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AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY 56:483-489(19811

The Development of Genetics and Population StudiesWILLIAM S. POLLITZERDepartment of Anatomy, University of North Carolina. Chapel Hill, NorthCarolina 27514

KEY WORDS Genetics, Selection, Population structure, Primates,Polymorphisms

ABSTRACT The contribution of genetics and population studies to physicalanthropology as reflected in the pages of our Journal is traced since it s establishment in 1918. Major trends include the use of more genetic polymorphisms, th esearch for natural selection and genetic drift, the unraveling of populationstructure in a wide variety of ecological niches, and the recognition of the role ofculture in human biology. Nonhuman primates have also been explored from theviewpoint of population genetics. Emphasis has been increasingly on processrather than classification.

It is appropriate that the American Association of Physical Anthropologists was foundedat the University of Virginia, that creation ofthe fertile mind of Thomas Jefferson, statesman and scientist. His wide range of interestsencompassed the peoples of the world and thediversity of their biology and culture. It isfitting that today we who are gathered herelook back over the years to the achievements ingenetics and population studies and the waysthey have enriched our discipline of physicalanthropology and contributed to an understanding of our variability.Modern genetics dates from the rediscovery \of Mendel's laws in 1900, and population gene- \tics from the principle of equilibrium by Hardy !and Weinberg in 1908. Blood groups, those!widely studied and useful monogenic traits,!were also discovered by Landsteiner in 1900.Our century has witnessed a fluorescence oftechniques from immunology to electrophoresis that have introduced a broad battery oftraits to the anthropologist for his investigation of human diversity. Unraveling the genetic component in continuous variables such asskit! and eye color has also challenged physicalanthropologists. Dermatoglyphics represent acomplex inherited trait that came into it s ownin this century as a handmaiden to medicineand anthropology. While chromosomes wereidentified 100 years ago, those of man wereaccurately counted in the 1950s when techniques of cell culture permitted that explosionof knowledge of karyotypes and their minutedeviations from the normal pattern.

00029483/81/5604-0483$02.50 1981 ALAN R. LISS, INC.

While natural selection was the keystone ofDarwin's great theory, and DeVries introducedmutation before the turn of the century, it wasHaldane and Fisher in England and Wright in \America who, in the years between the world .,,\)wars, blended gene frequencies and mating Ipatterns to provide a mathematical basis forthe factors of evolutionary change: mutation,selection, gene flow, genetic drift, and inbreed-ving.While the genetic basis of metabolic diseasewas first recognized in the early 1900s byGarrod, biochemical genetics has floweredonly in recent years with the discovery of newenzymes, their control by genes, and their rolein health and illness. The discovery of diseaseassociations with various normal or markergenes gave new impetus to the search fornatural selection. Cell culture techniques andhigh-speed computers facilitated linkagestudies and the assignment of genes to theirprecise location on chromosomes. Adaptability studies, from the heights of the Andes tothe heat of the Sahara, reminded us of the roleof the environment in shaping our biologicaldestiny.Social scientists no less than biologists havecontributed to the population genetics of mankind by recognizing the importance of culturein the flow of genes, whether by mating preferences and class structure or by economics andhostile tribes that governed migration routesDelivered at the Univer1iity of Virginia, Charlottesville, December11-12,1980.

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484 W.S. POLLITZERover time and space. Modern demography atlong last joined human genetics to permit moreprecise estimates of change and the opportunities of natural selection by utilizing birth anddeath records.Archaeologists and physical anthropologists share a common interest in the life andtimes of fossil man. Techniques for determining blood types and other biochemical traitsfrom his bones add a fourth dimension of timeto th e potential territory of populationgenetics and permit inferences on mortalityand family structure of our distant ancestors.Man has long been fascinated by other primates and their behavior. While observationsgo back over the centuries the systematicstudy of their biochemistry and behavior andit s relevance to our own species is a modern development. Quite recently enough data andtheory have accumulated to permit populationgenetics of nonhuman primates.Our Journal, which actually antedates ourassociation by a dozen years, reflects thegrowth of genetics and population studies andtheir influence upon physical anthropology. Ihave enjoyed following that development, for

