Evolution of the Dentition in Prehistoric Ohio Valley NativeAmericans: II. Morphology of the Deciduous Dentition

PAUL W. SCIULLI*Department of Anthropology, Ohio State University, Columbus, Ohio 43210

KEY WORDS deciduous teeth; morphology; Ohio Valley region

ABSTRACT In order to evaluate the microevolutionary dynamics ofmorphological features of the deciduous dentition, I collected data on thevariation of 57 features (33 crown and 24 root) from prehistoric Ohio Valleypopulations. I sampled a total of 370 individuals from 26 populationsrepresenting a lineage that inhabited the middle and upper Ohio valley regionfrom approximately 3000 to 350 BP.

Evolutionary changes in the frequencies of morphological features of thedeciduous teeth in this lineage were limited. Over 80% of the features show nosignificant differences among the populations. The relatively few featuresthat show consistent differences separate pre- and postmaize agriculturalpopulations. I discuss explanations for this change in terms of selectiondifferences or gene flow.

The general pattern of morphological trait expression in the deciduousteeth of this Ohio Valley lineage corresponds to what has been termed theMongoloid dental complex (sinodonty in the permanent teeth). I suggestadditional features that, with further study, may be added to this morphologi-cal complex. Am J Phys Anthropol 106:189–205, 1998. r 1998 Wiley-Liss, Inc.

Almost nothing is known of the microevo-lutionary dynamics of morphological fea-tures of the human deciduous dentition.This situation has arisen primarily as theresult of the difficulty in obtaining represen-tative samples from a sequence of ancestral-descendant populations. Demographic bi-ases, whether the result of differentialpreservation, poor recovery, or the culturalpractices of a population, often limit thenumber of children representing a skeletalsample. In addition, the timing of the devel-opment and replacement of the deciduousteeth usually results in the loss of morpho-logical information for many of the childrenavailable for study. However, while the docu-mentation of evolutionary change in thedeciduous dentition of any given lineage hasbeen sketchy and incomplete, variation inthe expression and frequency of morphologi-cal features of the deciduous teeth has beenrecorded for a number of human populations

(Jorgensen, 1956; Hanihara, 1963, 1967:Smith, 1976, 1978; Sciulli, 1977, 1990a;Lukacs and Walimbe, 1984; Grine, 1986).These studies have shown that the morphol-ogy of the deciduous dentition of humanpopulations exhibits significant within- andbetween-population variation and is thuspotentially liable to microevolutionary forces.

The purpose of this investigation is todocument and evaluate variation in the ex-pression and frequency of 57 morphologicalfeatures of the deciduous dentition in alineage of Native Americans from the middleand upper Ohio Valley. The populations fromwhich the samples are drawn come from anapproximately 2,500 year span of the culture-historic development of the lineage. During

*Correspondence to: Paul W. Sciulli, Department of Anthropol-ogy, Ohio State University, Columbus, OH 43210. E-mail:sciulli.1@osu.edu

Received 4 June 1997; accepted 26 February 1998.

AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY 106:189–205 (1998)

r 1998 WILEY-LISS, INC.

this time span, groups were initially rela-tively small and mobile without ceramicsand subsisted primarily as hunter-gather-ers. Through time, groups developed intolarger, more sedentary populations with awell-developed ceramic technology and anagricultural subsistence base, at first utiliz-ing plants native to the area and laterfocusing on introduced tropical cultigens.The populations representing this lineageduring the period under study have thusexperienced changes in subsistence, technol-ogy, and structure which are commonly asso-ciated with and have been hypothesized tocause microevolutionary changes in the per-manent dentition (Brace et al., 1991). One ofthe principal questions to be addressed hereis to what extent, if any, morphologicalvariation in the deciduous teeth is associ-ated with changes in the cultural accommo-dations of populations.

MATERIAL AND METHODSSamples

I scored a total of 370 individuals formorphological features of the deciduousteeth. The individuals are sampled from 26upper and middle Ohio Valley populationsrepresenting each culture-historic time pe-riod from the terminal LateArchaic (approxi-mately 3000 BP) to the Late Prehistoric(approximately 350 BP). Table 1 contains alisting of the populations by time period andthe number of individuals sampled fromeach. All populations are located in Ohioexcept for the Monongahela, located in adja-cent western Pennsylvania, and Buffalo, lo-cated in adjacent West Virginia. Because anumber of the samples are small and manyindividuals have incomplete data due to thenature of deciduous tooth development andreplacement, I combined the Late Archaicsamples into one group (n 5 64) and thethree Woodland samples into a second group(n 5 34). The Late Prehistoric samples, ex-cept for Anderson Village, are treated sepa-rately. The Anderson Village sample is in-cluded only in calculations and statisticsinvolving the total sample.

The general cultural features of the histori-cal periods from which the populations inTable 1 are sampled have recently beenpresented, so only a brief review is neces-

sary here (Sciulli, 1997). In the terminalLateArchaic period, populations can be char-acterized as relatively small, mobile, ac-eramic, primarily hunter-gatherers. Popula-tions are associated with specific mortuarypatterns, each of which has a generallyexclusive geographical distribution. How-ever, because Late Archaic habitation sitesare unknown, the significance of the varia-tion in mortuary behavior cannot be relatedto the general cultural context of the popula-tions. During the Late Archaic period, plantsnative to the region were domesticated(Smith, 1989) and may have contributed tothe diet of the populations under study. Thefrequencies of deciduous dental pathologieswhich relate most directly to diet (caries0.88%, number of teeth 5 NT 5 793; abscess0%, NL 5 number of loci 5 913; and antemor-tem loss 0%, NL 5 913) are minimal in these

TABLE 1. Population sampled from each culture-historical period of the middle and upper Ohio Valley

region1

Late Archaic (N 5 64) (approximately 3200BP to 2700 BP)

Kirian Treglia (8)Boose (16)Duff (19)Orleton (3)Scioto Country Homes (2)Davis (7)Williams Red Ocher (4)Muzzey Lake (5)

Early Woodland (N 5 20) (approximately2700 BP to 2000 BP)

Galbreath (2)McMurray (2)Sidner I (2)Sidner II (9)Cowan Creek (3)Niles Wolford (1)Teopfner (1)

Middle Woodland (N 5 9) (approximately1900 BP to 1500 BP)

Esch (3)Hopewell (3)Harness (1)Seip (2)

Late Woodland (N 5 5) (approximately 1500BP to 1000 BP)

Voss (1)Baker I (4)

Late Prehistoric (N 5 272) (approximately1000 BP to 350 BP)

Pearson Village (48)Anderson Village (13)Sun Watch (76)Monongahela (62)Buffalo (73)

1 Numbers in parentheses following site name is the sample size;N is the sample size for the period.

190 P.W. SCIULLI

populations, reflecting the presumably smallamount of simple carbohydrate in the diet.All samples included here are from theGlacial Kame or related cultural complexes(Griffin, 1983).

The beginning of the Early Woodland pe-riod in this area is defined by the introduc-tion of ceramics. At first very thick andcrude and probably used only for storage,pottery developed throughout the Woodlandperiod with varieties well designed for cook-ing appearing toward the end of the EarlyWoodland and characterizing all subsequentperiods. The Early Woodland period is alsocharacterized by the elaboration of mortu-ary ceremonialism in the form of conicalmound burial, often of an ornate nature, aswell as the continued use of native domesti-cates. Many populations of this period in thearea and all populations in the presentsample are included in the rather poorlydefined Adena mortuary complex which waswidely distributed in the middle and upperOhio Valley region (Webb and Baby, 1957).

