Human dento-gnathic metric variation in mesolithic/neolithic Ukraine: Possible evidence of demic...

AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY 95:1-26 (1994)

Human Dento-Gnathic Metric Variation in Mesolithic/Neolithic Ukraine: Possible Evidence of Demic Diffusion in the Dnieper Rapids Region

KENNETH JACOBS Departement d‘anthropologie, Uniuerszte de Montreal, Montreal PQ H3C 357, Canada

KEY WORDS zation, Gene flow, Metric variance

Dental dimensions, Eastern Europe, Neolithici-

ABSTRACT Dental and gnathic metrics from a series of Mesolithic and Neolithic cemetery samples in the Dnieper River valley (Ukraine) are compared. Both male and female Neolithic samples have larger dental dimensions, wider dental arcades, and a more robust mandibular corpus than do the Mesolithic samples. In addition, the relative variances (RVs) of bucco-lingual dental breadths (as measured by modified Levene’s Tests) show an intriguing pattern of change from the Mesolithic into and through the Neolithic. Female RVs show a clear tenency to increase through time, while male RVs show more mixed tendencies. Such a pattern indicates that the increases in Ukrai- nian Neolithic dento-gnathic dimensions are plausibly attributable to low intensity gene flow (demic diffusion). Seen in the light of new chronometric, paleodietary, and paleolinguistic information, as well as in the context of recent archaeological models for ago-pastoralist origins in the North Pontic, these data suggest that gene flow via population interactions originating in or transient through the circum-Caucasus may have played an important role in producing the Ukrainian Neolithic dental increase. o 1994 Wiley-Liss, Inc.

Paleoanthropologists have long been aware that in parallel with the diverse cultural transformations by which the Euro- pean Mesolithic-Neolithic transition is defined, the skeletal biology of European human populations underwent pari passu a similarly important set of modifications. The primary mode of analysis of these biological modifications usually has been em- bedded in the concept of “ethnogenesis,” that is, the idea that roots of extant European ethnic (i.e., “racial”) groups can be traced back to their proto- or even prehistoric origins. The resultant origins quest takes the form of a tracking of the geographic distribution of “ethnic markers” through progres- sively older archaeological and human skeletal horizons (see, e.g., Bernhard and Kandler-Palsson, 1986).

Usually accepted as the “ethnic markers” par excellence have been the myriad qualita-

tive and quantitative cranial features that have dominated scientific study of the skull in both its quasi- (Gould, 1981; Haller, 1971) and Neo-Darwinian (Howells, 1989; Brace and Hunt, 1990) modes. The intent of this paper is not to probe the weak conceptual foundations of the classic ethnogenetic approach (cf., Jacobs, 1994~1, nor to question the appropriateness of cranial features to either the definition or phylogenetic reconstruction of European “ethnic groups.” Rather, these dual themes are noted here due to their long having diverted attention away from analysis of the human masticatory complex during the European Meso- lithic-Neolithic transition. In contrast to the literally hundreds of publications in which this period‘s variations in cranial sizelshape

Received March 4, 1993; accepted March 4,1994.

0 1994 WILEY-LISS, INC.

2 K. JACOBS

are described and ethnically evaluated, only a very few analyses have focused explicitly on the coeval variation of dento-gnathic metrics (e.g., Cope, 1983; Meiklejohn and Schentag, 1988; y’Edynak, 1989).

This scientific neglect of the dento- gnathic record of human biological change during the European transition from food- gathering to food-producing economies stands in clear contrast to research tendencies that are evident in other global regions (e.g., Brace and Mahler, 1971; Brace et al., 1987; Calcagno, 1986; Carlson and Van Ger- ven, 1977; Larsen, 1982; Lukacs, 1982; Smith et al., 1984). Despite rare cases wherein either dento-gnathic dimensions do not change or their reduction commences well prior to significant adoption of nutri- tional domesticates, a notable reduction in human tooth and jaw size across the forager1 food-producer transition appears to be the most commonly documented pattern among groups dating from this period (Calcagno and Gibson, 1988). Moreover, such dento- gnathic reduction frequently appears to be associated with parallel transformations of cranio-facial size and shape, discrete aspects of which often have been accorded wholly ethnogenetic interpretations (Carl- son and Van Gerven, 1979). Seen in this context, the data regarding human tooth- and jaw-size variation during the Mesolithic- Neolithic transition in Ukraine assume a particular significance. While Ukraine traditionally has attracted a large and com- prehensive paleoanthropological research effort, most of this work has focused on the cranium, reflecting the prevailing ethnogenetic research agenda (Debets, 1948; Gokh- man, 1966; Konduktorova, 1973; Kruts, 1984; Zinevich, 1967). Thus the Ukrainian Mesolithic and Neolithic dento-gnathic sample, although sizeable and well-preserved, is largely unstudied.

The purpose here, therefore, is to describe the dental and gnathic metrics for a series of Ukrainian Mesolithic and Neolithic cemeteries. This will document the pattern of human dento-gnathic variation during this archaeological transition. The pattern of metric variation that emerges will be compared to that seen most commonly elsewhere during similar cultural shifts. The

nature of the observed differences between Ukraine and other regions then will serve to focus attention on the broader cultural context of human biological change during the Ukrainian Mesolithic-Neolithic. As will become clear, consideration of this larger context is critical to any meaningful discussion of ethnogenetic origins and interactions in the region.

MATERIALS The Ukrainian human skeletal samples

reported here derive from a series of sites along the middle reaches of the Dnieper River (Fig. 1) and were analyzed by the author during an extended research stay in 1985 in the former Soviet Union (Table 1). The designation here of the sites as “Meso- lithic” and “Neolithic” follows the scheme of the primary archaeological scholar of the North Pontic region (Telegin 1985, 1987b, inter alia). A brief discussion of the immedi- ately pertinent features of the regional Me- solithic and Neolithic is warranted here (see also Kozlowski, 1989; Zvelebil and Dolukha- nov, 1991), although concerns for the reli- ability of these terms qua socioeconomic la- bels will be considered more fully below (see also Anthony, 1994; cf., Jacobs, 1994a).

The chronological and archaeotypological structure of the Ukrainian Mesolithic has been reconstructed principally by Telegin (1982, 19851, from whom this account is derived. Microlith-dominated, aceramic assemblages in the North Pontic are documented from earliest postglacial times up to ca. 6,900 b.p. (uncalibrated) and are typically found in association with the remains of a broad spectrum of subsistence re- sources. An internal relative chronology for the local Mesolithic has been proposed that is centered on human grave morphology and shifting proportions of lithic tool types. In general, the most recent sites are said to be those preserving extended human burials and/or “trapezoidal” microliths, since the advent of these cultural features is assumed to coincide with heightened “pre-Neolithic” influences originating ultimately in the Bal- kans. It is in just this fashion that the three Mesolithic sites analyzed here have been se- riated. Voloshskoye is said to be the oldest, followed first by Vasilyevka I, and then, by

3 UKRAINIAN DENTO-GNATHIC METRIC VARIATION

Fig. 1. Map of the lower Dnieper region. Cemeteries are discussed in the text and include: 1, Igren; 2, Voloshskoye and Surskoy Island; 3, Vasilyevka I, 11, and 111; 4, Vovnigi 2; 5, Vovnigi 1; 6, Vilnyanka.

Vasilyevka 111. Several extended burials within this last site are taken to represent a presumably “proto-Neolithic’’ phase of cemetery use (Telegin, 1982,1985).

Recently obtained Oxford University ac- celerator 14C dates for human bone samples collected as part of the present research modify this picture, however. At the Vasi- lyevka 111 Mesolithic cemetery, three different adult individuals produced (uncalibrated) dates from 10,080 b.p. (2100) to 9,980 b.p. (2100). The relative sequence of the three cemetery sites must now be considered suspect, particularly in light of the fact that one of the 14C-dated individuals from Vasilyevka I11 was in a “proto-Neolithic’’ extended burial. These dates also have further implications for our understanding the Me-

TABLE 1 . Sample size, sex composition, and principal reference for Ukrainian site samples analyzed

Sample composition Mesolithic sites: Vasilyevka I: 9 males and 8 females

(Stolyar, 1959); Vasilyevka 111: 16 males and 17 females (Telegin, 1962); Voloshskoye: 6 males and 3 females (Danilenko, 1955).

Neolithic sites: Igren: 2 males and 4 females (Telegin, 1987a); Surskoy Island: 6 males and 3 females (Lagodovskaya, 1949); Vasilyevka 11: 9 males, 6 females, and 3 unsexable adults (Konduktorova, 1974; Telegin, 1987a); Vilnyanka: 11 males, 10 females, and 4 unsexable adults (Surnina, 1961; Telegin, 1987a); Vovnigi 1: 12 males and 7 females (Rudinskii, 1955); Vovnigi 2: 42 males, 28 females, and 4 unsexable adults (Rudinskii, 1955).

The samples were studied in the Otdel antropologii, Muzey antropologii i etnografii Leningrad (now St. Petersburg) (Vasilyevka I1 and 111; part of Vovnigi 2); the Muzey antropologii, Moskovskii gosudarstvennii universitet (Igren; Surskoy Island; Vasilyevka I; Volosh- skoye; Vovnigi 1; part of Vovnigi 2); and the Lahoratoriya plasticheskoy rekonstruktsii, Institut antropologii, ANISSSR Moskva (Vilnyanka).

K. JACOBS 4

solithic of the middle Dnieper within its larger regional context. At least the site of Vasilyevka I11 now must be considered roughly contemporaneous with epi-paleolithic “cultural entities” in the Crimea (Kozlowski, 1989) and circum-Caucasus (Dolukhanov, 1984). Lithic stylistic similari- ties between all three locales include the presence of “trapezoidal” microliths, the re- covery of which at Vasilyevka I11 had been evaluated previously only with respect to the “belated appearance of such items in central Europe (ca. 8,000 BP; Kozlowski, 1989). It may be instead the case that trape- zoids in the northeastern Black Sea region occur earlier than elsewhere in Europe (cf., Kozlowski, 1989)) as part of a distinct path of influence spreading out of the Near East (Domanska, 1990b), a possibility that will have considerable bearing on the later discussion.

Descriptions of the Ukrainian Neolithic sites analyzed here are presented in Telegin (1987a), where a detailed relative chronology incorporating these sites also may be found (Telegin, 1987a: 120 et passim). At the core of this relative chronological scheme is a seriation of cemeteries based on their overall morphology and on the form of their con- tained burials. Of the sites discussed here, Vasilyevka 11, Igren, Vovnigi 2, and one layer of the multicomponent Vilnyanka site fall within (the older) Burial Phase “A.” The small sample from Surskoy Island is roughly contemporaneous with this phase. The younger (“B”) phase includes the individuals from Vovnigi 1 and the middle burial layer atvilnyanka (with the upper- most, very much more recent layer a t Viln- yanka not being included in this study). Phase “B” cemetery horizons have been radiocarbon dated to 5,000-4,500 B.C. (Cali- brated; uncalibrated radiocarbon range from 6,075 2 400 b.p. to 5,540 2 400 b.p.; Telegin, 1987a). Only Phase “B” cemeteries are generally considered to be “Neolithic” in the classic sense of being heavily reliant on faunal and floral domesticates (e.g., An- thony, 1994). Preceding Phase “A” sites are taken as Neolithic solely by virtue of their preserved ceramics (Telegin, 1987b). Phase “ A is estimated to be only marginally older than Phase “B” (not older than 5,500 B.C.),

for layers of the two are in direct strati- graphic contact at some sites. Thus the entire range of Neolithic sites discussed here traditionally has been placed within the span of a single millenium (Telegin, 1987a).

Conflicting with such a view are accelera- tor 14C dates recently obtained for three human burials from Vasilyevka I1 that average 7850 b.p. (ca. 6,600 B.C. calibrated; the uncalibrated range is 7,620 k 80 b.p. to 8,020 2 90 b.p.). Although in keeping with a relative chronological scheme wherein Vasi- lyevka I1 belong to the very earliest sub- phase of the Dnieper Rapids “Neolithic” (Telegin, 1987a), this date significantly overlaps the range for Mesolithic sites in Ukraine. Nevertheless, carbon isotope and other elemental analyses of sizeable human bone samples from both Vasilyevka 111 (Me- solithic) and Vasilyevka I1 suggest that the latter’s occupants already had adopted to a significant extent certain aspects of a typically “neolithic” subsistence (Jacobs, 1994a; Price and Jacobs, n.d.). Moreover, that this was perhaps true of the other Phase “A individuals as well has been shown in a comparison of the patterns of postcranial variation within the Mesolithic and Neolithic samples from the same cemeteries discussed here (Jacobs, 1993, 1994a). Thus, despite the obvious divergence of the 14C dates from Vasilyevka I1 from the absolute chronology assumed by Telegin (1987a,b), there is am- ple reason to accept the “Neolithic” sample here as an internally coherent “post-Meso- lithic” group, which group also shows non- dento-gnathic evidence of important socioeconomic changes relative to the Mesolithic sample.

