Metric dental change in the European upper paleolithic and mesolithic

Metric Dental Change in the European Upper Paleolithic and Mesolithic

DAVID W. FRAYER Departmvnt ofilnthropoh~gy, Ihioersity of Knnsci,s, Luwrence, Kansas 6604.5

KEY WORDS Tooth size Dental reduction . Upper Paleolithic Mesolithic.

ABSTRACT Evolutionary trends for dental reduction are presented for Euro- pean Upper Paleolithic and Mesolithic samples. The analysis demonstrates that the greatest decrease in tooth size occurs between the two divisions of the Upper Pa- leolithic, while little and insignificant change characterizes the Late Upper Paleo- lithic/Mesolithic transition. Trends for tooth size over this period indicate that (1) human evolution does not stop with the appearance of “anatomically modern Homo sapiens,” (2) changes in tooth size fluctuate with increases in the efficiency and complexity of cultural systems, and (3) the Early Upper Paleolithic mmple should be considered transitional between Wurm 11 European Neanderthals and later Up- per Paleolithic and Mesolithic groups.

The late Pleistocene and Early Holocene prehistory of Europe, spanning the time between 35,000 and 6,000 R.P., has long in- terested archaeologists, particularly the evolution and succession of technological industries. Yet, the changes in human morphological and metric patterns over this same period have not commanded much attention by physical anthropologists. Even though the major innovations and improve- ments in cultural adaptations over the Up- per Paleolithic and Mesolithic must have had biological consequences, statements concerning this period, generally suggest that human evolution ceased with the appearance of “anatomically modern Homo sapiens” in the middle Wiirm interstadial (Brace and Mahler, ’71). Specimens dating to this period supposedly have “. , . little to distinguish them from modern Cextantl hominid populations” (Poirier, ’73: p. 1921, are “. . . not significantly different from . . . moderns humans” (Weiss and Mann, ’75: p. 215) or are “. . . modern in every respect” (Prideaux, ’73: p. 11). Only a few authors (Brace, ’64a, ’67; Brace and Mahler, ’71; Brose and Wolpoff, ’71; Bonnet, ’19; HrdliCka, ’30) have considered Upper Pa-

AM. J. PHYS. .&THROP., 46: 109-120

leolithic and Mesolithic specimens as representing something different from the contemporary Europeans.

It is ironic that a period associated with such rapid and significant cultural and technological changes, including the nearly complete substitution of blade for flake im- plements (Bordes, ’68; de Sonneville- Bordes, ’741, the parallel development of more varied, sophisticated and efficient stone and bone tool types (Semenov, ’64: Bordaz, ’701, the initial use and widespread incorporation of the spearthrower and bow and arrow (Semenov, ’64; Boule and Val- lois, ’57; Clark, ’521, and the improvement of cooking-and food-preparatory techniques, including the possible use of the earth oven (Brace, ’76; Perks, ’7.5) is simul- taneously characterized as one in which little or no biological evolution occixs. This paper attempts to demonstrate that the Upper- Paleolithic in Europe is a dynamic period in which significant metric changes in the size of teeth parallel and are associated with these cultural advances.

Previous studies of dental evolution in

’ Siippor-trrl i n part h y NSF Grmt GS-38067

109

110 DAVID W. FRAYRR

the Upper Paleolithic and Mesolithic have confirmed the existence of a trend for the overall reduction of tooth size, but have consistently underestimated the magnitude of dental change. Brabant and Twiesselmann (’64) and Brabant (’70) demonstrated that Upper Paleolithic tooth dimensions are larger than those from the Mesolithic and that both groups tend to have larger dimensions than more recent European populations. The samples utilized in their studies, however, represent only a partial inventory of the existing Up- per Paleolithic and Mesolithic specimens -consisting primarily of material from Predmost (Upper Paleolithic) and Rouffignac (Mesolithic?). Other studies by Brace (’62, ’67) and Brace and Mahler (’71) discuss Late Pleistocene and post-Pleisto- cene dental reduction in relation to technological and dietary changes, but use nearly the same sample as Brabant and Twiesselmann. Similarly, studies by Wol- poff (’71a) and Brose and Wolpoff (’71) have provided few, new additions. A recent analysis b y FlBchier and Verdene (’74) is also limited in its coverage of the Upper Paleolithic and Mesolithic, confined only to French and Portuguese material. The result of most of these studies has been the analysis of Upper Paleolithic and Mesolith- ic dental evolution without an adequate sample and the unavoidable tendency to group temporally and technologically diverse groups together within a single sample designated as “Upper Paleolithic” or “Mesolithic.”

Although gross overall changes have been observed when the samples com- prise “Upper Paleolithic” or “Mesolithic” categorizations, the micro-evolutionary changes occurring throughout this period or between the divisions of the Upper Pa- leolithic have not been examined in any detail. Since most of these studies are based on data culled from the literature, the results have been necessarily limited to a small number of published cases for the entire period and are subject to errors in- volving different measuring techniques, mis-identified teeth, and printing errors.

MATERIALS AKD METHODS

The Upper Paleolithic and Mesolithic samples used in this study include specimens from both Eastern and Western Europe. The cultural provenience, relative or absolute date, probable sex, and state of the remains are given in detail elsewhere (Frayer, ’76). Most of the specimens were measured by the author in various re- positories in Europe. A large portion of the French Mesolithic material, however, was unavailable at the time of my study and data for these specimens were generously supplied by Drs. Verdhe, Lefkvre and Flkchier, of the Institut de Pakontologie Humaine, Paris. Inter-observer error analysis among measurements made mutually by Verdene, Lefevre, Flechier, and myself indicates good concordance with no significant differences as shown by student’s t tests on means. Data for the Greek Meso- lithic material from Franchthi Cave were supplied by Angel (personal comment). A few other specimens, particularly those for the Upper Paleolithic, were extracted from the literature. Since all of the Pred- most material was destroyed at the end of World War 11, measurements by Matiegka (’34) and HrdliCka (’24) had to be used. Ob- vious printing errors in Matiegka’s appen- dix were not included. Similarly, since a few other specimens have been lost, mis- placed or destroyed over the years, published measurements were consulted. These included Combe-Capelle (Klatsch and Hauser, ’091, some specimens from MladeC (Szombathy, ’251, and all material from Rochette (Elmer, ’14). Dental material from Arcy-sur-Cure was taken from Leroi-Gourhan (’58).

