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1. Introduction
Most mammals stop producing lactase, the enzyme that hydrolyzes lactose, at weaning; thereafter, they are intolerant of milk (Johnson, Kretchmer, & Simoons, 1974). The cessation of lactase production at weaning and adult lactose malabsorption also predominate among humans, particularly peoples of Asian and African descent. However, some populations, particularly those of northern European and Scandanavian descent, exhibit high frequencies of lactase persistence. These people can continue to drink milk throughout life (Flatz, 1989, Flatz & Rotthauwe, 1971, Simoons, 1978, Simoons, 1982). Considerable information has recently become available about the physiological mechanisms and molecular bases of the adult lactase polymorphism (reviewed by Swallow, 2003, Swallow & Hollox, 2000).
However, the historical ecology and distribution of adult lactose absorption (LA) and malabsorption (LM) have, for decades, intrigued geneticists (e.g., Cavalli-Sforza, 1973, Flatz, 1987, Flatz, 1989, Flatz & Rotthauwe, 1971) and nutritional ecologists (Durham, 1991, Simoons, 1970, Simoons, 1971, Simoons, 1982). Genetic evidence indicates that the lactase polymorphism arose recently (5,000–10,000 years ago) and spread rapidly due to strong positive selection (Bersaglieri et al., 2004, Hollox et al., 2001). Today, lactase persistence occurs far north and south of the equator in Africa and Asia, far from its putative areas of origin (Flatz, 1987, Hollox & Swallow, 2002). Puzzlingly, in some parts of Africa and the Middle East, lactase-persistent populations have lived side-by-side with populations of lactose malabsorbers throughout recorded history (Flatz, 1987, Flatz, 1989, Simoons, 1978, Swallow, 2003).
The leading hypothesis to explain the geographical distribution of adult LA and LM is that nutritional advantages of lactose digestion confer a selective advantage on lactase-persistent phenotypes in areas where milk is reliably available throughout adult life, that is, where dairying is practiced regularly. This “culture–historical hypothesis” (McCracken, 1971, Simoons, 1970) has received considerable support (Durham, 1991, Hollox et al., 2001, Simoons, 1978, Simoons, 2001, Swallow, 2003), including, most recently, from (1) a maximum likelihood analysis that used genetic and linguistic trees to control for the effects of relatedness among subject populations (Holden & Mace, 1997) and (2) a study of single-nucleotide polymorphisms covering 3.2 Mb around the lactase gene, which indicated that alleles associated with lactase persistence have been subject to strong selection within the past several thousand years (i.e., in the setting of dairy farming: Bersaglieri et al., 2004).
The culture–historical hypothesis does not address the important antecedent question: Why is dairying practiced in some areas but not in others? Obviously, the geographical occurrence of dairying depends, proximally, on cultural traditions (McCracken, 1971, Simoons, 1970). We hypothesize that, ultimately, it depends on ecological factors that affect whether milk-producing ungulates, particularly cattle, can be raised in a given area. For example, extreme climates could make it impossible to keep dairy herds outdoors year-round, especially where forage is sparse, such as in extremely cold or hot climates (e.g., tundra, deserts, or rain forests). Moreover, in certain locales, endemic pathogens may debilitate or kill the animals, thereby making herding uneconomical, or even hazardous if the disease is transmissible to humans. Indeed, dairy herding is rare in Africa within the range of the tse-tse fly, which is the major vector of sleeping sickness (Simoons, 1982, Smith, 1992).
We evaluated this ecological “dairying barrier” hypothesis by synthesizing information on the frequencies of adult LA and LM among indigenous African, Asian, and European populations. Parallel information from indigenous peoples of North and South America and Oceania is not available. We also compiled data on latitude, climate (temperature), and the historical distributions (before 1900) of communicable and potentially fatal diseases of cattle. Relationships among these variables and lactase phenotypes were examined statistically.
2. Methods
We gathered and synthesized all published data on the frequencies of primary adult LA and LM throughout the world. Data in previous compilations by Flatz (1989), Holden and Mace (1997), Simoons, 1970, Simoons, 2001, and Swallow and Hollox (2000) were verified from original sources and augmented using online databases and searches of the primary literature. Our entire data set, which includes LA/LM information from 270 populations and their associated references, is presented as an electronic appendix (Appendix A) to this paper, to facilitate use by future investigators. We believe that it is the most comprehensive compilation of adult LA/LM information currently available.
For analyses, we subsampled the appendix database (Appendix A) to identify populations that met five criteria of accuracy, completeness, and adequacy of sample sizes: (i) n=≥20 subjects; (ii) subjects were ≥10 years old (i.e., weaned); (iii) subject population was not of “mixed” origin; (iv) subject population lived in its country of origin (i.e., nonimmigrants and nonnomadic); and (v) the country of origin was in Europe, Asia, or Africa. For each population that met all five criteria, we determined the latitude and mean annual temperature using the city where each study took place (or the closest major city) as the point of reference. Data were analyzed using Pearson correlations, partial correlations, and multiple regressions in Statview 2. Before analysis, LM frequency data were transformed (arcsine square root) to achieve normality.
Domesticated, milk-producing ungulates include goats, sheep, camels, and cattle; the latter are, by far, the most common sources of milk (Simoons, 1970, Simoons, 1971). We synthesized information on the historical geographical occurrence of communicable, deadly diseases of cattle, based on an exhaustive search of the veterinary literature, for each of the countries represented by the populations in our final sample. We used only sources that included a dated map showing the distribution of each disease and included only diseases whose geographical distribution, prior to 1900, was confirmed by ≥2 independent sources. Thus, we took into careful account the changes in the distribution of each disease due to recent spread (since 1900) and eradication efforts (generally beginning in the 1960s). We included in our analyses every disease for which it was possible to obtain appropriate and reliable information.
3. Results
LM data that fulfilled our five criteria of accuracy and completeness were available from 91 populations in 39 countries, ranging latitudinally from southern Africa to southern Greenland (Table 1). Overall, adult LM predominated (Fig. 1). In our sample, the mean frequency of LM was 61±30% (±S.D.; n=9940 subjects), with a range of 2% (Denmark) to 100% (Zambia; Fig. 2).
| Table 1.
