The origin of the Lapps in the light of recent genetic studies

A.W. ERIKSSON, J. FELLMAN, H. FORSIUS, W. LEHMANN, T. LEWIN, and P. LUUKKA

The Lapps deviate in many genetic and external characters from other peoples of the Finno-Ugrian language group, such as and . The relation between Lappish and other Finno-Ugrian languages (including Finnish) is still controversial (7). A considerable proportion of the ances­ tors of the present-day Lapps are supposed to have been spread over almost the whole of and the northern parts of . Five hundred years ago considerable areas of northern and eastern Finland were inhabited mainly by Lapps (Figure 1). Today there are about 35,000 Lapps in the whole of Fennoscandia: 20,000 in , 10,000 in SWeden, 3,500 in Finland, and 1,500 on the in the . One of the reasons why the commune of Inari was selected for International Biological Program Human Adaptability studies (9) was that this region is inhabited by three l70 Genetic considerations

LAPP TERRITORY f'ZZ2] ~ 100 8 c '~;.. :970 AO ---!LC..tt!~!_

~>-T.tr--.< I

Figure 1 A comparison of Lapp territory, around 100 BC and 1970 AD rather different Lapp populations: the Skolt Lapps in· the eastern parts , the Fisher Lapps around , and the Mountain Lapps in the western parts of the cc:mmune · After World War II the Sevettijarvi Skol t Lapps were evacuated from the village of Suenjel and the Skol t Lapps from the regions south of the River Pasvik (Figure 1). Since. d World War II, the proportion of Lapps in Inari has decline markedly' although their absolute number has not decreased. For the calculations of gene frequencies, the four groups of Sevettijarvi Skolt, Nellim Skolt, Fisher, and Mountain Lapps have been used as representatives of pure, unmixed Lapps, with only those whose ancestors over the last three generations are at least 75 per cent aboriginal Lapps being accepted. More than 95 per cent of the ' GO per cent of the Fisher Lapps, and more than 50 per ce~t of the Mountain Lapps have been studied. The Utsjoki series is a random sample investigated in 1973.

PHYSICAL ANTHROPOLOGY OF LAPPS

In external characteristics the majority of the Lapps deViate fran the surrounain~ populations. In contrast with Genetic studies on Lapps 171

other Scandinavian peoples, who are among the tallest in the world, the Lapps are short, with legs that are short both relatively and absolutely. The Lapps are also among the most brachycephalic (round-headed) peoples in the world, with a cephalic index of up to 85. They have rather prominent cheek­ bones and a low facial height (11). The flattening of the face, the prominence of the cheekbones, and the reduction of the nasal prominence are not nearly so marked in Lapps as in Mongolian populations. The inner eyefold, covering the inner margin of the eye and responsible for the half-moon slant­ eye of many Asiatic and Amerindian peoples, is rare in Lapps.

Pigmentation

Some Lapps may have a light complexion in childhood with quite fair hair. However, the darkening of the hair during adolescence is much IOC>re intense than in otper Nordic peoples. A screening of hair pigmentation with "Fisher­ Saller' s Haarfarbentafel," using 30 different hair colour shades (Figure 2), shows that the Lapps, particularly the Finnish Mountain Lapps, have considerably darker hair than the Finns, who are one of the blondest peoples in the world (6). Eye colours have been determined using Martin-Saller's scale (Figure 3). The pigmentation of the irises and the eye grounds (fundi) is much stronger than among the other Nordic populations. AlIIDst 30 per cent of the Fisher Lapps but hardly 5 per cent of the Skolt Lapps have deep brown eyes (7 and 8). On average, no less than 40-60 per cent of Fin- nish Lapps have blue eyes. Reflectance spectrophotometry has shown that among Fin­ nish Lapps skin colour is not yellow, as in Mongolians (un­ published observations). The Mongol spot can be found in most children of the Mongolian race but is rare among those of Lappish and other European populations.