Populations of the Middle Woodland pe-riod exhibited a further elaboration of mortu-ary ceremonialism. Large conical moundsoften with burials accompanied by exoticgoods are common in Middle Woodland con-texts. These populations further developednative domesticates even though they, likeEarly Woodland populations, still reliedheavily on hunting and gathering. The sizeof local populations appears to have gradu-ally increased in both the Early and MiddleWoodland periods. All of the Middle Wood-land samples included in the present studyare from Ohio Hopewell complex popula-tions (Griffin, 1983).

Late Woodland populations are perhapsthe most poorly known from any period. Thismay be because Late Woodland populationsabandoned the elaborate mortuary ceremo-nialism characteristic of the Early andMiddle Woodland periods (and even to someextent the Late Archaic period). Late Wood-land populations, however, for the first time,begin to inhabit, at least periodically, rela-tively large, fortified villages (Dancey, 1988).Subsistence practices in the Late Woodlandare very similar to those of the MiddleWoodland.

Frequencies of deciduous dental patholo-gies in each of the Woodland periods are lowand comparable to those in the Late Archaicperiod (Sciulli, 1997). These low frequenciesof dental pathologies reflect the continuedreliance on hunting and gathering and, de-spite the use of native cultigens, the lack of asource of high amounts of simple carbohy-drates in the diet. The frequencies of decidu-ous dental pathologies in the combined Wood-land sample are as follows: caries, 2.63%,NT 5 228; abscess, 0%, NL 5 280; antemor-tem loss, 0%, NL 5 280.

The Late Prehistoric period is defined onthe basis of the introduction to the region ofthe tropical domesticates maize and beans.Although populations now focused subsis-tence activities on these new plants, nativeplant use continued, and hunting and gath-ering still provided a significant contribu-tion to the diet. Populations, which appearto have been in general larger than in previ-ous periods, usually inhabited sizable, palli-saded villages near their fields for at leastpart of the year. In this study, I sampledpopulations of three Late Prehistoric cul-tural groups: Fort Ancient in southwestOhio and West Virginia (Anderson, Sun-Watch, and Buffalo), Monogahela in westernPennsylvania, and Sandusky in northwest-ern Ohio (Pearson) (Griffin, 1983; Stothersand Abel, 1989).

In the Late Prehistoric period, popula-tions suffered an extreme deterioration ofdental health (Sciulli, 1997). Presumablythis was the result of the inclusion of largeamounts of maize in the diet. In the decidu-ous dentition, the frequencies of pathologies,especially caries, increase dramatically com-pared to the previous periods: caries, 15.91%,NT 5 2237; abscess, 0.27%, NL 5 2191;antemortem loss, 0.41%, NL 5 2191.

The hypothesis that the samples includedhere represent an evolving population lin-eage is based on the analysis of adult cranialmetrics (Sciulli, 1990b; Sciulli and Mah-aney, 1990). It has been shown that forpopulations sampled from temporally adja-cent time periods (e.g., Archaic–MiddleWoodland, Middle Woodland–Late Wood-land, etc.) cranial metrics exhibit homoge-neous covariance structures. While there issome size difference among populations, the

191NATIVE AMERICAN DECIDUOUS TEETH

essentially identical shape of the craniaduring the period under consideration indi-cates strongly that these populations werepart of an evolving population or lineage.The pattern of evolution of these popula-tions, inferred from cranial metrics, appearsto be unilineal from the Late Archaic to theLate Woodland period. In the Late Prehis-toric period, three regional populations ap-pear (northwest, southwest, and southeastportions of Ohio), each, however, homoge-neous with the Late Woodland populationbut differing from each other. The structureof the Late Prehistoric population suggestsinstability which may have arisen as theresult of the introduction of maize accompa-nied by differential gene flow into the area(Tatarek and Sciulli, 1997).

Morphological features

From the samples described above I col-lected information on the nature and amountof variation of 57 morphological features ofthe deciduous dentition: 33 are features ofthe crown and 24 are root features. I scoredboth the left and right tooth of each indi-vidual for each feature if both teeth werepresent. However, if the expression of thefeature were symmetrical, only one scorerepresented the feature for the tooth in theindividual. If an individual exhibited asym-metry of expression in a feature, the morecomplex expression was used to representthe feature for a tooth. If only one of anantimeric pair were present, the score forthe feature of that tooth represented theindividual. Thus, all frequencies and aver-ages are based on counts per individual. Idid not collect data from individuals forwhom wear or pathologies affected the ex-pression of a feature. In all subsequentdescriptions of features and their variations,I use the following abbreviations: u, maxil-lary; l, mandibular; i, incisor; c, canine; 1,central; 2, lateral; m1, deciduous first molar;m2, deciduous second molar. The designa-tion of the postcanine deciduous teeth asmolars reflects only historical usage. Ontoge-netically these teeth are premolars. Below isthe list of the morphological features, theteeth for which each was scored, and thecriteria used in scoring variations of thefeatures. These explicit definitions should

facilitate interobserver comparisons, as thereis little agreement concerning the classifica-tion and interpretation of all morphologicalfeatures of the deciduous teeth.

Shovel shape (ui1-uc and li1-lc). All ante-rior teeth are scored for the relative develop-ment of lingual marginal ridges using Hani-hara’s (1963) criteria.

0. Absent: lingual surface smooth.1. Semishovel: slight elevation of marginal

ridges.2. Shovel: marginal ridges easily seen.3. Strong shovel: marginal ridges broad and

high.

Expressions 2 and 3 are defined to be thepresence of shoveling and 0 and 1 to beabsence of shoveling in calculating the fre-quency of this feature. This can be stated asp 5 2–3/0–3, where 2–3 is the number ofindividuals in these two classifications, 0–3is the total number of individuals scored,and p is the frequency of shoveling. Thefrequencies of all subsequent features willbe in this form, and the definition of p will besimply stated. A weighted average expres-sion, W, is also presented using all classes ofexpression: W 5 ocixi/oxi, where ci is theclass value and xi is the number of individu-als in the class.

Double shovel (ui1-uc). The maxillaryanterior teeth are scored for the presence oflabial marginal ridges.

0. Absent: labial surface smooth.1. Mesial ridge present.2. Distal ridge present.3. Mesial and distal ridge present.

The development of the labial ridges is notscored, as they are generally not stronglyexpressed. For double shovel, p 5 1–3/0–3.In calculating W, classes 1 and 2 are com-bined. Thus, for W, 0 5 absent, 1 5 oneridge, 2 5 both ridges.

Winging (ui1). Variations in the align-ment of the central incisors are scored usingDahlberg’s (1963) criteria.

0. Absent: ui1’s distal borders on straightline.

192 P.W. SCIULLI

1. Unilateral wing: distal border of one ui1toward labial.

2. Bilateral wing: both distal borders to-ward labial.

The frequency of winging is p 5 1–2/0–3.In the present sample, neither winging norcounterwinging, where the distal border(s)is rotated toward the lingual, occurred, andW is not calculated. Lack of winging in theui1 in these samples is likely due to the lackof crowding of the teeth (Dahlberg, 1963).

Interruption groove (ui1-ui2). The max-illary incisors are scored for the presence ofgrooves crossing the lingual cemento-enameljunction using the criteria of Turner et al.(1991).

0. Absent: no grooves present.1. Mesiolingual (ML): one groove at ML.2. Distolingual (DL): one groove at DL.3. Medial: one groove centrally located.4. ML and DL: two grooves, one ML and one

DL.

The frequency of interruption grooves isp 5 1–4/0–4. In the present samples, p 5 0,and W is not calculated.

Tuberculum dentale (ui1-uc and lc).The maxillary anterior teeth and lc arescored for the presence and degree of devel-opment of elaborations in the area of thelingual cingulum and lingual fossa usingGrine’s criteria (1986). In the present scor-ing, elaborations in this area are distin-guished from the size of the cingulum itself.

