: Surnames are a unique bio-cultural trait, which provide a convenient means of investigating microevolution in hu- man populations. The patrilineal mode of inheritance of surnames mimics highly polymorphic genes on the Y-chro- mosome; the non-biological nature of its dispersal is ex- pected to be independent of fertility and mortality differentials and therefore satisfies the expectations of the neutral theory of evolution (Kimura, 1980), which has been described by Karlin and MacGregor (1967) as the theoretical distribution of different mutant forms that are maintained in a population under the equilibrium between random genetic drift, mutation, and migration. The occur- rence of different surnames in human populations also conforms to Fisher’s logarithmic (Chakraborty et al, 1981) and Pareto’s discrete (Fox 1983) distributions. The earliest application of theoretical models to the sur- name distribution has been considered among the parishes in the Parma valley (Yasuda et al, 1974) and in island populations of Sardinia (Zei et al, 1983a, b). Since then, there has been progress in the use of isonymy studies to investigate the genetic structure among a wide variety of populations, particularly in Europe and Latin American countries (Barrai et al, 1987; Barrai et al, 2003; Colanto- nio et al, 2003). These studies have shown that the ____________________________________________________________ Address for correspondence: T. S. Vasulu, [email protected]Received: 26 Oct 2007; accepted: 05 Feb 2008. logarithmic distribution of surname frequency follows a linear trend (Barrai et al, 1987) in conformity with the neutral allele model (Kimura, 1983); however, a few studies have shown deviation from the expected linear trend (Barrai et al, 1987; Barrai et al, 2002; Barrai et al, 2003). These isonymy studies have revealed (a) geo- graphical patterns in surname distributions as a result of differential migration as demonstrated in Ferrara, an immigrant community, Italy (Barrai et al, 1987; Barrai et al, 1989; Barrai et al, 1990), Austria (Barrai et al, 2000), Perugia (Rodriguez et al, 1993), Spain (Rodriguez et al, 2003); (b) the reflection of social and natural selection in Denmark (Boldsen 1992); (c) devi- ation from linearity as a result of excesses of surname repetition especially, in Netherlands (Barrai et al, 2002), Sicily (Scapoli et al, 1997) and Belgium (Barrai et al, 2003). The majority of the cited studies are based on large sample sizes consisting of an entire nation or region; however, such studies among small isolated populations have rarely been conducted. In this regard, it will be of interest to examine the expectation of neutrality of surname distribution in small populations, especially among isolated tribes, since the demographic events and marriage practices bring rapid changes along the kinship lines influencing the surname struc- ture and in such situation the neutrality of surname distribution may not be expected. This study describes the surname distribution and ex- amines the neutral allele model in an isolated small tribe , Adi Panggi--one of the several sub-tribes of the Adi tribal cluster--inhabiting the Upper Siang River Valley in central Arunachal Pradesh State, India. Suvendu Maji and T. S. Vasulu 1
Transcript
Surnames are a unique bio-cultural trait which provide aconvenient means of investigating microevolution in hu-man populations The patrilineal mode of inheritance ofsurnames mimics highly polymorphic genes on the Y-chro-mosome the non-biological nature of its dispersal is ex-pected to be independent of fertility and mortalitydifferentials and therefore satisfies the expectations of theneutral theory of evolution (Kimura 1980) which hasbeen described by Karlin and MacGregor (1967) as thetheoretical distribution of different mutant forms that aremaintained in a population under the equilibrium betweenrandom genetic drift mutation and migration The occur-rence of different surnames in human populations alsoconforms to Fisherrsquos logarithmic (Chakraborty et al 1981)and Paretorsquos discrete (Fox 1983) distributions
The earliest application of theoretical models to the sur-name distribution has been considered among the parishesin the Parma valley (Yasuda et al 1974) and in islandpopulations of Sardinia (Zei et al 1983a b) Since thenthere has been progress in the use of isonymy studies toinvestigate the genetic structure among a wide variety ofpopulations particularly in Europe and Latin Americancountries (Barrai et al 1987 Barrai et al 2003 Colanto-nio et al 2003) These studies have shown that the
