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Newly recognized Pleistocene human teeth from Tabun Cave, Israel
Transcript of Newly recognized Pleistocene human teeth from Tabun Cave, Israel
Journal of Human Evolution 49 (2005) 301e315
Newly recognized Pleistocene human teethfrom Tabun Cave, Israel
Alfredo Coppa a,*, Rainer Grun b,c, Chris Stringer d,Stephen Eggins e, Rita Vargiu a
a Department of Animal and Human Biology, Section of Anthropology, University of Rome ‘‘La Sapienza’’,
Piazzale Aldo Moro, 5, 00185 Rome, Italyb Research School of Earth Sciences, The Australian National University, Canberra, ACT 0200, Australia
c Research School of Pacific and Asian Studies, The Australian National University, Canberra, ACT 0200, Australiad Department of Palaeontology, The Natural History Museum, London SW7 5BD, UK
e Research School of Earth Sciences, The Australian National University, Canberra, ACT 0200, Australia
Received 21 May 2004; accepted 20 April 2005
Abstract
Seven human teeth from Tabun Cave, Israel, curated at the Natural History Museum London since 1955, are ofuncertain provenance and identity. They are all from the upper dentition, without duplications, and are characterizedby a similar preservation. The Catalogue of Fossil Hominids (1975) suggested that they might have derived from Tabun
Layer A (Bronze Age to Recent). However, one of us (AC) noted some distinctive features of these teeth that warrantedfurther study. They are here assigned to a single individual, Tabun BC7. Their morphology and metrics were thencompared with the frequency of Late Pleistocene and Early Holocene groups from Europe, North Africa and Middle
East.A fragment of the right M3 crown of Tabun BC7 was removed for ESR and U-analysis, and it was determined that
only samples from Layer B have similar dose values. Using the sediment dose values of layer B, preliminary ageestimates of 82G 14 ka (early U-uptake) and 92G 18 ka (linear uptake) were obtained. U-series disequilibrium
determined from other samples attributed to Layer B resulted in a U-uptake history close to linear uptake, givinga very comparable age estimate of 90C30
�16 ka. The dose value previously obtained on an enamel fragment from theTabun C1 dentition is nearly double the value measured for BC7, and tentative age estimates for C1 were in the range
of 143G 37 ka. However, due to uncertainties in the exact provenance of the human fossils, we cannot confirm that C1is older than the new tooth sampled here, and both C1 and BC7 can be attributed to Layer B on chronologicalgrounds.
* Corresponding author. Tel.: C39 6 49912252 (Laboratory), C39 6 49912350 (Office); fax: C39 6 49912771.
E-mail addresses: [email protected] (A. Coppa), [email protected] (R. Grun), [email protected] (C. Stringer),
[email protected] (S. Eggins), [email protected] (R. Vargiu).
0047-2484/$ - see front matter � 2005 Elsevier Ltd. All rights reserved.
doi:10.1016/j.jhevol.2005.04.005
302 A. Coppa et al. / Journal of Human Evolution 49 (2005) 301e315
On the basis of chronology, dental morphology and metrics, the specimen named Tabun BC7 was identified asa probable Neanderthal.
� 2005 Elsevier Ltd. All rights reserved.
Keywords: Pleistocene human teeth; Neanderthal; Tabun; Israel; Middle Palaeolithic; Dating ESR U-analysis
Introduction
The cave of Tabun has become one of the mostimportant Palaeolithic sites in western Asia, notonly because of the important archaeological andfossil hominin remains found there over 60 yearsago (Garrod and Bate, 1937; McCown and Keith,1939), but also because of its more recent andmuch-discussed role in providing chronological,palaeontological, archaeological and environmen-tal sequences for the region as a whole (e.g. Grunet al., 1991; McDermott et al., 1993; Mercier et al.,1995; Schwarcz et al., 1998; Millard and Pike,1999; Alperson et al., 2000; Bar-Yosef, 2000; Grunand Stringer, 2000). After study by McCown andKeith, the fossil hominin collections from Tabunwere dispersed to three principal repositories in theUnited States (American School of PrehistoricResearch housed at the Peabody Museum, Har-vard), United Kingdom (Royal College of Sur-geons, London) and what was then Palestine (TheRockefeller Museum, Jerusalem). This paper con-cerns a partial upper dentition from Tabun thathas been in the collections of The Natural HistoryMuseum of London (NHM) since the transfer ofmaterial from the Royal College of Surgeons, in1955. AC examined the teeth in 1999, and notedinteresting features that warranted further study.
