Dental Wear in Immature Late Pleistocene European Hominines

Journal of Archaeological Science (1997) 24, 677–700

Dental Wear in Immature Late Pleistocene EuropeanHominines

Mark Skinner

Department of Archaeology, Simon Fraser University, Burnaby, B.C., V5A 1S6 Canada

(Received 28 June 1995, revised manuscript accepted 7 May 1996)

The relationship between dental attrition (nine stage scale) and specimen age, or functional age of teeth, is comparedbetween immature Middle Paleolithic (Neanderthal specimen count=28, tooth count=165) and Upper Paleolithic(anatomically modern specimen count=54, tooth count=338) samples. All specimens are from Western Europe.Maturation rate is treated as the same for both samples for analytical purposes and evaluated in terms of the results.Analysis of reduction in crown volume with age indicates that the wear rate for the posterior permanent (P4–M3) andprimary teeth (dm2) does not differ between the samples; however the anterior primary teeth (di1–dm1) are more wornin the UP sample with age held constant. By contrast, anterior permanent teeth (I1–P3) wear more rapidly in theNeanderthal sample. Similar results are obtained in a comparison of relative attrition.The MP sample of anterior primary teeth lags about a year behind the UP sample in the degree of accumulated wear

up to about age 5 years. Thereafter differences disappear. Analysis on the basis of individual specimens (ANOVA)confirms these results with the MP sample of anterior primary teeth significantly older (P=0·0498) at successive wearstages. While the rate of wear of the anterior primary teeth does not differ between the samples, the age of attritiononset differs by about 1 year; age 2 years in the UP sample and 3 in the MP sample. It is concluded that dietarysupplementation of infants commences 1 year earlier on average in the UP. Data are insufficient to determine whetherthe Late and Early Upper Paleolithic samples differ significantly in the study variables. Apart from the finding of earlieronset of dental wear in the UP, the lack of difference in the relationship between age and tooth wear supports a modelof similar maturation rates and dietary abrasiveness in the Middle and Upper Paleolithic. Significantly earlier dietarysupplementation of UP infants may have reduced birth spacing through diminished hormonal inhibition of ovulation.These results are compatible with demographic increase in the UP. ? 1997 Academic Press Limited

Keywords: LATE PLEISTOCENE, NEANDERTHAL, UPPER PALEOLITHIC, DENTAL ATTRITION,AGE AT DEATH, DIET.

Introduction

T he study of tooth wear as a guide to behaviourand life history has a long tradition in physicalanthropology (e.g. Molnar, 1972; Patterson,

1984 and contained references; Powell, 1985 and con-tained references; Dreier, 1994). However, fossil hom-inine of the Late Pleistocene have been little studied fordental attrition. The Gibraltar Neanderthal child wasreported by Carbonell (1965) to show more wear thanobserved in modern children. A study of microscopicenamel striations on the same specimen (Fox & Perez-Perez, 1993) led the authors to suggest a high meat-content diet like that of eskimos. A similar result isreported for the Middle Pleistocene individuals fromArago (Tautavel) (Puech, 1982) who wore their teethmuch like those of meat-consuming aborigines of thelast century; at a lesser rate than observed among purevegetarians. Wallace discussed incisor rounding in LaFerrassie 1 as an indication of industrial task wearrelated to clamping with the jaws (1975); while Smith(1977a,b) studied relative attrition and consequent oralpathology in specimens from Riss to Late Wurm.

6770305–4403/97/080677+24 $25.00/0/as960151

Beyond descriptions of single specimens, there are nointegrative studies of dental attrition among immaturefossil hominines from Europe.Wear on the teeth is a function primarily of the

interaction of the physical consistency of the diet andthe time during which a tooth has been functioning(industrial tasks are ignored for the moment) (Molnar,1971). With each variable alternately held constant, itis possible to compare developmental rates and dietaryconsistency between populations, such as those fromthe Middle and Upper Paleolithic, in which neithervariable is well understood. For example, a study ofdental attrition in adults by Smith (1977a) from theregion and time span reported here found that increas-ingly recent hominines with smaller teeth and facesshowed elevated levels of dental attrition; leading tothe conclusion that facial reduction was not linkedto reduced functional demand on the face. A possibleweakness with this conclusion is our inability to be sureof equivalent age compositions of samples composedof adults of indeterminate age.Furthermore, there are suggestions in the literature

that Neanderthals matured more quickly or, at least,

? 1997 Academic Press Limited

678 M. Skinner

erupted some teeth earlier than did anatomicallymodern humans (e.g. Legoux, 1966; Wolpoff, 1979;Stringer et al., 1990; Zollikofer et al., 1995). While it isinappropriate to attempt a complete discussion of thetopic here (see discussion in Skinner (submitted), it canbe pointed out that much of this viewpoint stems fromstudies of the Gibraltar child whose perikymata countis low and whose brain case is large (Dean et al., 1986;Zollikofer et al., 1995). Minugh-Purvis (1988) hasshown that the Gibraltar child is unusual amongNeanderthals for its size while the utility of perikymatacounting for age at death estimation has been stronglycalled into question by the results of Huda & Bowman(1994). Recently, Tompkins (1991), Trinkaus &Tompkins (1990) and Trinkaus (1995) have suggestedthat Neanderthal patterns of dental formation, erup-tion and developmental dental histology fall withinmodern human ranges of variation, although possiblyat the faster end, implying similar life history segments.Nevertheless, the suggestion of Neanderthal precocitycan be evaluated by assuming dietary equivalence andcomparing dental wear. Alternatively, maturation ratescan be assumed to be the same so as to compare dietaryabrasiveness. Some have maintained that the advent ofstone boiling and preparation of soup occurred in theUpper Paleolithic (Hadingham, 1979; Pfeiffer, 1986).Arguably this would soften the diet and reduce dentalwear.Several dental variables could potentially affect rates

of dental attrition; these include size, which is largeron average in Neanderthals (Brace et al., 1987) andenamel thickness. Two studies suggest that enamelwas thinner in Neanderthals (Zilberman et al., 1992;Molnar et al., 1993). The latter study found thatmodern permanent molar enamel was approximately1·3 times thicker than in Krapina Neanderthals whichthe authors tentatively attributed to systemic stress orenlarged pulp chambers among the latter. Furtherconsideration of this problem (Smith & Zilberman,1994) confirmed that Neanderthals had both thinnerenamel and enlarged pulp chambers.All else being equal the following predictions can be

made for MP populations:

(a) given accelerated dental (and presumably so-matic) maturation, dental wear will have anearlier onset but accrue at a lesser rate (or alter-natively, at equivalent wear stages, age will begreater);

(b) given the absence of stone boiling, wear will begreater;

(c) given larger teeth, wear will be less;(d) given thinner enamel, wear will be greater.

The variables of tooth size, enamel thickness, toothformation and wear are considered in this study.Attained tooth formation is used as a proxy for age atdeath using modern normative standards. While it isnot possible to control all the variables at once, analy-sis of primary and permanent waves and evaluation of

the magnitude of observed differences will enhance theinvestigator’s ability to judge the contribution thateach of the variables may be making. For example,if the pattern of differences between the MiddlePaleolithic and the Upper Paleolithic changes fromearly to late ontogeny (primary versus permanentteeth) it can be argued that innate features of differ-ences in developmental rate, tooth size and enamelthickness are not the explanation (unless these changetheir relative states during ontogeny).Immature skeletal remains in which teeth are pre-

served permit the investigator to determine individualage at death with more precision and accuracy than isnormally possible with adult remains (Ubelaker, 1978;Buikstra & Meikle, 1985). Physiological and behav-ioural phenomena which are time dependent such asintroduction of solid foods in infancy, tooth wear,and developmental rates can be calibrated from toothformation to investigate potential differences betweenpopulations. The research reported here compareswear on the teeth between immature remains from theMiddle and Upper Paleolithic of Europe.

MaterialsData were collected between May and July 1982. Someadditional data had been collected in the summer of1973 and some information is taken from the literature(Malegni & Ronchitelli, 1989). Eighty-two specimensare reported here. A brief description of each specimenwith its provenance and repository is available fromthe author.

Delimitation of samplesThe Middle Paleolithic (MP) of Europe occurred inOxygen Isotope Stage 3, while the Upper Paleolithic(UP) continues into Stage 2 (Straus, 1995). Althoughthe European Late Pleistocene artefactual record indi-cates fundamentally the hunting of large mammalsthroughout, with a largely undocumented but presum-ably critical contribution from gathered resources,there is general, but not total, agreement that the UPwith an emphasis on blade technology heralded aprofound cultural change (Mellars, 1973). This wasmarked, in addition, by bone working, concentrationon usually a single species of mammal, especiallyreindeer, increased social group size, increased popu-lation density, personal adornment, trade contacts, andartistic accomplishment, including cave art (White,1985). For this reason, it was felt acceptable to retainthe classic demarcation between the Middle and UpperPaleolithic as the basis for comparison of dental attri-tion. The temporal delimitation of samples was basedon the assumption that the major, but certainly notsole, factor in comparative dental attrition would likelybe changing subsistence strategies either in food baseor means of food processing. This is not meant to reifythe commonly held view of rapid cultural change

Dental Wear in Immature Late Pleistocene European Hominine 679

across the Middle/Upper Paleolithic boundary (seediscussion in Straus, 1995). Probably neither timeperiod was behaviourally and culturally static.The upper time limit excludes Mesolithic individuals

who are felt to have been adapted to a lifestyle basedmore on exploitation of smaller, more solitary forestgame and a wide variety of faunal resources (Frayer,1978). The lower time limit is deliberately set at thepenultimate glaciation. One reason for this is thepragmatic wish to include in a rather small MiddlePaleolithic sample, important pre-Neanderthalimmature specimens like La Chaise. The other is inrecognition of the apparent cultural continuity betweensites from this earlier time period and Early Wurmsites:

It is in fact difficult to think of a single well-characterizedtool-form found in middle Paleolithic industries whichdoes not have an ancestry extending back at least to thepenultimate (‘Riss’) glaciation . . . (Mellars, 1973, cited inWhite, 1985).

