-dentine (ppm) 5.05
U-sediment (ppm) 2.10 0.10
Th-sediment (ppm) 7.05 0.35
K-sediment (%) 1.52
EU (Gy/ka) 1.673 0.142
LU (Gy/ka) 1.316 0.124
EU age (ka) 34.78 3.28
LU age (ka) 44.18 4.54
W. Liu et al. / Quaternary International 211 (2010) 29–41 33
range between 92.9 and 97.2; the range of the same index in
Neandertal, European Middle Pleistocene humans, and modern
Europeans are 100.0–106.4 and 89.5–98.5 respectively. The data
indicate that both the shape and CSI of the Huanglong Cave upper
lateral incisor are more robust than modern humans.
4.6. Upper canine (C)
The tooth is robust. The crown and the root are at the same
longitudinal axis with only the root tip bending toward the distal
direction. The crown labial surface bulges in both horizontal and
longitudinal directions, more pronounced in the former direction.
Because of advanced wear, most of the crown’s morphological
features are absent. On the preserved portion, basal parts of mesial
and distal marginal ridges are well developed, and the lingual basal
prominence is complete. The basal prominence is partly separated
from the lingual surface and forms a weakly developed basal
tubercle. There is also a small finger-like projection coming from
the lower end of the basal prominence. Because of the severe crown
wear, the length and shape of the projection cannot be determined.
Both the crown MD and LL dimensions of the Huanglong Cave
upper canine are smaller than Chinese Homo erectus and archaic
Homo sapiens, but greater than those of Late Pleistocene and
Holocene Chinese populations (see Table 7).
The upper canines of Zhoukoudian Homo erectus are very robust
in both their crown and root morphology. The crowns are relatively
low and broad. There is a pronounced cingulum at the cervical
region and triangular prominence at the crown mesial and distal
sides. The structure of the crown lingual surface is complicated
with well developed basal tubercle and finger-shaped projections.
The labial surface bulges strongly in the horizontal direction. The
roots are big and robust with the root tip blunt. All dimensions of
Zhoukoudian Homo erectus upper canines are larger than modern
humans (Weidenreich, 1937). The morphology of Chinese archaic
Homo sapiens upper canines resembles that of Zhoukoudian spec-
imens in many aspects (Jia et al.,1979; Wu,1984). Although not well
studied (Li et al., 1984), the upper canine morphological features of
Late Pleistocene Chinese resemble those of modern Chinese. The
crowns of Late Pleistocene hominins thus far found in China are
robust with their CSI scores exceeding even Zhoukoudian Homo
erectus (see Table 7). The European Middle and Late Pleistocene
hominins share many upper canine feature patterns with Zhou-
koudian Homo erectus. However, the European hominin upper
canines have greater size variation (Wolpoff, 1979; Bermudez de
Table 5
Fauna compositions of the Huanglong Cave.
Order/genus /species Extinct Extant
Lamellibranchia Eulamellibbanchia
6
Gastropoda Order stylommatophora
6
Malacostraca Somanniathelphusa zhongshiensis
6
Osteichthyes Diptychus kaznakovi
6
Mylopharyngodon piceus
6
Elopichthys bambusa
6
Amphibia Rusa sp.
6
Reptilia Boidae
6
Mammalia Homo sapiens
6
Macaca robustus :
Macaca mulatta
6
Rhinopithecus iantianensis :
Trachypithecus phayrei
6
Hylobates sp.
6
Erinaceus sp.
6
Chimmarogale sp.
6
Anourosorex squamipes
6
Soriculus leucops
6
Uropsilus soricipes
6
Megaderm lyra
6
Hipposideros pratti
6
Hipposideros armiger
6
Rhinolophus ferrumequinum
6
Rhinolophus pearsoni
6
Rhinolophus macrotis
6
Rhinolophus cornutus
6
Murina leucogaster
6
Myotis formosus
6
Myotis daubentoni
6
Myotis sp.