Mean adult lactose malabsorption frequencies in 39 Eurasian countries (n=number of subjects), and information on latitude, temperature, number of deadly endemic cattle diseases found there historically (pre-1900), and sources of LM data |
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Fig. 1. Frequencies of primary adult LM among 91 indigenous populations (n=9940 people) from 39 countries in Africa, Asia, and Europe (see Table 1). |
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Fig. 2. Distribution of primary adult LM among 91 populations (n=9940 people; see Table 1) plus 9 nomadic, low-latitude, lactose-absorbing populations (n=1077 people; these nomads were analyzed separately). |
Historical (pre-1900) distributions of nine deadly cattle pathogens could be determined accurately (American Geographical Society of New York, 1951, Cliff & Haggett, 1988, Odend'hal, 1983, Rodenwaldt, 1952, Scott, 1981, World Anthrax Data Site, 2003). These diseases were anthrax, malaria, sleeping sickness, cholera, heartwater, Rift Valley fever, brucellosis, rinderpest, and theileriosis. The first four also are transmissible, and potentially fatal, to humans.
Our data revealed a significant negative correlation between adult LM frequencies and latitudes (Fig. 3a), with two extreme outliers (i.e., z scores >2.0): Siberia (61° N, in present day Russia) and Greenland (67° N). There was a significant, positive correlation between LM frequencies and mean temperatures (Fig. 3b), with Siberia and Greenland again appearing as statistical outliers. There also was a significant, positive correlation between LM frequencies and historical presence of deadly cattle diseases (Fig. 3c), again with the same two statistically identified outliers.
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Fig. 3. Occurrence of primary adult LM in relation to climate and historical occurrence of cattle diseases in 91 populations from 39 countries (see Table 1). (a) LM frequency vs. latitude. The negative correlation was highly significant whether the two statistically identified outliers, Siberia and Greenland (z scores >2.0), were included (r=−.56, p<.0002) or excluded (r=−.70, p<.0001). (b) LM frequency vs. mean annual temperature. The positive correlation was significant whether the two statistically identified outliers, Siberia and Greenland, were included (r=.42, p<.0086) or excluded (r=.61, p<.0001). (c) LM frequency vs. historical (before 1900) occurrence of nine cattle diseases (anthrax, brucellosis, cholera, heartwater, malaria, Rift Valley fever, rinderpest, sleeping sickness, and theileriosis). The positive correlation was significant whether the two statistically identified outliers, Siberia and Greenland, were included (r=.54, p<.0004) or excluded (r=.71, p<.0001). |
Multiple regression analyses indicated that adult LM frequencies were well predicted by latitude, temperature, and diseases together, whether Siberia and Greenland were included [r=.59, F(3,31)=6.363, p<.0015] or excluded [r=.74, F(3,29)=13.15, p<.0001]. Partial correlation analyses revealed that the numbers of diseases had an independent, significant effect on LM frequencies (t=2.14, p<.03), even when latitude and temperature were held constant statistically, whereas neither latitude (t=−.85, p=.40) nor temperature (t=.55, p=.58) significantly affected LM frequencies when diseases were held constant.
4. Discussion
Humans are the only mammals that regularly drink milk as adults. However, populations differ markedly in their physiological capabilities of digesting lactose (Fig. 1). Frequencies of adult LM in populations from Eurasia and Africa decrease with increasing latitude (Fig. 2) and increase with increasing temperature and, especially, with numbers of deadly cattle diseases that were present before 1900 (Table 1, Fig. 3). The implication is that harsh climates and dangerous diseases negatively impact dairy herding, thus also geographically restricting the availability of milk.
Populations from northern Siberia and Greenland are exceptions to the latitudinal trend: Although they live far north of the equator (>60° N), they are predominantly malabsorbers. In light of our dairying barrier hypothesis, both exceptions are instructive because neither cattle nor transmissable cattle diseases can thrive in such extreme climates. As a result, those populations did not predictably have access to fresh milk. Although the now-extinct Greenland Norse population attempted to maintain dairy herds, the animals and their owners always teetered on the brink of starvation (Simoons, 2001) and died out in <500 years. The latter-day Greenland Eskimos (the population in our database), which have persisted much longer, do not keep dairy cattle, and their diet is devoid of milk or dairy products (Simoons, 2001).
Previous investigators (Bayoumi et al., 1982, Durham, 1991, Flatz, 1989, Hussein & Ezzilarab, 1994, Simoons, 1978, Swallow, 2003) identified 13 additional outlier populations, but at the other end of the latitudinal spectrum. These peoples live at low latitudes in Africa and the mid-East, but they are predominantly lactose absorbers as adults (i.e., lactase persistent). Nine of these populations have been sampled adequately to be included in our database and analyses (Fig. 2). Among them, LM frequencies are indeed low (mean=35±25%), and milk and dairy products make up a significant fraction of their diet, raising the question of how these people surmounted ecological barriers to dairying.
The most likely explanation is nomadism. All 13 populations have historically been migratory pastoralists that inhabited borders between major climatic zones in Africa (Mali, Rwanda, Uganda, Congo, Egypt, and Sudan) and the mid-East (Jordan, Saudi Arabia; Griffiths, 1972, Johnson, 1969, Smith, 1992, Swallow, 2003). Seasonal movements of these peoples coincide with the advance and retreat of the intertropical (wind) convergence zone (the ITCZ: Waliser, 2003), enabling them to locate suitable cattle forage and avoid extreme temperatures year-round (Hayward & Ogentoyinbo, 1987, Kraus & Businger, 1994). Moreover, these pastoralists typically maintain small herds, spread their cattle out, and keep them moving (Johnson, 1969), all of which should reduce pathogen transmission rates.
In present-day Africa, some sedentary populations maintain domesticated cattle, particularly among the subequatorial Bantu tribes. Although few have been adequately sampled, those that have are predominantly lactose malabsorbers (Durham, 1991; Fig. 2). These patterns seem to run counter to both the culture–historical and dairying barrier hypotheses. However, these populations traditionally did not maintain dairy cattle or have access to fresh milk. Dairy herding in subequatorial Africa is a recent (20th century) phenomenon, facilitated by control or eradication of the most devastating cattle diseases and introduction of new breeds of cattle that are tolerant of extreme heat and draught (Coetzer, Thompson, & Tustin, 1994). Moreover, even today, adults in these populations rarely drink fresh milk, instead consuming fermented milk products, such as yogurt and cheese, which have minimal lactose and thus do not require lactase to digest (Durham, 1991).
