Dental Morphological Affinities Late Pleistocene Sub-saharan - Irish

8/7/2019 Dental Morphological Affinities Late Pleistocene Sub-saharan - Irish

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Joel D. Irish

Dental morphological affinities of Late Pleistocene through

recent sub-Saharan and north African peoplesIn: Bulletins et Mmoires de la Socit d'anthropologie de Paris, Nouvelle Srie, tome 10 fascicule 3-4, 1998. pp.

237-272.

Abstract

Summary. The utility of dental morphological analysis of Sub-Saharan and North African peoples to estimate population

origins and affinities has often been overlooked by anthropological researchers. The present report entails a summary review of

four recent publications by the author that have addressed this shortcoming m three principal ways. First, 36 morphological

features in 1 ,643 dentitions from over 30 samples throughout the continent have been recorded. The frequencies of trait

occurrence in each sample are provided here. Second, these traits were compared among samples with the multivanate Mean

Measure of Divergence (MMD) statistic; multidimensional scaling is employed to illustrate the MMD-based relationships.

Assuming phenetic similarity parallels genetic relatedness, these biological distance estimates reveal that: (a) significant

differences exist between most Sub-Saharan and North African samples; (b) dental homogeneity is evident within both regions;

and (c) North African samples exhibit the greatest homogeneity. Third, pooled Sub- Saharan and North African trait frequencies

were compared with published data from five non- African samples. It is shown that North Africans appear most like Europeans

and perhaps western Asians; Sub-Saharan Africans are unlike all other samples. These findings, in conjunction with additional

evidence presented in the recent publications, should contribute to an improved understanding of intra-African biological affinities

and African dental relatedness to other world populations.

Citer ce document / Cite this document :

Irish Joel D. Dental morphological affinities of Late Pleistocene through recent sub-Saharan and north African peoples. In:

Bulletins et Mmoires de la Socit d'anthropologie de Paris, Nouvelle Srie, tome 10 fascicule 3-4, 1998. pp. 237-272.

doi : 10.3406/bmsap.1998.2517

http://www.persee.fr/web/revues/home/prescript/article/bmsap_0037-8984_1998_num_10_3_2517
http://www.persee.fr/web/revues/home/prescript/author/auteur_bmsap_866http://dx.doi.org/10.3406/bmsap.1998.2517http://www.persee.fr/web/revues/home/prescript/article/bmsap_0037-8984_1998_num_10_3_2517http://www.persee.fr/web/revues/home/prescript/article/bmsap_0037-8984_1998_num_10_3_2517http://dx.doi.org/10.3406/bmsap.1998.2517http://www.persee.fr/web/revues/home/prescript/author/auteur_bmsap_866


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Bull, et Mm. de la Socit d'Anthropologie de Paris, n.s., t. 10, 1998, -, . 237-272.

DENTAL MORPHOLOGICAL AFFINITIES OF LATEPLEISTOCENE THROUGH RECENT SUB-SAHARAN

AND NORTH AFRICAN PEOPLES

Joel D. Irish 'Summary. The utility of dental morphological analysis of Sub-Saharan and North Africanpeoples to estimate population origins and affinities has often been overlooked by anthropologicalresearchers. The present report entails a summary review of four recent publications by the authorthat have addressed this shortcoming m three principal ways. First, 36 morphological features in

1 ,643 dentitions from over 30 samples throughout the continent have been recorded. The frequenciesof trait occurrence in each sample are provided here. Second, these traits were compared amongsamples with the multivanate Mean Measure of Divergence (MMD) statistic; multidimensionalscaling is employed to illustrate the MMD-based relationships. Assuming phenetic similarity parallelsgenetic relatedness, these biological distance estimates reveal that: (a) significant differences existbetween most Sub-Saharan and North African samples; (b) dental homogeneity is evident withinboth regions; and (c) North African samples exhibit the greatest homogeneity. Third, pooled Sub-Saharan and North African trait frequencies were compared with published data from five non-African samples. It is shown that North Africans appear most like Europeans and perhaps westernAsians; Sub-Saharan Africans are unlike all other samples. These findings, in conjunction withadditional evidence presented in the recent publications, should contribute to an improvedunderstanding of intra-African biological affinities and African dental relatedness to other worldpopulations.Key words: Dental anthropology, tooth crown and root morphology, biological affimty, Sub-Saharan and North African populations.

AFFINITS DES POPULATIONS D'AFRIQUE DU NORD ET D' AFRIQUE SUB-SAHARIENNE DEPUIS LA FIN DUPLEISTOCENE JUSQU' L' ACTUEL, D' APRS LA MORPHOLOGIE DENTAIRE

Rsum. L'intrt du recours la morphologie dentaire pour estimer les origines et les affinitsdes populations d'Afrique septentrionale et sub-sahanenne a souvent t nglig par lesanthropologues. Pour y remdier, cet article synthtise en trois tapes, quatre publications rcentesde l'auteur. Tout d'abord, 36 observations ont t faites sur 1643 dentures provenant de plus de 30i Department of Anthropology, University of Alaska Fairbanks, 310 Eielson Building, P.O Box 757720,Fairbanks, AK 99775-7720, USA. e-mail ffjdi@aurora alaska edu


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238 JOEL D. IRISH

populations africaines. Les frquences des diffrents caractres observs sont donnes ici. Ensuiteces traits on t t compars statistiquement entre populations, grce la MMD (multivariate MeanMeasure of Divergence) ; les affinits bases sur cette MMD sont illustres par une reprsentationmultidimensionnelle. En admettant que la ressemblance morphologique soit parallle aux affinitsgntiques, cette estimation de la distance biologique rvle que : (a) il existe des diffrencessignificatives entre la plupart des chantillons provenant d'Afrique du Nord et ceux d'Afrique subsaharienne ; (b) il y a l'vidence une homognit dentaire dans ces deux rgions ; et (c) ce sontles populations du Maghreb qui prsentent la plus grande homognit. Enfin, les frquences decaractres de l'ensemble des populations africaines ont t compares celles de cinq chantillonsnon-africains trouvs dans la littrature. Il apparat que les Maghrbins ressemblent surtout auxhabitants de l'Europe et peut-tre du Moyen-Orient, alors que les populations d'Afrique subsaharienne sont diffrentes de tous les autres chantillons. Ces rsultats, qui viennent s'ajouter ceux rcemment publis par d'autres, devraient contribuer mieux comprendre les affinitsbiologiques entre populations l'intrieur de l'Afrique, et les relations entre la dentition des Africainset celle des autres peuples du monde.Mots-cls : Anthropologie dentaire, morphologie de la racine et de la couronne dentaire, affinitsbiologiques, populations sub-sahariennes et d'Afrique du Nord.

INTRODUCTIONRelative to other world groups, few dental morphological data have been available fo rLate Pleistocene through recent peoples of Africa. South Africans (Drennan, 1929; Shaw,1931; Galloway, 1959; Van Reenen, 1966; Tobias, 1972; Jacobson, 1982; among others)and Nubians (Greene 1967, 1972, 1982; Greene et ai. 1967; Smith and Shegev 1988;Turner and Markowitz 1990; among others) have been studied to some extent. But only alimited number of published papers concern peoples from West (e.g., Watters, 1958),Central (Pales, 1938; Brabant, 1965; among others), East (Chagula, 1960; Barnes, 1969;Hassanali, 1982; Hassanali andMwaniki; 1984; Sakumaand Ogata, 1987, 1988; Sakumaet ai, 1991; among others), and the majority of North Africa (Bermudez de Castro 1989,1991; Johnson and Lovell, 1994; among others). Most native African populations remainunstudied by dental morphologists. Moreover, with few notable exceptions (e.g., Greene,

1967; Bermudez de Castro 1991; Johnson and Lovell, 1994), the existing studies rarelydescribe more than a handful of traits and samples; fewer still have presented inter-regionalcomparative analyses (e.g., Haeussler etal, 1989; Irish and Turner, 1990).Over the past ten years CG Turner II and the present author have supplemented thisdeficiency in dental data via several papers and presentations (e.g. , Irish and Turner, 1989,1990, 1992; Irish, 1993a), which ultimately led to four comprehensive reports (Irish 1993b,1997, 1998a,b) that incorporate analyses of many morphological traits in samplesthroughout the continent. The present summary article reviews three main aspects ofthese recent reports. First, morphological dental trait frequencies in a large number ofAfrican samples are tabulated. These data will aid in describing the peoples of this vast


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AFRICAN DENTAL AFCTNITIES 239

KABYLEBEDOUIN CARTHAGECANARYISLANDS LJSHT ELHESA

MESOLJTHICMEROITICX-GROUP

TOGO/BENINGABONPYGMYCONGO

continent, and provide acomparative baseline forinterested researchers.Second, samples of Sub-Saharan and North Africansare compared to oneanother. Results show thatmost samples from each ofthe two geographic regionsdiffer markedly from oneanother. However, despitesynchronie and diachronicdiversity these Sub-Saharanand North African samplesexhibit notable intra-regiondental homogeneity characterized by complexmass-additive, and simplemass-reduced teeth,respectively. Third, Sub-Saharan and North Africans are compared to published data fromseveral large non- African groups, to gain an understanding of these peoples from a globalperspective. North Africans are most similar to Europeans and perhaps western Asians;Sub-Saharan Africans are divergent from all others. These findings should lead to a betterdental characterization of African peoples, and provide data that may ultimately clarifyAfrican origins, and relationships to other world populations (e.g., Irish, 1998b).