Spatial delimitation of the samples is a functionprimarily of non-theoretical considerations. Themajority of accessible specimens are to be foundin western Europe; in terms of the immature LatePleistocene fossil record, a ‘‘francocentric’’ emphasis isunavoidable (cf. Straus, 1995). This and a 3-monthfield season curtailed collection somewhat. Quite anumber of specimens examined were excluded fromconsideration for reasons referred to above in thediscussion of suitability of specimens or because theywere deemed too ancient or recent (e.g. Badger’s Holeand Mother Grundy’s Parlour). I have excluded sevenimmature specimens from the MP sample because theyare either not Neanderthals, or are from North Africaor the Middle East and, arguably, form a separate MPclade of hominines (Stringer, 1994). These includeSkhul 1, Qafzeh 4, Qafzeh 11, Ihroud 3, Ksar ’Akil,Rabat 1 and Haua Fteah*. The comparisons drawnhere are between two samples which differ biologically,culturally and temporally.The total sample consists of 82 individuals with

503 teeth ranging in individual age from about 1 to18·6 years old. The earlier sample spans at least150,000 years, including 28 individuals with 165 teeth(mode=one tooth, six specimens), and the later samplespans only about 25,000 years, including 54 individualswith 338 teeth (mode=two teeth, 17 specimens). Since,for the majority of specimens, it was not possible toknow the sex of the individual, it was judged better touse mid-sex age determinations for each individualdespite acknowledged sex differences in dental matur-ation (Haavikko, 1970). The provisional nature of anyresults must be appreciated in light of these facts. Notall teeth were able to be included in all analyses.In summary, each sample contains individuals differ-

ing in largely uncontrolled ways in temporal and

*Details of dental formation and attrition for these specimens maybe obtained from the author.

spatial origin (some groupings of individuals are fromsingle sites, however), sex, dietary behaviour, facialconfiguration (at least in adults), and, possibly,maturation rate. Furthermore, the two samples containindividuals considered to differ at the subspecies or atleast ‘‘racial’’ level. The two samples are, however,demographically similar in that the mean ages at deathof the two samples do not differ significantly:MP=7·74 years, UP=7·02 years; t=0·142, P=0·5229(Figure 1). Interpretation of the significance of theresults must consider the role that each of thesevariables may play in having created any perceiveddifferences in attrition among these Late Pleistoceneimmature hominines.

MethodsThe basic relationship studied in this research isbetween occlusal tooth wear and functional age of thetooth. For a tooth to be included in the study it has topossess the following attributes:

(a) the tooth had to be sufficiently erupted that it wasprobably gingivally erupted;

(b) occlusal attrition, or the lack of it, had to bereliably scoreable, i.e. post-mortem enamel losscould not obscure attritional status;

(c) functional age of the tooth had to be ascertainableby reference to a normative eruption age for thattooth, i.e. age at death of the individual had tobe able to be determined, from which age theeruption age of the tooth was subtracted.

Age at death was found from tooth formation, or fromsome other age indicator such as root resorption wherenormative standards for such existed, or from theliterature. Put another way, the tooth had to be from anon-adult. For the purposes of this analysis, subadult-hood is defined dentally as a tooth (and individual) inwhich root formation (or that of another tooth fromthe individual) was not complete. Root formation isdefined as full closure of the apical canal (Fanning,1961). The age of socio-sexual maturation is notindicated in this definition.

Calculation of individual age at deathA Fisher Portable X-ray machine (65–90 kVp,20–10 mA) with collimator was transported to Europewhere dental film and powder developing chemicalswere purchased. A total of 258 radiographs were takenon approximately 59 specimens to supplement thoseearlier reported (Skinner & Sperber, 1982). All usefulradiographs were photographed and tooth formationwas determined from the photoradiographs and/ororiginal observations as detailed below. Age at deathwas derived by converting the degree of tooth for-mation as observed in radiographs to an equivalentdental age based on a combination of normativestandards. Nolla (1960) pioneered the use of dental

680 M. Skinner

formation for age determination but her standardshave been superseded by subsequent workers notedbelow. That tooth formation is a preferable means ofdetermining age at death in immature remains is widelyaccepted (Lewis & Garn, 1960; Demirjian, Goldstein &Tanner, 1973; Stewart, 1979; Buikstra & Mielke, 1985;Smith, 1991).Scoring tooth formation was based on criteria

supplied by Demirjian, Goldstein & Tanner (1973),modified for fossil hominine materials (Skinner, 1978).This is a nine-stage scale, including ‘‘0’’, which wasconverted into the scale provided by Haavikko (1970)as follows:

Haavikko’s normative standards are superior becausethey include both upper and lower jaws and the thirdmolars. However, since they do not include the pri-mary teeth, nor some of the beginning formation stagesfor the early maturing teeth, recourse was made to thefollowing additional sources:

(a) Fanning (1961): primary canine, primary firstmolar, primary second molar;

(b) Fanning & Brown (1971): female primarycanine—stage 8, male and female primary firstmolar and primary second molar—stage 8, lowerpermanent canine—stage 1, lower first (=third)premolar—stages 1 and 2, lower second (=fourth)premolar—stage 1, lower first permanentmolar—stages 1, 2 and 3;

(c) Aprile & Figun (1956, cited in Legoux, 1966):isolated upper primary teeth and lower primaryfirst and second incisors;

(d) Moorrees, Fanning & Hunt (1963): lower perma-nent canine—stages 2 and 3.

These studies provide age equivalents for stages ofdental formation for each stage of each tooth (withexceptions due to lack of detailed study) separated bysex. Consequently, it is possible to derive two ageestimates for a fossil hominine: as though it were male;and as though it were female. The ages at deathindicated by all teeth were combined and averaged anda mid-sex age calculated for analytical purposes.

Calculation of a tooth’s functional ageIdeally, in an analysis of this sort one would have alarge series of dentitions with no missing teeth which

Haavikko (1970) This study

Crown initiation (Ci) Crypt formation (1)Cusp coalescence (Cco) Initial crown formation (2)Crown 1/2 to 3/4 Crown 1/2 (3)Crown complete (Crc) Crown complete (4)Root 1/4 (R1/4) Root initiation (5)Root 1/2 to 3/4 Root advanced (6)Root complete, apex open (Rc) Root complete,

apical canal open (7)Apex closed (Ac) Apical canal closed (8)

would permit comparisons of dental attrition betweenindividuals of equivalent dental age. With fragmentaryfossil material this is not possible. To maximize samplesize, the tooth becomes the basis for comparison.However, since teeth erupt and start to wear at char-acteristically different times, one has to calculate afunctional age for a tooth. This allows teeth withequivalent functional ages, whether of different toothtypes or from different individuals, to be combinedfor analysis. There is no question that different popu-lations erupt their teeth at different times for acombination of genetic and environmental reasons(Haavikko, 1969). Since it is difficult to judge whichnormative standards would be most suitable forEuropean fossil hominids, I elected to continue to useFinnish data (which, after all, is northern European)because Haavikko provides gingival eruption schedulesfor the permanent teeth of the children whose toothformation was largely utilized in this study to age thefossil hominine children. Also, Nystrom (1977) haspublished gingival eruption standards for an additionalseries of Helsinki children. Utilization of a singlegeographical, and relatively genetically homogeneous,reference group for age determination and for calcu-lation of functional age of the teeth should provide ahigh degree of confidence in the uniformity of results.

0 2 4 6 8 10 12 14 16 18 20 years

7

Cou

nt

3

2

4

5

6MP, N = 28 individuals Mean age = 7.74 years

1

7

3

2

4

5

6 UP, N = 54 individuals Mean age = 7.02 years

1

Figure 1. Age at death distribution compared between the MiddlePaleolithic (MP) and Upper Paleolithic (UP). Mean age at death isalmost identical in both samples.


The Finnish data appear to accord with dental matu-ration studies from elsewhere (Moorrees, Fanning &Hunt, 1963; Demirjian, Goldstein & Tanner, 1973).The actual calculation of functional age for a tooth

is found by subtracting gingival eruption age for thattooth from the previously calculated individual age atdeath (see above). The latter figure is expressed asmid-sex age since for the majority of specimens sex wasunknown. This procedure provides an estimate of agein years during which a tooth has been gingivallyerupted and exposed to attritional forces, ie. attritioncan be compared to how long a tooth has beenwearing.

AttritionMammalian tooth size is assumed, generally, to beappropriate to diet (van Valen, 1972). Thus, regardlessof variables such as differences in life history, toothsize, enamel thickness, and dietary consistency, relativeattrition should not differ greatly between species.Natural selection would militate against any variationsin tooth size that were needlessly large or so small asto reduce reproductive success. Human choice couldpresumably override this force at least in the shortterm such that relative attrition could differ betweenpopulations.Attrition is defined as the progressive loss of occlusal

and interproximal contact tooth substance due tomasticatory behaviour. It embraces crown reductionfrom tooth–tooth contact and tooth–bolus contact;similarly it does not exclude tooth reduction due tooccasional or habitual non-dietary occlusion as in theperformance of industrial tasks or clamping of objectsbetween the jaws to aid in manipulation. The lattercause of tooth substance reduction, while not felt to bea significant factor in dentally immature individuals,was specifically countenanced in this study by treatingthe anterior and posterior teeth as separate analyticalunits (see below). This study does not address potentialdifferences in enamel quality. Thorough reviews ofboth ordinal and interval means of describing attritionare provided by Patterson (1984) and Powell (1985).The study reported here is designed to accomplishboth, and has built-in checks on replicability throughrecourse to sketches, photographs and tracings. Fur-thermore, it permits the amalgamation of differenttooth types, teeth of different size, and both primaryand permanent teeth.Attrition was recorded in four ways:

(a) by photographs of dentin exposure on occlusalsurfaces (see below);

(b) by sketches of dentin exposure on occlusal sur-faces. The function of these was to aid in tracingdentin exposure and crown outline onto acetatesheets from which to calibrate dentin exposure ona ratio scale (see below);

(c) by measurement of crown dimensions as follows:

height of cusp or crown from cemento-enameljunction at cervix, vertically to the worn or un-worn occlusal surface. This was taken labially onthe anterior teeth and buccally on the premolars.Crown height of molariform teeth was taken atthe following locations: mesio-buccal cusp (5–4,6–4); disto-buccal cusp (5–5, 1–6, 1–7, 1–8, andtheir antimeres); mesio-lingual cusp height (7–4,7–5, 3–6, 3–7, 3–8, and their antimeres). In someanalyses, crown heights were converted to z-scoresfor statistical analysis in which single observationsare expressed as standardized deviations from themean for a tooth group (e.g. MP anterior primaryteeth), so as to allow teeth of different innateheights to be compared for analysis of heightreduction as age increases. Cube root of crownvolume:

height#mesio-distal length#labio(bucco)-lingual breadth

was calculated for linear regression against age;(d) by assignment of an attrition score, subjectively

assessed at the time of examination. These scoreswere reviewed later in light of the photographsand sketches when all data were at hand forevaluation. Only rarely was it felt that an initialscore was in error. It should be noted that thenumbers are not meant to imply equal intervalsbetween stages although they do reflect thecontinuous nature of the variable. That is, theattrition scale is an ordinal description of wearon the teeth. Its advantage is that it expressesearly stages of enamel reduction (see below)which is necessary in immature remains but whichare usually ignored in attrition studies basedon adults (Molnar et al., 1983). While moredetailed schemes for scoring attrition exist (e.g.26 stages developed by Dreier, 1994), they cannotbe applied easily across tooth types as is neces-sary when working with quite fragmentaryremains.

The rating scale for attrition is described below.

Stage 0: No attrition. Teeth which have not pierced thegum or have only recently done so will be at this stage.Isolated (ageable) teeth with no attrition were excludedfrom the study unless there was sufficient root for-mation to be sure it was not still in its crypt at the timeof death. A tooth had to show substantial projectionabove the alveolar margin to qualify for inclusion inthe study when it was at this attrition stage.

