1.5 million homo erectus jaw sangiran 4

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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.

Y. Zaim et al. / Journal of Human Evolution 61 (2011) 363e376364


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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.

Y. Zaim et al. / Journal of Human Evolution 61 (2011) 363e376 365


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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.



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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.



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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.



13/14

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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1.5 million homo erectus jaw sangiran 4

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