blood group frequencies in northern west

Blood Group Frequencies in Northern WestNew Guinea (West Irian)

L. E. NIJENHUIS,l A. C. VAN DER GUGTEN,2 H. DEN BUTTER,2AND J. W. DOELAND2

WITH THE TECHNICAL ASSISTANCE OF KARI WOLTHUIS-BRATLIEIAND ELIN KLUKHUHN-HOFLAND1

'Central Laboratory of the Netherlands Red Cross BloodTransfusion Service, Amsterdam (Prof. Dr. J. J. van Loghem, Director).

2Public Health Service, New Guinea.

INTRODUCTION AND MATERIALS

IN PREVIOUS PUBLICATIONS, the blood group frequencies of some populationsof the northern part of West New Guinea (West Irian) have been described(Nijenhuis and van der Hoeven, 1956; de Vries and Nijenhuis, 1960; Nijen-huis and de Vries, 1960; Nijenhuis, 1961d). During the two years precedingthe incorporation of West New Guinea into the Republic of Indonesia, wewere able to carry out further investigations, enabling a complete survey tobe made of the blood group frequencies of the peoples of the MamberamoRiver and along the coast east of the Mamberamo to Hollandia (Fig. 1).Blood samples were collected along the Mamberamo River and along the

coast of the subdivision* of Sarmi by A. C. van der Gugten and in theeastern part of this area by J. W. Doeland (Fig. 2). H. den Butter collectedthe samples along the coast of the subdivision of Genjem (Fig. 3).

The Mamberamo District

Collection of blood samples was made possible by the courtesy of thegovernment in placing at our disposal, from 6 to 11 July 1961, a ship with arefrigerator. The samples were sent by air to Amsterdam, and blood group-ing tests were performed between 19 and 21 July.

In Pionier-Bivak, the administrative center of the district, 40 samples weretaken from persons visiting the locality. They belonged to a number ofclosely related tribes in the surrounding area, which speak the same languageand intermarry. The group name of these tribes is Kawerabidja, individualtribes being the Soromadja, Bagudja, Bilasse, Itjara, and Namenawedja,among others.

Further north, along Lake Rombebai, blood samples were collected fromtwo population groups: 56 from the Bagussa, an original people of the

Received July 14, 1965.Supported in part by grants from the Netherlands Organization for Pure Research

(Z. W. 0.) and the Organization for Scientific Research of New Guinea (W. 0. N. G.).*The government in West New Guinea was divided into regional administrative units

called divisions. These were divided into a number of subdivisions, each of which con-sisted of a number of districts.

39

AMERICAN JOURNAL OF HUMAN GENETICS, VOL. 18, No. 1 (JANUARY), 1966

BLOOD GROUPS IN NEW GUINEA

"'XmiakD, '7

hollandia

FiG. 1. Map of West New Guinea. Scale, 1:12,800,000.

Mamberamo living in and around a village at the junction of the lake andthe river, and 20 from the Kaipeso tribe, which originated from the inlandof the Apauwer region. The coastal region between the Waim River and Tebais inhabited by populations who may be considered to form more or less anentity with the peoples of the Mamberamo and Apauwer river areas.Somewhat south of Pionier-Bivak, in a kitchen garden along the river, a

group of Baudji were encountered from the Baudji settlement at Lake TelagaBiru. Their language is quite different from the Mamberamo-Apauwer lan-guages, and they have connections with the tribes in the Waren area farinland. The Baudji are an extensive group, consisting of three different tribes,but only members of the Telaga Biru tribe have on various occasions beencontacted by Europeans. Twenty Baudji blood samples were collected.

Fifty-two samples were obtained from Teba and 22 from Joke, two villagesalong the coast of the Mamberamo district. Teba is inhabited by people,probably originating from Seroei, who speak the language of Geelvink Bay.They differ from the true Mamberamo populations in skin color and hairtype, and some aspects of their material culture are the same as in Seroei.The same may be true in Joke, though to a lesser extent.

Foreign influence is known to have occurred in the Mamberamo region inrelatively recent times. Chinese crocodile hunters frequently visited the Mam-beramo River, and the Telaga Biru and Indonesians hunted paradise birdsalong the Mamberamo River and the coast towards the administrative centerat Sarmi, i.e., the coastal areas of the Mamberamo and Apauwer districts.

The Coast of the Apauwer District Between the Waim and Ferkami RiversIndonesian hunters of paradise birds are known to have visited this coastal

region in recent times. A study of the blood group frequencies of the popula-tion of this region has been published (Nijenhuis, 1961d).

40

NIJENHUIS ET AL.

