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models to describe the origin of modern humans are commonlyreferred to as the Replacement (or Out of Africa) Model and theMultiregional Evolution (or Regional Continuity) Model (for recentreviews published in the past decade see Pearson, 2004; Weaver

009; Conroy andodels exist, with

e modern humansEurasia admixtureleast occasionally

2004; Wu, 2004;ith the recent ge-man-Neanderthalrsement for someee Hodgson et al.,

ignicantly to thees ago a compre-

hensive volume edited by Fred Smith and Frank Spencer (1984)entitled The Origins of Modern Humans: A World Survey of the Fos-sil Evidence focused on the then state of the Old World homininpaleontological record and its contributions to the modern humanorigins debate. Follow up edited volumes appeared every few yearsafter that that also focused on the hominin fossil record orcontributed the archaeological and/or chronometric records to themodern human origins debate (e.g., Mellars and Stringer, 1989;Trinkaus, 1989; Akazawa et al., 1992; Aitken et al., 1993; Clark

* Corresponding authors.E-mail addresses: [email protected], [email protected] (C.J. Bae), wangwei@

Contents lists availab

Quaternary In

.e l

Quaternary International xxx (2014) 1e15gxmn.org (W. Wang).1. Introduction

The origin of modern humans has been one of the mostintensely debated topics in paleoanthropology over the past halfcentury. Within the origin of modern humans debate a variety oftopics have been deliberated, ranging from the hominin fossil re-cord (e.g., Howells, 1976; Thorne and Wolpoff, 1981; Wolpoff et al.,1984, 2001; Stringer and Andrews, 1988; Lahr, 1996; Trinkaus,2005; Gunz et al., 2009), to the genetics (e.g., Cann et al., 1987;Stoneking, 1993; Templeton, 1993, 2002, 2005; Relethford andHarpending, 1994, 1995; Stoneking and Krause, 2011), to thearchaeology (e.g., McBrearty and Brooks, 2000; Henshilwood andMarean, 2003; Shea, 2003, 2011; Klein, 2008). The two primary

and Roseman, 2008; Cartmill and Smith, 2Pontzer, 2012). Many variations of these two mseeming increasing support for a scenario wherrst arose in Africa, but with their dispersal intowith the indigenous populations taking place at(Brauer, 1989; Smith et al., 1989; Pearson,Trinkaus, 2005; Weaver and Roseman, 2008). Wnetic evidence of some degree of modern huadmixture (Green et al., 2010), growing endodegree of interbreeding appears to exist (but s2010).

The hominin fossil record has contributed smodern human origins question. Three decadArticle history:Available online xxx

Keywords:ChinaLate PleistoceneModern Homo sapiensTeethGeometric morphometricshttp://dx.doi.org/10.1016/j.quaint.2014.06.0511040-6182/ 2014 Elsevier Ltd and INQUA. All rights

Please cite this article in press as: Bae, C.International (2014), http://dx.doi.org/10.10a b s t r a c t

We present two previously unreported hominin permanent teeth [one right upper second molar (M2),one left lower second molar (m2)] from Lunadong (dong cave), Guangxi Zhuang AutonomousRegion, China. The teeth are important because: 1) they were found in situ; 2) at least one (M2) can becondently assigned to modern Homo sapiens, while the other (m2) is likely modern H. sapiens; and 3)the teeth can be securely dated between 126.9 1.5 ka and 70.2 1.4 ka, based on multiple MC-ICP-MSuranium-series dates of associated owstones in clear stratigraphic context. The Lunadong modernH. sapiens teeth contribute to growing evidence (e.g., Callao Cave, Huanglongdong, Zhirendong) thatmodern and/or transitional humans were likely in eastern Asia between the crucial 120e50 ka time span,a period that some researchers have suggested no hominins were present in the region.

2014 Elsevier Ltd and INQUA. All rights reserved.g College of Geographical Sciences, Nanjing Normal University, Nanjing, 210097, PR ChinaModern human teeth from Late PleistocChina)

Christopher J. Bae a, *, Wei Wang b, *, Jianxin Zhao c

Guanjun Shen g

a Department of Anthropology, University of Hawaii at Manoa, 2424 Maile Way, 346 Sb Guangxi Museum of Nationalities, Nanning, 530022, PR Chinac Radiogenic Isotope Facility, School of Earth Sciences, The University of Queensland, Brd Nanning Museum, Nanning, 530012, PR Chinae China University of Geosciences, Wuhan, 430074, PR Chinaf Tiandong County Museum, Tiandong, 531500, PR China

journal homepage: wwwreserved.

J., et al., Modern human tee16/j.quaint.2014.06.051e Luna Cave (Guangxi,

hengming Huang d, e, Feng Tian f,

ers Hall, Honolulu, HI, 96822, USA

e, Qld, 4072, Australia

le at ScienceDirect

ternational

sevier .com/locate/quaintth from Late Pleistocene Luna Cave (Guangxi, China), Quaternary

Inteand Willermet, 1997; Mellars et al., 2007). The diversity of in-terpretations of the hominin fossil record in support of one or theother model suggests that there have been variable results fromthese studies. However, there seems to be growing support thatminimally, modern humans and Neanderthals were morphologi-cally different enough to be considered distinct species (e.g., Bailey,2004; Harvati et al., 2004). The nature of modern humans and otherarchaic hominins (e.g., Neanderthals and mid-Pleistocene Homo)interactions (e.g., actual replacement or some degree of assimila-tion) remains a key area of research. Specimens assigned to mid-Pleistocene Homo are more or less the same fossils that are nor-mally included in archaic Homo sapiens (e.g., Homo antecessor,H. heidelbergensis, H. rhodesiensis, H. soloensis, etc. and excludingH. neanderthalensis and modern H. sapiens), without the nomen-clatural baggage associated with using the term archaic (forvarious discussion see Tattersall, 1986; Groves and Lahr, 1994;Rightmire, 1998, 2008; Cartmill and Smith, 2009; Bae, 2010, 2013;Xiao et al., in press).

