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New 1.5 million-year-old Homo erectus maxilla from Sangiran(Central Java, Indonesia)
Yahdi Zaim a, Russell L. Ciochon b,*, Joshua M. Polanski c, Frederick E. Grine d, E. Arthur Bettis III e,Yan Rizal a, Robert G. Franciscus f, Roy R. Larick g, Matthew Heizler h, Aswan a, K. Lindsay Eaves f,Hannah E. Marsh f
a Department of Geology, Institute of Technology Bandung, Bandung, Java 40132, Indonesiab Department of Anthropology and Museum of Natural History, Macbride Hall, University of Iowa, Iowa City, IA 52242, USAc Department of Anthropology, University of Arkansas, Fayetteville, AR 72701, USAd Departments of Anthropology and Anatomical Sciences, Stony Brook University, Stony Brook, NY 11794, USAe Department of Geoscience, University of Iowa, Iowa City, IA 52242, USAfDepartment of Anthropology, University of Iowa, Iowa City, IA 52242, USAg Helios Laser, 160 East 238th Street, Euclid, OH 44123, USAh New Mexico Bureau of Mines and Mineral Resources, Socorro, NM 87801, USA
a r t i c l e i n f o
Article history:
Received 9 January 2009
Accepted 23 April 2011
Keywords:
Southeast Asia
Hominin evolution
Homo habilis
Grenzbank ZoneBapang formation
Sangiran formation40Ar/39Ar dating
Zhoukoudian
Dmanisi
a b s t r a c t
Sangiran (Solo Basin, Central Java, Indonesia) is the singular Homo erectus fossil locale for Early Pleis-
tocene Southeast Asia. Sangiran is the source for more than 80 specimens in deposits with 40Ar/39Ar ages
of 1.51e0.9 Ma. In April 2001, we recovered a H. erectus left maxilla fragment (preserving P3- M2) from
the Sangiran site of Bapang. The nd spot lies at the base of the Bapang Formation type section in
cemented gravelly sands traditionally called the Grenzbank Zone. Two meters above the nd spot,
pumice hornblende has produced an 40Ar/39Ar age of 1.51 0.08 Ma. With the addition of Bpg 2001.04,
Sangiran now has ve H. erectus maxillae. We compare the new maxilla with homologs representing
SangiranH. erectus, ZhoukoudianH. erectus, WesternH. erectus(pooled African and Georgian specimens),
andHomo habilis. Greatest contrast is with the Zhoukoudian maxillae, which appear to exhibit a derived
pattern of premolar-molar relationships compared to Western and Sangiran H. erectus. The dental
patterns suggest distinct demic origins for the earlier H. erectus populations represented at Sangiran and
the later population represented at Zhoukoudian. These two east Asian populations, separated by
5000 km and nearly 800 k.yr., may have had separate origins from different African/west Eurasian
populations.
2011 Elsevier Ltd. All rights reserved.
Introduction
Originally published under various names, Homo erectus fossils
were rst found in the Far East at Trinil (in 1891), Zhoukoudian (in
1921), and Sangiran (in 1937). Initially, an East Asia origin for the
species was deemed probable, but as African paleoanthropology
ascended during the 1960s, the origins and taxonomy ofH. erectus
became largely an Afro-centric topic, with East Asian fossils rep-
resenting a dispersal endpoint. However, as Eurasian paleoan-
thropology resurged during the 1990s, the spatial center-of-gravity
forH. erectus has again shifted eastward. Three Eurasian sites now
account forthe vast majority ofH. erectus fossils: Dmanisi, Sangiran,
and Zhoukoudian. While the assumption remains that H. erectus
emerged from an African hominin, the species had its greatest, and,
possibly, earliest presence across southern Eurasia (see also Lepre
and Kent, 2010). Here, we present a new Sangiran maxilla,
increasing the number and known morphometric variation of Java
H. erectusspecimens.
In 1998, the Institute of Technology Bandung and the University
of Iowa began joint interdisciplinary research at Sangiran (Central
Java, Indonesia). The program has focused on the hominin-bearing
sedimentary sequence of the upper portion of the Sangiran
Formation and the overlying Bapang Formation. More than 80
H. erectus fossils have been found in this sequence. The earliest
H. erectus are found in sediments ranging in age from >1.5 Ma
through to about 0.9 Ma (Larick et al., 2001; Ciochon et al., 2001). At* Corresponding author.
E-mail address: [email protected] (R.L. Ciochon).
Contents lists available at ScienceDirect
Journal of Human Evolution
j o u r n a l h o m e p a g e : w w w . e l s e v i e r . c o m / l o ca t e / j h e v o l
0047-2484/$e see front matter 2011 Elsevier Ltd. All rights reserved.
doi:10.1016/j.jhevol.2011.04.009
Journal of Human Evolution 61 (2011) 363e376
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Sangiran, the research focus has since turned to understanding the
local conditions that sustained H. erectus on Sunda, the early
Pleistocene emergent landmass currently represented by the
Indonesian and Philippine archipelagos (Ciochon et al., 2003, 2007;
Bettis et al., 2004, 2009a; Dizon and Pawlik, 2010; Larick and
Ciochon, 2012). Sangiran is the only site that represents the early
PleistoceneH. erectus population across all Southeast Asia. Work
commenced with40
Ar/39
Ar age analysis of volcanic heavy minerals,and interpreting the stratigraphic sequence as a set of sedimentarycycles.