Our dairying barrier hypothesis is complementary, and not an alternative to, the culture–historical hypothesis (McCracken, 1971, Simoons, 1970), which posits that the mutations that maintain lactase into adulthood are positively selected only where dairying flourishes (Bersaglieri et al., 2004, Swallow, 2003). Results of our analyses imply that adults of Asian and African descent typically are lactose malabsorbers because dairy herding was precluded from their ancestral homes by ecological factors. Adult lactose malabsorbers or their recent ancestors were sedentary agriculturalists or hunter gatherers who lived in places that were not conducive to safe and economical maintenance of dairy herds, due especially to lethal pathogens, extreme climates or both. The historical distribution of dairying, and the resulting distribution of the adult LM phenotype, thus lend support to the assertion of Diamond (1999) that “the different historical trajectories of Africa and Europe stem ultimately from differences in their environments.”
Acknowledgments
We thank R. Booker, H. K. Reeve, S. Myles, R. J. Safran, J. S. Shellman, an anonymous reviewer, and members of the Animal Behavior Lunch Bunch at Cornell University for constructive comments. Financial support was provided by the College of Arts & Sciences and the Agricultural Experiment Station through the Hatch Grant Program (c/o D. J. Decker) at Cornell, and the Howard Hughes Medical Research Foundation (c/o L. E. Southard).
Appendix A.
 |
Index | Population | Country | Lat. | N/S | LM | Ages | #subj. | Reference | |
|
1 | Afghan–mixed urban | Afghanistan | 34.3 | N | 0.77 | 18–60 | 34 | Rahimi et al. 1976 | |
|
2 | Afghan–Pashtun | Afghanistan | 34.3 | N | 0.79 | 18–60 | 71 | Rahimi et al. 1976 | |
|
3 | Afghan–Hazara | Afghanistan | 34.3 | N | 0.8 | 18–60 | 10 | Rahimi et al. 1976 | |
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4 | Afghan–Tajik | Afghanistan | 34.3 | N | 0.82 | 18–60 | 79 | Rahimi et al. 1976 | |
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5 | Afgan–Pasha-I | Afghanistan | 38.5 | N | 0.87 | 18–60 | 60 | Rahimi et al. 1976 | |
|
6 | Afghan–Uzbek | Afghanistan | 34.3 | N | 1 | 18–60 | 16 | Rahimi et al. 1976 | |
|
7 | Indians and Eskimos | Alaska | 61.1 | N | 0.94 | adults | 36 | Duncan and Scott 1972 | |
|
8 | Australian whites | Australia | 33.53 | S | 0 | adults | 23 | Bolin and Davis 1969 | |
|
9 | Chinese in Australia | Australia | 33.53 | S | 0.56 | 3–34 | 34 | Bolin and Davis 1970b | |
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10 | Indians in Australia | Australia | 0.8 | adults | 5 | Bolin and Davis 1969 | | ||
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11 | Australian Aborigines | Australia | 17.2 | S | 0.84 | 15–75 | 45 | Brand et al. 1983 | |
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12 | Chinese in Australia | Australia | 22.2 | N | 0.9 | adults | 30 | Bolin and Davis 1969 | |
|
13 | New Guineans | Australia | 1 | adults | 8 | Bolin and Davis 1969 | | ||
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14 | Austria (west) | Austria | 47.2 | N | 0.15 | adults | 166 | Rosenkranz et al. 1982 | |
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15 | Austria (east) | Austria | 47.2 | N | 0.25 | adults | 181 | Rosenkranz et al. 1982 | |
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16 | Aymara (Bolivia)-children | Bolivia | 15.3 | S | 0.774 | 11–15 | 31 | Balanza and Taboada 1985 | |
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17 | Herero | Botswana | 0.9 | ≥15 | 10 | Currie et al. 1978 | | ||
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18 | ‡hua Bushmen | Botswana | 24.45 | S | 0.91 | adults | 22 | Nurse and Jenkins 1974 | |
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19 | Brazil–caucasoid | Brazil | 22.5 | S | 0.45 | 20–52 | 40 | Seva-Ereira et al. 1983 | |
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20 | Brazil–Negroid | Brazil | 22.5 | S | 0.85 | 20–52 | 20 | Seva-Ereira et al. 1983 | |
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21 | Brazil–Mongaloid | Brazil | 22.5 | S | 1 | 20–52 | 20 | Seva-Ereira et al. 1983 | |
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22 | British | Britain | 51.5 | N | 0.05 | 11–88 | 75 | Ho et al. 1982 | |
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23 | Jews in Britain | Britain | 51.3 | N | 0.8 | adults | 10 | Neal 1968 | |
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24 | Greek | Cypriots in Britain | Britain | 35.1 | N 0.88 | ad + child. | 17 | Mehta and Latham 1977 | |
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25 | Indians and pakastanis | Britain | 0.93 | adults | 16 | Neal 1968 | | ||
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26 | Bantu | Cameroon | 6 | N | 1 | adults | 6 | Elliott et al. 1973 | |
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27 | Czechs in Canada | Canada | 49 | N | 0.18 | adults | 17 | Leichter 1972 | |
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28 | Poles in Canada | Canada | 0.29 | adults | 21 | Leichter 1972 | | ||
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29 | Punjabis | Canada | 31 | N | 0.33 | adults | 9 | Murthy and Haworth 1970 | |