One of the primary problems in addressing many of the ques-tions raised by evaluations of the hominin fossils, genetics, andarchaeological records to either support or refute the Replacementor Multiregional models has been the nature of the eastern Asianrecord (Trinkaus, 2005; Norton and Jin, 2009; Bae, 2010; Demeteret al., 2012; Liu et al., 2013). Traditionally, replacement hypothesisadvocates found little to no evidence of continuity in the region,while multiregionalists identied a diversity of support for theirview. Following the Early and Late Paleolithic cultural sequence asdescribed by Gao and Norton (2002), the eastern Asian LatePaleolithic hominin fossils that have received the most attentionare those derived from Zhoukoudian Upper Cave (ZKD UC). Adiversity of studies that focus on or include the ZKD UC fossils (e.g.,Weidenreich, 1938/1939;Wu,1961;Wu and Zhang,1985; Habgood,1986; Brown, 1998; Neves and Pucciarelli, 1998; Cunningham andWescott, 2002; Cunningham and Jantz, 2003) has drawn variableconclusions. For instance, some researchers concluded that the ZKDUC human fossils can be used as evidence of continuity betweenHomo erectus, Late Paleolithic humans, and modern Chinese (e.g.,Weidenreich, 1938/1939; Wu, 1961; Wolpoff et al., 1984; Wu andZhang, 1985; Wu and Poirier, 1995). The argument for multire-gional evolution at Zhoukoudian and broader China has beenhighly contested (e.g., Kamminga and Wright, 1988; Wright, 1992;Lahr, 1995, 1996; Wolpoff, 1995; Wright, 1995). More recent studiesof the ZKD UC human crania found evidence of western Eurasiancharacters being present, possibly suggestive of population move-ments west to east (e.g., Harvati, 2009). The recent analysis of thenearby Tianyuandong human skeleton also suggested modernhumans originating from northwestern Eurasia may have movedinto the region during the Late Pleistocene (Shang et al., 2007; seealso; Norton and Jin, 2009).

One type of data that has contributed to various debates inpaleoanthropology is hominin teeth (Wood and Engleman, 1988;Ungar and Teaford, 2002; Ungar, 2007; Irish and Nelson, 2008).This is particularly because teeth are generally more resistant toperi- and postmortem destruction than bones and contain atremendous amount of information (Dahlberg, 1945, 1971; Turneret al., 1991; Scott and Turner, 1988, 1997; Hillson, 2005). Of thediversity of methods used to analyze teeth, non-metric, metric, andgeometric morphometricmethods receive themost attention. Mostof these studies have focused on analyses of hominin fossils fromEurope (e.g., Bailey, 2002, 2004; Bailey and Lynch, 2005; Martinon-Torres et al., 2006, 2012; Gomez-Robles et al., 2007, 2008, 2011,2012), Africa (e.g., Wood and Engelman, 1988; Irish, 1998; Irishand Guatelli-Steinberg, 2003); and Western and Central Asia (e.g.,Bailey et al., 2008; Martinon-Torres et al., 2008; Hershkovitz et al.,

C.J. Bae et al. / Quaternary22011). Fortunately, there are a growing number of studies that focus

Please cite this article in press as: Bae, C.J., et al., Modern human teeInternational (2014), http://dx.doi.org/10.1016/j.quaint.2014.06.051on or include data from eastern Asia (e.g., Brace et al., 1984; Turner,1987, 1990; Liu et al., 2000, 2010a, b; 2013; Xing et al., 2009; Baileyand Liu, 2010; Xiao et al., in press).

Recent morphometric analyses of Chinese hominin fossil teethare contributing to a better understanding of the variability of thelateMiddle and Late Pleistocene hominin fossil record of the region.For instance, Bailey and Liu (2010) analyzed a set of maxillary teethfrom the Middle Pleistocene Chaoxian site in eastern China todetermine if the hominin fossils could be allocated to H. nean-derthalensis or whether they should be retained in the more in-clusive mid-Pleistocene Homo group. Bailey and Liu (2010: 21)concluded that no derived morphological characters link the teethof Chaoxian specically with H. neanderthalensis. Xiao et al. (inpress) recently analyzed a set of previously unreported homininteeth (and a partial mandible) from the Middle-Late PleistoceneMaba site in southern China, a site that is best known for thepresence of a partial hominin calotte that at times has beenassigned to mid-Pleistocene Homo, H. neanderthalensis, and evenH. mabaensis (for review see Wu and Poirier, 1995; Bae, 2010). Xiaoet al. (in press) concluded that the new set of fossils should beassigned to modern H. sapiens and were likely deposited at the sitemore recently. In other words, the new fossils and the Maba mid-Pleistocene Homo calotte display little to no association. Anotherrecent metric and morphometric analysis of a new set of homininteeth from the Huanglongdong site in central China drew theconclusion that these teeth should be considered modern humanbecause they look gracile and lack the archaic features usuallyidentied on Middle and Late Pleistocene humans (Liu et al.,2010a: 40). The importance of the Huanglongdong human teethis that they appear to date between 100 ka and 80 ka (Liu et al.,2010a; Shen et al., 2013), with clear implications for the modernhuman origins debate. These recent studies focusing on the Chineserecord are clearly contributing to lling in empty spaces of what hastraditionally been considered a relatively sparse hominin fossilrecord when compared to better known regions like Africa andEurope (Trinkaus, 2005; Norton and Jin, 2009; Bae, 2010; Nortonand Braun, 2010; Liu et al., 2013).

Nevertheless, two long standing problems with the easternAsian record is the paucity of evidence dating between the Middle-Late Pleistocene transition and the upper limit of radiocarbondating (i.e., between 120 and 50 ka) and the association of thehominin fossils with various relative and numerical dates (Nortonand Jin, 2009; Bae, 2010; Shen et al., 2013). Jin and Su (2000)suggested that there is a complete absence of any hominins fromChina (and presumably the rest of eastern Asia as well) between100 and 50 ka. Jin and Su (2000) subsequently used that evidence tosupport their argument that modern humans arrived in southernChina after ~50 ka and spread north soon afterwards. However, asreviewed recently by Norton and Jin (2009) and Shen et al. (2013),there are a growing number of hominin fossils being reported inChina that likely date to this important time period (e.g., Xuchang,Laishui, and see above, Huanglongdong). Furthermore, Mijares et al.(2010) recently reported a modern human 3rd metatarsal fromCallao Cave, Luzon, Philippines, that has a direct U-series date of~66.1 ka. Thus, there seems to be growing evidence that modernhumans or at least mid-Pleistocene Homo were present in easternAsia prior to ~50 ka.

Another problem that has weakened the potential importanceof the eastern Asian hominin fossil record is that questions oftenexist about the relationship between the hominin fossils and thesamples used to derive the chronometric dates (Shen et al., 2002;Norton and Jin, 2009; Bae, 2010). The importance of having asolid chronology for hominin fossils cannot be overstated. Forinstance, Shen et al. (2002) state: The accurate dating of relevant

rnational xxx (2014) 1e15nds is basic to addressing this controversial topic [modern human

th from Late Pleistocene Luna Cave (Guangxi, China), Quaternary

origins]. Besides the famous Zhoukoudian Upper Cave humansthat have debated ages ranging between ~34 and 10 ka (Kammingaand Wright, 1988; Kamminga, 1992; Wu and Poirier, 1995; Nortonand Gao, 2008; Norton and Jin, 2009), another well-knowncontroversially-dated modern human fossil from China is the Liu-jiang skull (Wu and Poirier, 1995). Although the Liujiang humanskull is often considered to date to around 20 ka and possibly 67 ka,Shen et al. (2002) conducted a U-series analyses on travertine fromthe cave and concluded the age of the Liujiang skull should date tobetween 139 and 111 ka and possibly older than 153 ka. Shen et al.(2002) concluded that the Liujiang skull could be as old as some ofthe oldest modern H. sapiens in Africa (e.g., Herto, Klasies RiverMouth, Border Cave, but not Omo Kibish). Nevertheless, questionsdo exist regarding the exact provenance of the Liujiang fossil (Shenet al., 2002; Norton and Jin, 2009). Here, we hope to partiallyremedy these two long standing problems of the eastern Asianpaleoanthropological record by presenting a set of newly excavatedand previously unreported human teeth associated with a solidchronology from Lunadong, a Late Pleistocene cave site in GuangxiZhuang Autonomous Region, southern China.