Hirsute Traits of Lapps The Lapps have soft and not unconononly wavy hair, in con­ trast to the coarse, straight hair of the Mongolian race. Microscopic studies show that the hair texture and hair pig­ mentation of the Skolt Lapps should be classified as Euro­ pean (Caucasian), not as Mongoloid (14). In its ample growth of hair, the white race resembles monkeys more than any other human race. Among Lapps, beard growth is rather sparse, and sideburns on the cheeks are often missing. Among the Lapps, body and axillary hair 172 Genetic considerations

HAIR COLOUR (HAARFARBENTAFEL FIS CHER-SALLER)

FINNS (NE FINLAND) MOUNTAIN LAPPS 30 no322 d',20-55y n • 9 7 d', 20 - 59 y

20

10

°lo F"·L M·O P·T U v w x y F-L M·O P-T U 40

FINNS (OULU) SKOLT LAPPS 30 n " 249 d', 20 - 59 y n = 7,1, d', 20 - 4 9 Y

20

10

F"· L M·O P. T U V W X y y Figure 2 A comparison of hair coloration in Finns and black)Lapps (F-L, blond; M-o, fair; P-T, brown; U-Y, brown to growth too is commonly sparse, as among the Mongoloi· d race.ng The occurrence of lniddle-phalangeal hair, at least ~ Lapps in Finland, is as high as among other Fennoscandian P<>PUlations (SWedes 70 per cent, NE Finns 80 per cent~s "oki Skolt females 75 per cent, Skolt males 95 per cent, U ~ Lapp females 69 per cent, Utsjoki Lapp males 74 per cen. ns (unpublished observations). The majority of the populatio Genetic studies on Lapps 173

% 30

SKOLT LAPPS 20 <60yd',n=202

10

20 LAPPS < 60y d', n = 6l 10

30 FINNISH MOUNTAIN 20 LAPPS < 60 y d', n= 88

10

lO

30 FINNS <60yd', n= 675 20

10

1 2 3 l 5 6 7 8 IRIS COLOUR TYPES (MARTIN -SALL ER'S SCALE)

Figure 3 A comparison of iris colour types (Martin Saller's scale) for Lapps and Finns

of Mongoloid affinity so far studied seem to have consider­ ably lower frequencies of individuals with middle-phalangeal hair. 174 Genetic considerations

Dermatoglyphics

The patterns of skin ridges of Skol t Lapps are mainly of European type (8) . The frequency of congenital dislocation of the hip among Lapps is 20-30 times as high as among other North-European populations.

MONOGENIC TRAITS

High Gene Frequency in Lapps

The majority of the Lapp populations so far studied in Fennoscandia have some genetic traits in common. In compari - son with the Scandinavians, the Lapps have very high fre­ quencies of sane markers. The most striking is the gene A2, which is about 2 to 5 times as high as in the surrounding populations (Table 1). The Lapps have the highest frequency of the A2 gene in the world. The rather high frequency of this gene in other Scandinavian populations has been inter­ preted as a sign of Lappish gene flow to the surrounding populations (4).

TABLE l the High and low gene frequencies in Lapps; the arrows indicate that gene frequencies resemble eastern ( ~ ) and western (~) traits

Lapps Other Scandinavians

High frequency in Lapps A2 25 (14-37) 5 NS Gml,13 15 (5-22) 8 1 2 (0-4) l? Inv 18 (13-26) 7 Acf' 50 (40-61) 25 PGM2 pC l 40 (21-55) 20 10 (2-19) 8 ADA2 12 (10-16) 7 lm 60 (58-80) 30 w (dry cerumen) 30 (18-37) 17

Low frequency in Lapps B 6 (0-32) 11 r Gc2 17 (6-29) 38 14 (2-33) 24 se 2 16 (12-21) 48 ~F 1 (0-2) 5 6 (2-9) 19 t (PTC) 35 (26-55) 50 Genetic studies on Lapps 175

Until now, it has been believed that the Ai gene had a very low frequency among Siberian North-Asiatic populations. However, a high frequency of the A2 gene has been found re­ cently in a north-Altayan group (Sukernik, personal communi­ cation). It will be very interesting to see what continued subtyping of the blood group A will show in the north­ Siberian populations. A high frequency of A2 in the northern USSR could be interpreted in two ways, either as an indica­ tion of a relationship with the Lapps, or as evidence that the A2 gene is of selective advantage in arctic and sub­ arctic regions. However, the A2 gene seems to be ccmpletely missing among the Ainu and the pure Eskimos. The Mongolian ga:mniaglobulin haplotype Gml ,13 has been found in all Finnish Lapp populations except Fisher Lapps. This Mongoloid trait has, however, also been found in other European populations, e.g. in Aland Islanders, Finns, and Maris (13). Figure 4 shows that among Europeans there is a distinct increase of the Gm(l) allotype frequency in the northern direction (south-north cline). Mediterranean populations, e.g. Yugoslavs and Greeks, have frequencies of 30 to 40 per cent, but in north-European populations frequencies are 60 to 70 per cent. The Lapps living in the Far North, the BO