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30 | Jews, Canadian and American | Canada | 49.2 | 0.69 | adults | 32 | Leichter 1971 | | |
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31 | Indians in Canada | Canada | 0.93 | adults | 15 | Murthy and Haworth 1970 | | ||
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32 | Indians | Canada, w.coast | 52 | N | 0.63 | adoles. | 30 | Leichter and Lee 1971 | |
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33 | China–Kazakh | China | 43.5 | N | 0.76 | 16–28 | 195 | Yongfa et al. 1984 | |
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34 | China–Mongols | China | 40.5 | N | 0.88 | 17–46 | 198 | Yongfa et al. 1984 | |
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35 | Han (chinese) - general | China | 39.5 | N | 0.92 | 17–35 | 248 | Yongfa et al. 1984 | |
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36 | Chami Indians | Columbia | 5 | N | 1 | adults | 24 | Alzate et al. 1969 | |
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37 | Tussi in Congo | Congo | 1 | S | 0 | adults | 15 | Elliott et al. 1973 | |
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38 | Caucasian in Congo | Congo | 0.2 | adults | 10 | Elliott et al. 1973 | | ||
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39 | Danes | Denmark | 55.4 | N | 0.02 | adults | 670 | Gudmand-Huyer et al. 1969 | |
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40 | Danes | Denmark | 56.2 | N | 0.03 | adults | 91 | Busk et al. 1975 | |
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41 | North Sinai Egyptians (Nomadic) | Egypt | 30 | N | 0.11 | 72 | Hussein and Ezzilarab 1994 | | |
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42 | New Valley Egyptians | Egypt | 24 | N | 0.51 | 100 | Hussein and Ezzilarab 1994 | | |
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43 | Egypt–South Nile | Egypt | 26.1 | N | 0.6 | 14–34 | 85 | Hussein et al. 1982 | |
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44 | Egypt–Urban | Egypt | 30 | N | 0.67 | 14–34 | 67 | Hussein et al. 1982 | |
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45 | Egypt–suez canal | Egypt | 30 | N | 0.69 | 14–34 | 16 | Hussein et al. 1982 | |
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46 | Egypt–Nile | Delta | Egypt | 30.6 | N | 0.73 | 14–34 | 291 Hussein et al. 1982 | |
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47 | Egypt–North Nile | Egypt | 29.2 | N | 0.85 | 14–34 | 111 | Hussein et al. 1982 | |
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48 | Egyptian fellahin | Egypt | 30 | N | 0.93 | 16–75 | 14 | Halsted et al. 1969 | |
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49 | Ethiopian infants | Ethiopia | 9 | N | 0.61 | <1 yr | 26 | Habte et al. 1973 | |
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50 | Ethiopians/Eritreans | Ethiopia | 9 | N | 0.9 | 7–13 | 58 | Habte and Hjalmarsson 1973 | |
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51 | Ethiopian children | Ethiopia | 9 | N | 0.9 | 7–13 | 157 | Habte et al. 1973 | |
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52 | Fijians | Fiji | 18.1 | S | 1 | adults | 12 | Masarei et al. 1972 | |
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53 | Finns | Finland | 60.2 | N | 0.06 | 7–15 | 129 | Launiala et al. 1971; Sahi et al. 1972 | |
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54 | Swedes in Finland | Finland | 59.2 | N | 0.08 | adults | 91 | Sahi 1974 | |
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55 | Finns | Finland | 60.2 | N | 0.17 | adults | 159 | Jussila et al. 1970 | |
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56 | Finns | Finland | 60 | N | 0.18 | 50–70 | 134 | Jussila 1969 | |
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57 | French | France | 48.5 | N | 0.07 | adults | 14 | Gouin et al. 1972 | |
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58 | French–north | France | 48.5 | N | 0.23 | 18–21 | 62 | Cuddenec et al. 1982 | |
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59 | French–South | France | 43.4 | N | 0.42 | 20–75 | 55 | O'Morain et al. 1978 | |
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60 | French–south | France | 43.4 | N | 0.5 | 18–21 | 16 | Cuddenec et al. 1982 | |
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61 | Bantu | Gabon | 0.4 | N | 0.6 | adults | 20 | Gendrel et al. 1989 | |
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62 | Germany–North | Germany | 53.3 | N | 0.06 | adults | 100 | Flatz et al. 1982 | |
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63 | Germany–NW | Germany | 51.2 | N | 0.088 | adults | 341 | Flatz et al. 1982 | |
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64 | Germany–south | Germany | 48.1 | N | 0.136 | adults | 221 | Flatz et al. 1982 | |
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65 | Germany–west | Germany | 52.2 | N | 0.137 | adults | 182 | Flatz et al. 1982 | |
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66 | Germans from cent. Europe | Germany | 50.4 | N | 0.15 | adults | 55 | Rotthauwe et al. 1972 | |
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67 | Germany–East | Germany | 52.3 | N | 0.224 | adults | 246 | Flatz et al. 1982 | |