2. Background

Lunadong (dong cave) (233604800N, 106580100E) is locatedin the karst mountains of the southeastern part of the Bubing basinin Guangxi Zhuang Autonomous Region, southern China (Figs.1e2).The cave is about 30 m above the valley oor and 162 m above sealevel. Lunadong was initially found by our paleoanthropological

research team in March 2003. This cave is situated in Permianlimestone, with its southern part sloping into the karst mountainarea and the northern area facing the at Bubing valley. The caveentrance, facing east, is 1.4 m in height and 5 m in width. Theinterior of the cave is at, 3e4 m in depth, ~10 m in breadth (butpossibly extending back further), and 1e4m in height. The depositsare well preserved, about 0.5e2 m in thickness, and comprisedprimarily of light brown clay containing a few limestone breccia(Figs. 2e3).

The Natural History Museum of Guangxi Zhuang AutonomousRegion conducted the rst excavation in July 2004 and a secondexcavation in June 2008. Three 2 2 m squares were laid out in thewestern area of the cave and excavated down horizontally at 20 cmintervals. Squares A and Bwere excavated in 2004 and Square Cwasexcavated in 2008. As a result of the excavations, more than onehundred mammalian teeth and bones, two hominin teeth, and onedozen stone artifacts were recovered. Most of themammalian teethand bones, including the two hominin teeth, were unearthed fromSquare B (Figs. 2e3).

2.1. Stratigraphy

The Lunadong deposits are mainly distributed in the westernpart of the cave, with an area of about 30 m2 considered to beundisturbed. The sediment becomes thicker from east to west,varying in thickness from 10 to 130 cm. Five stratigraphic levelswere identied (from upper to lower) (Fig. 3):

C.J. Bae et al. / Quaternary International xxx (2014) 1e15 3Fig. 1. Map of Bubing Basin with locat

Please cite this article in press as: Bae, C.J., et al., Modern human teeInternational (2014), http://dx.doi.org/10.1016/j.quaint.2014.06.051ions of important cave localities.

th from Late Pleistocene Luna Cave (Guangxi, China), Quaternary

(1) Brown loose clay 0e5 cm;

ng d

Inte(2) Yellow-brown sandy clay with occasional breccia, containingstone artifacts 5e30 cm;

(3) Light brown sandy clay with breccia, yielding abundantmammalian fossils and the hominin teeth 30e100 cm;

(4) Brown sandy clay, with occasional breccia and somemammalian fossils 100e140 cm;

(5) Large breccia lled by sandy clay 140e160 cm.Fig. 2. a) North facing entrance of Lunadong. b) Interior of Lunado

C.J. Bae et al. / Quaternary42.2. Dating

The mammalian faunas identied at Lunadong include Hylo-bates sp., Macaca sp., Stegodon sp., Cervus sp., Ailuropoda melano-leuca baconi, Muntiacus sp., Sus scrofa, Sus sp., Hystrix sp., andBovidae. Because it is generally understood that Stegodon wentextinct in eastern Asia during the early Late Pleistocene (Huang,1979), the relative age of the deposits should be early LatePleistocene.

With the intent to better constrain the chronology of thestratigraphic sequence, we collected stratigraphically signicantowstone and stalagmite samples from the eastern wall of Exca-vation Square B. A total of ve speleothem samples (LN12-01-01,LN12-01-04, LN12-02, LN12-03, LN12-07), each consisting of care-fully hand-picked small fragments, were submitted to the Univer-sity of Queensland for U/Th dating using a Nu Plasma multi-collector Inductively Coupled Plasma Mass Spectrometer (MC-ICP-MS) following analytical procedures described in Zhou et al.(2011). The stratigraphic locations of these samples are presentedin Fig. 3. Among these ve samples, LN12-01-01, LN12-01-04 andLN12-02 are composed of impure calcite grains stained with clays.Their measurements show they contain elevated levels of detrital232Th (with Th/U ratios ranging from 1.4 to 3.3, similar to silicatesediments), reecting signicant silicate sediment contamination,rendering them unsuitable for dating. Samples LN12-03 and LN12-07 are much purer, with measured Th/U ratios of 0.21 and 0.17, andcorresponding 230Th/232Th activity ratios of 11.8 and 10.5. Ages

Please cite this article in press as: Bae, C.J., et al., Modern human teeInternational (2014), http://dx.doi.org/10.1016/j.quaint.2014.06.051derived from these two samples are reliable. As LN12-02 has ameasured 230Th/232Th activity ratio of 0.62, the initial 230Th/232Thratio in Lunadong cave must be < 0.62, otherwise LN12-02 willyield a negative corrected 230Th age (i.e. future age), which is un-realistic. Using this value for detrital 230Th correction, LN12-03 andLN12-07 give corrected 230Th ages of 126.9 1.5 ka and70.2 1.4 ka, which are only slightly younger than their corre-sponding uncorrected 230Th ages of 130.6 0.7 ka and 73.2 0.4 ka;the uncorrected 230Th ages being considered as the maximum ages

uring excavation. c) Exact position of the Lunadong human teeth.rnational xxx (2014) 1e15of the samples. Because the two hominin teeth were found in thehorizon slightly above LN12-03 and slightly below LN12-07, the ageof the teeth can be condently placed within the age range be-tween 126.9 1.5 ka and 70.2 1.4 ka. The biostratigraphy andnumerical dates corroborate each other nicely; the Lunadonghominin teeth date to between the end of the terminal MiddlePleistocene and the middle Late Pleistocene [between marineisotope stage (MIS) 6-5 transition and MIS 4].