The two measurements taken were maximum mesial-distal length as defmed and used by HrdliEka (’24) and maximum buc- cal (labial) -lingual breadth, taken perpen- dicular to the length. Intra-observer mea- surement error was less than 2% for both dimensions. Nearly all measurements were taken on original specimens, unless original specimens were destroyed or otherwise unavailable. Measurements from original

UPPER P.4LEOLITIIIC/MESOLITI1IC DENTAL CH 4UGE 111

plaster casts included only two specimens (Cheddar and Dolni Vektonice 1111, each checked against published cranial or dental measurements. Descriptive statistics reported in the tables represent left, right or averages of the left and right sides, when both antimeres were present. For specimens with a great deal of occlusal wear, only the breadth dimension was used since interproximal and occlusal wear affects the true length dimension of the tooth (Johan- son, '74).

For comparative purposes, an index of relative change is utilized which expresses the amount of reduction (or increase) from one period to the next. This percent difference (% diff) was calculated as the difference between the earlier and later group mean ( x 1001 divided by the mean of the earlier group. It represents the relative amount of reduction between two periods and allows for comparison between teeth of different dimensions. Also in each of the tables, two separate tests for statistically significant differences in means are given. Since the two Upper Paleolithic samples tended to show more variation than the Mesolithic group, the student's t test was of limited use due to concommitant significant differences in the variance as measured b y the F-test. For these cases, the Mann-Whitney U test, a distribution test less affected by differences in sample variances (Conover, ,711, was used. In each test, statistical significance is reported at the .05 level, even though many cases represent a higher level of significance.

The Upper Paleolithic sample is divided into early and late components. The Early Upper Paleolithic sample (EUP) consists of all material from Eastern and Western Europe dating to Lower Perigordian, Aurignacian, or Szeletian contexts. Late Upper Paleolithic (LUP) specimens were defined as originating from Upper Peri- gordian, Gravettian, Proto-magdalenian, Solutrean, Magdalenian and other Late Paleolithic industries in Eastern, Western and Southern Europe. The Mesolithic sample includes all material dated to the Early Holocene, encompassing all of Europe. Ap-

proximate dates and timespan for the Early Upper Paleolithic are 34,000-26,000 H.P., the Late Upper Paleolithic 26,000-10,500 B.P., and the Mesolithic 10,500-6,000 H.P. Radiocarbon dates for these three samples are based on a compilation of published figures presented in Oakley et al. ('711, Movius ('721, Riquet ('70), Delihrias and Evin ('73, and others.

RESULTS

Basic descriptive statistics for the Early Upper Paleolithic (EUP) Late Upper Pa- leolithic (LUP) and Mesolithic samples are given in tables 1, 2, and 3. Inspection of these tables indicates that the Early Upper Paleolithic sample has consistently larger tooth dimensions than either the Late Up- per Paleolithic or Mesolithic groups, while the Late Upper Paleolithic and Mesolithic samples more closely resemble each other for length, breadth and area measures.

In table 4, the differences between the EUP and LUP are greater than those between the LUP and Mesolithic samples, both in the amount of percent difference and in the number of statistically significant differences. In the mandibular anterior teeth, lengths and breadths show a substantial reduction in size over the Up- per Paleolithic and no change or a slight increase from the Late Upper Paleolithic to the Mesolithic. In the EUP-LUP comparison, teeth posterior to the canine show less overall reduction than those located more anteriorly, with fewer cases of statistical significance. Nevertheless, within the Upper Paleolithic, there is a consistent trend for the reduction of tooth size for every mandibular length and breadth dimension.

The pattern of this reduction and the difference in magnitude between EUP- LUP and LUP-Meso are well-illustrated when tooth areas are compared. The mandibular canine shows a 20% reduction within the Upper Paleolithic, while the same tooth shows minor and insignificant change between the LUP and Mesolithic. Similarly, all posterior tooth areas show a consistent reduction from the EUP to the

112 DAVID W. FRAYEn

TABLE 1

Mandibular dental dimensions (length arid hreadth) and range (r) for the Early Upper Paleolithic (EUP), Late Upper Paleolithic (LUP), and Mesolithic samples (MESO)

EUP LUP MESO (nl s.d. X in) s.d. x (11) S.d.