SOTHONGUNIKHOIKHOI

Figure 1 . Origin locationsof 28 African samples used inthe study (from Irish, 1997).

MATERIALSAND METHODSIn the previous studies (Irish 1993b, 1997, 1998a,b), 32 late Upper Pleistocene throughrecent Sub-Saharan and North African dental samples were examined. Twenty-eightsamples are from specific African cultures based on tribal and/or linguistic affiliations(Figure 1); the other four are pooled collections of miscellaneous individuals from thesame general geographic regions (i.e., North, South, East, and West Africa). For the sake


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240 JOEL D. IRISH

Sample nameSub-Saharan AfricaCongo (CON)Chad ()Gabon (GAB)Ghana (GHA)Kenya (KEN)Khoikhoi (KHO)Nguni (NGU)Nigena/Cameroon (NIC)Pygmy (PYG)San (SAN)Senegambia (SEN)Sotho (SOT)Tanzania (TAN)Togo/Benin (TOB)Tukulor (TUK)Mise South AfricaMise East AfricaMise West Africa

North AfricaAlgeria (ALG)Bedouin (BED)Canary Islands (CAN)Capstan (CAP)Carthage (CAR)Christian (CHR)El Hesa (HES)Kabyle (KAB)Kharga (KHA)Lisht (LIS)Meroitic (MER)Mesohthic (NUB)Pharonic (PHA)X-Group (XGR)Mise North Africa

ProvenienceCongoChadGabonGhanaKenya, TanzaniaSouth AfricaSouth AfricaNigeria, CameroonCongo, GabonBotswana, South AfricaSenegambiaSouth AfricaTanzania, ZanzibarTogo, BeninSenegambiaNamibia, Zaire, South AfricaZimbabwe, Ruanda, Kenya, etcCameroon, Ghana, Lbena, etc.AlgeriaMorocco, Tunisia, LibyaCanary IslandsAlgeria, TunisiaTunisiaNubia (Semna)EgyptAlgeriaEgyptEgyptNubia (Semna)Nubia (Jebel Sahaba)Nubia (Soleb)Nubia (Semna)Algeria, Northern Chad

Culture/tribeTeke, KongoToubou, Masalit, KanembuFang, Nkomi, Lumbo, MpongweAshanti, FantiKikuyu, Swahili, Chaga, PareNama, Koran(Nguni) Zulu, XhosaEfik, Ibibio?, Boki, AnyangBinga, Bongo'Kung, Naron, Tshakwe, MkaukauWolof, Balante, Serer(Sotho) , TauNyamwezi, Ngindo, Hehe, GogoEwe, FonTukulor?Masai, Nuer, 9Mabeya, Mandingo, Sanga, Golan

Shawia BerbersBedouin ArabsGuancheNeolithic CapsiansPunic Period PhoeniciansChristian Period NubiansRoman/Byzantine EgyptiansKabyle BerbersByzantine Egyptians12th Dynasty EgyptiansMeroitic Penod NubiansLate Paleo Period NubiansNew Kingdom Period NubiansX-Group Penod NubiansMdaga, Boudouma

Table I. The 33 Sub-Saharan and North African dental samples


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AFRICAN DENTAL AFFINITIES 241

PeriodXIXth CentXIX-XX"1 CentXIX-XX"1 CentXIX* CentXIX-XX"1 CentXIXth CentXIXth CentXIXth CentXIX-XX"1 CentXXth CentXIX-XX"1 CentXXth CentXIX-XX"1 CentXIX"1 CentXIX-XX* CentXIXth CentXIX-XX"1 CentPrehis/Hist

XIXth CentXIX-XXth CentAD 400-900?10,500-7,000 751-146 AD 550-1350AD 200-400XIX-XX111 CentAD 500-6001991-1783 100 -AD 35010-12,000 1575-1380 AD 350-550Prehis/Mod

LanguageCongo-KordofanNilo-SaharanCongo-KordofanCongo-KordofanCongo-KordofanKhoisanCongo-KordofanCongo-KordofanCongo-KordofanKhoisanCongo-KordofanCongo-KordofanCongo-KordofanCongo-KordofanCongo-KordofanCongo-KordofanC-K & N-SCongo-KordofanAfroasiaticAfroasiaticAfroasiatic??Afroasiatic7?Afroasiatic?AfroasiaticAfroasiatic''Afroasiatic??77?Afroasiatic

No.3029394711437225722994317844253992649

264916322281872322661916732398

Facility wAMNH, MHMHMH, NMNHAMNHAMNH, MH, NMNHAMNH, MHAMNH, NMNHAMNHAMNH, MHAMNH, ASUMH, NMNHASUAMNH, MHAMNHMHAMNH, MH, NMNHAMNH, MH, NMNHAMNH, MHMHMH.UMAMNH, MH, NMNHUM, IPHMHASUAMNHMHNMNHNMNHASUSMUMHASUMH, UM

()= Museum of Natural History, ASU=Arizona State University, IPH=Institutde Palontologie Humaine, MH=Muse de l'Homme, NMNH=National Museum of NaturalHistory, SMU=Southern Methodist University, UM=University of Minnesota.


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242 JOEL D. IRISH

MaxillaWingingLabial CurvaturePalatine TorusShovelDouble ShovelMidhne DiastemaInterruption GrooveTuberculum DentaleBushman CanineDistal Accessory RidgeHypoconeCusp 5 (Metaconule)Carabelh's TraitParastyleEnamel ExtensionRoot NumberRoot NumberPeg-ReducedOdontomeCongenital Absence

UI1UI1UI1UI1UI1UI2UI2UCUCUM2UM1UM1UM3UM1UP1UM2UI2P1-P2UM3

MandibleLingual Cusp NumberAnterior FoveaMandibular TorusGroove PatternRocker JawCusp NumberCusp NumberDeflecting WrinkleDistal Tngonid CrestProtostylidCusp 7 (Metaconulid)Tome'sRootRoot NumberRoot NumberRoot NumberTorsomolarAngle

LP2LM1LM2LM1LM2LM1LM1LM1LM1LP1LCLM1LM2LM3

Table II. The 36 dental and osseous traits used in the study. Traits from ASU DentalAnthropology System (Turner et al, 1991). (


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AFRICAN DENTAL AFHNITIES 243