70 EJ mTI 0 @~ 60 ~ l!i w 0 © 8 :::i © ® @©ii>22 ® '::: 50 >- <{ @ e @ __J ® 0 40 C0 C0 D LAPPS Q OTHER EUROPEANS

30

30 40 50 60 70 80 90 100% Gm (1) Figure 4 The relationship between latitude and frequency of occurrence of Gm(l) gene 176 Genetic considerations

Inari Fisher Lapps and the SWedish Lapps have frequencies in good agreement with an increasing frequency of the Gm{l) allotype with latitude. The other Lapp populations, however, deviate fran this rule, especially the Nellim Skolt Lapps, who have frequencies similar to those of Central Europeans· The advantage of having a Gm (1) constitution does not seem to be important, and in small isolated populations the effect of genetic drift, including founder effect, may be more illq:>Ortant than any selective factors. Among the Lapps the frequency of the gammaglobulin allele Invl is 13 to 28 per cent, i.e. 2 to 4 times as high as airong other Europeans. In other Finno-Ugrian populations, such as Finns and Maris, the frequency is the lowest in Europe, only 2 to 5 per cent (13). Among Finnish Lapps the inherited lipoprotein antigen allele Agx is about twice as high as in other Scandinavian populations (3). Mongoloid peoples seem to have a higher frequency of AgX than Caucasians. 2 The PGM1 allele seems to be another Lappish characteris­ tic· It has an average frequency about twice as high as among other Scandinavians and the highest among Europeans· The frequency of the PGM12 allele increases toward the north, both in Norway and in Finland (4). More studies are ~eeded to elucidate whether this increasing PGM 2 frequency in the northern direction should be in te :rpreted1 as a Lapp influence or as an effect of selection. The red cell acid phosphatase gene pc has a high fre­ quency an1mg Finnish Lapps. AlOOng Skol t Lapps and Fisher Lapps the pc frequency is the highest so far reported (13- 19 .pe7 cent)· The Pc allele is canpletely missing among all Asiatic Populations so far studied. This trait may there­ fore be int rp ted e re as a western gene. pk> ~t has been suggested that the acid phosphatase allele is of selective advantage in a tropical environment Cl)· Recent studie h · _ s ave shown, however that at least some cir CU!Dpolar p u1 ti _h ' Th op a ons have high !""' frequencies (4) • ~ f~equency of the adenosine deaminase gene ADA2 among the Fimush Lap · T<•- ps is the highest so far reported . .uuo;.e other Europe f - quency of the d ans' the Lapps have a rather low re ry ear wax type (Figure 5) • Th e frequency of 1 t p· ni. h La . ac ose malabsorption among adult :m- s PPs is the h" h (see Sah · l.g est so far reported among Europeans 1 et al.• this volume). Genetic studies on Lapps 177

1.0 N.CHINESE KOREANS MONGOLS .9 ·JAPANESE

.8

MONGOLOIDS . 7 - NAVAJO, SIOUX

- AINU 6

.5 - ESKIMOS (NWGr<"enlond) .I. - UTSJOKI LAPPS NELLIM SKOLT LAPPS SEVETTIJARVI SKOLT LAPPS .3 -MARI

FISHER LAPPS CAUCASOIDS .2 MOUNTAIN LAPPS WHITES .1 NE FINNS NEGROES U.S NEGROES

0 Figure 5 Frequencies of the allele for dry cerumen (w) in selected races

Low Gene Frequencies in Lapps Except for the marginal population, known as Skolts, the Lapps have a frequency of the B gene that is about a half of that found in surrounding populations. The frequency of Rhesus-negative individuals among wes­ tern Europeans is about 15 per cent, but among Lapps is only between 2 and 9 per cent. ABH non-secretors and Lewis(a+) subjects are only about one-third as frequent as among other Scandinavians (see Eskola et al. , this volume) . To exemplify what contrasting gene frequencies can be founa in small neighbouring isolates, we may mention that 2 the Gc gene frequency is extremely high in Nellim Skolt Lapps (33 per cent) with the lowest reported aioong Euro­ peans in their neighbours, the Fisher Lapps (2 per cent). As far as we know, there has not been any gene exchange be­ tween these Lapp populations in recent generations. 178 Genetic considerations