0. Absent: lingual surface smooth.1. Pit(s) or groove(s) present.2. One ridge present.3. Two ridges present.4. Free tubercle(s): strong ridge with free

apex.

The frequency of tuberculum dentale isp 5 1–4/0–4, and W uses all categories.

Mesial ridge (uc). The uc is scored for thepresence or absence of a distal deflection ofthe mesial marginal ridge (Irish and Morris,1996). P is the frequency of occurrence of themesial marginal ridge, and W is not calcu-lated.

Distal accessory ridge (uc, lc). The de-gree of expression of an accessory ridge onthe lingual surface of the canines betweenthe cusp apex and the distal marginal ridgeis scored using the criteria of Turner et al.(1991).

0. Absent: no ridge present.1. Faint.2. Weak.3. Moderate.4. Strong.

The frequency of canine distal accessoryridge is p 5 1–4/0–4 and W uses all catego-ries. In scoring this feature, Turner et al.(1991) use five categories. Here 5 is dropped,as no canine expressed a very pronounceddistal accessory ridge.

Cusp number, hypocone (um1). Cuspnumber and development of the upper ante-rior premolar are scored using Hanihara’s(1963) criteria.

2. Eocone (eo) and protocone (pr) present(eo 5 paracone).

3M. Eo, pr, and metacone (me) present.3H. Eo, pr, and hypocone (hy) present.42. All four cusps present but hy reduced.4. All four cusps present but hy not reduced.

For um1 cusp number, classes 42 and 4are presence of hy p 5 4 1 42/2–4. Threeclasses are used in calculating W: 2, 3 com-bining 3M and 3H, and 4 combining 4and 42.

Cusp number, hypocone (um2). The de-velopment of the posterior premolar’s hypo-cone is scored using Hanihara’s (1963) crite-ria.

3. Eo, pr, me, and a small distally placed hy.42. Eo, pr, and me attached to small hy by

distal ridge.4. Eo, pr, me, and large hy.

The frequency of um2 hypocone is p 5 4 142/3–4. Only two classes are used for W, 3and 4, where the latter combines 4 and 42.

Cusp 5 (um2). Accessory cusp located be-tween the metacone and hypocone of theupper posterior premolar is scored as pre-

193NATIVE AMERICAN DECIDUOUS TEETH

sent or absent. P is the frequency of occur-rence of this cusp, and W is not calculated.

Carabelli’s trait (um2). Elaborations ofthe mesiolingual surface of the upper poste-rior premolar are scored using Grine’s (1986)criteria.

0. Absent: mesiolingual surface smooth.1. Pit, groove: present.2. Two grooves: roughly parallel grooves.3. Welt: area between grooves raised, apex

not free.4. Cusp: same as 3 but apex free.

For Carabelli’s trait, p 5 2–4/0–4, and Wuses all categories.

Cusp number (lm2). This feature is sim-ply an enumeration of cusps present on thelower anterior premolar.

2. Eoconid (eod) and metaconid (med)(eod 5 protoconid).

3. Eod, med, and hypoconid (hyd).4. Eod, med, hyd, and entoconid (end).5. Eod, med, hyd, end, and hypoconulid

(hyld)6–8. Accessory cusps on distal marginal

ridge.

For lm1 cusp number, p 5 5–8/2–8, and Wuses all categories.

Groove pattern (lm2). The relationshipamong the principal cusps of the lower poste-rior premolar is scored. The relationshipsyield the following groove patterns:

1. 1: eod and end in contact.2. X: eod and end in contact.3. Y: med and hyd in contact.

For lm2 groove pattern, p 5 3/1–3, and Wuses all categories.

Hypoconulid (lm2). The distal cusp ofthe lower posterior premolar is scored aspresent or absent. P is the frequency of thepresence of the hypoconulid. W is not calcu-lated.

Deflecting wrinkle (lm2). The course ofthe medial ridge of the metaconid of thelower posterior premolar is scored as straightor deflected toward the distal. P is the

frequency of occurrence of the deflected me-dial ridge. W is not calculated.

Protostylid (lm2). The development ofelaborations of the mesiobuccal surface ofthe lower posterior premolar is scored usingGrine’s (1986) criteria.

0. Absent: mesiobuccal surface smooth andbuccal groove present.

1. Groove: horizontal or oblique fissure pre-sent.

2. Cusp: fissure(s) delineates swelling withfree apex.

For the protostylid, p 5 1–2/0–2, and Wuses all categories. The presence or absenceof the buccal pit is not considered in thesecriteria.

Cusp 6, tuberculum sextum (lm2). Thedevelopment of an accessory cusp (C6) lo-cated between the hyld and end is scoredusing the criteria of Turner et al. (1991).

0. Absent: no accessory cusp present.1. Hyld .. C6.2. Hyld . C6.3. Hyld 5 C6.4. Hyld , C6.5. Hyld ,, C6.

For cusp 6, p 5 1–5/0–5, and W uses allcategories.

Cusp 7, tuberculum intermedium (lm2).The development of an accessory cusp (C7)located between the med and end is scoredas follows:

0. Absent: no accessory cusp present.1. Hyld .. C7.2. Hyld . C7.3. Hyld 5 C7.4. Hyld , C7.5. Hyld ,, C7.

For cusp 7, p 5 1–5/0–5, and W uses allcategories.

Distal trigonid crest (lm2). The pres-ence or absence of a continuous ridge fromthe distal border of the eod to the distalborder of the med is noted (Turner et al.,

194 P.W. SCIULLI

1991). P is the frequency of the occurrence ofthe continuous ridge. W is not calculated.

Delta form (lm1). A triangular occlusaloutline of the lower anterior premolar isscored as present or absent (Dahlberg, 1949).When present, the triangular or delta formanterior premolar is broad distally, taperingto a mesial apex. In the present sample, thisfeature is accompanied by a ridge extendingfrom the distal marginal ridge into the talonidbasin (Fig. 1C). P is the frequency of occurrenceof the delta form. W is not calculated.

Double teeth (li1-lc). The presence orabsence of fusion-gemination of the loweranterior teeth is noted. Because it is notalways apparent whether fusion or gemina-tion occurred, these cases will be referred toas double teeth (Stevenson, 1985). To becounted, an individual must have at least a li2since virtually all cases included li1-li2 or li2-lc.

Root number (all teeth). All observa-tions of the roots of the deciduous teeth weremade on teeth extracted from the jaws. Thenumber of roots is counted for each tooth inindividuals whose roots were completed (api-ces may have been open). For anterior toothroots to be counted as double, separationbetween the roots must have extended for atleast one-quarter of the distance from theapical end. Anterior teeth are scored as 1 or2 rooted, and p 5 2/1–2. Maxillary premo-lars (um1 and um2) are scored as 2, 3, or 4rooted and mandibular premolars (lm1 andlm2) as 2 or 3 rooted. The 3 rooted mandibu-lar premolars exhibit an accessory distolon-gual root (Turner, 1971). For maxillary pre-molar root number, p 5 2/2–4, and formandibular premolar root number, p 53/2–3. W is not calculated for root numbers.

Root groove (anterior teeth). The ante-rior teeth exhibit a variation wherein in-

Fig. 1. Morphological variants of the deciduous dentition. A: Root grooves,left to right: right li1, left li1, left ui1, and left ui2; ui2 shows minimalexpression of root groove scored as absent. B: Root sheath left um2.Monongahela individual FC 3765. C: Delta form right lpa. Monongahelaindividual FC 2215. SunWatch individual 3/72.

195NATIVE AMERICAN DECIDUOUS TEETH

stead of a simple conical root the root isexpanded in the mesiodistal direction (flat-tened) and exhibits a marked vertical buccalgroove (Fig. 1A). The roots of these teethoften have two apices (Sciulli, 1990a). For allanterior teeth, p is the frequency of occur-rence of the buccal groove. W is not calcu-lated.