personal communication, 1987; Haeussler etal, 1989), and thus, could only be examinedfo r 28 traits; root traits, UM1 enamel extension, and rocker jaw can not be reproduced incasts (refer to Table III).All traits, except UI1 midline diastema, are included in the Arizona State University(ASU) Dental Anthropology System. Procedures used in the ASU System are based onwell-established criteria fo r scoring intra-trait variation, and have been successfullyemployed in many previous dental morphological studies {e.g., Hassanali, 1982; Scott etal, 1983;Turner, 1984, 1985, 1987, 1992a,b;Sakuma?a/., 1987, 1988, 1991; Haeussleret al, 1989; Irish and Turner, 1990; Turner and Markowitz, 1990; Lukacs and Hemphill,1991; Roler, 1992; Johnson and Lovell, 1994; Lipschultz, 1996; Irish, 1997, 1998a,bamong others). The traits in each African individual were recorded with the aid of 23ASU rank-scaled standard reference plaques, that are available for purchase by all interestedresearchers from theArizona State University Laboratory of Dental Anthropology(2). Onlythe specimen's highest antimere expressions were analyzed to maximize the geneticpotential fo r these polygenic features. Because of a documented lack of trait sexualdimorphism (Scott 1973, 1980; Smith and Shegev, 1988; Bermudez de Castro, 1989;Turner et al, 1991; Hanihara, 1992; Irish 1993b; among others), it is standard Systemprotocol to pool the sexes (Irish, 1997); this protocol is followed here. For a completedescription of ASU System procedures and dental traits see Turner etal (1991). Cursorydescriptions of those traits used in the present study are also listed in the Appendix.Following data recording, trait frequencies were determined for the 33 samples, andC.A.B. Smith's Mean Measure of Divergence (MMD) statistic, using the Freeman andTukey angular transformation fo r small sample sizes (Berry and Berry, 1967; Sj0vold,1973, 1977; Green and Suchey, 1976), was employed on the 29 samples of known culturalaffiliation (incl. Capsian). This multivariate technique provides a quantitative estimate ofbiological divergence among samples based on the degree of phenetic similarity. A lowerMMD value indicates greater similarity, and vice versa. It is assumed that pheneticsimilarity approximates cladistic relationship (Sokal and Sneath, 1 963 ; Scott et al , 1 983).Prior work suggests that as many discrete traits as possible should be used indetermining MMDs (see Sj0vold, 1977). However, these traits should not be correlatedwith one another, otherwise differential weighting of the underlying dimensions can leadto erroneous results (Sj0vold, 1977). In order to test for unwanted correlation, Kendall'stau-b and Spearman's rho rank-order correlation coefficient statistics were employed onthe ranked dental variables. The greatest correlation (r) is only -0.332 (tau-b) to -0.357(rho) between double shoveling UI1 and labial curvature UI1. Further, the total amountof variance (r2) is 0.11-0.13, meaning that only 11-13% of the variance in double shovelingUI1 is directly related to that of labial curvature UI1 . The remaining trait pair correlations

2 Th e Arizona State University Dental Anthropology System plaques, instructions, and scoring sheets areavailable fo r purchase to all interested researchers For details, contact the Laboratory of Dental Anthropology,Department of Anthropology, Arizona State University, Tempe, AZ 85287-2402, USA.


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244 JOEL D. IRISH

are all much closer to zero. Thus, there are no significant correlations among any of the36 dental morphological traits (Carr, personal communication, 1991).

RESULTS AND DISCUSSION

The African Dental Trait FrequenciesTable III lists the occurrence of the 36 traits in the Sub-Saharan samples of knownaffiliation. Table IV lists the trait incidences fo r the North Africans. In both cases, thenumber of individuals with a particular trait is presented, along with the total number ofindividuals fo r whom the trait was scored; from this, the percentage of each trait'soccurrence in the sample was calculated. In some cases, the number of individuals scoredfo r a trait is small (e.g., Capsian, Pygmy, etc.); this unavoidable shortcoming, inherentwith the study of archaeological specimens, should be taken into account when comparingtrait percentages among samples. Trait presence/absence dichotomies corresponding tothe ASU System are listed under the trait names on the left of each table.The 15 Sub-Saharan African samples share some measure of dental trait uniformity,as evidenced by similar high frequencies of complex traits like Bushman Canine, two-rooted UP1, three-rooted UM2, LM2 Y-groove, LP1 Tome's root, two-rooted LM2, and

M3 presence. However, despite similarities there are four trends that suggest specificregional, linguistic, and therefore, likely biological relationships. First, Khoisan-speakingSan (SAN) and Khoikhoi (KHO) from southern Africa share the highest frequencies ofBushman Canine, and lowest frequencies of UC distal accessory ridge and LM1 anteriorfovea. Second, Niger-Congo-speaking (Congo-Khordofanian language superfamily)samples such as Ghana (GHA), Nigeria/ Cameroon (NIC), and Togo/Benin (TOB) fromnear the Gulf of Guinea in western Africa, have the highest frequencies of UI 1 shoveling,UM1 enamel extension, LM1 anterior fovea, and six-cusped LM1. Third, the Bantu-speaking (Congo- Khordofanian superfamily) eastern, western, and southern Africa groups(i.e., Kenya (KEN), Tanzania (TAN), Congo (CON), Gabon (GAB), Nguni (NGU), andSotho (SOT)) exhibit intermediate frequencies for many of these traits. And fourth,dentitions of Sub-Saharan/North African boundary groups (i.e., Senegambia (SEN),Tukulor (TUK), Chad ()) indicate probable North African genetic input based onlower frequencies of LM1 deflecting wrinkle and LM1 cusp 7, and higher UM3 agenesis.As seen in Table IV, the late Upper Pleistocene sample of Mesolithic (Late Paleolithic)Nubians (NUB) clearly differs from the remaining North Africans by its high frequenciesof complex, mass-additive morphology, including: UI1 shoveling, Bushman Canine, UCdistal accessory ridge, UM1 cusp 5, six-cusped LM1, five-cusped LM2, LM1 deflectingwrinkle, and LP1 Tome's root. Further, it has comparatively low frequencies of UI2interruption groove, UM3 agenesis, and rocker jaw. This trait combination is more


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AFRICAN DENTAL AFFINITIES 25 1

indicative of a Sub-Saharan-like dental pattern (see Table III and below). The remaining13 samples exhibit few mass-additive traits, and are instead characterized by higherfrequencies of UI2 interruption groove, UM3 agenesis, and rocker jaw. Moreover, withfew exceptions (e.g., high UI1 shoveling in Meroitic Nubians (MER), high BushmanCanine in Christian (CHR) and Meroitic Nubians, high palatine torus in Canary Islanders(CAN), and low Carabelli's in Pharonic Nubians (PHA)), there is obvious uniformityamong samples.Sub-Saharan and North African Statistical Comparisons

Mean Measure of Divergence analysis yielded results that support the patterns andrelationships suggested by the trait frequency data. A large table of these MMD valuesamong samples is presented elsewhere (Irish, 1993b), and is not included in this briefpaper. Instead, the dental relationships are visualized in a plot of inter-sample MMDvalues using multi-dimensional scaling (MDS) (Kruskal and Wish, 1978). As seen inFigure 2, there is an obvious separation of Sub-Saharan and North African samples, yetapparent homogeneity within regions particularly North Africa. These findings aresupported by previous affinity estimates based on African genetic, skeletal, dermatoglyphic,anthropomtrie, linguistic, and cultural data (see Mourant 1954, 1983; Greenberg 1959,1966; Murdock 1959; Hiernaux 1975; Nurse et al, 1985; Sanchez-Mazas et al, 1986;Excoffier et al, 1987; Roychoudhury and Nei 1988; Howells 1989; Froment, 1992a,b;Franciscus 1995; Holliday 1995; among others).The 15 dentally-complex Sub-Saharan samples are grouped together on the left of theMDS configuration. However, the intra-region trends noted above are evident. SouthAfrican San and Khoikhoi are distinct from other Sub-Saharan samples, yet show anassociation with each other. The Ghana, Nigeria/Cameroon, and Togo/Benin samples fromthe Gulf of Guinea area are loosely clustered; this geographic region is considered thesource of Bantu-speaking agriculturalists, who around 3000-2000 BP began a movementtoward eastern and southern sub-Saharan Africa (Greenberg, 1959, 1966; Hiernaux, 1975;Ehret, 1982; Phillipson, 1985; Nurse etal, 1985). Bantu dental samples from these latterareas (Gabon, Congo, Kenya, Tanzania, Nguni, Sotho) all show an affinity to the WestAfricans and to one another. Moreover, there is evidence for a dental cline rangingfrom more to less trait complexity between West and East/South Africa (refer again toTable III) (see Irish, 1993b fo r a complete discussion). The Sub-Saharan/North Africanboundary groups, Chad, Senegambia, Tukulor, and perhaps Tanzania, are grouped togethernearest the cluster of North Africans (center of Figure 2). And not surprisingly, MesolithicNubians are plotted in close proximity to the Sub-Saharan Africans; this dental affinity,which is discussed in detail elsewhere and may involve an ancestral relationship (Irishand Turner, 1990; Irish, 1993b), is supported by numerous cranial and other morphometrichard tissue studies (see Wendorf 1968; Charon etal, 1974; Hiernaux 1975; Petit-Maire1979; Franciscus 1995, personal communication, 1995; Holliday 1995; among others).


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252 JOEL D. IRISH

H 1 1 1 1 1 1 1 1 h H 1 1 1 1 1 1 1 1 h20 _

1,5 -lesolithic Nubia1.0 -

0, 5 _.

0, 0

Ghana

Togo +BninCongo Sotho

Senegambia Capstan Kharga Kabyle * Canary Is.

-05 -

-10 -

-1.5 _.

-2.0 _ .