A footnote to p. 146 of Oakley et al. (1975)reports the following: ‘‘There are 7 isolated teethfrom Tabun..now housed in the British Museum(Natural History). They appear to be Recent as themolars show marked compression-reduction,which is a feature of the dentition of modernman. These teeth are probably those derived fromTabun Layer A (Bronze Age to Recent) referredto in Garrod and Bate, 1937: 64’’. Given thepreservation and morphology of the teeth, ACsurveyed the existing Tabun human dental collec-tions, in comparison with the published records, in
order to establish their provenance and identitymore accurately with the following conclusions. Inorder of stratigraphic attribution (Garrod andBate, 1937; McCown and Keith, 1939; Oakleyet al., 1975), the dental collections located andidentified are shown in Table 1. In the table, welisted also the direct observations made by AC. Itcan be noted that some teeth were differentlyidentified and others are apparently missing. Froma general view, four main points were highlightedas follows.
1) The seven teeth examined at NHM (right I1, I2,M1, M2, M3 and left M2, M3) cannot be thosefrom Layer A (Bronze Age to Historic), men-tioned in McCown and Keith (1939). These so-called ‘‘recent teeth’’ were located at the PeabodyMuseum (although with minor differences inidentification), and they relate to ‘‘Series V’’.
2) The teeth belonging to Series I to IV (fromChimney II and Layer B) were almost alllocated except for three teeth: one from TabunBC2 (‘‘Series II’’, left I2), one from Tabun B1(‘‘Series I’’, right M1) and one from Tabun B4(‘‘Series III’’, left M1). However, Tabun BC6(right I1, left M2) is apparently missing. On thebasis of morphology and contact surfaces, itwas deduced that the deciduous teeth attributedto Tabun B5 ‘‘Series IV’’ and the permanentteeth of BC2 ‘‘Series II’’ belonged to the sameindividual.
3) Most of the missing teeth do not match withthe seven of our series (they are mandibular, orfrom the opposite side). Some that potentiallycould match, like the right M1 from Series I,were in the socket before it was lost, and in anycase, from Plate XIV in McCown and Keith(1939), it is clear that this is a different tooth.Thus, only the right I1 of Tabun BC6 poten-tially matches.
303A. Coppa et al. / Journal of Human Evolution 49 (2005) 301e315
Table
1
Listofthedentalcollectionslocatedandidentified,withtheobservationsmadebyAC.Theteethnew
lydescribed
hereare
inbold
type
GarrodandBate,1937
McC
ownandKeith,1939
Oakleyet
al.,1975
AC
Left
Right
Left
Right
Left
Right
Layer
AA
humanmilk
incisor
SeriesV
I1,C1,M
2m
2,I2,C1,P3,
C1,M
1,M
1
SeriesV
C1
I1,C1,P3,
M1,M
2,M
3
Chim
ney
II9teeth
SeriesII
I2,M
1P3,M
1BC2SeriesII
I2,M
1P3,M
1BC2SeriesII
M1
P3,M
1
SeriesIV
m2
i1BC6
M2
I1
Layer
B9decidousand
7isolatedteeth
SeriesI
I2,C1,P3,
P4,M
1,M
2B1SeriesI
I2,C1,P3,P4,
M1,M
2B1SeriesI
I2,C1,P3,
P4,M
2
SeriesII
I 2B3SeriesII
I 2B3SeriesII
I 2SeriesIII
I1,I2,M
1M
3B4SeriesIII
I1,I2,M
1M
3B4SeriesIII
I1,I2
M3
SeriesIV
m2
m2
B5SeriesIV
M2
M2
B5SeriesIV
m2,m
2i1,m
2
Layer
CSometeethalm
ost
certainly
derived
from
Layer
A
(see
text)
7isolated
teeth(see
text)
TabunBC7
M2,M
3I1,I2
M1,
M2,M
3
TabunITabunII
TabunC1‘‘TabunI’’
TabunC2‘‘TabunII’’
TabunC3-C
7
TabunC1‘‘TabunI’’
TabunC2‘‘TabunII’’
4) Garrod and Bate (1937, bottom of page 64)state that there are human teeth from theoutermost fringe of C, which they then insteadattribute to layer A. It is evident that theseotherwise unlocated teeth are almost certainlythe seven indicated in the Oakley et al. (1975)footnote. Oakley et al. (1975) confused theseteeth with the ones from ‘‘Series V’’ becausethe molar shape evidently implied that theywere Recent. It is certainly true that the secondmolars show mesio-distal compression (due to‘‘metacone reduction’’), but the other teeth donot show reduction (neither for MD or BLdiameters, nor occlusal surface areas).
Description of the teeth
The seven teeth are characterized by the samedegree of fossilization, and represent only upperteeth, without duplication: right I1 and I2, M1, M2,M3; left M2 and M3 (Figs. 1e3). The first questionis to establish whether all these teeth belong to thesame individual, and then to decide whether it ispossible to attribute them to a Neanderthal (as theC1 skeleton) specimen or not. In order to answerthese questions, we have analysed the dentalremains morphologically and metrically.