Stage 1: Enamel faceting (trace). On individual cuspsof the primary molars and permanent molars thisearliest stage of attrition is visible as tiny planes orfacets which reflect light from their flat surfaces. Oncanine teeth, faceting can be seen on the side or tip of

682 M. Skinner

the crown while incisors show initial reduction ofmammelons. Otherwise there is no reduction in crownheight at this attrition stage.

Stage 2: Enamel rounding (mid). Cusp tips are slightlysmoothed and rounded with loss of angulated faceting.Main fissures and crenulations are largely pristine. Onincisors, the mammelons are worn away while theincisal edge is still narrow and unflattened. There isonly minimal loss of crown height. (There was oneindividual with primary molars best scored at this stagebut with a trace of dentin exposure—La Chaise 14,7–4, 8–5.)

Stage 3: Enamel flattening (advanced). There is appre-ciable reduction in crown height resulting in broad,flattish, low occlusal elevation. On cheek teeth themajority of the occlusal surface is involved althoughdeeper fissures may be little affected. Cusp tips areobviously rounded. There is trace, or typically no,dentin exposure but dentin may be discernible througha thin enamel layer. Canines may tend to show rela-tively more dentin than the other teeth at this stage.Incisors exhibit a broad, flat incisal edge with crownreduction and often darkly staining dentin visiblethrough a thin enamel veneer.

Stage 4: Slight dentin exposure. On molariform teeth,this stage is differentiated from the next by the fact thatattrition tends to be angled such that dentin is exposedfirst on one side of the tooth and only later on the otheras well. At this stage, one or two (rarely more) islandsof dentin are exposed on one side of the tooth (buccalin lowers, lingual in uppers). On premolars one cusp, asopposed to both, shows slight dentin exposure. Forcanines, only a small spot of dentin is exposed—aboutthe size of a pencil dot. Incisors show a thin strip ofdentin, tapering mesially and distally—about the widthof a thin pencil line.

Stage 5: Dentin advanced. Dentin islands show on bothsides of molariform teeth of a size exceeding that of theprevious stage. There is no coalescence of dentinislands. Larger spots and strips of dentin may be seenon canines and incisors, respectively.

Stage 6: Strong dentin exposure. On molariform teeththere is coalescence of two or more islands of dentineven to the point where enamel remnants may onlyremain on the central occlusal surface. There ismarked crown height reduction. Canine teeth showlarge dentin exposure with crowns about half originalheight. Incisors now resemble canine teeth with anexpanding circle of dentin within the dentin strip,due to encroachment of attrition on the deep pulpchamber.

Stage 7: Enamel ring. All occlusal enamel is worn awayon molariform teeth leaving only an enamel ring of

fairly uniform width circumferentially. There may bedarkly stained islands of secondary dentin. Canines arejudged to be at this stage by having circumferentialenamel of a width similar to that of posterior teeth.Incisors show very strong height reduction with lossof interproximal contact and round or oval dentinexposure to a marked degree.

Stage 8: Root involvement. While self-explanatory, thisoccurs on the labial and buccal side of mandibularteeth and on the lingual side of maxillary teeth. Thisstage was not observed in the current study.

Calculation of proportion of dentin exposureIt has proved difficult to quantify attrition on enamel,which is why studies of attrition usually employ anordinal scale. However, it is feasible to measure thedegree of dentin exposure in a fairly replicable fashion(see discussion in Powell, 1985). The procedureadopted in this study is to trace the outline of exposeddentin within the crown outline similarly traced fromenlarged photographs of occlusal surfaces. The amountof dentin exposure is expressed as a proportion (%) ofcrown outline. This method does not include enamelattrition at all; a not insignificant factor when dealingwith infants. This, however, is accomplished throughthe ordinal scale approach described above.Given the variable appearance of worn occlusal

surfaces and the inherent difficulties of consistentlyscoring dental attrition when travelling from collectionto collection, all occlusal surfaces were photographedand sketched to record the extent of dentin exposure.This was in addition to a system of scoring relativedental attrition assessed visually at the time of exam-ination. Close-up photography was done with a200 mm f/5·6 Medical-Nikkor lens equipped with aring-light and a series of screw-on supplementarylenses which permit magnifications ranging from1/15# to 3#, i.e. the size of the image created on thenegative. More than 700 photographs were reproducedbased on about 82 specimens. These form the basis forthe study described below of proportion of dentinexposure, plus they served as a check on the attritionscoring. Ilford Pan F ASA 50 black-and-white film wasused with the camera set on ASA 25 followed byprolonged development for extra high quality detail.

Statistical analysisSimilar analyses were made of ‘‘teeth in combination’’(regardless of source) and ‘‘individuals’’. The formerwere undertaken to search only for patterns, in recog-nition of the possibility of making a Type I error due tostatistical redundancy among teeth from single jaws.The latter were made to test for statistically significantdifferences. Most specimens are very incomplete andmay not contain the same teeth. Hence, the systemsfor scoring dental attrition (classes) and dental height


(z-scores) were created so as to be comparable acrosstooth types. By this means the probability of making aType II error was minimized. Specifically, individualsare represented only by single teeth. These were foundby random selection of one tooth per individual.In those analyses of teeth in combination, regardless

of the individuals from which they derived, analysiswas performed separately by location and wave. Theseanalytical categories were created to allow for thepossibility of differential (possibly non-dietary) wearon the anterior teeth and the possibility of markedlydifferent attrition environments in infancy and laterchildhood (as well as the difference in tooth sizebetween baby and adult teeth). The permanent teethwere divided into an anterior and posterior moiety offour teeth each composed of the I1 to P3, and P4 toM3, respectively. Similarly, the primary teeth weredivided into an anterior group (i1, i2, c, m1) and aposterior tooth (m2). The latter grouping divided theprimary wave into an early erupting group (c. 6–18months) and a later erupting group (c. 27 months).All statistical procedures were carried out usingStatView 4.01.

ResultsThe role of enamel thickness

While others have shown that Neanderthals havesignificantly thinner enamel (Zilberman et al., 1992;Molnar et al., 1993), further analysis is undertaken soas to compare this attribute directly between the twofossil samples under consideration here. Measurementsof enamel thickness (and mesio-distal crown length)were taken directly on radiophotographs and scaled byreference to crown length measures taken on originalspecimens. Measures were taken at the visibly thickest

enamel. Analysis was undertaken only on the mostcommonly preserved tooth types (mandibular primarysecond molar and permanent first molar) and only onthose specimens with attrition stage 2 (enamel round-ing) or less. The results are shown in Table 1. Thethicknesses shown are (scaled) absolute values; the MPprimary tooth enamel is 0·27 mm thinner (c. 22%)while the permanent molar enamel is approximately0·49 mm thinner (c. 30%). A larger sample reported byZilberman et al. (1992) showed the thin permanentmolar enamel of Neanderthals. The value obtainedfor the UP permanent enamel sample (1·6 mm) is thesame as that reported for modern humans by Puech(1982).

Absolute wearCrown volume was calculated by multiplying the threelinear dimensions and finding the cube root so as togenerate a linear variable for regression against func-tional age. The results are shown in Figure 2(a)–(d).Not surprisingly, given the contributing variables, thecorrelation values are very low. Consequently, theapparent differences among tooth groups and betweenthe MP and UP samples should be interpreted withcaution. Nevertheless, the figure is a good guide todifferences detected in subsequent analyses.It seems that, apart from the obviously larger teeth

on average of the MP sample, there are few differences.Apparently, the anterior primary and posterior perma-nent teeth wear at the same absolute rate while thesecond primary molar may wear faster in the UPsample while the anterior permanent teeth wear moreslowly in the UP sample. These are merely indicationswhose validity and significance have to be determinedfrom more exacting analyses.

Table 1. Comparison of enamel thickness between the MP and UP samples based on teeth in combination with minimal or no attrition

Culture

Second primary molar First permanent molar

CountMean enamelthickness (mm) t-value P-value Count

Mean enamelthickness (mm) t-value P-value

MP 4 0·92 "0·270 0·0172 1 1·12 — —UP 11 1·19 4 1·61

Table 2. Comparison of interaction of dental attrition and functional age between the Middle and Upper Paleolithic samples based on teeth incombination (ANOVA—see Figure 3(a)–(d))

Tooth groupMean agedifference

Number Culture+functional age Culture+attr. class

MP UP F-value P-value F-value P-value

Anterior primary 0·82 69 113 2·364 0·1261 3·432 0·0019Anterior primary (sans Att. Stage 6,7) 1·40 61 91 16·278 <0·0001 2·617 0·0270Posterior primary "0·73 33 62 0·471 0·4944 3·422 0·0073Anterior permanent "0·61 31 82 4·901 0·0291 3·486 0·0060Posterior permanent "0·55 32 71 <0·0001 0·9927 1·683 0·1446

684 M. Skinner

Relative wearFigure 3(a)–(d) shows the relationship between attri-tion wear stage and functional age of a tooth for eachtooth group. This type of analysis does not assessattrition rate per se, but rather compares the differencein mean functional age at each successive wear stagefor the two samples (ANOVA) (Table 2). All suchanalyses based on ‘‘teeth in combination’’ may showType I error due to statistical redundancy. Resultsconform fairly well to the previous analysis. Becausethe anterior primary teeth show the most consistentand statistically significant difference in attritionbetween the MP and UP samples, this tooth group willbe dealt with last, and in more detail.

Posterior primary toothUnlike the previous analysis, there is no indicationfrom ANOVA that the two samples differ consistentlyin the functional ages at successive wear stages of theposterior primary (dm2) tooth. While the rate of crown

volume reduction (Figure 2(b)) is faster in the UPsample, a comparison (ANOVA) of mean functionalages over successive attrition stages (Figure 3(b)) doesnot differ between the two samples. Indeed, it is clearfrom Figure 4 that this tooth does not differ in thedevelopmental age required to reach slight dentinexposure (good large sample) nearly as much as do therest of the primary teeth. It may be concluded that inthis part of the mouth at the developmental agesinvolved (¢3 years or so), what was a large differencebetween the samples (UP showing more worn teeththan the MP sample at similar ages) is starting todisappear. It is distinctly possible that the thinnerprimary enamel of Neanderthals is causing their teethto wear faster once it is breached.

Anterior permanent teethNeanderthals show (Figure 3(c)) a significantlyyounger mean age (ANOVA) at successive wearstages for these teeth (P=0·0291). In other words,

CubRootVol. = 9.132 – 0.199 * FunctAge; R2 = 0.178 (MP)

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CubRootVol. = 9.247 – 0.069 * FunctAge; R2 = 0.092 (UP)

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CubRootVol. = 8.133 – 0.083 * FunctAge; R2 = 0.35 (UP)CubRootVol. = 6.146 – 0.055 * FunctAge; R2 = 0.015 (UP)CubRootVol. = 6.823 – 0.074 * FunctAge; R2 = 0.026 (MP)

CubRootVol. = 9.547 – 0.058 * FunctAge; R2 = 0.055 (MP)

Figure 2. Regression of cube root of crown volume against functional age of teeth in combination: (a) anterior primary (di1–dm1); (b)posterior primary (dm2); (c) anterior permanent (I1–P3); (d) posterior permanent (P4–M3) teeth. In this analysis, the anterior primary andposterior permanent teeth wear at the same rate in both samples, whereas the posterior primary teeth wear more slowly and the anteriorpermanent teeth wear more rapidly in the MP sample. There is considerable overlap between the two distributions. ,: Middle Paleolithic;+: Upper Paleolithic.