contains an appeal by the editor Ales Hrdlicka,for "intensive studies of human heredity." TheReports of Current Literature include studieson heredity and eugenics, such as Davenport'swork on stature.An article on inher itance of eye color in manappears in volume 2 (Boas, 1919). In 1922 ourstill-active and much-honored first lady,Mildred Trotter, published, with Danforth, acareful study of the incidence and heredity offacial hypertrichosis in 1,696 white women(Trotter and Danforth, 1922). Two years laterwe read of racial differences in papillary linesof the palm (Keith, 1924), an early venture intodermatoglyphics soon to be followed by themany contributions of Cummins and Midlo.Of far-reaching importance is an article byCastle (1926) in which he questions any biological harm from race crossing, and considerssocial factors of surpassing importance. In thesame year Snyder (1926) published on thei human blood groups, their inheritance, and; racial significance, including techniques, dis\ tribution, and his own survey of 250 Cherokee\ Indians in North Carolina. Subsequent issueswere filled increasingly with such data on thefrequency of blood factors in various peoples ofthe earth. Some 118 articles on serologyappeared in our first 21 volumes. Kroeber

(1934), whose name is more often linked with iJ1cultural anthropology, suggested a scheme of Aclassifying these distributions by plotting the saratio of 0 to A plus B. 1lUVolume 14 stands ou t in our heritage, forit nerecords the founding of our association here atCharlottesville, that first meeting from April wi17 through 19, 1930, the adoption ofAJPA 89 qUour official journal, and a list of founding SOmembers. faLThe earliest article potentially related to denatural selection and demography in mankind Alwas one on the fecundity of Sioux women by gI QHrdlicka (1932). filCandela (1936) wrote on blood group reae- sigtions of 11 ancient human skeletons of Egyp- biotians of 1500 B.C., generously crediting Boyd onwith similar and earlier investigations. bleOur Book Notes often present as much his dYtory of our discipline as our articles. One in foU1939 said that Dobzhansky's book, Genetic. deand the Origin of Species (1937), was "re- hefreshing and well worth a perusal by the 8II" E;hropologist." Not long after Dobzhansky as~ 1 9 4 4 ) himself wrote on species and races of ch s!living and fossil man. Grounding his GefiDi:. fo rit s lineage can be traced by turning t h r o u g h ~ I tions in genetics, he saw living mankind 89 . drUthese pages. Even the firs t volume in 1918l('. single polytyp ic species and found "no ason giQJ: to suppose that more than a single ho . . V. species has existed on any time level ,. an. Pleistocene." Whether one agrees or not, his traviewpoint united genetics with paleoant sigJ. pology. - f lOIn that same volume Wiener with his ourcolleagues (1944) contributed the first of many Tarticles on the frequency of blood factors, in pricluding th e recently discovered Rh. cell." In 1950 geneticists and anthropologists met ha pat Cold Spring Harbor for a watershed sym G6Jsium on the origin and evolution of man. Sub- of ssequent issues of our Journal echo and refine thathe important ideas generated at that confer- hisence. Boyd's seminal book, Genetics and tIN theRaces ofMan, was favorably reviewed by Bini- gensell (1951). The following year Birdsell (19521 (l9Eexplained in a brief communication why selefrequencies need not coincide with raCli M8Iphenotypes. Sanghvi (1953) demonstra impthat blood types indicate a degree of divi- isollgence of castes of Bombay, India, sirnilllrto hthat shown by their morphology. tionThe 1950s saw an increasing shift from 8B pop1interest in racia l classification for its own sake sucJto a focus on the process of change; races WII8 Yanseen as fleeting stages in the broad sweep rl so nevolution. Coon et al. (1950) published their ch81slender bu t provocative book on races, review- and

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- -------- - - - - - - - -

485EVELOPMENT OF GENETICS AND POPULATION STUDIESwith

scheme ofby plotting theheritage, for itat

of AJPA asfounding

related toin mankindwomen by

group reacEgyp-

crediting Boydpresent as much his

One inGenetics(1937), was "reby th e anafter Dobzhanskyspecies and races ofhis definimankind as areasonthan a single hominid

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in the or not, hispaleoanthnF.

