Arctic Medical Research, Vol. 47: Suppl. 1, pp. 218-225, 1988

GENETIC COMPOSITION AND VARIATION IN SAAMI PC)PULATIONS IN NORTHERN COMPARED WITH NORDIC POPULATIONS IN MIDDLE NORWAY

A study of non-metric skull variants

E. lregren (1) and P.-E. lsberg (2)

Institute of Archaeology (1) and Department of Statistics (2), University of Lund, Lund, Sweden

Abstract. Skulls of six Saami populations from as well as of two Nor­ dic groups from Middle Norway were studied concerning non-metric traits. All the in­ dividuals derive from church-yards of Post-Medieval origin. The frequencies and combinations of non-metric traits were evaluated. Several traits 122/791 showed significant differencies between the two ethnic groups at the 5 % level. 146 individuals were used to construct a model in order to discriminate be­ tween the Saami and the Nordic group. The ethnic origin of a total of 475 individuals was predicted. Of the individuals from the Saami church-yards 92. 7% were grouped as Saamis. Of the skeletal sample from Middle Norway 52.3% were grouped as Nordic.

Arctic Medical Research, vol. 47: suppl. 1, pp. 218-225, 1988.

The Saa.mis (Lapps) have for a long period of time the South-Saami research project in which archaeol­ been of interest to anthropologists. A research history ogists from Norway and Sweden take part as well. Fur­ covering the period 1808-1931 has been published by ther, odontological and palaeopathological investiga­ Schreiner (1). In 1971 more recent works were compiled tions have also been performed. by Lewin & Hedegard (2,3). The studies have been per­ formed in different disciplines: physical anthropology, MATERIAL odontology and several fields of genetic research. The skeletal collections studied are available at the Anatomi­ Some titles are mentioned here in which further refer­ cal Institute of the University of Oslo. ences can be found. Investigations of the physical anthropology of living 1. Sumi populations Saamis as well as of skeletons have been of interest Six different Saami populations from , Northern Norway were investigated concerning non-metric traits. Ex­ (l-14). Odontological research has been performed as cavations from 1913 to 1935 have yielded the vast bulk of well (10,13,15-20). The blood groups and serum material. Further a few skeletons found during investigations groups of Saamis have been explored, especially dur­ in the 1950s (27 ,29) are also included in the present study. The ing the last decades (2,13,21,22). collections are dated to the 18th and 19th centuries. A total of 327 individuals were studied. The distribution of the burial Further genetic investigations regarding different grounds is seen at figure 1:1-6. An introduction to the biological and physiological aspects in living Saamis material can be found in two articles by Iregren (13,14). have been performed (2,21,23-25).Some articles deal These populations have been thoroughly examined by pro­ with the anthropology and the origin of the Saamis in fessor K.E. Schreiner, Oslo (l,4,5). Crania and skeletal general (26,27,28). Literature concerning diseases, elements were measured by him and pathological finds were described. The children's skeletons were analysed as well. The nutrition and health among Saami populations can publications of Schreiner constitute the most detailed and also be found (2,13,21). elaborate work on the physical anthropology of the Saainis The aim of the present study is to map the frequen­ performed in the Nordic countries. cies of non-metric traits of the cranium in populations 2. Nordic popnla_tions of different ethnic affiliation and to try to separate the To be able to evaluate the genetic composition and variation populations by using this biological information. of the Saami populations three populations of Nordic origin These anthropological studies constitute a part of were studied for comparison. One sample, possibly Nordic,

Cimnnpolar Health 87 lregren & lsberg: Genetic composition in Saami Populations in Northern Norway 219

Fig. I. Map of Norway showing the position of the church-yards. Saamish; l=, 2=, 3=Angsnes, 4=Kistrand, 5=Neiden, 6=Pasvik. Possibly Nordic; 7=Vard0. Nordic; 8=Prestgard, 9=Bj0lstad.