Neanderthal anterior permanent teeth are wearingfaster or commence wear earlier. This could beexplained if Neanderthals erupted these teeth earlier;but this would not explain their faster rate of wear(Figure 2(c)). Thinner enamel could be a factor herebut, if so, would also have produced the same effectamong the posterior permanent teeth, for which thereis no evidence.The same kind of analysis, avoiding type I error,

allows only one anterior permanent tooth to representthe individual, but suffers from insufficient sample sizesto show significant differences. Given the relativelysmall sample for this tooth group and the possibility ofmaking a Type I error using teeth in combination, onecan only very tentatively conclude that anterior toothusage in late childhood/early adolescence may havediffered between the two samples such that Neander-thals wore their front teeth down more quickly at thisstage of life.

Posterior permanent teeth

The posterior permanent teeth wear at virtually thesame rate in both samples whether one is consideringthe absolute reduction in crown substance (Figure2(d)) or relative wear (Figure 3(d)). The latter is to beexpected because dental attributes of size, shape,enamel hardness and thickness should be appropriatefor the growth rates and dietary abrasiveness of anyparticular taxon. Virtually all studies of dental attritionin traditional or prehistoric peoples concentrate onadults (e.g. Richards, 1984; Dreier, 1994). An excep-tion is the study by Molnar et al. (1983) of wear amongcontemporary Australian aborigines eating a tran-sitional diet. The first molar is reported to lose 0·5 mmof enamel between the ages of 6 and 18 years. In theresults reported here for the same tooth, both MP andUP samples lose 1·1 mm over the same period of time(more than twice the rate).

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Error bars: +/– 2 S.E.

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Figure 3. ANOVA of relationship between functional age and attrition class for teeth in combination: (a) anterior primary (di1–dm1); (b)posterior primary (dm2); (c) anterior permanent (I1–P3); (d) posterior permanent (P4–M3). Statistically significant differences are shown in thefigure; viz. MP is significantly older than the UP sample of the earlier stages of wear in the anterior primary teeth in combination, as well asfor anterior permanent teeth in combination. There are no differences between the two samples for the posterior primary and posteriorpermanent teeth in combination. .: Middle Paleolithic; /: Upper Paleolithic.

686 M. Skinner

Anterior primary teethThe anterior primary teeth show the most obvious andconsistent difference across several analyses betweenthe two samples. Figure 2(a), showing reduction incrown volume with increasing age, shows that theanterior primary teeth appear to wear at the same ratein both samples. There is no statistically significantdifference in age at death at successive wear stages(ANOVA) between the MP and UP samples, consider-ing all wear stages (Figure 3(a)). However, it is clearthat until the most severe wear stages (¢strong dentinexposure) are attained, the mean functional age of theMP sample is consistently older than the UP sample(P=0·0001). It seems the MP sample lags consistentlybehind the UP sample in wear of these teeth up to thelast wear stages. In other words, the MP children startto wear their anterior primary teeth later, but at muchthe same rate. This would suggest a delay in the onsetof wear in the MP sample (or, equally, earlier onset ofwear in the UP sample).In order to test for real differences, with the effect of

statistical redundancy removed, the analysis shown inFigure 5 compares the mean functional ages acrossattrition classes (ANOVA); in this analysis each indi-vidual contributes only a single, randomly chosen,tooth (increasing chances of a Type II error). Never-theless, the ANOVA shows a statistically significantdifference (P=0·0498) such that the UP sample isconsistently younger at equivalent wear stages up to‘‘dentin advanced’’ stage of wear.The following section is designed to examine the

meaning of the difference in the timing of anteriorprimary tooth wear between the MP and UP samples.A potential criticism of previous analyses (e.g. Figure3(a)) is that the wear stages are so broad (crude) that

each stage could embrace statistically significant meanfunctional age differences with no real difference be-tween the two samples. This can be tested by using amore precise estimate of attrition, viz. tooth height anddentin exposure.A regression of functional age against anterior

primary crown height (expressed as standardized devi-ations (z-scores) so as to remove the effects of innatedifferences in crown height) is similar between the twosamples except that the intercept difference (value of Y(age) when X (z-score)=0) is 0·9 years later in the MPsample (Figure 6(a)). In other words, about halfwaythrough the process of reduction in crown height dueto attrition, the MP sample lags just under a yearbehind. Restriction of wear to those teeth scored atstages less than or equal to dentin advanced (forcomparability to Figure 3(a)) shows a similar relation-ship except that the intercept difference has increasedto 1·4 years (Figure 6(b)).Figure 7(a) illustrates the regression of dentin

exposure against functional age. Unlike previousanalyses, this one, perforce, ignores earlier stages ofdental attrition. At equivalent ages, the MP sampleshows much less worn teeth. A more informative, ifless conventional, analysis is to regress functional ageagainst dentin exposure (Figure 7(b)). (This is not sobackward as it sounds since neither independent vari-able in these cases (7(a), (b)) is measured without error,as strictly speaking it should be for linear regression.)The value of this analysis is that it shows clearly thatthe rate of wear is the same in both samples, but thatthe MP sample lags behind the UP sample by about 1·1years (difference in intercept values in Figure 7(b)). Inother words, the onset of attrition of the anterior

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#1 #2 #3 #4 #5

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Figure 4. ANOVA of relationship between developmental age andprimary tooth position at a specific stage of dental attrition (slightdentin exposure) chosen to maximize sample size. The MP sample ofprimary teeth is significantly older at this attrition stage by anaverage of 1·06 years. .: Middle Paleolithic; /: Upper Paleolithic.

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2 1 7 45 7431 2


Figure 5. ANOVA of the relationship between functional age andsuccessive attrition stages for MP and UP individuals for the anteriorprimary teeth (di1–dm1). Overall, the MP sample is significantlyolder throughout wear..: Middle Paleolithic;/: Upper Paleolithic.


primary teeth appears to be delayed by about 1 yearon average in the MP sample, but they wear at indis-tinguishable rates thereafter.All three means of assessing attrition (loss of crown

substance volume (Figure 2(a)) and height (Figure 6),wear stage (Figure 3(a)), and dentin exposure (Figure7), indicate close to a 1-year lag in the MP sample.Thus the intercept values of Figures 6(a) & 7, whichprovide the difference in Y (functional age) when X(attrition)=0, are 0·9 and 1·1 years respectively. Simi-larly, the average difference in functional age withattrition held constant (Figure 3(a)) is 0·9 years.An alternative analysis, which includes the primary

second molar, is shown in Figure 4. Here, the relation-ship between individual age and primary tooth number

at just one attrition stage (slight dentin exposure forwhich a reasonably large sample of teeth can beamassed) is examined using ANOVA. The mean differ-ence in functional age at slight dentin exposureaveraged across all five primary teeth is 1·06 years(P<0·0001).Another analysis based on teeth in combination is

designed to disclose the developmental age at which theMP sample is demonstrably starting to lag behind(Figure 8). This analysis requires that developmentalage be treated as a factor variable (age classes as shownin the figure) with an ANOVA of percentage dentinexposure at successive age classes compared betweenthe samples. The delay in dentin exposure in theMP sample, while statistically significant (P=0·0097),

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NegFunctAge = –3.24 – 1.005 * z-score; R2 = 0.308 (MP)NegFunctAge = –2.343 – 1.299 * z-score; R2 = 0.405 (UP)

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NegFunctAge = –3.028 – 0.827 * z-score; R2 = 0.245 (MP)NegFunctAge = –1.65 – 0.396 * z-score; R2 = 0.105 (UP)

0.5

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Figure 6. Linear regression of functional age of anterior primaryteeth (di1–dm1) against tooth height (expressed as a z-score). Withinnate tooth height differences removed, the MP sample clearly lagsbehind the UP sample by 1 year or more (difference in intercept valueof Y when X=0): (a) all wear stages; (b) wear stages up to andincluding advanced dentin exposure (cf. Fig. 3(a)). ,: MiddlePaleolithic; +: Upper Paleolithic.

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Figure 7. Linear regression using all attrition stages of (a) pro-portion (%) of crown surface showing dentin exposure againstfunctional age of anterior primary teeth in combination; (b) thereverse. The UP sample has markedly more worn teeth; with the MPlagging behind by about 1 year (cf. intercept values in (b)). Thesedifferences probably reflect early onset of attrition in the UP sample.,: Middle Paleolithic; +: Upper Paleolithic.

688 M. Skinner

is only mildly expressed in infants aged less than2·5 years, but is present by age 2·5–3·4 years andmaintained into older childhood. It would seem thatby age 2·5 years the UP sample is starting to wear itsanterior teeth quite significantly (already into dentineexposure) whereas the MP sample is still no more wornby age 2·5–3·4 years than it was in the previous ageclass. A similar analysis compares average individualage at the beginning of enamel attrition (trace wear).Although samples are rather small (MP=three teeth,UP=13 teeth), the difference in mean age is statisticallydifferent: 2·0 years in the UP and 3·3 years in theMP (t=2·787, P=0·0145). An obvious interpretationto be placed on these results is that the introduction ofsolid foods occurred 1 year earlier on average in theUP sample, commencing by age 2 years.

DiscussionThe essential findings of this study are that, overall,statistically significant differences in attrition are fewand those that do exist are found in the anteriordentition, most particularly the primary teeth whichare markedly more worn in the UP sample with ageheld constant. The working assumption of this researchwas that developmental rates did not differ between theMP and UP hominines. Tompkins (1991) andTrinkaus & Tompkins (1990) have reasoned that sincemodern variation in eruption sequences and relativetooth formation embraces those patterns seen in theMP, there is no need to assume Neanderthals maturedat significantly different rates from anatomic moderns.This view is not universal. Wolpoff (1979) hassuggested that third molar eruption in Krapina

Neanderthals occurred relatively early, at age 15 years(assuming the rest of the dentition matured at modernrates) while Stringer et al. (1990) reason that theGibraltar child died at age 3·9 years (as opposed toc. 5·1 years reconstructed by Skinner & Sperber, 1982).The Stringer study suggests that MP populations couldhave matured in approximately 80% of the timerequired for more anatomically modern samples. Sub-stituting this maturation rate steepens the wear slopefor the MP sample, but has very little effect on thepreviously disclosed patterns for all tooth groups,except the anterior permanent teeth which wear evenfaster in the MP sample.Given the lack of significant differences between the

MP and UP tooth groups (except for the 1 year delayin onset of wear on the anterior primary tooth group inthe MP sample), the simplest interpretation is thatneither the diet nor the overall growth rate differedbetween the two samples during ontogeny. If oneassumed the validity of a faster growth rate forNeanderthals, say 80%, then their diet must havebeen proportionately more wearing (ie. 1·25#). If afaster growth rate included the anterior primary teeth,the difference is not enough to remove the effect ofthe large lag in wear of these teeth in Neanderthals(P changes from 0·0001 to 0·0274). A more economicalscenario is that the growth rates were the same in bothsamples, and so was the abrasiveness of the diet.The effect of thin enamel in the MP sample needs to

be considered. Clearly, this is not the explanation forthe relatively reduced wear of their primary dentition.At all but the last two stages of wear for the anteriorprimary dentition, the MP sample shows higher func-tional ages. In other words, it is taking the MP primarytooth sample longer to attain these wear stages. Thinenamel in the MP sample would tend to reduce theobserved difference, or, put another way, the UPsample shows markedly more wear on the primarydentition. As for wear of the permanent teeth, Figures2(d) and 3(d) show no difference between the twosamples. Consideration of the thin enamel in the MPsample indicates that the UP diet must have been ashard as, if not harder than, that of the MP. The effectof relative enamel thickness is less than that due toother causes such as delay in onset of wear (anteriorprimary teeth) or tooth usage (anterior permanentteeth).MP samples had larger primary and permanent

teeth on average. All else being equal, this wouldtend to reduce their rate of wear. One implication ofthis study is that ultimately the MP sample will runout of anterior permanent tooth substance earlier;thus root wear would commence a bit earlier in theMP sample, which could conceivably lead to compro-mised health and/or productivity in older adults and,in a demographic study, to their being over-agedslightly. The difference seems mild. However, perhapsthe thinner enamel of MP teeth would exacerbate thedifference.