Among all the Lapp populations studied so far, the adenylate kinase gene AK2 is so extremely low that it may be supposed to have been extinct among the aboriginal Lapps· If genealogical studies can show that subjects with a rare allele are not pure Lapps and that the presence of this allele depends entirely on gene flow from surrounding popu­ lations, this gene frequency may be used as an indicator of gene flow from surrounding populations. Estimates based 2 on the frequency of AK and other rare genes, such as B or r, among Lapps suggest that the gene flow from the Finns to the Lapps in Inari has been between 20 and 40 per cent. Studies of the genetic polymorphism of the third compo­ nent of complement (C1 3) in serum have shown that the gene 1 c3F (C' 3 ) is considerably lower among Lapps than among other Scandinavians (2) • The frequency of PTC non-tasters is low among Lapps in the central Lapp region, while the proportion of PTC tas­ ters among the Skolt Lapps is the same as among other Euro­ peans (5).

CONCLUSICNS

In SOll\e morphological characteristics, the Lapps are differ­ ent from other Nordic populations. However, Swedes, Nor- weg· p· ians, inns, and Russians are among the tallest and least · ~igmented peoples in the world. The Lapps are not more pigmented than many Central-European populations and have a considerably lighter complexion than other long­ resident circumpolar populations. Some ronogenic traits of the Lapps suggest a trace of th~ Mongolian race· However, the Diego factor, Di (a+) , res­ tricted to the Mongolian race, has not been found in the LaEps. studied so far. The transferrin alleles TfBo-1 and ~ Chi are characteristic of Mongoloids and occur in fairly high frequenci · · Lapps (4). es in Finns and Estonians , but are rare among The Skolt Lapp de · . · t · f s viate in many hereditary characteris- ~cs r: other Lapp popUlations. In contrast to other r ppGcs 2 e Skol ts have high frequencies of the genes B, M, , , and t (PTc n -ta . . how that the Jna. . on sters) . There is evidence to s Lap d Jority of the ancestors (founders) of the Skolt ps escended from 1 . . . th century (lQ). on Y a few fanulies in the sixteen ab The Lapps have always been a small ethnic group, prob- ly never la:r:qer than t d domn ess must have had o ay . The founder effect and ran- a great influence on the gene pool. Genetic studies on Lapps 179

However, this, in combination with inbreeding (16) and selection, can hardly explain all the peculiar genetic characteristics common to the majority of the Lapp popula­ tions so far investigated. It is surprising that so many of the Lapp populations have such a unique "Lappish" gene pattern, for according to history and verbal tradition there has been a consider­ able gene flow from the nearby majority populations (see von Bonsdorff et al., this volume). The fact that the Lapps differ to such a high degree from other Nordic peoples in both monogenic markers and external characteristics suggests that in their ancestry they differed markedly from the present Finns, Swedes, and Norwegians. Certainly some of the morphological traits, such as stature and craniofacial dimensions, have been influenced by the hard conditions of the subarctic areas. The younger generations of Lapps have not such extreme somatometric characteristics (11). For the linguist it has been a problem why the Lapps speak a Finno-Ugrian language in spite of the fact that they deviate so markedly from Finns and Estonians. However, Baltic Finnish, spoken by Finns and Estonians, lacks the dual which is typical of Lappish, samoyedish, and Obugrish (7). The culture of the Lapps as a whole also differs sharply from that of the surrounding peoples, e.g. the troll-drtnn used to evoke ecstasy, skin-covered skis, and breeding. These factors indicate contact with peoples of the northern USSR. Certain physical features and osteological traits have been established as similar in Lapps and Samoyeds (12) . It remains uncertain whether the ancestors of the Lapps may have comprised relics of one or more proto-European Populations (15) and whether they may have had any contact With peoples of the cultures of the Mesolithic periods, about 8000 to 4500 BC, the cultures of Kunda, Suomusjarvi, and Kansa (Ruija). Recent studies have confirmed that the majority of the Lapp populations still have a distinct distribution of many genetic traits and deviate from the surrounding majori­ ty populations in some external characteristics. One is in­ clined to consider the Lapps, if not as a separate race, then at least as a population descended from an original stock. Some of the ancestral populations of the Lapps may have had some extreme gene frequencies, which today are Still reflected in the majority of Lapp populations. 180 Genetic considerations

To reach DOre definite conclusions about th: orig~n of the Lapps, more Lapps and neighbouring populations w~ll have to be studied. Of particular interest are the circum­ polar populations in the USSR,

ACKNOWLEDGMENTS

'Ibis work was aided by grants from the Nordiska Kultur­ fonden, the Finnish National Man and Biosphere Comni ttee' and the Deutsche Forschungsgemeinschaft.

