Genetic Support for Proposed Patterns of Relationship among Lowland South American Languages

Previous articleNext article FreeReportsGenetic Support for Proposed Patterns of Relationship among Lowland South American Languages1FranciscoMauroSalzano, MaraHelenaHutz, SabrinaPintoSalamoni, PaulaRohr, and SidiaMariaCallegariJacquesFranciscoMauroSalzanoDepartamento de Gentica, Instituto de Biocincias, Universidade Federal do Rio Grande do Sul, Caixa Postal 15053, 91501970 Porto Alegre, RS, Brazil (all authors)/Departamento de Estatstica, Instituto de Matemtica, Universidade Federal do Rio Grande do Sul, Campus do Vale, 91540000 Porto Alegre, RS, Brazil ([email protected]) (CallegariJacques). 25 v 05, MaraHelenaHutzDepartamento de Gentica, Instituto de Biocincias, Universidade Federal do Rio Grande do Sul, Caixa Postal 15053, 91501970 Porto Alegre, RS, Brazil (all authors)/Departamento de Estatstica, Instituto de Matemtica, Universidade Federal do Rio Grande do Sul, Campus do Vale, 91540000 Porto Alegre, RS, Brazil ([email protected]) (CallegariJacques). 25 v 05, SabrinaPintoSalamoniDepartamento de Gentica, Instituto de Biocincias, Universidade Federal do Rio Grande do Sul, Caixa Postal 15053, 91501970 Porto Alegre, RS, Brazil (all authors)/Departamento de Estatstica, Instituto de Matemtica, Universidade Federal do Rio Grande do Sul, Campus do Vale, 91540000 Porto Alegre, RS, Brazil ([email protected]) (CallegariJacques). 25 v 05, PaulaRohrDepartamento de Gentica, Instituto de Biocincias, Universidade Federal do Rio Grande do Sul, Caixa Postal 15053, 91501970 Porto Alegre, RS, Brazil (all authors)/Departamento de Estatstica, Instituto de Matemtica, Universidade Federal do Rio Grande do Sul, Campus do Vale, 91540000 Porto Alegre, RS, Brazil ([email protected]) (CallegariJacques). 25 v 05, and SidiaMariaCallegariJacquesDepartamento de Gentica, Instituto de Biocincias, Universidade Federal do Rio Grande do Sul, Caixa Postal 15053, 91501970 Porto Alegre, RS, Brazil (all authors)/Departamento de Estatstica, Instituto de Matemtica, Universidade Federal do Rio Grande do Sul, Campus do Vale, 91540000 Porto Alegre, RS, Brazil ([email protected]) (CallegariJacques). 25 v 05Departamento de Gentica, Instituto de Biocincias, Universidade Federal do Rio Grande do Sul, Caixa Postal 15053, 91501970 Porto Alegre, RS, Brazil (all authors)/Departamento de Estatstica, Instituto de Matemtica, Universidade Federal do Rio Grande do Sul, Campus do Vale, 91540000 Porto Alegre, RS, Brazil ([email protected]) (CallegariJacques). 25 v 05PDFPDF PLUSFull Text Add to favoritesDownload CitationTrack CitationsPermissionsReprints Share onFacebookTwitterLinked InRedditEmailQR Code SectionsMoreComparison of different sets of markers to unravel the past of human populations is an established procedure in both anthropology and genetics. Language characteristics can be easily quantified, and the field of comparative linguistics has a respectable past (Trask 1996; Nichols 1997, 2002; Campbell 1997). Therefore, it is only natural that evolutionary geneticists have turned to linguistics to evaluate the population relationships that they have been obtaining with genetic markers.Quantitative geneticlinguistic population comparisons have been conducted for some three decades. A landmark paper for South American populations was that of Spielman, Migliazza, and Neel (1974), which extensively considered the regional differences among Yanomama from both the linguistic and the genetic point of view. At the worldwide level the studies of CavalliSforza et al. (1988; CavalliSforza, Minch, and Mountain 1992; review in CavalliSforza 1997) and recent reanalysis in Nettle and Harris (2003) are notable. Although these investigations have been severely criticized (Bateman et al. 1990a,b), their validity has been confirmed in some cases (see discussion in Nettle and Harris 2003). All of these studies have involved blood group and protein genetic markers. At the DNA level the patterns of agreement are not so clear, but a strong correspondence has been reported at the world level between Ychromosome markers and linguistics (Poloni et al. 1997, Barbujani 1997).There is no consensus among linguists about the relationships among South American Indian languages. The classification of these languages presents several difficulties. First, this subcontinent exhibits considerably more linguistic diversity than North and Central America combined. Second, significant historical linguistic research has been conducted on only a few of these families and isolates. Third, the dominant scientific tendency has been to present broad, largescale classifications, while historical research on individual languages families has received less attention (Campbell 1997:170).Our group has been interested in aspects of evolutionary genetics in South America for a long time (Salzano et al. 1977, Black et al. 1983, CallegariJacques and Salzano 1989, CallegariJacques et al. 1993, Fagundes et al. 2002), and we have often relied on the linguistic classifications proposed by various scholars, aware that they disagree with regard to the relationships between