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><

Figure 8. ANOVA of relationship between proportion (%) of dentinexposure and individual age class in anterior primary teeth. By ageclass ‘‘2·5–3·4 years’’ the MP sample has not increased dentinexposure while the UP sample has; this probably reflects the earlieronset of weaning in the latter sample. .: Middle Paleolithic; /:Upper Paleolithic.


VariationIn order to examine Hayden’s proposition (1993) thatin the UP there was an increase in social heterogeneity,the variance ratio for mean individual age with attri-tion class held constant is compared in Table 3. Theunit of analysis is the tooth, increasing the chance ofType I error. At younger ages (c.<6·3 years old) thereare two instances out of seven in which the F-valuediffers significantly between the MP and UP. In bothcases the UP sample is more variable. In the older ageclass (>6·2 years), in one instance (out of seven), theUP sample is more variable, and in one the MP sampleis more variable. The results provide mild support forHayden’s model that variation in the individual’sexperience with dietary attrition is increased in the UP.

Changes within the Upper PaleolithicJustification was provided earlier for an analyticaldistinction between the Middle and Upper Paleolithic.Frayer (1984) demonstrated that during the LatePleistocene of Europe there is progressive decline indental areas with the Early Upper Paleolithic (EUP)sample intermediate between the MP and Late UpperPaleolithic (LUP). Consequently, it is appropriate tocompare the timing of dental attrition within theUpper Paleolithic. The EUP sample consists of 77 teethfrom 15 individuals (i.e. about one-quarter of the totalUP sample) which is really too small for satisfactorystatistical analysis. Nevertheless, examination of theresult (Table 3) shows that the EUP sample, like theMP sample, shows higher mean ages at successive wearstages for the anterior and posterior primary dentition(F=10·534, P=0·0019, and F=4·012, P=0·0518) and

no significant differences for the permanent dentition.This analysis contains a heightened chance of Type Ierror. It may be concluded that the apparent differ-ences in dental attrition that exist between the MP andUP arose, possibly, in the LUP.

Summary and ConclusionsA comparison of the interaction of tooth age and toothwear between immature cohorts of Middle and UpperPaleolithic specimens shows that, apart from earlieronset of wear of the anterior primary teeth in the UPsample, and faster rate of wear for the anteriorpermanent teeth of the MP sample, the other two toothgroups do not differ significantly. With the possibleexception, then, of the anterior permanent teeth therates for the other tooth groups are indistinguishable.Therefore, if one can assume dietary equivalence dur-ing the growing period, there is no evidence for theirmaturing at different rates (contra Stringer et al., 1990).If, on the other hand, the MP sample is deemed tomature more quickly, then the UP diet must have beenless abrasive to an equivalent degree to maintainsimilar wear rates. While soups may be an UpperPaleolithic invention (see descriptions in Hadingham,1979 and Pfeiffer, 1986), the advent of storage(Soffer, 1985; Hayden, 1993) may have included thesmoking and drying of meats, rendering them moreabrasive. Consequently, it remains problematicwhether one can characterize the UP diet as generallysofter or harder than that of the MP.Further discussion is limited to the anterior primary

teeth which commence wear significantly earlier in theUP sample. Different interpretations can be placed on

Table 3. Comparison between MP and UP samples of variance ratios for specimen age at successive attrition classes

Attrition

¡6·2 years ¢6·3 years

F-value (MPn:UPn) P-value F-value (MPn:UPn) P-value

None 0·107 (5:19) 0·0218 1·121 (9:16) NSEnamel trace 0·156 (11:25) 0·0022 1·601 (11:31) NSEnamel mid 0·632 (10:17) NS 2·052 (14:27) NSEnamel advanced 1·531 (18:29) NS 1·489 (11:38) NSDentin slight 0·719 (25:34) NS 2·803 (22:34) 0·0117Dentin advanced 0·753 (8:18) NS 0·655 (12:18) NSDentin strong 0·000 (2:6) — 0·069 (6:19) 0·0039

Table 4. Comparison of interaction of dental attrition and functional age between the Early (EUP) and Late Upper Paleolithic (LUP) samplesbased on teeth in combination (ANOVA)

Tooth groupMean agedifference

Number Culture+functional age Culture+attr. class

EUP LUP F-value P-value F-value P-value

Anterior primary 1·94 11 53 10·534 0·0019 0·225 0·7990Posterior primary 0·83 17 34 4·012 0·0518 0·491 0·7424Anterior permanent "0·72 26 53 0·0067 0·9349 1·087 0·3701Posterior permanent "1·25 23 43 0·027 0·8707 1·952 0·1145

690 M. Skinner

the significance of this observation. Supplementationmay occur as early as the first few days after birth, asamong the Apsaroke Eskimo who give the baby a pieceof partially dried blubber to suck until the mother’sflow of milk is adequate (Gidley, 1976); and cancontinue for years as among the !Kung who introduceadult foods at 6 months or earlier but suckle veryfrequently for 3–4 years (Gaulin & Konner, 1977).Suckling frequency can override the potential effects ofdietary supplementation on ovulation (Gaulin &Konner, 1977; Konner & Worthman, 1980). Thus, theadvent of dental attrition does not necessarily signalweaning. For example, plants and pieces of meat aregiven to !Kung infants as pacifiers and objects tomouth (ibid.). Similarly, on the Pacific Northwest somenursing infants were also given fat meat on a stringtied to their big toe (to prevent choking)(McKechnie, 1972). Bullington (1987) found that ageat onset of dental microwear did not differ among threeprehistoric groups thought likely to wean at differentages due to differences in group diet. Clearly, theadvent of dental attrition very likely signals theintroduction of solid foods and initiation of a weaningprocess which could be abrupt or gradual.A major determinant of fertility in non-

contracepting mothers is suckling frequency(McNeilly, 1977; Jain & Bongaarts, 1981). Humanreproductive ecology is a complex phenomenon ofwhich suckling frequency is only one of many import-ant variables that influence ovarian function (Ellison,1990). The contraceptive effect of breast-feeding isdebatable. Clearly, when breast-feeding continuesfor more than a few months menses (and ovulation)returns. Indeed weaning is often precipitated by a newpregnancy (Santow, 1987). Nevertheless, dietarysupplementation of infants has been shown to haveimportant effects on suckling frequency, prolactinlevels and return of ovulation. Prolactin is secretedwithin minutes of nipple mechanical stimulation; thishormone in turn inhibits gonadal hormones whichcontrol ovulation (Konner & Worthman, 1980; Wood,1994). Use of a pacifier or the introduction of water orsolid foods can diminish prolactin levels (Jeliffe &Jeliffe, 1979; Zeller, 1987). In other words, while sup-plementation need not herald weaning, it will probablyhasten the process.This study echoes earlier work undertaken by

Brennan (1991) which concluded that UP infants mayhave been weaned relatively early. We can ask howlikely is this conclusion. A recent study of enamelhypoplasia among the same specimens reported herefound that the UP sample showed significantly moredevelopmental stress in infancy (Skinner, 1996). Thiswas tentatively attributed to combined effects of socialcrowding, vitamin A malnutrition and infection. Earlydietary supplementation could contribute to thisproblem through the oral introduction of pathogens.This begs the question, however, of why UP mothers

would have felt able to, or had to, wean a year earlier

than was the case in the MP. Conceivably, infantsupplementation led to renewed pregnancies which inturn would favour accelerated weaning. There is noparticular advantage in traditional societies to intro-duce early weaning. In fact the disadvantages are wellknown; for example, loss of passive immunity, loss oflactational amennorrhoea, loss of maternal–infantbonding, and dietary stress for the weaned infant.Rather, it seems likely that under traditional con-ditions the ideal would be prolonged lactation andthat a departure from this should be viewed asunintentional and possibly deleterious.Large mammal hunting, accompanied by an un-

known contribution from plant foods, prevailedthroughout the Late Pleistocene (Gamble, 1986).Climatic conditions probably limited the growingseason to about 3 months. Notably, climatic conditionsdeteriorated during Oxygen Isotope Stage 2; the glacialmaximum occurred in the middle Upper Paleolithic(Straus, 1995). If weaning at these times was tied toseasonality, then inter-population differences at age ofweaning would be separated by yearly intervals, assuggested by this study. Evidence of a delay in weaningamong Neanderthals, in the form of a 1-year lag indental attrition, may signal longer average birthspacing in comparison to the UP sample. Birth spacingis the most important variable affecting fertility(Scott & Johnston, 1985). Profound demographicand social changes ensue from reduced birth spacing(MacCormack, 1982; Constandse-Westermann,Newell & Meiklejohn, 1984). Thus, the finding ofrelatively early onset dental attrition provides a plaus-ible mechanism for population increase in the UP.