with histhe first of manyfactors, inanthropologists metwatershed sympoman. SubJournal echo and refineat that conferGenetics and thereview ed by Bird

why genewith racial ...demonstrateddegree of diverIndia, similar toshift from anit s own sakeess of change ; races werebroad sweep ofpublished theiron races, review

ing the nineteenth century zoological rules ofAllen, Bergmann, and Gloger to apply theseselective pressures to th e formation ofmankind. Did no t hemolytic disease of thenewborn, with the loss of one rarer Rh negativeallele and one more common Rh posit ive allelewith each infant death, mean that allele frequencies must change over time? Did no t Allison demonstrate the profound influence offalciparum malaria on the fluctuating incidence of sickle cell hemoglobin? Studies onABO disease associations, briefly explored ageneration earlier and then shelved, began tofill the pages of many journals. In her farsighted article in AJPA on selection and ABOblood groups, Brues (1954) showed that onlyone-fifth of the phenotype combinations possible actually occurred, an argument fordynamically balanced selection factors. In thefollowing year Boyd (1955) wrestled with thedetection of selective advantages of theheterozygotes in human genetic traits.But Sewall Wright had stressed genetic driftas well as selection as a factor of evolutionarychange. Glass (1956) presented in our Journalfor the first time evidence of random geneticdrift in a human population, the Dunker religious isolate in Pennsylvania.We have noted the recognition amonganthropologists of the influence of culturaltraits in gene flow. Hulse (1957) published asignificant paper. linguistic barriers to geneflow, a theme that has several later echoets inour pages.

Through the sixties our Journal continued toprint population surveys using no t only redcell antigens, bu t such serological traits ashaptoglobins, group-specific component, andG6PD deficiency, as well as art icles on aspectsof selection and drift. Thus, in the last year ofthat decade Livingstone (1969l, well known forhis studies of hemoglobin variants, wrote onthe founder effect- that twin brother ofgenetic drift - and deleterious genes, and Sever(19691 discussed ABO hemolytic disease as aselective mechanism. In the following yearMartin (19701 considered the evolutionaryimplications of the founder effect in a humanisolate.Increasingly in the seventies we find evolutionary mechan isms related to demography inpopulation studies. Chagnon et al. (1970) foundsuch influences on gene flow into theYanomama Indians of South America. Erickson and colleagues (19701 related geneticchange to demography in Chiapas, Mexico,and the studies of Halberstein and Crawford

(1972) in Tlaxcala, Mexico, approached naturalselection through demographic data.One recurrent theme in population studieshas been a measure of genetic distance, ordegree of dissimilarity among people. II haveeven been guilty of it myself.) A more recentand more fruitful inquiry has been understand-ing the causes for the diversity observed: Howmuch is due to drift because of isolation, howmuch reflects admixture, and what is the roleof mating patterns? Leaders in these investigations have been primarily geneticists whosework overlapped with and contributed tophysical anthropology. Our discipline has beenenriched because Neel pioneered in populationstudies of such primitive people as theXavante and Yanomama Indians. He contributed the idea of a lineal effect - a fission of avillage along certain kinship linets. When twovillages combine their inhabitants we mayspeak of fusion. Such a fission-fusion model isconsidered potential ly important in its geneticeffects no t only among uncivilized tribes oftoday but over the millenia of prehistory aswell. Moreover, one head-man, endowed withattributes of leadership, may through polygamy contribute a disproportionate share ofgenes to succeeding generations - a specialkind of founder effect. Concepts and calculations generated by these ideas often foundtheir way into our Journal. Thus, Spielman(1973) published on th e differences inYanomama Indian villages, and Fix (1975)cited a similar model of fission-fusion andlineal effect in it s investigation of the SemaiSenoi of Malaysia.One topic increasingly explored by physicalanthropologists in the past decade is the genetic structure of human populations. Even halfa century ago Sewall Wright investigated systems of mating and their effects on gene frequencies. Recognizing that preferential matings, along with immigration, produce deviations from Hardy-Weinberg equilibrium, hecontributed two models of the inbreeding co-efficient, the genealogical and the hierarchical.The latter especially was designed to explorethe opportunity for drift in a branchingprocess of semi-isolated populations. Hedemonstrated that the genetic variance between such islands depends on the effectivepopulation size and migration from outside. Alater spatial model devised by Malecot utilizedthe covariance of gene frequencies to reflectthe probability of identity by descent. Such acoefficient of kinship tends to decline with increasing distance between the birthplace of