METHODS Cranial, including all dento-gnathic parts,

and postcranial materials have undergone separate curation (i.e., storage, and often, even cataloging) in all of the museum collec- tions studied here. For myriad reasons, not the least of which is its differential treat- ment during excavation and subsequent curation, the cranial portion of any cemetery sample tends to be best preserved. As a result, this subsample usually approximates most closely the number of burials originally

UKRAINIAN DENTO-GNATHIC METRIC VARIATION 5

recorded as having been discovered at the site. For all these reasons, a preliminary sex determination was done by the author on all cranially represented individuals using the seriation method.

The cranial remains of each entire site sample was studied prior to the determination of sex for any particular specimen from that site. Then, among all cranial remains, a site-specific “perfect male” and “perfect female” were chosen, based on the standard array of size- and robusticity-related features. Finally, and beginning with those specimens most similar to one of the two “type specimens,” the other individuals in the site sample were sorted into the “male” or “female” category, based on their greater similarity to the growing number of specimens in either of the two sex groups. This process stopped either when all adults had been sexed, or when the few remaining individuals were equally similar to both groups of sorted specimens. These latter individuals were classed as “unsexable.” When the postcranial remains from each site were studied (usually much later), a sex diagnosis was made when possible, based solely on the postcranial criteria and without reference to any prior, skull-based determination. When all results were combined for the definitive sexing of each site sample, in no case was a skull-based sex designation found to be in- consistent with the postcranial sex diagnosis.

This time-consuming process is preferable to the standard “forensic” approach in that it is sensitive to each site’s particular degree and mode of expression of sexual dimorphism (White and Folkens, 1991). This is especially important during a period of tech- nological and economic change (e.g., the Me- solithic-Neolithic transition), wherein quali- tative and quantitative changes in the nature of human sexual dimorphism are to be expected (Stini, 1985). By first establishing the maximal extent and overall pattern of sexual dimorphism that is evident in a given site sample and by only then sorting individuals into sex categories, the seriation method avoids the errors inherent in the application of forensically based diagnostic criteria that are perhaps inappropriate to that site. A further advantage of this approach is

that by establishing site-specific patterns of sexual dimorphism in osteological features not treated by traditional, whole-skull (or pelvis) methods, a sex diagnosis could often be reached for a less skeletally complete, previously unsexed individual.

All measurements of the Ukrainian material reported here were taken by the author using finely pointed, electronic digital-read- out calipers to the nearest 0.01 mm. All cal- culations were performed at this level of measurement. Dental measurements represent maximum dimensions in the bucco-lingual (breadth = perpendicular to the mesiodistal axis of the tooth) and mesiodistal (length = parallel to the mesiodistal axis of the tooth) directions. Measurements of teeth worn beyond the point of maximal “swelling” in the pertinent dental plane were not taken. Intraobserver measurement error is estimated at 1.6% for length measures and 1.5% for breadth measures. To calculate these values, a complete set of dental measurements was taken twice, at an interval of two months, for the St. Petersburg portion of the Vovnigi 2 sample. The error figures represent the mean of the percent differences between the original and the repeated values for the sets of all mesio-distal and bucco- lingual measurements, respectively.

RESULTS Table 2 presents the summary dental di-

mensions for the Mesolithic and Neolithic combined-sex (male, female, and unsexable adult) samples. Sample size ranges from a minimum of N = 23 (Mesolithic mandibular I1 length) to a maximum of N = 111 (Neo- lithic mandibular M1 breadth). As a rule, bucco-lingual (breadth) sample sizes exceed those for mesio-distal (length) dimensions, the latter being much more prone to dental (interproximal) wear and thus exclusion of the worn tooth from measurement. Immedi- ately apparent is that Neolithic dental dimensions are virtually always larger than those of the Mesolithic. In fact, values for 13 of 16 (81%) Neolithic maxillary dental dimensions and 12 of 16 (75%) Neolithic mandibular dimensions exceed those of the Me- solithic. Of the maxillary differences, five (38%) are statistically significant (Mann- Whitney U-test a t P < 0.05 or better), while

K. JACOBS

TABLE 2. Mean breadths (BL) and lengths (MD) plus standard deviation (SD) and sample size (N) for maxillary and mandibular teeth of Ukrainian Mesolithic and Neolithic samples

6

I1 BL Meso Neo

Meso Neo

Meso Neo

Meso Neo

Meso Neo

Meso Neo

Meso Neo

Meso Neo

Meso Neo

Meso Neo

Meso Neo

M1 MD Meso Neo

Meso Neo

M 2 M D Meso Neo

Meso Neo

M3 MD Meso

I1 MD

I2 BL

I2 MD

C BL

CMD

P3 BL

P3 MD

P4 BL

P4 MD

M1 BL

M2 BL

M3 BL

Neo

__ Mean __

7.40 7.54

8.13 8.69

6.59 6.77

6.41 6.90

8.52 8.53

7.87 7.93

9.53 9.30

6.69 6.74

9.56 9.38

6.33 6.37

11.63 11.98

10.11 10.83

11.89 12.05

9.75 9.63

11.29 11.32

8.46 8.66

Maxilla Mandible ~~

SD N

0.42 0.45

0.72 0.70

0.40 0.43

0.56 0.47

0.58 0.53

0.48 0.43

0.58 0.55

0.45 0.49

0.52 0.54

0.38 0.46

0.58 0.50

0.68 0.67

0.69 0.68

0.75 0.81

0.82 0.75

0.70 0.69

37 70

27 47

38 69

29 49

42 89

31 64

40 97

34 71

39 102

35 69

45 105

35 83

45 107

33 74

43 82

31 56

%diff Mean SD N %diff

+1.9

+6.9$

+2.7*

+7.6**

+0.1

+0.8

-2.41

+0.8

-1.9

+0.6

+3.0**

+7.1**

+1.4

-1.2

+0.3

+2.4

6.01 6.28

4.66 5.13

6.32 6.58

5.30 5.75

7.54 7.84

6.63 6.95

7.94 7.78

6.75 6.77

8.42 8.25

6.80 6.81

11.02 11.15

11.19 11.51

10.69 10.71

10.76 10.70

10.18 10.32

10.61 10.81

0.47 0.57

0.50 0.49

0.47 0.39

0.48 0.43

0.51 0.46

0.61 0.40

0.63 0.44

0.40 0.36

0.53 0.54

0.46 0.44

0.52 0.46

0.61 0.62

0.51 0.57

0.47 0.57

0.64 0.61

0.71 0.68

35 65

23 47

42 84

30 60

51 91

40 66

50 101

39 76

50 103

41 75

53 111

43 82

53 109

43 81

52 100

40 68

+4.5$

+10.1**

+4.1*

+8.5**

+4.0**

+4.81

-2.0

+ 0.3

-2.0

+0.2

+1.8

+2.9$

+0.2

-0.6

+1.4

+1.9

Site composition of samples is as described in the text. All males, females, and unsexable adults are included in the samples. Significant percent differences (%dim between the means (Mann-Whitney U-test) are indicated as fallows: *P < 0.05; tP < 0.025; W' < 0.01; **P < 0.001, The percent difference between the means is calculated as (Mesolithic mean - Neolithic mean) x 100IMesolithic mean.

six (50%) of the larger Neolithic mandibular dimensions were significantly different. By contrast, of the seven dimensions for which the Mesolithic sample possessed the larger value, for only one (14%) of these (maxillary P3 breadth) was the difference statistically significant.

The pattern of larger dental dimensions in the Neolithic, rather than in the Mesolithic

where they might be expected, is all the more striking when the percent differences in the various dimensions between the two Ukrainian samples are compared to those evident in other Mesolithic-Neolithic con- texts. Table 3 presents the percent differences €or the Ukrainian samples, as compared to two other Mesolithic-Neolithic sample series, one from Nubia (data from


TABLE 3. Percent difference in mean maxillary and mandibular breadths (BL) and lengths (MD) between the Ukraninian combined-sex (male, female, and unsexable adults) Mesolithic and Neolithic samples compared to those

obtained in two similar MesolithiclNeolithic, combined-sex comparisons

Maxilla Mandible Nubia Danube Ukraine Nubia Danube Ukraine

(%) f%) l%) (%) (%‘) __ I1

I2

C

P3

P4

M1

M2

M3

Mea

BL MD BL MD BL MD BL MD BL MD BL MD BL MD BL MD

.n %

-6.8 -5.6 -7.5 -6.7 -6.2 -5.8 -6.5 -6.4 -7.3 -2.8 -6.1 -5.0 -6.4 -7.6 -6.1 -5.2 -6.1

-3.7 -6.2 -3.3 -10.3 -0.4 -4.5 -6.1 -2.4 -4.5 -2.3 -3.9 -3.4 -4.3 -3.0 -4.2 -0.6 -3.9

+1.9 +6.9 +2.7 f7 .6 +0.1 +0.8 -2.4 +0.8 -1.9 +0.6 +3.0 f7.1 +1.4 -1.2 +0.3 +2.4 f1.9

-8.3 -5.9 -8.6 -3.9 -9.3 -8.5 -9.1 -3.6 -7.2 -1.5 -6.9 -2.8 -7.2 -3.9 -8.1 -4.6 -6.2

-5.2 -7.8 -3.3 -7.7 -1.7 -5.6 -3.2 -5.5 -4.2 -2.4 -4.8 -7.5 -5.7 -2.9 -3.3 -2.2 -4.6

+4.5 +10.1 +4.1 +8.5 +4.0 +4.8 -2.0 +0.3 -2.0 +0.2 +1.8 +2.9 +0.2 -0.6 +1.4 +1.9 +2.5

The nubian values are from a comparison of composite Mesolithic and Neolithic site samples and are derived from Calcagno (1986). Nubian samples sizes range from 20 (Mesolithic mandibular I1 MD) to 204 (Neolithic mandibular M2 MD). The Danube basin values derive from unpublished data (courtesy of Dr. David Frayer). The comparison is between the Mesolithic dental sample from Vlasac (Iron Gates, Yugoslavia) and the early Neolithic (LBK) dental samples from Krs Krany and Vedrovice (Czechoslovakia). The Danubian Mesolithic sample size averages 24 (range 3-32). The Neolithic sample size ranges from 44 to 90.

Calcagno, 1986) and the other from central Europe (DW Frayer, unpublished data). Not surprisingly, the Nubian sample, the Neo- lithic part of which represents populations with a long history of participation in food- producing economies, exhibits the greatest degree of dental reduction. Both maxillary and mandibular dimensions reduce significantly at an average of slightly more than 6%. This figure is in keeping with adaptive/ functional models that predict a reduction in tooth and jaw size as diet becomes increasingly agriculturally based (Calcagno, 1986; Carlson and Van Gerven, 1977).

The Mesolithic component of the central European material derives from Vlasac, a hunter-fisher-forager site in the Iron Gates region of Serbia (Prinz, 1987). (This use of “central Europe” thus does not conform en- tirely to the historical sense of Mitteleuropa. However, if true Osteuropa is that part from the Carpathians to the Urals, the relative geographic centrality of the Balkans is evident). The Mesolithic at Vlasac is dated to 8,000-7,500 b.p. (uncalibrated; Voytek and Tringham, 1989: 494). The central Euro- pean Neolithic component consists of material from Vedrovice and Krs Krany, two Lin- ear Pottery (LBK) sites in Moravia. Given

their location near the southern edge of the rapidly northward-expanding LBK system, an age of ca. 6,300 b.p. is not improbable (Gimbutas, 1991).

A somewhat lesser degree of dental reduction is apparent in the central European series compared to that in Nubia. Maxillary dental reduction averages 3.9%, while mandibular dental reduction averages 4.6% for all dimensions. This difference from the Nu- bian values may reflect the relatively small time gap between the two central European samples, as well as a persistent hunter-forager component in LBK subsistence (Whit- tle, 1985). Nevertheless, standing in high contrast to both the Nubian and the central European series is the actual dental increase that is seen in Ukraine. Maxillary dental dimensions here are nearly 2% greater in the Neolithic, while Neolithic mandibular dental dimensions show an average 2.5% increase.