On April 22, 2001, local team member, Samingan, recovereda partialH. erectusmaxilla (Bpg 2001.04) at the base of the Bapang
Formation reference section, in a unit traditionally known as the
Grenzbank Zone (Larick et al., 2000). The nd spot lay about 2 m
belowa level fromwhich pumice hornblendeproduced an 40Ar/39Ar
age of 1.51 0.08 Ma (Ciochon et al., 2005). The incontestable
provenience linked directly with datedvolcanic material makesthis
maxilla one of the oldest Sangiran dentate specimens. It lies within
the geochronological rangeofH. erectus in Africa, and istwice as old
as theoldest from Zhoukoudianin NortheastAsia (Shen et al., 2009).
Along with specimens S4, S17, S27 and Tjg 1993.05, Bpg 2001.04
representsthefth H. erectus maxilla recovered fromSangiran. Here,
we describethe specimenand its localgeological andenvironmentalcontext. Wethen turn to inter-site quantitative comparisons of basic
maxillary dental morphology. Bpg 2001.04 is compared with the
other Sangiran maxillary specimens, and with homologous mate-rials from Northeast Asia, Southwest Eurasia, and Africa.
Geological and paleoclimate background
H. erectus fossils are found in a long succession of lowland
deposits exposed in the Sangiran area of Central Java (Figure 1a).
The upper reaches of the Sangiran Formation contain the oldest
H. erectus fossils in Southeast Asia, dating to 1.6 Ma, while the
overlying Bapang Formation has yielded a large number ofH. erec-
tus remains dating to 1.5e
0.9 Ma (Larick et al., 2001). During thisperiod, depositional environments change from lake margin andmarsh to riverine. The oldest H. erectus fossils occur as one
component of the fully terrestrial and endemic island-type Ci Saatfauna, which occurs within dark-colored siltstones and mudstones
in the upper reaches of the Sangiran Formation (Watanabe and
Kadar, 1985; de Vos et al., 1994; Larick et al., 2001; Ciochon et al.,
2003; Bettis et al., 2004). When H. erectus rst arrived in the San-
giran area, sometime between 1.66 Ma and 1.57 Ma, streams
draining nearby volcanic highlands intermittently ooded the lake
margins and marshes, and occasional volcanic eruptions deposited
thin blankets of ash (Swisher, 1997, 1999; Bettis et al., 2004).
Freshwater lake-edge and marsh environments supported sedges,
ferns, water-tolerant grasses, and trees (Smah, 1984; Tonkunaga
et al., 1985), in addition to a variety of aquatic and semi-aquatic
vertebrate (Hexaprotodon, various cervids, crocodiles, turtles, andsh) and invertebrate species. Wet grasslands with scattered
shrubs occupied slightly higher parts of the Early Pleistocene
landscape, where water tables uctuated from near the land
surface to a depth of about 1 m on an annual basis (Bettis et al.,
Figure 1. (a) Plan view showing the location and regional geology of the Sangiran Dome in Central Java. The formations within the large dome map are delineated by the colors as
explained in the key. Hominin nd spots are depicted with a skull, calvaria, or mandible, as appropriate. (b) Satellite image of the Bapang site and the Bpg 2001.04 nd spot.
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2009a). Still higher, better-drained parts of the landscape sup-
ported savanna vegetation, which was most likely dominated by
sedges, grass, and ferns with scattered trees.
Between 1.6 Ma and 1.5 Ma, large streams draining volcanic
highlands to the northwest and southeast began to locally scour
and ll in the lowland in theSangiran area (Larick et al., 2001; Bettis
et al., 2004, 2009a). Local variations in stream ow and channel
characteristics and distance from river channels were importantelements that inuenced the depth of scour, the nature of sedimentthat accumulated and that controlled edaphic conditions and
vegetation patterns during this time. Riparian forest occupied theactive channel belt where shifting channels left many sandbars and
shallow abandoned channels, low-lying and frequently ooded
areas supported a moist savanna with scattered trees and shrubs,
and well-drained terraces and valley slopes were covered with
open woodlands (Ciochon et al., 2007; Bettis et al., 2009a). It was in
this setting that the hominin reported here lived at about 1.5 Ma. In
addition to H. erectus, these environments supported the Trinil H.K.
fauna represented by Panthera, various cervids, Sus branchygnatus,
and primates (Larick et al., 2000, 2001; van den Bergh et al., 2001).
Locally, the Bapang Formation is conformably overlain by uvial
deposits of the Pohjajar Formation that have a higher proportion of
air-fall tuffs, uvially reworked ash fall, and lahar-formeddiamictons.