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486 W.S. POLLITZERparents. Morton especially applied Malecot'sideas to human populations, from Switzerlandand Brazil to Pacific Islands, in a process hecalls bioassay. I t can be shown that inbreedingdeclines rapidly only at small distances, anindication that consanguineous marriagesoccur preferentially at large distances.These approaches to population analysis areexpressed in the pages of our Journal. Friedlaender (1971) discovered barr iers to matingsboth from distance and from kinship as indicated in gene frequencies in BougainvilleIsland, and, equally important, he found differences in the population structure of horticulturalists and hunter-gatherers. Workman alsovigorously applied such techniques to severalpopulations. He and his colleagues (1974,1975)found isolation by distance a most importantfactor in the pattern of regional differentiationamong Zuni Indian tribes and also among 14villages of Sardinia. More recently he exploredthe genetic structure of Finland (1976), notinggene flow from Sweden, the role of settlementpatterns, and the gradient in certain allelefrequencies.In other areas and differen t ecological nichesinvestigators reported varying results of population structure analysis. For example, Kirk etal. (1977) found no systematic relationship between genetic distance and geographic location in the Caspian littoral of Northern Iran.Crawford and his associates contributedgreatly to an understanding of populationstructure in several lands. In Mexico, they(1974) found geographical distance and hybridization to be the major determinants ofgene frequency distribution.Since surnames are culturally inherited fromfather to son, just like the biological transmission of the Y chromosome, the frequency ofthe same last name has been useful in population analysis. Lasker was particularly responsible for developing this method of isonymy,applying it extensively to the people of GreatBritain. Morton and Lalouel (1973), in theirbioassay in Micronesia, find some agreement.lIl1ong estimations of kinship from phenotypes, anthropometrics, and isonymy.Azevedo and colleagues (Tavares-Neto andAzevedo, 1978; Oliveira and Azevedo, 1977) inBrazil demonstrated the correlation of thedegree of black, white, and Indian admixturewith last names, physical measurements, andgene frequencies.The quest for detecting natural selection viapopulation studies from several sources is mirrored in our Journal in the seventies. Thedemographic approach is expanded, direct

measures of fertility are attempted, anddisease associations are further explored.Palmarino and associates (1975) reported thatred cell acid phosphatase may be another poly-morphism correlated with malaria. Of enor-mous promise are investigations that combineadaptation with specific genetic traits. In thiscurrent year, Constans and colleagues \1980),in a study of Gc (Vitamin D binding protein) inthe Sahara, find the lowest Gc ' frequencieswhere sunlight is strongest and a gradient ofGC1F that parallels a skin color cline. Similarly,Moore and Brewer (1980) explore the biochemi-cal mechanisms that make RBC 2,3 Diphos-phoglycerate increased at high altitudes.Variations in the number and structure ofchromosomes are increasingly important tomedicine and anthropology. Literally hun-dreds of deletions, duplications, inversions,translocations. and other alterations can nowbe identified precisely thanks to banding tech-niques. While many of them are distinctly rareand abnormal, others are common and harm'less polymorphisms useful for the growing sci-ence of human population cytogenetics.Heltne and Singer (1971) reported on themorphology and distribution of the chromo-somes of the Hottentot population. Curiously,very little additional human cytogenetics hasappeared in our Journal since.The application of serological techniques toancient tissues expanded the time depth ofgenetics and physical anthropology. We havenoted the initial work of Candela and Boyd onblood groups of bones. In the intervening yearsgreat improvements were made in their tech-niques. Allison and colleagues (1976) appliedsuch improved procedures to the typing ofPeruvian mummies. Micle and colleagues(1977) at Tel Aviv University reported resultsof typing 55 ancient skeletons of Israel. In thesame year Facchini and Pettener (1977) pre-sented new techniques for dating skeletalremains: benzidine reaction, ultraviolet fluor-escence, specific gravity, and especially super-sonic conductivity. These and related ad-vances hold great promise for extendingpopu-lation genetics of our species far into the past.The idea dawned slowly that as man is apri-mate, the serology and genetics, like the mor-phology, of other primates should be of con-cern to the physical anthropologists. Weineretal. (1953) published the first of a series ofarticles on blood factors in apes and monkeys,including Rh, in our Journal and Butts (19531described the hemagglutinins of chimpanzeesin the same year. Shortly after, Gartler et aL(1956) wrote on inherited biochemical traits in