One possible explanation for this unex- pected pattern would lie in a sex-based bias in the Ukrainian Mesolithic and Neolithic samples. Yet, as can be seen in Tables 4 and 5, this is not the case. In fact, it is among the females that the more consistent pattern of dental increase is evident. Fully 14 of 16

K. JACOBS

TABLE 4. Mean dental breadths and dental lengths for Ukrainian Mesolithic us. Neolithic maxillary and

8

I1 BL Meso Neo

Meso Neo

Meso Neo

Meso Neo

Meso Neo

Meso Neo

Meso Neo

Meso Neo

Meso Neo

Meso Neo

Meso Neo

M1 MD Meso Neo

Meso Neo

M 2 MD Meso Neo

Meso Neo

M3 MD Meso Neo

I1 MD

I2 BL

I2 MD

C BL

C MD

P3 BL

P3 MD

P4 BL

P4 MD

M1 BL

M2 BL

M3 BL

Mean

7.46 7.64

8.19 8.76

6.69 6.83

6.46 6.88

8.68 8.60

7.89 7.89

9.64 9.29

6.81 6.77

9.69 9.47

6.38 6.36

11.74 12.09

10.24 10.72

12.00 12.18

10.02 9.58

11.58 11.45

8.59 8.58

mandibular samples

Maxilla Mandible SD N %diff Mean SD N %diff

0.46 0.43

0.69 0.66

0.42 0.43

0.45 0.50

0.54 0.47

0.48 0.40

0.64 0.55

0.49 0.45

0.58 0.52

0.41 0.43

0.64 0.45

0.71 0.61

0.53 0.62

0.77 0.70

0.76 0.60

0.78 0.61

25 38

17 25

25 43

20 28

28 48

21 32

26 53

21 33

24 55

22 34

28 50

21 38

29 56

20 34

27 43

19 26

+2.4

+7.0t

+2.1

+6.5$

-0.9

0

-3.67

-0.6

-2.3

-0.3

+3.0t

+4.7$

+1.5

-4.4t

-1.1

6.04 6.35

4.66 5.14

6.43 6.63

5.36 5.71

7.75 8.00

6.82 7.00

8.11 7.81

6.87 6.77

8.58 8.29

6.94 6.88

11.13 11.19

11.33 11.65

10.87 10.85

10.85 10.78

10.48 10.40

10.78

0.54 0.66

0.52 0.42

0.52 0.31

0.30 0.48

0.45 0.42

0.46 0.41

0.64 0.38

0.35 0.35

0.56 0.51

0.35 0.40

0.56 0.46

0.58 0.55

0.51 0.49

0.44 0.50

0.64 0.59

0.72

22 37

15 25

15 47

18 29

28 51

23 32

28 55

23 36

26 57

23 37

30 59

25 37

29 57

25 38

29 57

23

+5.1*

+10.3$

+3.1

+6.5f

+3.2*

+2.6

-3.7

-1.5

-3.4*

-0.9

+0.5

+2.8*

-0.2

-0.6

-0.8

~~

~~ -0.1 10.93 0.68 34 +1.4

Males only included in these samples. Significant percent differences (%dim between the means (Mann-Whitney U-test) are indicated as follows: *P < 0.05; t P < 0.025; $.P ‘I 0.01; **P < 0.001.

(88%) dimensions in both their maxillary and mandibular dentitions are higher in the Neolithic. Three of the maxillary and six of the mandibular Neolithic female increases show statistical significance, while none of the four Neolithic decreases is significant. For the males, only 7 of 16 (44%) maxillary dimensions show an increase in the Neo- lithic (with two at statistical significance),

with nine (56%) mandibular dimensions (five at statistical significance) also increasing in the Neolithic. A total of eight maxillary and seven mandibular male dental dimensions show a decrease from the Meso- lithic to the Neolithic, although none of these changes is statistically significant, Thus the overall pattern is one of an evident increase in Neolithic dental size in both


TABLE 5. Mean dental breadths and dental lengths for Ukrainian Mesolithic us. Neolithic maxillary and mandibular samules

Maxilla Mandible SD N %difT Mean SD N %diff Mean

I1 BL Meso Neo

Meso Neo

Meso Neo

Meso Neo

Meso Neo

Meso Neo

Meso Neo

Meso Neo

Meso Neo

Meso Neo

Meso Neo

M1 MD Meso Neo

Meso Neo

M2 MD Meso Neo

Meso Neo

M3 MD Meso Neo

I1 MD

I2 BL

I2 MD

C BL

CMD

P3 BL

P3 MD

P4 BL

P4 MD

M1 BL

M2 BL

M3 BL

7.29 7.45

8.04 8.48

6.39 6.71

6.31 6.81

8.19 8.38

7.81 7.89

9.33 9.30

6.49 6.66

9.35 9.29

6.25 6.32

11.46 11.86

9.92 10.83

11.70 11.86

9.34 9.62

10.81 11.15

8.26 8.67

0.31 0.45

0.79 0.56

0.28 0.41

0.79 0.38

0.54 0.61

0.49 0.46

0.38 0.53

0.31 0.49

0.32 0.54

0.32 0.43

0.46 0.53

0.62 0.63

0.89 0.74

0.52 0.87

0.70 0.83

0.55 0.62

12 25

10 16

13 21

9 18

14 34

10 26

14 37

13 31

15 42

13 30

17 45

14 35

16 43

13 33

16 36

12 27

+2.2

+5.5

+5.01

+7.9

+2.3

+1.0

-0.3

+2.6

-0.6

+1.1

+3.5t

+9.2**

+1.4

+3.0

+3.2

+5.0

5.96 6.16

4.65 5.00

6.18 6.52

5.21 5.73

7.28 7.63

6.38 6.85

7.72 1.74

6.57 6.74

8.25 8.23

6.63 6.68

10.87 11.07

10.99 11.32

10.49 10.53

10.64 10.53

9.81 10.18

10.36 10.68

0.31 0.39

0.49 0.42

0.36 0.42

0.67 0.34

0.46 0.42

0.71 0.38

0.57 0.47

0.40 0.35

0.44 0.53

0.54 0.42

0.44 0.47

0.62 0.64

0.42 0.55

0.51 0.53

0.43 0.63

0.64 0.68

13 24

8 18

18 32

12 26

23 35

17 29

22 41

16 35

24 41

18 33

23 47

18 40

24 48

18 39

23 41

17 32

+3.4

+7.5

+5.5$

+10.0t

+4.8$

+7.4t

+0.3

+2.6

-0.2

+0.8

+1.8

+3.0*

+0.4

-1.0

+3.8t

+3.1

Females only included in these samples. Significant percent differences (%dim between the means (Mann-Whitney U-test) are indicated as follows: *P < 0.05; tP < 0.025; W < 0.01; **P < 0.001.

sexes, albeit to a seemingly less consistent degree in the males.

This differentially greater increase in females relative to the males is further in evidence when sex-specific percent differences in dental dimensions are considered (Table 6) . For maxillary dental size, the average male increase in the Neolithic is 1.2%, compared to the average female maxillary tooth-

size increase of 3.2%. The respective mandibular dental values are comparable, with males showing an average 1.5% increase in tooth size, while females again document a 3.2% average increase. As a result of this male-female differential in degree of dental change, a concomitant Neolithic reduction in dental sexual dimorphism is also evident (Table 7). Mesolithic dental sexual dimor-

10 K. JACOBS

TABLE 6. Mean percent differences between the Ukrainian Mesolithic and Neolithic samples for all

maxillary and mandibular dental dimensions

Maxilla Mandible Male Female Male Female

I1

I2

C

P3

P4

M1

M2

M3

Mes

BL MD BL MD BL MD BL MD BL MD BL MD BL MD BL MD

in %

+2.4 +2.2 +7.0 +5.5 t2 .1 +5.0 +6.5 +7.9 -0.9 +2.3

0 +1.0 -3.6 -0.3 -0.6 +2.6 -2.3 -0.6 -0.3 +1.1 +3.0 +3.5 +4.7 +9.2 +1.5 +1.4 +4.4 +3.0 -1.1 +3.2 -0.1 +5.0 +1.2 +3.2

+5.1 +3.4 +10.3 +7.5 +3.1 +5.5 +6.5 +10.0 +3.2 +4.8 +2.6 +7.4 -3.7 -0.3 -1.5 +2.6 -3.4 -0.2 -0.9 +0.8 +0.5 f1.8 +2.8 +3.0 -0.2 -0.4 -0.6 -1.0 -0.8 +3.8 +1.4 +3.1 +1.5 +3.2

Male and female differences calculated separately and as in Table 2.

TABLE 7. Percentage dental sexual dimorphism for Ukrainian Mesolithic and Neolithic maxillary and

mandibular breadths and lengths

Maxilla Mandible Meso Neo Meso Neo

I1 BL 2.3 2.5 1.3 3.0 MD 1.8 3.2 0.2 2.7

I2 BL 4.5 1.8 3.9 1.7 MD 2.3 1.0 2.8 -0.4

C BL 5.6 2.6 6.1 4.6 MD 1.0 0 6.4 2.1

P3 BL 3.2 0.1 4.8 0.9 MD 4.7 1.6 4.4 0.4

P4 BL 3.5 1.9 3.8 0.7 MD 2.0 0.6 4.5 2.9

M1 BL 2.4 1.9 2.3 1.1 MD 3.1 -1.0 3.0 2.8

M2 BL 2.5 2.6 3.5 3.0 MD 6.8 -0.4 1.9 2.3

M3 BL 6.6 2.6 6.4 2.1 MD 3.8 -1.0 3.9 2.3

Mean 9i 3.4 1.1 3.7 2.0

Percent dimorphism calculated a6 (male mean - female mean) x loo/ male mean.

phism averages 3.4% for the maxillary teeth and 3.7% for the mandibular teeth. For the Neolithic, dental sexual dimorphism values are 1.1% (maxillary) and 2.0% (mandibular). Reduction in dental sexual dimorphism is widely reported as accompanying the transition from the Mesolithic to the Neolithic, but this is usually the product of a greater degree of dental reduction in the males, with the dental dimensions of this sex thereby converging on those of the lesser reducing

females (e.g., Calcagno, 1986). In the Ukrai- nian samples, by contrast, dental sexual dimorphism reduction between the Mesolithic and Neolithic must be seen as the result of a broader trend to larger teeth, with the greater average dental metric increase occurring in the females.

It must be recalled that the total Neolithic sample derives from six spatially and chro- nologically distinct sites. The apparent Neo- lithic dental increase, therefore, could be merely the statistical artifact of the artifi- cial conjoining of these site samples. The initial possibility to be tested in this regard was that of a single, exceptionally megadont site sample having skewed the results. The individual Neolithic sites were compared using one-factor analysis of variance tests (ANOVA) on males and females separately, and for all dental dimensions (a total of 32 per sex). Of the 64 calculated ANOVAs, only in six cases was the F-test P < 0.05, suggestive of intersite differences at the usually accepted significance threshold. It must be noted, however, that in none of these cases was P < 0.001, the critical value required in order to safely reject the null hypothesis when such a large number of redundant tests are conducted (Cope, 1989). In each of these six cases, pairwise comparisons showed that the values for the “outlying” site were actually smaller than those from the other sites. Moreover, they were significantly smaller only relative to the single largest site sample in the comparison, not to the other sites. The dental metric homogeneity of the Ukrainian Neolithic sample thus would appear to be confirmed and the dimensional increase in this sample relative to the Mesolithic sample cannot be ascribed to the skewing influence of a single site.