I, Lt r Hr I‘

I, Lt r Br r

c Lt r 5r r

P, Lt r Br r

P, Lt r Rr r

M, Lt r Br r

r Br r

M, 1.t

M, Lt r Br r

5.8112) 0.67

6.4(14) 0.44 5.9- 7.1

6.415) 0.68 5.0- 7.3

7.0(18) 0.60 6.0- 8.5

7.6(11) 0.45 7.0- 8.3

9.0(13) 0.62

7.3(12) 0.65 6.0- 8.1

8.5(13) 0.54 7.8- 9.3

7.4(10) 0.51 6.6- 8.5

8.7( 1 1) 0.58

11.6(26) 0.87 10.0-13.0

11.0(27) 0.61 10.0-12.0

11.3(22) 1.00 9.5-12.8

10.8(22) 0.82 9.8-12.0

11.1(12) 1.24

10.7(12) 0.95

4.6- 6.6

8.0-10.0

8.0-10.1

9.5-13.0

9.3-12.4

5.2(101 0.55 4.6- 6.1

6.1(15) 0.54 5 . 3 - 7.1

5.6(13) 0.55 4.5- 6.5

6.5(19) 0.56 5.7- 7.7

6.8(18) 0.62 6.0- 8.4

8.0(20) 0.61 6.9- 9.5

7.0(22) 0.57 6.0- 8.0

8.1(23) 0.77 6.1- 9.7

6.9(21) 0.56 6.0- 8.1

8.3(20) 0.46 7.5- 9.2

11.1(30) 0.71 9.4- 12.1

10.9(35) 0.56 9.2-1 1.9

10.9(33) 0.70 9.8-12.1

10.7(33) 0.61 9.4-12.0

10.6(21) 0.76 9.2-12.2

10.4(19) 0.93 8.7-12.8

5.3(67) 0.39 4.5- 6.2

6.1(71) 0.38 4.6- 6.9

5.9(82) 0.46 4.7- 6.9

6.5(91) 0.38 5 . 5 - 7.4

6.9(87) 0.49 5.2- 8.1

7.9(92) 0.58 6.2- 9.1

7.0(96) 0.46 5.1- 8.2

7.9(97) 0.42 7.0- 9.0

6.9(102) 0.46 5.8- 8.1

8.4(103) 0.48 6.9- 9.4

11.3(116) 0.67 9.7-12.9

11.0(118) 0.52

10.7(103) 0.65 9.3- 12.3

10.7(103) 0.59

10.3(81) 0.61

10.3(31) 0.61

9.2- 12.5

8.8-1 1.8

9.3-12.0

8.8- 1 1.9

All ~nca\urernmts arc in millimeterr

LUP, representing a decrease in the summed posterior tooth area of 7.1%. The same teeth in the LUP-Meso comparison show an overall reduction of only 0.2% with some teeth (P, and M,) increasing in the Mesolithic. Figures for the EUP-Meso comparison, indicate that even though some tooth dimensions increase in the Mesolithic, the overall trend for all tooth dimensions is for reduction in sizc. Tooth size expansion in the Mesolithic never sur- passes the size found in the EUP sample.

Trends for maxillary tooth dimensions essentially parallel those in the mandible, with the Early Upper Paleolithic sample in

all but one case (P3 breadth) exhibiting the largest average tooth size. The greatest amount of maxillary dental reduction occurs within the Upper Paleolithic, not between the Late Upper Paleolithic and Mesolithic. Even though the same trend for overall reduction is apparent in the maxilla, there are some interesting differences in the pattern of tooth size changes. For example, whereas the mandibular anterior teeth show a great deal of reduction in the EUP-LUP comparison, in the maxilla these same teeth reduce much less. In the Upper Paleolithic, mandibular canine length and breadth reduced 10.5% and 11.5% (both

UPPER PALEOLITIIIC/?VlESOLIT€Il~~ UEYl AL CHA\GE 113

T4ULE 2

Maxillary dental dzmenstons (length and breadth) und runge (r) f o r Early Upper Palvolzthic (EUP), Late Upper Paleolithic (LUP), and Mesolithic (XIESO) samples

EI'P S 111) \.d

I' Lt r Br r

I' Lt r Br r

c Lt r Rr r

P3 Lt

Rr r

F Lt r Br r

M' Lt r Br

M2 Lt r Rr r

?# Lt r Br r

1'

r

9.3(10) 0.51 8.5-10.2

7.5(13) 0.28

7.6(6) 0.86 6.4- 8.8

6.8(10) 0.52 6.0- 7.4

8.0(12) 0.62 7.1- 9 1

9.0(12) 0.91 7.8-10.8

7.2(15) 0.59

9.6(15) 0.59

7.0(14) 0.57

9.6(16) 0.75 8.5-1 1.2

10.'7(24) 0.79 9.1- 12.0

12.3(24) 0.73 11.0-14.0

10.6(20) 0.80 8.8-11.8

12.3(20) 0.95

9.5(12) 1.00

11.4(14) 1.15

7.1- 8.0

6.0- 8.0

8.7-10 6

6.1- 7.9

10.8-13.8

7.7-11.0

9.2-13.1

8,7112) 0.63 7.1- 9.3

7.5(16) 0.52 6.8- 8.6

7 . lN ) 0.52 6.1- 7.7

6.6(15) 0.62 5.8- 7.6

8.0(14) 0.60 7.3- 9.5

8.8(15) 0.89 7.8-10.5

7.0(18) 0.76 6.0- 8.7

9.7(19) 0.7.5 8.2-10.9

6.8114) 0.71

9.6(15) 0.55 9.0-10.8

10.4124) 0.49 9.2-11.0

11.8(24) 0.58 10.6- 12.8 9.7(21) 0.63

8.2-11.1 12.0(21) 0.80

9.0(14) 0.83

11.4(15) 0.75

5.9- 8.1

10.3-13.6

7.6-10.5

10.2-12.8

9.2(49) 0.64 8.0-10.5

7.4(56) 0.46 6.3- 8.4

7.0(46) 0.60 5.2- 8.2

6.5(61) 0.48 5.4- 7.2

7.8(68) 0.lil 6.5- 8.9

8.7(74) 0.60 7.4-10.4

6.9(71) 0.43 5.6- 7.9

9.4(75) 0.61 6.5-10.4

6.61783 0.41 5.8- 7.8

Y.6(83) 0.54 8.0-10.8

10.4(Y9) 0.58 9.0-11s

11.9(100) 0.57 10.1-13.1

9.7193) 0.69 7.4-1 1.5

11.9(93) 0.73 9.8-13.5

8.8164) 0.69 7.4-10.5

11.4(63) 0.80 9.6- 13.1

All rrirasurernent5 itre in niilli1neter5.

statistically significant), while the maxillary counterparts show no change for length and only a 2.2% reduction in breadth. Viewed with respect to areas, over the Up- per Paleolithic the mandibular canines decrease in size at a rate almost five times that of the maxillary canine. Premolar dimensions also reduce more in the mandible than in the maxilla which may be the result of a canine field effect (Garn et al., '67; Turner, '69). At the same time, the first two molars exhibit a greater amount of reduction in the maxilla, while the third molar exhibits slightly more reduction in the mandible. The overall reduction of the

posterior teeth (EP3 -M3) is about the samt. in both jaws (7.1% in the mandible; 6.6% in the maxilla). Even though the specific teeth responsible for the reduction in summed posterior area are different in the two jaws, the amount of occlusal area avail- able for mastication decreases harmonious- ly in the mandible and maxilla.