3. Materials and methods

3.1. Materials (Lunadong)

The Lunadong (LND) hominin fossil assemblage is comprised ofone left upper second molar (M2) and one right lower secondmolar (m2). Both specimens are permanent teeth. These aredescribed in more detail as follows with general mesial-distal andbuccal-lingual dimensions presented inTable 2.We relied onTurneret al. (1991), Scott and Turner (1997), andWhite and Folkens (2000)for the morphological descriptions of the hominin teeth. Followingstandard paleontological practice (see author guide for the Journalof Vertebrate Paleontology), upper case M represents maxillaryteeth, while lower case m represents mandibular teeth. We esti-mated the cusp sizes. We focused our analysis on the linear metric(size variation) and geometric morphometric (shape variation) as-pects of the LND teeth. Although additional analyses (e.g., micro-CTstudy) could strengthen the evaluation of the LND hominin teeth,they are currently beyond the scope of the current investigation.

th from Late Pleistocene Luna Cave (Guangxi, China), Quaternary

Table 1MC-ICP-MS U-series isotopic data for owstone and stalagmite samples from Lunadong.

Sample name U (ppm) 232Th (ppb) (230Th/232Th) (230Th/238U) (234U/238U) Uncorrected230Th age (ka)

Corrected age-I (ka)(230Th/232Th)0 0.6

Corrected age-II (ka)(230Th/232Th)0 0.825

Corr. Initial (234U/238U)

LN12-01-01 0.09197 0.00006 207.4 0.7 0.669 0.006 0.4969 0.0043 1.1841 0.0035 58.4 0.7 8 32 Negative ge NALN12-01-04 0.10675 0.00004 153.5 0.3 0.809 0.006 0.3834 0.0030 1.1713 0.0015 42.8 0.4 13 17 Negative ge NALN12-02 0.15826 0.00007 522.8 1.9 0.617 0.003 0.6722 0.0022 1.2466 0.0013 81.9 0.4 4 59 Negative ge NALN12-03 0.11957 0.00006 25.51 0.02 11.88 0.03 0.8356 0.0021 1.1694 0.0013 130.6 0.7 126.9 1.5 125.4 2.0 1.2564 0.0073LN12-07 0.11989 0.00004 20.18 0.05 10.50 0.05 0.5825 0.0022 1.1751 0.0010 73.2 0.4 70.2 1.4 69.0 1.8 1.2230 0.0054

Note: Ratios in parentheses are activity ratios calculated from the atomic ratios. Errors are at 2s level for the least signicant digits. The ages are calculated using Isoplot 3.0 Program of Ludwig (2003) with decay constants fromCheng et al. (2000). Corrected Age-I and Age-II were calculated assuming initial/detrital 230Th/232Th activity ratio 0.6 50% (slightly lower thanmeasured 230Th/232Th activity for LN12-2) and 0.825 50% (the bulk-Earth value,which is the most commonly used), respectively. For Luna Cave samples, Corrected Age-I should be a better estimate, because LN12-01-1, LN12-01-4, LN12-2 cannot have negative ages, as is the case if the bulk-Earth initial/detrital 230Th/232Th activity ratio is used for correction. Only LN12-03 and LN12-07 are pure enough to return reliable age estimates. The other three samples are too dirty and severely altered, not suitable for dating. See text fordiscussion.

Table 2Linear metric data of M2 and m2 teeth (see text for references). Measurements given in mm.

M2 m2

Specimen/population Buccal-lingual (s.d.) Mesial-distal (s.d.) Specimen/population Buccal-lingual (s.d.) Mesial-distal (s.d.)

Lunadong (LN0031) 11.83 10.65 Lunadong (LN0030) 11.11 12.05

A. afarensis (n 8) 14.64 (0.76) 13.11 (0.70) A. afarensis (n 21) 13.39 (1.10) 14.03 (1.28)A. africanus (n 11) 15.60 (0.76) 13.80 (0.71) A. africanus (n 5) 14.10 (0.87) 15.22 (1.00)P. robustus (n 12) 15.84 (0.97) 14.73 (0.53) P. robustus (n 10) 14.90 (0.79) 16.23 (0.85)P. boisei (n 4) 18.38 (1.87) 16.05 (1.03) P. boisei (n 10) 16.80 (1.56) 18.51 (1.82)Early Homo (n 6) 14.82 (1.71) 12.77 (1.03) Early Homo (n 6) 13.53 (1.07) 15.08 (0.98)H. erectus (n 14) 13.09 (0.95) 11.86 (1.10) H. erectus (n 32) 12.55 (0.95) 13.30 (1.01)Neanderthal (n 17) 12.83 (0.85) 10.82 (0.89) Neanderthal (n 20) 11.29 (0.82) 12.34 (0.93)Middle Paleolithic Modern Human (n 3) 12.60 (0.40) 10.93 (0.75) Middle Paleolithic Modern Human (n 3) 11.53 (0.25) 11.27 (0.67)Upper Paleolithic Modern Human (n 128) 12.30 (0.77) 10.05 (0.79) Upper Paleolithic Modern Human (n 155) 10.97 (0.91) 11.13 (0.82)Mesolithic Modern Human (n 206) 11.81 (0.78) 9.62 (0.64) Mesolithic Modern Human (n 198) 10.49 (0.59) 10.65 (0.65)Neolithic Modern Human (n 148) 10.92 (0.88) 8.99 (0.61) Neolithic Modern Human (n 160) 9.79 (0.60) 10.11 (0.65)

C.J.Baeet

al./Quaternary

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(2014)1e15

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Pleasecite

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as:Bae,

C.J.,et

al.,Modern

human

teethfrom

LatePleistocene

LunaCave

(Guangxi,

China),Quaternary

International(2014),http://dx.doi.org/10.1016/j.quaint.2014.06.051

InteC.J. Bae et al. / Quaternary63.1.1. M2The upper left second molar (LN0031, Fig. 4) is milky white in

color and generally well preserved. The crown is well preserved,most of the neck is present, but the roots are absent. The crown ofLN0031 has four well-developed cusps arranged in a rhombicshape, with the occlusal surface slightly worn. The four cusps areprotocone > paracone >metacone > hypocone. The protocone andparacone are distinctly larger than the metacone and hypocone.The protocone is connected to the paracone by a gentle valley,separated from the hypocone with a ne lingual groove, and islinked to the metacone by a small ridge. The paracone is markedlyconvex mesially and buccally and is separated from the metaconeby a buccal groove. The paracone extends in the mesial-buccal di-rection. Given the wear pattern, a chip on the distal-lingual side ofthe metacone probably occurred premortem (during chewing?)rather than postmortem.

Fig. 3. Plan and section map. Locations of speleothem and hominin fossils given (LN04-07;bottom prole represent breaks in the stratigraphy.

Please cite this article in press as: Bae, C.J., et al., Modern human teeInternational (2014), http://dx.doi.org/10.1016/j.quaint.2014.06.051rnational xxx (2014) 1e15The lingual surface is rectangular, convex vertically and hori-zontally, with the lingual groove narrow and terminating in themiddle. The mesial side is rectangular, with a fairly pronouncedinterproximal contact facet present that occupies half of the mesialsurface. A transverse elliptical interproximal contact facet is pre-sent on the distal surface. The roots aremostly absent, with only thebasal portion remaining. Evidence of rodent gnawing marks ispresent; thus, the roots were likely chewed off.