language families. Because of our efforts and those of others, a large amount of information on protein genetic markers in these populations has been compiled, and we wondered whether these data could help clarify the issue of native language classification in this subcontinent. The objective of this report, is distinct from that of our previous work, is to evaluate the agreement of the available genetic data with the alternatives proposed by three eminent linguists (C. Loukotka, J. H. Greenberg, and A. D. Rodrigues) for the relationships among the four most important lowland South American native language families, namely, Maipure, Carib, Tupi, and Ge.The Maipure or Arawak family is the largest in the New World. It covers a wide geographic region, including languages spoken in Central America and the Gran Chaco and the area from the Andes to the mouth of the Amazon River. Maipure used to be thought of as a major subgroup of Arawak, but Campbell (1997) has suggested that at present the terms are interchangeable. Carib designates a family found on the northern coast of South America and the Lesser Antilles. The majority of the groups that speak languages of the Carib family live in the Guiana region, but there are also speakers of it south of the Amazon (Rodrigues 1994). Tupi languages occur in a vast region across South America, but their dispersion may have begun 3,0005,000 years ago in the region between the Madeira and Xingu Rivers in what is now centralwestern Brazil (Urban 1992). The Ge languages are spoken mainly in the eastern and central areas of the Brazilian Central Plateau. Their point of dispersion may have been somewhere between the headwaters of the San Francisco and the Araguaia River, with a split leading to southern and northern expansions (Urban 1992).Loukotka (1968) proposed that the Maipure family was more similar to the Tupi and that Ge was more remotely related to this pair of families than the Carib family. Rodrigues (1985, 1994, 2000, personal communication) has linked Carib with Tupi and regards Maipure as less closely related to this pair than Ge. Greenberg (1987) groups Maipure with Tupi and Carib with Ge. A schematic representation of these ideas is displayed in figure 1.Fig. 1. Schematic representation of the relationships among the four main lowland South American native linguistic families according to Greenberg, Loukotka, and Rodrigues.View Large ImageDownload PowerPointSubjects, Methods, and AnalysisA databank for the evaluation of South American Native genetic variability was started by one of us (SMCJ) in 1982, using information from previous decades, and has been kept up to date ever since. We began by identifying the populations studied for blood group and protein genetic traits that fell into the four linguistic families of interest. The available data consisted of 37 genetic systems (including blood groups, electrophoretic markers, immunoglobulins, and histocompatibility lociHLA) that we will call classical and 13 autosomal shorttandemrepeat polymorphisms (STRPs), which directly assess the variability at the DNA level. The amount of information related to the latter set of markers was much more limited, but we decided to see whether the STRP results would support the conclusions obtained with the first set. The testing of the hypotheses concerning language relationship patterns of Greenberg, Loukotka, and Rodrigues was performed using genetic data by means of a method developed by CavalliSforza and Piazza (1975) to reconstruct the evolutionary history of populations, as follows: Suppose that a population (e.g., South American Indians) has several levels of nested subdivisions (e.g., language classifications). Individuals are nested within languages, which in turn can be grouped into larger groups, for example, stocks or families. The hierarchy can be represented by a bifurcating tree in which the terminal nodes are populations defined according to the same criterion (e.g., linguistic families) and the internal nodes (bifurcating points) are hypothetical populations from which the terminal nodes could have been derived.Consider now that allele frequencies of several loci have been determined in the populations defined as terminal nodes of the tree. CavalliSforza and Piazza (1975) showed that the expected covariance of gene frequencies between two existing populations equals the expected variance associated with the highest internal node that links them. This theorem allows estimation of the variance associated with any internal node of the hierarchy, whether or not the population at that node has been observed. They also showed that the variance of an extant population is equal to a sum of certain quantities defined as the amounts of evolution that have occurred in the transitions from an ancestor common to all populations through all the intermediate nodes in the line to the population considered. An assumption related to this theory is independence of effects across hierarchical levels, but no assumptions are made about gene frequency transformations. The latter have been used to stabilize the variance (for instance, arcsine