AcknowledgementsThe research for this article was completed during mytenure as Senior Visiting Research Fellow, Departmentof Archaeology and Prehistory, University of Sheffield.I would like to express gratitude for assistance towardscompletion of this project to the persons and insti-tutions listed below: Chris Stringer (British MuseumNatural History), Alan Bilsborough, Bernard Denston(Duckworth Laboratory, University of Cambridge),M. Leguebe (Institut Royal des Sciences Naturelles deBelgique), G. Ubaghs (Laboratoire de PaleontologieAnimale, Universite de Liege), Anne-Marie Tiller,Jean-Jaques Hublin, B. Vandermeersch (Laboratoirede Paleontologie des Vertebres et de PaleontologieHumaine, Universite Pierre et Marie Curie), YvesCoppens (Laboratoire d’Anthropologie, Musee del’Homme), Henri Delporte (Musee des AntiquitesNationales, Chateau de Saint Germain-en-Laye), ProfPiveteau (Institut de Paleontologie), Jean-Louis Heim(Institute de Paleontologie Humaine), Prof de Bonis(Laboratoire de Paleontologie des Vertebres, Univer-site de Poitiers), A. Roussot, G. Chauvin (Museed’Aquitaine), Mme Prudhomme (Musee d’Histoire


Naturelle, Bordeaux), Jean Guichard, AndreMorala (Musee National de Prehistoire, Les Eyzies),Mme Edmee Lauder (Musee d’Histoire Naturelle,Montauban), Michel Philippe (Museum d’HistoireNaturelle, Lyon), Jean-Francois Bussiere (Museed’Anthropologie Prehistorique, Monaco), JakovRadovcic (Hrvatski Prirodoslovni Muzej, Zagreb),Mirko Malez (Zavod za Paleotologiju i GeologijuKvartara).Particularly I am grateful to the often unnamed

assistant curators who helped me cope with installingthe radiographic apparatus and tolerated my brokenFrench. Social Sciences and Humanities ResearchCouncil of Canada provided funding. Student assist-ants were Anja Streich, Arne Carlson and BrianStrongman. Jack Nance, Richard Lazenby andAndrew Chamberlain gave me valuable advice. JeanMcKendry, my wife, was my very able research assist-ant and companion throughout the data collection andanalytical phases. Thank you.

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AppendixList of specimens with attributes of age, height, and attrition

Group SpecName Spec IndAge Tooth* FunctAge TthHt AttScor AttClass %Dent

EUP* Patau 26242 1 1·0 uldi1 0·20 6·30 4 DentSlght 4·30LUP* Ranis 2 1·0 lldm1 "0·30 - 1 EnamTrace -

lrdm1 "0·30 - 1 EnamTrace -

LUP Bruniqel545 3 1·4 lrdm1 0·10 - 0 None -

LUP EnfantsYoun 4 1·4 lrdm1 0·10 5·15 1 EnamTrace -

lrdm2 "0·80 - 0 None -

lldi1 0·80 4·90 5 DentAdv 14·53lldm2 "0·80 5·60 0 None -

lldm1 0·10 5·25 0 None -

urdm2 "0·90 - 0 None -

urdm1 0·20 5·65 1 EnamTrace -

uldm1 0·20 5·95 1 EnamTrace -

urdc "0·10 7·55 0 None -

uldi1 0·60 5·95 4 DentSlght 3·39lrdi1 0·80 5·00 4 DentSlght 14·23lrdi2 0·40 6·60 4 DentSlght 6·91lrdc "0·20 7·65 0 None -

urdi2 0·60 6·15 1 EnamTrace -

lldc "0·20 7·75 0 None -

uldc "0·10 7·85 0 None -

lldi2 0·40 6·00 3 EnamAdv 2·22urdi1 0·60 6·20 4 DentSlght 3·78uldi2 0·60 5·85 1 EnamTrace -

LUP Figuier 5 1·6 lldi2 0·60 6·30 4 DentSlght 8·00uldm1 0·40 4·95 4 DentSlght 2·27uldm2 "0·70 5·90 0 None -

lldc 0·00 7·60 3 EnamAdv -

lrdi1 1·00 5·10 5 DentAdv 15·38lldi1 1·00 4·55 5 DentAdv 18·31lrdi2 0·60 6·25 4 DentSlght 7·33lldm2 "0·60 5·95 1 EnamTrace -

lldm1 0·30 5·25 4 DentSlght 1·66lrdc 0·00 7·55 3 EnamAdv -

urdm1 0·40 5·00 4 DentSlght 1·47lrdm1 0·30 5·10 4 DentSlght 1·11

LUP Fees 6 1·8 lldm1 0·50 4·85 2 EnamMid -

lldm2 "0·40 5·50 1 EnamTrace -

LUP Laugerie-5 7 2·0 uldm1 0·80 4·95 1 EnamTrace -

uldm2 "0·30 6·20 0 None -

LUP LaugeriTooth 8 2·0 lldm2 "0·20 5·00 2 EnamMid -

LUP Isturitz-7 9 2·4 lrdm1 1·10 5·20 3 EnamAdv 1·79lrdm2 0·20 5·80 1 EnamTrace -

LUP Brun540/541 10 2·8 lldm2 0·60 5·60 2 EnamMid -

urdm1 1·60 5·45 3 EnamAdv -

uldm1 1·60 4·95 3 EnamAdv -

uldm2 0·50 - 2 EnamMid -

lrdm1 1·50 4·70 3 EnamAdv -

lrdm2 0·60 5·65 2 EnamMid -

urdm2 0·50 6·50 2 EnamMid -

lldm1 1·50 4·85 3 EnamAdv -

LUP Rochereil 11 2·8 uldi2 2·00 6·10 2 EnamMid -

urdm2 0·50 6·95 0 None -


uldm1 1·60 6·65 0 None -

urdm1 1·60 6·55 0 None -

lrdm2 0·60 5·95 1 EnamTrace -

uldm2 0·50 - 0 None -

lldm2 0·60 6·45 1 EnamTrace -

uldc 1·30 7·75 0 None -

urdi2 2·00 6·00 2 EnamMid -

uldi1 2·00 5·65 5 DentAdv 11·46urdi1 2·00 6·55 4 DentSlght 7·42

694 M. Skinner

Appendix continued

Group SpecName Spec IndAge FDI FunctAge TthHt AttScor AttClass %Dent

LUP Bruniquel 25 12 2·8 lldc 1·20 7·25 1 EnamTrace -

urdi1 2·00 5·35 4 DentSlght 9·47urdi2 2·00 5·75 3 EnamAdv -

uldi1 2·00 5·15 4 DentSlght 15·32uldi2 2·00 6·40 3 EnamAdv -

lrdm2 0·60 - 1 EnamTrace -

lrdm1 1·50 - 1 EnamTrace -

lldm2 0·60 - 1 EnamTrace -

lldm1 1·50 5·35 2 EnamMid -

LUP EnfantsOldr 13 3·0 urdm2 0·70 5·70 1 EnamTrace -

urdm1 1·80 5·55 2 EnamMid -



uldm2 0·70 6·05 1 EnamTrace -

uldi1 2·20 5·35 5 DentAdv 16·63uldm1 1·80 5·80 1 EnamTrace -



lrdc 1·40 7·35 3 EnamAdv -

lrdi2 2·00 6·00 4 DentSlght 4·90lrdi1 2·40 4·35 5 DentAdv 16·67lldc 1·40 7·40 3 EnamAdv -

urdi1 2·20 5·35 5 DentAdv 15·25uldi2 2·20 5·55 4 DentSlght 8·04uldc 1·50 7·40 4 DentSlght 2·04urdc 1·50 7·00 4 DentSlght 2·32

EUP Baousse 14 3·2 lrdm1 1·90 5·35 3 EnamAdv -


LUP Plac61401a 15 3·3 urdm1 2·10 5·45 2 EnamMid -

LUP Madeleine 18 3·4 urdm1 2·20 6·15 3 EnamAdv -

uldm1 2·20 6·15 3 EnamAdv -

uldm2 1·10 6·60 2 EnamMid -

lldm1 2·10 5·45 3 EnamAdv 0·81lldm2 1·20 6·15 2 EnamMid -

lrdm1 2·10 4·35 3 EnamAdv 0·77lrdm2 1·20 - 2 EnamMid -

lrdi2 2·40 6·20 4 DentSlght 4·60lrdi1 2·80 4·60 5 DentAdv 28·29lldc 1·80 7·65 3 EnamAdv -

lldi2 2·40 5·80 4 DentSlght 7·90lldi1 2·80 4·30 5 DentAdv 23·92uldc 1·90 6·90 3 EnamAdv 2·37uldi2 2·60 6·05 4 DentSlght 5·27uldi1 2·60 5·70 6 DentStrng 24·15urdc 1·90 7·30 3 EnamAdv 1·75urdi2 2·60 6·50 4 DentSlght 4·82urdi1 2·60 5·65 6 DentStrng 22·75

EUP Rouset 19 3·6 lrdm2 1·40 - 3 EnamAdv -

lrdm1 2·30 - 4 DentSlght -

LUP Laugr2/4/6 20 3·8 lrdm1 2·50 4·45 5 DentAdv 10·93lldm2 1·60 5·50 3 EnamAdv 0·43uldm2 1·50 5·90 3 EnamAdv -

lldm1 2·50 4·45 5 DentAdv 12·75lrdm2 1·60 6·80 3 EnamAdv 0·67uldm1 2·60 5·15 5 DentAdv 7·11

EUP Magrite2878 21 4·2 lrdm1 2·90 4·75 4 DentSlght 3·02lrdm2 2·00 5·60 4 DentSlght 2·07

EUP Patau26236 23 4·7 urdm1 3·50 5·55 4 DentSlght 1·04urdm2 2·40 6·65 3 EnamAdv 0·27

LUP Plac61401b 24 4·8 lldm1 3·50 4·20 6 DentStrng 12·75lldm2 2·60 5·80 4 DentSlght 1·10

LUP Isturitz-6 25 5·0 lrdm2 2·80 5·05 4 DentSlght 0·86lldm2 2·80 5·15 4 DentSlght 0·70

LUP St. Germ5–7 26 5·3 uldm2 3·00 5·40 4 DentSlght 0·75urM1 "1·10 7·25 0 None -

ulM1 "1·10 7·65 0 None -

urdm1 4·10 5·15 5 DentAdv 10·85urdm2 3·00 5·75 4 DentSlght 1·56llM1 "1·00 6·70 1 EnamTrace -

lldc 3·70 6·75 5 DentAdv 13·08uldm1 4·10 4·95 5 DentAdv 13·09lldm2 3·10 5·10 4 DentSlght 1·13lldm1 4·00 - 5 DentAdv 4·61


Appendix continued


EUP Solutre 27 5·4 lldm1 4·10 4·80 3 EnamAdv -

lrdm2 3·20 5·25 3 EnamAdv 0·06lldm2 3·20 5·70 3 EnamAdv 0·67

LUP Mas D’Azil 28 5·8 uldm2 3·50 5·60 4 DentSlght 0·96EUP Fontechvade 29 5·8 lrdm2 3·60 5·90 3 EnamAdv 0·22LUP Laugerie-1 30 5·8 lldm1 4·50 3·10 6 DentRing 35·56

lldm2 3·60 4·55 5 DentAdv 9·15LUP Laugerie-3 31 5·8 lrdm1 4·50 4·20 6 DentStrng 16·02

lrdm2 3·60 5·25 5 DentAdv 13·27EUP Roches 32 6·0 lrM1 "0·30 - 0 None -

lrdm2 3·80 - 2 EnamMid -

LUP Isturitz-5 33 6·2 lrdm2 4·00 4·90 4 DentSlght 1·68lrdm1 4·90 4·20 6 DentStrng 15·90