The further possibility was tested of there having been a chronological bias introduced through the lumping of temporally distinct Neolithic sites, a bias overlooked by virtue of the fractionation of sample sizes inherent in the ANOVA. The Neolithic sample thus was divided into “early” and “late” components, corresponding to Burial Phases “A” and “B” as outlined previously. The chronological pattern of Mesolithic through Neolithic change in all dental dimensions, for both single- and combined-sex samples was then


Ukr MESO Ukr Neo ”A”

12

mm

11

10

9

8

7

6

5 I1 I2 C P3 P4 M1 M2 M3 I1 I2 C P3 P4 M1 M2 M3

A B

Fig. 2. A Histograms showing mean mandibular dental (bucco-lingual) breadths for the Ukrainian Mesolithic, Neolithic “A,” and Neolithic “B” samples. Samples are with sexes combined, and their composition is as defined in the text. B: Same comparison as in A, except that the remains from Vasilyevka I1 have been classified here as “Mesolithic.” See the text for discussion.

evaluated. All such evaluations produced equivalent results, but only mean mandibular dental breadths for the combined-sex Mesolithic, Neolithic “A,” and later Neolithic “B” samples are shown here (Fig. 2A). It can be seen that for none of the teeth are either of the Neolithic phases distinctly inconsis- tent with the overall pattern of Mesolithic- to-Neolithic change for that tooth. Even in the enigmatic cases of dental dimensional decrease (P3, P4), the pattern across the three time periods is consistent. This also held true in the final test for any bias possi- bly introduced by the lumping of site samples into larger time periods. Here, the Vas- ilyevka I1 material, potentially problematic due to its early radiocarbon date, was classed with the Mesolithic material. The pattern of change in dental dimensions does not differ substantially from that seen when the site sample is classed in the earlier Neo- lithic phase (Fig. 2B).

The changes in dental dimensions already noted are accompanied by changes in the

size of the gnathic skeleton. The breadth of the palate increases significantly from the Mesolithic to the Neolithic in both males and females (Table 8). These increases average 5.8% and 6.2%, respectively. Palate length appears to decrease from the earlier to the later period, although very small sample sizes, especially in the females, render meaningless any judgment regarding the statistical significance of this trend. Man- dibular sample sizes are more adequate, however, and a number of significant tendencies are evident, particularly in breadth measurements. In both sexes, bi-mandibular breadth is greater in the Neolithic than in the Mesolithic, although statistical significance occurs solely in the males (Mann- Whitney P < 0.05). For mandibular breadth measured at the coronoid notch, however, a greater Neolithic breadth is present and is statistically significant in both males and females. Mandibular length, on the other hand, while showing a slight decrease in the Neolithic for both sexes, does not show sta-

K. JACOBS

TABLE 8. Comparison of Ukrainian Mesolithic and Neolithic male and female averages for selected gnathic dimensions

12

Male Mean

Palate breadth at M2 Meso Neo

Meso Neo

Meso Neo

notch (2) Meso Neo

Palate length (1)

Bi-mandibular breadth at M1

Mandibular breadth at coronoid

Mandibular length (gonion-gnathion)

Meso Neo

Mandibular bi-M2 breadthlmandibular length

Meso Neo

Mandibular corpus breadth at Ml/M2

Meso Neo

Meso Neo

height (at MlIM2)

Mandibular corpus height at MUM2

Mandibular corpus breadthlcorpus

Meso Neo

63.3 67.0

38.4 36.8

58.2 59.9

101.2 107.1

87.4 85.2

74.6 78.4

13.1 14.5

31.9 33.2

41.3 44.0

SD

4.20 3.23

4.04 2.87

1.89 2.29

4.72 6.46

5.12 5.42

6.60 4.71

1.23 1.72

2.58 3.00

5.73 6.53

__ Mean __

59.2 62.9

40.4 35.3

55.6 57.6

93.2 100.3

82.5 80.1

76.3 80.6

12.1 13.9

29.2 29.8

42.6 47.0

Female SD (N)

1.83 3.28

-

1.92

5.03 2.68

5.55 4.72

6.32 5.46

7.61 5.52

1.28 1.25

2.04 2.39

5.21 5.42

Notes on measurements: (1) Palate length is taken in midline from Orale to the intersection of the maxillary and palatine sutures. ( 2 ) Distance between the vertical rami of the mandible, taken at the lowest point in each of the coronoid notches. Correlation of this measurement with bi-coronoid and bi-condylar mandibular breadths invariably exceeds r = +0.9. Thus use of this measurement yields comparable results, but at larger sample sizes, due to typically greater preservation of this part of the ramus. Significant differences between Mesolithic and Neolithic means for males and/or females are indicated. Symbols as in Table 2.

tistical significance despite relatively large samples for this measure. The mandibular breadtMength index, being larger in the Neolithic for both sexes (significantly so for males), thus would seem to indicate that the Neolithic mandibular arcade has become relatively wider (as opposed to relatively shorter). Associated with this wider mandible is a broader (both absolutely and relatively) mandibular corpus. Mandibular corpus breadth increases from Mesolithic to Neolithic by 10.7% in males and by 14.9% in females. Corpus height in males increases only 4.1% during this same time, while the female corpus height increase is barely 2%. The result of these changes in terms of relative corpus breadth is that the male Neo- lithic mandibular corpus is 6.5% more robust than in the Mesolithic, while the

Neolithic female mandibular corpus is fully 10.3% more robust than in the Mesolithic.

Thus, the dental increase from the Ukrai- nian Mesolithic to the Neolithic is not an isolated phenomenon, but is instead part of a larger transformation of the entire dento- gnathic apparatus. In the wake of function- ally oriented studies elsewhere (Carlson and Van Gerven, 19771, this transformation might be expected to be associated with more general changes in cranio-facial andor cranial vault features. For the Ukrainian Mesolithic-Neolithic skeletal series, a number of such cranial sizehhape changes have been discussed by others (Debets, 1948; Gokhman, 1966; Konduktorova, 1973; Kruts, 1984; Zinevich, 1967) and have been used as evidence in reconstructions of the ethnogenetic prehistory of Ukraine. For this


reason, an analysis exploring the extent to which these numerous cranial changes co- vary with the dento-gnathic transformations described here is currently in progress.

DISCUSSION Many previous studies of the human bio-

logical changes coincident with sundry “Me- solithic-Neolithic” transitions have found that the passage from a foraging to a food- producing economy often is associated with a concomitant decrease in dental dimensions and in the general robusticity of the facial architecture (Brace et al., 1991; Cal- cagno, 1986; Larsen, 1982; Smith et al., 1984). While there is debate over the precise evolutionary mechanisms effecting this change (see, e.g., Calcagno and Gibson, 1991 on “selective compromises” vs. “probable mutations”), and while any Neolithic dental reduction may be in simple fact a mere con- tinuation of a process already underway in the Upper Paleolithic (e.g., Frayer, 1978), it is nonetheless a reasonable expectation that the dentition should get smaller during any given Mesolithic-Neolithic transition. Yet quite the opposite trend is evident here among the Mesolithic and Neolithic cemetery samples of the Dnieper Rapids region. Four different, but not perforce mutually ex- clusive, explanations for this counterintui- tive result suggest themselves: The cemeteries do not represent a Mesolithic-Neolithic transition comparable to those studied elsewhere; larger teeth were a side effect of a body size increase in Neolithic groups relative to their antecedents; the dental changes resulted from selective pressures for larger teeth that acted upon local populations during the Mesolithic-Neolithic transition, and dental dimensions increased as a result of gene flow into Ukraine.

To reject an expectation of dental reduction during this region’s Mesolithic-Neo- lithic transition as unjustified by virtue of the Ukrainian Neolithic’s idiosyncratic nature does appear at least initially warranted. The prevailing interpretation of the Phase “A” cemeteries here considers them to represent societies that are “Neolithic” only in the minimalist sense of possessing archaeological assemblages with pottery, but without metal artifacts (Telegin, 1987b).

The subsistence base assumed for such societies is said to be little different from that reconstructed for the Mesolithic-broad- spectrum hunting of cervids, ovi-caprids, and equids, but with a major focus on fresh- water fish and molluscs (Dolukhanov, 1986). Only later, during the period of the Phase “B” cemeteries, is subsistence assumed to have changed significantly, when elements of the Near East agropastoralist complex were introduced by Tripolye farmers whose roots lay in the Carpathian piedmont and, ultimately, the Balkans (Zvelebil and Dolukhanov, 1991). If this scenario is wholly accurate, the Ukrainian “Neolithic” would differ markedly from the other Neo- lithic samples considered (A- and C-Group Horizon Nubia; LBK Central Europe), where the epithet denotes a major subsistence shift.

To reject the Neolithic of Ukraine as some- how sui generis merely begs the question, however. Since the dental increase clearly begins with the initial shift from the Meso- lithic to Neolithic Phase “ A samples (Fig. 2), one still must explain why the ostensible persistence of Mesolithic-like subsistence would foment a dental dimension increase. Also in need of explanation is why both Ukrainian Neolithic males and females show patterns of increased articular and di- aphyseal robustness in their postcranial skeletons (Jacobs, 1993, 1994a), which are elsewhere associated with the shift from mo- bile forager to more sedentary food producer (Bridges, 1989). Partial resolution of at least this latter derivative issue may be offered by evidence that an important dietary transition already had occurred in central Ukraine by Burial Phase “ A times, well before the Phase “B” cemeteries and their well- documented interactions with Danubian-in- fluenced agropastoralists.

Indirect evidence derives from extensive edgewear and replication studies of Meso- lithic assemblages in the North Pontic (Ko- robkova, 1993). These show that cereal gathering became of steadily increasing- and, ultimately, prime-subsistence importance throughout the period that culmi- nated in the earliest Phase “A” cemeteries, leaving such groups “on the threshold of food production” (Korobkova, 1993: 170). Direct

14 K. JACOBS

evidence of this is provided by osteochemis- try analyses of human bone samples from Vasilyevka I11 and Vasilyevka I1 (Jacobs, 1994a; Price and Jacobs, n.d.). In addition to important shifts in barium and strontium concentrations, increasing carbon isotope (S13C) values between the older and younger site suggest a trend toward greater consumption of C, plants in the group buried at Vasilyevka 11. At present, a likely candidate for such a plant is wild millet, which was cultivated in this region by at least 5,000 B.C. as a domesticated crop (Zohary and Hopf, 1978). Interestingly, another plant of possible relevance here is lambs quarters. A variety of chenopodium that may belong in the category of C, plants (JH Burton, pers. comm.), its intensive use is attested by its palynological abundance in the Mesolithic horizons of several North Pontic sites (Ko- robkova, 1993). Thus the presence of “Neo- lithic-like” postcranial robustness may be explained by the advent of more Neolithic- like subsistence activities by Phase “ A times (Jacobs 1994a). This, of course, re- news the question: Why then a dental increase?

Here, the potential confounding effect of body size must be considered briefly, for if Neolithic individuals were simply larger than those of the Mesolithic, it would not be unusual for their teeth also to be somewhat larger. For the Mesolithic, male femur length averages 470 mm (N = 20), while average female femur length is 439 mm (N = 14). For the Neolithic, the averages are 471 mm for the males (N = 31) and 441 mm for the females (N = 23). These values differ from those reported elsewhere (e.g., Debets, 1966; Gokhman, 1966) due to larger sample sizes made available through use of a re- gression technique for estimating the original lengths of fragmentary femora and tib- iae (Jacobs, 1992b). The estimated mean statures derived from these values (after Feldesman et al., 1990) are males, 175.7 cm (Mesolithic) and 176.2 cm (Neolithic); females, 164.3 cm (Mesolithic) and 165.0 cm (Neolithic). Neolithic stature thus shows less than a 0.05% increase from the Meso- lithic. Larger Neolithic teeth cannot be dis- missed as a correlate of larger body size in general. Neolithic teeth are both absolutely

and relatively larger than those of the Meso- lithic.

Of the remaining two explanations for a Neolithic dental increase-natural selection for larger teeth or gene flow into Ukraine- the former appears the less probable. If selection were driving the dental increase, the dental dimension curves for the two Neo- lithic burial phases should show some ap- preciable change in their shape and skew relative to the Mesolithic tooth size curve. Specifically, the Neolithic curves ought to envince some degree of peaking, along with a definite positive skew. Yet consideration of the summary kurtosis and skewness data (Table 9) reveals that in virtually all cases (the sole exception being Neolithic Phase “B” maxillary dental dimensions) all three sample phases tend heavily toward platykurtic distributions. The pattern of skewness is more mixed. It does not, however, tend to support a selection hypothesis, for while there is some slight increase in the fre- quency of positively skewed dimensions in the first Neolithic phase, there is an equivalent predominance of negatively skewed dimensions in the last Neolithic sample. Be- yond these statistical contraindications, a selection scenario is difficult to reconcile with other, ancillary data.