1. Antllropological metllods Since the late 1950s and the 1960s (33,34) minor mor­ phological variants have been studied in the human cranium. From animal studies (35-38) and a few investigations on man (39-43) it is known that these morphological traits are inherited. The exact heredity pattern is seldom known, how­ ever. It is ;lSsumed that the presence or absence of a cenain non-metric trait does not influence tile survival of the in­ dividual. In consequence they are uot subject to selective forces. Thus, it is probable that the traits are passed on from parents to children without changes and it is assumed that the frequencies of the traits remain the same in a certain popula­ tion. By studying the traits it should be possible to explore the genetic relationship between populations and to detect changes due to e.g. immigration or genetic drift. Non-metric traits can been studied at skulls as well as 8,9 skeletal elements (44). In this study only crania have been sub­ NORWAY ject to investigation. The tr.lits are unilateral as well as bilateral and a number of 44 different traits have been registered for skulls and mandibles (Definitions see (14)). A total of 79 observations were made on each individual.

2. Statistical metllods The data were analysed using a lo(listic regression model (45). This model can be used to analyze how a probability, in this case the probability of being a Saami, depends on different variables. The model states: ebo+b1x1+ ... + bkxk P=Prob. of being a Saami=------l+ebo+b1x1 + ... +~xk 0 2ookm where x1... x-. are background variables and b0 ••• bk are coefficients that are to be estimated from data. If the back­ ground variables are continuous this function is an S-shaped curve (in one dimension). emanates from Vard0 in Nonhern Norway (Figure l :7) found Another way of writing this relation is: in 1925. The date is somewhat uncenain, but the find is not LOG (P/(l-P))=bo+b1X1 ... +bkxk older than from the 14th century (30). This population has been investigated by other scientists (31,32). The other skele­ This may give a clearer interpretation of the model, in which tal materials were uneanhened in Middle Norway, where two one sees the natural logarithm of the odds of an event as a church-yards were excavated by Schreiner in 1925. One of the linear function of a number of variables. churches was built in 1531 and the other during the 1630s (30). After the model bas been fit it can be used to predict the A total of 148 individuals were examined in the present study. probability of being a Saami, when in our case a non-metric In figure 1:8-9 the position of the church-yards is shown. trait is present or non-present. 220 Anthropology and demography

Table I. Frequencies of non-metric skull variants in Saami populations, Nordic populations and Varde respectively

Population Saami Varde Nordic Used in Traits no. % no. % no. % log.reg.

V2 371290 12.8 6/35 17.1 11/88 12.5 V3 1231272 45.2 17/35 48.6 43/79 54.4 H3 1401275 50.9 20135 57.1 46/75 61.3 V4 171/317 53.9 23137 62.2 58/111 52.3 H4 173/317 54.6 19/37 51.4 61/110 55.5 V5 4/301 13.3 1/32 3.1 11108 0.9 V6 10/322 3.1 5/37 13.5 13/111 11.7 + V7 2/192 1.0 6/35 17.1 5/73 6.8 + H7 5/188 2.7 4/35 11.4 10/75 13.3 + vs 27/198 13.6 8/33 2.4 9/91 9.9 HS 36/202 17.8 7/35 20.0 8/82 9.8 V9 10/203 4.9 2/35 5.7 1/93 1.1 H9 6/213 2.8 1/35 2.9 2/84 2.4 V10 8/314 5.7 1/37 2.7 8/104 7.7 H10 26/312 8.3 6/37 16.2 13/99 13.1 V11 45/307 14.7 3137 8.1 6/105 5.7 + H11 62/306 20.3 6137 16.2 6/98 6.1 + V13 43/313 13.7 1/33 3.0 13195 13.2 H13 37/310 11.9 1/35 2.9 7/94 7.4 V14 105/294 35.7 15/36 41.7 33/92 35.9 H14 96/292 32.9 20/37 54.1 41/88 46.6 + V16 103/309 33.3 14/35 40.0 16/77 20.8 + H16 111/312 35.6 14/34 41.2 23/74 31.1 V17 14/319 4.4 2/34 5.9 0/73 0 H17 19/318 6.0 3/34 8.S 1/73 1.4 V19 168/322 52.2 20/36 55.6 45/93 48.4 H19 1331322 41.3 9136 25.0 43/92 46.7 V20 9/303 3.0 0/35 0 0189 0 H20 12/299 4.0 0/34 0 O/S5 0 V2i 33/295 11.2 3134 8.8 7/82 8.5 H21 33/303 10.9 5/34 14.7 5/81 6.2 V22 224/301 74.4 27/35 77.1 43/70 61.4 + H22 215/293 73.4 27/35 77.1 45/73 61.6 V23 215/313 68.7 6/36 16.7 5/89 5.6 + V24 105/257 40.9 2/35 5.7 1/70 1.4 + H24 106/257 41.2 2/36 5.6 2 el 3/74 2.7/4.1 + V25A 258/292 88.3 26/34 76.5 65/82 79.3 H25A 256/285 89.8 25/35 71.4 71/86 82.6 V258 104/285 36.5 10/34 29.4 35/81 43.2 H25B 104/277 37.5 10/35 28.6 28/86 32.6 V26 116/307 37.8 11/37 29.7 31/106 29.2 H26 125/310 40.3 13/36 36.1 26/102 25.5 + V27 9/307 2.9 3137 8.1 5/105 4.8 H27 14/308 4.5 0136 0 2/102 2.0 V30 12/266 4.5 9/34 26.5 18/76 23.7 + H30 4/259 1.5 4/35 11.4 12/80 15.0 + V31 0/316 0 0/36 0 0/83 0 V34 12/275 4.4 3/31 9.7 2/67 3.0 H34 11/263 4.2 3/27 9.0 1/73 1.4 V35 111/283 39.2 5/36 13.9 10/87 11.5 + V36 21/312 6.7 2/37 5.4 13/107 12.1 V37 8/191 4.2 4/29 13.8 2/47 4.3 V38 83/273 30.4 10/36 27.8 34/109 31.2 H38 68/276 24.6 6/36 16.7 39/105 37.1 + V39 24/309 7.8 1/35 2.9 4/89 4.5 H39 25/307 8.1 1/35 2.9 3/88 3.4 V40 3S/300 12.7 0/35 0 3/88 3.4 + H40 41/305 13.4 2136 5.6 4/81 4.9 V41 1781310 57.4 34/36 94.4 59/88 67.0 V42 1/314 0.3 0/36 0 0/85 0 V43 77/308 25.0 9/37 2.4 18/109 16.5