Nigeria + 'TanzanCameroon Tukulor * ChadGabonPygmyNguni Kenya

-25 -20 -15 -1.0 -05 0, 0 05 10 15 20 25

Figure 2. Multidimensional scaling of MMD values between 29 African samplesbased on the dental morphological traits.

The average MMD among the 16 Sub-Saharan-affiliated samples is 0.051 (sum of allpairwise MMD comparison values divided by total number of comparisons).The remaining 13 North African samples (excluding Mesolithic Nubia) arecharacterized by morphologically-simple, mass-reduced teeth; they are plotted in a tightcluster on the right of Figure 2. With the exception of Meroitic, X-Group, and ChristianNubians (top of North African cluster), no strong intra-region trends are evident. Theaverage North African MMD is only 0.019, which suggests much greater intra-regiondental homogeneity than among the Sub-Saharan samples. Finally, a noticeable dichotomy


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is apparent between the Sub-Saharan and North African MDS groupings. The much higherMMD value between samples from the two regions is 0.123.African and Non-African Sample Comparisons

Using the observed regional dental differentiation as a baseline, the next step in analysisinvolved combining the homogeneous individual samples into two respective large groups:Sub-Saharan and North Africa. This pooling was effected to: (1) determine collectiveregional dental frequencies, (2) identify the most diagnostic traits, and (3) provide acorresponding level of comparison with published data from several large, pooled non-African groups (see below).The pooled Sub-Saharan group (n=976) consists of the original 15 samples, plusMesolithic Nubia and the regional Sub-Saharan samples from West, East, and South Africa(refer to Table I). The pooled North African group (n=667) consists of the 13 originalsamples (excluding Mesolithic Nubia) and the regional North African sample. The pooledsamples' trait frequencies were then compared to each other and to published data recordedby CG Turner (1985, 1987, 1992a,b) of five large non-African groups: Europe, SoutheastAsia (Sundadonts), Northeast Asia/New World (Sinodonts), Melanesia, and Australia/Tasmania (see Tables V and VI). Although only qualitative comparisons are presented inthis section, all African/non-African dental relationships summarized below are supportedby inter-sample univariate and multivariate statistical analyses (see Irish, 1993b, 1997,1998b).

Table V lists frequencies of 27 dental traits for the Sub-Saharan and North Africans,as well as Europeans, Sundadonts, and Sinodonts. Table VI lists 14 traits fo r the Africansamples and Australia/Tasmania and Melanesia. The total 36 traits are not comparedbecause Turner did not report the other features in his articles. The ASU System wasemployed in all studies, and the presence/absence breakpoints are listed under each trait'sname on the left of the two tables. Several breakpoints vary between tables, and thisshould be taken into account when comparing trait frequencies (Irish 1997, 1998a).The dentally-complex Sub-Saharan Africans are divergent from all groups. Forexample, a Continuity Chi-square analysis (see Irish, 1993b, 1998a) revealed that 23 of

the 36 traits differ significantly from the North Africans. There are a few similarities toAustralia/Tasmania and perhaps Melanesia (e.g., root traits). Turner (1992a) also notedsome dental similarity between Sub-Saharan Africa and Australia, and others (Giblett,1969; Nurse et ai, 1985; Howells, 1989; Brace and Tracer, 1990) found seeming skeletaland genetic links between the two regions and Melanesia. Cavalli-Sforza et al. (1993)even suggested that after 60,000-55,000 BP, Africans may have developed sea-going skillsthat allowed them to contact Australia. However, a direct (recent) relationship seemsunlikely, and any similarity may instead be due to a shared retention of ancestral features(symplesiomorphy) in the populations (Stringer et ai, 1997; Irish, 1998b).


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254 JOEL D. IRISH

Trait:2Winging UI1(+ = ASU1)Shovel UI1(+ = ASU2-6)Dbl Shovel UI1(+ = ASU2-6)Tuber Dent UI2(+ = ASUl-6)Bushman Can UC(+ = ASUl-3)DistAccRidUC(+ = ASU 1-5)

Hypocone UM2(+ = ASU 1-5)Cusp 5 UM1(+ = ASUl-5)Carabelh's UM1(+ = ASU2-7)Parastyle UM3(+ = ASUl-5)Enamel Ex t UM1(+=ASUl-3)Root No UP1(+ = ASU2+)

Root No. UM2(+ = ASU3+)OdontomePl-2(+=ASU+)Peg-red abs UM3(+ = ASUP,R,C)Ling Cusp LP2(+ = ASU2-9)

Groove Pat LM2(+ = ASUY)Cusp No LM1(+=ASU6+)

Sample:%N%N%N%N%N%N%N%N%N%N%N%N%N%N%N%N%N%N

SSA6.674 2

28.141 31.143 761.245 4

18 158671.848 399 077 232 861951268 32.055094574

58 957083 7503047565470868 553052.4617166561

NAF7.4460

19 51548.6175

58 71896.126134.919595 744618 535754.73311233268503

57 146878 637402441

18.354572.6270

30 64027.7352

EUR9.0129

17.014123.313738 11524.8146

51.78979.422813.721247.42304.513443 728634.023853.8197

1.217120.724162 915922 921479178

SUN22.821979 220214 920158 11862.0245

65 013992 041430 037030.6427

1.315357 5388

40 844173 9333

1.414644030079 127819634235 5282

SIN414195498 8192271.0178164 2199012228073 9107390.2363919 0281732 1319448240068 5513512 247575183718462738

22.5462347.2239310 9378347 82947

Table V. Comparison of 27 dental crown and root traits in Sub-Saharan Africa (SSA), NorthAfrica (NAF), Europe (EUR), Sundadonts (SUN), and Sinodonts (SIN)1


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Trait:2Cusp No. LM2(+ = ASU4)DefWrinkle LM1(+ = ASUl-3)C1-C2 Crest LM1(+=ASU+)Protostylid LM1(+ = ASUl-6)Cusp 7 LM1(+=ASUl-4)Tome's Root LP1(+ = ASU3-5)Root No. LC(+ = ASU2+)Root No LM1(+=ASU3+)Root No LM2(+=ASU2+)

Sample:%N%N%N%N%N%N%N%N%N

SSA24.158530.1432

1.344721.055638 559822.436100333

1.740993.3388

NAF66 438124.726 73.327632.53519.44148.63722.33471.2337

88.3333

EUR65.118930.91498.6185

20.0200582237.41484.82070825471.4224

SUN40.431755.31617.39630 03377.43679.8133002079.3343

81.5297

SIN7.9358370.718175.4282534 737399839988228830.53722

13 8519265 54346

Table V. (Cont.). uData for SUN (Southeast Asia) and SIN (Northeast Asia and all New Worldnatives) and standard rank-scaled breakpoints from Turner (1985). Data for EUR from Turner (1985)and from Turner's unpublished Poundbury data. All frequencies are of individual counts.The Sub-Saharan Africans are least like Sinodonts, despite the fact that both groupshave morphologically-complex, mass-additive teeth relative to other world populations.Indeed, the two appear to be at opposite ends of a dental morphological spectrum formany traits. Based on the present data (Table V), Sub-Saharan Africans exhibit the highestfrequencies of Bushman Canine, two-rooted UP1, three-rooted UM2, LM2 Y-groove,LP1 Tome'sroot, two-rooted LM2, and M3 presence. Sinodonts have the lowest or second

lowest frequencies for these traits. Conversely, Sub-Saharan Africans have the lowest oramong the lowest frequencies of such complex traits as UI1 winging, UI1 double shovel,UM1 enamel extension, premolar odontomes, LM1 deflecting wrinkle, and-three-rootedLM1; Sinodonts have the highest frequencies.Lastly, with few exceptions (e.g., Europe fo r UM2 hypocone, 4-cusped LM2, 2-rootedLC), the 27 North African, European, and Sundadont trait frequencies are intermediate tothose of extreme Sub-Saharan Africans and Sinodonts (Table V). Those groups that aregeographically closer to one another display greater dental similarity, and vice versa. Forexample, disparate Sub-Saharan and Sinodont samples are farthest apart geographically.Conversely, those samples that are closer to Sub-Saharan Africa are more dentally similarto the latter group. Thus, these relationships may identify an expansive dental


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256 JOEL D. IRISH

Trait (* >Shovel UI1(+ = ASU3-6)Dbl Shovel UI1(+ = ASU2-6)Int Groove UI2(+ = ASU+)Bushman Can UC(+ = ASU 1-3)Cusp 5 UM1(+ = ASU 1-5)Enamel Ex t UM1(+ = ASU2-3)Root No UM2(+ = ASU3+)Peg-red abs UM3(+ = ASUP,R,C)Cusp No LM2(+ = ASU4)Def Wrinkle LM1(+ = ASU3)Cusp 7 LM1(+ = ASUl-4)Tome's Root LP1(+=ASU3-5)Root No LM1(+=ASU3+)Root No LM2(+=ASU2+)