6
Tylonycteris pachypus
6
Miniopterus schreibersi
6
Scotomanes emarginatus
6
Pipistrellus coromandra
6
Ochotona thibetana
6
Ochotona huangensis
6
Collosciurus erythraeus
6
Sciurotamias forreti
6
Belomys pearsoni
6
Petaurista xanthotis
6
Petaurista alborufus
6
Caryomys inez
6
Leopoldamys edwardsi
6
Niviventer andersoni :
Niviventer fulvescens
6
Niviventer confucianus
6
Mus sp.
6
Hapalomys sp.
6
Hystrix magna :
Hystrix subcristata
6
Atherurus macrourus
6
T.cf. fasciculata
6
Rhizomys sinensis
6
Rhizomys sp.
6
Stegodon orientalis :
Cuon javanicus antiquus :
Canis variabilisb :
Crocuta ultima :
Selenarctos thibetanus :
Helarctos malayanus
6
Ailuropoda melanoleuca baconi :
Mustela kathiah
6
Arctonyx collaris
6
Herpestes urva
6
Lutra lutra
6
Viverra zibetha
6
Paguma larvata
6
Felis sinensis :
Neofelis nebulosa
6
Panthera tigris amoyensis
6
Rhinoceros sinensis :
Dicerorhinus kirchbergensis :
Megatapirus augustus :
Sus scrofa
6
Table 5 (continued )
Order/genus /species Extinct Extant
Sus xiaozhu :
Moschus moschiferus plicodon :
Muntiacus muntjak
6
Muntiacus reevesi
6
Elaphoudus cephalophus
6
Rusa unicolor
6
Hydropotes inermis
6
Budorcas cf. taxicolor
6
Capricornis sumatraensis
6
Naemorhedus goral
6
Bubalus bubalis :
Leptobos sp. :
Aves Aviceda jerdoni
6
Cygnus
6
Ciconia boyciana
6
Phasianidae indet
6
Scolopacidae indet
6
Total 91 18 (20%) 73(80%)
W. Liu et al. / Quaternary International 211 (2010) 29–4134
Castro, 1986, 1988, 1993). Compared with the Pleistocene hominins
mentioned above, the Huanglong Cave upper canine looks robust
and possesses some primitive features of archaic hominins. Taking
crown area (CBA) as the tooth size, the Huanglong Cave upper
canine is smaller than the averages of Chinese Homo erectus and
archaic Homo sapiens, but lager than the averages of Chinese Late
Pleistocene hominins and modern Chinese. The CSI of the Huang-
long Cave canine is close to those of nearly all Late Pleistocene
hominins and modern humans. In modern humans, both the crown
and roots of the canines are gracile with the bulging of the crown
labial surface not pronounced. All dimensions of the upper canines
in modern humans are small. The root is short relative to the crown.
The root length (from cervical line to root tip) is 56.3% of the whole
tooth length in the canines of modern Chinese, while this measure
is 62.5% in the Huanglong Cave upper canine, which is very close to
62.0% of Zhoukoudian Homo erectus (Liu and Yang, 1999).
Fig. 5. The human teeth found in the Huanglong Cave (from Liu et al., 2009a,b) (A: right I
1
; B: Left I
2
; C: Left UC; D: left M
3
; E: Right I
2
; F: right M
2
; G: left M3 Lingual, buccal, mesial,
distal sides, and occusal surface are represented from 1 to 5 respectively) (from Liu et al., 2009a).
W. Liu et al. / Quaternary International 211 (2010) 29–41 35
4.7. Upper third molar (M
3
)
The crown is triangular-shaped with its base long in the mesial
side. Among the four cusps of paracone, metacone, protocone and
hypocone on the occlusal surface, the paracone is the largest fol-
lowed by the protocone and metacone, with hypocone the smallest.
The tooth has three roots diverging at about 3 mm from the cervical
region. All of the roots have not fully developed with their root
canals open at the tips. Around the root canals, the cement is fused
with the surface flat; thus indicating the roots were still developing.