EUP LaQuina25 34 6·3 lldm2 4·10 5·85 3 EnamAdv 2·42llM1 0·00 7·70 1 EnamTrace -

LUP Bruniqel538 35 7·1 lldm2 4·90 - 5 DentAdv -

LUP StGerm-3 36 7·2 lrI2 0·20 10·35 1 EnamTrace -

lrI1 1·00 10·25 2 EnamMid -

lldc 5·60 4·40 6 DentStrng 30·40lrdc 5·60 3·60 6 DentStrng 46·49lrdm2 5·00 4·40 5 DentAdv 19·43llI1 1·00 10·05 2 EnamMid -

llI2 0·20 10·35 1 EnamTrace -

lldm1 5·90 4·00 6 DentStrng 22·54llM1 0·90 6·85 3 EnamAdv -

lrM1 0·90 6·70 2 EnamMid -

lldm2 5·00 5·25 5 DentAdv 10·56lrdm1 5·90 4·00 6 DentStrng 32·30

LUP Plac56029 37 7·5 lrdm2 5·30 4·55 5 DentAdv 17·02lrdm1 6·20 3·25 6 DentStrng 37·51

EUP Magrite2426 38 7·6 uldm1 6·40 4·90 6 DentStrng 22·94uldm2 5·30 5·65 4 DentSlght 4·39urdm2 5·30 5·85 5 DentAdv 5·14urM1 1·20 8·15 2 EnamMid -

ulM1 1·20 7·85 2 EnamMid -

EUP Vindija 39 7·6 lrdm1 6·30 4·10 6 DentStrng 14·40lrdm2 5·40 4·90 5 DentAdv 14·44lrM1 1·30 7·00 1 EnamTrace -

EUP Miesslingtal 40 8·0 lldm2 5·80 4·90 4 DentSlght -

llM1 1·70 6·00 2 EnamMid -

lldm1 6·70 3·45 6 DentStrng -

lrdm2 5·80 4·75 4 DentSlght -

lrM1 1·70 6·75 2 EnamMid -

lrdc 6·40 3·45 6 DentStrng -

lrI1 1·80 7·60 1 EnamTrace -

llI2 1·00 - 0 None -

lldc 6·40 3·40 6 DentStrng -

lrI2 1·00 - 0 None -

LUP StGerm-4 41 8·3 lldm1 7·00 3·50 7 EnamRing 54·51lldm2 6·10 4·55 6 DentStrng 32·02lrdm1 7·00 4·15 6 DentStrng 19·53lrdm2 6·10 4·55 5 DentAdv 16·51lrM1 2·00 5·95 3 EnamAdv -

llM1 2·00 5·95 3 EnamAdv 0·21llI1 2·10 8·05 4 DentSlght 8·68lrI1 2·10 8·30 4 DentSlght 4·54lrI2 1·30 9·20 3 EnamAdv 2·33llI2 1·30 8·65 3 EnamAdv 0·96

LUP Laugri15107 42 8·9 urI2 0·90 9·95 2 EnamMid -

LUP Brun536/537 43 9·5 urdm2 7·20 - 5 DentAdv -

urM1 3·10 - 1 EnamTrace -

lrM1 3·20 - 1 EnamTrace -

urdm1 8·30 - 6 DentStrng -

lrdm1 8·20 - 7 EnamRing -

lrdm2 7·30 - 5 DentAdv -

LUP Laugerie-31 44 9·8 uldm2 7·50 5·55 4 DentSlght 8·94LUP Plac61401c 45 10·0 urdm2 7·70 4·35 6 DentStrng 22·08

lrdm2 7·80 4·85 7 EnamRing 75·61lrdm1 8·70 4·15 7 EnamRing 49·01lrI2 3·00 9·15 4 DentSlght 3·00urdm1 8·80 4·80 7 EnamRing 15·70urM1 3·60 6·45 3 EnamAdv -

696 M. Skinner

Appendix continued


EUP Rois 46 11·1 lrC 1·30 12·65 2 EnamMid -

llP1 1·10 5·15 1 EnamTrace -

llM1 4·80 - 3 EnamAdv -

lldm2 8·90 3·70 6 DentStrng -

lrdm2 8·90 3·95 6 DentStrng 54·51llC 1·30 13·20 2 EnamMid -

lrM1 4·80 7·80 3 EnamAdv 0·28lrP1 1·10 - 0 None -

LUP LaugeriAdol 47 11·2 urI1 4·40 11·90 4 DentSlght 1·77urI2 3·20 9·85 2 EnamMid -

llM1 4·90 6·50 4 DentSlght 1·57ulM1 4·80 7·20 4 DentSlght 1·05llP1 1·20 3·70 0 None -

llI2 4·20 9·65 3 EnamAdv 2·17llC 1·40 12·85 0 None -

ulM2 "1·40 8·00 0 None -

lrM1 4·90 - 4 DentSlght 1·36ulP1 1·30 8·30 2 EnamMid -

lrM2 "0·60 7·05 1 EnamTrace -

lrP1 1·20 - 0 None -

llM2 "0·60 7·25 1 EnamTrace -

llI1 5·00 9·60 4 DentSlght 2·72lldm2 9·00 4·45 7 EnamRing 70·61uldm2 8·90 4·25 6 DentStrng 45·44ulI1 4·40 11·80 3 EnamAdv 1·41lrdm2 9·00 3·75 7 EnamRing 73·63lrI2 4·20 10·20 3 EnamAdv 1·77lrC 1·40 12·95 0 None -

ulI2 3·20 10·00 2 EnamMid -

lrI1 5·00 9·40 4 DentSlght 3·62LUP Morin 48 11·6 lrdm2 9·40 - 5 DentAdv -

lrM1 5·30 - 2 EnamMid -

LUP Farincourt 49 11·8 ulC 0·40 11·65 0 None -

EUP Parpallo 50 13·3 ulC 1·80 - 2 EnamMid -

urI1 6·40 - 4 DentSlght -

ulP1 3·30 - 2 EnamMid -

ulP2 2·40 - 2 EnamMid -

llM2 1·40 - 1 EnamTrace -


EUP GrEnfants-6 51 13·8 llI1 7·60 8·35 4 DentSlght 14·08lrI2 6·80 9·85 4 DentSlght 5·10urC 2·40 11·80 2 EnamMid -

urI1 7·00 11·25 4 DentSlght 7·05urI2 5·80 - 3 EnamAdv -

lrC 4·00 - 2 EnamMid -

lrI1 7·60 8·80 4 DentSlght 11·81uldm2 11·50 4·70 6 DentStrng 42·83llC 4·00 12·65 1 EnamTrace -

urP1 3·90 9·20 1 EnamTrace -

llI2 6·80 9·20 4 DentSlght 7·57ulP1 3·90 - 1 EnamTrace -

urM2 1·20 - 0 None -

lrP2 3·20 6·10 1 EnamTrace -

llP2 3·20 - 0 None -

ulM2 1·20 6·90 0 None -

ulM1 7·40 6·60 4 DentSlght 1·37urM1 7·40 7·05 3 EnamAdv 0·39llP1 3·80 - 0 None -

lrM1 7·50 - 3 EnamAdv 1·34ulI1 7·00 11·05 4 DentSlght 9·44urP2 3·00 - 0 None -

ulC 2·40 - 0 None -

ulI2 5·80 11·40 3 EnamAdv -

llM1 7·50 - 4 DentSlght 3·44lrM2 2·00 - 1 EnamTrace -

lrP1 3·80 - 1 EnamTrace -

llM2 2·00 6·55 2 EnamMid -

LUP RocDeSers 52 14·4 llP1 4·40 4·20 2 EnamMid -

lrdm2 12·20 5·05 6 DentStrng 37·74llP2 3·80 5·30 1 EnamTrace -

lrP1 4·40 4·50 2 EnamMid -

lrM1 8·10 7·05 4 DentSlght 1·51


Appendix continued


lrM2 2·60 - 2 EnamMid -

lrC 4·60 12·65 3 EnamAdv -

llC 4·60 12·65 2 EnamMid -

lrI1 8·20 9·05 4 DentSlght 8·42llM1 8·10 6·10 4 DentSlght 1·46llI1 8·20 9·00 3 EnamAdv 2·11lrI2 7·40 10·40 4 DentSlght 3·69

LUP Cap Blanc 53 15·6 ulI2 7·60 - 3 EnamAdv -

ulI1 8·80 - 4 DentSlght -

lrI1 9·40 - 4 DentSlght -

lrC 5·80 - 1 EnamTrace -

lrI2 8·60 - 1 EnamTrace -

ulI1 8·80 - 4 DentSlght -

urI2 7·60 - 3 EnamAdv -

urC 4·20 - 4 DentSlght -

llP1 5·60 - 1 EnamTrace -

ulM2 3·00 - 2 EnamMid -

llC 5·80 - 1 EnamTrace -

urM2 3·00 - 2 EnamMid -


llP2 5·00 - 1 EnamTrace -

llM2 3·80 - 1 EnamTrace -

ulP2 4·80 - 3 EnamAdv -

ulP1 5·70 - 3 EnamAdv -


lrM1 9·30 - 4 DentSlght -

llM1 9·30 - 4 DentSlght -

llI2 8·60 - 1 EnamTrace -

ulM1 9·20 - 3 EnamAdv -

ulC 4·20 - 1 EnamTrace -

llI1 9·40 - 4 DentSlght -


urP1 5·70 - 3 EnamAdv -

urP2 4·80 - 3 EnamAdv -

urM1 9·20 - 3 EnamAdv -

LUP Lachaud-5 54 15·8 llP1 5·80 4·60 3 EnamAdv -

lrI2 8·80 7·60 5 DentAdv 7·91lrI1 9·60 6·95 5 DentAdv 6·44llC 6·00 10·15 3 EnamAdv -

llP2 5·20 6·55 3 EnamAdv -

lrP1 5·80 4·65 3 EnamAdv -

lrM1 9·50 6·30 5 DentAdv 2·52llM1 9·50 5·30 5 DentAdv 3·79llM2 4·00 6·90 3 EnamAdv -

llI1 9·60 7·00 5 DentAdv 5·50lrC 6·00 9·65 3 EnamAdv 1·67llI2 8·80 8·75 4 DentSlght 13·11lrM2 4·00 6·60 3 EnamAdv -

lrP2 5·20 5·95 3 EnamAdv -

LUP Plac61397 55 18·0 llM2 6·20 - 1 EnamTrace -

llM1 11·70 6·20 3 EnamAdv 0·42llP2 7·40 4·70 2 EnamMid -

llC 8·20 11·05 3 EnamAdv 1·58LUP LaugeriNoNo 56 18·0 lrM1 11·70 6·35 5 DentAdv 5·67LUP Lachaud-3 57 18·6 lrM3 "0·40 10·70 0 None -

lrM2 6·80 6·15 3 EnamAdv -

lrM1 12·30 5·55 5 DentAdv 2·84llM2 6·80 6·05 3 EnamAdv -

llM1 12·30 5·65 5 DentAdv 3·21MP* Chatoneuf1 60 2·0 lrdc 0·40 8·00 0 None -

lrdm1 0·70 6·35 3 EnamAdv 1·41MP PechdeL’Aze 61 2·4 urdi1 1·60 6·00 4 DentSlght 10·58


urdm1 1·20 6·05 4 DentSlght 1·61urdc 0·90 7·95 0 None -

lrdm1 1·10 5·00 3 EnamAdv 0·74lrdc 0·80 6·50 2 EnamMid -

lldi1 1·80 5·20 5 DentAdv 11·11lrdi2 1·40 6·95 3 EnamAdv 1·88uldi1 1·60 6·30 4 DentSlght 4·95uldc 0·90 7·60 0 None -

uldm2 0·10 6·05 2 EnamMid -

698 M. Skinner

Appendix continued


lrdi1 1·80 5·05 5 DentAdv 10·58lrdm2 0·20 5·65 2 EnamMid 0·06uldm1 1·20 6·45 3 EnamAdv 1·09