transformation [CavalliSforza and Piazza 1975) or to enhance the interpretation of the variancecovariance matrix (Urbanek, Goldman, and Long 1996), rendering its elements equal to coefficients of gene identities (Nei 1972).These are the general bases for establishing tests of different models of bifurcating trees using genetic information. The adjustment of the data to a given tree can be evaluated by a goodnessoffit test using a likelihood ratio statistic (lambda), which involves the observed variancecovariance matrix and the corresponding theoretical matrix estimated by maximum likelihood (CavalliSforza and Piazza 1975). With a large number of traits, lambda follows a chisquare distribution with degrees of freedom equal to s(s + 1)/2 minus the number of fitted parameters, where s is the number of external nodes (populations for which there is actual genetic information). A specific tree (e.g., a particular pattern proposed for the linguistic relationships among languages) constitutes a null hypothesis that will be rejected if the tests P value is equal to or less than an arbitrary figure (in this study 0.05).In order to have a better general picture of the fit of other models to our data, we tested all possible bifurcating trees for four sets of populations (language families). This number happens to be 15 (CavalliSforza and Piazza 1975), 3 of them being the models suggested by Greenberg, Loukotka, and Rodrigues. Two of the remaining 12 are the trees inferred using Neis (1972) standard and Nei et al.s (1983) DA genetic distances and the neighborjoining algorithm (Saitou and Nei 1987) for our data. These two topologies yield elements for a test of the treeness of the genetic data, which is an assumption for the whole analysis. In other words, if the treeness quality is rejected for the genetic markers, no tree will fit the genetic data well (e.g., Hunley and Long 2005).Details about the classical genetic systems employed are provided in Table 1, the populations considered being those that speak languages classified as Maipure, Carib, Ge, or Tupi. Gene frequencies were obtained by direct counting or using maximum likelihood procedures in the case of dominance. For the HLA data the squareroot method was used, followed by appropriate adjustments. The average gene frequency obtained from a variable number of tribes, according to the information available for each system, was used to represent each language family. The use of an average allows a better genetic characterization of the language family, since it reduces the drift effect that would occur if individual tribes were considered.Table 1 Number of Samples and Individuals Studied According to Four Linguistic Families and 37 Blood Group and Protein SystemsGenetic SystemaMaipureCaribGeTupiNo. of IndividualsNo. of SamplesNo. of IndividualsNo. of SamplesNo. of IndividualsNo. of SamplesNo. of IndividualsNo. of SamplesABO3,622287,675432,543143,24333ACP1,23262,74291,08382,38621AK1,25671,86281,06492,39721ALB1,32062,814112,181102,41220CA21,09941,49061,35181,52213CHE1118194551,06592,03619CP1,14151,69871,646102,09918DIEGO2,261174,083352,091101,40422DUFFY2,272175,172372,127122,08729ESA1,09241,54661,21481,45214ESD1,18351,79881,466101,83417G6PD23457851040581,01121GC1,15972,67192,3501097310GLO21131,094736151,28313GM2,291112,805111,15231,70816GPI1,03641,093371622682HBA2,468193,981322,206122,68731HBA276511,01471,16801,75616HBFET751320249411502HLAA401439549851,14412HLAB401439549851,14412HLAC401302349851,07811HP2,297144,396242,392132,56327KELL2,211164,687272,242122,18028KIDD1,772123,125191,423688613KM2,024102,961141,64761,62818LEWIS1,30682,17381,19332776LUTHERAN41841,8101017411522MDH1,05051,173465922682MNSs2,192164,508241,95391,81823P2,141164,985302,044102,20427PEPA1,18751,496678831,46215PGD1,46272,91411238102,29321PGM11,24462,50991,452102,31420PGM21,19051,84071,03441,90017RH2,361185,283342,374132,31828TF2,446164,085232,354132,86327 Mean1,35882,560141,35581,65417Median1,23262,17391,21491,75618Modal class 15 610 610 1620Range403,6221283027,6752431742,5431141503,243233No. of tribes 14 15 5 24a Information about these systems can be found in Roychoudhury and Nei (1988) and Vogel and Motulsky (1996). The allele frequency data can be obtained from S. M. CallegariJacques ([email protected]) by request.View Table ImageInformation about variability at the DNA level in Native Americans is much more limited, but allele frequencies for 13 STRP loci, namely, CSF1PO, D3S1358, D5S818, D7S820, D8S1179, D13S317, D16S539, D18S51, D21S11, FGA, TH01, TPOX, and VWA, were available for one Carib (Wai Wai, n = 29), one Ge (Xavante, n = 33), and three Tupi (Gavio, Suru, and Zor, total n = 77) tribes (Hutz et al. 2002). An additional Maipurean tribe, the Wayuu, had been studied in relation to STRPs (Guarino et al. 1999), but because the number of loci determined did not match the above set this sample was not considered in the study.All the calculations necessary to test the different hypotheses were done with the COVAR.EXE and TREES.EXE programs, kindly provided by J. C. Long of the Department of Human Genetics and Center for Statistical Genetics, University