Thus, this tooth is from a sub-adult. All of the crown dimensions of
the Huanglong Cave upper third molar are smaller than those of
Pleistocene hominins in both China and other regions of the world.
Compared with the upper third molars of archaic hominins, the
Huanglong Cave upper third molar has simple crown occlusal
patterns and lacks the features often present in Pleistocene homi-
nins upper M3, such as wrinkles and oblique ridges. The paracone
and protocone are similar in size. Compared with the other cusps,
the metacone is a little smaller. All of the tooth dimensions of the
Huanglong Cave upper third molar are within the range of modern
human humans. The roots of the Huanglong Cave upper M3 are
substantially branched, which is different from the modern human
pattern.
4.8. Lower lateral incisor (I
2
)
A weak shovel-shaped lingual side (equal grade 1–2 of ASUDAS)
is present. The lingual surface of the crown is very smooth. The
region bordering the crown base and root slightly bulges, but no
obvious tubercle-like structure is present. The whole crown labial
side is flat and inclines toward the lingual direction. Compared with
the crown, the root looks slender. Among the lower lateral incisors
of modern Chinese, the shovel-shaped crown lingual sides are more
obvious than the Huanglong Cave specimen. Compared with
Pleistocene hominin lower lateral incisors, the morphological
patterns of the Huanglong Cave specimen more resembles modern
humans because of the following characteristics: gracile tooth; less
bulged labial surface; the longitudinal groove occurred in only one
side of the root and smaller dimensions. All of the crown dimen-
sions of the Huanglong Cave lower lateral incisor are smaller than
those of Chinese Homo erectus and archaic Homo sapiens, and also
smaller than those of the early modern humans of Skhul and Qaf-
zeh and European Pleistocene humans (see Table 7). However, the
root tip of the Huanglong Cave specimen is rounded, which differs
from modern humans.
4.9. Lower second molar (M
2
)
Because of the occlusal wear, nearly all of the morphological
features are missing. The maximum BL dimension is located at the
slightly mesial to the middle part of the crown, indicating that the
trigonid is bigger than the talonid. Judging from the remaining
occlusal profile, there appears to be five cusps on the occlusal
surface. The hypoconid and hypoconulid are much smaller than the
other three cusps. The dimensions of the Huanglong Cave lower
second molar are smaller than those of Chinese Homo erectus, and
also smaller than those of Krapina and the Neandertals. But the
tooth dimensions of the Huanglong Cave specimen is larger than
those of Atapuerca SH and European Late Pleistocene hominins,
and closer to those of the Near East early modern humans from
Skhul. Compared with both Chinese and European modern
humans, all of the dimensions of the Huanglong Cave lower second
molar are larger (see Table 7). In most cases, the lower second
Table 6
List of isolated hominin teeth from the Huanglong Cave.
Specimen No. Tooth Side Wear category Estimated age
(years old)
Upper
N2E6:X34 I1 R 2–3 20–30
I2 L 4 30–40
C L 5 30–40
M3 L 1 <20
Lower
I2 R 3 20–25
M2 R 5 35–45
M3 L 2 20–30
Tooth wears were graded after Smith (1984); Aging was made with methods
provided by Alt et al. (1998) and Hillson (1996).
Table 7
Tooth measurements of the Huanglong Cave hominin and comparative samples (mm).