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68 | Germany–SW | Germany | 49 | N | 0.235 | adults | 136 | Flatz et al. 1982 | |
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69 | Arabs from Jordan,Syria,Saudi Arabia,Egypt,Iraq,Tunisia | Germany | 50.4 | N | 0.81 | adults | 26 | Rotthauwe et al. 1971 | |
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70 | Children | Ghana | 5.3 | N | 0.73 | children | 100 | White and Latham 1973 | |
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71 | Greeks, | mainland | Greece 37.6 | N | 0.38 | adults | 16 | Spanidou and Petrakis 1972 | |
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72 | Greeks, mainland | Greece | 37.6 | N | 0.45 | adults | 600 | Kanaghinis et.el. 1974 | |
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73 | Greeks | Greece | 37.6 | N | 0.54 | 7–13 | 24 | Doxiadis and Papageorgiadis 197 | |
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74 | Greek Cretans | Greece | 35.2 | N | 0.56 | adults | 50 | Kanaghinis et el. 1974 | |
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75 | Greek Cypriots | Greece | 35.1 | N | 0.67 | adults | 50 | Kanaghinis et.el. 1974 | |
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76 | Greeks | Greece | 37.6 | N | 0.67 | 7–13 | 82 | Kattamis et al. 1973 | |
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77 | Greenland Eskimos/nw eur.mixed | Greenland | 64.1 | N | 0.14 | adults | 7 | Gudmand-Hoyer and Jarnum 1969 | |
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78 | Greenland Eskimos/nw eur.mixed | Greenland | 64.1 | N | 0.5 | adults | 4 | Asp et al. 1975 | |
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79 | Eskimos | Greenland | 69.2 | N | 0.84 | 81 | Gudmand-Hoyer et al. 1973 | | |
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80 | Eskimos | Greenland | 64.1 | N | 0.85 | adults | 13 | Asp et al. 1975 | |
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81 | Eskimos | Greenland | 64.1 | N | 0.88 | adults | 25 | Gudmand-Huyer et al. 1973 | |
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82 | Eskimos, greenland | Greenland | 64.1 | N | 0.94 | 18–60 | 19 | Asp et al. 1975 | |
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83 | Hungarians–west | Hungary | 47.3 | N | 0.28 | 17–39 | 100 | Czeizel et al. 1983 | |
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84 | Hungarians–east | Hungary | 47.3 | N | 0.29 | 17–39 | 70 | Czeizel et al. 1983 | |
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85 | Hungary–Matyo | Hungary | 47.5 | N | 0.37 | 16–54 | 172 | Czeizel et al. 1983 | |
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86 | Hungarians–mixed | Hungary | 47.3 | N | 0.41 | 17–39 | 262 | Czeizel et al. 1983 | |
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87 | Hungarians–NE | Hungary | 47.3 | N | 0.42 | 17–39 | 103 | Czeizel et al. 1983 | |
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88 | Hungary–Romai (Gypsies) | Hungary | 47.6 | N | 0.56 | 16–47 | 113 | Czeizel et al. 1983 | |
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89 | “Mohajirs” (mixed pkastani/dravidians) | India | 24.5 | N | 0.2 | adults | 15 | Rab and Baseer 1976 | |
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90 | Indians in Bombay | India | 18.6 | N | 0.24 | adults | 17 | Desai et al. 1967 | |
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91 | Indians in Hyderabad (Deccan) | India | 17.2 | N | 0.61 | adults | 18 | Reddy and Pershad 1972 | |
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92 | Indians in Bombay | India | 18.6 | N | 0.64 | adults | 100 | Desai et al. 1970 | |
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93 | Iranians | Iran | 35.4 | N | 0.86 | 20–25 | 21 | Sadre and Karbasi 1979 | |
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94 | Iraqi Jews | Iraq | 15 | N | 0.84 | 17–65 | 38 | Gilat et al. 1970 | |
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95 | Irish | Ireland | 53.2 | N | 0.04 | 16–68 | 50 | Fielding et al. 1981 | |
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96 | Jews in Israel | Israel | 32 | N | 0.54 | 8–46 | 50 | Gilat et al. 1974 | |
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97 | Jews in Israel | Israel | 32 | N | 0.6 | adults | 58 | Gilat et al. 1973 | |
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98 | Jews in Israel | Israel | 31.5 | N | 0.61 | adults | 93 | Rozen and Shafrir 1968 | |
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99 | Jews in israel, summary of following: | Israel | 32 | N | 0.71 | 17–70 | 215 | Gilat et al. 1970 | |
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100 | Sephardi, other | Israel | 32 | N | 0.72 | 17–69 | 36 | Gilat et al. 1970 | |
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101 | Ashkenazi | Israel | 32 | N | 0.79 | 20–70 | 53 | Gilat et al. 1970 | |
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102 | Arab villagers in Israel | Israel | 32 | N | 0.81 | adults | 67 | Gilat et al. 1971 | |
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103 | Jews, other orientals | Israel | 33 | N | 0.85 | 24–64 | 20 | Gilat et.al 1970 | |
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104 | Sicilians | Italy | 37.3 | N | 0.29 | adults | 100 | Burgio et al. 1984 | |