& Q kumamba islands I

baudji hi %'

lake telaga

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FIG. 2. Detailed map of the subdivision of Sarmi. Scale, 1:2,500,000.

FIG. 3. Detailed map of the coastal region of the subdivision of Genjem. Scale, 1:1,250,000.

The autochthonous population originally came from inland but has livedalong the coast for many generations. In language and culture, the peopleform an entity with the inland population between the two rivers.

The Coast of the Districts of Sarmi and Sarmi East CoastBlood group frequencies of the coast of Sarmi and of the western part of

the East Coast district have already been published for Amsira, Sarmi, Ku-

41


mamba Islands, Island Wakde, and Takar (Nijenhuis, 1961d). The peopleof the Kumamba Islands (220 inhabitants) came originally from the islandof Biak and from a village east of the town of Sarmi. The town of Sarmi has350 inhabitants of Papuan origin and an Indonesian element which originates,at least in part, from former prisoners. (Before the second world war, therewas a prison for Indonesians in Sarmi, and some intermarriage between thetwo groups has occurred.) The island of Wakde was the original home of thepopulation of the village bearing the same name on the mainland oppositethe island (500 inhabitants). Takar (500 inhabitants) has a population re-lated to that of the Tor district, but they have a relatively long traditionas people of the coast.

In the east of the East Coast district of the Sarmi subdivision a few bloodsamples were collected between 25 and 27 August 1961 (35 samples fromArmopa and 25 from Mawesdai), and blood grouping tests were performedon 12 and 13 September 1961. All along this coast, marriage within thevillage of origin is the rule, although it is not invariable. Some intermarriagebetween localities does occur.

The Coastal Region Between Tarfia and Humbold Bay

This region is inhabited by four major language groups:1. The population of Tarfia, composed of two groups, one originating from

Berap in Nimboran and the other with its own language related to that ofKaptiau, which is situated 40 kilometers along the coast to the west.

2. The Muris group, inhabiting the coastal region between Ambora andMuris. Their common origin is probably a coastal village near the more re-cently established administrative center of Demta. Some intermarriage mayhave occurred with the population of Bukisi and, especially in Muris Ketjiland Muris Besar, with people from the region of Berap. The main contactof this group with the inhabitants of the western areas (Tarfia) has beenhostile.

3. The group between Bukisi and Jongsu Ketjil, which may be dividedinto three subgroups:

(a) The Bukisi group, occupying the region between Bukisi and Sorojena,probably originated from Bukisi and migrated eastward along the coast. Itis not quite clear if some elements from group (c) (Tanah Merah) havebeen absorbed into the population, but it clearly contains Kamtuk elements.

(b) The population of Kendate, which seems to have come from theKamtuk-Gressie village of Jansu. The village dialect is still closely related tothe Kamtuk language, but, in their contacts with other populations of theBukisi-Jongsu group, the inhabitants use the "great" language. For example,the local word for "coconut" in Kendate is kim, as in Kamtuk. In the rest ofthe area, extending from Bukisi to Ormu, the Humbold Bay, and Lake Sen-tani, this word is to'ko. "Foot" is berente in Kamtuk and Kendate, oto in theregion between Bukisi and Jongsu.

(c) The villages between Tablanusu and Jongsu Ketjil, where the TanahMerah language is spoken. The main components of the language come froman original population in the region of Cape Tanah Merah; other influences

42

NIJENHUIS ET AL.

migrated along the shore from Humbold Bay or even Australian New Guinea(trading connections with these areas still exist) and via Lake Sentani fromthe southeast. The three groups have fused completely, and it seems prob-able that no mixing with western groups has occurred. People of Tablasufa(between Depapre and Cape Tanah Merah) still own kitchen gardens aroundLake Sentani, and it seems likely that people from Gressie, a region be-tween Lake Sentani and Nimboran, must have been settled between Tabla-sufa and Dormena.

4. The Ormu group, composed of an original local element and two groupsof immigrants: one from the south via Lake Sentani and one from the eastvia the coast of Humbold Bay.