The pattern of change in the mandibular and maxillary molars during the Upper Pa- leolithic is similar to that recently observed by LeRlanc and Black ('741, in that two of the three maxillary molars decrease at a faster rate than the mandibular molars. Inclusion of the premolars as a functioning

114 DAVID W. FHhYER

T4BJ,E 3

Mandibular and maxillary tooth areas (C-M3) and range (r) for Early Upper Paleolithic (EUP), Late Lpper Paleolithic (LUP), and Mesolithic (MESO) samples

MESO h (n) s.d

68.5(11) 7.4 58.8- 80.0

62.1(12) 8.4 48.0- 73.7 65.5(10) 8.6 52.8- 85.9

127.4(26) 14.9 101 .O- 14Cj.6 122.3(21) 19.0

95.0-153.6 120.0(12) 23.5

88.4-152.5

497.3

72.7(12) 12.5

68.7(15) 9.3 54.0- 84.0 68.3(14) 10.2 Fj4.3- 88.4

132.1(23) 16.2 102.1-165.2 130.5(20) 18.0

95.0-159.3 108.6(12) 20.7

78.2-144.1

55.4- 98.3

508.2

54.4(18) 8.7 43.9- 75.6

56.8(22) 9.5 38.4- 75.7

57.1(20) 6.7 48.0- 73.7

120.1(33) 12.2 92.1-144.0

ll6.4(33) 13.2 97.8-141.6

111.7(19) 17.8 80.0-156.2

462. I

69.6(1.4) 11.1 57.7- 93.5 68.6(18) 11.4 52.5- 88.7 65 6(14 10.4 53.1- 87.4

121.8(24) 8.3 109.2-138.2

116.0(21) 13.8

102.6(13) 15.4 87.6-142.8

83.6-127.1

474.6

35.1(87) 7.0 38.ij- 72.1

55.3(96) 5.4 35.7- 72.2

58.4(102) 6.2 41.9- 7 3 5

124.1(116) 11.6 92.9-153.5

114.7(101) 11.9 81.8-140.4

108.8(79) 11.2 81.8-135 6

461.3

68.0(67) 7.8 53.3- 87.1

64.9(69) 7.1

64.0(78) 5.9 50.7- 77.2

124.2(98) 11.2

115.9(92) 12.1

100.7(61) 11.0

43.6- 80.6

9 1.9- 146.9

85.3-193.9

75.0-134.2

469.7

Al l i ~ w ~ i ~ ~ u r ~ ~ i e ~ i t ~ in millirnetrr! ' Sum of group mcwis.

part of the total occlusal area (Wolpoff, '71b1, however, eliminates the paradox in- troduced by LeRlanc and Black. Although they do not include data on premolar changes, these data show that in Europe both jaws show evidence of a similar amount of reduction in summed posterior area, but that different teeth are resporisi- ble for the decrease in total masticatory area. As Wolpoff ('71b) suggested, the unit of selection for the teeth posterior to the canine is probably not any individual tooth, but the summed areas of the two premolars and three molars.

The transition from the Late Upper

Paleolithic to the Mesolithic exhibits no comparable amount of tooth size reduction as demonstrated for the change within the Upper Paleolithic (tables 1, 2, 3) . In mandibular lengths and breadths, five dimensions decrease between 2.5% and 0.9%, five show no change, and six increase between 5.4% and 0.9% (table 4). Only one represents a significant change (12 length) and none are greater than those occurring in the Upper Paleolithic comparison. For mandibular tooth areas, three Mesolithic averages are greater than the LUP, while three are less. Whereas the mandibular summed posterior areas reduced 7.1% in

CPPER PALEOLITIIlC/MESOLITHIC DENTAL CH 4KCE 11.5

TABLE 4

Percentage change (9 difference) in tooth dimensions from Early Upper Paleolithic (EUP) through Late Upper Paleolithic (LUP) to Mesolithic (MESO)

Mmdildc %fdXlll.i

EUP LUP IXP-hIESO EL'P-MESO ECP-LCP Lt'P MESO FUP h lEW

% dltl" % dlff % dlff %dlH c/r dlft '% diff ___ ~- ~ _ _ _ _ _ _

I1 Lt Br

12 Lt Br

c Lt Br

P3 Lt Br

P4 Lt Br

M1 Lt Br

M2 Lt Br

M3 Lt Hr

Areas C P3 P4 hl 1 M2 M3 tP3-M3

- 10.3 1 2 - 4.7 1.2

- 12.5 ',* - 7.1 2

- 10.5 - 11.1 - 4.1 - 4.7 - 6.8 - 4.6 I , 2

- 4.3 1 J

- 0.9 - 3.52 - 0.9 - 4.5 - 2.8

- 20.6 I,' - 8.5 - 12.8 '7'

- 5.7 1 2

- 4.8 - 6.9 - 7.1

+1.9 - 8.62 0 - 4.7 1 2

+5.4 - 7.82 0 - 7.1

+I .5 - 9.2 1x2

- 1.3 - 12.2 112

0 - 4.12 - 2.5 - 7.1 l:2

0 - 6.8 ' 5 '

+1.2 - 3.4' +1.8 - 2.6 +0.9 0 - 1.8 - 5.32

0 - 0.9 - 0.9 - 5.4 - 1.0 - 3.7

+1.3 - 19.6 ' 2 '