Stratigraphy and dating of the Bpg 2001.04 locality
The Bpg 2001.04 H. erectus maxilla was found in the basal
Bapang Formation eroding from a carbonate-cemented, pebbly
channel sand, which has traditionally been called the Grenzbank
Zone, near the low-relief erosional contact between the Bapang and
Sangiran Formations (Figures 1b and 2aeb). At the nd spot, this
cemented zone occupies the central portion of a shallow uvial
channel locally cut into a 2040 cm-thick basal Bapang Formation
matrix-supported conglomerate, dominated by rip-up clasts
derived from the underlying Sangiran Formation siltstones and
mudstones. The maxilla was located in the upper meter of the
channel sand approximately 20 cm above the contact with the
Sangiran Formation (Figure 2a; UTM Zone 49 483990E 9174670N).A modern intermittent stream has cut through overlying deposits
and exposed the channel sands just north of the nd spot. The
cemented channel sands grade laterally into uncemented troughcrossbedded pebbly sand that is overlain by a silt-lled trough that
contains lenses of reworked tuff (Figure 2a). The silts are overlain
by trough crossbedded pebbly sand with lenses of epiclastic
pumice. The 40Ar/39Ar date of 1.51 0.08 Ma was obtained on
hornblende extracted from pumice in one of these lenses strati-
graphically about 2 m above and 45 m northwest of the maxilla nd
site. This date agrees well with a 40Ar/39Ar age of 1.58 0.02 Ma
from the basal Bapang Formation in the same stratigraphic section
(Swisher, 1997, 1999) as well as other 40Ar/39Ar ages from the
H. erectus-bearing interval in Sangiran (Larick et al., 2001; Ciochon
et al., 2001; Bettis et al., 2004).
The sediment at the discovery site is a poorly sorted, subangularto subrounded, carbonate-cemented, volcaniclastic sandstone that
contains a few granules. The sandstone is composed of a variety of
intermediate igneous rock fragments (granodiorite, diorite, anddacite, but dominantly andesite), as well as welded tuff andpumice,
with subordinate amounts of sedimentary rock fragments such assandstone, limestone, quartzite, phyllite, and fragments of eroded
carbonate nodules. The sand and granule matrix is cemented by
sparry calcite with euhedral to subeuhedral crystal development.
Figure 2. (a) Bapang Formation stratigraphy, dating, and sedimentary cycles. The Bpg 2001.04 nd spot, at the base of the section, is depicted with a white star. (b) Detailed
stratigraphic section showing the precise Bpg 2001.04 nd spot in the Grenzbank Zone, 2 m below the pumice horizon dated to 1.51 0.08 Ma.
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Similar sedimentlls the maxillary sinusoor of Bpg 2001.04, tying
the specimen to this localized stratigraphic horizon (see Figure 3d
and comparative description below).
The composition of this uvial sandstone indicates that its
clastic grains were derived dominantly from volcanic sources. The
fresh appearance of individual feldspar crystals indicates that the
grains underwent little to no chemical weathering prior to, during,
and subsequent to transport from volcanic highlands into the SoloBasin. Cementation by carbonate occurred at a later date asa precipitate from calcite-saturated groundwater preferentially
moving through the coarse channel sands at the Bapang/Sangirancontact. Cementation, the primary distinguishing characteristic of
the Grenzbank Zone, is a horizontally discontinuous, post-
depositional phenomenon formed in a geochemical environment
that postdates the affected sediment. The cementation has no
relationship to the age of the affected sediment, and we have
observed similar cemented sediments in other stratigraphic posi-
tions in Pleistocene deposits elsewhere in Central Java. Thus, the
Grenzbank Zone should not be considered a stratigraphic marker
bed as has been done in the past (e.g., Sudijono, 1985).
Approximately 15% of the 80 H. erectusspecimens recovered in
Sangiran have provenience in the thin zone of cemented Grenzbank
Zone sediments at the base of the Bapang Formation (Larick et al.,
2004). Though uvial reworking that concentrated fossilsoccurred throughout accumulation of the Bapang Formation, theprocess was enhanced during development of the uvial erosion
surface that marks the contact between the Sangiran and BapangFormations. This erosion surface, and another marking the contact
between the Bapang and overlying Pohjajar Formation, represent
periods of net sediment removal from the Sangiran area (Bettis
et al., 2009b). During these periods, heavy clasts, including large
vertebrate fossils, were concentrated and subjected to multiple
Figure 3. Bpg 2001.04 maxilla: (a) occlusal aspect; (b) buccal aspect; (c) lingual aspect; (d) superior aspect; (e) anterior aspect; (f) posterior aspect. Scale bar is 1 cm.
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Dental morphology and crown size
The P3 crown has an ovorectangular occlusal outline, with
a slightly dominant paracone (Figure 3a). The mesial marginal
ridge, which is slightly thicker than the distal marginal ridge, is
incised by a narrow furrow such that the mesial end of the longi-
tudinal furrow between the paracone and protocone is open. The
mesial surface preserves a 3 mm long, oblong contact facet for the
canine. Occlusal wear is slight to moderate; both cusps are blunted,and there is a small dentine exposure in the center of the paracone.
Its mesio-distal (MD) diameter is 8.1 mm. The bucco-lingual (BL)
diameter is 11.3 mm.
The crown of P4 also presents an ovorectangular outline, though
the lingual face of the protocone is skewed very slightly mesially
(Figure 3a). The protoconeand paracone are approximately equal in
size. The mesial and distal marginal ridges are complete, but low,
and the longitudinal furrow between the paracone and protocone
cuts through the lingual end of the principal paracone crest, thus
separating it from the protocone. Occlusal wear is slight; both cusps
are blunted, but dentine is not exposed. It measures 7.6 mm MD.The BL diameter is 12.2 mm.