Before the second world war, Tarfia was mainly endogamous. There weresome contacts with Kaptiau and occasionally with the Muris area. In thelatter area, there was contact between the four villages and some with Berapand Tarfia. The villages between Bukisi and Sorojena had contacts with thosebetween Bukisi and Dormena and with the region between Dormena andthe western part of Lake Sentani (the Maribu-Sabron region) but none withthe Kamtuk.The people of Kendate had contact with the villages along the shore be-

tween Dormena and Bukisi and with the Kamtuk. Depapre and Tablanusu,on the other hand, had relations with the villages between Dormena andBukisi and with the Maribu-Sabron area. Contacts also existed betweenTablasufa, the area between jongsu and Bukisi, Humbold Bay, and theMaribu-Sabron area. Dormena and other villages at Cape Tanah Merah alsohad contact with this area and along the shore between Ormu and Sufa. Thepeople of Jongsu, however, had contact only with villages along the coastbetween Ormu and Sufa, those of Ormu along the coast between Tablastufaand the Humbold Bay, and with the people of Sentani.Throughout the area, as far west as Tarfia, the canoes of the coastal people

closely resemble those of Lake Sentani but differ from those of the inhabitantsof Geelvink Bay. Each canoe bears a mnukaper-ahut, a figure of a bird or afish, and each family possesses its own sign; these ornaments are not foundfurther west than Tarfia.

Marriage customs are also similar within the same area and are identicalwith those along Humbold Bay and Lake Sentani. They differ from those inNimboran.A blood group survey of the coastal population between Tarfia and Dor-

mena was carried out in May, 1961. Fourteen blood samples were collectedin Tarfia, 27 in the Muris region (8 Ambora, 5 Jaugapsa, 3 Muris Ketjil, and11 Muris Besar) and 137 in the Bukisi-jongsu group comprised of 60 Sorojena-Bukisi, 10 Bukisi, 10 Andowatufu, 9 Wanja, 9 Nasuna, 13 Kantu Milena, 2Bona, 6 Demoi, 1 Sorojena, 20 Kendate, and 57 Tanah Merah (20 Tablanusu,16 Depapre, and 21 Dormena). The material is representative of a numberof populations totaling somewhat over 4,000.

METHODS

The blood samples were collected as follows: Four drops of blood were

43


taken from a finger tip into tubes with rubber stoppers, each containing 0.5ml of a sterile trisodium citrate-glucose solution with "cialit" 1:20,000 (fromHoechst Farbwerke, Frankfurt/M, Germany) as a preservative. All bloodsamples were stored in a thermos vacuum container with ice water. Freshice was added at every opportunity, and the containers were shipped as soonas possible by air to Amsterdam, where blood grouping tests were carriedout. In all cases, blood was collected from adults only, and in the Tarfia-Dormena area an attempt was made to exclude close relationship betweenthe donors.The samples were investigated by tube tests with anti-A, anti-B, anti-A+B,

anti-M, anti-N, anti-S, anti-C, anti-c, anti-D, and anti-E antisera and withtwo specimens of anti-P1 sera. Seventy-five blood samples, all of groups Aand 0 from the series taken in the area between Tarfia and Dormena, weretested with anti-I; all M-positive samples of this series were tested with asecond anti-M serum known to react negatively with the M, factor.As Papuan blood samples taken in the same way are known to show weak,

very weak, or even negative reactions with saline-agglutinating anti-D sera(Nijenhuis, 1961d), the test was modified as follows in order to obtain clear-cut agglutinations: After incubation for one hour, one drop of 20% bovinealbumin was added to the tubes, which were then incubated for a further30 minutes before the reactions were read. In other cases, very weakly re-acting red cell suspensions were retested with another anti-D serum or byan enzyme technique using bromelin (series Armopa and Mawesday). Testsfor the E and c antigens were in all cases performed by the bromelin tech-nique, and in the series from Armopa and Mawesday a second (saline ag-glutinating) anti-c serum was used as a control, in order to verify that noneof the blood samples were nonspecifically agglutinated by the bromeiin.Some of the tests for S. which were mostly carried out with an incomplete

anti-S serum by the indirect antiglobulin test, were controlled by retestingthe positive samples with a second (saline-agglutinating) anti-S serum inorder to check the reliability of the incomplete anti-S. All the tests per-formed in parallel gave identical results, including the tests with two anti-P1 sera. In the case of the Pionier-Bivak series, some doubt remains as tothe reliability of the S tests, since for these samples an anti-S serum was usedwhich is known to produce false-positive reactions with some samples afterprolonged storage.Gene frequencies were calculated by the usual formulae, as summarized

by Mourant (1954), and these preliminary values were corrected by recalcu-lation using the gene-count method of Ceppellini, Siniscalco, and Smith(1955).