3.1.2. m2The lower right second molar (LN0030, Fig. 4) is milky white in

color and heavily fossilized. However, the crown is well preserved.In occlusal view, the crown outline is square, with a fair degree ofwear on the four intact cusps (metaconid, protoconid, entoconid,and hypoconid). The protoconid is the largest cusp. The entoconidand hypoconid are the smallest, but similar in size. A hypoconulid is

LN12-01; LN12-02; LN12-03; LN0030; LN0031). The unbroken horizontal lines in the

th from Late Pleistocene Luna Cave (Guangxi, China), Quaternary

absent. The buccal and lingual surfaces are slightly verticallyconvex, while the distal surface is vertical and horizontally convex.The mesial surface is at due to heavy interproximal wear (a largeinterproximal contact facet is present at the upper edge of themesial surface). The buccal and lingual grooves are short and weakbecause of wear. On the distal surface there is a smaller ovalinterproximal contact facet. The roots, relatively slender, divide intotwo branches, mesially and distally. The buccal-lingual dimensionof the mesial branch is markedly larger than the same dimension ofthe distal branch.

3.2. Materials (comparative)

The comparative linear data for the mesial-distal and buccal-lingual measurements include Neolithic, Mesolithic, Upper Paleo-lithic, and Middle Paleolithic modern humans and Neanderthals.These data are from Voisin et al. (2012), while data from olderhominins (e.g., H. erectus, H. habilis, gracile and robust Austral-opiths) were culled from published literature (Robinson, 1956;Tobias, 1967; Wood, 1991; Wu and Poirier, 1995; Kimbel et al.,2004; Kaifu et al., 2005; Bailey and Liu, 2010; Liu et al., 2010a;

of this effect on the analysis of crania), these specimens were alsoleft out of the analyses. Although we admit that there may be somedegree of error with using photographs of hominin teeth taken byothers, the comparative database was created without any notice-able errors (see also Xiao et al., in press).

3.3. Methods

We took the linear measurement data using Mitutoyo digitalcalipers (model no. CD-20C). The mesial-distal and buccal-lingualmeasurements were taken at their widest points. The linear mea-surement data were graphed on bivariate scatterplots. In order toevaluate the degree of overlap among the different populations,convex hulls were created. Because of the large comparative sam-ples of modern humans (Neolithic, Mesolithic, Upper Paleolithic),only the outline of the convex hulls are graphed for those respectivepopulations. For the older hominins, each individual datapoint isgraphed.

We collected digital images of the teeth occlusal surfaces using a

C.J. Bae et al. / Quaternary InteMacaluso, 2010, Table 2).We conducted a two dimensional geometric morphometric

(GM) analysis of the tooth occlusal surfaces on a fairly robustcollection of hominin right M2s (N 113) and left m2s (N 87)(Tables 3a, b). The comparative samples for the GM analyses arefrom photographs of various hominins (e.g., Paranthropus, Austral-opithecus afarensis, Au. africanus, H. habilis, H. erectus, mid-Pleistocene Homo, H. neanderthalensis, etc.) contributed primarilyby Eric Delson (ED) and Erik Trinkaus (ET) and their colleagues.In discussions with ED and ET, the images they provided wereproduced using fairly standard photographing methods in paleo-anthropology (see also Xiao et al., in press). Other hominin fossilteeth derive from publications that include high resolution imagesof tooth occlusal surfaces (e.g., Walker et al., 2008; Martinon-Torreset al., 2012).When possible, we collapsed various hominin taxa intomore inclusive categories in order to increase sample size withinthat particular group (e.g., Au. afarensis and Au. africanus weregrouped together, and H. antecessor, H. heidelbergensis, and archaicH. sapiens were grouped into the more inclusive category of mid-Pleistocene Homo).

Table 3aM2s used in this study (n 114).

Genus/species Specimens

Australopithecus afarensis(n 5)

AL 199-1; AL 200; AL 333-x; AL 417-1; AL 442-2

Australopithecus africanus(n 2)

Sts 52a; Stw 252

Homo habilis s.l. (n 4) ER 1813; OH 13, OH 16, OH 65Homo erectus s.l. (n 6) Dmanisi D2700, Dmanisi D2882; KNM WT 15000;

Sangiran 4; Isolated Trinil; ZhoukoudianMid-Pleistocene Homo

(n 22)Arago 14, 21; AT 12, 15, 46, 170, 407, 588, 815, 817,821, 824, 960, 2175, 2179, 4319, 4326, 4336;Chaoxian; Gran Dolina TD 6; Petralona; Steinhem

Homo neanderthalensis(n 12)

Amud; Arcy sur Cure 9; Feldhofer NN 16;Monsempron 3; Moustier 1; Quina 5; Saccopastore1; Shanidar 1, 2; Sima Palomas 36; St. Cesaire 1;Tabun 1

Early modern human(n 6)

Mumba XXI; Qafzeh 6, 7, 9, 11; Skhul 5

Upper Paleolithic modernhuman (n 9)

Brassempouy 542; Dolni Vestonice 13, 14; Kostenki3, 4; Mladec 2, 8; Oase 2; Ohalo 2

Modern human (n 47) Chinese Neolithic (n 11); Modern Koreans(n 36)Lunadong (n 1)

Please cite this article in press as: Bae, C.J., et al., Modern human teeInternational (2014), http://dx.doi.org/10.1016/j.quaint.2014.06.051The modern human teeth used in the GM analyses are sampleswewere able to study directly. Themodern human samples includea small skeletal collection fromDushandong, a Neolithic burial cavesite from Guangxi, China and excavated by some of the coauthors.The modern Korean samples are from Chosun Dynasty (1392e1897A.D.) cemeteries, housed and curated at Hanyang University andChungbuk University, and studied with the permission of KidongBae and Sunjoo Park, the principal curators of those respectivecollections. In cases (for both hominins and modern humans)where a right M2 or a left m2was absent, but a left M2 or a right m2was present, we mirror imaged the latter specimens. In no instancedid we knowingly use two maxillary or two mandibular teeth fromthe same individual.

In general, we only used high resolution tooth images that hadfour cusps and intact occlusal surface outlines (see also Xiao et al.,in press). Following many previous studies (e.g., Martinon-Torreset al., 2006; Gomez-Robles et al., 2007, 2008, 2011; Bailey andLiu, 2010; Xiao et al., in press), we excluded samples for a num-ber of reasons. For instance, if the tooth was too worn or chippedand we could not easily locate the cusps and/or the outer edge ofthe occlusal surface, the tooth was not included in the subsequentanalyses. Deciduous teeth and incomplete teeth were excluded. Ifthe photograph was too grainy or of too low resolution, thatparticular specimen was also excluded from the study. Because ithas been shown that photographs taken at an angle can skew theoverall shape of the object (e.g., see Gharaibeh, 2005 for discussion

Table 3bm2s used in this study (n 87).