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105 | Italians – northern | Italy | 45.4 | N | 0.49 | adults | 208 | Burgio et al. 1984 | |
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106 | Italians | Italy | 45.3 | N | 0.8 | adults | 20 | Zuccato et al. 1983 | |
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107 | Sardinians–mts | Italy | 40.3 | N | 0.81 | 20–51 | 38 | Meloni et al. 1998 | |
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108 | Italians, Neapolitans | Italy | 40.5 | N | 0.84 | 27–70 | 37 | Rinaldi et al. 1984 | |
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109 | Sardinians–lowlands | Italy | 39.4 | N | 0.85 | 20–55 | 47 | Meloni et al. 1998 | |
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110 | Sardinians–north | Italy | 40.4 | N | 0.86 | adults | 50 | Meloni et al. 2001 | |
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111 | Sardinians–mts | Italy | 40.1 | N | 0.88 | 19–59 | 53 | Meloni et al. 1998 | |
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112 | Italians in Naples | Italy | 40.5 | N | 1 | adults | 9 | De Ritis et al. 1970 | |
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113 | Japanese | Japan | 40.34 | N | 0.73 | adults | 40 | Yoshida et al. 1975 | |
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114 | Jordanians–bedouins | Jordan | ? | ? | 0.24 | 17–46 | 162 | Hijazi et al. 1983 | |
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115 | Jordanians–urban | Jordan | 31.4 | N | 0.75 | 18–33 | 148 | Hijazi et al. 1983 | |
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116 | Jordanian Arabs | Jordan | 31.4 | N | 0.77 | adults | 56 | Snook et al. 1976 | |
|
117 | Kenyans (bantu agricultural tribes) | Kenya | 1.17 | S | 0.73 | 5–15 | 71 | Pieters and Van Rens 1973 | |
|
118 | Lebanese | Lebanon | 33.5 | N | 0.78 | 17–43 | 74 | Nasrallah 1979 | |
|
119 | Niger–nomads (Tuareg valley) | Mali | ? | ? | 0.127 | adults | 118 | Flatz et al. 1986b | |
|
120 | Mexican Mestizos | Mexico | 19.2 | N | 0.74 | 18–72 | 100 | Lisker et al. 1974 | |
|
121 | N. African Sephardi | Morocco | 32 | N | 0.63 | 19–65 | 32 | Gilat et al. 1970 | |
|
122 | !Kung Bushmen | Namibia | 19.6 | S | 0.92 | adults | 40 | Jenkins et al. 1974 | |
|
123 | Herero | Namibia | ? | ? | 0.978 | ≥11 | 46 | Currie et al. 1978 | |
|
124 | New guineans | New Guinea | 3.5 | S | 0.75 | adults | 32 | Arnhold et al. 1981 | |
|
125 | Massim(NW Guinea) | New Guinea | 10.2 | S | 0.83 | adults? | 35 | Gibney et al. 1981 | |
|
126 | Anglo–saxons in Nigeria | Nigeria | ? | 0.13 | adults | 8 | Kretchmer et al. 1971 | | |
|
127 | Europeans in Nigeria | Nigeria | ? | 0.22 | ≥3 | 9 | Ransome-Kuti et al. 1975 | | |
|
128 | Yoruba/European | Nigeria | 6.3 | N | 0.44 | ≥3 | 43 | Ransome-Kuti et al. 1975 | |
|
129 | Hausa/Fulani | Nigeria | 7.2 | N | 0.6 | adults | 15 | Olatunboson and Adadevoh 1971 | |
|
130 | Fulani (fulani/hausa), urban | Nigeria | 10.5 | N | 0.71 | adults | 24 | Kretchmer et al. 1971 | |
|
131 | Hausa | Nigeria | 12 | N | 0.76 | ad.+child | 17 | Kretchmer et al. 1971 | |
|
132 | Ibo | Nigeria | 7.2 | N | 0.82 | adults | 11 | Olatunboson and Adadevoh 1971 | |
|
133 | Yoruba | Nigeria | 7.22 | N | 0.84 | adults | 48 | Olatunboson and Adadevoh 1971 | |
|
134 | Yoruba | Nigeria | 6.3 | N | 0.98 | ≥4 | 41 | Kretchmer et al. 1971 | |
|
135 | S. Nigerians (mostly) | Nigeria | 7.2 | N | 1 | adults | 9 | Olatunboson and Adadevoh 1971 | |
|
136 | Yoruba | Nigeria | 6.3 | N | 1 | ≥3 | 11 | Ransome-Kuti et al. 1975 | |
|
137 | Punjabis | Pakistan | 24.5 | N | 0 | adults | 9 | Rab and Baseer 1976 | |
|
138 | Sindhis | Pakistan | 24.5 | N | 0 | adults | 12 | Rab and Baseer 1976 | |
|
139 | Baloochis | Pakistan | 24.5 | N | 0 | adults | 4 | Rab and Baseer 1976 | |
|
140 | Pathans | Pakistan | 24.5 | N | 0 | adults | 15 | Rab and Baseer 1976 | |
|
141 | Pakistani | Pakistan | 33.4 | N | 0.516 | adults | 66 | Abbas 1984 | |
|
142 | Pakistani–Sindh | Pakistan | 25.2 | N | 0.58 | 18–48 | 33 | Ahmad and Flatz 1984 | |
|
143 | Pakastani–punjab | Pakistan | 31 | N | 0.59 | 18–48 | 322 | Ahmad and Flatz 1984 | |
|
144 | Pakistani–Baluchistan | Pakistan | 28 | N | 0.62 | 18–48 | 32 | Ahmad and Flatz 1984 | |
|
145 | Kashmir (pakistan) | Pakistan | 34 | N | 0.7 | 18–48 | 27 | Ahmad and Flatz 1984 | |
|
146 | Peruvian students | Peru | 12 | S | 0.63 | 21–29 | 44 | Calderon-Viacava et al. 1971 | |
|
147 | Peruvian Mestizos | Peru | 12 | S | 0.8 | adults | 50 | Figueroa et al. 1971 | |
|
148 | Polish–mixed | Poland | 52.1 | N | 0.36 | 16–59 | 85 | Socha and Ksiazyk 1984 | |
|
149 | Polish–east | Poland | 51.3 | N | 0.37 | 16–59 | 35 | Socha and Ksiazyk 1984 | |
|
150 | Polish–central | Poland | 51.5 | N | 0.37 | 16–59 | 92 | Socha and Ksiazyk 1984 | |
|
151 | Polish–south | Poland | 50 | N | 0.38 | 16–59 | 29 | Socha and Ksiazyk 1984 | |
|
152 | Polish–northeast | Poland | 53 | N | 0.41 | 16–59 | 34 | Socha and Ksiazyk 1984 | |
|
153 | Tussi in Rwanda | Rwanda | 2 | S | 0.08 | adults | 27 | Cox and Elliot 1974 | |
|
154 | Hutu/Tussi mixed | Rwanda | 2 | S | 0.55 | adults | 11 | Cox and Elliot 1974 | |
|
155 | Hutu | Rwanda | 2 | S | 0.58 | adults | 36 | Cox and Elliot 1974 | |
|