- 2.6 - 11.02 f 2 . 3 - 10.82 +3.3 1 - 2.6 - 1.5 - 6.2 - 2.6 - 9.3 - 0.2 - 7.2

- 6.5 1

0 - 6.6 - 2.9

0 - 2.2 - 2.8 + 1.0 - 2.9

0 - 2.82 - 4.1 112

- 2.4 - 5.3

0

- 4.3 - 0.1 - 4.0 - 7.8 1.2

- 11.1 1,: - 5.52 - 6.6

- 8.5 lJ

+5.7 ' - 1.1 - 1.3 - 1.3 - 1.4 - 7.9 I - 1.5 - 4.4 - 2.5 - 2.5 - 1.1 - 3.3 - 1.4' - 4.2 - 3.1 - 2.1 - 2.9 - 5.7 2

0 0 0 - 2.82

+0.8 - 3.3' 0 - 8.5 1,'

- 0.8 - 3.3 ~~ 2.2 - 7.4 2

0 0

- 2.3 - 6.5 - 5.4 - 5.52 - 2.4 - 4.3 +2.0 - 6.02 - 0.8 - 11.22 - 1.9 - 7.3 - 1.0 - 7.6

~ denotes reduction; + increase. I Significant difference in means (one-tailed student's t - 0.05 level). 2 Significant difference in means (one-tailed bfmn Whitney U. - 0.05 level).

the Upper Paleolithic, the maxillary total occlusal area reduces only 0.2% from the Late Upper Paleolithic to the Mesolithic.

In the maxilla 10 of the 16 length and breadth dimensions are larger in the Late Upper Paleolithic sample, but again, the amount of reduction is much less than the differences in the two Upper Paleolithic samples. The tooth dimension representing the greatest percent reduction between the LUP and Mesolithic is P3 breadth (3.1%) which is the only case where a statistically significant reduction has occurred. The Mesolithic sample has two maxillary dimensions larger than the LUP, but only one (1' length) is significantly larger. Maxillary tooth areas, except for MI,

are all smaller in the Mesolithic, representing percent reductions from the LUP between 0.8% and 5.4%. The overall reduction in maxillary summed posterior areas amounts to only 196, compared to a 6.6% reduction from the Early to Late Upper Paleolithic summed posterior areas.

Table 4 also gives the percentage change in tooth size from the Early Upper Paleo- lithic to the Mesolithic, summarizing the magnitude of dental change in Europe over approximately 25,000 years. There are no cases where the Mesolithic has an average tooth size larger than the EUP, and only three cases (MI p4, and M3 breadths) where the dimensions are identical. Anterior teeth show a greater rate of reduction in

116 DAVID W. FKAYEK

the mandible than in the maxilla, particularly the central incisor and canine. Posterior tooth dimensions also reduce significantly through time with the premolars reducing more in the mandible. Molars, however, show more reduction in the maxilla, although when summed posterior areas are considered, the amount of reduction in both jaws is about the same.

DISCUSSION

The pattern of tooth size changes in the three samples from the Late Pleistocene and Early Holocene suggests a period of substantial reduction in the Upper Paleo- lithic with only niinor changes in tooth size from the Late Upper Paleolithic to Meso- lithic. In addition, there is a clear pattern in the posterior teeth of a greater amount of reduction in lengths as opposed to breadths. This observation would not seem to be due to the environmental effects of interproximal wear, since heavily worn teeth were not included in the samples, but may relate to reduction in jaw length.

To further illustrate the extent of dental reduction in the late Pleistocene and post- Pleistocene and the magnitude of differences in tooth size between Upper Paleo- lithic specimens and more recent groups, comparison of the EUP and hlerovingean (Vexin) sample means are given in table 5. The Merovingean data are from Menard ('75) and represent tooth measurements for 104-265 individuals. Tooth size for this group is similar to the Medieval Danes (Lunt, '69) and the Belgian Medieval site of Coxyde (Brabant and Twiesselmann, '67). Sample means for all three Medieval groups are slightly smaller than found in most extant European populations (La- velle, '68; Alvesalo, '701, which may be re- lated to secular changes in body size over the recent historic period.

The data presented in table 5 clearly demonstrate the magnitude of dental change in Europe over the past 30,000 years. The Early Upper Paleolithic sample has tooth dimensions larger than the Vexin group -generally by more than 10% in each length, breadth or area measure. In

"t: l'! t -10 3, k 9 9

a m m

the mandible the teeth showing the most marked decrease are the incisors and canine (the mandibular canine area has reduced nearly 30961, while in the maxilla, with some exceptions, the molars show a more pronounced reduction. In both jaws summed posterior areas reduce at approximately the same magnitude, and the reduction of the third premolar closely par- allels the reduction of the canine. These data demonstrate the scale of tooth size reduction from the late Pleistocene to the modern era indicating the significant and substantial changes in tooth size which have occurred after the appearance of “anatomically modern Homo sapiens.”

Dental changes presented here do not deviate substantially from the results of previous studies in which only “Upper Pa- leolithic” or “Mesolithic” categorizations were used. They do indicate, however, that the main reason Upper Paleolithic specimens have consistently larger dimensions than found in the Mesolithic relates more to large tooth size in the Early Upper Paleolithic division, rather than large tooth dimensions in all Upper Paleolithic periods. Changes in tooth size over the Upper Paleolithic and Mesolithic, then, are not evenly graded, but rather are characterized by a period of accelerated dental reduction, followed by a period of only minor change.