greatpresevarievariaAt a;(1967ledgethatsomelappeaMOl

J a n u ~andpited tprosir.chromof semonk!I t iscepts

One ,a m o n ~heardternit)Steifrom t4an d suquencidu e toones. Ientiation Caycolleagmigratcan exMorenpointedgeneticdiffereamongTodageneticprovideonly deevolutiprimatThistion st1tendedtions obeen omcover a

devotean d it svelopmtheir wAdvaThe exc

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- - - - - - - - - -

487EVELOPMENT OF GENETICS AND POPULATION STUDIESgreat apes. Succeeding years saw many similar tics have much to contribute to physical an

further explored. #II presentations on monogenic traits in a wide thropology. There is no reason why all of the(1975) reported thatattempted, and

variety of primates: blood types, hemoglobin known biochemical traits should no t be exbe another poly variants, serum proteins, and red cell enzymes. plored in varied populations, with one eye onOf enor At a symposium in Chapel Hill Barnicot et al. social structure and another on adaptation to

that combinetraits. In this(1980),binding protein) in ,.Gc ' frequenciesand a gradient of

Similarly,explore th e biochemiDiphoshigh altitudes.and structure ofimportant toLiterally huninversions,

alterations can nowto banding techare distinctly rarean d harmthe growing sci

cytogenetics.reported on theth e chromoCuriously,hassince. techniques to

the time depth ofWe haveand Boyd onthe intervening yearsmade in their tech(1976) appliedto the typing ofand colleaguesreported results

Israel. In thePettener (1977) predating skeletalultraviolet fluorand especially superand related adextending popuinto the past.that as man is a prith e mor

should be of conWeiner ete first of a series ofand monkeys.

and Butts (1953)of chimpanzeesGartler et al.traits in

(1967) summarized the then-current knowledge of protein variations and primatology. Inthat same year the first article on chromosomes in the lemuridae (Egozcue, 1967)appeared in our Journal.More than any other person, BuettnerJanusch has made us aware, from his researchand publications, of the importance of inherited traits in nonhuman primates, especiallyprosimians. He and his followers published onchromosomes in lemurs (1973) as well as a hostof serological traits in other prosimians,monkeys, baboons, and apes.It is now possible to apply the data and concepts of population genetics to such primates.One can determine paternity exclusionsamong monkeys, although I have not ye theard of any resulting lawsuits from nonpaternity.Steinberg et al. (1977) reviewed Gm samplesfrom ten troops of baboons in Northern Kenyaand suggested that variation in haplotype frequencies is the result of founder effect largelydue to fission of a large troop into two smallerones. Duggleby (1978) reported genetic differentiation of social groups among the macaqueson Cayo Santiago. In the same year Ober andcolleagues (1978) showed that recurrent malemigration between troops of baboons in Kenyacan explain intertroop microdifferentiation.More recently Cheverud and associates (1978)pointed to lineal fusion, male migration, andgenetic drift as explanations for the degree ofdifferences observed at the transferrin locusamong macaques on Cayo Santiago.Today the various techniques of moleculargenetics supplement traditional serology andprovide a valuable tool for constructing no tonly dendrograms of living populations bu tevolutionary trees of the phylogeny of otherprimates as well.This brief overview of genetics and population studies, as voiced in AJPA, is no t intended to be complete. Significant contributions of many physical anthropologists havebeen omitted. We have no t even attempted tocover a dozen textbooks of our science thatdevote up to half of their contents to geneticsand it s impact. Moreover, many important developments in genetics have no t as ye t foundtheir way into our discipline.Advances on all fronts would be welcome.The exciting new discoveries of modern gene-