156 | Twa | Rwanda | 2 | S | 0.77 | adults | 22 | Cox and Elliot 1974 | |
|
157 | Shi, Bantu of Lake Kivu area | Rwanda | 2 | S | 0.96 | adults | 28 | Cox and Elliot 1974 | |
|
158 | Arabs(Saudi), urban | Saudi Arabia | 24.41 | N | 0.13 | adults | 8 | Cook and Al-Torki 1975 | |
|
159 | Arabs, bedouins | Saudi Arabia | 0.14 | adults | 14 | Cook and Al-Torki 1975 | | ||
|
160 | Yemen Arabs | Saudi Arabia | 24.4 | N | 0.25 | 16–40 | 8 | Cook and Al-Torki 1975 | |
|
161 | “Khadiry” (mixed african/arab) | Saudi Arabia | 24.41 | N | 0.78 | 14–60 | 9 | Cook and Al-Torki 1975 | |
|
162 | Khants(west. Siberia) | Siberia | 61.1 | N | 0.869 | 14–57 | 61 | Lember et al. 1995 | |
|
163 | Indians in Singapore | Singapore | 1.17 | N | 0.8 | 3–10 | 5 | Chua and Seah 1973 | |
|
164 | Chinese | Singapore | 1.17 | N | 0.9 | 3–16 | 10 | Chua and Seah 1973 | |
|
165 | Chinese, Malays,Indians | Singapore | 1.17 | N | 1 | adults | 22 | Bolin et al. 1970a | |
|
166 | Sotho | South | Africa | 26.2 | S | 0.65 | 23 | Segal et al. 1983 | |
|
167 | Zulu | South Africa | 26.2 | S | 0.81 | 32 | Segal et al. 1983 | | |
|
168 | Xhosa | South Africa | 26.2 | S | 0.82 | 17 | Segal et al. 1983 | | |
|
169 | Tswana | South Africa | 26.2 | S | 0.83 | 24 | Segal et al. 1983 | | |
|
170 | Swazi | South Africa | 26.2 | S | 0.86 | 12 | Segal et al. 1983 | | |
|
171 | Bantu of South Africa | South Africa | 26.1 | S | 0.9 | N/A | 31 | Jersky and Kinsley 1967 | |
|
172 | Spaniards | Spain | 40.2 | N | 0.15 | adults | 265 | Pena-Yanez et al. 1971 | |
|
173 | Ceylonese | Sri Lanka | 7 | N | 0.73 | adults | 200 | Senewiratne et al. 1977 | |
|
174 | Bedja | Sudan | 15 | N | 0.111 | adults | 9 | Bayoumi et al. 1981 | |
|
175 | Beja | Sudan | 19 | N | 0.17 | 303 | Bayoumi et al. 1982 | | |
|
176 | Gomocia | Sudan | 15 | N | 0.323 | adults | 31 | Bayoumi et al. 1981 | |
|
177 | Kahli | Sudan | 15 | N | 0.381 | adults | 21 | Bayoumi et al. 1981 | |
|
178 | Jaali | Sudan | 15 | N | 0.469 | adults | 113 | Bayoumi et al. 1981 | |
|
179 | Baggara (sudan) | Sudan | 11 | N | 0.49 | adults | 53 | Bayoumi et al. 1981 | |
|
180 | Sudanese, central | Sudan | 15 | N | 0.5 | adults | 56 | Bayoumi et al. 1981 | |
|
181 | Habbani | Sudan | 14.5 | N | 0.526 | adults | 19 | Bayoumi et al. 1981 | |
|
182 | Sudanese, North Nile | Sudan | 16.5 | N | 0.542 | adults | 24 | Bayoumi et al. 1981 | |
|
183 | Sudanese, northern nomadic | Sudan | 15 | N | 0.565 | adults | 23 | Bayoumi et al. 1981 | |
|
184 | Sydanese, southern | Sudan | 5.5 | N | 0.857 | adults | 13 | Bayoumi et al. 1981 | |
|
185 | Misseri | Sudan | 14.5 | N | 0.6 | adults | 20 | Bayoumi et al. 1981 | |
|
186 | Shaygi | Sudan | 16.5 | N | 0.619 | adults | 42 | Bayoumi et al. 1981 | |
|
187 | Nilotic (Sudan) | Sudan | 5.5 | N | 0.667 | adults | 18 | Bayoumi et al. 1981 | |
|
188 | Nubians | Sudan | 16.5 | N | 0.667 | adults | 21 | Bayoumi et al. 1981 | |
|
189 | sudanese(central), aboriginal negroid | Sudan | 14 | N | 0.692 | adults | 26 | Bayoumi et al. 1981 | |
|
190 | Nilotes (Dinka) | Sudan | 6.5 | N | 0.745 | 282 | Bayoumi et al. 1982 | | |
|
191 | Nuba | Sudan | 14 | N | 0.793 | adults | 20 | Bayoumi et al. 1981 | |
|
192 | Dongolawi | Sudan | 16.5 | N | 0.813 | adults | 16 | Bayoumi et al. 1981 | |
|
193 | Bush negroes | Surinam | 4 | N | 1 | adults | 29 | Luyken et al. 1970 | |
|
194 | Indians in Surinam | Surinam | 0.66 | 27 | Luyken et al. 1970 | | |||
|
195 | Surinam creole adults | Surinam | 0.71 | 31 | Luyken et al. 1970 | | |||
|
196 | Swedes in Sweden | Sweden | 59.2 | N | 0.01 | adults | 400 | Dahlqvist and Linquist 1971 | |
|
197 | Chinese in Taiwan | Taiwan | ? | 1 | adults | 71 | Sung et al. 1972 | | |
|
198 | Thai/nw europ. | Thailand | 18.47 | N | 0.5 | adults | 6 | Flatz and Rotthauwe 1971 | |
|
199 | Thai | Thailand | 13.45 | N | 0.96 | adults | 100 | Flatz and Saengudom 1969 | |
|
200 | Thai | Thailand | 13.45 | N | 0.97 | adults | 140 | Keusch et al. 1969 | |
|
201 | Thai | Thailand | 18.47 | N | 0.99 | adults | 75 | Flatz et al. 1969 | |
|
202 | Thai | Thailand | 18.47 | N | 1 | ad + child | 9 | Flatz and Rotthauwe 1971 | |
|
203 | Thai | Thailand | 18.47 | N | 1 | 4–12 | 24 | Flatz et al. 1969 | |
|
204 | Thai | Thailand | 13.45 | N | 1 | adults | 39 | Troncale et al. 1967 | |
|
205 | Thai | Thailand | 13.45 | N | 1 | 2–4 | 16 | Varavithya et al. 1976 | |
|
206 | Indians from trinidad | Trinidad | ? | 0.6 | adults | 25 | Bartholomew and Young Pong 1976 | | |
|
207 | Tunisians | Tunisia | 34 | N | 0.83 | adults | 43 | Filali et al. 1987 | |
|
208 | Turkish | Turkey | 39.5 | N | 0.71 | 18–28 | 480 | Flatz et al.1986a | |
|
209 | American whites | U.S. | 44.6 | N | 0.06 | adults | 100 | Newcomer et al. 1967 | |
|
210 | American whites | U.S. | 37.5 | N | 0.08 | adults | 12 | Calloway et al. 1969 | |
|
211 | American whites | U.S. | 29.2 | N | 0.09 | 2–14 | 17 | Woteki et al. 1976 | |
|
212 | American whites | U.S. | 39.2 | N | 0.1 | 18–54 | 20 | Bayless and Rosensweig 1966 | |
|
213 | Non–Jewish American whites | U.S. | 41.4 | N | 0.11 | adults | 53 | Tandon et al. 1971 | |
|
214 | American whites | U.S. | 29.2 | N | 0.13 | 19–57 | 8 | Dill et al. 1972 | |
|
215 | Anglo–american whites | U.S. | 0.15 | 18–82 | 142 | Woteki et al. 1977 | | ||
|