of Michigan Medical School. For these softwares, inputs of the allele frequencies and the desired tree models are introduced by the user. The programs allow for two kinds of data transformation (arcsine and geneidentity). Constant or variable rates of evolution are also available options of the programs.Table 1 presents the number of individuals and of samples tested for the classical systems in populations of the four linguistic families (see appendix for list). There is wide variation in the number of individuals or samples studied for each of the 37 genetic systems considered, but the averages ranged from 8 samples and 1,355 subjects (Ge) to 14 samples and 2,560 individuals (Carib). The maximum number of individuals considered was 17,083, and each language family was represented by the weighted average of the gene frequencies obtained in the samples. The populations were spread all over the continent (fig. 2) without obvious differences in range, although the Maipure and Carib occupy a more northern area.Fig. 2. Geographic distribution of the populations studied for 37 classical genetic systems, according to linguistic classification. Triangles, Maipure; circles, Carib; crosses, Ge; inverted triangles, Tupi.View Large ImageDownload PowerPointIn comparing the people of these four language families, it is important to verify that they do not significantly differ in relation to interethnic admixture. The relevant data are presented in Table 2. The calculations were performed using three methods. The one developed by Szathmary and Reed (1978) relies on individual markers that are present in nonIndian ethnic groups only. Long and Smouses (1983) estimates the amount of admixture using a weightedleastsquares solution with multinomial sampling schemes considered in all populations involved. Chakraborty (1985) estimates the contribution of the parental populations to the hybrid based on the expected relationship between gene identities (in Neis [1972] sense) among populations. Different combinations of systems were used depending on the particular characteristics of the method employed. The estimates obtained with the different approaches do not differ significantly, indicating an admixture rate of at most 7%.Table 2 Estimated Genetic Contribution (G) and Poisson Distribution 95% Confidence Intervals of Different Parental Populations to Four South American Linguistic Groups as Measured by Three Alternative Methods and Different MarkersMethodProportion of Admixture in Different Language GroupsMaipureCaribGeTupiSzathmary and Reed (1978) 8 informative systemsa Native0.962 (0.9550.967)0.948 (0.9430.953)0.978 (0.9730.982)0.980 (0.9760.984)European0.038 (0.0330.045)0.052 (0.0470.057)0.022 (0.0180.027)0.020 (0.0160.024)10 informative systemsb Native0.975 (0.9710.979)0.959 (0.9550.963)0.987 (0.9840.990)0.980 (0.9760.983)African0.025 (0.0210.029)0.041 (0.0370.045)0.013 (0.0100.016)0.020 (0.0170.024)Long and Smouse (1983) 15 polymorphic systemsc Native0.986 (0.9661.000)0.979 (0.9501.000)0.992 (0.9571.000)0.993 (0.9711.000)African0.014 (0.0000.034)0.021 (0.0000.050)0.008 (0.0000.043)0.007 (0.0000.029)Chakraborty (1985) 29 systemsd Native0.980 (0.9291.000)0.979 (0.9361.000)0.934 (0.8071.000)0.980 (0.8881.000)European0.000 (0.0000.000)0.008 (0.0000.039)0.066 (0.0000.193)0.000 (0.0000.000)African0.020 (0.0000.071)0.013 (0.0000.074)0.000 (0.0000.000)0.020 (0.0000.112)a ABO, ACP, AK, G6PD, GM, KELL, LUTHERAN, and RH, in a total of 10 markers.b ABO, CA2, CP, GM, G6PD, HBA, LUTHERAN, PEPA, PGD, and RH, in total of 12 markers.c ACP, DI, ESD, DUFFY, GC, GLO, GM, HP, KIDD, KM, LEWIS, MNSs, P, PGM1, and RH. No trihybrid admixture was identified with this method.d Those listed in Table 1 except ALB, CHE1, ESA, GPI, HBA2, HBFET, HLAC, and MDH.View Table ImageResultsThe P values obtained in the tests of the hypotheses of Greenberg, Loukotka, and Rodrigues using classical genetic systems data are given in Table 3. Three approaches were used in the analysis: (1) the more comprehensive set of 37 genetic systems; (2) the 18 more polymorphic loci (namely, those in which the frequency of the most common allele is less than 0.90), to minimize the effect of genetic drift; and (3) a more reduced set of 15 systems, designed to reduce both the effects of drift and the eventual effect of selection on HLA. For each set of data, four alternative calculations were performed according to the type of gene frequency transformation (arcsine or geneidentity) and the evolutionary rate (variable or constant) assumed.Table 3 P Values Obtained in Testing Three Alternative Patterns of Relationships among South American Native Linguistic Families Using Blood Group and Protein Genetic DataGenetic Systems and Data Transformation ConsideredProposed Linguistic Relationship and Evolutionary RateGreenbergLoukotkaRodriguesVariableConstantVariableConstantVariableConstantAll 37 systemsa Arcsine0.42070.00990.47770.47560.59930.0202Gene identity0.4012<0.00010.48350.05150.5270<0.000118 polymorphic systemsb polymorphic The genetic systems are listed in Table