On the standard reconstruction of regional culture history (see, e.g., Anthony, 1994), the cemetery samples analyzed here derive from populations wherein diet (the most likely source of any putative selective pressures on dental size) initially was one typical of temperate-latitude European for- agers (for the Mesolithic and Neolithic Phase “ A groups). Diet then came to be dominated by (at least some of) the cereal and herd-animal products typical of Central European farmers. Precisely this pattern of dietary change was accompanied by significant dental reduction in Central European skeletal samples (as documented above), a reduction readily explicable by reference to evolutionarily plausible selective-pressure models (e.g., Calcagno and Gibson, 1991). In Ukraine, however, both the earlier period of ostensible dietary stasis and the later phase of dietary change are clearly associated with notable increases in dental dimensions. It is not readily apparent how a selection-based


TABLE 9. Number of cases ofpositive (i) and negative (-1 kurtosis and skewness in the sample distributions of dental lengths and breadths bv time veriod (combined sex samvles)

Neo “A” Neo “B” ..~ - Meso + ~ + - + -

Kurtosis Maxilla

Lengths Breadths Subtotal


Mandible

Total Skewness

Maxilla Lengths Breadths Subtotal


Mandible

Total

3 2 5

4 2 6

11

2 6 8

2 7 9

17

5 6

11

4 6

10 21

6 2 8

6 1 7

15

4 2 6

4 2 6

12

5 5

10

6 3 9

19

4 6

10

4 6

10 20

3 3 6

2 5 7

13

4 5 9

2 4 6

15

5 3 8

4 3 7 15

~

4 3 7

6 4

10 17

3 5 8

4 5 9

17

Composition of samples defined in text

model might reconcile Ukraine’s combina- tion of dietary convergence on the Central European pattern with trends in dental dimensions of a diametrically opposite nature.

A far more conciliant accomodation of these data can be accomplished by consider- ing the possible effects of gene flow, particularly given recent models of the cultural ecology of “neolithicization” in Europe (e.g., Zvelebil and Rowley-Conwy, 1984; Zvelebil, 1986; Zvelebil and Dolukhanov, 1991). Ac- cording to one such heuristically useful model (Zvelebil and Dolukhanov, 1991), forager populations peripheral to established agriculturalist “centers” pass through a series of predictable phases once they enter a trajectory leading towards total integration into a food-producing economy. The first of these is the “availability” phase, in which “farming is known to the foraging groups and there is some exchange of materials and information between them and the farming communities” (Zvelebil and Dolukhanov, 1991: 239). The subsequent phase is the “substitution phase,” during which the economy incorporates an increasing number of agropastoralist elements, but is not yet wholly dependent on these.

The osteochemical comparison of the Vas- ilyekva I1 and I11 cemeteries, as well as comparison of the larger set of Mesolithic and

Neolithic postcranial data (Jacobs, 1993, 1994a), make it reasonable to consider the Ukrainian Neolithic populations analyzed here as having been between the availability and substitution phases. While most dietary sources perhaps remained undomesticated, significant interactions between sites of the Dnieper Rapids Neolithic and those representing more fully agropastoralist economies is well documented, particularly in sites dating to cemetery Phase “B” (Doluk- hanov, 1979; Telegin, 1968). At the very least, the populations with which the Dnieper Rapids groups were in contact were well into the “consolidation phase” of neolithicization, which is to say that they were groups increasingly dependent on the man- agement of domesticated species. Indeed, in many regards these groups warrant consideration as the actual agricultural “center.” For a variety of reasons (Armelagos et al., 1991; Meiklejohn and Zvelebil, 19911, a demographic feature of populations in or near such center would have been a rate of population growth higher than that found in populations further away from the center. The archaeological visibility of this phenomenon in neolithicizing Europe is quite high, with many sites documenting that substitution and consolidation phases typically show a doubling and tripling of village numbers and

16 K. JACOBS

size relative to those of earlier periods (Gimbutas, 1991).

The high intrinsic population growth of neolithicizing groups does not necessarily suggest that the spread of agropastoralism was accompanied by major population dis- placements (cf., Ammerman and Cavalli- Sforza, 1984). In fact, the actual rate of population growth and the resultant level of migratory pressure implied by the archaeological data are not dramatic (Milisauskas and Kruk, 1989). These are less suggestive of a series of demic replacements than of a gradual infiltration into already occupied areas by individuals or very small groups (Zvelebil and Zvelebil, 1990twhat has been called demic diffusion (Sokal and Menozzi, 1982). Returning then to the context of the Ukrainian Neolithic, where groups clearly were involved in the “exchange of materials and information” with consolidated agriculturalist populations, three questions can be posed. First, might not such “information exchange” have included gene flow as a result of demic diffusion? Further, if the previous answer is positive, would such gene flow most probably have resulted in an increase in dental size? Finally, from where might such gene flow have occurred?

Dental morphology (cf., Scott and Turner, 1988) and osteological discrete traits (e.g., Kozintsev, 1992; Molto, 1983; Saunders, 1978) often provide data of great utility in assessing the biogeographic affinities of human skeletal populations. Unfortunately, such data are only inadequately available for the prehistoric Central Eurasian samples with which the Neolithic sample in Ukraine might be expected to demonstrate population ties. Some of the Dnieper Rapids material analyzed here was included recently in a broader dental morphological comparison that concluded, in part, that these teeth were typically European in their morphology (Haeussler, 1994). Clearly, any derivation of more detailed and useful inferences of gene flow patterns from the skeleto- dental data must be considered problematic until finer scaled, population-specific genetic markers can be identified. This caveat notwithstanding, it may be helpful to recall that the genetic variance within a popula-

tion will be altered if there is any change in the normal level of gene flow into that population from a source (or sources) with non- equivalent gene frequencies (Cavalli-Sforza and Bodmer, 1971). For this reason, to the extent that one could infer genetic variance from the phenotypic variance seen in a population’s skeleto-dental data, it would then be possible to infer changes in gene flow parameters over a time series of such populations. The phenotypic variables best suited to such an analysis are bucco-lingual dental breadths, in that these show relatively high genetic heritability values (Alvesalo and Ti- gerstedt, 1974) and are less prone to distortion by dental attrition than are other dental dimensions.

Thus the relative variances of dental breadths will be adopted here as a “proxy” variable for genetic variance, with the assumption that a sample’s increase in dental breadth variance signals a connate increase in the source population’s genetic variance.’ This procedure was prompted by Cope (19831, wherein can be found a fuller explanation and justification of the methodology,

‘This assumption, particularly with respect to its application here, has not been directly tested, but it appears nonetheless reasonable. Across numerous human populations, over a wide range of environments, dental breadths are found to exhibit a high heritability; that is, the percentage of populational phenotypic variance that is due to underlying genotypic variance (Alve- salo and Tigerstedt, 1974). In that teeth as a whole do not re- model during life, and because dental breadths do not suffer from interproximal wear over the course of tooth use, these consis- tently high heritabilities must strongly imply a large genetic component in the determination of phenotypic expression. Thus higher phenotypic variances (in dental breadths) in one population relative to another are most likely to be the product of higher genotypic variance in that population, particularly when differences between the populations’ environments are not pronounced (such as is the case here). Contrary claims (reviewed in Bailit, 1966) are based either on variance differences among highly inbred strains of laboratory animals or on inferences from an apparent positive correlation between the coefficient of variation and inbreeding coefficients in certain comparisons of small, semi-isolated human demes (e.g., Tristan da Cunha) to larger population samples. In the former case, the extreme homozygos- ity of inbred labortory animals makes them dubious models for understanding the behavior of phenotypic variances in human populations. In the latter analyses, inadequate consideration is given to the confounding influence of extreme initial homogeneity (due to founder effects) in the formation of human isolates. Similarly ignored is the coefficient of variation’s susceptibility to distortion due to the disproportionate effect on the variance of a single, greatly different individual (such as might occur, for in- stance, with an immigrant joining a small, isolated population). For these reasons, and in view of the strong results obtained elsewhere by virtue of this assumption’s adoption (Cope, 19831, no reason is seen to not continue this practice here.


plus an application to a western and central European prehistoric data set. Briefly, however, differences between samples in the relative variances of their dental measurements are evaluated using modified Levene’s tests (Schultz, 1985). For each separate sample to be compared, the absolute deviation of each individual observation from the sample median is calculated. The result then is divided by the sample median, which thus gives each individual a relative variance value for each measurement. As calculated in this manner, the population’s mean relative variance value, therefore, is analogous to the more traditional coefficient of variation, but it is less prone to the prob- lems that inhere to the use of this latter variable when small, non-normally distrib- uted samples are being analyzed (Van Valen, 1978). This procedure thereby allows a robust comparison among any number of samples of the relative variabilities that characterize any particular metric features of interest.

The goal here, however, simply was to evaluate whether any differences between the intrasample dental breadth variances of the Mesolithic, Neolithic Phase “ A and Neo- lithic Phase “B” groups exhibited patterns such as might result from changes in the gene flow parameters over the three time periods. More precisely, the aim of the procedure used here was to estimate the relative diversity of the gene pools that served as the “catchment areas” for the cemeteries in this study. To this end, relative variance values (RVs) for dental breadths were calculated as described above. Because of the multisite composition of the whole sample and the multicomponent nature of Vilnyanka, two different methods for pooling of the site samples were evaluated. First, site-, sex-, and, for Vilnyanka, horizon-specific RVs were derived by calculating the RVs for males and females (separately) at most of the sites (ex- clusive of Surskoy, Igren, and the middle Vilnyanka layer). All individuals (and their RVs) then were sorted into the appropriate one of the three chronological samples, al- lowing mean RVs for each sex and time period to be compared. The second procedure was to calculate sex-specific RVs with all individuals already sorted into their appro-

TABLE 10. Mean “relative variance” values for Mesolithic, Neolithic Phase ‘a,” and Neolithic Phase “B” mazillary

and mandibular dental (bucco-lingual) breadths

I1 Meso Neo “A” Neo “B”

Neo “A Neo “B”

Neo “A Neo “ B

Neo “A” Neo “B’

Neo “A” Neo “B”

Neo “A” Neo “ B

Neo “A” Neo “B”

Neo “A Neo “B”

Meso Neo “A Neo “B”

Mean all teeth Meso Neo “A

I2 Meso

C Meso

P3 Meso

P4 Meso

M1 Meso

M2 Meso

M3 Meso

Mean

Maxilla Male Female

5.13 4.24 4.01 4.89 5.61 4.10 4.86 4.67 4.41 5.25 4.16 5.01 4.86 4.79 4.48 4.65 2.64 3.20 3.69 4.40 3.87 5.51 3.73 3.87

4.86 4.28 4.12

5.12 4.30

Neo “B” 4.22

3.59 4.78 5.28 3.67 3.71 4.82 5.04 4.35 7.19 3.15 3.89 3.96 2.60 4.38 4.63 2.97 3.45 3.95 4.73 5.24 4.37 5.73 6.27 5.62

3.94 4.51 4.98

4.10 4.36 5.01

Male

7.16 7.39 5.37 6.72 3.13 4.15 4.76 4.11 4.22 6.63 3.75 4.00 5.54 4.66 5.01 3.57 3.29 3.35 3.51 3.74 3.28 5.07 4.47 5.11

5.37 4.32 4.31

Mandible - 1 - ____ Female

4.09 4.21 6.24 5.05 4.04 6.32 5.18 3.30 5.20 5.89 4.37 5.19 4.09 4.94 5.27 2.91 3.25 3.74 3.09 4.50 3.89 3.73 5.02 4.37

4.25 4.20 5.03

~

Relative variance calculated as described in the text. Composition of samples defined in the text.

priate temporal group. This permitted the inclusion of the material that, for reasons of extremely small sample size, had to be ex- cluded from the first procedure. Since the pattern of results was not found to vary between the two pooling methods, the results of the second method are reported here (Ta- ble 10).

On the whole, average male RVs appear to drop over the three time periods, while female RVs seem to increase on average. For the former, the mean RV for all dental breadths decreases by 16% from the Meso- lithic to the Neolithic “ A phase, then drops a further 2% into the Neolithic “ B phase [all percentages calculated as (earlier value - later value)/earlier value]. Female RVs show an increase of over 6% from the Meso- lithic to the Neolithic “ A period and then increase another 15% between the two Neo- lithic phases. Interpreting these differences

K. JACOBS 18

TABLE 11. Sex-specific distribution of increase us. decrease in relative variances of maxillary and

mandibular dental breadths across three time periods

Male Female Change in RV + - + -

Maxilla Meso = > Neo “A” 2 6 7 1 Neo “A => Neo “B 3 5 6 2 Subtotal 5 11 13 3

Mandible Meso => Neo “A” 2 6 5 3 Neo “A” => Neo “ B 6 2 6 2 Subtotal 8 8 11 5

Total 13 19 24 8

is made problematic, however, by the fact that only four of the tooth-specific relative variances show a statistically significant change (male M1, I,, and P,; female M,; Kruskal-Wallis P < 0.051, with the change occurring between the Mesolithic and Neo- lithic Phase “A” in each case. Thus of consid- erably greater interest in this case are the patterns of RV increases and decreases that may be seen for the separate sexes across the three time periods (Table 11). Despite their average RV decrease, males actually showed a variance increase in 13 of the 32 possible changes (i.e., 16 dental breadths over two temporal transitions). In contrast, females showed a far more consistent pattern of change, with an increase in relative variance in 24 of 32 cases. While the male pattern is statistically indistinguishable from that which chance alone would create, the female pattern is such that its binomial probability of occurrence due solely to chance is P = 0.004 (cf., sign test; Siege1 and Castellan, 1988: 324).