Root sheath (um1, um2). A thin sheet ofcementum may connect the distobuccal andlingual roots of the upper premolars (Jor-gensen, 1956) (Fig. 1B). This feature can benoted prior to root completion as the connec-tion begins at the base of the roots (andusually extends almost to the apex). Forboth upper premolars, p is the frequency ofoccurrence of the root sheath. W is notcalculated.

Labial deflection (anterior teeth). Thepresence or absence of a labial orientation ofthe roots of the anterior teeth (Jorgensen,1956) was scored if the roots were complete(see above). To be considered present, theapical half of the root at least must exhibitany degree of labial tilt. P is the frequency ofoccurrence of labial deflection, and W is notcalculated.

Heterogeneity of frequencies among thesamples is tested by the G-test (Sokal andRohlf, 1981) using the dichotomized data(the presence of a feature is the same as thedefinition of the numerator given for p forthe feature). All features are tested for asso-ciation using X2 or Fisher’s Exact Test (So-kal and Rohlf, 1981). For the majority offeatures, the entire sample is used for test-ing associations (see below). Finally, themean measure of divergence (mmd) betweensamples is calculated using independent fea-tures (de Souza and Houghton, 1977). Thematrix of mmds is graphically displayedusing principal coordinates analysis (Maridaet al., 1979).

RESULTS

Tables 2 and 3 contain the frequencies andweighted average expressions for 41 of the57 morphological features of the deciduousdentition in the Ohio Valley samples. Ninefeatures are not included in these tablesbecause they exhibit no variation in the total

sample, while another five features are notincluded because they exhibit extremely littlevariation. These 14 features with little or novariation are listed in Table 4. Root numbersof um1 and um2 are not listed in Table 2because the presence of a root sheath isequivalent to the expression of two roots andits absence is equivalent to the expression ofthree roots.

Shoveling

Shoveling of the anterior teeth is bothfrequent and well expressed in the OhioValley samples. Among the six samples, onlyli1 (G5 5 12.18) and li2 (G5 5 16.64) shovel-ing exhibit heterogeneity. In both cases thedifferences are due to the relatively highfrequencies of shoveling in the Woodlandsample and the low frequency of shoveling inthe Pearson sample. Because the remaininganterior teeth show no differences in thefrequency of shoveling and because the sizesof the Woodland and Pearson samples arerelatively small for li1 and li2, there is astrong possibility that the observed heteroge-neity for shoveling in these two teeth is theresult of sampling effects and not indicativeof true differences.

Shoveling in the total sample is mostfrequent and strongly expressed in ui2, uc,and lc. The mandibular incisors, especiallyli1, exhibit the lowest frequency of shovelingand the weakest expression, while ui1 isintermediate in both measures.

For the total sample and in the Woodlandand Pearson samples, shoveling showsstrong associations between anterior teeth.The maxillary incisors are significantly asso-ciated with each other but independent ofthe canines, while the mandibular incisorsare associated with each other, the maxil-lary incisors, and the mandibular canine.Shoveling of the maxillary canine is the onlyfeature independent of shoveling in all otheranterior teeth. Associations between shovel-ing and other features in the total sampleare as follows: ui1-ui1 double shovel (nega-tive association), li1-ui2 tuberculum dentaleand um1 cusp number (negative associa-tion), li2-um2 Carabelli trait, lc-lm2 rootnumber (negative association), lc root groove,and um2 root sheath.

196 P.W. SCIULLI

Double shovel

Labial shoveling of the maxillary anteriorteeth is much less frequent than lingualshoveling and even when present is notstrongly expressed. The most frequent mani-festation of this feature is a single mesialridge. The presence of a mesial and distalridge is the least common expression ofdouble shoveling (,2%).

Among the six samples, ui1 (G5 5 13.72)and ui2 (G5 5 15.04) show significantly het-erogeneity. In the former case, heterogeneityis due to the low frequency of double shovelin the SunWatch sample, while in the lattercase it is due to the high frequencies in theWoodland and Monongahela samples.

In the total sample and in the combinedWoodland and Monongahela samples, double

shovel of each anterior tooth is associatedwith each other anterior tooth. Associationsbetween double shovel and other features inthe total sample are as follows: ui1-uc distalaccessory ridge, um1 cusp number, um2Carabelli trait and ui2-uc distal accessoryridge, um2 Carabelli trait.

Tuberculum dentale

There are no differences among thesamples for the frequency of this feature. Inthe total sample, the maxillary canine ex-presses tuberculum dentale most frequentlyand strongly. Tuberculum dentale is rareand weakly expressed on the mandibularcanine, while the upper incisors show asomewhat higher frequency and strongerexpression. Only the upper canine (14.6%)

TABLE 2. Frequencies of morphological features of the maxillary deciduous dentition in prehistoricOhio Valley samples1

Tooth Trait

Archaic Woodland Pearson SunWatch Monongahela Buffalo Total

N % W N % W N % W N % W N % W N % W N % CI W

ui1 Shovel 34 73.5 1.97 20 75.0 2.10 22 72.7 1.95 37 78.4 2.19 25 80.0 1.88 17 82.4 1.94 163 77.3 70.983.7

2.01

Double shovel 32 15.6 0.16 20 35.0 0.40 20 15.0 0.15 34 5.9 0.06 27 33.3 0.41 16 37.5 0.44 157 20.4 14.126.7

0.23

Tuberculumdentate

33 15.1 0.42 20 5.0 0.10 19 5.3 0.05 33 12.1 0.24 26 15.4 0.27 17 17.6 0.35 155 11.6 6.616.6

0.25

Root groove 24 58.3 — 13 84.6 — 16 18.8 — 17 52.9 — 13 69.2 — 14 35.7 — 105 54.3 44.863.8

—

Labial deflec-tion

24 54.1 — 13 76.9 — 16 25.0 — 17 64.7 — 12 25.0 — 13 38.5 — 103 47.6 38.057.2

—

ui2 Shovel 33 87.9 2.06 18 100 2.17 19 84.2 2.00 37 91.9 2.38 18 94.4 2.22 14 100 2.21 147 92.5 88.296.8

2.18

Double shovel 32 3.1 0.03 18 33.3 0.33 19 5.3 0.05 34 2.9 0.03 18 22.2 0.39 14 14.3 0.14 143 10.5 5.515.5

0.13

Tuberculumdentate

32 15.6 0.44 18 5.6 0.22 17 17.6 0.35 34 14.7 0.32 18 22.2 0.28 13 23.1 0.46 139 15.1 9.121.1

0.33

Labial deflec-tion

23 4.3 — 12 25.0 — 16 6.2 — 16 6.2 — 9 0 — 12 16.7 — 96 9.4 3.615.2

—

uc Shovel 34 88.2 2.02 23 82.6 2.04 18 83.3 2.11 33 90.9 2.27 27 88.9 2.11 20 90.0 2.15 162 87.7 82.692.8

2.12

Double shovel 34 14.7 0.24 23 13.0 0.13 18 11.1 0.11 31 3.2 0.03 27 3.7 0.04 18 16.7 0.28 158 9.5 4.914.1

0.13

Tuberculumdentate

34 35.3 1.00 23 26.1 0.74 23 52.2 1.09 33 54.5 1.00 28 57.1 1.07 23 52.2 0.96 171 45.6 38.153.1

1.02

Distal acces-sory ridge

33 27.3 0.73 15 40.0 0.93 20 35.0 0.95 34 20.6 0.53 21 52.4 1.67 21 9.5 0.24 150 30.0 22.737.3

0.82

Root groove 25 60.0 — 17 64.7 — 21 14.3 — 15 40.0 — 23 47.8 — 16 31.3 — 123 44.7 35.953.5