The identity of the teeth
All the molars share corresponding mesial anddistal surfaces of contact, and they are compatiblewith each other (cf Wolpoff, 1979; Radov�cic et al.,1988). The patterns of wear are also comparable,but with the incisors more worn than the molars,as generally observed in Neanderthals, includingwestern Asian examples such as Tabun C1, Amud1, Kebara 2 and Shanidar sample (Wolpoff, 1980;E. Trinkaus pers. comm. 2004). The similar degreeof fossilization and the compatibility of the teethboth physically and in wear suggest that theyprobably represent the same individual. Given thatthe seven teeth were reportedly found in the thin,outermost fringe of Layer C, but that dating workreported below suggests alternative attribution toLayer B, we will refer to these seven teeth as
304 A. Coppa et al. / Journal of Human Evolution 49 (2005) 301e315
Fig. 1. Occlusal view of Tabun BC7 teeth in anatomical position adapted from Gray’s Anatomy (35th British edition, 1973).
representing individual BC7 for the rest of thispaper (continuing the numbering system of Oakleyet al., 1975).
Morphological traits
We scored 23 morphological traits on the seventeeth attributed to Tabun BC7 according to theArizona State University Dental AnthropologySystem (ASUDAS) using rank-scale referenceplaques (Turner et al., 1991). Trait expressionwas dichotomized into presence/absence (1/0).This was done based upon each trait’s thresholdusing standard procedures (Turner, 1985, 1987;Irish, 1993; Scott and Turner, 1997). The score ofeach trait in Tabun BC7 was compared with thefrequencies of 9 late Pleistocene and early Holo-cene samples from Europe, North Africa and theMiddle East (Coppa et al., 2001; Lucci, 2001)(Tables 2 and 3). The groups are not exactly the
same as those used for the subsequent morpho-metric analysis because of sampling differences.The European Neanderthals were separated in twosubgroups on assigned stratigraphic age: early (lastinterglacial and older) and late (last glaciation).The European Upper Palaeolithic specimens werealso split in two: early (Aurignacian and Gravet-tian) and late (Solutrean, Magdalenian and Epi-Gravettian). In addition, late Upper PalaeolithicNatufians and Iberomaurusians were also includedin the comparison (Coppa et al., 2001; Lucci,2001).
Labial curvature
The labial surface of the upper incisors, canshow a certain degree of convexity when viewedfrom the occlusal aspect. From a general pre-liminary analysis of the data, it can be seen that I1
curvature is a very frequent trait among the
305A. Coppa et al. / Journal of Human Evolution 49 (2005) 301e315
Fig. 2. Tabun BC7 upper incisors.
Neanderthals and the Skhul-Qafzeh (SQ) samples,while it was scarce in the Upper Palaeolithicgroups. Tabun BC7 shows this feature.
Shoveling (Table 3)
The feature of this trait is the presence of mesialand distal marginal ridges on the lingual surface ofthe upper incisors. Presence of this trait on I1 isconsidered to be grades 3-6 (semi-shovel to shovel),on I2 is considered to be grades 3-7. The presence ofI1 and I2 shovel shape made BC7 more similar tothe Neanderthal samples than to SQ, with the latternow closer to the Upper Palaeolithics.
Double-shoveling
Double-shoveling refers to the development ofmesial and distal ridges on the labial surface ofupper incisors. This trait was absent in bothcentral and lateral incisors in BC7 and in theNeanderthal samples. However, it is present on I1
in the later groups (e.g. Natufian 5.4% andEuropean Mesolithic 22.2%), on I2 in SQ (9.1%)and subsequent anatomically modern samples.
Tuberculum dentale
This trait is present as ridges, tubercles, or cuspsthat occur on the lingual surfaces of the upper
306 A. Coppa et al. / Journal of Human Evolution 49 (2005) 301e315
Fig. 3. Tabun BC7 upper incisors compared with anterior upper dentition of Tabun C1.
incisors and canines. The tuberculum dentale wasscored only on the I2. It occurred very frequentlyin almost all the groups, including BC7, with theexception of the European Mesolithics (25.0%).