Thus a reasonable interpretation would envision male variance as somewhat reduced or unchanged, while females, despite the lack of a statistically significant difference for any single tooth, show a strong pattern of increased variance. These different variance patterns of the two sexes provide yet another reason to doubt that natural selective pressures are the cause of the dimensional changes seen here. Any change in the intensity of such selective pressures would indeed alter intrapopulational dental metric variances (Calcagno and Gibson, 1988), but it is difficult to imagine a plausible selection-mediated scenario involving sex-dis-

tinctive and opposite shifts. A more par- simonious interpretation is that the evident changes in relative variance values reflect underlying alteration of the variance of the respective-sex gene pools from which are derived the Mesolithic and Neolithic skeletal samples. The demographic dynamics creating such a situation in the Dnieper Rapids context are easily imagined.

Chapman (1989) has argued that in the Iron Gates region of Serbia and Romania, during an “availability-substitution” phase similar to that of the middle Dnieper region, the average spatial extent of “late Meso- lithic” breeding networks steadily decreased. This resulted from both growth in local population size and a heightened population density imposed by expanding and adjacent socioeconomic systems, which were modifying and preempting access to ever larger portions of the local habitat. Under such circumstances, a relative reduction in local genetic variability might be expected as increased local endogamy became more pronounced for at least a segment of the population. Among late Mesolithic groups of the middle Dnieper region, the highly endoga- mous population segment most likely would have differentially comprised males, judg- ing from the not-infrequent evidence of ho- micide in cemeteries of the period (Telegin, 19821. Such evidence of interpersonal vio- lence is quite plausibly seen as reflecting recurrent conflicts over (hereditary?) rights to resource procurement territories (Mal- lory, 19901, a process perhaps even exacer- bated by loss of such territories as a result of the region’s incipient neolithicization.

A high degree of male local exogamy in such a context would be, at the least, unusual. Instead, a strong increase in local male homogeneity, that is, a decreased male variance, would be expected. This tendency would be strongest in the earlier stages of the economic transformation, as local Meso- lithic groups contracted and before the demographic landscape was wholly dominated by agropastoralists. It therefore is interest- ing that nearly all of the average decrease in male relative variances seen here occurs between the Mesolithic and Neolithic “ A samples, with little change as the Neolithic later


consolidated. Moreover, that the overall male pattern of variance decrease is not particularly strong (indeed, as noted, it is quite mixed) may be attributable to countervail- ing demographic tendencies among females, for connate with increasing male genetic centripetality there would have been an intensifying degree of female exogamy.

Territorially expanding agropastoralist economies are argued to have suffered a demographic imbalance whereby there would have been a predominance of young males (Chapman, 1989; Dennell, 1985). One therefore might expect such conditions to foment a movement of females to “out-group~~’ at a level equaling or even exceeding that seen among extant horticulturalists. Such a tendency would be particularly pronounced during later stages, when domestic production is intensifying and consolidating. Exog- enous females, upon entering the local mat- ing network, clearly would increase the biological variability of local populations. Yet this change should be evident not just among the females in any subsequent cemetery sample, but also among the males, since the variance-reducing effects of the latter’s greater local endogamy would be at least partly mitigated by the male infants being born to exogenous females. “Frontier” agropastoralist population dynamics of this sort may very well underlie the pattern seen here wherein a female relative variance increase-an increase that is indeed greater in the latter part of the temporal transition-is combined with a somewhat attenuated decrease in male relative variance values.

Thus the response to the first of the questions previously noted is that it appears quite likely that the pattern of dental variance changes observed over the three time periods here reflects, at least in part, altered patterns of gene flow both within and from without the region. A subsequent question then must be whether the apparent Ukrai- nian Neolithic dental increase constitutes a probable consequence of such gene flow changes. To respond fully to this query would require a consideration of the biogeographic pattern of Mesolithic human variation in west and central Eurasia far exceeding the purview of this paper (cf., Jacobs, 1992b, 1993, 1994~; Meiklejohn et al., 1994;

Meiklejohn and Zvelebil, 1991). However, it warrants noting briefly that one aspect of this pattern entails a tendency for Meso- lithic populations of East Europe-including those of the middle Dnieper region-to have dental dimensions that are reduced relative to roughly contemporaneous populations found elsewhere in Europe or southwesthouth central Asia (Table 12). Given this larger biogeographic context, the Ukrainian Neolithic dental increase perhaps should be seen as wholly expectable, rather than enigmatic. Virtually any increase in gene flow into East Europe would have been accompanied by at least some increase in the dental dimensions of the subsequent East European populations.

A feature common to most scenarios for the MesolithicAVeolithic transition in the North Pontic is the role of allochthonous peoples in this transformation. A usual concomitant of presumed exogenous influence in the region is some degree of gene flow, whether at a moderate level associated with low-intensity demic diffusion, or on a more massive scale associated with major population migration (cf., Zvelebil and Zvelebil, 1990). The primary zone of origination for any such population interaction is usually argued to be within an area encompassing the middle Danube basin and Carpathian piedmont, extending, perhaps, into the Bal- kans more generally (Dolukhanov, 1986; Gimbutas, 1991; Renfrew, 1988; Zvelebil and Dolukhanov, 1991). Yet two discrepen- cies arise on this view of the matter, at least with respect to dental dimensions. First, the available data for the teeth of the southeast European Mesolithic (i.e., Vlasac; Table 12) do not demonstrate a size differential relative to the Ukrainian Mesolithic of an order likely to produce the Ukrainian Neolithic dental increase. A second, and more telling, dilemma arises from the archaeological observation that extensive contacts between the populations of the middle Dnieper and those further west in Europe do not reach important levels until the onset of Burial Phase “B” (Telegin, 1987a,b), while the Ukrainian dental increase as documented here was well underway by the time of Burial Phase “A.”

This dental change thus is not compatible

20 K. JACOBS

TABLE 12. Summed dental areas (all maxillary and mandibular teeth included) for selected west and central Eurasian Dooulations rouphlv contemuoraneous with the Ukrainian Mesolithic samule

Sample H dental area (mm’) Source

North India Mesolithic composite

Ganj Dareh (Iran) early Neolithic

Skateholm (Sweden) Mesolithic

European coastal Mesolithic

Natufian Epipaleolithic composite

Levantine Neolithic composite

European inland Mesolithic

Vlasac (Serbia) Mesolithic cemetery

sample

sample

cemetery sample’

composite sample

sample

sample

composite sample

1,366

1,363

1,285

Lukacs and Pal, 1993

Lambed, 1980

Frayer (pers. comm.)

1,260 Frayer, 1978

1,260

1,258

1,215

Smith et al., 1984

Smith et al., 1984

Frayer, 1978

1,199 Frayer (pres. comm.) sample’

composite sample

cemetery sample

Dnieper Rapids Mesolithic 1,192 this study

Olenii ostrov (Karelia) Mesolithic 1,186 Jacobs, 199213

‘Unpublished samples analyzed after publication of Frayer (1978). Thanks are extended to DW Frayer for graciously sharing these data.

with the classic “out-of-Danubia” model, by the light of which all significant influences in the North Pontic are traceable to sources in the greater Danube Basin. Yet it does coincide with other major changes in the behavior of North Pontic populations. As men- tioned earlier in the present paper, osteochemical analyses of human bones from Vasilyevka I11 and Vasilyevka 11 indicate a noteworthy dietary shift between the two sites. The more recent site shows evidence of a type of cereal consumption, which perhaps presages more fully “Neolithic” patterns, much as lithic assemblages elsewhere in the region bear witness to an ever-increasing exploitation of cereal grains (Ko- robkova, 1993). Moreover, changes in the postcranial skeleton of the same Mesolithic and Neolithic cemetery samples analyzed here are strongly in the direction predicted to accompany a subsistence shift of this type (cf., Bridges, 1989). Neolithic males and females, though equivalent to their Mesolithic counterparts in long bone lengths (and thus stature), nevertheless, exhibit markedly greater long bone robustness. With the increase in upper body robustness exceeding that of the lower limb, and with female robustness generally increasing more than that of males, an intensification of the sorts

of subsistence activities more usually associated with greater sedentism and food production is clearly indicated (Jacobs, 1993).

Nothing would preclude all of the above changes representing more than a local, en- dogenously stimulated subsistence intensification. Yet the occurrence of shifts in the earliest North Pontic Neolithic gene pools-if indeed this is what the dental Rel- ative Variances imply-would argue that influences exogenous to the region were of importance, influences unlikely to have originated in greater Danubia. This returns then to the question of where the source(s) of such influence might plausible lie. In this regard, the Ponto-Caspian intersea area (the “circum-Caucasus”) and the Irano-Tu- ranian zone still further south must be considered probable. Indirect support for this derives from a recent paleolinguistic analysis (Colarusso, 19921, wherein the common ancestry of the Proto-Indo-European (PIE) and Proto-Northwest Caucasian languages is posited and reconstructed. The stem language-termed Pontic-has a maximal time depth (established relative to other accepted paleolinguistic branching times) roughly coincident with the start of the Dnieper Rap- ids Mesolithic. Moreover, that PIE (with its own, much later core area in the North


Ponto-Caspian; cf., Anthony, 1991) and Proto-Northwest-Cacuasian diverged from a common source strongly implies a common “homeland within which population contacts remained extensive over most of the period represented by the cemetery samples analyzed here.

Archaeological evidence corroborates this notion, for important shared archaeotypological features among their respective assemblages suggest that there were indeed continued population interactions between the circum-Caucasus region (and further south) and the North Pontic, extending perhps even into the greater east-central European Plain (Domanska, 1990a; Koz- lowski, 1989). These interactions began in at least the terminal Pleistocene, with the very early presence of trapezoidal forms at Vasi- lyevka III (as noted above, dated ca. 10,000 b.p.) perhaps being but one manifestation of this larger interaction sphere. Population contacts much have remained of continued importance throughout the Mesolithic and into the earliest Neolithic, given the sugges- tion by some specialists (e.g., Shnirelman, 1989, 1992) that domesticated ovi-caprids and perhaps even millet spread from the circum-Caucasus into the North Pontic by a time coincident with the Burial Phase “A” of the latter area. On this view, the socioeconomic influences of southeast European agro-pastoralist traditions (e.g., Tripolyel Cucuteni) only later came into a dominant position in the North Pontic (Zvelebil and Dolukhanov, 1991).

Yet because Neolithic developments in the Caucasus were only partly autochthonous- and in the interest of finding the true impetus for their consequent effects in the North Pontic-it is essential not to lose sight of important contemporaneous developments in the Irano-Turanian zone or in zones still further southwest. Here, among the many neolithicizing populations represented by the Natufian and similar coeval/successor traditions, may lie the ultimate source for many of the population dynamics which af- fected the early Neolithic North Pontic (Bar- Yosef and Belfer-Cohen, 1989; Domanska, 1990b). Significantly, the dental metric data for one appropriately dated and located

sample are not incompatible with such a view, as may be seen in Figure 3.

Illustrated are the mandibular dental areas for the Ukrainian Mesolithic and combined Neolithic samples. These are con- trasted with the dental areas for an early Neolithic sample from Ganj Dareh Tepe, Iran (Smith, 1978; dental data from Lam- bert, 1980). The Ganj Dareh material is intriguing for many reasons, among which are its early dates (7,5004,500 B.C.: Meikle- john, pers. comm.)-slightly preceding the earliest beginnings of the Ukrainian Neo- lithic (as used here)-and its possible evidence for domesticated ovicaprids, which, as noted above, may have entered Europe via the circum-Caucasus. It would be absurd to posit some manner of direct biological rela- tionship between the Ganj Dareh population and those of the Dnieper Rapids for a myriad of reasons, not the least of which is simply the geographic distance involved (roughly 2,000 km on a straight line). Still, if the Iranian sample is a t all typical of those populations to be found closer to or in the circum-Caucasus during that region’s neolithicization process (whose populations are unfortunately unrecorded by their skeletal remains), then it is notable that the Ukrai- nian Neolithic dental increase gives a t least the impression of a convergence on a much larger dentition as typified by that at Ganj Dareh. Recalling the archaeological and paleolinguistic evidence for population interactions between the North Pontic and more southern regions at the time of the Ukrai- nian Mesolithic-Neolithic, this apparent dental convergence therefore suggests that some degree of gene flow between the two areas might be plausibly viewed as adding yet another dimension to such interactions.