Cin:lunpolar Health 87 /regren & /sberg: Genetic composition in Saami Populations in Northern Norway 221

Population

Saami Varde Nordic Used in Traits no. % no. % no. % log.reg. V44 204/316 64.6 010 0 5/57 8.8 + H44 203/316 64.2 010 0 5/57 8.8 + V45 142/317 44.8 010 0 10/57 17.5 + H45 117/319 36.7 010 0 9/56 16.1 + V46 19/309 6.1 010 0 3/58 5.2 H46 18/312 5.8 010 0 1/57 1.8 V47 14/308 4.5 4136 11.1 4/80 5.0 H47 17/307 5.5 5/37 13.5 2176 2.6 V48 2271263 86.3 26/35 74.3 53/65 81.5 H48 221/256 86.3 27/35 77.1 53/69 76.8 V49 213/275 77.5 010 0 44/54 81.5 H49 217/271 80.1 010 0 44/52 84.6 V50 631177 35.6 6/18 33.3 21/40 52.5 H50 53/159 33.3 10/18 55.6 26/44 59.1 + V51 10/300 3.3 1/37 2.7 3/110 2.7 H51 14/302 4.6 1/37 2.7 6/110 5.5 V52 8/301 2.7 2/37 5.4 3/110 2.7 H52 5/303 1. 7 0/37 0 1/107 0.9

All multivariate statistical methods demand a complete 3. After reaching a final model. this model is used to obtain data set for every individual. This is of course unrealistic in the probability for each i.mfividual being a Saami. our case working with archaeological skeletal material. 4. This individual probability is then used for classifica­ Therefore we have chosen the following strategy to maximize tion. If an individual has a probability greater than 0.5 of the number of individuals in the analysis: being a Saami, he is predicted to be a Saami. Otherwise I. As preliminary analysis all the 79 traits in the two popu­ he is predicted as being of Nordic.origin. lations were compared using a chi-square-test (Yates cor­ 5. This is done for the total material (n=475). Missing rection). Only those traits that showed a significant dif­ values were then treated in accordance with the actual in­ ference at the 5 'lo level were used in the logistic formation about the frequencies of ttaits, or combita­ regression. tions of traits, in the two populations. If the true un­ 2. At each step in the logistic regression analysis the most known value of ibe missing observatioJ\ could have insignificant variables were omitted, thereby increasing changed the prediction (Saami/Nordic), the prediction is the number of cases in each step. classified as "possible".