Peg shaped I2
The peg shaped I2 is a variant which refers toa very reduced size and lacking the normal crown
Table 2
Scores for morphological characters in Tabun BC7 and comparative samples. Presence/absence breakpoints for each trait are in
parenthesis. Tabun BC7 traits expression were dichotomized into presence/absence (1/0)
Upper
I1
Curvature
(2-4/0-1)
Upper
I1
Shoveling
(3-6/0-2)
Upper
I2
Shoveling
(3-7/0-2)
Upper I1
Double
Shoveling
(3-6/0-2)
Upper I2
Double
Shoveling
(3-6/0-2)
Upper I2
Tuberculum
Dentale
(2-6/0-1)
Upper I2
Peg
Shape
(2/0-1)
Upper
M1
Metacone
(5/0-4)
Upper
M2
Metacone
(4-5/0-3)
Upper
M3
Metacone
(4-5/0-3)
Early Neanderthal EU 3.9 63.2 70.6 0.0 0.0 100.0 0.0 100.0 95.8 85.0
Neanderthal EU 100.0 88.9 90.9 0.0 0.0 75.0 0.0 100.0 100.0 100.0
Neanderthal ME with
Tabun BC7
83.3 83.3 100.0 0.0 0.0 100.0 0.0 88.9 85.7 100.0
Neanderthal ME without
Tabun BC7
80.0 80.0 100.0 0.0 0.0 100.0 0.0 75.0 83.3 100.0
AMH Palestine 90.9 9.1 25.0 0.0 9.1 88.9 0.0 100.0 100.0 100.0
Early Upper Palaeolithic EU 45.0 5.3 6.2 0.0 6.2 54.5 0.0 97.3 97.6 92.6
Late Upper Palaeolithic EU 40.0 0.0 0.0 0.0 0.0 64.0 2.2 96.0 94.1 93.2
Natufian 24.3 0.0 18.4 5.4 2.0 80.4 0.0 100.0 100.0 86.1
Iberomaurusian 0.0 0.0 33.3 0.0 10.0 100.0 0.0 100.0 100.0 91.8
Mesolithic EU 33.3 0.0 0.0 22.2 0.0 25.0 0.0 100.0 100.0 100.0
Tabun BC7 1 1 1 0 0 1 0 1 0 0
307A. Coppa et al. / Journal of Human Evolution 49 (2005) 301e315
morphology. A peg shaped I2 was absent in almostall the groups, including BC7.
Metacone reduction
The upper molars distobuccal cusp or cusp 3may show certain variability in size. Absence andweaker forms of expression are rare for M1 andM2, but occasionally occur on M3. Tabun BC7shows metacone reduction only on M2. Generally,a well-developed M1 metacone was present in allthe samples with only the Middle East Neander-thals showing slight M1 and M2 metacone re-duction. Metacone reduction on M2 in Tabun BC7produced a smaller mesiodistal diameter and widerbuccolingual diameter due to the shift of thehypocone towards the lingual surface, giving thetooth a broader shape. This morphology is alsopresent in a more marked expression in theKrapina isolated tooth D168 (Radov�cic et al.,1978: figure 171, page 71).
Hypocone reduction
Like the metacone, this trait refers to theobservation of the variability in size of the uppermolars distolingual cusp or cusp 4. This cusp mayvary from large to a reduced form. Tabun BC7shows no M1 or M2 hypocone reduction. M1 hypo-cone reduction was present only in the European
Mesolithics (20.0%), while M2 hypocone reductionwas absent in the European and Middle EastNeanderthals and in SQ. However, it was presentin the early Neanderthals (35.0%) and in the latersamples. BC7 shows hypocone reduction onM3, andthis was shared with the Middle East Neanderthals.
Metaconule (Cusp 5)
A fifth cusp that can occur in the distal foveabetween the metacone and the hypocone. This traitcould only be scored in Tabun BC7 on M3, whereit is present. It was also present in the Middle EastNeanderthals and SQ (65.0-85.0%). On the contrary,there were low percentages in the European Nean-derthals, both early (60.0%) and late (42.9%).