SUMMARY

In ex>ntrast to the tall, blond, and doliocephalic Nordic people, the Lapps are relatively short and dark, and amonq the most brachycephalic people in the world. The majority .of the Lapp populations deviate from Finnish and other populations in northwestern Europe in having the highest noted frequencies of the blood group ~ene A2, high frequencies of the alleles Fya, PGM2, pc, AbA , and low fre­ 2 2 quencies of the alleles B, r, P1 , Jka, Lea, se, Gc , AK ' t (PTC non-tasters). Of the gammaglobulin allotypes, Lapps have essentially Caucasoid haplotypes, with a small portion of the Gml,13 haplotype characteristic of Mongoloid popula­ tions. :I'he frequency of the gene Invl in Finnish Lapps is the highest known among Europeans, and w (dry cerumen) is rare· The frequency of congenital dislocation of the hip is 20 to 30 times as high among Lapps as in other North­ European Populations. In spite of gene flow from the sur­ rounding majority populations and genetic drift and in­ breeding in the rather small and isolated groups, most Lapp populations show similarities in several distinct gene fre­ q~ncies · The inference is that the Lapps originated from different ancient peoples inhabiting northwestern Europe and at least some of these seem to have had some extreme gene frequencies that are still reflected in the present­ day Lapps.

REFERENCES

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3. Berg, K. and Eriksson, A. W., "Genetic marker systems in Arctic populations, V. The inherited Ag(x) sermn lipo­ protein antigen in Finnish Lapps," Human Hered., 23: 241-6 (1973) 4. Eriksson, A. W., "Genetic polymorphisms in Finno-Ugrian populations: Finns, Lapps and Maris," Israel J. med. Sci., 9: 1156-70 (1973) 5. Eriksson, A.W., Fellman, J., Forsius, H., and Lehmann, W., "Phenyl thiocarbarnide tasting ability among Lapps and Finns, " Hmnan Hered., 20: 623-30 (1970) 6. Forsius, H., Luukka, H., Lehmann, W., Fellman, J., and Eriksson, A. W. , "Ophthalrnogenetical studies on Skol t Lapps and Finns: corneal thickness, corneal refraction and iris pigmentation," Nord. Med., 84: 1559-61 (1970) 7. Itkonen, E., "Die Herkunft und Vorgeschichte der Lappen im Lichte der Sprachwissenschaft," Ural-Altaische Jb., 27: 32-44 (1955) 8. Lehmann, W., Jiirgens, H.w., Forsius, H., Eriksson, A.W., Haack, M., Junkelmann, B., Bahlmann, E., and Pape, G., "Ober das Hautleistensystem der Skoltlappen," Z. Morph. Anthrop., 62: 61-99 (1970) 9. Lewin, T. and Eriksson, A.W., "The Scandinavian Inter­ national Biological Program, Section for Human Adapta­ bility, IBP/HA, Scandinavian IBP/HA Investigations in 1967-1969," Arct. Anthrop. 7: 63-9 (1970) 10. Lewin, T., Rundgren, A., Forsius, H., and Eriksson, A.'W., "Demography of the Skolt Lapps in Northern Fin­ land," Finska TandlakSallsk.Fi.'Srhandl., 67: Suppl. 1: 24-38 (1971) 11. Lewin, T., Skrobak-Kaczynski, J., and Sigholrn, G., "Secular changes in craniofacial dimensions in a homo­ geneous population," Acta morph. neerl. -scand., 11: 289-319 (1973) 12. Schreiner, K.E., Zur Osteologie der Lappen (Oslo: A.W. Br~ger, 1935) 13. Steinberg, A.G., Tiilikainen, A., Eskola, M.-R., and Erikson, A. W., "Ganmaglobulin allotypes in Finnish Lapps, Finns, Aland Islanders, Maris (Cheremis) , and Greenland Eskimos," Am. J. hum. Genet., 26: 223-43 (1974) 14 · Stybalkowski, M., "Mikroskopische Untersuchungen an Querschnitten von Kopfhaaren der Sevettijarvi-Skolt­ Lappen," Anthrop. Anz., 33: 219-32 (1972) 15. Torgersen, J., Getz, B., and Simonsen, P., "Varanger­ funnene. I. Funn av menniskeskjeletter," Troms¢ Mus· Skr., 7: 1-28 (1959)