The crown of M1 has a nearly square occlusal outline, witha more prominent mesio-buccal corner compared to the dis-
tobuccal corner (Figure 3a). The BL breadth across the trigon(13.5 mm) is slightly greater than across the talon (12.4 mm). The
four principal cusps are well developed: the protocone is the
largest, and the paracone, metacone, and hypocone are of approx-
imately equal size. The distal marginal ridge is thin and incised at
the base of the metacone by a very narrow ssure. The crista
obliqua appears to have been complete, although constricted. The
mesio-lingual and lingual aspects of the protocone preserve short,
slightly oblique ssures at the occlusal margin that represent the
remnants of the Carabelli trait. This feature would likely have been
fairly large based on the distance between the furrows (3.7 mm).
Occlusal wear is moderate. The lingual cusps have been reduced to
a at platform and the buccal cusps are rounded. The paracone,
metacone, and hypocone exhibit small dentine islands, and there is
moderate, concave exposure on the protocone. The MD diameter of
the crown is 12.5 mm. The maximum BL diameter is 13.5 mm.
The crown of M2 has a square occlusal outline, although the
distobuccal corner is very slightly reduced (Figure 3a). The BL
breadth of the trigon (13.7 mm) is somewhat greater than across
the talon (12.3 mm). The four principal cusps are present, and the
protocone is the largest, followed closely by the paracone. The
hypocone is considerably smaller than the paracone, and the met-
acone is reduced, making it the smallest cusp. The mesial marginalridge, while worn, appears to have been thick and is complete. The
thinner distal marginal ridge is complete, and encloses a fovea
posterior (talon basin) that takes the form of a narrow T-shaped
ssure, the tines of which comprise a transverse ssure distal to the
principal crests of the metacone and hypocone. The crista obliqua is
incised by a narrow furrow. There is no evidence of the presence of
the Carabelli trait. Occlusal wear is slight, with all cusps reduced
and rounded, though dentine is not exposed. The distal surface
preserves a slightly concave, moderately broad (c. 5 mm) contact
facet for the M3 (Figure 3f). The MD diameter of the crown is
12.5 mm. The maximum BL diameter is 13.8 mm.
Materials and methods
Odontometric comparisons
The MD, BL, and crown area (CA MD BL) dimensions of Bpg
2001.04, and means for these measurements in several compara-
tive samples, were used to create individual tooth prole graphs to
investigate trends in size differences of P3M2 following the
methods advocated by Rosas and Bermdez de Castro (1998). As
such, most of the comparisons involve shape using Interdental
Indices to characterize crown shape relationships (Rosas and
Bermdez de Castro, 1998; Bermdez de Castro et al., 1999) and
Average Dental Ratios (Bermdez de Castroet al., 1999). To this end,
we calculated WF distances (F) to assess phenetic afnities
following Rosas and Bermdez de Castro (1998) and Bermdez de
OH 65s AVG 9.3 12.7 9.1 13.1 12.8 13.6 13.1 14.4
Omo L894-1f AVG 8.8 12.7 9.2 12.2 12.7 13.2 11.8 12.6
KNM-ER 42703s R 9.0 12.0 8.8 12.4 12.6 13.3 13.0 14.1
*For this study, we deneHomo habilisas an eastern African early hominin with a temporal range between 1.9 and 1.6 Ma, thus A.L. 666-1 is excluded from our sample. There
are specimensfrom South Africa that have been described as Homo aff. H. habilis oras Homo erectus. At Swartkrans, SK 27 and SKX 268 have been suggested by someto belong
to a taxon with closer afnities toH. habilis(sensu stricto) than to H. erectus(e.g., Howell, 1978; Grine et al., 2009). However, for the purpose of this study, we have followed
others (e.g., Rightmire, 1990) who have included SK 27 and SKX 268 in Homo erectus. Similarly, at Sterkfontein, SE 255 derives from Member 5 Westand while it might
representH. erectus, many workers have considered it to represent the same species as Stw 53 (i.e., something with closer afnities toHomo habilis). Of course, the Member 5
deposit is not a single deposit, and it does contain material that is most likely attributable to H. erectus(e.g., the Stw 80 mandible from Member 5 West[Kuman and Clarke,2000; Moggi-Cecchi et al., 2006]). Thus, while it is not unreasonable to place SE 255 in the H. erectussample as we have done, we wish to point out that there is a controversy
over the species attribution in Sterkfontein Member 5. That is,Australopithecussp. for some of the material (Kuman and Clarke, 2000), Homoaff.H. habilis (Tobias, 1978) for
some of the material (at least that from Member 5A), and H. erectusfor some of the material (at least that from Member 5B, or Member 5 West) (Kuman and Clarke, 2000).a Grine and Franzen, 1994.b Tobias and von Koenigswald, 1964.c Wolpoff, 1971.d Dean, 2007.e Blumenberg and Lloyd, 1993.f Wood, 1991.g Jacob, 1975.h Indriati and Antn, 2008.i Arif, 1998; Arif et al., 2001.
j Weidenreich, 1937a.k Weidenreich, 1937b.l Walker and Leakey, 1993.
m Rightmire et al., 2006n Martnon-Torres et al., 2008.o Leakey et al., 1978.p Tobias, 1968.q Grine, 1989.r Boaz and Howell, 1977.s Spoor et al., 2007.
Y. Zaim et al. / Journal of Human Evolution 61 (2011) 363e376 369
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Castro et al. (1999), and analyzed the WF distances by cluster
analysis using the statistical software package NCSS (Hintze, 2001).