Fig. 2. Classification tree from the estimated model. Start-point at V24. Traits; V24, H7 etc. l =trait present, 0= trait not pre­ sent. S,s indicate prediction Saami. N,n indicate prediction Nordic. Capitals show certain prediction. Minusc:uJes show possible prediction. 222 Anthropology and demography

RESULTS Table II. Classifying the individuals by means of the model compared to their population respectively. Description of the the Saami population from Nor­ compared to their population, respectively thern Norway and the Nordic population from Middle Norway. Prognosis In Table I the frequencies of all non-metric traits Possible Possible Population Saami Saami Nordic Nordic Total studied are presented. The individuals are divided into the groups: Saarni, Vard0 (possibly Nordic) and Nor­ Saamish dic respectively. The relative frequencies of the traits church-yards Kautokeino 1 6 0 1 0 7 are exposed as well. The traits used in the step-wise Kautokeino 2 17 12 1 0 30 logistic regression are indicated ( + ). Kautokeino 3 17 8 2 0 27 Karasjok 18 3 2 0 23 Angsnes 45 3 0 0 48 Comparison between Saami and Nordic populations Ki strand 82 5 2 2 91 Neiden 52 10 5 2 69 From the frequencies of the minor skull variants in Pasvik 1 3 5 6 0 14 Tuble I it is evident that differencies exist between the Pasvik 2 8 9 1 0 18 Saami and the Nordic populations investigated. The Nordic? variables of the Saami and Nordic populations respec­ church-yard Varde 9 4 0 24 37 tively were tested and twenty-two traits showed dif­ Nordic ferences that were significant at the 50fo level. A step­ church-yards wise logistic regression was used to find out which of Prestgard 4 13 32 5 54 8jelstad 5 31 57 the non-metric traits were most useful to separate the 6 15 populations in the final step. 146 individuals could be Total 266 103 58 48 475 used. The best traits were H7 (Coronal ossicle present at the right side), V23 (Palatine torus present), V24 (Left It appears from Tuble II that of 327 individuals from maxillary torus present), V35 (Infantile nose aperture) the Saami church-yards in Northern Norway, 303 and H44 (Right mandibular torus present). When (92. 70fo) were classified as Saamis or possibly of Saami either of four traits was present it turned out that the origin. In the Nordic church-yards in Middle Norway, probability of being a Saami increased. This concern­ 111 individuals could be used in the grouping. Of these ed trait V23, V24, V35 and H44. A presence of trait H7 only 58 were grouped as Nordic or possibly Nordic increased the probability of beeing of Nordic origin. (52.30fo). Thus the prognosis turned out to be most useful in distinguishing the Saami individuals in these Trait coefficient populations. A majority (24/37) of the individuals buried at Vard0 turned out to be of possibly Nordic H7 - 3.68 origin. V23 3.37 V24 14.78 A check was made on how the predictions were in­ V35 3.52 fluenced by the sex of the individual. Among the H44 3.74 women at the Saami church-yards, 145 of 154 were These traits were used to create a model. classified as Saamis or uncertain Saamis (94.lOfo ). The corresponding figures of men were 148 of 162 (91.30fo ). The figures of the church-yards in Middle Norway Discussion and comparison between the present genet­ were the following: Women, 25 of 50 were classified as ic imestigation and earlier anthropological stndies of Nordic or uncertain Nordic origin (50%) and of A model using the non-metric traits that were good at men, 33 of 59 (55.90fo). differentiating the Saan1i and Nordic populations The occurrence/absence of tori was useful in respectively was created (N = 146). This model was distinguishing these Saami and Nordic populations. It used for grouping a total of 475 individuals. In­ has, though, been known for a long time that popula­ dividuals not previously used for creating the model tions living in an arctic or subarctic surrounding show were especially valuable as their values were indepen­ high frequencies of tori of the palate. the upper and the dent of those building up the model. How an indi­ lower jaw (torus palatinus: 1,46,47,48,49) (torus man­ vidual was classified can be seen from figure 2. dibularis: 1,49,50,51,52,53).