China Neanderthals Skhul Qafzeh Herto European
Early UP
European
Later UP
Atapuerca
SH
Modern
Europeans
HC H. erectus A rchaic
H. sapiens
Late
Pleistocene
Neolithic
moderns
Living
moderns
I
1
MD 8.5 10.7 9.8 8.2 8.6 8.6 9.3 9.4 9.9 9.3 8.7 9.6 7.8
LL 6.5 8.1 8.0 7.2 7.3 7.3 8.2 8.0 8.1 7.5 7.5 7.7 6.6
CSI 76.5 75.7 81.8 87.8 84.8 84.8 87.3 85.7 81.8 80.6 86.2 80.5 84.6
I
2
MD 7.3 8.3 7.9 7.1 7.1 6.9 7.8 7.0 8.0 7.6 7.1 7.9 6.2
BL 7.0 8.1 7.6 6.9 6.6 6.5 8.3 7.3 7.7 6.8 6.6 7.9 6.0
CSI 95.3 97.5 95.8 97.2 93.0 94.2 106.4 105.5 96.3 89.5 93.0 100.0 96.8
UC MD 9.0 9.4 9.5 8.0 7.9 7.9 8.2 8.5 8.6 7.0 8.0 8.0 8.6 7.4
BL 9.8 10.2 9.6 8.9 8.4 8.3 9.5 9.1 9.5 9.0 9.0 8.8 9.7 8.2
CSI 108.9 108.5 103.9 111.3 106.3 106.4 115.9 107.6 110.5 120.0 112.5 110.0 112.8 110.8
M
3
MD 8.5 9.6 8.5 9.4 9.0 8.9 9.6 9.9 9.8 8.6 9.5 9.0 8.7 8.5
BL 11.2 11.9 9.9 11.0 10.8 10.9 12.0 11.7 12.6 11.9 11.4 11.4 11.7 10.5
CSI 131.8 121.1 116.5 117.0 120.0 122.5 125.0 119.8 128.6 138.4 120.0 126.7 134.5 123.5
I
2
MD 6.1 6.9 6.0 6.0 6.7 6.5 6.8 6.4 5.6 6.6 5.8
LL 6.0 7.1 6.3 6.3 7.8 7.4 7.2 7.0 6.5 7.3 6.3
CSI 98.4 102.9 105.0 105.0 116.4 112.9 105.9 109.4 116.0 110.7 108.6
M
2
MD 11.0 12.6 11.2 11.4 10.9 10.9 12.0 10.9 11.4 11.3 10.9 11.0 10.8
BL 11.0 12.7 10.1 10 .7 10.7 10.4 11.1 10.9 11.7 10.8 10.7 10.2 10.3
CSI 100.0 100.8 90.2 93.9 98.2 95.4 92.5 100. 4 102.6 95.6 98.2 93.3 95.3
M
3
MD 10.0 11.7 10.9 10.7 10.7 11.7 10.9 12.6 11.1 10.6 11.3 10.8
BL 10.5 11.2 10.4 10.3 10.1 10.9 10.3 11.5 10.7 10.4 9.9 10.2
CSI 105.0 95.7 95.4 96.3 94.4 93.2 94.2 91.3 96.4 98.1 87.9 94.4
W. Liu et al. / Quaternary International 211 (2010) 29–4136
molars of modern humans are more gracile with simple occlusal
structure and much smaller dimensions. Both the length and
breadth of the trigonid decrease in modern human lower second
molars, while the talonid increases to be comparable to the trig-
onid. The occlusal pattern tends to be simple and the cusp surface is
flat and smooth lacking the accessory ridges. There are only four or
five cusps in modern human lower second molars. Modern Chinese
lower second molars are round–square-shaped with four cusps.
Based on these comparisons, the Huanglong Cave specimen
resembles modern humans in most of the features, but still
preserves some primitive morphological features of Late Pleisto-
cene hominins.
4.10. Lower third molar (M
3
)
The occlusal surface is square-shaped with its four corners
blunt. The crown MD dimension is greater than the BL dimension.
The crown has pronounced bulges on all sides. There is an obvious
enamel extension at the crown buccal side that reaches the roots
branching region (Fig. 5). Among the six cusps on the occlusal
surface, the protoconid, endoconid and metaconid are similar in
size and much larger than the hypoconid, hypoconulid and endo-
conulid. Except for the cusps and grooves mentioned above, there
are no other minor groove, fissure or accessory ridges on the
occlusal surface. Thus, the occlusal structure is simple. The
Huanglong Cave lower third molar has two roots on the mesial and
distal sides. The two roots become narrower and sharper toward
the tips.