216 | American whites | U.S. | 39.2 | N | 0.16 | adults | 19 | Cuatrecasas et al. 1965 | |
|
217 | American whites | U.S. | 31 | N | 0.17 | adults | 18 | Knudsen et al. 1968 | |
|
218 | American whites | U.S. | 0.19 | 93 | Marenco et al. 1970 | | |||
|
219 | American whites | U.S. | 35.3 | N | 0.19 | 3–80 | 145 | Welsh and Rohrer 1967 | |
|
220 | American blacks | U.S. | 39.2 | N | 0.24 | children | 25 | Paige et al. 1977 | |
|
221 | American whites | U.S. | 61.1 | N | 0.25 | adults | 16 | Duncan and Scott 1972 | |
|
222 | American whites | U.S. | 42 | N | 0.25 | >5 yrs | 65 | Lebenthal et al. 1975 | |
|
223 | Chippewa/nw europ.mixed | U.S. | 47.5 | N | 0.36 | 5–73 | 39 | Newcomer et al. 1977b | |
|
224 | American blacks | U.S. | 39.2 | N | 0.45 | children | 31 | Paige et al. 1971 | |
|
225 | Mexican Americans | U.S. | 35.3 | N | 0.47 | adults | 17 | Sowers and Winterfeldt 1975 | |
|
226 | Mexican Americans | U.S. | 29.2 | N | 0.53 | 18–94 | 277 | Woteki et al. 1977 | |
|
227 | American blacks | U.S. | 39.2 | N | 0.54 | 6–13 | 89 | Paige et al. 1975 | |
|
228 | Mexican Americans | U.S. | 29.2 | N | 0.55 | 18–57 | 11 | Dill et al. 1972 | |
|
229 | Mexican Americans | U.S. | 29.2 | N | 0.56 | 2–14 | 75 | Woteki et al. 1976 | |
|
230 | American blacks | U.S. | 39.2 | N | 0.59 | adoles. | 32 | Mitchell et al. 1975 | |
|
231 | American Indian/anglo mixed | U.S. | 33.3 | N | 0.61 | ≥4 | 41 | Johnson et al. 1977 | |
|
232 | American indians mixed | U.S | 35.3 | N | 0.63 | 3mo–57 | 16 | Bose and Welsh 1973 | |
|
233 | Orientals in US (chinese and filipino) | U.S. | 0.65 | adults | 20 | Hua and Bayless 1968 | | ||
|
234 | American Jews | U.S. | 41.4 | N | 0.71 | adults | 41 | Tandon et al. 1971 | |
|
235 | American blacks | U.S. | 39.2 | N | 0.73 | adults | 41 | Cuatrecasas et al. 1965 | |
|
236 | American blacks | U.S. | 39.2 | N | 0.75 | 18–52 | 20 | Bayless and Rosensweig 1966 | |
|
237 | American blacks | U.S. | 41.5 | N | 0.75 | adults | 24 | Littman et al. 1968 | |
|
238 | American blacks | U.S. | 44.6 | N | 0.75 | adults | 8 | Knudsen et al. 1968 | |
|
239 | American blacks | U.S. | 35.3 | N | 0.77 | 3–82 | 22 | Welsh and Rohrer 1967 | |
|
240 | Indians in US | U.S. | 0.83 | adults | 18 | Mehta and Latham 1977 | | ||
|
241 | Chippewa, Minnesota | U.S. | 47.5 | N | 0.93 | 5–73 | 15 | Newcomer et al. 1977b | |
|
242 | Popago | U.S. | 33.27 | N | 0.93 | adults | 14 | Johnson et al. 1978 | |
|
243 | Pima | U.S. | 33.27 | N | 0.95 | ≥4 | 62 | Johnson et al. 1977 | |
|
244 | Indians, Oklahoma | U.S. | 35.3 | N | 0.95 | adults | 20 | Bose and Welsh 1973 | |
|
245 | Orientals in US | U.S. | 1 | adults | 11 | Chung and McGill 1968 | | ||
|
246 | Pima | U.S. | 33.27 | N | 1 | adults | 4 | Johnson et al. 1978 | |
|
247 | Hopi | U.S. | 33.27 | N | 1 | adults | 21 | Johnson et al. 1978 | |
|
248 | Apache | U.S. | 33.27 | N | 1 | adults | 22 | Johnson et al. 1978 | |
|
249 | Chinese in US | U.S. | 39.5 | N | 1 | adults | 6 | Calloway et al. 1969 | |
|
250 | Vietnamese in US | U.S. | 16 | N | 1 | adults | 31 | Anh et al. 1977 | |
|
251 | Tussi | Uganda | 0.2 | N | 0 | adults | 5 | Cook and Dahlqvist 1968 | |
|
252 | Hima Pastoralists | Uganda | 0.4 | S | 0.09 | adults | 11 | Cook and Kajubi 1966 | |
|
253 | Tussi Pastoralists in Uganda | Uganda | 0.2 | N | 0.17 | adults | 12 | Cook and Kajubi 1966 | |
|
254 | Iru (mixed Bantu/Hamitic) | Uganda | 0.2 | N | 0.39 | 13 | Cook and Dahlqvist 1968 | | |
|
255 | Nilotes, nilo–hamites in Uganda | Uganda | 0.2 | N | 0.44 | adults | 9 | Cook and Kajubi 1966 | |
|
256 | Ganda | Uganda | 0.2 | N | 0.67 | 5–12 | 6 | Cook et al. 1967 | |
|
257 | Uganda(agricultural Bantu, I.e.Ganda) | Uganda | 0.2 | N | 0.96 | adults | 52 | Cook and Kajubi 1966 | |
|
258 | Ganda and others in Uganda | Uganda | 0.2 | N | 1 | adults | 12 | Cook and Dahlqvist 1968 | |
|
259 | Yemen Jews | Yemen | 32.1 | N | 0.44 | 20–70 | 36 | Gilat et al. 1970 | |
|
260 | Ibo | Zaire | 4.2 | S | 0.75 | adults | 4 | Elliott et al. 1973 | |
|
261 | Bantu of various types | Zaire | 4.2 | S | 0.95 | adults | 52 | Elliott et al. 1973 | |
|
262 | Bantu of Zambia | Zambia | 15.28 | S | 1 | adults | 26 | Cook et al. 1973 | |
|
263 | canadian whites | ? | 0.06 | 16 | Cox and Elliot 1974 | | |||
|
264 | Indians from trinidad | ? | 0.2 | 5 | Murthy and Haworth 1970 | | |||
|
265 | Fulani, nomadic | ? | 10.5 | N | 0.22 | ≥4 | 9 | Kretchmer et al. 1971 | |
|
266 | Hutu (mixed Bantu/Hamitic) | ? | 0 | N/S | 0.33 | 15 | Cook and Dahlqvist 1968 | | |
|
267 | Greenland Eskimos/nw eur.mixed | ? | 69.2 | N | 0.39 | 1–30 | 97 | Gouin et al. 1972 | |
|
268 | American Indian/anglo mixed | ? | 0.5 | 6 | Gilat et al. 1974 | | |||
|
269 | Africans–north (Maghreb) | ? | 0.78 | 20–70 | 55 | O'Morain et al. 1978 | | ||
|
270 | Dinka | ? | 1 | 5 | Elliott et al. 1973 | | |||
|
Complete data of all populations for which LM frequencies are available. ?=information could not be determined. |
Appendix A. References
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Department of Neurobiology and Behavior, Cornell University, Ithaca, NY 14853, USA
Corresponding author. Tel.: +1 607 254 4333; fax: +1 607 254 4308.
PII: S1090-5138(04)00100-X
doi:10.1016/j.evolhumbehav.2004.10.002
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