the natural environment for clues to selection.I would also like to see the gap closed betweenthe gene and the chromosome with further investigation of the ultrastructure of cytogenetic variations.Fa r more information can ye t be wrung fromthe interface of disease and genetics. Diseaseassociations of polymorphisms are one dimension. HLA, the human leucocyte antigens,with their wide range of variations in populations, and the striking association of somehaplotypes with specific diseases, offer th egreatest insights for the future. The more precise measure of predisposition to diseaseagainst a background of varying climates andcultures opens up vistas of knowledge as important for anthropology as for medicine.Presumably many so-called polygenic traitsmay prove to be governed by one or a fewmajor genes, whose presence can be detectedby segregation and linkage analysis.How little we still know about the genetic influence on our utilization of food - no t just ofproteins bu t a multitude of other nutrients including important trace elements, with a resultant effect on the growth and adaptation ofpopulations with different ecologies. In th econtribution of genes to succeeding generations, we still have much to understand aboutdifferential fertili ty and it s causes. What is theinfluence of birth control in different cultureson the actual fecundity of the people?One new discipline, behavior genetics, hasbeen relatively neglected by physical anthropologists-perhaps for good and healthy reasons. We do not wish to return to the eugenicsof half a century ago with it s excessive andmisplaced zeal for human betterment by eliminating the allegedly poor germ plasm, it sracism, and it s trail of human degrad'ltion. Butthere is the proper place for the careful,objective investigation of th e possible geneticand environmental influences on humanbehavior, quite independent of any classes orcategori es of people and totally devoid of valuejudgements.

What are the major trends and lessons seenas we look back across these decades? Not justthe dramatic growth of genetic markersutilized in population surveys. Nor the development of theory of kinship structure alone. I tis also the increasing recognition of culture andattitudes in shaping society and directing the

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488 W.S. POLLITZERflow of genes over the centuries. The expansionof th e genetics of populations to embrace allprimates is another step in the right direction.To see ourselves as part of the great unfoldingof evolution, to understand better thosefactors of change that have produced allspecies of life is a supreme intellectualchallenge. We must continue with courage andhonesty to learn all we can about the naturalcauses of natural phenomena. Debates aboutpunctuated versus Darwinian evolution orselectionists versus neutralists must be seenas a healthy science refining it s concepts in thelight of evidence and no t as any rejection ofevolution.Perhaps the most important lesson fromgenetics. and its greatest influence upon ourdiscipline, has been the shift from a static to adynamic viewpoint. I t sounded the deathknell of typology and rang in the birth of populations. Whether one discards racial groups entirely to delineate only clines, or still findsutility in identifying clusters of mankind, wecan recognize that such groups are relative,labile. and quantitative rather than absolute,stable, and qualitative. Our human perceptions influence the very formation of suchgroups as well as their definitions. Weare moreconcerned today with processes than withclassifications. We recognize that each individual is unique in genetic endowment and environmental heritage. We can appreciate andcheri sh people in all of their splendid diversit y.

And I believe that Thomas Jefferson wouldhave welcomed that democratic viewpoint.LITERATURE CITED

Allison, MJ , Hossaini, AA, Castro, N, Munizaga, J, an dPezzia, A 11976) ABO blood groups in Peruvian mummies.1. An evaluation of teclmiques. Am. J. Phys. Anthropo1.44:55-61.Barnicot, NA, Jolly. CJ, and Wade, PT 119671 Protein variations and primatology. Am. J. Phys. Anthropo1.27:343-355.Birdsell, JB (19511 Genetics and the Races of Man by WCBoyd. Am. J. Phys. Anthropo l. 19:219-233 (book review).Birdsell, JB (19521 On various levels of objectivity in genetical anthropology. Am. J. Phys. Anthropo1. 10:355--362.Boas, HM 119191 Inheritance of eye-color in man. Am. J.Phys. Anthropol. (0.8.)2:15-20.Boyd, WC 119551 Detection of selective advantages of the

heterozygotes in man. Am. J. Phys. Anthropo1. 13:37 -52.Brues, AM 11954) Selection and polymorphism in the A-B-Oblood groups. Am. J. Phys. Anthropo1. 12:559-597.Buettner-Janusch, J, Hamilton, AE, and Bergeron, JA(1973) Chromosomes of lemuriformes. 1. A chromosome

complement of Lepilemur mustelinus 11. Geoffroy 1851)Am. J. Phys. Anthropo1. 39:1-5.Butts, DCA (1953! Hemagglutinogens of the chimpanzee.Am. J. Phys. Anthropol. 11:215-224.Candela, PB (19361 Bloodgroup reactions in ancient humanskeletons. Am. J. Phys. Anthropo1. (0.8.121:429-432.Castle, WE (19261 Biological and social consequences of race

crossing. Am. J. Phys. Anthropol. 10.S.) 9:145-156.Chagnon. NA, Neel, JV, Weitkamp. L, Gershowitz, H, andAyres, M 11970) The influence of cultural factors on thedemography and pattern of gene flow from the Makiritareto the Yanomama Indians. Am. J. Phys. Anthropol.32:339-349.Cheverud, JM, Buettner'Janusch, J, and Sade, D (l97BISocial group fission and the origin of intergroup geneticdifferentiation among the rhesus monkeys of CayoSantiago. Am. J. Phys. Anthropo1. 49:449-456.