Genus/species Specimens

Paranthropus (n 5) L62e17, L157e35, L427-7; OMO 47-46; PeninjHomo habilis (n 1) OH 13Homo erectus s.l. (n 9) Dmanisi 211, D2735; Meganthropus; Sangiran

B; Sangiran 'Pith dubius'; Ternine 1, 2, 3; TrinilMid-Pleistocene Homo

(n 23)AT-1, 142, 169, 271, 273, 284, 300, 557, 607,941, 946, 1761, 2193, 2270, 2272, 2396, 2763,3179, 4147; ATD6-96; Arago 2, 13; Mauer

Homo neanderthalensis(n 11)

Amud 1; Arcy-sur-Cure 5, 21; Krapina D, E, H, L;Sima Palomas 29, 59, 80; Tabun 1

Early modern human(n 2)

Mumba XXI; Qafzeh 9

Upper Paleolithic modernhuman (n 1)

Dolni Vestonice 3

Modern human (n 34) Chinese Neolithic (n 6); Modern Koreans(n 28)

Lunadong (n 1)

rnational xxx (2014) 1e15 7Dino-Lite Digital Microscope [model AM413T Dino-Lite Pro

th from Late Pleistocene Luna Cave (Guangxi, China), Quaternary

Catalyst and Base).

from

Inte3.4. Geometric morphometrics

Geometric (landmark based) morphometrics (GM) methodolo-gies is widely used to explore within- and between-taxa variationin paleoanthropology (for recent reviews of applications toanthropological research see: O'Higgins, 2000; Richtsmeier et al.,2002; Slice, 2005, 2007; Perez et al., 2006; Baab et al., 2012). TheGM methodology employed here for the analysis of the Lunadonghominin teeth follows closely previous studies (e.g., Martinon-Torres et al., 2006; Perez et al., 2006; Gomez-Robles et al., 2007,(20e230 magnication)] with stand. The methodology followsclosely many other recent dental studies (e.g., Bailey, 2002;Martinon-Torres et al., 2006; Gomez-Robles et al., 2007, 2008,2011, 2012; Bailey et al., 2008; Bailey and Liu, 2010), except thatinstead of using a digital camera andmacrolens, we used the digitalmicroscope. In general, we found it easier to identify the toothcusps and general outline of the tooth occlusal surface from theimages produced by the Dino-Lite (see also Xiao et al., in press). Inorder to help hold the tooth level and parallel to the Dino-Lite, weused polyvinylsiloxane impression material (Exane Putty Type

Fig. 4. Two human teeth from Lunadong: upper (LN0031); lower (LN0030). Views

C.J. Bae et al. / Quaternary82008, 2011, 2012; Xing et al., 2009; Liu et al., 2010a, 2013; Xiaoet al., In Press). Prior to beginning the digitizing process, eachtooth was oriented in the same direction with the mesial side to-ward the top of the computer screen and the buccal side to the right(see Fig. 5 for example). The programMakeFan7 (Sheets, 2001) wasused to create the centroid necessary to generate a series of equi-angular fans. In order to create the centroid, four cusps on eachtooth was digitized on the occlusal surface of the M2s [protocone(red), paracone (blue), metacone (green), hypocone (yellow)] andm2s [metaconid (red); protoconid (blue); hypoconid (green);entoconid (yellow)]. Because MakeFan7 generates a centroid fromfour datapoints, in cases where a fth cusp may have been presentit was not marked and included in the analyses (e.g., m2's hypo-conulid). As described above, if the specimen did not have fourcusps (e.g., M2's hypocone was absent), the specimenwas excludedfrom the analysis. As with previous studies (e.g., Martinon-Torreset al., 2006; Gomez-Robles et al., 2007, 2008, 2011; Xiao et al., inpress), the center of the dentinal facet of each cusp was chosen asthe landmark location. If no dentinal facet was present, the highestpoint of the cusp was utilized. Once the centroid was calculated, 30equiangular fan lines were created in MakeFan7 with an exagger-ation of 3 to ensure that the fan lines crossed the edge of the tooth

Please cite this article in press as: Bae, C.J., et al., Modern human teeInternational (2014), http://dx.doi.org/10.1016/j.quaint.2014.06.051occlusal surface. Then, using TpsDig2, version 2.16 (Rohlf, 2010a),each of the 30 equiangular fan lines were digitized at the pointwhere the fan line crossed the edge of the occlusal surface. Thepoints were always digitized in the same clockwise order, with therst point digitized always the fan line that ran through the redlandmark which occurred in every case (M2: protocone; m2:metaconid). In order tominimize the bending energy between eachlandmark and target form (Bookstein, 1997; Gunz et al., 2005) eachof the 30 semilandmarks were then slid using TpsUtil and tpsRelw(Rohlf, 2010b, c). A Generalized Procrustes Analysis (GPA) was thenperformed in tpsRelw (Rohlf, 2010c) in order to remove the po-tential effects of translation, rotation, and scaling. The resultantshapes were then analyzed using the Relative Warps Analysisfunction in tpsRelw. In this context, the relative warp output is thesame as the principal component output. TPS-grids were alsoevaluated to better observe the degree of variation at the edges ofthe different principal components (Bookstein,1997). The TPS-gridsand principal components plots were generated in tpsRelw andMicrosoft Excel (for previous examples see Martinon-Torres et al.,2006; Gomez-Robles et al., 2007, 2008, 2011, 2012; Xiao et al., InPress). Minimum spanning trees were created using the programPAST (Hammer et al., 2001) to determine the closest links betweenthe Lunadong teeth and the comparative samples.

left to right: occlusal view; mesial view; distal view; buccal view; lingual view.

rnational xxx (2014) 1e154. Results

4.1. M2

4.1.1. Size variationFig. 6 plots the mesial-distal and buccal-lingual measurements

of the upper second molars. In general, the older hominins havemuch larger teeth than the Holocene modern human samples andeach group tends to cluster together. Homo erectus andH. neanderthalensis fall in between, overlapping somewhat at theupper range of theMP and UPmodernH. sapiens and at the low endof the range of the Australopiths and early Homo populations. TheLND M2 falls squarely in the middle of the Mesolithic and UPmodern humans, at the upper end of the range of Neolithic modernhumans, and at the low end of the range of H. neanderthalensis.

4.1.2. Shape variationResults were obtained using the slid semilandmark congura-

tions (tpsRelw). The results of the principal components analysisare presented in Figs. 7e8, with the rst ve principal components,

th from Late Pleistocene Luna Cave (Guangxi, China), Quaternary

totaling 81.40% of the variation, presented in Table 4; here andbelow only the principal components that can explain at least 5% ofthe variation are offered. The rst three principal components total65.80% of the explained variance (PC1 28.95%; PC2 20.83%;PC3 16.02%). Because a great deal of overlap exists between thedifferent hominin fossil samples, we evaluated two separate prin-cipal components charts [PC1-PC2 (Fig. 7), PC1-PC3 (Fig. 8)].