CONCLUSIONS A generalized increase in dental and

gnathic dimensions between Mesolithic and Neolithic cemetery samples in the Dnieper Rapids region of Ukraine has been demon- strated. The extent of the metric increase across the large majority of all dimensions in both sexes and the rarity of any statistically significant metric decrease between

K. JACOBS 22

M1

Ganj Dareh

Ukr Neo P4

P3 Ukr Meso

C

u

I1

20 40 60 80 100 120 140

sq. mm

Fig. 3. Histograms showing combined-sex mean mandibular dental areas for the Ukrainian Meso- lithic, Ukrainian Neolithic (Phases “A” and “B” combined), and Ganj Dareh (Iran) early Neolithic. See the text for discussion.

the two samples combine to suggest that these results warrant serious examination. No body-size effect, nor sex skewing of the samples, nor chronological or other impreci- sion resulting from the aggregation of separate site samples, can be invoked so as to reject the Neolithic dental increase as a mere statistical Poltergeist. Moreover, changes in the relative variance values of dental breadths through the Mesolithic- Neolithic transition suggest that some gene flow effect may underlie the observed changes in absolute dental dimensions.

Within the context of skeletal biological changes in prehistoric Ukraine, any discussion of gene flow perforce evokes an ambiva- lent reaction in light of the recurring role this “historical materialist process” traditionally has played within Soviet paleoanthropological orthodoxy (Alekseev, 1974, 1978). In a series of important articles, De- bets (1936,1948,1961) articulated the theo- retically canonical (and, to be fair, not wholly non-empirical) expectation of contin-

uous skeletal gracilization over the entire course of human evolution. From this Grun- dregel, it is a logical imperative to diagnose genetic input from foreign, vestigially retro- grade populations whenever a more recent group possesses a robusticity greater than that seen in the autochthonous, antecedent populations. Exemplary of this process would be the interpretations accorded to the widespread and problematic recognition that the Ukrainian Neolithic skeletal sample evinces somewhat greater cranial and postcranial robusticity than does the Meso- lithic sample (Debets, 1948; Gokhman, 1966; Konduktorova, 1973; Kruts, 1984; Zinevich, 1967).

Invariably, the solution to the problem has been to refer to the “primitivization” that results when there is mixing of the local, more evolved groups with intrusive, usually “Cro-Magnoid” elements (e.g., Bunak, 1976; plus others previously cited). As regards Ukraine, the source of the intrusive l‘robus”’ genes responsible for the “more


primitive’’ Neolithic material most often is said to have been the northern reaches of the Russian Plain (e.g., Konduktorova, 1974; cf., Gokhman, 1966). However, late Mesolithic populations in this region were characterized by very small dental dimensions (Jacobs, 1992b; see also Olenii ostrov in Table 12 on page 20, above). Any gene flow from the north during the period of the Ukrainian Mesolithic-Neolithic transition would have had an effect quite the opposite of that described here, to wit, a Neolithic dental reduction. Nonetheless, an element of truth is to be found in the standard explanation of Ukrainian Mesolithic-Neolithic distinctions, if only to the extent that gene flow in fact was a likely factor in the development of such distinctions.

The interpretive appeal to gene flow here, however, is not grounded in expectations regarding the long-term, orthogenetic tendencies that do (or, more correctly, do not) drive human biological change. Rather, the invo- cation of gene flow via demic diffusion is warranted in that it forms part of a coherent attempt to understand the cultural ecologi- cal dynamics of the Ukrainian Mesolithic- Neolithic transition. In this, the present study shares some features with numerous others that have developed and tested models of demic diffusion in an effort to associate the geographic pattern of human biological variation with diverse possible modes of Eu- ropean neolithicization (e.g., Ammerman and Cavalli-Sforza, 1984; Harding et al., 1989; Sokal and Menozzi, 1982; Sokal et al., 1989). The results obtained in these other studies often have been mixed, with the degree of congruence between the archaeologi- cally predicted spatial patterning and the actual spatial behavior of variation in the biological data depending in large extent on which biological variables were analyzed. In contrast to studies of this sort, however, the geographic region considered here is more circumscribed, and the phenotypic variance of the biological features being analyzed is less subject to multiple short-term influences. To this extent, then, and as compared to previous efforts, it is difficult in the present context to ascribe the Ukrainian dental increase to factors other than diffu-

sion into the Dnieper Rapids area of larger toothed individuals.

In short then, as has long been suggested, evidence for gene flow into the Dnieper Rap- ids area of Ukraine during the initial phases of the local Neolithic does exist in the paleoanthropological record. However, the odontometric data suggest that the source of this gene flow is not to be found among attenuated “Cro-Magnoid” populations of more northerly origin. In addition, while the archaeological evidence is clear that the major sociocultural transformation of the North Pontic Neolithic (the Dnieper-Donets I1 stage) involved interactions with populations with their roots further west in “greater Danubia,” these latter are unlikely to have been the source of the Ukrainian Neolithic dental increase. Instead, it is probable that the economic and demographic transformation of the Dnieper Rapids region began somewhat earlier than the classic Dnieper-Donets Neolithic, partly due to en- dogenous processes and partly as a consequence of interactions with populations located in the circum-Caucasus and Irano- Turanian zones. The impetus for the increase in dental dimensions among the early Neolithic populations of the North Pontic thus are most logically to be sought among these latter interactions.

ACKNOWLEDGMENTS Museum research in the former Soviet

Union (1985) was supported by IREX. My gratitude for their boundless help goes to all of the personnel of the Otdel antropologii, Muzey antropologii i etnografii Leningrad (St. Petersburg), the Muzey antropologii, Moskovskii gosudarstvennii universitet, and the Laboratoriya plasticheskoy rekonstruktsii, Institut antropologii, AN/SSSR Moskva. My thanks go to David Frayer (Uni- versity of Kansas) for graciously sharing, once again, some of his unpublished data; to Dana Cope and Sue Haeussler for stimulat- ing and enlightening discussion; and to Pier- rette Hart for the hardware. Comments from two anonymous reviewers led to important improvements in the final version of this paper and are gratefully acknowledged. The Departement d’anthropologie, Univer-

24 K. JACOBS

site de Montreal, underwrote technical help by Luc Litwinionek in the very early part of this work. An SSHRC Institutional Grant to the Universite facilitated the completion of the work. Finally, as always, to Kenna, Ian, and Cheryl, for being there.

LITERATURE CITED Alekseev VP (1974) Geografiya Chelovecheskikh Ras.

Moscow: Mysl. Alekseev VF' (1978) Paleoantropologiya Zemnogo Shara

i Formirovanie Chelovecheskikh Rac. Moscow: Nauka.

Alvesalo L, and Tigerstedt P (1974) Heritabilities of human tooth dimensions. Hereditas 77r311-318.

Ammerman AJ, and Cavalli-Sforza L (1984) The Neo- lithic Transition and the Genetics of Populations in Europe. Princeton, NJ: Princeton University Press.

Armelagos GJ, Goodman AH, and Jacobs K (1991) The origins of agriculture: Population growth during a period of declining health. Popul. Environm. 13:9-22.

Anthony DW (1991) The archaeology of Indo-European origins. J . Indo-Europ. Stud. 19:193-222.

Anthony DW (1994) On subsistence change at the Meso- lithic-Neolithic transition. Curr. Anthrop. 35:49-52.

Bailit HL (1966) Tooth size variability, inbreeding and evolution. Ann. NYAcad Sci 134:61&623.

Bar-Yosef, 0, and A Belfer-Cohen (1989) The origins of sedentism and farming communities in the Levant. J . World Prehist 3t447-498.

Bernhard W, and Kandler-Palsson A (1986) Ethnogen- ese europaischer Volker. (Aus der Sicht der Anthro- pologie und Vor- und Friihgeschichte). Stuttgart: Gustav Fischer.

Brace CL, and Hunt KD (1990) A non-racial cranio-fa- cia1 perspective on human variation: A(ustra1ia) to Z(uni). Am. J . Phys. Anthropol. 83:341-460.

Brace CL, and Mahler PE (1971) Post-Pleistocene changes in the human dentition. Am. J. Phys. Anthro- pol. 34: 191-204.

Brace CL, Rosenberg K, and Hunt KD (1987) Gradual change in human tooth size in the late Pleistocene and post-Pleistocene. Evolution 41 :70&720.

Brace CL, Smith SL, and Hunt KD (1991) What big teeth you had Grandma! Human tooth size, past and present. In MA Kelley and CS Larsen (eds.): Advances in Dental Anthropology. New York Wiley-Liss, pp. 33-57.

Bridges PS (1989) Changes in activities with the shift to agriculture in the southeastern United States. Curr. Anthropol. 30:385-394.

BunakV (1976) Rassengeschichte Osteuropas. In I Sch- widetzky (ed.): Rassengeschichte der Menschheit Vol. 4: Europa 11- Ost- und Nordeuropa. Munschen: Olden- berg, pp. 7-101.

Calcagno JM (1986) Dental reduction in Post-Pleis- tocene Nubia. Am. J. Phys. Anthropol. 70:349-363.

Calcagno, JM, and Gibson, KR (1988) Human dental reduction: Natural selection or the probable mutation effect. Am. J. Phys. Anthropol. 77:505-517.

Calcagno JM, and Gibson KR (1991) Selective compro- mise: Evolutionary trends and mechanisms in homi-

nid tooth size. In MA Kelley and CS Larsen (eds.): Advances in Dental Anthropology. New York: Wiley- Liss, pp. 59-76.

Carlson DS, and Van Gerven DP (1977): Masticatory function and post-Pleistocene evlolution in Nubia. Am. J. Phys. Anthropol. 46:495-506.

Carlson DS, and Van Gerven DP (1979) Diffusion, biological determinism, and biocultural adaptation in the Nubian corridor. Am. Anthropol. 81 :561-580.

Cavalli-Sforza LL, and WF Bodmer (1971) The Genetics of Human Populations. San Francisco: WH Freeman.

Chapman J (1989) Demographic trends in Neothermal south-east Europe. In C Bonsall (ed.): The Mesolithic in Europe. Edinburgh: John Donald, pp. 500-515.

Colarusso J (1992) Phyletic links between Proto-Indo- European and Proto-Northwest Caucasian. In HI Aronson (ed.): The non-Slavic languages of the USSR- Linguistic Studies (second series). Chicago: Chicago Lingustic Society, University of Chicago, pp. 19-54

Cope DA (1983) Geographic variation of dental metrics in the Mesolithic and Neolithic of Europe. Master's Thesis, University of Kansas, Lawrence.

Cope DA (1989) Systematic Variation in Cercopzthecus Dental Samples. Ph.D. Dissertation, University of Texas, Austin.

Danilenko VI (1955) Voloshskii epipaleolithicheskii mogilnik. Sovet. Etnograf. 3:56-61.

Debets GF (1936) Bryunn-Pschedmosti, Kro-Manyon i sovremennye rasy Evropy. Antropologisch. Zhournal 3:316322.

Debets GF (1948) Paleoantropologiya SSSR. [Trudy In- stituta Etnografii 4(n.s.)]. Moscow: Nauka.

Debets GF (1961) 0 nekotorykh napravleniax izmenenii v stroenii cheloveka sovremennogo vida. Sovets. Et- nograf. 2.9-23.

Debets GF (1966) 0 fizicheskom tipe naseleniya dnepro- donetskoy kultury. Sovetsk. Arkheol. 1:14-22.

Dennell R (1985) The hunter-gatherer/agricultural frontier in prehistoric Europe. In SW Green and SM Perl- man (eds.): The Archaeology of Frontiers and Bound- aries. Orlando: Academic, pp. 113-139.

Dolukhanov PM (1979) Ecology and Economy in Neo- lithic Eastern Europe. London: Duckworth.