Parastyle (Table 3)
This cusp commonly occurs on the buccalsurface of the Paracone (Cusp 2), and less oftenon the buccal surface of the Metacone (Cusp 3).Sometimes it is referred to as a paramolar cusp.Presence of this trait is considered to be grades 1-5.Parastyle was scored in Tabun BC7 only on M2
and M3, with the parastyle being absent in boththese molars. In the comparative samples, thepresence of the parastyle on M2 was chronologi-cally distributed, ranging from 35.5% in theearly Neanderthals to 0.0% in the European
Upper
M1
Hypocone
(5/0-4)
Upper
M2
Hypocone
(4-5/0-3)
Upper
M3
Hypocone
(4-5/0-3)
Upper
M3
Cusp 5
(1-5/0)
Upper
M2
Parastyle
(1-5/0)
Upper
M3
Parastyle
(1-5/0)
Upper M1
Enamel
Extension
(2-3/0-1)
Upper M2
Enamel
Extension
(2-3/0-1)
Upper M3
Enamel
Extension
(2-3/0-1)
Upper
M3 Peg
Shape
(2/0-1)
Upper
M1 Root
Number
(3/1-2)
Upper
M2 Root
Number
(3/1-2)
Upper
M3 Root
Number
(3/1-2)
100.0 65.0 6.7 60.0 35.3 28.6 0.0 6.7 0.0 0.0 44.4 0.0 11.1
100.0 100.0 57.1 42.9 16.7 0.0 7.1 10.0 0.0 0.0 66.7 40.0 40.0
100.0 100.0 0.0 100.0 16.7 40.0 0.0 0.0 0.0 11.1 0.0 0.0 0.0
71.4 60.0 0.0 100.0 20.0 50.0 0.0 0.0 0.0 12.5 0.0
100.0 100.0 100.0 83.3 14.3 14.3 0.0 0.0 0.0 0.0 100.0 0.0
100.0 65.5 38.1 72.7 13.8 15.8 0.0 9.4 0.0 9.7 80.0 66.7 11.1
93.6 67.4 28.2 63.9 4.4 2.5 6.7 21.7 14.3 2.2 78.3 80.0 16.7
100.0 70.6 37.7 74.2 2.5 6.1 0.0 1.3 0.0 1.4 81.2 92.3 28.6
97.7 86.0 69.2 71.7 2.1 9.3 0.0 0.0 1.9 3.2 100.0 84.6 54.5
80.0 64.7 30.0 72.7 0.0 0.0 5.9 8.3 33.3 0.0 50.0 16.7 0.0
1 1 0 1 0 0 0 0 0 0 0 0 0
308 A. Coppa et al. / Journal of Human Evolution 49 (2005) 301e315
Table 3
Scores for I1 and I2 shoveling, M2 and M3 Parastyle in Neanderthal and SQ samples
Specimen I1 Shoveling I2 Shoveling M2 Parastyle M3 Parastyle
Present Absent Present Absent Present Absent Present Absent
Arcy 9 *
Biache A * *
Gibraltar 2 *
Hortus II *
Hortus III *
Hortus VII * *
Hortus VIII * *
Hortus IX *
Hortus X *
Krapina 1 *
Krapina 2 * * *
Krapina 3 * *
Krapina 4 * * * *
Krapina 5 * *
Krapina 6 * * *
Krapina 17 * *
Krapina 18 * *
Krapina 19 * * *
Krapina 21 *
Krapina 22 * *
Krapina 23 *
Krapina 24 * *
Krapina 25 * *
Krapina 29 * *
Krapina 30 * *
Krapina 32 * *
Krapina 33 *
Krapina 35 * *
Krapina 97 *
Krapina 99 *
Krapina 109 *
Krapina 132 *
Krapina 148 *
Krapina 173 *
Krapina 175 *
Krapina 180 *
La Quina 5 * *
La Quina 18 * *
Le Fate XI *
Le Moustier 1 * * *
Marillac 1 *
Montsempron 2 *
Montsempron 3 * *
Montsempron 4 *
Peyrards 1 *
Petit Puymoyen 2 * *
Pontnewydd 1 *
Pontnewydd 17 *
Pontnewydd 19 *
Saccopastore 2 * *
Saint-Cesaire 1 * * * *
Sclayn A *
309A. Coppa et al. / Journal of Human Evolution 49 (2005) 301e315
Table 3 (continued)
Specimen I1 Shoveling I2 Shoveling M2 Parastyle M3 Parastyle
Present Absent Present Absent Present Absent Present Absent
Sima de los Huesos 2 *
Spy 1 * *
Spy 2 * *
Subalyuk 1 * *
Subalyuk 2 * *
Vindija 259 *
Vindija 299 *
Amud 1 * * * *
Dederiyeh 2 *
Kebara 2 *
Kebara 27 *
Shanidar 2 * * *
Shanidar 6 * *
Tabun B1 * *
Tabun B4 * *
Tabun C1 * * * *
Tabun BC7 * * * *
Teshik-Tash 1 * *
Qafzeh 3 *
Qafzeh 4 * *
Qafzeh 5 * *
Qafzeh 6 * * *
Qafzeh 7 * * * *
Qafzeh 8 *
Qafzeh 8b * * * *
Qafzeh 9 * * * *
Qafzeh 11 * * * *
Qafzeh 15 * *
Qafzeh 20 * *
Qafzeh 27 * * * *
Skhul 1 * *
Skhul 4 * *
Skhul 5 * * * *
Mesolithics. On M3 a higher frequency was shownby the Middle East Neanderthals (50.0%), while itwas completely absent in the European Neander-thals and Mesolithics. The presence or absence ofthe parastyle scored on M2 and M3 in the MiddlePalaeolithic samples (both Neanderthals and SQ)is shown in Table 3. In most specimens the traitwas absent, but it did occur in some Middle EastNeanderthals, such as Tabun B1 (on M2), TabunC1 and Shanidar 6 (on M3).