Sample:%N%N%N%N%N%N%N%N%N%N%N%N%N%N

SSA534131 1437

13 447118 158632 86190357483 7503547082415852343238 559822.4361

1740993 3388

NAF321548.6175

36 12086126118.535730503

78 637418354566 4381042679.4414863721233788 3333

AUT159447043216515060

68.28881208

82 6178642207297

17 1358.297216835.7157914152

MEL9311850118

18.41473215662.42214425073 324012 627053 021917 613611724012 11163222294 7209

Table VI. Comparison of 14 dental crown and root traits in Sub-Saharan Africa (SSA), NorthAfrica (NAF), Australia-Tasmania (AUT), and Melanesia (MEL). Data for AUT and MEL,and standard rank-scaled breakpoints from Turner (1987, 1992a).

to the latter group. Thus, these relationships may identify an expansive dentalmorphological cline that extends from Sub-Saharan Africa through North Africa intoEurope, Southeast Asia, and Northeast Asia and the New World (Irish, 1993b, 1997,1998a,b). This finding, based on dental data, supports a similar conclusion by Cavalli-Sforza et al. (1993) and others using gene frequency data.


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North Africans exhibit a marked divergence in several traits from the Sundadonts andSinodonts (e.g., UI1 winging, UI1 shoveling, UC distal accessory ridge, six-cusped LM1,etc.), as well as Australia/Tasmania and Melanesia samples (e.g., UM1 cusp 5, UM3agenesis, LM1 deflecting wrinkle, four-cusped LM2). However, they show a strong affinityto Europeans, and perhaps West Asians based on recent work by Lipschultz (1996), sharingmany traits involving dental simplification and mass reduction. These traits (Table V)include comparable frequencies of four-cusped LM2 and M3 agenesis, as well as UM1Carabelli's trait, two-rooted LC, and relatively low frequencies of UI1 shoveling, UCdistal accessory ridge, UM1 cusp 5, premolar odontomes, six-cusped LM1, LM1 deflectingwrinkle, and LP1 Tome's root. North Africans also exhibit rocker jaw (see Table IV), atrait known to commonly occur in Europeans (Turner and Markowitz, 1990).

Any North African deviations away from a simple dental pattern are in the directionof complex Sub-Saharan traits (e.g., UI2 tuberculum dentale, Bushman Canine, two-rootedUP1, three-rooted UM2), suggesting likely gene flow from these peoples. These findingsagree with genetic-based results that link North Africans to Europeans and western Asians,yet record many Sub-Saharan influenced traits (e.g., Mourant 1954, 1983; Hiernaux 1975;Nurse et al., 1985; Sanchez-Mazas et ai, 1986; Excoffier et ai, 1987; Roy choudhury andNei 1988).Characteristic Sub-Saharan African Dental Traits

Based on this seven-sample dental comparison, I proposed (Irish, 1997) that a tentative"Sub-Saharan African Dental Complex" (suite of traits that best differentiates them fromother world populations) consists of a recurrent combination of two low- and nine high-frequency traits (see below). A few traits are present in comparable frequencies elsewhere(e.g., North Africa and Europe fo r UM1 Carabelli's; Australia/Tasmania and Melanesiafo r LM2 root number), but the prevalence of most differs significantly relative to otherworld groups. Thus, the relevant combination of all 1 1 traits clearly denotes a Sub-Saharanpattern.This Complex includes among the world's lowest frequencies of UI1 double shovelingand UM1 enamel extension, and among the world's highest frequencies of BushmanCanine, two-rooted UP1, UM1 Carabelli's trait, three-rooted UM2, LM2 Y-groove, LM1cusp 7, LP1 Tome's root, two-rooted LM2, and M3 presence (or correspondingly, thelowest frequency of M3 absence). In a few cases, LM1 cusp 7, LP1 Tome's root, andBushman Canine may not be expressed in all Sub-Saharan samples (e.g. , Nguni, San, andCongo) (refer back to Table III). However, these exceptions involve small sample sizeswhich are likely not representative of the larger populations. These traits are ubiquitous inall remaining Sub-Saharan samples.Sub-Saharan Africans also have notable frequencies of UI1 labial curvature and UI1midline diastema. Unfortunately, these features are not routinely recorded in other dentalstudies, so an African/non-African comparison can not be attempted to determine if theytoo represent diagnostic African traits.


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258 JOEL D. IRISH

Characteristic North African Dental TraitsAs described by others (Hiernaux, 1975; Excoffier et al, 1987; Roychoudhury andNei, 1988; Lipschultz, 1996; among others) and as noted in this and previous studies(Irish, 1993b, 1997, 1998a), North Africans are genetically and phenetically allied withEuropeans and western Asians. North African dental frequencies are similar to those ofEuropeans, except for some traits that show apparent sub-Saharan influence. Such a North/Sub-Saharan African combination is also evident in many genetic systems (e.g.,Roychoudhury and Nei, 1988). Thus, I proposed (Irish, 1993b, 1998a) that the NorthAfrican dental trait complex is one which parallels that of Europeans, yet displays higherfrequencies of Bushman Canine, two-rooted UP1, three-rooted UM2, LM2 Y- groove,LM1 cusp 7, LP1 Tome's root, two-rooted LM2, and lower frequencies of UM1 enamel

extension and peg/reduced or absent UM3. North Africans also exhibit a higher frequencyof UM1 Carabelli's trait than sub-Saharan Africans or Europeans.

CONCLUSIONThe findings summarized in this article have been provided to better dentallycharacterize the peoples of Sub-Saharan and North Africa, and allow potential new insightinto their phenetic relationships. Sub-Saharan and North African intra-region homogeneitycontrasts sharply with significant inter-regional differences. From a global perspective,dentally-simple North Africans resemble Europeans and western Asians to some degree,whereas Sub-Saharan-affiliated Africans differ greatly from all other world groups with only superficial similarities to Australian/Tasmanians and perhaps Melanesians. Sub-Saharan peoples are particularly distinctive in their expression of numerous,morphologically-complex dental crown and root traits that are absent or found in lowfrequencies elsewhere.Future analyses will benefit by the examination of additional African and non- Africansamples particularly if the latter are subjected to a more specific level of inquiry. Inother words, the same type of investigation as that applied to the individual African samples,or to individual Asian and New World samples (per Turner, 1984, 1985, 1987, 1990,1992a,b), will provide much greater detail concerning African/non-African dentalrelationships. Comparing large pooled samples, as necessitated by the use of publisheddata, provides an initial idea of world-wide biological affiliation, but the use of manysmaller, regional samples will allow finer resolution. Such an approach should also affordadditional evidence to (1) test the credibility of the proposed African dental complexes,and (2) perhaps help to further our understanding of modern human migrations and originsbased on recent population relationships (see Stringer, 1993; Irish, 1998b).


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AFRICAN DENTAL AFRNITIES 259

AcknowledgmentsI am indebted to Alain Froment from the Laboratoire ORSTOM-ERMES, Orleans fo rthe article's French summary and for his encouragement. My appreciation is also extendedto Christy G. Turner II from Arizona State University; the present dental inquiry wasmodeled after his studies of populations from Europe, Asia, the New World, and theSouth Pacific. Lastly, I thank the many people at the seven institutions I collected datafrom, including: Charles Merbs and Donald Morris from Arizona State University, Tempe,AZ; Guy Gibbon and the late Elden Johnson from the University of Minnesota,Minneapolis, MN; Douglas Ubelaker and David Hunt from the National Museum ofNatural History, Washington, DC; Ian Tattersall, Jaymie Brauer, and Gary Sawyer fromthe American Museum of Natural History, New York; Andre Langaney, Frances Roville-

Sausse, and Miya AwazuPeriera da Silva from the Muse de l'Homme, Paris; Fred Wendorfand Sue Linder-Linsley from Southern Methodist University, Dallas, TX, and Henry deLumley and Dominique Grimaud-Herve from the Institut de Palontologie Humaine, Paris.Research was supported by grants from the National Science Foundation (BNS-90 13942),the Arizona State University Research Development Program, and the American Museumof Natural History.