MP Krapina 68 62 2·7 lrdm2 0·50 - 1 EnamTrace -

MP Archi 63 3·0 lrdm2 0·80 - 1 EnamTrace -

lrdm1 1·70 - 2 EnamMid -

lldm1 1·70 - 2 EnamMid -

lldm2 0·80 - 1 EnamTrace -

lldc 1·40 - 0 None -

MP RocDeMarsal 64 3·2 lldi1 2·60 5·40 5 DentAdv 13·26uldc 1·70 7·55 2 EnamMid -

uldi2 2·40 5·85 4 DentSlght 8·21uldi1 2·40 6·15 4 DentSlght 5·79urdc 1·70 7·40 1 EnamTrace -

urdi2 2·40 6·30 3 EnamAdv 1·75urdi1 2·40 6·85 4 DentSlght 2·55lrdc 1·60 7·85 1 EnamTrace -


uldm1 2·00 6·55 4 DentSlght 1·62uldm2 0·90 6·15 1 EnamTrace -

lldm1 1·90 5·60 4 DentSlght 1·38lldm2 1·00 5·80 2 EnamMid -

urdm1 2·00 6·25 3 EnamAdv 1·18lrdm2 1·00 5·60 2 EnamMid -

lrdm1 1·90 5·50 3 EnamAdv 0·46lrdi2 2·20 6·50 4 DentSlght 6·46lldi2 2·20 6·60 4 DentSlght 6·04lldc 1·60 7·80 0 None -

lrdi1 2·60 4·95 5 DentAdv 11·99MP La Chaise 13 65 3·2 lrdm2 1·00 5·75 3 EnamAdv 0·23

lrdm1 1·90 5·75 4 DentSlght 1·51lldm2 1·00 5·95 3 EnamAdv 0·27lldm1 1·90 5·40 5 DentAdv -

MP Molare 94 3·6 lldm2 1·40 6·40 1 EnamTrace -



MP Engis 66 3·6 lrdm1 2·30 5·20 4 DentSlght -

urdm1 2·40 5·90 4 DentSlght -

urdm2 1·30 5·75 3 EnamAdv -

lrdm2 1·40 6·15 3 EnamAdv -

MP La Chaise 14 67 4·3 lldm1 3·00 5·20 2 EnamMid -

lrdm2 2·10 6·15 2 EnamMid 1·23lrdi2 3·30 6·50 4 DentSlght 8·38lrdi1 3·70 4·35 5 DentAdv 27·78

MP Gibraltar 2 68 5·0 lldm1 3·70 5·15 4 DentSlght 9·45urdm2 2·70 5·80 3 EnamAdv 2·76lldm2 2·80 6·00 3 EnamAdv 2·57urdm1 3·80 5·75 4 DentSlght 13·32

MP Krapina 67 69 5·3 lldm1 4·00 4·95 5 DentAdv -

MP KrapMand A 70 5·4 lldm1 4·10 - 6 DentStrng 16·85lldc 3·80 5·70 6 DentStrng 15·36

MP Chatoneuf 2 71 5·7 uldi1 4·90 5·15 4 DentSlght 14·60uldi2 4·90 5·40 4 DentSlght 11·22urdi2 4·90 5·45 4 DentSlght 12·92urdi1 4·90 5·35 4 DentSlght 11·61uldc 4·20 6·30 4 DentSlght 7·38lrdi1 5·10 4·85 5 DentAdv 18·80lldm2 3·50 5·45 3 EnamAdv 1·27uldm1 4·50 5·40 4 DentSlght 9·06uldm2 3·40 5·50 3 EnamAdv 1·26lldm1 4·40 4·50 4 DentSlght 7·09lrdm1 4·40 4·90 3 EnamAdv 1·22urdm1 4·50 5·65 4 DentSlght 5·06urdc 4·20 6·80 4 DentSlght 3·62lrdm2 3·50 5·50 3 EnamAdv 1·57urdm2 3·40 5·35 3 EnamAdv 1·64

MP Krapina 11 72 6·3 lrdi2 5·30 5·70 5 DentAdv -

MP KrapMax B 73 6·5 urdi1 5·70 4·70 6 DentStrng 15·55urdm1 5·30 - 4 DentSlght 3·81urdi2 5·70 4·65 6 DentStrng 23·25urdc 5·00 6·45 4 DentSlght 3·68urdm2 4·20 - 3 EnamAdv 0·29


Appendix continued


uldm1 5·30 5·80 5 DentAdv 8·43uldm2 4·20 6·15 4 DentSlght 1·84urM1 0·10 8·10 1 EnamTrace -

ulM1 0·10 7·80 0 None -

uldc 5·00 6·30 4 DentSlght 5·80uldc 5·20 7·30 4 DentSlght 2·67ulM1 0·30 7·65 2 EnamMid -

llM1 0·40 6·30 2 EnamMid -

urdm1 5·50 5·35 5 DentAdv 13·75lrdm1 5·40 4·60 4 DentSlght 1·88lldm2 4·50 6·20 4 DentSlght 1·93lldm1 5·40 5·35 4 DentSlght 1·56lrdm2 4·50 5·65 4 DentSlght 2·47uldm2 4·40 5·90 3 EnamAdv 0·55uldm1 5·50 5·40 5 DentAdv 4·19urdm2 4·40 5·50 4 DentSlght 1·59lrI1 0·50 10·60 0 None -

llI1 0·50 10·65 0 None -

ulI1 "0·10 12·65 0 None -

urdc 5·20 6·70 4 DentSlght 3·12MP La Quina 18 75 7·0 uldm1 5·80 4·50 6 DentStrng 19·87

urdm2 4·70 4·65 6 DentStrng 13·50ulM1 0·60 6·75 1 EnamTrace -

urM1 0·60 6·35 2 EnamMid -

uldm2 4·70 4·00 5 DentAdv 9·04urdm1 5·80 4·90 7 EnamRing 38·26

MP Krapina 63 77 7·2 lldm2 5·00 7·25 4 DentSlght -

MP KrMdBMxA 79 7·8 lrI1 1·60 10·05 3 EnamAdv 2·04lrI2 0·80 11·75 1 EnamTrace -

lrdc 6·20 5·70 6 DentStrng 17·89urdi2 7·00 5·40 6 DentStrng 26·12uldm2 5·50 5·75 4 DentSlght 1·00urdm2 5·50 5·95 4 DentSlght -

uldm1 6·60 5·75 5 DentAdv 7·34llM1 1·50 6·45 2 EnamMid -

ulM1 1·40 7·35 2 EnamMid -

urM1 1·40 7·15 1 EnamTrace -

MP Le Fate 2 80 8·0 llM1 1·70 - 2 EnamMid -

MP Rene Simard 81 8·8 lrI1 2·60 9·10 2 EnamMid -

urI2 0·80 10·85 1 EnamTrace -

urI1 2·00 11·60 3 EnamAdv -

uldi2 8·00 5·05 5 DentAdv 19·13llI1 2·60 9·10 3 EnamAdv -

ulI1 2·00 12·35 2 EnamMid -

MP KrMdCMxC 82 10·4 ulM1 4·00 - 2 EnamMid -

urI2 2·40 11·35 0 None -

ulM2 "2·20 - 0 None -

llC 0·60 13·25 0 None -

llI2 3·40 11·00 4 DentSlght 5·48urI1 3·60 12·70 0 None -

uldm2 8·10 6·50 5 DentAdv 2·97lrdm2 8·20 6·25 5 DentAdv 8·65lrM2 "1·40 7·60 1 EnamTrace -

lldm2 8·20 - 5 DentAdv 10·96llI1 4·20 10·15 4 DentSlght 6·85ulC "1·00 14·50 0 None -

lrM1 4·10 8·10 3 EnamAdv -

MP Ehringsdorf4 84 12·2 lrC 2·40 - 0 None -

lrP1 2·20 - 0 None -





MP Krapina MdD 85 12·4 llP2 1·80 6·15 2 EnamMid -

llM1 6·10 6·10 3 EnamAdv 0·25llI2 5·40 9·75 4 DentSlght 11·19llP1 2·40 4·90 1 EnamTrace -

llM2 0·60 6·40 1 EnamTrace -

llC 2·60 11·15 4 DentSlght 2·06MP Malarnaud 86 13·2 lrM1 6·90 6·15 4 DentSlght 0·55MP Krapina MdE 87 13·5 llM1 7·20 6·95 3 EnamAdv 0·52

llM2 1·70 7·55 1 EnamTrace -

700 M. Skinner

Appendix continued


llP1 3·50 4·85 3 EnamAdv -

lrI1 7·30 9·35 5 DentAdv 7·27llC 3·70 12·50 3 EnamAdv 1·50llI1 7·30 9·05 5 DentAdv 9·79llP2 2·90 6·30 2 EnamMid -

llI2 6·50 9·90 5 DentAdv 7·18MP Puymoyen 1 89 16·8 ulM3 "2·20 6·35 0 None -

llP1 6·80 4·95 2 EnamMid -

llM1 10·50 - 4 DentSlght 1·26ulM1 10·40 6·30 4 DentSlght 0·62ulM2 4·20 7·50 2 EnamMid -

llP2 6·20 6·25 1 EnamTrace -

llM2 5·00 6·40 2 EnamMid -

MP Puymoyen 2 90 18·6 lrM2 6·80 7·00 2 EnamMid -

lrM3 "0·40 6·80 1 EnamTrace -

lrM1 12·30 6·55 4 DentSlght 1·90lrP1 8·60 5·10 2 EnamMid -

lrC 8·80 12·70 4 DentSlght 1·35MP La Quina 9 92 18·6 llM2 6·80 6·85 3 EnamAdv -

llM1 12·30 6·00 5 DentAdv 11·02llM3 "0·40 6·70 3 EnamAdv -

llP2 8·00 6·75 4 DentSlght 2·88llP1 8·60 5·00 4 DentSlght 3·93

*EUP: Early Upper Paleolithic; LUP: Late Upper Paleolithic; MP: Middle Paleolithic; ul: upper left; ur: upper right; ll: lower left; lr: lowerright; i: primary incisor; c: primary canine; m: primary molar; I: permanent incisor; C: permanent canine; M: permanent molar; 1–8: toothnumber.

Dental Wear in Immature Late Pleistocene European Hominines

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