Address for correspondence T S Vasulu vasulugmailcom
Received 26 Oct 2007 accepted 05 Feb 2008
logarithmic distribution of surname frequency followsa linear trend (Barrai et al 1987) in conformity with theneutral allele model (Kimura 1983) however a fewstudies have shown deviation from the expected lineartrend (Barrai et al 1987 Barrai et al 2002 Barrai et al2003) These isonymy studies have revealed (a) geo-graphical patterns in surname distributions as a resultof differential migration as demonstrated in Ferrara animmigrant community Italy (Barrai et al 1987 Barraiet al 1989 Barrai et al 1990) Austria (Barrai et al2000) Perugia (Rodriguez et al 1993) Spain(Rodriguez et al 2003) (b) the reflection of social andnatural selection in Denmark (Boldsen 1992) (c) devi-ation from linearity as a result of excesses of surnamerepetition especially in Netherlands (Barrai et al2002) Sicily (Scapoli et al 1997) and Belgium (Barraiet al 2003) The majority of the cited studies are basedon large sample sizes consisting of an entire nation orregion however such studies among small isolatedpopulations have rarely been conducted In this regardit will be of interest to examine the expectation ofneutrality of surname distribution in small populationsespecially among isolated tribes since the demographicevents and marriage practices bring rapid changesalong the kinship lines influencing the surname struc-ture and in such situation the neutrality of surnamedistribution may not be expected
This study describes the surname distribution and ex-amines the neutral allele model in an isolated smalltribe Adi Panggi--one of the several sub-tribes ofthe Adi tribal cluster--inhabiting the Upper Siang RiverValley in central Arunachal Pradesh State India
Suvendu Maji and T S Vasulu
1
2
The Adi Tribe consists of several sub-tribes inhabitingdifferent altitudes of the southeastern part of Himalayanmountain terrain along the Siang river valley in the cen-tral region of the State of Arunachal Pradesh (Roy 1960Singh 1994 Blackburn 2004 Lego 2005) In general anAdi tribal village consists of a group of families belongingto a few specific clans living together at different locationsof the valley Adi Panggi is one of the smallest isolatedsub-tribes They reside in 7 villages situated in differentvalleys or hill slopes over an area of about 50 square kmin Geku Circle Upper Siang district and number aboutfour thousand individuals (Koley 2005) The sub-tribespeaks the Adi language a member of the Tibeto-Burmanlinguistic family The northern part of the region sharesa border with China Although the Adi sub-tribes sharea common historical migration possible common originand linguistic and cultural affiliations each sub-tribeforms a separate group that maintains its identity Eachsub-tribe is geographically isolated practices high endog-amy and has specific clans and surname structure differ-ent from other sub-tribes of the Adi (Roy 1960 Lego2005 Koley 2005) Like other tribes in India they arepatrilocal patrilineal and patriarchal sons tend to stayin the village or nearby villages while daughters migrateto their husbandrsquos village after their marriage and adopttheir husbandrsquos surname (Das 1953)
The clan and surname is indicative of their putativeorigin from their possible common ancestral stock andindicates genetic kinship The surname structure playsan important role in their marriage warfare and inhunting and cultural activities and are stable over gener-ations These factors make surname analysis useful forinvestigating the genetic structure of the populationThere are only a few studies on the Adi Panggi egABO polymorphism (Bhattacharjee 1954 Krithika etal 2006) cultural aspects (Sharma 1960) anthropo-metric variation (Roy 1966) and a recent ethnographicstudy (Koley 2005) have been reported
Demographic data and blood samples of Adi Panggi(Pangi) tribal population were collected from six vil-lages in Geku circle ( ) for a molecular popula-tion genetics study among the Adi tribe of ArunachalPradesh India The study was approved by the ethicalcommittee of Indian Statistical Institute Kolkata Thegenetic analysis will be the subject of an upcomingarticle
For the surname analysis the surname of the husbandand the lsquomaiden surnamersquo of the wife were collectedthrough pedigree data from a field survey in 2006 (Majiet al 2007) The surname data were collected from 154husbands and 130 wives Of the seven villages sixvillages were studied for surname distribution Sumsing