The morphology of the Huanglong Cave lower third molar is
characterized by a lower crown, six cusps, decreased talonid and
separate roots. Although these characteristics are not specific to
modern humans, the variation in the lower third molar
morphology in modern humans is quite large. Thus the pattern in
the Huanglong Cave specimen can also be found in modern
humans. Additional features that align the Huanglong Cave spec-
imen with modern humans include a simple crown structure, lack
of accessory ridges and grooves on the occlusal surface, and overall
tooth size that is smaller than fossil hominins and within range of
Table 8
The main diagnostic morphological features of the Huanglong Cave hominin teeth, which share with modern humans and similar to archaic hominins.
Tooth Features shared with modern humans Features similar to archaic hominins
I
1
The crown lingual surface is smooth without basal tubercle and linger-shaped
projection. Both the lingual and labial sides are shovel-shaped. All the crown
dimensions are within the modern human ranges.
The MD diameter is relatively big with CSI as 76.5 close to
archaic hominins. The root is a little robust.
I
2
The main morphology and metric dimensions are within the variation ranges of
the modern humans. The structure of crown lingual surface is simple and
smooth without basal tubercle and finger-shaped projection. The bulging of
crown labial surface is not obvious. The root is gracile with its
long axis meeting the crown in an angle.
The crown looks robust with its labiolingual diameter relatively
big and CSI closer to archaic hominins.
UC The crown dimensions and CSI are closer to modern humans. The whole tooth looks robust with its crown and root in the
same axis. The crown labial surface bulges pronouncedly. The
crown lingual surface has well developed basal tubercle and
finger-shaped projection. The root tip is blunt. The tooth
dimensions are bigger than the average of modern humans.
M
3
The pattern of occlusal surface is simple without wrinkles. The paracone and
protocone are similar in size. Metacone does not reduce very much. All the
metric dimensions are within the variation ranges of modern humans.
The root bifurcates pronouncedly, which is different from the
fusing trend of modern humans.
I
2
The whole tooth looks gracile. The crown lingual surface is concave and smooth
slightly with marginal ridges weakly developed and no basal tubercle. The
crown labial surface is flat and inclines lingually. There is a vertical groove at one
side of the root. All the tooth dimensions are small and within the variation
ranges of modern humans.
The root tip is blunt.
M
2
The structure of the crown occlusal surface is simple. The trigonid is bigger than telanid. Root is robust. Tooth
dimensions are bigger than the average of modern humans.
M
3
The pattern of occlusal surface is simple lacking the accessory
ridge and fissures. The tooth dimensions are closer to those of
modern humans. Enamel extension is identified.
The crown is low. There are six cusps. The reduction of telanid is
not obvious. The two roots separate.
Fig. 6. Scatter plots of the UI2 and UC crown MD and BL dimensions. Left: UI2; right: UC.
W. Liu et al. / Quaternary International 211 (2010) 29–41 37
modern humans. Perhaps most importantly, the enamel extension
is widely regarded as a modern human feature (Turner, 1990;
Turner and Scott, 1997).
Based on all of these morphological and metric observations
(see Table 8 for a summary), all of the Huanglong Cave hominin
teeth most closely align with modern humans. However, analysis
indicates that the Huanglong Cave teeth retain a few symplesio-
morphic features that characterize Late Pleistocene hominins (see
Table 8 and Fig. 6).
The Huanglong Cave anterior teeth display some evidence of
activity-induced patterns of dental abrasion including obvious
enamel chipping along the incisal region (Fig. 7), interproximal
grooves, and damage on the labial surface of the lateral upper
incisor. These unusual patterns of tooth use-marks suggest the
Huanglong Cave humans frequently used their anterior teeth for
non-masticatory utilization (see Larsen, 1997 for discussion of this).