Dolukhanov PM (1984) Upper Pleistocene and Holocene cultures of the Russian Plain and Caucasus: Ecology, economy, and settlement pattern. In F Wendorf and AE Close (eds.): Advances in World Archaeology, Vol. 1. New York: Academic Press, pp. 323-358.

Dolukhanov PM (1986) The Late Mesolithic and the transition to food production in eastern Europe. In M. Zvelebil (ed.): Hunters in Transition. Cambridge: Cambridge University Press, pp. 109-119.

Domanska L (1990a) Kaukasko-Nadczarnomorskie Wzorce Kulturowe w Rozwoju Poznomezolitycznych Spoleczenstw Nizu Strefy Pogranicza Europy Wschodniej i Srodkowej. (Studia i Materialy do Dzie- jow Kujaw, No. 5) Wroclaw.

Domanska L (1990b) The Role of the Near East factor in the development of late Mesolithic communities of the central and eastern part of the European Plain. In P Vermeersch and P Van Peer (eds.): Contributions to the Mesolithic in Europe. Leuven: Leuven University Press, pp. 323-333.

Feldesman MR, Kleckner JG, and Lundy JK (1990)

Femurhtature ratio and estimates of stature in mid- and late-Pleistocene fossil hominids. Am. J . Phys. An- thropol. 83~359-372.

Frayer DW (1978) Evolution of the Dentition in Upper Paleolithic and Mesolithic Europe. Lawrence KS: Uni- versity of Kansas Publications in Anthropology (#lo).

Gimbutas M (1991) The Civilisation of the Goddess. San Francisco: Harper.

Gokhman I1 (1966) Naseleniye Ukrainy v epokhu mezolita i neolita (antropologicheskii ocherk). Moscow: Nauka.

Gould SJ (1981) The Mismeasure of Man. New York: Norton.

Haeussler AM (1994) Upper Paleolithic teeth from the Kostenki sites on the Don River, Russia. In J Moggi- Cecchi and WP Luckett (eds.): Teeth: Form, Function and Evolution (Ninth International Symposium on Dental Morphology). In press.

Haller JS, Jr 1971 Outcasts from Evolution. Urbana: University of Illinois Press

Harding RM, Rosing FW, and Sokal RR (1989) Cranial measurements do not support Neolithization of Eu- rope by demic expansion. Homno 40:45-58.

Howells WW (1989) Skull shapes and the map: Cranio- metric analyses in the dispersion of modern Homo. Cambridge, MA: Harvard University Press.

Jacobs K (1992a) Estimating femur and tibia length from fragmentary bones: An evaluation of Steele’s (1970) method using a prehistoric European sample. Am. J . Phys. Anthropol. 89:333-345.

Jacobs K (199213) Human population differentiation in the peri-Baltic Mesolithic: The odontometrics of Ole- neostrovskii mogilnik. Hum. Evol. 7:3343.

Jacobs K (1993) Human postcranial variation in the Me- solithic-Neolithic of the Ukraine. Curr. Anthropol. 34:

Jacobs K (1994a) On subsistence change at the Meso- lithic-Neolithic transition. Curr. Anthropol. 35.52-59.

Jacobs K (1994b) Oleneostrovskii social organization re- visited Skeletal biology and social differentiation in a boreal forest Mesolithic cemetery. J . Anthropol. Ar- chaeol., In press.

Jacobs K (1994~) Multi-ethnicity in pre-Indo-European northeast Europe: Theoretical and empirical con- straints on the interpretation of prehistoric human biodiversity. Paper presented at International Con- ference. The Indo-Europeanization of Northern Eu- rope. September 1994, Vilnius, Lithuania.

Konduktorova TS (1973) Antropologiya naseleniya Ukrainy mezolita, neolita, i epokhu bronzy. Moscow: Nauka.

Konduktorova TS (1974) The Ancient population of the Ukraine (from the Mesolithic Age to the first centu- ries of our era). Anthropologie (Brno) 12:5-149.

Korobkova GF (1993) The technology and function of tools in the context of regional adaptations: A case study of the Upper Paleolithic and Mesolithic of the northwestern Black Sea region. In 0. Soffer and N.D. Praslov (eds.): From Kostenki to Clovis: Upper Paleo- lithic-Paleoindian Adaptations. New York: Plenum Press, pp. 159-173.

Kozintesev A (1992) Ethnic epigenetics: A new approach. Homo 43t213-244.

311-324.

of the western part of the Russian Plain. In C Bonsall (ed.): The Mesolithic in Europe. Edinburgh: John Don- ald, pp. 424-442.

Kruts SI (1984) Paleoantropologicheskie Issledovaniya Stepnogo Pridneprovya. Kiev: Naukova Dumka.

Lagodovskaya OF (1949) Kamyani zakladki Na- dropizhzhya (za materialami doclidzhen 1945- 1946gg). Arkheol. Pamyat (UkrRSR) 2t159-179.

Lambert PJB (1980) An Osteological Analysis of the Ne- olithic Skeletal Population from Ganj Dareh Tepe, Iran. Master’s Thesis, University of Manitoba, Win- nipeg.

Larsen CS (1982) The Anthropology of St. Catherines Island. 3. Prehistoric human biological adaptation. Anthropol. Pap. Am. Mus. Nat. Hist. 57t155-270.

Lukacs J R (1982) Cultural variation and the evolution of dental reduction: An interpretation of the evidence from South Asia. In A Basu and KC Malhotra (eds.): Human Genetics and Adaptation. Calcutta: Indian Statistical Institute, pp. 252-269.

Lukacs JR and Pal J N (1993) Mesolithic subsistence in North India: Inferences from dental attributes. Curr. Anthropol. 34:745-765.

Mallory JP (1990) Social structure in the Pontic-Cas- pian Eneolithic: A preliminary review. J . Indo-Eur.

Meiklejohn C, and Zvelebil M (1991) Health status of European populations at the agricultural transition and the implications for the adoption of farming. In H Bush and M Zvelebil (eds.): Health in Past Societies. Oxford: British Archaeological Reports, pp. 129-145.

Meiklejohn C, and Schentag CT (1988) Dental size in two Portuguese Mesolithic series and their implications. Am. J. Phys. Anthropol. 75:249 (abstract).

Meiklejohn C, Wyman J, Alexandersen V, and Jacobs K (1994) Craniometric evidence for population differentiation in the Mesolithic peri-Baltic. Paper presented in Invited Session, “Language, Culture and Biology in Prehistoric Central Eurasia,” Annual Meetings, American Anthropological Association, December, 1994, Atlanta, GA.

Milisauskas S, and Kruk J (1989) Neolithic economy in Central Europe. J . World Prehist. 3:403-446.

Molto E J (1983) Biological relationships of Southern Ontario Woodland peoples: The evidence of discontinuous cranial morphology. National Museum of Man, Mercury Series, No. 117: Ottawa, Canada.

Price TD, and Jacobs K (n.d.) Bone chemistry and dietary composition during the Mesolithic-Neolithic of Ukraine. Manuscript in preparation.

Prinz B (1987) Mesolithic Adaptations of the Lower Danube: Vlasac and the Iron Gates Gorge. Oxford: British Archaeological Reports.

Renfrew C (1988) Archaeology and Language: The Puz- zle of the Indo-Europeans. Cambridge: Cambridge University Press.

Rudinskii MYa (1955) Vovnigskie pozdneneolithich- eskie mogilniki (k voprosu ob oformlenii mogilnikov mariupolskogo tipa). Kratkie Soobscheniya (AN- SSSR, Kiev)4:147-151.

Saunders SR (1978) The development and distribution of discontinuous morphological variation of the human infracranial skeleton. National Museum of Man.

Stud. 18:15-57.


Kozlowski SK(1989) A survey of early Holocene cultures Mercury Series No. 81: Ottawa, Canada.

26 K. JACOBS

Schultz, BB (1985) Levene’s test for relative variation. Syst. Zool. 34:449456.

Schwidetzky I, and Rosing FW (1989) Vergleichend- statistische Untersuchungen zur Anthropologie von Neolithikum und Bronzezeit. Homo 40:545.

Scott GR, and Turner CG I1 (1988) Dental anthropology. Ann. Rev. Anthropol. 17:99-126.

Shnirelman VA (1989) Voznikovenie Proizvodyashego Khozyaistva. Moscow: Nauka.

Shnirelman VA (1992) The emergence of food-producing economy in the steppe and forest-steppe zones of East- ern Europe. J . Indo-Europ. Stud. 20:123-143.

Siege1 S and NJ Castellan Jr. (1988) Nonparametric statistics for the behavioral sciences. New York: Mc- Graw Hill.

Smith P, Bar-Yosef 0, and Sillen A (1984) Archaeologi- cal and skeletal evidence for dietary change during the late Pleistocendearly Holocene in the Levant. In MN Cohen and GJ Armelagos (eds.): Paleopathology at the Origins of Agriculture. New York Academic Press, pp. 101-136.

Smith PEL (1978) An interim report on Ganj Dareh Tepe, Iran. Am. J . Archaeol. 84:538-540.

Sokal RR, and Menozzi P (1982) Spatial autocor- relations of HLA frequencies in Europe support demic diffusion of early farmers. Am. Natural. 119:l- 17.

Sokal RR, Harding RM, and Oden NL (1989) Spatial patterns of human gene frequencies in Europe. Am. J. Phys. Anthropol. 80:267-294.

Stini WA (1985) Growth rates and sexual dimorphism in evolutionary perspective. In B Gilbert and J Mielke (eds.): The Analysis of Prehistoric Diets. Orlando: Ac- ademic Press, pp. 191-226.

Stolyar AD (1959) Pervyi Vasilyevskii mezolithicheskii mogilnik. Arkheol. Sborn. 1:78-158.

Surnina TS (1961) Paleoantropologicheskie materialy iz Volnenskogo mogilnika. Trudy Instituta etnografii

Telegin DYa (1957) Tretti Vasilyevskii mogilnik. Krat-

Telegin DYa (1962) Vasilivski tretiy nekropol v nad-

(MOSCOW) 71:3-25.

kie Soobscheniya (Kiev) 7:9-12.

porozhzhi. Arkheologiya (Kiev) 13:3-19.

Telegin DYa (1968) Dnipro-Donetska Kultura. Kiev: Naukova Dumka.

Telegin DYa (1982) Mesolithichni Pamyatki Ukraini (9-6 tisyacholitta do n.e.1. Kiev: Naukova Dumka.

Telegm, DYa (1985) Pamyatniki Epokhi Mezolita na Territorii Ukrainskoy SSR. Kiev: Naukova Dumka.

Telegin DYa (1987a) The archaeological remains. In DYa Telegin and ID Potekhina (eds.): Neolithic Ceme- teries and Populations in the Dnieper Basin. Oxford British Archaeological Reports, pp. 3-147.

Telegin DYa (1987b) Neolithic cultures of the Ukraine and adjacent areas and their chronology. J. World Prehist. 1:307-331.

Van Valen L (1978) The statistics of variation. Evol. Theory 4:3343.

Voytek BA, and Tringham R (1989) Rethinking the Me- solithic: The case of south-east Europe. In C. Bonsall (ed.): The Mesolithic in Europe. Edinburgh: John Don- ald, pp. 492499.

White TD, and Folkens PA (1991) Human Osteology. Orlando: Academic Press.

Whittle A (1985) Nelithic Europe: A survey. Cambridge: Cambridge University Press.

y’Edynak G (1989) Yugoslav Mesolithic dental reduction. Am. J . Phys. Anthropol. 78:17-36.

Zinevich GP (1967) Ocherki Paleoantropologii Ukrainy. Kiev: Naukova Dumka.

Zohary D., and M. Hopf (1988) Domestication of plants in the Old World. Oxford: Clarendon Press.

Zvelebil M (1986) Mesolithic prelude and Neolithic revo- lution. In M Zvelebil (ed.): Hunters in Transition. Cambridge: Cambridge University Press, pp. 5-15.

Zvelebil M, and Dolukhanov PM (1991) The transition to farming in eastern and northern Europe. J . World Prehist. 5:233-278.

Zvelebil M, and Rowley-Conwy P (1984) The transition to farming in northern Europe: A hunter-gatherer perspective. Norweg. Archaeol. Rev. 17t104-127.

Zvelebil M, and Zvelebil KV (1990) Agricultural transition, “Indo-European origins” and the spread of farming. In TL Markey and JAC Greppin (eds.): When Worlds Collide: Indo-Europeans and pre-Indo-Euro- peans. Ann Arbor, MI: Karoma, pp. 237-266.

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