Enamel Extension
The enamel extension is the projection of theenamel border of the upper molars that extends
towards the tooth root. Tabun BC7 lacked theenamel extension in all the molars. The same wastrue for all the Middle East Neanderthals and SQ.However, the trait was present at low frequencieson M1 in the European Neanderthals (7.1%), andon M2 in both the early (6.7%) and late EuropeanNeanderthals (10.0%).
Peg shaped M3
This refers to a third molar that is reduced insize (7 to 10 mm buccolingual diameter), or peg-like (cone-shaped with its buccolingual diameterless than 7 mm). A peg shaped M3 was absent inTabun BC7, as well as in all the European
310 A. Coppa et al. / Journal of Human Evolution 49 (2005) 301e315
Neanderthals and SQ. However, it was present inthe Middle East Neanderthal specimen Amud 1.
Root number (three-rooted upper molars)
Root number is counted and considered bi-furcated when the individual root length is greaterthan one-quarter to one-third of the total rootlength. The upper molars root number may varyfrom 1 to 3, rarely four or five third molar rootsare present. In this study we considered asthreshold the presence of three roots. This traitwas completely absent in Tabun BC7, since the M1
and M2 were two-rooted and M3 was single-rooted. Tabun BC7 was the only specimen amongthe Middle East Neanderthals where it waspossible to record this trait on both M1 and M2.In general, the frequency of this character waslower in the European Neanderthals than in SQ.In the Middle East samples it was not possible toscore the trait on M1, while in the only specimenwhere M2 was available (Qafzeh 11), this wasthree-rooted. In the case of M3, in the two Middle
East specimens available (Kebara 2 and Shanidar3), they were two-rooted. In SQ, only Qafzeh 9had the M3 available and this was single-rooted.Among the European Neanderthals this trait wasmore frequent (40.0%).
Metrical data
Metric data are tabulated as mesio-distal (MD)and bucco-lingual (BL) diameters, and occlusalsurface areas (Tables 4a,b,c). The Tabun BC7 datawere compared with Neanderthal samples fromEurope and the Middle East (Amud, Kebara,Tabun and Shanidar), and with late Pleistoceneand early Holocene modern human samples fromthe same regions (Tables 4a,b,c). All metric datawere taken on original fossils and were scored byone of the authors (AC, unpublished data).
MD diameter
As far as the MD diameter is concerned, wehave scored this only on molars because of the
Table 4a
Scores for molars Mesio-Distal diameters in Tabun BC7 and comparative samples
M-D
M1 M2 M3
N Mean S.D. N Mean S.D. N Mean S.D.
Neanderthal EU 9 10.5 0.5 6 10.2 0.7 8 9.1 0.6
Neanderthal Middle East with Tabun BC7 4 11.0 0.4 4 9.9 0.7 5 8.9 0.6
Neanderthal Middle East without Tabun BC7 3 10.9 0.4 3 10.3 0.2 4 8.9 0.7
Amud 1 10.5 10.1 8.2
Kebara 2 9.3
Tabun B1 10.5
Tabun B4
Tabun BC2 11.3
Tabun C1 10.9 10.2 8.5
Shanidar 1 9.6
Shanidar 3
AMH Palestine 10 11.3 0.5 5 10.8 0.7 5 9.4 0.5
Early Upper Pal. France 5 10.7 0.7 3 10.3 1.3 4 8.8 0.3
Early Upper Pal. Italy 6 10.5 0.5 7 9.6 0.5 5 8.6 0.6
Early Upper Pal. Cent. Europe 13 10.4 0.5 14 9.8 0.9 10 8.9 0.6
Late Upper Pal. France 15 10.3 0.4 14 9.4 0.5 10 8.8 0.6
Late Upper Pal. Italy 14 11.0 0.6 13 10.2 0.7 9 8.6 0.5
Late Upper Pal. Cen. Europe 1 9.4 1 7.9
Mesolithic EU 12 10.1 0.3 10 9.8 1.2 9 8.5 1.3
Tabun BC7 11.2 9.0 9.1
311A. Coppa et al. / Journal of Human Evolution 49 (2005) 301e315
Tale 4b
Scores for incisors and molars Bucco-Lingual diameters in Tabun BC7 and comparative samples
B-L
I1 I2 M1 M2 M3
N Mean S.D. N Mean S.D. N Mean S.D. N Mean S.D. N Mean S.D.