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BIBLIOGRAPHY

Barnes (D.S.), 1969. Tooth morphology and other aspects of the Teso dentition. Am. J. Phys.Anthropol. 30: 183-194.Bermudez de Castro (J.M.), 1991. La denture de la population Msolithique d'Afrique du Nord etl'hypothse "Afro-Europen Sapiens". L'Anthropologie 95: 201-218.Bermudez de Castro (J.M.), 1989. The Carabelli trait in human prehistoric populations of theCanary Islands. Hum. Biol. 61: 117-131.Berry (A.C.), Berry (R.J.), 1967. Epigenetic variation in the human cranium. J. Anat. 101: 361-379.Brabant (H.), 1965. Observations sur la denture des Pygmes de l'Afrique centrale. Bull. Group.Int. Rech. Se. Stomat. 8: 27-49.Brace (C.L.), Tracer (D.P.), 1990. Craniofacial change and continuity: A comparison of latePleistocene and recent Europe and Asia. Presented at University of Tokyo Symposium: TheEvolution and Dispersal of Modern Humans in Asia, November 14-17.Camps (G.), 1975. The prehistoric cultures of North Africa: Radiocarbon chronology. In: F. Wendorf,A.E. Marks (eds.) Problems in Prehistory: North Africa and The Levant. Southern MethodistUniversity Press, Dallas, pp . 181-192.Cavalli-Sforza (L.L.), Menozzi (P.), Piazza (A.), 1993. Demie expansions and human evolution.Science 259: 639-646.Chagula (W.K.), 1960. The cusps on the mandibularmolars of east Africans. Am. J. Phys. Anthropol.18: 83-90.Charon (M.), Hugot (H.), Petit-Maire (N.), 1974. Les restes humains de Tagdait (Ahaggar). Bull.Mm. Soc. Anthropol. Paris, 13: 151-155.Drennan (M.R.), 1929. The dentition of a Bushman tribe. 5. Afr. Mus., Cape TownAnnals. 24: 61-87.Ehret (), 1982. Linguistic inferences about early Bantu history. In: Ehret and M Posnansky(eds.) The Archaeological and Linguistic Reconstruction ofAfrican History. Univ. of Cal.Press, Berkeley, pp . 57-65.Excofher (L.), Pellegrini (.), Sanchez-Mazas (A.), Simon (C), Langaney (A.), 1987. Geneticsand history of sub-Saharan Africa. Yrbk. Phys. Anthropol. 30: 151-194.


26/37

AFRICAN DENTAL AHNITIES 26 1

Franciscus (R.G.), 1995. Later Pleistocene Nasofacial Variation in Western Eurasia and Africaand Modern Human Origins. Ph.D. Diss., Univ. of New Mexico, Albuquerque.Froment (A.), 1992a. La diffrenciation morphologique de l'Homme moderne: Congruence entreforme du crne et rpartition gographique du peuplement. Comptes-Rendus Acad. Sci. 3:323-329.

Froment (A.), 1992b. Origines du peuplement de l'Egypte ancienne: L'apport de anthropobiologie.Archo-Nil 2: 79-98.Galloway (A.), 1959. The Skeletal Remains of Bambandyanalo. Witwatersrand University Press,Johannesburg.Giblett (E.R.), 1969. Genetic Markers in Human Blood. Blackwell, Oxford.Green (R.), Suchey (J.), 1 976. The use of inverse sine transformation in the analysis of non-metricaldata. Am . J. Phys. Anthropol. 45: 61-68.Greenberg (J.H.), 1959. Africa as a linguistic area. In: W.R. Bascom, M.J. Herskovits (eds.)Continuity and Change in African Cultures. Chicago: University of Chicago, pp. 15-27.Greenberg (J.H.), 1966. The Languages ofAfrica. Indiana University, Bloomington.Greene (D.L.), 1967. Dentition of Meroitic, X-Group, and Christian Populations From Wadi Haifa,Sudan. Anthropol. Paper 85 , Nubian Ser. J. University of Utah Press, Salt Lake City.Greene (D.L.), 1972. Dental anthropology of early Egypt and Nubia. /. Hum. Evol. 1: 315-324.Greene (D.L.), 1982. Discrete dental variations and biological distances of Nubian populations.Am. J . Phys. Anthropol. 58: 75-79.Greene (D.L.), Ewing (G.H.), Armelagos (G.J.), 1967. Dentition of a Mesohthic population fromWadi Haifa, Sudan. Am. J. Phys. Anthropol. 27: 41-56.HaeussLER (A.M.), Irish (J.D.), Morris (D.H.), Turner (C.G. II), 1989. Morphological and metricalcomparison of San and Central Sotho dentitions from southern Africa. Am . J . Phys. Anthropol.78: 115-122.Hanihara (T.), 1992. Dental and cranial affinities among populations of East Asia and the Pacific:The basic populations in East Asia, IV. Am. J. Phys. Anthropol. 88: 163-182.Hassanali (J.), 1982. Incidence of Carabelli's trait in Kenyan Africans and Asians. Am . J. Phys.Anthropol. 59:317-319.Hassanali (J.), Mwaniki (D.), 1984. Palatal analysis and osteology of the hard palate of the KenyanAfrican skulls. Anat. Rec. 209: 273-280.


27/37

262 JOEL D. IRISH

Hiernaux (J.), 1975. The People ofAfrica. Charles Scribner's Sons, New York.Holliday (T.W.), 1995. Body Size and Proportions in the Late Pleistocene Western Old World andthe Origins ofModern Humans. Ph.D. Diss, Univ. of New Mexico, Albuquerque.Howells (W.W.), 1989. Skull Shapes and the Map: Craniometric Analyses in the Dispersion ofModern Homo. Papers of the Peabody Museum of Archaeology and Ethnology, HarvardUniversity, Vol. 79, Cambridge.Irish, (J.D.), 1998a. Diachronic and synchronie dental trait affinities of Late and post-Pleistocenepeoples from North Africa. Homo 49: 138-155.Irish (J.D.), 1 998b. Ancestral dental traits in recent Sub-Saharan Africans and the origins of modernhumans. J. Hum. Evol. 34: 81-98.Irish (J.D.), 1997. Characteristic high- and low-frequency dental traits in Sub-Saharan Africanpopulations. Am. J. Phys. Anthropol. 102: 455-467.Irish (J.D.), 1994. The African dental complex: Diagnostic morphological variants of modern sub-Saharan populations. Am. J. Phys. Anthropol. Supplement 18 : 112 (Abstract).Irish (J.D.), 1993a. Dental morphometric affinity of Canary Islanders with North African Maghrebpopulations. Am. /. Phys. Anthropol. Supplement 16: 114 (Abstract).Irish (J.D.), 1993b. Biological Affinities of Late Pleistocene through Modern African AboriginalPopulations: The Dental Evidence. Ph.D. Dissertation, Arizona State University, Tempe.Ann Arbor: University Microfilms.Irish (J.D), Turner (C.G. II), 1992. Further evidence of dental discontinuity between Mesolithicand recent Nubians. Am. J. Phys. Anthropol. Suppl. 14: 93-94.Irish (J.D.), Turner (C.G. II), 1990. West African dental affinity of late Pleistocene Nubians: Peoplingof the Eurafncan-South Asian triangle II. Homo 41: 42-53.Irish (J.D.), Turner (C.G. II), 1989. Dental affinity of late Pleistocene Nubians and historic WestAfricans. Am. J . Phys. Anthropol. 78: 245.Jacobson (A.), 1982. The Dentition of the South African Negro. Higgenbotham, Inc, Anniston,AL.Johnson (A.L.), Lovell (NC), 1994. Biological differentiation at Predynastic Naqada, Egypt: Ananalysis of dental morphological traits. Am . J. Phys. Anthropol. 93: 427-433.Kruskal (J.B.), Wish (M.), 1978. Multidimensional Scaling. Sage Publications, Beverly Hills, CA.Lipschultz (J.G.), 1996. Who Were the Natufians? A Dental Assessment of their PopulationAffinities.

M.A. Thesis, Arizona State University, Tempe.


28/37


Lubell (D.), 1984. Paleoenvironments and Epi-Paleolithic economies in the Maghreb (ca. 20,000-5,000 B.P.). In- : JD Clark and SA Brandt (eds.) From Hunters to Farmers: The Causes andConsequences of Food Production in Africa. University of California Press, Berkeley, pp.41-56.