We evaluated the variation in the morphology of the toothocclusal surface at the extreme edges of the buccal, lingual, mesial,and distal sides using the thin plate spline function in tpsRelw(Figs. 7e8). On the negative end of PC1 the occlusal surface isroughly rectangular shaped, while at the positive extreme of PC1 itis more squarish. The occlusal surfaces at the extremes of PC2 arerectangular. However, the negative end is longer on the mesial anddistal sides, while the positive end has longer buccal and lingualsides. The extreme ends of PC3 are similar to those of PC2, with

Table 4Principal components of M2s and m2s (only PCs that explain >5% of the variationpresented).

Principal component M2 m2

1 28.95% 35.38%2 20.83% 27.64%3 16.02% 12.75%4 10.08% 5.77%5 5.52%

C.J. Bae et al. / Quaternary InteFig. 5. Example of the location of the marked cusps [metaconid (red); protoconid(blue); hypoconid (green); entoconid (pink)] on the LND m2 used to create the 30equiangular landmarks around the edge of the tooth occlusal surface that were digi-both extremes rectangular and longer on themesial and distal sides(negative end) or buccal and lingual sides (positive end).

In Fig. 7 (PC1 vs. PC2), there appears to be substantial overlapamong the modern human and hominin samples. In general, mostof the older hominins (e.g., Australopiths, H. habilis, H. erectus) falltized using MakeFan7 and tpsDig2. (For interpretation of the references to colour inthis gure legend, the reader is referred to the web version of this article.)

Please cite this article in press as: Bae, C.J., et al., Modern human teeInternational (2014), http://dx.doi.org/10.1016/j.quaint.2014.06.051on the negative side of PC1 (the exception is OH 16). Mid-Pleistocene Homo and the Neanderthals display some degree ofseparation, with the former plotting on the positive side of PC2 andthe majority of the latter specimens falling in the negative side ofPC2. Modern humans (early, UP, and recent) generally fall awayfrom the older hominins and overlap fairly extensively with mid-Pleistocene Homo and the Neanderthals. The LND M2 plots on thepositive sides of PC1 and PC2 falling most closely to several RecentModern Humans (Koreans), an Early Modern Human (Mumba XXI),and a Neanderthal (Amud). The minimum spanning tree connectsLND M2 most closely to a Recent Modern Human (Korean).

The PC1 vs. PC3 chart (Fig. 8) presents a slightly clearer pattern.For instance, in Fig. 7 the mid-Pleistocene Homo and Neanderthalsspecimens display a great deal of variation, plotting in all fourquadrants. However, in Fig. 8 the same specimens are much moretightly grouped, with the majority of them falling on the positiveside of PC3 and displaying a good deal of overlap with each other. Inthe PC1 vs. PC3 chart, LND M2 falls closest to an Early ModernHuman (Mumba XXI) and two Recent Modern Humans (Koreans).Not surprisingly, minimum spanning trees link LND M2 directlywith those specimens.

4.2. m2

4.2.1. Size variationFig. 9 plots the mesial-distal and buccal-lingual measurements

of the lower second molars. In general, the older hominins havelarger teeth, particularly the robust Australopiths, and each grouptends to cluster together. Although the older hominins generallyhave larger teeth, there does appear to be a substantial degree ofoverlap between the different populations. As with the M2s, the H.erectus and H. neanderthalensis populations generally fall in be-tween, overlapping with the Mesolithic and Neolithic modernhumans (falling at the upper range of those populations) and theolder hominins (falling at the lower range of those populations).The LNDm2 falls squarely in themiddle of the Neanderthals and UPmodern human convex hulls.

4.2.2. Shape variationResults were obtained using the slid semilandmark congura-

tions (tpsRelw). The results of the principal components analysisare presented in Figs. 10e11, with the rst four principal compo-nents, totaling 81.54% of the variation, presented in Table 4. The rstthree principal components total 75.77% of the explained variance(PC1 35.38%; PC2 27.64%; PC3 12.75%). We present twoseparate principal components graphs [PC1-PC2 (Fig. 10), PC1-PC3(Fig. 11)].

We evaluated the variation in the morphology of the toothocclusal surface at the extreme edges of the buccal, lingual, mesial,and distal sides using the thin plate spline function in tpsRelw(Figs. 10e11). Overall, when moving from one extreme to the other(e.g., moving from the positive extreme of PC1 to the negativeextreme) the morphology of the m2 occlusal surfaces does notpresent as much variation as the M2 (described above) does.

A great deal of overlap exists between the different populationswhen comparing PC1 vs. PC2 (Fig. 10). Only the mid-PleistoceneHomo specimens separate somewhat from the rest of the data-points. LND m2 falls very close to the zero center of PC1 and on theslightly positive side of PC2 and almost squarely on a RecentModern Human (Korean). Not surprisingly, the minimum spanningtree indicates LND m2 is most closely linked to two Recent ModernHumans (Koreans).

Compared to the PC1-PC2 chart, the PC1-PC3 chart (Fig. 11)presents somewhat overall clearer separation among the different

rnational xxx (2014) 1e15 9hominin populations. For instance, almost all of the Neanderthals

th from Late Pleistocene Luna Cave (Guangxi, China), Quaternary

Lunadong

7

9

11

13

15

17

19

21

23

8 10 12 14 16 18 20

Mesial-D

istal

Buccal-Lingual

M2s

A. afarensis

A. africanus

P. robustus

P. boisei

H. erectus

Neanderthal

MP modern human

UP modern human

Mesolithic modern human

Neolithic modern human

Lunadong

Early Homo

Fig. 6. Scatterplot of mesial-distal length and buccal-lingual width for Lunadong and other M2s. Raw data presented in Table 3a (see text for references). Measurements in mm.

-0.08

-0.06

-0.04

-0.02

0

0.02

0.04

0.06

-0.14 -0.12 -0.1 -0.08 -0.06 -0.04 -0.02 0 0.02 0.04 0.06 0.08

A. afarensis/ africanusH. habilisH. erectusmid-Pleistocene Homo

Neanderthal

Early modern human

UP modern human

Recent modern human

Lunadong

PC1: 28.95%

PC2:

20.

83%

Fig. 7. Principal components plot (PC1-PC2) of LND M2 and other hominin M2s. Minimum spanning tree shows links to LND M2 as a solid black line. For comparative purposes, thebranches between all of the other hominins are presented as light lines.