Constans, J, L eFevre-Wi tier, Ph, Richard, P, and Jaeger,G(1980) Gc (Vitamin D Binding Proteinl subtype polymorphism and variants distribution among Saharan, MiddleEast, and African populations. Am. J. Phys. Anthropol.52:435-441.Coon, CS, Garn, SM, and Birdsell, JB 119501 Races: AStudyof the Problems of Race Formation in Man. Springfield,Ill.: CC Thomas.Crawford, MH, Leyshon, WC, Brown, K. Lees, F, andTaylor, L (1974) Human biology in Mexico. 11. Acomparison of blood groups, serum and red cell enzymefrequencies, and genetic distances of the Indian populations of Mexico. Am. J. Phys. Anthropo1. 41:251-268.Dobzhansky, T (1937) Genetics and the Origin of S p e c i e ~ New York: Columbia University Press.

Dobzhansky, T 11944) On species an d races of living and f o sil man. Am. J. Phys. Anthropol. 2:251-265.Duggleby, CR 11978) Blood group antigens and the population genetics of Macaca mulatta on Cayo Santiago. I.Genetic differentiation of social groups. Am. J. P h y ~ Anthropol. 48:35-40.Egozcue, J (1967) Chromosome variability in theLemuridae. Am. J. Phys. Ant hropol. 26:341-347.Erickson, RP, Nerlove, S, Creger, WP, and Romney, AK119701 Comparison of genetic and anthropological inter'pretations of population isolates in Aguacatenango,Chiapas, Mexico. Am. J. Phys. Anthropo1. 32:105-120.Facchini, F, and Pettener. D (1977) Chemical and physicalmethods in dating human skeletal remains. Am. J. Phys,Anthropol. 47:65-70.Fix, AG (19751 Fission-fusion and lineal effect: Aspects ofthe population structure of the Semai Senoi of Malaysia.Am. J. Phys. Anthropo1. 43:295-302.Friedlaender, JS 11971) The population structure of

South-Central Bougainville. Am. J. Phys. Anthropol.35:1.3-21i.Gartler. SM, Firschein, L, and Dobzhansky, TI19561 Achromatographic investigation of urinary amino-acids in thegreat apes. Am. J. Phys. Anthropo1. 14:41-57.Glass, HB 119561 On the evidence of random genetic drift in

human populations. Am. J. Phys. Anthropo1.14:541-555.Halberstein, RA, and Crawford, MH 11972) Human biologyin Tlaxcala, Mexico: Demography. Am. J. Phys.Anthropol. 36:199-212.Heltne, PG. and Singer, R 11971) Cytogenet ic studies in theHottentot population: Count distribution, report of a

fragment, and preliminary description of morphology.Am. J. Phys. Anthropol. 34:1-36.Hrdlicka, A (] 932) Frecundity in the Sioux women. Am. J.Phys. Anthropo1. 10.S.) 16:81-90.Hulse, FS 11957) Linguistic barriers to gene flow. The bloodgroups of the Yakima, Okanagan and Swinomish IndillJ1ll.Am. J. Phys. Anthropol. 1 5 : ~ 3 5 - 2 4 6 . Keith, HH 119241 Racial differences in the papillary lines ofthe palm. Am. J. Phys. Anthropol. (0.8.1 7:165-206.Kirk, RL, Keats, B, Blake, NM, McDermid, EM , Ala, F,Karimi, M, Nickbin, B, Shabazi, H, and Kmet, J 119771Genes and people in the Caspian littoral: A population

genetic study in Northern Iran. Am. J. Phys. AnthropoL46:377 -390 .Kroeber, AL 119341 Blood-group classification. Am. J.Phys. Anthropo1. 10.S.) 18:377-393.

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Oliveira,in antBrazil.

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Sever, LEselectiothropol

Snyder. Lan d ra