Cimmrpo/4T Health 87 Jregren & Jsberg: Genetic composition in Saami Populations in Northern Norway 223

Table III shows the frequencies of these tori of some results indicate that genetic factors play a major role in populations previously studied. As several scientists determining the morphology of the mandibular are involved and as tori may develop to different size or torus." at somewhat different location the figures can not be discussed in detail. However, the general trend is ob­ vious. SUMMARY The aim of this investigation was to describe and Table Ill. Frequencies of tori in the maxilla, palate separate populations of different ethnic origin using and the mandible in human groups living .in the biological information from the cranium. Six Saami arctic or subarctic populations from Northern Norway formed the Saami Frequencies(%) of tori sample while two populations from Middle Norway constitute the Nordic sample. Another population Man- Population Maxillary Palatine dibular References from Northern Norway was of possible Nordic origin. The latter can be used for a test of possible environ­ Finland mental effects on populations of different ethnic af­ lnari Saamis 30 (48) lnari Saamis 40 (54) filiation living in an area with the same climate. The Norway materials are dated from to the 14th to the 19th cen­ Saam is 32.5 60.3 ( 1 I tury. Saa mis 41.1 68.6 64.4 lregren Statistical methods were used for evaluating the fre­ Nordic 17.0 38.0 ( 1 I Nordic quencies and combinations of minor morphological Heida I 2.1/8 5.6 8.8 lregren traits of the skull. Several (22/79) traits showed dif­ Nordic? Varde 5.7 16.7 lregren ferencies at the 50Je level (Chi-square tests) between the ethnic groups. Further. five non-metric traits were Greenland Eskimos 47.1 23.8 ( 1) found to be the best fonlifferentiating the Saami and Eskimos 80 (55) Nordic group respect:iVely. Three of these concerned Norsemen 60 (56) the presence of tori. As mentioned, it is an established Denmark fact that Saami populations show high frequencies of Nordic 13 (55) tori occurrence. Thus our results agree with those of Schreiner and others. High tori occurrences have been found in Mongoloid 146 individuals were used for constructing a model populations and much lower in Caucasoid popula­ to discriminate between the groups. The ethnic origin tions. This fact has suggested a genetic background of of a total of 475 individuals was predicted. Of the in­ the traits. Some familial studies regarding the man­ dividuals from the Saami church-yards 92.70/e were dibular torus support the hereditary hypothesis grouped as Saamis. Of the skeletal sample from Mid­ (33,41,42). Other scientists point out the environment dle Norway, 52.30/o were grouped as being of Nordic in a broad sense is the most important factor behind origin. New information about this ethnic groups is the occurrence of the traits. Mayhall & Mayhall (57) also given, as the frequencies of a large number of have found that groups of the same population when quasi-continuous morphological variants. living in different environments show different fre­ quencies of torus mandibularis. Further, different racial groups living in roughly the same environment REFERENCES have similar tori frequencies (58,59). l. Schreiner KE. Zur Osteologie der Lappen.Skr.Inst.f. Sellevold (60) in a recent study compared Greenland Sammenlign. Kulturforskn. B, Oslo 1935; XVIII(!): 292. 2. Lewin T, HedegArd B. 197la Human Biological Studies Eskimos with Norsemen living in Greenland. Selle­ Among Skolts and Other Lapps. A survey of earlier Iit­ vold's (60:p.572) results regarding the mandibular terature. In: I..ewin T. Introduction to the Biological char­ torus: "show that two different racial groups from acteristics of the Skolt Lapps. Proc Finnish Dent Soc roughly the same physical environment, both with 1971; 67: suppl 1: 63-70. high frequencies of occurrence of mandibular torus, 3. Lewin T, HedegArd R 1971b Longitudinal Studies on Age Changes in Craniofacial Dimensions of Skolt Lapps with possess different phenotypes of the trait. While the no Access to Dental Attention. In: Lewin. T. Introduc­ role of the environment cannot be disregarded as a fac­ tion to the Biological characteristics of the Skok Lapps. tor in determining the presence of the trait, the pRSent Proc Finnish Dent Soc 1971; 67: 161-170. 224 Anthropology and demography

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