Lukacs (J.R.), Hemphill (B.E.), 1991. The dental anthropology of prehistoric baluchistan: Amorphometnc approach to the peopling of South Asia. In: M.A. Kelley, C.L. Larsen (eds.)Advances in Dental Anthropology. Wiley-Liss, New York, pp . 77-119.Mourant (A.E.), 1954. The Distribution of the Human Blood Groups. Charles Thomas, Publisher,Springfield, 111.Mourant (A.E.), 1983. BloodRelations: Blood Groups and Anthropology. Oxford University Press,Oxford.Murdock (G.P.), 1959. Africa: Its Peoples and Their Culture History. McGraw-Hill, New York.Nurse (G.T.), Weiner (J.S.), Jenkins (T.), 1985. The Peoples ofSouthern Africa and TheirAffinities.Clarendon Press, Oxford.Pales (L.), 1938. Contribution l'Etude anthropologique des Babinga de l'Afrique quatorialefranaise. L'Anthropologie 48: 503-520.Petit-Maire (N.), 1979. Le Sahara Atlantique l'Holocne. Peuplement et Ecologie. Mmoires duCentre de Recherches Anthropologiques, Prhistoriques et Ethnographiques, No. 28. Algiers:Socit Nationale d'Editions et de Diffusion.Phillipson (D.W.), 1994. African Archaeology. Cambridge University Press, New York.Roler (K.L.), 1992. Near Eastern Dental Variation Past and Present. M.A. Thesis. Arizona StateUniversity, Tempe, AZ.Roychoudhury (A.K.), Nei (M.), 1988. Human Polymorphic Genes World Distribution. OxfordUniversity Press, New York.Sakuma (M.), Ogata (T.), 1987. Sixth and seventh cusp on lower molar teeth of Malawians in east-central Africa. Japan. J. Oral. . 29: 738-745.Sakuma (M.), Ogata (T.), 1988. The protostylid on teeth of Malawians, and the Mongoloid dentalcomplex. Japan. J. Oral Biol. 30: 70-74.Sakuma (M.), Irish (J.D.), Morris (D.H.), 1991. The Bushman maxillary canine of the ChewaTribe in east-central Africa. J. Anthropol. Soc. Nippon. 99: 411-417.Sakuma (M.), Mine (K.), Ogata (T.), 1988. The incidence and expression of Carabelli complex inMalawians and Japanese. Japan. J. Oral Biol. 30: 545-549.


29/37

264 JOEL D. IRISH

Sakuma (M.), Yamamoto (M.), Ogata (T.), 1987. On the deflecting wrinkle in the lower molars ofMalawians derived from pureblooded Negroid racial stock. Japan. J. Oral Biol. 29: 371-377.Sanchez-Mazas (A.), Excoffier (L.), Langaney (A.), 1986. Measurement and representation of thegenetic similarity between populations by the percentage of isoactive genes. Theona 4: 143-154.Scott (G.R.), 1973. Dental Morphology:A Genetic Study ofAmerican White Families and Variationin Living Southwest Indians. Ph.D. Dissertation. Arizona State University, Tempe, AZ.Scott (G.R.), 1980. Population variation of Carabelli's trait. Hum. Biol. 52: 63-78.Scott (G.R.), Yap Potter (R.H.), Noss (J.F.), Dahlberg (A.A.), Dahlberg (T.), 1983. The dentalmorphology of Pima Indians. Am. J. Phys. Anthropol. 61: 13-31.Shaw (J.C.M.), 1 93 1 . The Teeth, the Bony Palate and the Mandible in Bantu Races of South Africa.John Bale, Sons and Danielson, Ltd, London.Sheppard (P.J.), 1987. The Capsian ofNorth Africa: Stylistic Variation in Stone ToolAssemblages.BAR International Press, Oxford.Sj0vold (.), 1973. Occurrence of minor non-metrical variants in the skeleton and their quantitative

treatment for population comparisons. Homo 24: 204-233.Sj0vold (.), 1977. Non-metrical divergence between skeletal populations. Ossa 4: 1-133.Smith (P.), Shegev (M.), 1988. The dentition of Nubians from Wadi Haifa, Sudan: An evolutionaryperspective. /. Dent. Assoc. S. Afr. 43: 539-541.Sokal (R.R.), Sneath (P.A.), 1963. Principles of Numerical Taxonomy. W.H. Freeman and Company,San Francisco.Stringer (C.B.), Humphrey (L.T.), Compton (T.), 1997. Cladistic analysis of dental traits in recent

humans using a fossil outgroup. J . hum. Evol. 32: 389-402.Stringer (C.B.), 1993. Understanding the fossil human record: Past, present and future. Riv.Anthropol. [Roma] 71: 91-100.Tobias (P.V.), 1972. Recent human biological studies in southern Africa, with special reference toNegros and Khoisans. Trans. Roy. Soc. S. Afr., Part 3 40: 109-133.Turner (C.G. II), 1992a. The dental bridge between Australia and Asia: Following Macintosh intothe East Asian hearth of humanity. Persp. Hum. Biol. 2/Archaeol. Oceania 27: 120-127.


30/37


Turner, (C.G. II), 1992b. Microevolution of East Asian and European populations: A dentalperspective. In: T. Akaszawa, K. Aoki, T. Kimura (eds.) The Evolution and Dispersal ofModern Humans in Asia. Hokusen-Sha Publishing Co., Tokyo, pp. 415-438.

Turner (C.G. II), 1990. Major features of Sundadonty and Sinodonty, including suggestions aboutEast Asian microevolution, population history, and late Pleistocene relationships withAustralian Aboriginals. /n. J. Phys. Anthropol. 82: 295-318.Turner (C.G. II), 1987. Late Pleistocene and Holocene population history of East Asia based ondental variation. Am. J. Phys. Anthropol. 73: 305-322.Turner (C.G. II), 1986. Dentochronological separation estimates for Pacific Rim populations. Science

232: 1140-1142.Turner (C.G. II), 1985. The dental search for Native American origins. In: R. Kirk, E. Szathmary(eds.) Out of Asia: Peopling the Americas and the Pacific. The Journal of Pacific History,Canberra, pp . 31-78.Turner (C.G. II), 1984. Advances in the dental search for Native American origins. AdaAnthropogenetwa 8: 23-78.Turner (C.G. II), Markowitz (M.), 1990. Dental discontinuity between late Pleistocene and recentNubians. I. Peopling of the Eurafncan-South Asian Tnangle. Homo 41 : 42-53.Turner (C.G. II), Nichol (C.R.), Scott (G.R.), 1991. Scoring procedures for key morphologicaltraits of the permanent dentition: The Arizona State University dental anthropology system.In: M.A. Kelley, C.S. Larsen (eds.) Advances in Dental Anthropology. Wiley-Liss, NewYork, pp. 13-32.Van Reenen (J.F.), 1966. Dental features of a low-caries primitive population. J. Dent. Res. Sup. toNo. 3 45: 703-713.Vaufrey (R.), 1933. Notes sur le Capsien. L'Anthropologie 43: 457-483.Waiters (J.), 1962. Extra dentition in the west African Negro. Mankind Quarterly. 2: 263-264.Wendorf (E), 1 968. A Nubian final Paleolithic graveyard near Jebel Sahaba, Sudan. In: F. Wendorf(ed.) The Prehistory of Nubia, Volume Two. Fort Burgwin Research Center, Dallas, TX, pp.954-995.Wulsin (F.R.) 1 941 . Th e PrehistoricArchaeology of Northw est Africa. Peabody Museum of AmericanArchaeology and Ethnology, Harvard University, Cambridge MA.

Reu le 08/01/1998, accept le 15/06/1998.


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APPENDIXARIZONA STATE UNIVERSITY DENTAL ANTHROPOLOGY SYSTEMAND OTHER TRAITS USED IN THE PRESENT STUDY

Maxillary TraitsWinging UI1Upper central incisors may be rotated mesiolingualward, giving a V-shaped appearance whenviewed from the occlusal surface. No reference plaque. Four possible ASU grades may occur:1. Bilateral winging2. Unilateral winging3. No expression4. Counter wingingLabial Curvature UI1Labial surface of the tooth may display a notable convex curvature. Reference plaque ASU UI1labial curvature grades scored as:0. No expression1 . Trace curvature2. Weak curvature3. Moderate curvature4. Strong curvaturePalatine TorusLinear bony exostosis that may develop along the palatine suture (in adults only). No referenceplaque. Five possible ASU grades are:0. No expression1. Trace (1-2 mm elevation)2. Medium (2-5 mm)3. Marked (> 5 mm)4. Very marked (may be as high as 10 mm)Shovel UUPossible presence of mesial and distal vertical ridges on lingual surface, giving the tooth ashovel-like appearance. Six grades may be scored with reference plaque ASU UI1 shovel:0. No expression1. Faint ridges2. Trace ridges3. Semi-shovel shaped4. Shovel-shaped5. Marked shovelling


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Double-Shovel UI1Mesial and distal marginal ridges may be present on the labial surface. Six possible grades havebeen established on reference plaque ASU UI1 double-shovel:0. No expression1. Trace ridge on one margin2. Trace ridges on both margins3. One moderate and one trace ridge4. Two moderate ridges5. One large and one moderate ridge6. Two large ridges

Interruption Groove UI2Groove on lingual borders of teeth. No reference plaque. Graded as absent or present (andlocation).