C.J. Bae et al. / Quaternary International xxx (2014) 1e1510

Please cite this article in press as: Bae, C.J., et al., Modern human teeth from Late Pleistocene Luna Cave (Guangxi, China), QuaternaryInternational (2014), http://dx.doi.org/10.1016/j.quaint.2014.06.051

Fig. 8. Principal components plot (PC1-PC3) of LND M2 and other hominin M2s. Minimum spanning tree shows links to LND M2 as a solid black line. For comparative purposes, thebranches between all of the other hominins are presented as light lines.

Fig. 9. Scatterplot of mesial-distal length and buccal-lingual width for Lunadong and other m2s. Raw data presented in Table 3b (see text for references). Measurements in mm.

C.J. Bae et al. / Quaternary International xxx (2014) 1e15 11

Please cite this article in press as: Bae, C.J., et al., Modern human teeth from Late Pleistocene Luna Cave (Guangxi, China), QuaternaryInternational (2014), http://dx.doi.org/10.1016/j.quaint.2014.06.051

Fig. 10. Principal components plot (PC1-PC2) of LND m2 and other hominin m2s. Minimum spanning tree shows links to LNDm2 as a solid black line. For comparative purposes, thebranches between all of the other hominins are presented as light lines.

Fig. 11. Principal components plot (PC1-PC3) of LND m2 and other hominin m2s. Minimum spanning tree shows links to LND m2 as a solid black line. For comparative purposes, thebranches between all of the other hominins are presented as light lines.

C.J. Bae et al. / Quaternary International xxx (2014) 1e1512

Please cite this article in press as: Bae, C.J., et al., Modern human teeth from Late Pleistocene Luna Cave (Guangxi, China), QuaternaryInternational (2014), http://dx.doi.org/10.1016/j.quaint.2014.06.051

Brauer, G., 1989. The evolution of modern humans: a comparison of the African and

Inteplot on the negative side of PC3, while most H. erectus plot on thepositive side. The majority of the Recent Modern Humans fall onthe positive side of PC3, while most of the mid-Pleistocene Homospecimens fall on the negative side. When compared to othermembers of their group, the only apparent outlier is one H. erectusspecimen (Dmanisi D2735), which falls far away from the rest of theH. erectus specimens. LNDm2 falls very close to the zeroezero pointfor both PC1 and PC3. However, unlike PC1-PC2 where LND m2plotted almost directly on a RecentModern Human, PC1-PC3 showsthat the closest datapoints to LND m2 are a H. erectus (Ternine 3)and a mid-Pleistocene Homo (AT 4147). Not surprisingly, the min-imum spanning trees links LND m2 to Ternine 3 and AT 4147.

5. Discussion

LND M2 metrically falls closest to the Mesolithic and Neolithicmodern humans, though it does fall at the low end of the range ofthe Neanderthals. The LND M2 falls outside the range of the olderhominin populations. In terms of the geometric morphometrics,LND M2 plots closely with Recent Modern Humans (Koreans in allcases) and an Early Modern Human (Mumba XXI) on both the PC1-PC2 (Fig. 7) and PC1-PC3 (Fig. 8) plots. The Minimum SpanningTrees for both plots indicate the closest relationship for LND M2 isto Chosun Dynasty Koreans and Mumba XXI, corroborating the sizeanalysis.

In general, the results for the LNDm2 are not as clear as they arefor the LND M2. There is clear distinction metrically between theolder and younger hominin populations' m2s (with the H. erectusand Neanderthal populations falling in between). The LNDm2 doesfall most comfortably within the UP and Mesolithic modernH. sapiens convex hulls. In reference to the geometric morpho-metrics, the PC1-PC2 plot (Fig. 10) and the PC1-PC3 plot (Fig. 11)present contrasting results. In the PC1-PC2 plot LNDm2 falls almostdirectly on a Recent Modern Human, while in the PC1-PC3 plot LNDm2 falls closest to Ternine 3 (H. erectus) and AT 4147 (mid-Pleis-tocene Homo). The minimum spanning trees clearly link LNDm2 toa Recent Modern Human (Fig. 10), but also to Ternine 3 and AT4147 (Fig. 11).

Overall, the metric and GM analyses of the LND M2 suggest itmost closely aligns with modern H. sapiens. Although the results ofthe LND m2 are less clear, the metric and geometric morphometricanalyses suggests likely afliation with modern H. sapiens, Theabsence of a hypoconulid and relatively slender roots also suggestLND m2 represents a modern human. Nevertheless, given therelatively robust sample sizes utilized in this study, it is possible them2 is more difcult to distinguish taxonomically than is the M2.

6. Conclusion

The Lunadong modern human M2 and the probable modernhuman m2 found in situ, given their chronometric age, haveimportant implications to the modern human origins debate,particularly because some researchers (e.g., Jin and Su, 2000) havesuggested there is a complete or nearly complete absence ofhominin occupation in China during the rst half of the LatePleistocene. Simply stated, the Lunadong human teeth add to agrowing list of modern and/or transitional humans that appear inthis spatio-temporal point (e.g., Callao Cave, Huanglongdong,Zhirendong). The Lunadong human fossils can contribute to variousaspects of the modern human origins debate.

Acknowledgements

Funding for this research was provided by the Wenner-Gren

C.J. Bae et al. / QuaternaryFoundation (ICRG #82) and the National Geographic Society

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Demeter, F., Shackleford, L.L., Bacon, A.M., Duringer, P., Westaway, K.,(#8372-07). We are grateful to Eric Delson and Erik Trinkaus forallowing us to measure and analyze the hominin fossil photographsthey generously provided and to Kidong Bae and Sunjoo Park forpermission to study the modern human skeletal collections undertheir care at Hanyang University and Chungbuk University in Korea.We thank Youngsun Shin for taking photographs of the ChosunDynasty modern human dental occlusal surfaces stored at Chung-buk University, and Maureen Moft, Kitae Park, and Robert Benitezfor help with putting the gures and tables together. An earlierversion of this paper was presented at the 2010 American Associ-ation of Physical Anthropologists (AAPA) meeting in Albuquerque,New Mexico. We acknowledge the University of Hawaii UniversityResearch Council and the Center for Chinese Studies Chung-fongand Grace Ning Chinese Studies Travel Awards provided to CJB topresent this paper at the AAPA meeting. We appreciate the com-ments from the anonymous reviewers that helped to tighten up themanuscript. We take full responsibility for any errors that may bepresent.

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Modern human teeth from Late Pleistocene Luna Cave (Guangxi, China)1 Introduction2 Background2.1 Stratigraphy2.2 Dating

3 Materials and methods3.1 Materials (Lunadong)3.1.1 M23.1.2 m2

3.2 Materials (comparative)3.3 Methods3.4 Geometric morphometrics

4 Results4.1 M24.1.1 Size variation4.1.2 Shape variation

4.2 m24.2.1 Size variation4.2.2 Shape variation

5 Discussion6 ConclusionAcknowledgementsReferences