GekuGeku
Map showing the location of different Panggi villages ( bull ) in the Upper Siang Districtalong the Siang River Valley in Arunachal Pradesh India
Maji Selective Neutrality of Surname Distribution in Adi Panggi 3
for the total sample was considered separately in thepresent study This distribution is expected to show alinear trend under the assumptions of the neutral allelemodel (Kimura 1983) We also have considered thePareto distribution (Fox 1983) where the relationshipbetween the logarithms of and is expected to be
Sur-Name
ID
Sur-Name
Hus-bands
WivesPanggiOnly
WivesPanggi
and Non-Panggi
N=154 N=130 N=147
1 Paron 1558 1692 1497
2 Panyang 2208 2923 2585
3 Taku 584 1000 884
4 Panggeng 844 385 340
5 Tagi 779 385 340
6 Mongku 1039 308 272
7 Tosang 260 231 204
8 Jopir 065 000 000
9 Taying 195 231 204
10 Tatung 130 154 136
11 Ejing 844 385 340
12 Paloh 390 308 272
13 Padun 130 538 476
14 Tateh 390 308 272
15 Tayom 130 154 136
16 Aje 195 000 000
17 Tarang 195 231 204
18 Taruk 065 385 340
19 Teksin - 077 068
20 Kirom - 077 068
21 Gete - 154 136
22 Tangu - 077 068
23 Non-Panggi
(15)
- - 1156
2ipI or 2
iqI and ii qpI
(SS) and Sibum (SB) are remotely located around 15-30km away from Geku Town (GT) whereas the remainingthree villages Ramku (RK) Kumku (KK) and Peram(PR) are located close to GT
Random isonymy (I) was calculated from the frequencydistribution of the abundance of each surname separatelyin males ( p i ) and females ( q i ) and for both in each( i t h ) village If p i and q i are the frequencies of aparticular surname in males (husbands) and females(wives) respectively then the random isonymy estimateamong males females and both males and females is
Since the random isonymy is biased for the samplesrsquo sizevariations an unbiased isonymy and its variance wasestimated (Dyke et al 1983 Relethford 1988 Barrai etal 1989 Barrai et al 1991) If there are Ni males and Nj
females in the samples the unbiased random isonomy I i jis defined as
I i j = ( n i s bull njs) Ni bull Nj
where nis and njs are the numbers of individuals withsurname in populations i and j respectively and Ni andNj are the total number of surnames in populations i andj and summation is over all s surnames
Since the surnames mimic a genetic trait on the Y-chro-mosome and the surname distribution in a large popula-tion also correlates with genetic diversity (Barrai et al1996) it is expected to conform under certain assump-tions to the selection neutrality of an infinite allele modeldescribed by Karlin and MacGregor (1967) which can beaccounted for by the logarithmic distribution first givenby Fisher That is if we let S represent the number oftimes that a surname occurs in a population and let Krepresent the frequency of S (ie K is the number ofsurnames occurring this same number of S times) thenwe may characterize the distribution by graphing log Kversus log S Karlin and MacGregorrsquos
proposed by Zei et al (1983a 1983b) and Barrai et al(1992) respectively where
I i j ( N i )
and I is random isonymy N i is the total number ofindividuals and
Further the l o g 2 S - l o g 2 K distribution of surnameoccurrence (S) and its frequency K for husband wife and
4
linear where is the number of individuals and is thenumber of surnames but the results are presented for thelog-log distribution only
Occurrence of surname with respect to variation in prog-eny size distribution (separately for male female andtotal children) among the post-reproductive women andtheir husbands were analyzed to investigate the pattern ofsurname distribution in a system of patrilocal marriagewhere males (kin group especially brothers) reside in thesame village and females move out of the village aftertheir marriage Similar analysis was carried out for thecase of reproductive wives and their husbandsrsquo sample
The occurrence of different surnames among husbandsand wives of the Adi Panggi tribe are shown in There are 18 different surnames among 154 husbandsand 22 surnames among 130 wives among the Panggi
GT PR RK KM SS SB
Husband01144 01606 00966 00560 01083 02750
42612 44005 12832 10085 49796 106207
Wife(Panggi)
00967 03444 06965 01467 01334 03131
25678 94574 252379 25790 53877 117380
Wife(Panggi andNon-Panggi)
01679 04706 07450 01827 01607 03444
56482 213309 408934 35767 70828 147090