—

Labial deflec-tion

24 12.5 — 17 47.1 — 21 42.9 — 14 64.3 — 23 30.4 — 16 25.0 — 120 33.3 24.941.7

—

um1 Cusp number 44 25.0 3.00 27 40.7 3.30 25 40.0 3.28 34 64.7 3.65 30 73.3 3.73 19 68.4 3.68 188 49.5 42.456.6

3.22

Root sheath 38 57.9 — 17 52.9 — 28 64.3 — 19 84.2 — 29 55.2 — 26 53.8 — 165 59.4 51.966.9

—

um2 Cusp 5 42 38.1 — 26 23.1 — 29 24.1 — 32 25.0 — 36 22.2 — 32 25.0 — 206 26.7 20.732.7

—

Carabelli 44 13.6 0.82 27 11.1 0.93 30 30.0 1.17 30 26.7 1.17 40 25.0 1.15 36 33.3 1.17 217 22.6 17.028.2

1.05

Root sheath 28 32.1 — 16 25.0 — 26 30.8 — 13 30.8 — 27 22.2 — 19 0 — 133 25.6 18.233.0

—

1 CI, confidence interval; N, sample size; W, weighted expression; %, frequency.

197NATIVE AMERICAN DECIDUOUS TEETH

exhibits a free tubercle at a frequency greaterthan 5%.

The upper canine and ui1 show a signifi-cant association for the presence of thetuberculum dentale. Associations betweentuberculum dentale and other features areas follows: ui2-uc root groove, uc-uc distalaccessory ridge, um1 cusp number, lm1 cuspnumber (negative association), lm2 cusp 7,lm1 root number, lc-uc distal accessory ridge,lc distal accessory ridge, um1 cusp number,lm2 cusp 7, ui1 root groove.

Distal accessory ridge

The frequency of the canine distal acces-sory ridge (uc, lc) is homogeneous among the

TABLE 3. Frequencies of morphological traits of the mandibular deciduous dentition in prehistoricOhio Valley samples1

Tooth Trait

Archaic Woodland Pearson SunWatch Monongahela Buffalo Total

N % W N % W N % W N % W N % W N % W N % CI W

li1 Shovel 29 48.3 1.28 16 62.5 1.44 12 8.3 1.00 31 45.2 1.42 14 28.6 1.21 11 27.3 1.18 118 38.9 30.147.7

1.25

Root groove 21 4.8 — 16 6.2 — 9 11.1 — 18 5.6 — 12 16.7 — 10 0 — 92 6.5 1.511.5

—

Labialdeflection

22 18.2 — 15 6.7 — 9 0 — 18 0 — 12 0 — 10 0 — 91 5.5 0.8010.2

—

il2 Shovel 33 66.7 1.55 20 95.0 2.20 18 38.9 1.39 35 68.6 1.74 20 65.0 1.75 15 53.3 1.53 147 65.3 57.673.0

1.67

Root groove 23 13.0 — 15 6.7 — 16 18.8 — 17 23.5 — 16 25.0 — 8 37.5 — 100 18.0 10.525.5

—

Labialdeflection

23 13.0 — 15 0 — 16 12.5 — 17 5.9 — 16 0 — 8 0 — 101 5.9 1.310.5

—

lc Shovel 41 87.8 2.12 20 100 2.15 15 93.3 2.47 33 97.0 2.55 20 100 2.60 14 64.3 1.79 150 90.7 86.195.3

2.25

Tuberculumdentate

39 5.1 0.13 20 10.0 0.15 16 0 0 31 3.2 0.06 16 6.2 0.06 15 6.7 0.13 144 5.6 1.89.4

0.10

Distal acces-sory ridge

38 5.3 0.16 18 22.2 0.44 16 12.5 0.44 31 3.2 0.13 17 5.9 0.18 17 23.5 0.47 144 11.1 6.016.2

0.28

Root groove 24 20.8 — 15 26.7 — 19 42.1 — 13 38.5 — 20 65.0 — 11 36.4 — 106 38.7 29.448.0

—

Labialdeflection

23 26.1 — 15 20.0 — 19 36.8 — 13 53.8 — 21 33.3 — 11 9.1 — 107 29.0 20.437.6

—

lm1 Cuspnumber

40 77.5 5.02 24 70.8 4.96 26 61.5 4.69 40 80.0 5.25 40 70.0 4.80 35 71.4 4.80 214 72.0 66.078.0

4.94

Delta form 41 24.3 — 25 24.0 — 26 15.4 — 40 22.5 — 43 30.2 — 35 11.4 — 219 21.9 16.427.4

—

Disto-lingual root

31 9.7 — 15 6.7 — 28 14.3 — 18 0 — 34 8.8 — 28 0 — 159 6.9 3.010.8

—

lm2 Deflectingwrinkle

44 61.4 — 23 69.6 — 35 91.4 — 35 88.6 — 38 92.1 — 39 89.7 — 223 82.1 77.187.1

—

Protostylid 47 34.0 0.45 26 38.5 0.50 36 30.6 0.42 35 25.7 0.40 38 15.8 0.18 42 33.3 0.48 233 28.3 22.534.1

0.39

C6 46 45.6 1.41 26 46.2 1.27 35 34.3 0.60 36 41.7 0.72 38 44.7 0.89 40 47.5 1.12 230 43.9 37.550.3

1.03

C7 45 24.4 0.42 26 11.5 0.31 35 48.6 1.43 36 72.2 2.06 38 47.4 1.13 41 70.7 1.73 230 47.4 40.953.9

1.20

Groovepattern

37 91.9 2.84 24 100 3.00 34 94.1 2.88 30 93.3 2.90 36 91.7 2.86 35 88.6 2.83 203 93.1 89.696.6

2.88

Disto-lingual root

30 13.3 — 15 6.7 — 32 9.4 — 13 0 — 32 0 — 32 9.4 — 159 6.9 3.010.8

—

li1-li2-lcdoubleteeth

35 2.9 — 20 0 — 22 4.5 — 69 7.2 — 28 0 — 49 2.0 — 231 4.3 1.76.9

—

1 CI, confidence interval; N, sample size; W, weighted expression; %, frequency.

TABLE 4. Features of the deciduous dentitionexhibiting little or no variation in the

Ohio valley samples1

Morphological feature Tooth N %

Winging (1) ui1 137 0Interruption groove (1) ui1 161 0

ui2 149 0Mesial ridge (1) uc 154 0Root groove (1) ui2 98 0Root number (.1) ui1 105 0

ui2 98 0li1 91 0li2 100 0

Cusp number (,4) um2 222 0.5Hypoconulid (2) lm2 233 0.4Distal trigonid crest (1) lm2 188 1.1Root number (2) uc 122 1.6

lc 109 0.91 (1), present; (2), absent.

198 P.W. SCIULLI

six samples. In general, uc exhibits both ahigher frequency and stronger expression ofthis accessory ridge than the lower canine.However, the presence of this feature isstrongly associated in the upper and lowercanines.

Associations between the distal accessoryridge and other features are as follows:uc-um1 cusp number, lm1 cusp number,um2 Carabelli trait, lc-lm1 cusp number,lm2 deflecting wrinkle (negative associa-tion).

Cusp number um1

Cusp number of um1 exhibits significantheterogeneity among the six samples(G5 5 25.86). In this case, the pattern ofvariation suggests the differences may notbe due to sampling. The frequency of fourcusps is relatively low in the Archaic, Wood-land, and early (approximately 950–850 BP)Late Prehistoric Pearson samples and ap-proximately doubles in frequency in samplesfrom later agricultural populations. In boththe earlier and later groups, the presence offour cusps on um1 is associated with lm1cusp number, um2 Carabelli trait, and lm2protostylid.