ACP, DUFFY, ESD, GC, GLO, GM, HLAC, HP, KIDD, KM, LEWIS, MNSs, P, PGM1, ACP, DUFFY, ESD, GC, GLO, GM, HP, KIDD, KM, LEWIS, MNSs, P, PGM1, Table null hypothesis was rejected in 15 of 18 tests evolutionary There is a fit for all three alternative linguistic the genetic data are considered values from to a case of This with previous results obtained for markers in South American (e.g., et al. that the evolutionary rate is variable for different the tests were done variable a better fit was observed of the type of transformation (arcsine or geneidentity) used in the allele For the first and sets of data, of the three linguistic hypotheses is rejected values from to but a different picture is found the set of genetic systems is considered. both and are rejected at the level of of the type of transformation used in the gene frequencies values from to hypothesis is not rejected with the arcsine = or the = transformation of the agreement of the linguistic models with the much more variable 13 STRP data was with a sample of loci and In this the Carib and are represented by a while the three Tupi populations studied live in to each other and are classified in the same linguistic Information about all these systems was for the Maipure, us from between the hypotheses proposed by and Rodrigues. rates of evolution was rejected for both of transformation from to of the other models were rejected values from to the agreement of tree models other than the three considered we tested the hypotheses for the remaining 12 using the set of classical genetic systems and the arcsine For all 12 tree the P values were larger than that obtained for values above P value observed for tree above and were larger than P = the figure observed for trees and to the obtained using two genetic distances and the neighborjoining These models were not rejected = and which means that there is a tree that the genetic data P values for the same models using 37 genetic systems were and same calculations were done for the STRP data, available for three language families Carib, Ge, and Tupi. there was only one additional bifurcating trees for these language families, one that linked Tupi and Ge, with Carib in a more For this not by these the P values obtained with the arcsine and were and and between linguistic and genetic variability be performed with For instance, the type of variation The is follows while language is that it can be both to and differences in the of also an important in linguistic an of this being the of a whole group by with the total of a The rate of of genetic and linguistic characteristics is much in the latter than in the these there are common to linguistic and genetic evolution that are for the observed in Human populations, in often The of two or more groups of both and and the of this is a of the time since the languages, some stocks have been identified in the there are families and languages in the region Central America South America In relation to the four South American linguistic families considered Rodrigues (1985) has in of a strong on of and as well as related to personal case and other and words, and of elements of and and and bifurcating pattern of for both genetic and linguistic evolution us to a method for treeness testing developed by CavalliSforza and Piazza (1975) to the of of different patterns of linguistic relationships to the available genetic data. This method is based on the assumption that populations that split according to a given pattern of and the have a variancecovariance matrix that the history of these as this one can be used to test whether an observed matrix is with a given using any of data. The assumption that the genetic data treeness is because if they do there is no for testing models suggested by other kinds of data. The treeness of the genetic markers considered in this study was evaluated and not the of the of genetic systems to minimize the effects of genetic drift and selection in large samples of four lowland language families, hypothesis was the only one not possible tree the relationships among the language families not considered by the three linguists were identified but are to the present (1987) classified all Native American languages into three large by other of is and rejected The of an has been criticized by several linguists (e.g., Campbell and and by geneticists as well et al. on the of Ychromosome variation Hunley and Long have the relationship between genetics and language classification of Native North American populations using DNA data and by classification. has been criticized by Campbell but by found classification as as genetic are do not to have received much interpretation of these be as follows: Genetic data support other tree those proposed by Greenberg, Loukotka, and Rodrigues. This is not because different kinds of data may different estimates of the history of the populations. the three proposed by these scholars, that of Rodrigues has the genetic In The and of Maipure Carib Ge Tupi Table DNA variation and language American of Human Genetics in A. J. and of The of human and the history of anthropology in human and linguistic to

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