Neanderthal EU 8 8.1 0.4 7 8.1 0.5 11 12.6 0.5 6 12.8 0.7 9 12.6 1.0
Neanderthal Middle East
with Tabun BC7
4 8.5 0.4 5 8.0 0.2 4 12.4 0.3 4 12.4 0.6 5 11.7 1.1
Neanderthal Middle East
without Tabun BC7
3 8.6 0.5 4 8.0 0.2 3 12.4 0.4 3 12.1 0.4 4 11.8 1.3
Amud 1 8.6 8.3 12.5 12.2 11.1
Kebara 2 13.0
Tabun B1 7.9 12.4
Tabun B4 9.1 8.0
Tabun BC2 12.7
Tabun C1 8.1 7.8 11.9 11.7 10.4
Shanidar 1
Shanidar 3 12.7
AMH Palestine 9 8.1 0.7 8 7.5 0.4 10 12.6 0.7 5 12.4 0.9 5 11.7 1.2
Early Upper Pal. France 4 7.4 0.3 1 7.3 4 12.8 0.9 3 12.3 1.4 4 10.8 1.1
Early Upper Pal. Italy 4 7.4 0.4 5 7.0 0.2 7 12.4 0.6 8 12.4 0.6 6 11.0 1.0
Early Upper Pal. Cent. Europe 7 7.4 0.7 8 7.0 0.8 13 11.8 0.9 15 12.4 1.0 10 11.7 1.0
Late Upper Pal. France 7 7.5 0.6 9 6.6 0.4 17 12.1 0.6 15 12.1 0.9 11 11.6 0.8
Late Upper Pal. Italy 3 7.0 0.5 11 6.6 0.4 15 12.5 0.7 13 12.7 0.8 12 11.7 0.7
Late Upper Pal. Cen. Europe 1 6.7 1 6.2 1 11.2 1 10.6
Mesolithic EU 8 7.4 0.5 6 6.6 0.3 13 11.8 0.6 9 11.9 0.9 9 10.7 1.1
Tabun BC7 8.3 8.2 12.5 13.2 11.5
high degree of attrition on the incisors of TabunBC7. The M1 value for Tabun BC7 is the largest,along with SQ and Tabun BC2. The other samplesshowed lower diameters, particularly the Meso-lithics. SQ and Tabun B1 showed the highestvalues for M2, while Tabun BC7 had the lowest ofall. This is due to metacone reduction, as discussedearlier. The three highest values for M3 were forShanidar 1, Kebara 2 and SQ. Tabun BC7 and theEuropean Neanderthals are slightly smaller. Con-trary to the data of Smith (1989), second and thirdmolar MD diameters in our samples were larger inSQ than in the European and Middle EastNeanderthals.
BL diameter
With reference to BL diameter, I1 and I2
showed a marked difference between the Middleand Upper Palaeolithic series, but with SQ in-termediate, especially for I2. M1 values do notshow a very high degree of variability among the
samples. Tabun BC7 shows the highest M2 value,followed by the European Neanderthals andItalian late Upper Palaeolithics. Conversely, Ta-bun BC7 shows a smaller M3. Another peculiarityof our specimen is that the M2 BL diameter isgreater than that of the M1, but this characteristicmight be due to its unusual M2 morphology.
Occlusal surface area (molars only)
As far as molar occlusal surface area is con-cerned, Tabun BC7, Tabun BC2 and SQ show thehighest values for M1. As discussed, Tabun BC7has the smallest M2, due to its small MD dimen-sion, while it has an intermediate value for M3.
Dating Analysis of Tabun BC7: experimental
procedures and results
A fragment of the tooth was removed for ESRand U-analysis. The uranium distribution on
312 A. Coppa et al. / Journal of Human Evolution 49 (2005) 301e315
Table 4c
Scores for molars areas in Tabun BC7 and comparative samples
Areas
M1 M2 M3
N Mean S.D. N Mean S.D. N Mean S.D.