Metric changes in tooth size and different rates of reduction between the three periods are closely correlated with the evolution of technological and cidtiiral systems. Technologically the EUP and LUP show major qualitative differences, while differences in tool types, functions and cultural behavior between the LUP and Mesolithic are less marked. For example, the earliest Upper Paleolithic traditions in Eastern Europe (Aurignacian and Szele- tian) and Western Europe (Chfitelperon- nean) bear many affinities to the earlier in- digenous Mousterian industries (Hahn, ’70, ’73; LaPlace, ’70; Bordes, ’68; Delporte, ’70). The earliest Upper Paleolithic phase represents primarily a shift in the frequen- cy of production of blade tools (Pradel, ’66)

and probably the initial widespread use of handled utensils (Movius, ’69; Leroi- Gourhan, ’64). Burins also are more pre- dominant in Early Upper Paleolithic industries than in the underlying Mousterian complexes which may indicate a greater reliance on non-lithic tools (David, ’73). The major technological and cultural features cited as the hallmarks of the Upper Paleolithic period, however, do not lie- come widespread until the Late Upper Pa- leolithic. Undoubtedly, some of these forms were present in the Early Upper Pa- leolithic division, but most of these tcch- nological and cultural features had either yet to appear or occurred in low enough frequencies that they are not readily identified in Early Upper Paleolithic contexts. Howevcr, by the beginning of the Late Up- per Paleolithic (Upper Perigordian in Western Europe; Gravettian or Pavlovian in Eastern Europe) the cultural markers of the Upper Paleolithic appear with morc and more regularity. These include the first evidence of the spearthrower (Solu- trean: Mellars, ’731, the development of composite stone and osseous tools (Middle Magdalcnian: Semenov, ‘64; de Sonneville- Rordes, ’60, ’631, the initial use of the bow and arrow (Late Magdalenian: Rust, ’37, ’431, the first appearance of eyed needles (Solutrean: Bordes, ’681, thermal pretreat- ment of lithic materials (Solutrean: Rordes, ’691, development of the pressure flaking technique (Solutrean: Bordes, ’681, and the common use of stone-boiling (Upper Peri- gordian: Movius, ’65, ’66) or possibly earth ovens (Upper Perigordian:J: Brace, ’76; Eastern Gravettian?: Klima, ’63: Late Mag- daleriian: Leroi-Gourhan and Brkzillon, ’66, ’72; Perks, ’75). Each of these represent a significant advance in the level of technological efficiency.

Most of these same features characterize the Early Mesolithic (hilian, epi-Gravett- ian, etc.) and continue into Late Mesolithic contexts. The major differences between Late Upper Paleolithic and Mesolithic techno-cultural systcms relate more to the climatic and faimal shift, than to the in- troduction or radically new and more effi-

118 DAVID W.

cient tool types (Bordes, '68; Escalon de Fonton, '74). The period of the greatest change in techno-cultural systems (EUP- LUP transition), then, is also the period of greatest reduction in tooth size.

These inter-relationships between cultural and biological evolution have at least two implications. (1) Since the change in tooth size seems to be correlated with the increasing specialization of techno-cultural systems, it is unlikely that the mechanism for the reduction is the Probable Mutation Effect (PME) (Brace, '63, '64b; Wolpoff, '75). If this principle were in operation, Mesolithic dentitions should be smaller than the LUP group, since mutations for smaller tooth size would continue to be in- corporated in these later populations. Also? with PME, the rate of change for dental dimensions should accelerate through time as tools take on functions previously ac- complished by the dentition. With sclec- tion, the rate and magnitude of change should be more closely correlated with the specific technological changes. Therefore, reductions due to selection will fluctuate through time in accordance with the technological changes, rather than accelerate. Also, variation in tooth size, measured by the variance and the coefficient of variation, decreases through the EUP-LUP- Mesolithic sequence (Frayer, '761, exactly opposite of what is expected if PME were the agent of change (Wolpoff, '69).

(2) Analysis of the relationship between Upper Paleolithic descendants and their possible Middle Paleolithic ancestors should concentrate on the Early Upper Pa- leolithic sample as stressed by Brace ('64a) more than ten years ago. Nearly every recent study of the inter-relationship between Neanderthals and Modern forms has either lumped together all the Upper Pa- leolithic specimens (including as Howells C'701 states a "hodge-podge" of more recent material, [see Stringer, '741) or has compared the Wiirm Neanderthals with a recent Homo sapiens group (Howells, '67; '74a,b) in the tradition of Boule ('131, Bode and Vallois ('571, Morant ('27) and Hooton ('47). Since the Early Upper Paleolithic

FRAYER

group represents (at least for the dentition) a more robust morphotype, it may be more appropriate to consider this group as a transitional phase between the European Wiirm Neanderthals and the Late Upper Paleolithic sample. It is interesting that the only multivariate study which has main- tained the possibility of a close relationship between European Mousterian and Upper Paleolithic types, only considered specimens dated to the Early Upper Paleolithic in the analysis (Bilsborough, '72). Loading the Early Upper Paleolithic sample with a series of Late Upper Paleolithic and more recent specimens only serves to confuse the issue of possible Neanderthal ancestry.

ACKNOWLEDGMENTS

Funds for museum fieldwork in Europe were provided by an NSF Doctoral Disser- tation Research Grant (GS-38067) and a University of Michigan Graduate School stipend. It is impossible to acknowledge all the European scholars who granted me permission to examine the Upper Paleo- lithic and Mesolithic skeletal material under their care. These are listed elsewhere (Frayer, '76). Drs. Verdene, Flkchier, and Lefkvre, however, deserve special mention for graciously allowing me access to their measurements. I would also like to thank D. O'Rourke, who read and critically commented on the manuscript and J. Perry who typed it.

LITERATURE CITED Alvesalo, I.. 1970 The Influence of Sex-Chromo-

some Genes on Tooth Size in Man. Academic! Dis- sertation, University of Turku, Finland.

Bilsborough, A. 1972 Cranial morphology of Ne- anderthal man. Nature, 237: 351-352.