The interproximal grooves on the upper anterior teeth suggest the
Huanglong Cave humans often practiced tooth-pickings. Consid-
ering these observations, the Huanglong Cave humans used their
teeth to perform a series of work-related activities (e.g., holding
skin with their teeth during skin preparation), similar to what
Neandertals may have been doing (Wallace, 1975; Bermudez de
Castro et al., 1988). Thus tooth morphology may be related to the
functional adaptation of the anterior tooth use.
5. Archaeology
5.1. Evidence for fire use
During the 2006 excavation, in the same layer as the human
teeth, a few patches of black material were identified embedded in
the deposit. To explore the possibility that the materials were the
remains of fire, comparative laboratory analyses of these black
patches were conducted, along with samples taken from several
other places in the cave. A variety of methods were used: micro-
morphology; element content determination; and deposit
temperature analysis. The results indicate that the carbon content
in the black deposit ranged between 64.59 and 73.29%, which was
much greater than the 5.82–9.49% derived from the comparative
samples (Table 9). The micromorphology analysis on the black
deposit samples reveals plant structures like axial parenchyma,
fibrocyte, uniseriate ray and vessel (Fig. 8). Deposit temperature
analysis confirms that the blackened materials were the result of
a high temperature event. Based on the result of these lab analyses,
the black patches are the remains of fire.
5.2. Artifacts
The artifact assemblage is comprised of lithic and bone
implements [representative samples are shown in Fig. 9; detailed
descriptions and analysis of the artifacts can be found in Pei et al.,
2008]. The lithic assemblage includes thirty-six artifacts: two stone
hammers, one core, eight flakes, fourteen retouched artifacts, four
block fragments, five waste flakes (debris) and two bipolar cores.
Many of the stone artifacts (78%) were manufactured using the
local vein quartz embedded in the Upper Sinian limestone and
marlite from which the cave developed. Only 22% of the lithic
artifacts were made on pebbles available from the ancient riverbeds
outside the cave. The hominins probably knapped inside the cave,
as deduced from the different lithic types identified. In general, the
artifacts were detached by direct hard-hammer percussion, though
the existence of two bipolar (one core and one flake) indicates that
the bipolar technique was also used. Knapping strategies were
simple, flakes show plan rather than cortical butts, and dorsal scars
are parallel to the flake extraction axis, which indicates unidirec-
tional knapping.
Stone tools with secondary retouch include scrapers, picks,
chopper-chopping tools, a burin and an awl. Six scrapers, made of
quartz and flint, represent the greatest percentage of stone tool
types. All of the scrapers display retouch on the flakes. Three picks,
made of silicarenite, volcanic rock, and quartzite were identified.
The blanks for the picks are pebbles. One finished pick has a thick
and blunt point at one end, which may have been used for digging.
Two of the three picks exhibit deliberate and standardized modi-
fication: a pointed end was carefully produced from the original
curved surface of the pebble and the other end was purposely
Fig. 7. The tooth wear and chippings of upper incisor from the Huanglong Cave hominin. A: incisal view; B: lingual view; C: labial view.
Table 9
The element content analysis of the Huanglong Cave samples (wt%).
Specimen No. C O Cu Na Mg Al Si P K Ca Ti Fe
HLD 1(1) 73.29 23.99 / / 0.88 0.05 0.06 0.09 0.06 1.58 / /
HLD 1(2) 71.62 23.03 0.51 / 1.19 0.49 0.16 0.09 / 2.91 / /
HLD 1(3) 70.75 25.79 / / 0.92 / / 0.07 / 2.47 / /
HLD 1(4) 72.02 22.13 0.28 / 1.24 0.42 0.12 0.11 / 3.69 / /
HLD 1(5) 62.44 25.41 / 1.13 / / 0.56 / 10.47 / /
HLD 2 64.59 29.35 / 0.60 0.67 0.96 0.89 / 2.94 / /
HLD 3 9.49 48.38 0.69 1.96 7.76 20.13 / 1.57 4.24 / 5.78
HLD 4 5.82 43.75 2.10 8.29 22.09 1.18 / 1.69 4.94 1.28 8.87
HLD 5 6.52 44.42 1.51 1.90 7.84 23.59 / 1.69 2.72 / 9.82
HLD 6 5.82 46.83 1.06 2.03 8.43 25.66 / 2.32 0.97 / 6.86
HLD 1: sample of black layer.