Neanderthal EU 9 131.5 7.5 6 130.9 11.4 8 115.3 12.8
Neanderthal Middle East with Tabun BC7 4 136.0 6.9 4 122.6 5.5 5 105.3 15.9
Neanderthal Middle East without Tabun BC7 3 134.6 7.8 3 123.9 6.0 4 105.6 18.3
Amud 1 131.3 123.2 91.0
Kebara 2 120.9
Tabun B1 130.2
Tabun B4
Tabun BC2 143.5
Tabun C1 129.1 118.3 88.4
AMH Palestine 10 142.6 12.4 5 134.7 17.0 5 110.2 16.7
Early Upper Pal. France 4 139.0 18.4 3 128.5 29.4 4 94.8 6.9
Early Upper Pal. Italy 6 130.0 11.4 7 117.9 10.3 5 95.0 12.6
Early Upper Pal. Cent. Europe 12 122.8 12.5 14 120.6 16.8 10 104.3 14.1
Late Upper Pal. France 15 124.1 11.0 14 113.3 12.7 10 102.9 13.4
Late Upper Pal. Italy 14 137.7 14.0 13 130.0 15.6 9 101.1 10.8
Late Upper Pal. Cen. Europe 1 105.3 1 83.2
Mesolithic EU 12 118.1 7.4 9 116.6 14.6 9 91.1 14.3
Tabun BC7 140.0 118.8 104.2
dentine and enamel was analysed by laser ablationICP-MS (for details on the method, see Egginset al., 2003). Figure 4a shows the cross section ofthe sample along with the scan which has a widthof about 100 mm and a depth of about 10 to 20 mm.The distribution of uranium in dentine is quitehomogeneous and estimated to be 19.8G0.7 ppm.The uranium in the enamel decreases from about0.27 ppm close to the dentine to background levels(�1 ppb) close to the enamel surface. The averageenamel concentration is 0.08 ppm. After removalof the dentine, the enamel fragment was mountedin a Bruker ER 218PG1 programmable goniom-eter and measured at each dose step at 10E angleintervals for 290E (additional spectra to 360E werelost in a transfer process). ESR measurementswere carried out on a Bruker ECS 106 spectro-meter with a 15 kG magnet and a rectangular 4102ST cavity. The samples were recorded with themeasurement parameters routinely applied in thislaboratory: accumulation of between 200 (naturalsample) and 100 scans (for the higher dosedsamples) with 1.015 Gpp modulation amplitude,10.24 ms conversion factor, 20.48 ms time con-stant, 2048 bit spectrum resolution (resulting in
a total sweep time of 20.972 s), 120 G sweep widthand 2mW microwave power. The enamel piecewas successively irradiated with the followingcumulative doses: 0, 11.1, 22.2, 44.3, 88.8, 133,222, 310 and 488 Gy. ESR intensity values wereobtained by natural spectrum fitting (see Grun,2002), dose values were obtained by applyinga single saturating function with linear conversion,errors were obtained by Monte Carlo simulation(for more details see Grun and Brumby, 1994).Figure 4b shows the angular dose measurements,which yielded an average value of 87.6G2.1 Gy.
Discussion of dating results
When the dose value of the BC7 molar iscompared to ESR analysis of faunal samples fromGarrod’s Tabun collection (Grun et al., 1991) onlythe samples from Layer B (548 and 550, see Table 2in Grun et al., 1991) have similar dose values.All other samples from the lower levels hadconsiderably higher values (O 130 Gy). If the toothis not intrusive, it is most likely contemporaneouswith the faunal samples from layer B. Using thesediment values of layer B, preliminary age
313A. Coppa et al. / Journal of Human Evolution 49 (2005) 301e315
Fig. 4. a. Uranium and Thorium profiles of Tabun BC7. Thin vertical lines indicate where the laser tracks cross the dentine/enamel
interface and the outer surface of the tooth. U and Th are given in ppm whereas P (used to correct for ablation efficiency) is the
measured raw signal intensity. b. ESR dose estimation at different angles. Natural spectrum fitting (Grun 2002) yields an average dose
of 87.6G 2.1 Gy (mean and standard deviation of the repeated measurements).
estimates of 82G14 ka (early U-uptake) and92G18 ka (linear uptake) are obtained. The bestestimate for BC7 is 90C 30-16 ka when using theU-series data obtained on the other faunal material
from layer B (see Grun and Stringer, 2000). Theearly uptake age, being the minimum possible age,precludes an intrusion of BC7 into Layer B ata much later stage than the deposition of this layer.
314 A. Coppa et al. / Journal of Human Evolution 49 (2005) 301e315
The dose value obtained on an enamel fragmentfrom C1, 172G8 Gy, is nearly double the valuemeasured for this sample and tentative ageestimates were in the range of 143G37 ka. Becauseof the overlapping error bars, which are the resultof the inhomogeneity of the sediment and theuncertainties of the exact provenance of the humanspecimens, we cannot conclude that C1 is definitelyolder than BC7. It may well be that both are ofapproximately the same age, but C1 was buried ina high dose rate environment and BC7 in a lowdose rate environment.
Concluding remarks: dating and affinities
of the Tabun BC7 dentition
The preservation, morphological and metricalanalyses, and the new direct age estimates, are allin contrast with the old hypothesis that the seventeeth were intrusive in the Middle Palaeolithicsediments at Tabun. The present study suggeststhat they derived from the thin marginal sedimentsof Layers B-C and chronologically derive fromLayer B. Morphologically, we believe that TabunBC7 most probably represents a Neanderthal fromthe same population as Tabun C1.
Acknowledgements
We would like to thank the Photographic Unitof The Natural History Museum, London for thedigital images of the Tabun teeth, and threereviewers for their considerable help in improvingour manuscript. This research was supported inpart by MURST COFIN03.
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