Cusp 5 um2

The frequency of this feature is homoge-neous among the samples and achieves amoderate frequency in each. The occurrenceof cusp 5 is independent of all other features.

Carabelli trait

Carabelli’s trait tends to increase in fre-quency and show a stronger expression overtime in the Ohio Valley samples. Much of thevariation between samples is, however, lim-ited to the expression of the double fissurewhich becomes more frequent in the LatePrehistoric populations. While there is atendency for an increase in frequency of thisfeature over time, there are no significantdifferences among the samples. Carabellitrait is associated with lm1 cusp number.

In the present scoring scheme, Carabellitrait presence is defined as double fissure,welt, or cusp, and this classification has alow to moderate frequency in the samples.However, in all samples the pit-groove ex-pression is much more frequent (50.7% in

the total sample) and shows no tendency tochange over time.

Cusp number lm1

Five is the modal cusp number for lm1 inthe Ohio Valley samples. Reduction of cuspnumber (28%) is slightly more frequent thanaugmentation (20%). The frequency of lm1cusp number is homogeneous in the samples.

Lm1 cusp number is associated with thefollowing features: lm2 protostylid, lm1 deltaform (negative association), lm1 root num-ber (negative association), ui1 root groove,and root sheath of both um1 and um2.

Groove pattern lm2

The Y pattern (Y-5) is the dominant expres-sion of this feature. The plus (1) pattern(5%) is somewhat more frequent than the Xpattern (2%), but both are sporadic in occur-rence. This feature is homogeneous amongthe samples. Because of the virtual fixationof the Y pattern, I did not test this featurefor associations.

Deflecting wrinkle

The presence of a deflecting wrinkle in-creases significantly among the Late Prehis-toric agricultural groups (G5 5 21.06). It isonly in this latter grouping that the deflect-ing wrinkle shows an association with lm2cusp 7.

Protostylid

This feature is homogeneous among thesix samples. In each sample the frequency ofthe horizontal-oblique fissure is greater thanthat of the cusp. In the total sample, thefrequency of the fissure is 19.7%, while thefrequency of the cusp is 8.6%.

The presence of the protostylid is nega-tively associated with the uc and li2 rootgroove.

Tuberculum sextum

While the frequency of C6 shows no signifi-cant differences among the samples, thedegree of expression of this feature is gener-ally greater in the Archaic and Woodlandsamples than among the Late Prehistoricsamples. C6 is independent of all otherfeatures.

199NATIVE AMERICAN DECIDUOUS TEETH

Tuberculum intermedium

The frequency of C7 is heterogeneousamong the samples (G5 5 43.78). The earlierArchaic and Woodland samples have fre-quencies of C7 less than 25%, while the LatePrehistoric samples show at least twice thisfrequency. In both the earlier and lattergroups, however, the presence of C7 is nega-tively associated with ui1 root groove.

Delta form

The frequency of the delta form lm1 ishomogeneous in the six samples. This fea-ture achieves a moderate frequency in eachsample and is associated with lm1 rootnumber.

Double teeth

The occurrence of double teeth is sporadicin most of the Ohio Valley samples. Theexceptions appear to be the SunWatch (5/69)and Anderson Village (2/6) samples, bothrepresenting the Fort Ancient Tradition andboth located in southwest Ohio. When com-pared to all other samples (1.99%), the com-bined SunWatch–Anderson Village sample(9.33%) exhibits a significantly higher fre-quency of double teeth. Because of the gener-ally low frequency of this feature, tests forassociations were not performed.

Root number

As stated above, root number for the de-ciduous incisors shows no variation (all singlerooted), and the canines only rarely exhibit adouble root. Root number for the upperpremolars (um1, um2) will be consideredbelow under root sheath.

The mandibular premolars (lm1, lm2) eachexhibit an accessory distolingual root at afrequency of 6.9% in the total sample. Be-cause of the reduced sample sizes for thisfeature and the low frequency, tests forheterogeneity or associations were not per-formed.

Root sheath

The maxillary premolars (um1, um2) eachexhibit the root sheath, with um1 sheath(59.4%) being about twice as frequent asum2 sheath (25.1%). There is no heterogene-ity among the samples for this feature. As

the presence of the sheath is equivalent totwo roots and its absence is equivalent tothree roots, root number of the upper premo-lars is also homogeneous. Four roots (and nosheath) occurs sporadically in um1 (0.6%)and um2 (4.5%).

Um1 and um2 are strongly associated forthe presence of the root sheath. Um1 rootsheath also shows a negative associationwith lc root groove.

Root groove

The presence of the buccal root groove ismost common on ui1 and the canines. Thisfeature is generally rare on the lower inci-sors and is absent on ui2. Ui1 and uc exhibitheterogeneity for this feature (G5 5 17.20and G5 5 15.41, respectively). In both casesthe frequency of the root groove is somewhathigher in theArchaic and Woodland samples,but the source of heterogeneity is the rela-tively small Pearson sample.

In the total sample and in the combinedArchaic-Woodland sample, ui1 and uc rootgroove are strongly associated.

Labial deflection

As with the buccal groove, the presence oflabial deflection is most common for ui1 andthe canines. Ui1 labial deflection exhibitsheterogeneity among the samples, but againthe source of the heterogeneity is the smallPearson (and Monongahela) sample. Be-cause sample sizes are reduced for thisfeature, tests for associations were not per-formed.

Tables of the counts of each expression forall traits in each sample are available onrequest.

In order to evaluate patterns of variationin the frequency of deciduous morphologicaltraits in the Ohio Valley samples, I calcu-lated the mean measure of divergence be-tween each sample and performed principalcoordinates analysis on the resulting dis-tance matrix. For this analysis, 13 indepen-dent features were used: ui1 and uc shovel,ui2 double shovel, ui1 tuberculum dentale,lc distal accessory ridge, um1 cusp numberand root sheath, um2 C5, lm2 C6, C7, androot number (distolingual root), lm1 deltaform, and uc root groove. This set of featuresincludes two which show systematic differ-

200 P.W. SCIULLI

ences among the samples (um1 cusp numberand lm2 C7) and two features which showdifferences that may be due to sampling (ui2double shovel and uc root groove). In this setof 13 features, approximately 30% of thefeatures shows some differences among thesamples. This proportion of differencesshould reflect the variation in the total set offeatures in which about 20% exhibit somedifferences among the samples.

Figure 2 is the plot of the first two axesfrom the principal coordinates analysis. Inthis figure, a basic dichotomy appears tooccurs along axis 1: a contrast between LatePrehistoric samples on the left and earlierArchaic and Woodland samples toward theright. Axis 1 also shows that among the LatePrehistoric samples, SunWatch and Buffalo,both representing Fort Ancient populations,cluster very closely. Axis 2 separates Archaicfrom Woodland populations and non–FortAncient Late Prehistoric populations fromthe Fort Ancient cluster. Although not pre-sented, a correspondence analysis (Greena-

cre, 1984) of the 13 frequencies produced asimilar pattern of differences among thesamples, and the first two axes accounted fora similar amount of the total variation(73.1%). As might be expected from Tables 1and 2, the first axis of the correspondenceanalysis revealed that the Late Prehistoricpopulations are most closely associated withlm2 C7, while the Archaic and Woodlandpopulations are associated with uc rootgroove. Axis 2 shows an association of theWoodland sample with ui2 double shoveland the Archaic sample with lm2 root num-ber and um1 root sheath.

Figure 3 contains the result of a correspon-dence analysis based on six features and 12samples: the six Ohio Valley samples and sixcomparative samples. This analysis is lim-ited to six features, as these features havebeen scored in a common manner (Grine,1986). The data for the comparative samplesare from American white, American Black,and Japanese (Hanihara, 1963), MedievalDanes (Jorgensen, 1956), South African

Fig. 2. Principle coordinates analysis of mean measures of divergence between Ohio Valley popula-tions.