HLD 2–6: comparative samples taken from other parts of Huanglong Cave.
W. Liu et al. / Quaternary International 211 (2010) 29–4138
Fig. 8. The SMC picture of charcoal from the Huanglong Cave black layer shows plant structure (ap: axial parenchyma; fc: fibrocyte; ur: uniseriate ray; vl: vessel) from Liu., et al,
2009b.
Fig. 9. Some stone and bone tools found in Huanglong Cave 1 Bone unifacial spade; 2 Bone point; 3 Bone scraper; 4 Bone point; 5 Flake; 6 Pick; 7 Pick; 8 Burin; 9 Core from Pei.,
et al, 2008.
W. Liu et al. / Quaternary International 211 (2010) 29–41 39
blunted, making it comfortable to be held. One stone awl, made of
vein quartz, has two sharp edges that meet each other to form
a short and blunt point edge. One stone burin, made of vein quartz,
displays unidirectional retouch. Three chopper-chopping tools, two
made of vein quartz and one made of jasper, have been retouched
on several edges. In addition, one stone hammerstone has
a depressed region with small pits. The depression developed over
time through consistent breaking of nuts. In the third excavation,
six bone artifacts were found, which include three bone points, two
bone scrapers and one bone unifacial spade.
5.3. Bone surface modifications
Although a complete taphonomic study utilizing modern
methodologies has yet to be undertaken, evidence of butchering
was found on forty of the 3000 bones. Cut marks, percussion marks,
and scraping marks were identified, primarily on the long bones.
Cut marks and scraping marks are usually considered to be
evidence of defleshing activities (Lyman, 1994). Percussion marks
on long bone midshafts are the result of marrow processing (Nor-
ton and Gao, 2008a). Based on the current evidence, butchered
bones are present in the cave, a case not unlike that at Zhoukoudian
Upper Cave (Norton and Gao, 2008b). Huanglong Cave differs from
Zhoukoudian Upper Cave, however, because roughly 50% of the
former faunal assemblage displays evidence of rodent gnawing.
More detailed taphonomic studies should clarify the formation of
the Huanglong Cave faunal assemblage.
6. Discussion and conclusions
Because of the age disparity between the U-series (103–79 ka)
and ESR (44–37 ka) analyses, the chronometric dates of the
Huanglong Cave should be interpreted cautiously. At the minimum,
based on the biostratigraphy, the age of the Huanglong Cave
deposits is Late Pleistocene. Further detailed dating analysis will
hopefully give a more exact age for the Huanglong Cave hominin
occupation. Because of the paucity of reported early modern
human fossils from East Asia (Shang et al., 2007), the Huanglong
Cave human fossils could be of potential importance to addressing
issues related to the emergence and dispersal of early modern
humans in East Asia.
This analysis indicates that most of the morphologic and metric
features of the seven Huanglong Cave human teeth resemble those
of modern Homo sapiens. Generally speaking, the Huanglong Cave
human teeth look gracile and lack the archaic features usually
identified on Middle and Late Pleistocene humans. Furthermore,
this study also indicates that the Huanglong Cave human teeth
already possess some dental features of modern East Asian pop-
ulations. Studies of worldwide dental collections indicate there are
eight major dental morphological features that characterize
modern East Asian human populations (Turner, 1990). These eight
features have higher frequencies and pronounced expressions in
modern East Asian Sinodont populations (Turner, 1990). Three of
these traits (shovel-shaped incisors, double shovel-shaped upper
central incisors, and enamel extension of the upper molars) were
identified in the Huanglong Cave human teeth. However, the
Huanglong Cave human teeth still retain a few features of Late
Pleistocene humans (mainly the robust upper anterior teeth) (see
Table 8 for summary).