Only 13 specimens ofH. habilisandH. erectuspreserve the same
four teeth as Bpg 2001.04(see boldface entries inTable 1).In orderto
increase the comparative base, site samples using the mean values
foreach dimensionof eachtooth were constructed. Weincludedany
specimen that possessed at least one of the teeth preserved in Bpg
2001.04, as long as it preserved both BL and MD dimensions. Spec-imens were assigned to one of four comparative samples, eachrepresenting a different Old World hominin population by site or
region: Sangiran H. erectus (Early Pleistocene Southeast Asia);Zhoukoudian H. erectus (Middle Pleistocene Northeast Asia);
Western H. erectus (Early Pleistocene of Africa and Georgia [Dma-
nisi]), andH. habilis(Plio-Pleistocene East Africa). The comparative
sample groups range temporally from the Late Pliocene (1.9 Ma) to
the Middle Pleistocene (0.4e0.78 Ma) (Ciochon and Bettis, 2009;
Shen et al., 2009) and derive from all areas of the Old World occu-
pied by H. erectus (Kramer, 1993). Antn (2002) and Kaifu (2006)
have emphasized the need to recognize regional and temporal
variation among Asian H. erectus samples. Although there is
a decrease in mandibular post-canine tooth size during the entire
600 k.yr.of Sangiran H. erectus (Kaifu, 2006), oursample of maxillary
dentition is derived from a 300 k.yr. portion of the total H. erectustemporal range; therefore we feel justied in including all relevant
individuals for comparison in order to maximize sample size.
The goals of the metric analyses performed here are descriptive
and do not test for statistical signicance. This is because the indi-
vidual comparative samples are too small to avoid Type-II compar-ison errors. Under these conditions, descriptive results that identify
metric patterns between samples and specimens and are suggestive
of overall trends are preferable to error-prone statistical tests.
Methodology
Raw measurements for the MD, BL, and crown area
(CA MD BL) dimensions in Bpg 2001.04, and means for these
measurements in the comparative samples (Table 1), are used tocreate individual tooth prole graphs to investigate trends in size
differences of P3M2. It has been argued, however, that shape is
a more useful indicator than size when taxonomic afnity is con-
cerned (Rosas and Bermdez de Castro,1998); as such, most of the
comparisons in the present study involve shape using the followingmeasurements.
Interdental indices Interdental indices (6 in total) were con-structed using the CAs for the preserved teeth (e.g., P3/P4) to
characterize crown shape differences or similarities (Rosas and
Bermdez de Castro, 1998; Bermdez de Castro et al., 1999) of
each comparative sample and Bpg 2001.04.
Average dental ratios The relative differences in the proportions of
Bpg 2001.04 relative to the comparative samples were investigated
using
average dental ratios
(ADR) following Bermdez de Castroet al. (1999). This methodology allows for comparison of the
relative proportions of each dimension of each tooth in Bpg
2001.04 to each dimension of each tooth in the comparative
sample means. We use the following procedures. First, the raw
data are scaled relative to Bpg 2001.04 using the formula:
Bpg 2001:04Vi$2=Bpg 2001:04Vi Si
whereVi isthe rawdimensionof Bpg 2001.04(e.g., P3 BL)andSi isthe
same dimension of the comparative sample mean (e.g., P3 BL for
WesternH. erectus). The resulting value of this formula is therefore
a size ratio between the comparative sample and Bpg 2001.04. The
formula hasa valueof 1.0if theVi forBpg2001.04is equalto thevalue
of that same variable for the comparative sample (Si). If Siis larger
than that of Bpg 2001.04, the formula will yield a value of
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index of slightly less than 1.0, revealing that Bpg 2001.04 has
a slight increase in overall crown area from P3 to P4. This rela-
tionship is not shared by any of the other comparative samples,
save for H. habilis, which shows an even larger increase in crown
area. The three H. erectus samples are characterized by slight
decreases in P4 crown area, with Western and Zhoukoudian
H. erectus showing a larger decrease than Sangiran. The molar
interdental index for Bpg 2001.04 shows a slight increase in crown
area from M1 to M2; this condition is shared with the other
samples, and is seen in the most extreme with Western H. erectus
andH. habilis.
Inthe P3/M1 index, Bpg 2001.04 displays a third premolar crownarea which is roughly half the area of its M 1. This same relationship
can be seen with Sangiran H. erectus. Western H. erectus andH. habilisshare similar values with each other for this index, with
a P3 crown area roughly two-thirds the size of their M1 crown area.
Zhoukoudian is the most disparate, displaying a crown area for P 3
that is roughly three-quarters the size of its M 1 crown area.
Specimen Bpg 2001.04 has a P3/M2 index of 0.53. The Sangiran
sample has a nearly identical ratio between these two teeth, and
H. habilishas a similar value (0.58). Both the Western H. erectus and
Zhoukoudian samples have larger ratios, with Zhoukoudian having
the largest P3 crown area relative to its M2 crown area of any
sample.