201NATIVE AMERICAN DECIDUOUS TEETH

blacks (Grine, 1986), and Prehistoric Inga-moan (Luckas and Walimbe, 1984). For theMedieval Danes, ui1 shovel is not recorded,and a frequency of 0% is assumed for thissample.

Axis 1 in Figure 3 separates East Asianand derived populations (Ohio Valley) to-ward the right of the plot, associated withui1 shovel, lm2 C6, and um1 cusp number(hypocone), while the European and African(and African-derived) populations are lo-cated to the left of the plot and associatedwith Carabelli’s trait. Lm2 C7 and um2 cuspnumber (hypocone) are similar in frequencyamong the samples and are thus locatedtoward the center of the plot not associatedwith any divergence among populations.Axis2 appears to separate recent populations,toward the top of the plot, from populationsearlier in time.

DISCUSSION AND CONCLUSIONS

The frequencies and degree of expressionof morphological features of the deciduousteeth in the Ohio Valley lineage were gener-

ally conservative. Within the 2,500 yearspan considered, 46 of the 57 features inves-tigated show no statistically significant dif-ferences among the samples. Of the 11 fea-tures that do show significant differences,eight exhibit deviations expressed by one ortwo populations with small samples for thefeature, and these deviations are thus likelyto be the result of sampling effects.

This observed stasis or in fact minor fluc-tuations around average frequencies for mostfeatures could have been the result of stabi-lizing selection acting within small popula-tions or genetic drift acting within largerpopulations. Since archaeological evidenceindicates that the lineage during this periodwas composed of relatively large, interact-ing populations and since the deciduousteeth have a limited functional exposure tothe environment and thus a limited liabilityto the direct effects of selection, drift is themore likely cause of the minor fluctuationsin the frequencies of most of the features ofthe deciduous teeth in the Ohio Valley lin-eage.

Fig. 3. Correspondence analysis of frequencies of morphological features of the deciduous dentition inOhio Valley and comparative populations.

202 P.W. SCIULLI

Although the vast majority of morphologi-cal variations exhibits only minor fluctua-tions in frequency over time in this lineage,three features (um1 cusp number and lm2deflecting wrinkle and C7) show significant,consistent differences among the sampleseach increasing in frequency in the LatePrehistoric period. The increase in fre-quency of these three features is associatedwith the transition (approximately 1000 BP)during which populations changed from asubsistence base in which native cultigensmade a major contribution to a subsistencebase dominated by the introduced tropicaldomesticates maize and beans. The magni-tude of the differences in these three fea-tures is sufficient to allow discriminationbetween pre- and post-1000 BP populations.

As will be shown in a future report, thesize of the deciduous teeth in this lineage didnot change significantly from the Late Ar-chaic to the Late Prehistoric period. Thereare thus two likely explanations for thechanges in the frequencies of morphologicalvariations which occurred at this transi-tional period: changes in selective forces andgene flow. Postulating changes in selectionfor this period is not unreasonable, as popu-lations were relatively large and major envi-ronmental changes (diet, settlement, ecol-ogy) were occurring (Sciulli, 1997; Tatarekand Sciulli, 1997; Fritz, 1990). If selectionwere the cause of the change in frequencies,the increase in frequency of four cusped um1and lm2 deflecting wrinkle and C7 in theLate Prehistoric populations may have beenthe result of selection against these expres-sions. All individuals in the samples diedprior to reproduction. The samples thuscontain individuals representing backgroundmortality and potentially individuals uponwhom selection was acting based on charac-teristics other than the dentition as well asindividuals upon whom selection was actingbased on dental morphology either directlyor indirectly. The morphological variationsin the sample may thus represent to somedegree the features conferring a lower fit-ness. Other scenarios based on selectionmay also be postulated, but at present anyhypotheses based on these ideas cannot betested, as the morphology of the deciduousteeth of individuals who survived childhood

cannot be determined. However, since thesechanges occurred very quickly (compared tothe total time span) and are limited to threefeatures, differences in selection may not bethe most likely explanation.

The second mechanism that may explainthe morphological changes in the deciduousteeth at the transition to the Late Prehis-toric period is gene flow. The introduction ofmaize and beans to the middle and upperOhio Valley (approximately 1000 BP) mayhave been accompanied by the introductionof genes either by diffusion or by migrationand diffusion from a population(s) to thewest or south of the region. Evidence indicat-ing an introduction rather than a gradualdevelopment includes the fact that the Earlyagricultural populations of the region, repre-senting the Fort Ancient Tradition, show anabrupt change in agricultural and settle-ment patterns at this time. From the outset,Fort Ancient assemblages contain little evi-dence of the native seed crops common inMiddle and Late Woodland assemblages ofthe area but rather an abundance of maizeand beans (Fritz, 1990). This change insubsistence is accompanied by the presenceof large villages and a marked degree ofnucleation, both features distinguishingthese populations from earlier groups aswell as from many contemporaneous groups.In addition, the uniformity among Fort An-cient maize assemblages (Eastern Eight-row maize) indicates a single source of seedrather than a series of genetic sources (Wag-ner, 1987). The clustering of the Fort An-cient samples in Figure 2 (SunWatch andBuffalo) and the similarity in the archaeo-logical assemblages of Fort Ancient sitessuggest that both maize and genes may havebeen derived from a single source.

The incidence of double teeth also appearsto support this scenario. Double teeth aregenerally rare in human populations. Sur-veys of European and North American chil-dren have determined that in these popula-tions the frequency of double teeth neverexceeded 1% and averaged about 0.5% (Brookand Winter, 1970). An exception to thisgeneral low frequency was found howeveramong Japanese children, who expresseddouble teeth at a frequency of 4.95% (Saito,1959).

203NATIVE AMERICAN DECIDUOUS TEETH

The frequency of double teeth in OhioValley populations is low (approximately2%) until the Late Prehistoric period, whenthis feature appears in southwestern Ohioat frequencies approaching 10%. But, awayfrom southwest Ohio in the Late Prehistoricperiod, toward the north and east, the fre-quency of double teeth remains low (approxi-mately 2%). Stevenson (1985) has docu-mented a relatively high frequency of doubleteeth (6.38%) in the deciduous dentition atAverbuch, a Mississippian population fromthe area around Memphis, Tennessee. Sincepopulations to the west and south of themiddle Ohio Valley are a likely source fromwhich maize and beans were introduced,they may also be the source of the increasedfrequencies of double teeth and other mor-phological variants of the deciduous denti-tion which appear at this time. This idea ofgene flow may be evaluated more easily thanselection alternatives by surveying popula-tions to the west and south of the middleOhio Valley for the frequency of doubleteeth, um1 cusp number, and lm2 deflectingwrinkle and C7.

The hypotheses stated above, while basedon the present evidence, must be consideredtentative. Additional samples from the re-gion, especially samples representing Wood-land populations, are necessary for a morecomplete evaluation of microevolutionarychanges in this lineage. Particularly criticalto this endeavor are studies of Late Wood-land populations. Many of the significantchanges in morphological variation of thedeciduous teeth of this lineage are associ-ated with changes in subsistence practicesthat occurred at about 1000 BP. However,both the archaeological and biological recordof this lineage is at best sketchy just prior tothis time. Additional Late Woodland sitesand samples are necessary in order to evalu-ate the specific causes of the biological andcultural changes made manifest at this time.