RESULTS

The results of the investigations are summarized in Table 1. In general,the figures are given apart for all distinct populations, and the data areadded only in cases of no statistical differences between the observed fre-quencies in related populations or in populations of adjacent regions. These

44

NIJENHUIS ET AL.

combinations of local or regional series are shown in Table 1 in two in-stances: total Mamberamo (Pionier-Bivak, Bagussa, and Kaipeso) and totalBukisi-Jongsu.The results of the statistical tests are summarized in Table 2. Wherever

possible, x2 testing has been performed; if, however, any of the expectednumbers was below 5, P values have been calculated directly from 2 X 2tables.The observed frequency of the S factor in Pionier Bivak was found to be

significantly higher than in the two other Mamberamo populations. How-ever, as previously indicated, there is a possibility that the S test on thePionier-Bivak series was not completely reliable, and therefore the threeMamberamo series have been combined.The calculated gene-frequencies are given in Table 3, together with re-

sults previously reported for the northern areas of New Guinea. The ex-pected numbers of the various blood group phenotypes were calculated fromthe gene frequencies. In no case were significant differences found betweenthe expected and observed numbers.No weak, very weak, or negative reactions were observed in the Tarfia-

Dormena series, but a few samples from the Mamberamo district reactedweakly with saline anti-D, even after the addition of albumin as previouslydescribed (3 samples in the Pionier-Bivak series, none from Kaipeso, nonefrom Joke, and 2 from Teba). No albumin was added when testing theBaudji and BagLissa series with saline anti-D, and in the latter particularly ahigh frequency of weak and very weak reactions were observed. The Armopa-Mawesday series contained two very weakly reacting samples, which gavestrong reactions when another anti-D serum was used by the enzymetechnique.The I antigen, which has been found to have very high frequencies in

Europeans (negative reactions being very rare in adults), proved to bepresent in all 74 Papuan samples tested for this factor. 'm, could not bedemonstrated in any of the 29 N-positive blood samples from the Tarfia-Dormena series. The antigen had been found previously in three out ofnine MN bloods from Merauke (Nijenhuis, 1961d), in four out of six MNsamples from Muyu Papuans, and in one out of five MN samples from theStar Mountains (Nijenhuis, 1961a). It could not be demonstrated in 18 M-positive samples from the Mimika district, in eight MN Dani samples fromTiom, in 31 M and MN Dani samples from the Great Balim Valley, or in 73M-positive blood samples from the Sarmi subdivision, taken mainly from thecoastal populations of the Apauwer district (Nijenhuis, 1961d).

DISCUSSION

The gene frequencies of the present series, of the series from the Sarmisubdivision previously examined, and of other series in northern West NewGuinea and the western part of East New Guinea, are listed in Table 3. Inview of the small numbers of most of the local or regional series, it wasthought advisable to carry the figures to two decimals only. The Apauwer

45o


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TABLE 2. SIGNIFICANCE TESTS ON DIFFERENCES BETWEEN POPULATIONSPopulations Factors x'[df 1 P

Pionier-Bivak vs. Bagussa S factor 8.60 [1] <0.01+ Kaipeso

Mamberamo vs. Baudji Rh (CCDee vs. other types) <0.004

Mamberamo vs. Joke Rh (CCDee vs. other types) 0.005ABO (0-A-B, AB) 4.61 [21 0.10

Mamberamo vs. Teba ABO (0-A-B, AB) 10.97 [2] <0.01MNS, (MMS+, MNS+ vs. 9.6 [1] <0.01NNS+)

Joke vs. Teba MNS (MMS+, MNS+ vs. 0.091NNS+)

Rh (CCDee vs. other types) 0.057P1 2.10 [1] 0.1-0.2

Armopa vs. Mawesdai ABO (0-A-B, AB) 6.84 [21 <0.05

Tarfia vs. Muris MN vs. NN 0.00061

Kendate vs. Tanah Merah ABO (0-A-B, AB) 4.31 [2] 0.1-0.2+ Bukisi groups P1 2.53 [1] 0.1-0.2

Rh (CCDee vs. other types) 1.71 [1] 0.1-0.2

Muris vs. Bukisi/Jongsu ABO (O vs. non-O) 2.84 [1] 0.1-0.2S factor 0.13MM, MN vs. NN 0.07

coastal region was subdivided into a main group, containing the villages ofWebro, Arbais, Niwerawar, Aruswar, Wari, and Martewar, and two minorgroups-Samarokema and Ferkami river-in order to prevent disturbance ofthe relative homogeneity of the main group by the figures for the minorgroups, which probably deviate, although this has not been clearly demon-strated. The differences between the main group and Samarokema are sta-tistically significant for the P1 frequencies (x21l = 4.65, P < 0.05), but nosignificant differences were found between the main group and Ferkami.Between the Martewar Webro group and Amsira, the differences for ABOand Rh are more marked.