Bonnet, R. 1919 Die Skelett. In: Der Diluviale Menschenfund von Oberkassel bei Bonn. M. Ver- worn, R. Bonnet, and G. Steinmann, eds. J. F. Berg- mann Verlag, Weisbaden, pp. 11-185.

Bordaz, J. 1970 Tools of the Old and New Stone Age. Natural History Press, New York.

Bordes, F. 1968 The Old Stone Age. McGraw-Hill, New York.

1969 Traitement thermique du silex ail So- IutrCen. Bull. Soc. PrPhist. Fr., 66: 197.

Bode, M. 1913 L'Homme Fossile de La Chapelle- aux-Saints. Mason, Paris.

Bode, M., and H. V. Vallois 1957 Fossil Men. Dryden Press, New York.

UPPEH PALE(>LITHIC/~~ESOI,ITHlC DENTAL CHAhGE 119

Brabant, H. 1970 La denture humaine as Paleo- lithiqrie superieur d’Europe. In: L’Homme de Cro- Magnon. G. Camps and G . Olivier, eds. Arts et Metiers Graphiques, Paris, pp. 99-120.

Brabant, H., and F. Twiesselmann 1964 Observations sur l’dvolution de la denture permanente humaine en Europe occidentale. Bull. Group. Int. Rech. Sc. Stom., 7: 11 -84. - 1967 Nouvelles observations sur la denture

d’une population ancienne d g e franc de Coxyde, Belgique. Bull. Group Int. Rech. Sc. Stom., 10;

Brace, C. L. 1962 Cultural factors in the evolution of the human dentition. In: Culture and the Evolu- tion of Man. M. F. A. Montague, ed. Oxford Univer- sity Press, New York, pp. 343-354.

1963 Structural reduction in evolution. Am. Nat., 97: 39-49.

1964a The fate of classic Neanderthals: a consideration of hominid catastrophism. Curr. Anthrop., 5: 3-43.

1964b The probable mutation effect. Am. Nat., 98: 453-455.

1967 Environment, tooth form, arid size in the Pleistocene. J. Dent. Res., 46: 809-816.

1976 Tooth reduction in the Orient. Asian Perspect, in press.

Brace, C . L., and P. E. Mahler 1971 Post-Pleistocene changes in the human dentition. Am. J. Phys. Anthrop., 34: 191-203.

Brose, D. S., and M. H. Wolpoff 1972 Early Upper Paleolithic man and Late Upper Paleolithic tools. Am. Anthrop., 7,3; 1156-1194.

Clark, J. G . I). 1952 Prehistoric Europe. Stanford University Press, Stanford.

Conover, W. J. 1971 Practical Nonparametric Statistics. John Wiley, New York.

David, N. 1973 On the upper Paleolithic society, ecology, and technological change: the Noaillian case. In: The Explanation of Cultural Change. C. Renfrew, ed. University of Pittsburgh Press, Pitts- burgh, pp. 277-330.

Delibrias, G., and J. Evin 1975 Sommaire des data- tions 14C concernant la prehistoire en France. Bull. Soc. Prehist. Fr., 72: 93-96.

Dclporte, El. 1970 Le passage du Moustkrien au Palblithique supkrieur. In: LHomme de Cro-Mag- non. G. Camps and G. Olivier, eds. Arts et Metiers Graphiques, Paris, pp. 129-140.

Elmer, F. W. 1914 Die Zahne von La Rochette. Archiv fiir Anthrop., 41: 127-129.

Escalon de Fonton, M. 1974 From the end of the Ice Age to the first agriculturalists: 10,000-4,000 B.C. In: France Before the Romans. S. Piggott and G. Daniels, eds. Thames and Hudson, London, pp.

Flkhier, J-P., and J. Verdene 1974 Quelques con- siderations sur la denture des MCsolithiques franpis. Bull. Mkm. Soc. dAnthrop. (Paris), 2: (skrie XIII); 237-269.

Frayer, D. W. 1976 Evolutionary Dental Changes in Upper Paleolithic and Mesolithic Human Popula-

5-180.

61-101.

tions. Ph.D. Dissertation, Univ. of Michigan. Univer- sity Microfilms, Ann Arbor.

Garn, S. M., D. R. Swindler and R. S. Kerewsky 1966 Canine “field” in sexual dimorphism o f tooth size. Nature, 212: 1501-1502.

Hahn, J. 1970 Recherches sur I’Aurignacien en Europe centrale et orientale. L’hth. , 74: 195-220.

Das Aurignacien in Mittel- und Ost- europa. Acta Praehist. et Arch., 3: 77-107.

Hooton, E. A. 1947 Up From the Ape. Revised ed. MacMillan, New York.

Howells, W. W. 1967 Mankind in the Making. Doiihleday, Garden City, N.J.

1970 Mount Carmel man. morphological relationships. Proc. 8th Int. Cong. Anthrop. and Ethno. Sci., Tokyo and Kyoto, I: 269-272.

1974a Neanderthals: names, hypotheses, and scientific method. Am. Anthrop., 76; 24-38.

1974b L’Homine de Neanderthal. La Re- cherche., 5: 634-643.

Hrdlieka, A. 1924 New data on the teeth of early man and certain European anthropoid apes. Am. J. Phys. Anthrop., 7: 109-137.

1930 The skeletal remains of early man. Smithsonian Misc. Coll., whole volume: 83.

Johanson, D. C. 1974 Some metric aspects of the permanent and deciduous dentition of the Pygmy Chimpanzee (Pun puniscus). Am. J. Phys. Anthrop.,

Klaatsch, H., and 0. Hauser 1909 Homo aurignacen- s i s hauseri. Prahist. Zeit. 1: 273-338.

Klima, €3. 1963 Dolni Vestonice. CeskoslovmskC Akademie Ved, Praha.

LaPlace, G . 1970 Le niveaux aurignaciens et l’hypothese du synthktotype. In: L’Homme de Cro- Magnon. G . Camps and G. Olivier, eds. Arts et Metiers Graphiques, Paris, pp. 141-164.