The intentional production and controlled use of fire is one of
the most important events in human evolution. However, the
advent and nature of fire use by Pleistocene hominins has been
strongly debated for many decades (James, 1989). To date, the
evidence of fire use by Pleistocene hominins in China is limited to
reports from Zhoukoudian Locality 1, Lantian, Xihoudu, Yuanmou
and Jinniushan. Among these sites, only the sediment from Zhou-
koudian locality 1 has undergone lab analysis, the results of which
cast doubt on controlled use of fire there (Weiner et al., 1998). More
recently, new methods of lab analyses have been applied to
examine the evidence of fire use, including element analysis,
micromorphology analysis, and high temperature of deposit anal-
ysis (Hrouda et al., 2003; Goren-Inbar et al., 2004). In the analysis of
the fire use by the Huanglong Cave humans, all three methods were
used (Liu et al., 2009b). The results indicate: 1) that carbon contents
in the samples from the black deposit are over 64%, much higher
than those of the comparative samples which are less than 10%; 2)
micromorphology analysis shows the plant structure suggesting
the black deposit layer is the charred remains of plants; and 3) the
deposit experienced high temperatures. The results of the three lab
analyses indicate that fire did occur in the cave. To determine
whether it is a natural fire or human-induced fire, further research
is necessary. Indirect evidence of a human-induced fire may be
from the location of the charred sediment, which is found about
w100 m inside the cave. Natural fire is nearly impossible to occur
so deep within a cave (Wu et al., 2007b).
The Huanglong Cave stone tools were primarily produced with
local vein quartz, sandstone, and flint. The main stone tools are
scrapers and picks, which were made by hard-hammer direct
percussion. Both large and small lithics are represented in the tool
assemblage. Although at present the number of stone artifacts found
in Huanglong Cave is insufficient to reconstruct the whole picture of
hominin behavior, the Huanglong Cave hominins used both direct
hammerstone percussion and bipolar flaking to produce flakes and
other stone tools. The existence of hammerstones, cores, flakes and
debitage further suggests that stone knapping activities occurred
inside the cave. Butchered bones found in association with the lithics
are suggestive of what the stone tools may have been used for.
In the west Hubei and Three Gorge region, where Huanglong
Cave is located, several hominin fossil sites ranging from Early to
Late Pleistocene have been discovered. These include the Yunxian
Homo erectus crania, Meipu Homo sp. teeth in Yunxian, H. erectus
teeth from Bailong Cave in Yunxi, early Homo sapiens teeth from
Xinglong Cave in Fengjie, the newly found early Homo sp. teeth
from Jianshi, and the debated mandible from Longgupo in Wushan
(Wu et al., 1989; Li and Etler, 1992; Huang et al., 1995; Gao et al.,
2004; Zhang et al., 2004; Liu et al., 2006). The Huanglong Cave
human teeth contribute to this growing list of Pleistocene hominin
fossils from this region. Ongoing and future multi-disciplinary
research at Huanglong Cave will serve to determine the role the site
and materials play in any reconstructions of human evolution from
this time period and geographic region.
Acknowledgments
The studies of Huanglong Cave have received support from many
institutions and individuals. The dating analysis was conducted
Dr. Yin Gongming and Dr. Cheng Hai. Dr. Shen Guanjun of Nanjing
Normal University assisted with the measurements of uranium
content. The field andlab analyses were supported by the Knowledge
Innovation Program of the Chinese Academy of Sciences (Grant No.
kzcx2-yw-106), the National Natural Science Foundation of China
(Grant No. 40772016) and the International Cooperation Program of
MST of China (Grant Nos. 2007DFB20330 and 2009DFB20580). We
thank the anonymous reviewers for many thoughtful and detailed
comments on an earlier draft of this manuscript.
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