In general, the ABO frequencies in the area vary greatly, as they dothroughout the whole of New Guinea. The Mamberamo district also appearsto have a relatively high frequency of Ms, comparable to the frequencies ofSorong and Biak and to those along the south coast of the eastern part of theAustralian half of the island (Dunn et al., 1956; see also the reviews of Sim-mons et al., 1961, and Nijenhuis, 1961d).The S frequency is comparable to that of Sorong and Biak; MS and NS

genes are both present. The original inhabitants of the Mamberamo-Apauwerarea have very high frequencies of CDe; lower frequencies of CDe andhigher cDE and cDe occur in the minor series of different or mixed origin(Baudji, Joke). The same very high frequency of CDe is found in the coastal

48

NIJENHUIS ET AL.

region of Apauwer. Here, however, the Ms frequency is low and the S fre-quency high; MS has a higher frequency than NS.Along the coast between Sarmi and Tarfia, gene frequencies vary con-

siderably, confirming that endogamy is the rule in the villages of this area.In general, Ms ranges from relatively high to very high, and S is low in com-parison to the coast of Apauwer; it is probable that both MS and NS arepresent. cDE and cDe are somewhat more frequent than in Apauwer.The Muris series is in good agreement with the Nimboran series inland

from Muris: Ms is very low, S is absent, and the Rh frequencies agree rea-sonably well.The Jongsu-Bukisi series and Sentani have slightly higher Ms frequencies,

and S. probably exclusively as NS, is present though with a low percentage.Rh frequencies are comparable with those of Muris and Nimboran (Jongsu-Bukisi) or diverge owing to a relatively low frequency of CDe (Sentani).Sekoli, some distance away around the western end of Lake Sentani, hasabout the same Ms frequency but lacks S and has a much higher percentageCDe.The interpretation of the occurrence and frequencies of MS, Ms, and NS

in the pattern of MNS frequencies in New Guinea and the surrounding areasis particularly interesting. It appears that over an extensive area in WestNew Guinea the S factor is absent. The Muris series, Nimboran, and Sekolifall within this area, the boundary of which is found westward along thecoast from Bukisi and south from Bukisi via Lake Sentani. West of Tarfia,along the coast to Apauwer, relatively low or very low S frequencies werefound. The area with no or low S extends to the south, into the CentralMountains of West New Guinea. S was found to be absent in the StarMountains (Nijenhuis, 1961a), in the Dani of Mulia (Simmons et al., 1961),and in the Dani of the Balim area (Nijenhuis, 1961d). In another Dani seriesfrom the great Balim valley, only two S-positive specimens were found in atotal of 266 (Nijenhuis, 1961d). In the Ekari (= Kapauka), Graydon et al.(1958) observed 0.7% S genes, and Groves et al. (1958) reported 2% in thesame region (Wissel Lakes). In the westernmost areas of the highlands ofAustralian New Guinea, S gene frequencies so far investigated are above 10%(Macintosh, Walsh, and Kooptzoff, 1958), so that the eastern boundary ofthe area with no S must run somewhere between the Star Mountains andthe regions studied by Macintosh, Walsh, and Kooptzoff. A little to thesouth of the Star Mountains, S was observed in the Muyu (9% of S genes)and somewhat farther to the southwest in the Jair (16%) and the Mandobo(29-20%) (Nijenhuis, 1961a).

In the Marinds on the south coast of New Guinea, the frequency of the Sgene was found to be 10% (Nijenhuis, 1961d); along the coast to the north-west in Asmat, 7.5% (Nijenhuis, Beckers, and de Vries, 1960); and in theMimika region, 5-11% (Nijenhuis, 1961d). These observations lead one tosuppose that the population of New Guinea originally lacked S (Nijenhuis,1961a, b). S will thus have been introduced from outside, and natural selec-tion will have raised its frequency to higher percentages than those found in

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Z;0

, iDaH ,f

._

t- 5¢t:t ;= NC

51


the surrounding areas (Indonesia, Melanesia, Micronesia, and Australia). Itis improbable that the population of New Guinea consists of two components,one with and one without S, in view of the typical blood group frequencypatterns throughout the island, which has the highest N and CDe frequenciesin the world and 100% of Duffya positives.

Further evidence for the selection hypothesis is obtained by examiningthe segregation of S, comparing the Ms frequencies in the various popula-tions, and the MNS frequencies in the Papuan populations with those ofsurrounding areas (Nijenhuis, 1961c, d). MS is rare in New Guinea; it hasbeen demonstrated along the southern coast of eastern Australian NewGuinea (Sanger, Walsh, and Kay, 1951) and in the Sepik region of north-western Australian New Guinea (Simmons et al., 1965). In some other cases,the MNS+/MNS- and NNS+/NNS- ratios make it probable that the MSgene is present, although it has not been directly demonstrated as MMS+.In this connection, as far as West New Guinea is concerned, it is probablethat MS occurs in Mimika in the southwest (Nijenhuis, 1961d) and perhapsin Biak and Sorong. In most of the populations of New Guinea, S, if presentat all, occurs as NS, while MS is completely absent.