The P4/M1 of Bpg 2001.04 equals its P3/M1 index. This result is
not surprising given that the premolar crown area index was nearly1.0. It shares the same value for this index with the Sangiran
H. erectussample, and the WesternH. erectussample.H. habilisandthe Zhoukoudian sample have similar index values, both of which
are larger than the Bpg 2001.04 index value.The ratios for P4/M2 show that Bpg 2001.04, the Sangiran
sample, and WesternH. erectussample are characterized by similar
relationships, that is, a fourth premolar with a crown area roughly
half the size of the crown area of the second molar. H. habilis and
the Zhoukoudian sample have a larger index for these two teeth,
with a fourth premolar roughly two-thirds the size of the second
molar.
Average dental ratios
P3 Bucco-lingually (Figure 6a), the distribution of the regional
samples relative to Bpg 2001.04 shows both the Western
H. erectusand Zhoukoudian samples falling furthest from the line
Figure 5. Individual tooth measurement (raw data) proles for Bpg 2001.04 and mean values forH. erectus(early African, Sangiran, and Zhoukoudian) andH. habilis. Fromtop left to
bottom right, they include (a) bucco-lingual length (BL), (b) mesio-distal length (MD), and (c) computed crown area.
Table 2
Interdental indices created from computed crown areas of Bpg 2001.04 and the
comparative samples.
Specimen/Sampl e P3/P4 M1/M2 P3/M1 P3/M2 P4/M1 P4/M2
Bpg 2001.04 0.99 0.98 0.54 0.53 0.55 0.54
SangiranH. erectus 1.04 0.94 0.58 0.55 0.56 0.53
ZhoukoudianH. erectus 1.09 0.98 0.72 0.70 0.66 0.64
WesternH. erectus 1.23 0.99 0.66 0.66 0.53 0.53
H. habilis 0.94 0.94 0.61 0.58 0.65 0.61
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in the lower quadrant, indicating that they are relatively larger in
this dimension than Bpg 2001.04. While H. habilis is close to the
X Y line, Sangiran H. erectus falls directly on the line, indicating
that Bpg 2001.04 and the Sangiran sample possess similar BL
proportions.
In the MD dimension (Figure 6b), all samples fall below the
X Y line, indicating that all three regional samples are pro-
portionately larger in this dimension than Bpg 2001.04. The sampleclosest to the X Y line, and therefore most similar to Bpg 2001.04,is H. habilis, followed closely by the Sangiran H. erectus sample.
ZhoukoudianH. erectusand Western H. erectusare further from theline, possessing relatively larger MD dimensions.
P4 The BL dimension of Bpg 2001.04 (Figure 6c) is relatively larger
than the comparative samples, all of which fall above the X Y line.
While the comparative samples do fall close to the line, the
Zhoukoudian sample is closest, followed by H. habilis and
Western H. erectus. Sangiran H. erectus is furthest from the line,
indicating that in this dimension, Bpg 2001.04 least resembles its
local analogs.
Bpg 2001.04s MD dimension (Figure 6d) is relatively smaller
than all the comparative samples. However, WesternH. erectusand
Sangiran H. erectus fall closest to the X Y line, indicating that
these two regional samples are generally similar to Bpg 2001.04 inthis dimension. TheH. habilissample, on the other hand, is located
far from the line, revealing that in the MD dimension, H. habilis is
considerably larger than Bpg 2001.04. ZhoukoudianH. erectusfallsbelow the line, as well, and is relatively larger in this dimension
than Bpg 2001.04.M1 Bpg 2001.04 is relatively larger than the comparative samples
in the BL dimension (Figure 7a), all of which fall above the X Y
line. Western H. erectus and H. habilis are furthest above the line,
indicating that these specimens are relatively smaller than Bpg
2001.04, with H. habilis being furthest from the line. Sangiran and
Zhoukoudian H. erectus are closer to the line than the Western
samples, with Sangiran falling almost directly on the X Y line.
The results show an Asian/African dichotomy, with the Asian
specimens having larger BL dimensions.
Like the BL relationship, the MD dimension (Figure 7b) of M1
shows that Bpg 2001.04 is relatively larger than most of the
comparative samples. Sangiran H. erectus, Western H. erectus andH. habilis all cluster tightly around the X Y line, with WesternH. erectusfalling slightly under the X Y line, and therefore being
slightly larger in this dimension to Bpg 2001.04. Zhoukoudian isconsiderably smaller in this M1 dimension than Bpg 2001.04 or any
of the other samples, falling quite far above the line in the upper
quadrant.
M2 The results for BL analysis (Figure 7c) show that Bpg 2001.04
possesses a relatively larger dimension, with the exception of
Sangiran H. erectus. All of the samples show tight clustering
around the line. Sangiran H. erectus is on the X Y line,
indicating that this population is very similar in its BL dimension
to Bpg 2001.04. The next closest sample to the X Y line is
H. habilis, followed by Zhoukoudian H. erectus, and nally,
Western H. erectus, all of which are relatively smaller than Bpg
2001.04 in the BL dimension.The analysis of the MD dimension of M2 (Figure 7d) shows
a nearly identical pattern to that of the MD dimension of M1.
Again, H. habilis, Western H. erectus, and the Sangiran H. erectus
samples cluster close to the line, with the placement of the
H. habilis and Western H. erectus indicating that they are slightlysmaller than Bpg 2001.04. Sangiran is nearly identical to Bpg
2001.04, being located just below the X Y line. Zhoukoudian falls
far above the line, and its M2 is much smaller in the MD dimension
than Bpg 2001.04.