Lavelle, C. L. B. 1968 Anglo-Saxon and !vlodcrn British teeth. J. Dent. Res., 47. 811-815.

LeBlanr, S. A,, and B. Black 1974 A long term trend in tooth size in the Eastern Mediterranean. Am. J. Phys. Anthrop., 41; 417-422.

Leroi-Courhan, A. 1958 Etude des resteb humains fossiles prouvenant des Grottes d’Arcy-sur-Cure. Ann. Pakont. (Vertebres), 44: 87-148.

1964 Chronologie des Grottes d’Arcy-sur- Cure (Yonne). Gallia-Prehist., 8: 1-64.

Leroi-Gourhan, A,, and bl. Brezillon 1966 Lhabitation magdalenienne no. 1 de Pincevent pres Montcreau (Seine-et -Marne). Gallia-Prkhist . , 9;

Fouilles de Pincevent. Essai d’analyse ethnographique d’un habitat magdalknien (la Sec- tion 36). Gallia-Prkhist. (Suppl. 71, whole volume.

Lunt, D. A. 1969 An ondontometric study of Mediaeval Danes. Acta Odont. Scand. 27 (Suppl. 55), whole volume.

Matiegka, J, 1934 Homo Pfedmostensis: Fosilni Clovek z Pfedmosti na Morave. Publ. Czech. Acad. Sci. Art., pp. 105-145. (French summaryl.

Mellars, P. 1973 The character of the middle-upper Paleolithic transition in southwest France. In:

1973

41: 39-48.

266-385. 1972

120 DAVID w.

The Explanation of Cultural Change. C. Renfrew, c:d. Uniwrsity of Pittsburgh Press, Pittsburgh, pp.

Menard, J . 1975 L'evolution des dents des Franyais. Bull. et Mkm. Soc. d 'hthrop. (Paris), Zskrie XII1): 45-59.

Morant, G. hi. 1927 Studies of Palaeolithic man 11: A biornetric stiidy of Keanderthaloid skulls and their relationships to modern racial types. Ann.

Wovius, H. L. Jr, 1Y65 Upper Perigordian and L4urignnrian lir~artlis at thc: .4lrri Pataud, Les Eyzies (Dordognc). In: Miscelhea en Homenaje a1 Abate Hanri Breuil. E. R. PerellO, ed. 2: 181-189.

The hearths of the Upper Perigordian and Aurignacien at the Atrri Pataud, Les Eyzies (Dordognc.), and their possible significance. Am. Anthrop., 68 @t. 2, # 21: 246-325

19F9 The ChAteIperronian in French Ar- chaeology: the evidence of Arcy-sur-Cure. Anti- quit>-, 4 3 : 111-123.

___ 1972 Radiocarbon dating of the Upper Pa- leolithic sequence at the Abri Patand, Les Eyzies (Dordogne). In: Thc, Origin of Honm sapiens. F. Bordes, ed. Unesco. Paris, pp. 253-260.

Oakley. 6. P . . H. G. Campbell and T. 1. Molleson 1971 Catalogue of Fossil Hominids, Part 11: Europe. Trustees of the Rritish Miiseum (Natural History), London.

Perks, C. 1975 L'llomme prehistorique et le feu. La Rerhcrchc, 60: 829-839.

Poirier, F. E. 197'3 Fossil Man: .4n Ewliitionary Jonrncy. C. V. Mosby Compai

Pradel. L. 1966 Transition Perigordian: skeletal and industrial. Curr. Anthrop.. 7: 33-50,

Prideaux, T. 1973 Cro-Magnon Man. 'Time-Life, New York.

253-276.

EuF;., 2: 318-380.

1966

FHAYER

Riqiiet, R. 1970 La race de Cro-Magnon, ahus de langage ou rkalitk objective? In: L'Homme de Cro- Magnon. G. Camps and G. Olivier, eds. Arts et Metiers Graphiques, Paris.

Rust, R. 1937 Das altsteinzttitliche Rentierjagcr- lager hleicndorf. K. Wachholtz Verlag, Neu- miulster.

1943 Die At- untl Mittolsteinzeitlichcn von Stellmoor. K. Wachholtz Verlag, Neumiinster.

Semenov, S. A . 1964 Prehistoric Technology. Barnes and Noblc, London.

de Sonneville-Bordes, D. 1960 Le Paleolithique Supkrieur en Pkrigord. CNRS, Bordeaux.

. 1963 Upper Paleolithic cultures in West- ern Europe. Science. 142: 347-355.

1974 The Upper Paleolithic: 33,000- 10,000 B.C. In: France before the Romans. S. Piggott and G . Daniel, cds. Tharnes and Hudson, London,

Stringer, C. B. 1974 Population relationships of later Pleistocene honiinids: a inultivariate studs; of av:iiIable crania. J. Arch. Sci., I : 317-342.

Szonibathy, J. 1925 Die diluvinlen Menschcnreste ails tlw Furst-Johaiins-Hohle bei Lautsch in Mabren. Die Eiszeit., 2: 1-34, 73-95.

Turner, C. G. 1969 Directionality in the canine field model. J. Dent. Res., 48: 1310.

Weiss, M. L., and A. E. Mann 1975 Human Biology and Behavior. Little, Brown and Co., Boston.

Wolpoff, $1. H. 1969 The effect of mutations under conditions of reduced selection. Soc. Biol.,

1971a Metric Trends in Hominid Dental Evolution. Case Western Rescrve University Press, Cleveland.

h functional measure of tooth size. Southw. J. Anthrop., 27: 279-386.

Dental reduction and the probable mutation effect, Am. J. Phys. Anthrop., 43: 307-308.

pp. 30-60.

16; 11-23.

l97lh

1975

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Metric dental change in the European upper paleolithic and mesolithic