If the evidence for the presence of MS is accepted, its frequency musteverywhere be much lower than that of NS, except in the populations of theSarmi area. The geographical situation of Sarmi is such that it is bordered ontwo sides by the low S areas (south and east). If the S gene reached NewGuinea from the east, it would have penetrated from east to west; thus itmust have stopped in the north and center near the boundary between Westand East New Guinea. In the south, it continued its route, at least as faras Mimika. If Sarmi received its S genes from the same source, S will havecrossed to the northeast of the island at its narrowest point, leaving theWissel lakes with hardly any S, and passing on to the northwest coast ofGeelvink Bay, the islands, and the extreme west. This does not explain, how-ever, why MS is higher than NS in Sarmi. A more probable explanation isthat the S factor in Sarmi, and probably also on Biak and in the extremewest of New Guinea, originates from direct mixing of the local populationswith Indonesians. This in fact agrees well with the high M frequencies inSorong, Biak, Mamberamo district, Sarmi district, and Sarmi East Coastdistrict. However, Ms frequencies are very high in Indonesia, but MS andNS relatively low, and the highest MS and NS frequencies in Sarmi havebeen found mainly in the coastal region of Apauwer, with very low Msfrequencies.

In this connection, it must be mentioned that the Rhesus gene Rz (CDE)has been observed in West Guinea only on five occasions. Its local occur-rence in the Star Mountains (Nijenhuis, 1961a) is difficult to explain; theother four cases were in coastal populations, as in Australian New Guinea.This would mean that Rz is an imported gene. It has been found in Biaktogether with high Ms, in the Apauwer coastal region (where M'S has beendemonstrated), in Mimika (where the occurrence of MS is very probable),and in the Marinds in the region of Merauke. This could be an indication that

52

NIJENHUIS ET AL.

the occurrence of MS, even if not accompanied by a high frequency of Ms,is due to "infection" of the population with genes from overseas.

It is reasonable to think of Polynesia as the origin of the S genes in thegreater part of New Guinea. Simmons et al. (1951) found 0.8% of MS genesand 5.5% of NS genes in Polynesians (Maoris), which may explain why S inNew Guinea occurs mainly as NS and why MS occurs in East New Guinea,especially in the coastal areas and not inland. Very ancient contacts withPolynesians are also a reasonable explanation of the cultivation throughoutNew Guinea of some native American plants such as sweet potatoes andtobacco.The areas of West New Guinea with high Ms frequencies or with MS and

NS, however, must have received these genes from other sources, probablyIndonesian. As previously mentioned, there have been recent contacts withIndonesians (and Chinese), but much earlier contacts with Indonesia areknown to have existed with West Irian in historical times. In southeast Asia,the frequency of the S factor is relatively low; in the majority of observa-tions, MS appears to have a higher frequency than NS (Malaya: Polunin andSneath, 1953; China: Miller et al., 1951; North Borneo: Graydon et al., 1952,and Sweetman et al., 1951. See Mourant, 1954). However, in eastern In-donesia, which must be considered as an area with a mixed population, Nijen-huis (1961d) observed only a very low frequency of S, exclusively in theform of NS.One explanation of the MNS frequencies in southeast Asia and Oceania

may be that the genetical composition of the Polynesians is in a state ofequilibrium when Ms and Ns are present in high and about equal proportions,with a low percentage of NS and very few MS genes. In Indonesia, the gen-eral genetical background of the population differs from that of Polynesia,the MNS system reaching equilibrium only when there is a preponderanceof Ms over Ns, low MS frequencies, and an even lower level of NS.

In Papuans, considered as a third "race" between the Indonesians andPolynesians (and Australians), S was absent but was introduced in smallamounts. Against the general genetical background of the Papuans, veryhigh Ns was in balance with low Ms; once the S gene was introduced, a newpattern of equilibrium developed, in which Ms is still low to very low (genefrequencies in most of the pure Papuan populations being 10% or less), NSreaches frequencies higher than in the surrounding "races" (up to 20 to 25%and sometimes even higher), and Ns retains the highest frequency.