Figure 6. Bivariate scatter-plots of the average dental ratios (ADR) versus the values of equation 1 of the bucco-lingual (BL) and mesio-distal (MD) dimensions of P 3eP4. P3 is
presented on top from left (a) bucco-lingual length, to right (b) mesio-distal length. P4 is presented on the bottom from left (c) bucco-lingual length, to right (d) mesio-distal length.
Y. Zaim et al. / Journal of Human Evolution 61 (2011) 363e376372
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hominin arrival in Sangiran commenced before1.6 Ma and lasted at
least 700 k.yr. The occupation at Zhoukoudian began at about
0.78 Ma and lasted approximately 400 k.yr. (Shen et al., 2009). The
basal occupations at Sangiran precede those of Zhoukoudian by as
much as 800 k.yr. Historically, all H. erectus fossils from Sangiran
and Zhoukoudian have been lumped together and considered the
result of a singular African exodus. Under this model, Bpg 2001.04
and the other Sangiran maxillae should be most similar to Zhou-koudianH. erectusand display lower phenetic resemblance to theWestern fossils. Our analysis, however, supports the research of
Kaifu et al. (2005a, b), which shows that the Sangiran specimensaremorphologically more similar to Western H. erectus populations
than they are to the Zhoukoudian H. erectus sample, due to the
absolutely and relatively smaller M1 and M2 dimensions of the
Zhoukoudian sample.
Given that later populations of Sangiran H. erectus have signi-
cantly smaller tooth crowns (Kaifu, 2006), similar to those seen in
the Zhoukoudian sample, it is possible that Zhoukoudian was colo-
nized by later hominins from Sangiran, or that Java experienced
a second colonization from the samepopulationas did Zhoukoudian.
Our results cannot answer that question. They do support the
hypothesis, based on the differences in tooth dimensions (particu-
larly the dissimilarity in the preserved molars of Sangiran andZhoukoudian H. erectus), that Zhoukoudian was not part of theinitial
wave of hominin dispersal that entered Java approximately 1.6 Ma.
In overall morphology of the zygomatic arch root, Bpg 2001.04and S17 share numerous features with contemporary (Early Pleis-
tocene) Western H. erectus (e.g., D2282 and D2700, SK 847, KNM-WT 15000) and the later (Middle Pleistocene) Zhoukoudian
sample (e.g., Skulls IX and XIII). However, while the Sangiran
sample shares aspects of the zygomatic arch root with the WesternH. erectusspecimens to the exclusion of the Zhoukoudian sample,
the reverse is not true.
The similarity between Bpg 2001.04 and other SangiranH. erectusmaxillary fossils is especially strong in the dentition. Bpg
2001.04 and the Sangiran H. erectusfossils cluster together in ADR
results. In only one dental dimension in the ADR results (P
4
BLdiameter) does Bpg 2001.04 cluster more with Zhoukoudian than
the Sangiran sample (Figures 6 and 7).
Similarities between Bpg 2001.04 and the Sangiran sample can
also be seen in the interdental indices (Table 2). While all H. erectus
samples possess similar interdental indices in the P 3/P4 and M1/M2
crown area ratios, in the premolar/molar indices, the Bpg 2001.04
values are most similar to those of other Sangiran fossils, wherein
the premolars are roughly half the size of the molars.
The similarity between Bpg 2001.04 and the Sangiran sample
can be seen in the WF distance values, where these two samples are
the most similar (Table 3). Bpg 2001.04 is next most similar to theWestern H. erectus sample, then the H. habilis sample, and nally
the Zhoukoudian sample is the most dissimilar to Bpg 2001.04 forthis value. The relative similarities between Bpg 2001.04 and San-
giranH. erectuswith the WesternH. erectussample is likely due totheir close temporal proximity, and the Western H. erectussample
serving as the ancestral stock for the population of Java byH. erectus.
As can be seen in Table 2, ZhoukoudianH. erectus has the largest
WF distance value from the Sangiran sample. The interdental
indices of P3 to M1 and M2 also show this separation between
ZhoukoudianH. erectus and Sangiran, with the former having a P3
crown area roughly seventy-ve percent the size of its molar crown
areas. This is, in large part, accounted for by the absolutely shorter
molars in Zhoukoudian compared to the other samples. This result
and others reecting the dissimilarity of the Zhoukoudian and
SangiranH. erectus samples can be understood in a chronological
framework.
The Sangiran sample is chronologically much closer to the
Western H. erectussample than it is to Zhoukoudian, for which the
most current age analysis suggests a range of 400e780 ka (Shen
et al., 2009; Ciochon and Bettis, 2009). With a radiometric age of
1.5 Ma, Bpg 2001.04 is chronologically closer to some of the
younger H. habilis specimens included in this study than to the
Zhoukoudian fossils. It is therefore likely that the similarities
between the Sangiran hominins and both Western H. erectus andH. habilissamples seen in the WF distances (Table 3), the WF-baseddendrogram (Figure 8), and the ADR scatter-plots (Figures 6 and 7)
reect ancestral retentions.The clustering of WesternH. erectusand the SangiranH. erectus
sample (including Bpg 2001.04) with H. habilis (and not with the
ZhoukoudianH. erectussample) is more likely due to the primitive
(basal) morphology of the rst hominin to disperse to Southeast
Asia. After all, H. habilis-like cranial and dental features are found in
the Dmanisi H. erectus population (Rightmire et al., 2006; Macaluso,
2006; Martinn-Torres et al., 2008; Margvelashvili, 2008;
Rightmire and Lordkipanidze, 2009). Most authorities now
considerH. habilisandH. erectusas sister group species (Lieberman
et al., 1996; Spoor et al., 2007). Some even consider these taxa to
represent a single chronospecies (Howell,1982; Tobias,1989,1991).