Tuberculum Dentale UI2Ridging or cusp formation may occur on the mediolingual surface. There are eight possiblegrades using plaques ASU UC tuberculum dentale (grades 1 -4), and ASU UC distal accessory ridge(grades 5-6):0. No expression1 . Faint ridging2. Trace ridging3. Strong ridging4. Pronounced ridging5-. A weakly developed cuspule5. Weakly developed cuspule with free tip6. Strong cusp with free tip

Mesial Ridge UC (Bushman Canine)Mesiohngual ridge which may be notably larger than the distolingual ridge may incorporatethe tuberculum dentale. Called "Bushman Canine" in the present study after Morris (1975). Fourpossible grades may be scored with reference plaque ASU UC mesial ridge:0. No expression1 . ML ridge larger than DL, and weakly attached to the tuberculum dentale2. ML ridge larger than DL, and moderately attached to the tuberculum dentale3. ML ridge is much larger than the DL, and is fully incorporated into the tuberculumdentale

Distal Accessory Ridge UCAnterior to upper canine distal marginal ridge, another disto-lingual ridge can be found. Thisfeature can be very pronounced. The six possible ASU grades on reference plaque DAR UC are:0. No expression1. Ridge is very faint2. Ridge is weakly developed3. Ridge is moderately developed


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4. Ridge is strongly developed5. Ridge is very largeHypocone UM2Cusp 4 may range from absent to large and developed. Seven possible grades exist on referenceplaque ASU UM hypocone:0. No expression1 . Faint ridge present2. Faint cuspule present3. Small cusp present3.5.Moderate-sized cusp present4. Large cusp present5. Very large cusp presentCusp 5 (Metaconule) UM1Possible presence of a fifth cusp between the third and fourth cusps. There are six possiblegrades on reference plaque ASU UM cusp 5:0. No expression

1 . Tiny round cusp2. Tiny wedge-shaped cusp3. Small cusp4. Medium-sized cusp5. Large cuspCarabelli's Trait UM1

If present, the mesiolingual aspect of upper molars may display a range of variation from afurrow to a large free cusp. An eight-grade classification, originated by Dahlberg (1956), is usedwith reference plaque Zoller Laboratory UM Carabelli cusp:0. No expression1 . Furrow2. Pi t3. Double furrow4. Small attached cusp5. Large attached cusp6. Small free cusp7. Large free cuspParastyle UM3If present, the buccal surface may display variation from a pit to a free cusp. There are sixgrades on the reference plaque ASU UM parastyle:0. No expression1. Pit2. Small attached cusp3. Small free cusp4. Medium-sized free cusp5. Large free cusp


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Enamel Extension UM1An extension of the enamel border may be present and extend toward the root apex. No referenceplaque. Four possible ASU grades may be scored:0. No expression1. A short extension (up to 1 mm)2. A medium extension (up to 2 mm)3. A lengthy extension (up to 4 mm +)

Root Number UP1Number of free roots. No reference plaque. Graded according to number of roots present.Root Number UM2

Number of free roots. No reference plaque. Graded according to number of roots present.Peg-Reduced UI2The tooth may be very reduced in size and display very simple morphology. No referenceplaque. This trait is graded as normal or peg-shaped.OdontomePl-P2Any pin-sized, spike-shaped enamel and dentine projection occurring on occlusal surface. Noreference plaque. Scored as either present or absent.Congenital Absence UM3

Tooth may not be formed in adult individuals. No ASU plaque. Scored as tooth present orabsent.Midline DiastemaIn addition to the Arizona State University dental anthropology system traits, the occurrence ofthe UI1 midline diastema was also recorded. Previous research by the author suggests that a present/absent level of dichotomization is sufficient to record this metric feature, which is based on ameasurement between the upper central incisors:0. No diastema (space < .5 mm)1. Diastema (space > .5 mm)The midline diastema has been shown to occur in high frequencies in many aboriginal Africanpopulations (Shaw, 1931; Deverall, 1949; Sperber, 1958; Van Reenen, 1964; Hassanali, 1982;Jacobson, 1982), yet is unusual outside Africa. Thus, the feature may prove to be a useful Africanmarker.Mandibular TraitsLingual Cusp Number LP2The number of lingual cusps present are recorded. Four possible grades exist on referenceplaque ASU LP2: 0-3 cusps.


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Anterior Fovea LM1A depression that can occur anterior to cusps 1 and 2. It can range in expression from absent, toa large depression with a ridge connecting the mesial margins of the tw o cusps. There are fivepossible grades on reference plaque ASU LM1 anterior fovea:0. No expression1 . Faint depression anterior to cusps2. Small depression3. Medium depression4. Large depressionMandibular TorusA nodular bony exostosis that may develop on the lingual side of the mandible in region of thecanine and premolars. No reference plaque. Four possible ASU grades exist:

0. No expression1. Trace elevation2. Elevation between 2 to 5 mm3. Elevation greater than 5 mmGroove Pattern LM2Pattern created on the occlusal surface from cusps. No reference plaque. There are three possiblegrades: Y. Cusps 2 and 3 touchX. Cusps 1 and 4 touch+. Cusps 1 through 4 touchRocker JawInferior surface curvature of the mandible's horizontal ramus. This age-dependent feature occursonly in adults. No reference plaque. There are three possible ASU grades:0. No expression1. Slight curvature of the jaw2. Extreme curvature, allowing the jaw to rock back and forth when placed on a flat surfaceCusp Number LM1Number of cusps present, excluding the metaconulid (cusp 7). No reference plaque. Threepossible grades (4-6 cusps).Cusp Number LM2Number of cusps present, excluding the metaconulid. No referenceplaque. Three possible grades(4-6 cusps).Deflecting Wrinkle LM1Medial ridge on occlusal surface may be present and vary on cusp 2. Expression can range fromabsent, to a large ridge which may make contact with cusp 3. There are four possible grades onreference plaque ASU LM deflecting wrinkle:0. No expression1. Ridge extends 1/2 way across the cusp


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2. Ridge extends completely across the cusp3. Ridge extends into the central grooveC1-C2 (Distal Trigonid) Crest LM1A ndge or loph may connect the distal borders of cusps 1 and 2. This trait is scored as present orabsent with the aid of a reference plaque developed by Hanihara (1961) for deciduous teeth.

ProtostylidLMlA paramolar cusp that may occur on the mesiobuccal surface of cusp 1 . The trait is often associatedwith the buccal groove, and can range from a pit to a free cusp. Eight possible ASU grades existusing the reference plaque Zoller Laboratory LM protostylid:0. No expression1. Buccal pit2. Distal deviation of the buccal groove3. Secondary mesial groove occurs4. Secondary groove is larger than 35. Secondary groove is larger than 46. Small cusp7. Large cuspCusp 7 (Metaconulid) LM1Cusp may be present in the lingual groove between cusps 2 and 4. Six possible grades can bescored with reference plaque ASU LM cusp 7:0. No expression

1 . Faint cusp1 A. Faint bulge on the lingual surface of cusp 22. Small cusp3. Medium-sized cusp4. Large cuspTome's Root LP1Condition when the mesial and distal root surfaces may be deeply grooved. There are six possiblegrades on reference plaque ASU LP Tome's root:0. No expression1. Shallow groove is present

2. Moderate groove is present3. Deep groove is present4. Very deep groove is present5. Two free roots are presentRoot Number LCNumber of free roots. No reference plaque. Graded according to number of roots present.Root Number LM1Number of free roots. No reference plaque. Graded according to number of roots present.


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Root Number LM2Number of free roots. No reference plaque. Graded according to number of roots present.TorsomolarAngle LM3The tooth may be rotated relative to a line drawn through the middle of the first and secondmolars. No reference plaque. Three possible ASU grades exist: straight, buccal rotation, and lingualrotation. The degree of rotation is also noted (Turner et ai, 1991; Irish, 1993).

Dental Morphological Affinities Late Pleistocene Sub-saharan - Irish

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