It therefore appears that the Papuan genome is a more favorable geneticalbackground for the S factor than that of Polynesians and Indonesians. Thatit is indeed the S factor in general which is relatively favorable in Papuansand not only the combination NS is shown in the Apauwer coastal popula-tions of the Sarmi subdivision, where the introduction of small numbers ofIndonesian genes has caused an explosive increase not only of NS but evenmore of MS. As the Indonesian genome is relatively unfavorable to MS andNS, it is clear that any widespread introduction of Indonesian genes intoPapuan populations, altering the favorable genome, will result in an equilib-

53


rium in which MS and NS have much lower frequencies than after intro-duction into a wholly Papuan genome. This may be the explanation of theS frequencies in Sorong, Biak, the Mamberamo district, Sarmi district, andSarmi East Coast district, which are in general much lower than those ofother Papuan populations where S is found, i.e. eastern and southern NewGuinea. At the same time, wholesale mixing with Indonesians has changedthe genetical background of the population in such a way that against thebackground of the new genome Ms is no longer an unfavorable combinationand its frequency is raised to 20% or higher. It is not certain that the increasein Ms is the result of the direct introduction of Indonesian genes. The amountof foreign genes introduced may in fact have been larger or smaller than isnow indicated by the higher frequencies. The change in the genome of thepopulation would have produced a new pattern of equilibrium, and theobserved MNS frequencies would be a reflection of this equilibrium moreas an indirect than a direct result of the racial mixing.

It is worth noting that variation in the frequencies of genetical charactersbetween populations in a given race or within a given area are not neces-sarily the reversible effects of chance. Even regional populations may reachan equilibrium, which may be different from those of adjacent regions, dif-ferences between gene frequencies representing minor differences in thegenetical background of the local population. These minor differences maybe caused by chance (genetic drift) or by the assimilation of small numbersof foreign genes in the gene pool, with the result that such assimilation mayhave quite different effects when spread over more generations or broughtabout suddenly in one or two generations.The MNS frequencies in the Jongsu-Bukisi and Sentani regions may per-

haps be explained by the advancing penetration of the S gene into areaswhere it was previously absent. S genes are still being introduced in smallnumbers from eastern areas, for instance (either directly or indirectly) fromthe Sepik area. S frequencies are still very low, and it may be supposed thatgenetical equilibrium has not yet been reached, so that the frequency willcontinue to increase in the next few generations. As is shown in Table 3,MacLennan, Kooptzoff, and Walsh (1930) and Simmons et al. (1965) dem-onstrated the presence of MS in the Sepik area. In some of the componentseries, the total M frequency is high and the frequency of Ms sometimeshigher than 10%. Dunn et al. (1956) found 10% of Ms in the Sepik district,and in the districts of Madang and Marobe further east they found about thesame frequencies: 11.5% and 10%. It seems therefore reasonable that thepopulations with high Ms and with some MS represent the results of mixingon a regional scale. As a whole it would appear that the M gene frequenciesalong the north coast of New Guinea generally reach 10%. Since M frequenciesunder 10% are more the rule than the exception in the central areas of NewGuinea and also along the southern coast between Mimika and Kikori, itmay be supposed that almost the entire northern coast of New Guinea hasbeen influenced in varying degrees by non-Papuan elements.

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NIJENHUIS ET AL.

SUMMARY

A study has been carried out on the ABO, MNS, Rh (CcDE), and P1 bloodgroup frequencies in West New Guinea along the Mamberamo River andalong the coast between the Mamberamo and the Humbold Bay. Blood groupsand gene frequencies of the various populations in the area have been listed.No M variants or I-negatives were found. Weak reactions with anti-D insaline and also after addition of albumin were observed in some samples. Inthe western part of the Sarmi subdivision in particular, the observed CDe fre-quencies are very high.

In the Mamberamo dlistrict, the Apauwer coastal region, and the coastalregion of Demta, some regional populations appear more or less homogeneous.In the eastern coastal region of the Sarmi subdivision, however, considerablelocal variations are the rule. In the Sarmi subdivision, M frequencies are high,indicating mixing of the population with foreign elements, most probably In-donesians. S is absent from the Muris population, as from the Nimborans andthe inhabitants of the Sekoli region; east and northeast of this area, NS isfound but with low frequencies.The S factor in New Guinea is supposed to have originated from foreign

influences from the east, natural selection having subsequently caused it toincrease to frequencies which are higher than in other southeastern Asiaticand Pacific populations. In the Sarmi region, unlike most of the other Papuantribes, MS sometimes occurs more often than NS, indicating a difference inthe origin of S in the area; this observation agrees very well with the hy-pothesis that the action of natural selection against the MNS genes dependsupon the composition of the genome of the populations. There is some evi-dence from the literature that along the whole of the north coast of NewGuinea M frequencies are higher than is usual for pure Papuan populations.