If that is the case, then the basal Western H. erectuspopulation thatrst dispersed to Southeast Asia may have retained someH. habilis-
like features in its morphology. Rightmire and Lordkipanidze
(2009: 47) reached a similar conclusion in that . the Dmanisi
hominins individuals had a habilis-like ancestor. Indeed, Homo
oresiensis, thought by many to be a descendant of the foundingH. erectusdeme in Southeast Asia, appears to retain a H. habilis-like
wrist and foot bone morphology (Tocheri et al., 2007; Jungers et al.,2009).
The west-east links and south-north contrasts evident in the
morphometric data considered here may have implications for
dispersal patterns in H. erectus. One contributing factor to the
south-north contrasts may have been an ecological barrier to
latitudinal displacement in East Asia. There are two lines of
evidence in support of this scenario of dual dispersal. First, theregions numerous hominin-like fossil teeth may, in reality,
represent a small, as yet unidentied hominoid member of the
fauna (Ciochon, 2009). Second, all undisputed mainland
H. erectus remains lie north of the Stegodon-Ailuropoda zone:
Zhoukoudian (Hebei province), Gongwangling (Shaanxi prov-
ince), Hexian (Anhui province), and Nanjing (Jiangsu province)
(Ciochon, 2010).
IfH. erectuscould not penetrate the Stegodon-Ailuropodafaunal
complex, the equatorial and north temperate populations may have
had two separate geographic origins and little genetic connection
(Ciochon, 2009, 2010). This preliminary hypothesis envisionsdispersals from two Western H. erectus populations. One held an
early H. erectus/H. habilis-like premolar-molar pattern anddispersedearlyalong a southern route to equatorial Southeast Asia:
the result we know as Sangiran (Java) H. erectus. Another pop-ulation held the more derived premolar-molar pattern and
dispersed later along a northerly route toward Northeast Asia.
These we know as Zhoukoudian H. erectus. Finally, we note the
implications of the Late Pleistocene nuclear genome extracted from
a nger bone of an archaic hominin found in Denisova Cave in
southern Siberia (Krause et al., 2010; Reich et al., 2010). While the
signicance of this discovery is not yet fully digested, the
Denisovan-Melanesian link reported by Reich et al. (2010) estab-
lishes that, by the Late Pleistocene, there were complex population
dynamics between mainland and island Southeast Asia. If one
applies the implications of this later dynamic to the earlier Pleis-
tocene, we can no longer regard H. erectus as an undifferentiated
paleodeme.
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Conclusion
Resurgence in Eurasian paleoanthropology has broadened our
knowledge ofH. erectus beyond Africa. This species, whose conti-
nent of origin remains unknown, quickly came to occupy a diverse
set of open-land resources across southern Eurasia. The site of
Sangiran represents the Early Pleistocene emplacement of
H. erectus in equatorial Southeast Asia. The Bpg 2001.04 partialmaxilla is a signicant addition to the Sangiran collection,providing a sample ofve specimens that preserve this part of the
face and dentition. Our analysis of the Sangiran maxillary dentitionsuggests that an east-dispersing H. erectus paleodeme may have
carried a H. habilis/early Western H. erectus-like premolar-molar
pattern along a southern route to the equatorial zone, while
a separate H. erectus paleodeme may have carried the more derived
premolar-molar pattern along a northerly route to the temperate
zone of China. In any event, the dental patterns revealed here
suggest distinct and separate demic origins for the only two true
(and highly disparate) H. erectus population samples in East Asia. As
additional fossil and genetic evidence accumulates for Early and
Middle Pleistocene East Asia, the H. erectus paleodeme may split
further into a number of regionally distinct, but still geographically
uid populations.
Acknowledgments
The Institute of Technology Bandung (ITB) and the University ofIowa (UI) collaborated in this research, with assistance from the
Indonesian Geological Research and Development Centre (GRDC) in
Bandung. This research was carried out under the following eld
research permits from the Indonesian Institute of Sciences and
RISTEK: 7450/V3/KS/1998, 3174/V3/KS/1999, 4301/1.3/KS/2001,
4212/SU/KU/2003, 03799/SU/KS/2006, 1718/FRP/SM/VII/08, and
04/TKPIPA/FRP/SM/IV/2010. Providingeld assistance were Johan
Arif (ITB), Suminto, Sutikno Bronto, the late Sudijono (GRDC), and
Sujatmiko (National Archaeological Research Centre, Jakarta). At UI,
James W. Rogers rened the digital graphics while Anna Watermanassisted with editing and referencing. Fieldwork funds were
provided by the L.S.B. Leakey Foundation and the following UI
sources: Center for Global and Regional Environmental Research,
Central Investment Fund for Research Enhancement, Ofce of the
Vice-President for Research, the Ofce of the Dean of the College of
Liberal Arts and Sciences, and the Human Evolution Research Fund
at the University of Iowa Foundation.
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