tribe distributed over six villages There are 15 sur-names of non-Panggi origin that have filtered into thepopulation through marriages with non-Panggi wivesHusbands represent a smaller number of surnameswhereas wives have more diversity of surnames Onaverage a single surname is shared by about 85 ofhusbands while in the case of wives it is about 6 Thethree most common surnames among husbands includeParon Panyang and Mongku which represent about47 of the husbands In case of wives the surnamesPanyang Paron and Taku are the three most commonand occur among 57 of Panggi wives These sevensurnames (which have a frequency above 5) represent785 of the husbands and 615 of thei wives Themost frequent surnames occur among both husbandsand wives with similar percentages and the seven mostfrequent (above 5 occurrence) constitute 707 ofthe sample The sample contains four singly occurringPanggi surnames representing 13 percent of the totalindividuals Considering both Panggi and non-Panggisurnames there are 18 singletons that constitute 6 of
Sample Size F F
Husband 154 00344 54830 00513 83278
Wife(Panggi) 130 00555 76390 00483 66020
Wife(Panggi andNon-Panggi)
147 09010 147215 00559 87113
F and F are distributions based upon random isonymy
Maji Selective Neutrality of Surname Distribution in Adi Panggi 5
Log-log distribution of surnames in Adi Panggi husbands and wives
A) Distribution of surnames among Adi Panggi Husbands
the sample The unbiased random isonymy I i j for theAdi Panggi samples is 01025
shows the estimates the two parametersof Karlin-MacGregor and Fishersrsquo distributions that de-scribe the differential migration rates and surname diver-sity among six villages of Adi Panggi Both the husbandsand wives show wide variation in the rate of migrationand surname diversity between the villages The twovillages KM and RK show the least migration rates andsurname diversity whereas SB shows the highest values of
in case of the husband samples
Panggi wives show higher surname diversity and migra-tion rates than husbands except in case of GT Panggiand non-Panggi wives show higher values than husbands( ) The estimates of F F based on randomisonymy show lesser values than from the Karlin-MacGregor and Fishersrsquo distributions The results showa similar pattern among husbands (n F = 00513 and a F
= 83278) and Panggi plus non-Panggi wives (n F = 00559and a F = 87113)
The logarithmic distribution of surname occurrenceamong husbands and wives and in the total sample for thePanggi is shown in
Both the figures show a non-linear trend suggesting devi-ation from the expected neutral allele model The shapeof the distribution is different in husbands and wives it is
an inverted parabolic curve in males and a slightlyconcave curve in females The best non-linear fit of thecurve with polynomial degree four show a variance ofabout 60 (Rsup2 = 059) in the case of the husbandssample while for wives a polynomial degree threeshows a good fit to the distribution (Rsup2 = 080) whichis better than the linear fit (Rsup2 = 074) In the case ofall Adi Panggi (both husbands and wives) the distribu-tions show a nonlinear trend which is different fromthe trend observed in the case of husband and wifesamples ( ) The best fit to the curve is apolynomial of degree four which accounts for about96 of the variance The non-linear distribution is dueto the higher frequency of occurrence of rare surnames(especially those that occur only once and are contrib-uted by wives) However the truncated log-log distri-bution (with the rare surnames excluded) shows alinear trend
The surname distribution observed among husbandsand wives has also been considered with respect to theirprogeny size to investigate variation of fertility differ-entials with respect to surnames among the post-repro-ductive wives and their husbands separately for malefemale and total progeny size The results of thedistribution show a non-linear trend suggesting a dif-ferential reproductive fitness for different surnamesamong husbands and wives ( ) In the case ofhusbands the rare surnames (occurring once or twice)and common surnames (occurring more than 6-9
Maji Selective Neutrality of Surname Distribution in Adi Panggi 7
Occurrence of surname distribution (s) total number of male ( ) female ( ) andtotal children (times) among Adi Panggi husbands and wives (above 45 Years) and progeny sizedistribution (--)
times) show a trend toward higher progeny size (malesfemales and total children) than those surnames thatoccur less frequently (between 3-5 times) The best fitcurve in case of total children (polynomial degree three)accounts for about 945 percent of the variance (Rsup2 =0945) In case of females the trend shows a negativeassociation between occurrence of surnames and theprogeny size and the best fit curve for the total childrenaccounts (polynomial degree three) about 99 percent ofthe variance (Rsup2 = 0989) Progeny size shows a decreas-