ACKNOWLEDGMENTS

Our thanks are due to Dr. A. E. Mourant for his valuable comments.

REFERENCESCEPPELLINI, R., SINISCALCO, NI., AND SMITH, C. A. B. 1955. The estimation of gene

frequencies in a random-mating population. Ann. Hum. Genet. (Lond.) 20: 97-115.DUNN, D., KOOPTZOFF, O., PRICE, A. V. G., AND WALSH, R. J. 1956. The blood groups of

a third series of New Guinea natives from Port Moresby. Oceania 27: 56-63.GRAYDON, J. J., SEMPLE, N. NI., SIMMONS, R. T., AND FRANKEN, S. 1958. Blood groups

in pygmies of the Wissel Lakes in Netherlands New Guinea, with anthropologicalnotes by H. J. T. Bijlmer. Amer. J. Phy.s. Anthrop. 16: 149-172.

GROVES, NI., PRICE, A. V. G., WALSH, R. J., AND KOopPTZOFF, 0. 1958. Blood groups ofthe NIotu and Koita peoples. Oceania 28: 222-238.

MACINTOSH, N. W. G., WALSH, R. J., AND KOOPTZOFF, 0. 1958. The blood groups of thenative inhabitants of the Western Highlands. New Guinea. Oceania 28:173-189.

MACLENNAN, R., KOOPTZOFF, O., AND WALSH, R. J. 1960. A survey of the blood groupsand haemoglogin values in the Sepik river district. Oceania 30: 305-312.

MOURANT, A. E. 1954. The Distribution of the Human Blood Groups. Oxford: Blackwell.NIJENHUIS, L. E. 1961a. Blood group frequencies in the Upper Digul and Muyu districts

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and in the Star Mountains of Netherlands New Guinea. Nova Guinea, 1 (n. s.): 1-14.

NIJENHUIS, L. E. 1961b. Investigation of blood groups as a key to the relationship ofPapuans. Trop. Geogr. Med. (Amsterdam) 13: 267-272.

NIJENHUIS, L. E. 1961c. Some aspects of blood groups and natural selection. In Proceed-irngs of the Second International Congress of Human Genetics, Vol. II. Rome: IstitutoG. Mendel, pp. 903-906.

NIJENHUIS, L. E. 1961d. Blood Group Frequencies in the Netherlands, Curapao, Surinamand New Guinea. A Study in Population Genetics. Thesis, Amsterdam. (Availablefrom the Library, Central Laboratory of the NRC Blood Transfusion Service, POB200, Amsterdam.)

NIJENHUiS, L. E., BECKERS, T., AND DE VRIES, J. L. 1960. Blood group frequencies in NewGuinea. II. The blood groups of the Asmat Papuans. Amer. J. Phys. Anthrop. 18:189-191.

NIJENHUIS, L. E., AND DE VRIES, J. L. 1960. Blood group frequencies in New Guinea. III.Blood groups of the Nimboran Papuans. Amer. J. Phys. Anthrop. 18: 193-196.

NIJENHUIS, L. E., AND VAN DER HOEVEN, J. A. 1956. Blood group frequencies in Papuansfrom Biak (Isles of Schouten). Vox Sang. 1: 241-249.

SANGER, R., WALSH, R. J., AND KAY, M. P. 1951. Blood types of natives of Australia andNew Guinea. Amer. J. Phys. Anthrop. 9: 71-78.

SIMMONS, R. T., GRAYDON, J. J., GAJDUSEK, D. C., SCHOFIELD, F. D., AND PARKINSON, A.D. 1965. Blood group genetic data from the Maprik area of the Sepik district, NewGuinea. Oceania 35:218-232.

SIMMONS, R. T., GRAYDON, J. J., ZIGAS, V., BAKER, L. L., AND GAJDUSEK, D. C. 1961.Studies on Kuru. V. A blood group genetical survey of the Kuru region and otherparts of Papua, New Guinea. Amer. J. Trop. Med. Hyg. 10: 639-664.

SIMMONS, R. T., GRAYDON, J. J., SEMPLE, N. M., AND TAYLOR, C. N. D. 1951. Blood, tasteand secretion: a genetical survey in Maoris. Med. J. Australia 1: 423-431.

DE VRIES, J. L., AND NIJENHUIS, L. E. 1960. Blood group frequencies in New Guinea. I.The Sentani Papuans. Amer. J. Phys. Anthrop. 18: 125-130.

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blood group frequencies in northern west

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