Even with the modest overall changes infrequency of some of the morphological vari-ants in Ohio Valley populations, the generalpattern of expression of morphological fea-tures of the deciduous dentition correspondsto what has been called the ‘‘MongoloidDental Complex’’ (Hanihara, 1967). Thissuite of features includes high frequencies of

ui1 and ui2 shovel and lm2 deflecting wrinkleand moderate frequencies of lm2 proto-stylid, C7, and um2 C5. In a cladistic analy-sis of morphological features of the perma-nent dentition among human populations,Stringer et al. (1997) suggest that manyfeatures of the Sinodont Pattern (Turner,1987), including I1 shovel and M1 deflectingwrinkle, are highly derived. If this is true forthe permanent dentition, then it appearsthat these features as well as others mayshow the same character state polarity inthe deciduous dentition. Delta form lm1may be another highly derived feature of theMongoloid Dental Complex of the deciduousdentition. Among European and relatedpopulations, this variant achieves a fre-quency of only 2–3% (Dahlberg, 1949; Jor-gensen, 1956), and it was not found in aSouthAfrican black population (Grine, 1986).However, delta form lm1 reaches moderatefrequencies (22–57%) in some East Asianand derived populations (Hanihara, 1956;Dahlberg, 1949), including the Ohio Valleypopulations. The presence of an accessorydistolingual root on lm1 and lm2 and doubleteeth may also be derived features, as theyappear to be rare at least in Europeanpopulations (.1%) while reaching appre-ciable frequencies in the Ohio Valley popula-tions and some East Asian populations.

Finally, variants of root morphology, whichare not generally considered in discussionsof the deciduous dentition, appear to bequite common in human populations. Rootgrooves of the anterior teeth and the rootsheath of um1 and um2 achieve moderatefrequencies in both European and Ohio Val-ley populations (Jorgensen, 1956; Black,1902). This distribution suggests these fea-tures may be primitive characteristics andthus common in most human populations.

LITERATURE CITED

Black GV (1902) Descriptive Anatomy of the HumanTeeth. Philadelphia: SS White Dental ManufacturingCo.

Brace CL, Smith SL, and Hunt KD (1991) What bigteeth you had grandma! Human tooth size past andpresent. In MA Kelley and CS Larsen (eds.): Advancesin Dental Anthropology. New York: Wiley-Liss, pp.33–58.

Brook AH, and Winter SB (1970) Double teeth: A retro-spective study of ‘‘geminated’’ and ‘‘fused’’ teeth inchildren. Br. Dent. J. 129:123–130.

204 P.W. SCIULLI

Dahlberg AA (1949) The dentition of the AmericanIndian. In WS Laughlin (ed.): Papers on the PhysicalAnthropology of the American Indian. New York:Viking Fund, pp. 138–176.

Dahlberg AA (1963) Analysis of the American Indiandentition. In DR Brothwell (ed.): Dental Anthropology.London: Pergamon Press, pp. 149–177.

Dancey WS (1988) The community plan of an early LateWoodland village in the middle Scioto river valley.Midcont. J. Arch. 13:233–258.

de Souza P, and Houghton P (1977) The mean measureof divergence and the use of non-metric data in theestimation of biological distances. J. Arch. Sci. 4:163–196.

Fritz GJ (1990) Multiple pathways to farming in precon-tact eastern NorthAmerica. J. World Prehistory 4:387–435.

Greenacre MJ (1984) Theory and Applications of Corre-spondence Analysis. New York: Academic Press.

Griffin JB (1983) The Midlands. In JD Jennings (ed.):Ancient North Americans. San Francisco: WH Free-man, pp. 243–301.

Grine FE (1986) Anthropological aspects of the decidu-ous teeth of South African blacks. In R Singer and JKLundy (eds.): Variation, Culture and Evolution inAfrican Populations. Johannesburg: WitwatersrandUniversity Press, pp. 47–83.

Hanihara K (1956) Studies on the deciduous dentition ofthe Japanese and the Japanese-American hybrids. IIIDeciduous lower molars. J. Anthropol. Soc. Nippon64:95.

Hanihara K (1963) Crown characters of the deciduousdentition of the Japanese-American hybrids. In DRBrothwell (ed.): Dental Anthropology. London: Per-gamon Press, pp. 104–124.

Hanihara K (1967) Racial characteristics in the denti-tion. J. Dent. Res. 46:923–926.

Irish JD, and Morris DH (1996) Canine mesial ridge(Bushman Canine) dental trait definition. Am. J.Phys. Anthropol. 99:357–359.

Jorgensen K (1956) The deciduous dentition: A descrip-tive and comparative anatomical study. Acta Odontol.Scand. 14:1–202.

Lukacs JR, and Walimbe SR (1984) Deciduous dentalmorphology and the biological affinities of a lateChalcolithic skeletal series from western India. Am. J.Phys. Anthropol. 65:23–30.

Mardia KV, Kent JT, and Bibby JM (1979) MultivariateAnalysis. New York: Academic Press.

Saito T (1959) A genetic study of the degenerativeanomalies of deciduous teeth. Jpn. J. Hum. Gen.4:27–54.

Sciulli PW (1977) A descriptive and comparative study ofthe deciduous dentition of prehistoric Ohio ValleyAmerindians. Am. J. Phys. Anthropol. 47:71–80.

Sciulli PW (1990a) Deciduous dentition of a Late Ar-chaic population of Ohio. Hum. Biol. 62:221–245.

Sciulli PW (1990b) Cranial metric and discrete traitvariation and biological differentiation in terminalLate Archaic populations of Ohio: The Duff site. Am. J.Phys. Anthorpol. 82:19–29.

Sciulli PW (1997) Dental evolution in prehistoric NativeAmericans of the Ohio Valley area: I Wear and pathol-ogy. Int. J. Osteoarch. 7:507–524.

Sciulli PW, and Mahaney MC (1990) Evidence of localbiological continuity for an Ohio Hopewell complexpopulation. Midcont. J. Arch. 12:117–114.

Smith BD (1989) Origins of agriculture in EasternNorth America. Science 246:1566–1571.

Smith P (1976) Evolutionary changes in the deciduousdentition of Near Eastern populations. Bull. Group.Int. Rech. Sci. Stomatol. 19:187–198.

Smith P (1978) Evolutionary changes in the deciduousdentition of Near East populations. J. Hum. Evol.7:401–408.

Sokal RR, and Rohlf FJ (1981) Biometry. San Francisco:WH Freeman.

Stevenson DR (1985) Prevalence rate of double teeth indeciduous dentition at Averbuck (40DV60). Tenn. An-thropol. 10:134–155.

Stothers DM, and Abel TJ (1989) The position of the‘‘Pearson Complex’’ in the late prehistory of Ohio. ArchOf E. North America 17:109–141.

Stringer CB, Humphrey LT, and Compton T (1997)Cladistic analysis of dental traits in recent humansusing a fossil outgroup. J. Hum. Evol. 32:389–402.

Tatarek NE, and Sciulli PW (1997) Comparisons ofpopulation structure in Ohio’s Late Archaic and LatePrehistoric period. Amer. J. Phys. Anthropol.24(Suppl.):225–226.

Turner CG II (1971) Three-rooted mandibular firstpermanent molars and the question of AmericanIndian origins. Am. J. Phys. Anthropol. 34:229–242.

Turner CG II (1987) Late Pleistocene and Holocenepopulation history of East Asia based on dental varia-tion. Am. J. Phys. Anthropol. 73:305–322.

Turner CG II, Nichol CR, and Scott GR (1991) Scoringprocedures for bey morphological traits of the perma-nent dentition: The Arizona State University dentalanthropology system. In MA Kelley and CS Larsen(eds.): Advances in Dental Anthropology. New York:Wiley-Liss, pp. 13–32.

Wagner GE (1987) Uses of Plants by the Fort AncientIndians. Ph.D. dissertation, Washington University,St. Louis. Ann Arbor: University Microfilms.

Webb WS, and Baby RS (1957) The Adena People No. 2.Columbus, OH: The Ohio Historical Society.

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