ing trend with the rare surnames that occur once ortwice and a trend of lesser progeny in case of morecommon surnames (except in case the most commonsurnames that occur ten times among wives) Theseresults are a reflection of the population structure of thetribe especially the patrilocal system of marriage whichtends to increase the prevalence of some specific sur-names as a result of preferential marriages among a fewspecific clans or surnames
Husband ( gt 4 5 )
05
101520253035
0 2 4 6 8 10
Occurrence (s)
FrequencyW i f e ( gt 4 5 )
0
10
20
30
40
0 2 4 6 8 10
Occurrence (s)
Frequency
8
imilar distribution of progeny size with respect to sur-name distribution was also considered in case of repro-ductive wives and their husbandsrsquo sample ( )The pattern of distributions is quite varied compared tothe case of post-reproductive samples In the case of thewives the progeny size shows an increasing trend fromthe rare surnames to the most frequent surnamesthough there is some perturbance at the initial case ofrarer ones Among the husband samples a similar initialperturbance of progeny size is noticed with the rarersurnames but the progeny size shows stability with themore commonly occurring surnames of the tribe
Occurrence of surname distribution(s) total number of male ( ) female ( ) and totalchildren (times) among Adi Panggi husbands and wives (below 45 years) and progeny size distribution (--)
Husband (lt45)
0
10
20
30
40
50
60
0 5 10 15 20 25 30
Occurrence (s)
Frequency
W i f e ( lt 4 5 )
0
20
40
60
80
100
0 5 10 15 20 25 30
Occurrence (s)
Frequency
Though selective neutrality of surname distribution hasbeen validated in a number of populations in Europe andLatin American countries a few studies have showndeviation from neutrality implicating the role of migra-tion and operation of selection with respect to certainsurnames The results of the present study indicate thatsuch trends are also observed in small isolated popula-tions Apart from the expected influences of migrationand drift there could be other related factors of popula-tion structure that could cause the deviation from neutral-
Maji Selective Neutrality of Surname Distribution in Adi Panggi 9
ity In small tribal populations the structure of thesurname distribution might vary widely depending on thehistory of settlement prevailing marriage patterns andother socio-cultural and environmental reasons Furtherthe demographic parameters can vary drastically owing totheir subsistence pattern epidemiological factors andsocial organization such as warfare history Also internaltribal factions which are formed based on kinship or clanor lineages involving several closely knit families (eg inSouth Amerindian tribes Neel 1968 Salzano et al 1967)may play a role This can significantly influence thepopulation structure and can lead to perturbations in thesurname distribution in short time periods and in suchcases the neutral theory predictions may not hold Someof the above factors are important in explaining thedeviation from neutrality in surname distribution amongthe Adi Panggi
In general the Adi tribe consists of several sub-tribes whooccupy different terrains in the upper Himalayan moun-tain ranges and some in plains in the northeastern part ofthe country (Roy 1960 Lego 2005) The formation of thesub-tribes followed the fission-fusion model of popula-tion structure and was triggered by intertribal factionscompeting for survival necessities in this hunting-gather-ing population (Neel 1968 Salzano et al 1967) Theformation of Adi Panggi sub-tribes occurred a couple ofgenerations ago and was a result of conflict between amajor subgroup of the tribe and some other close-kinfactions over hunting rights and other resources (Roy1960 Lego 2005 Lego 2006) The sub-tribes now occu-py a recognized territory and constitute a few close affinaland consanguineous lineages and their affiliated clansSince they have a patrilocal society a majority of theirmen stay in the home village whereas females come fromdifferent villages and from other Adi sub-tribes Thisresults in more female marital migration and greatersurname diversity among wives We therefore observe ahigher proportion of homonymy of surname diversityamong males For example the five most frequent sur-names represent about 65 of total husbands while raresurnames (that occur either once or twice) constituteabout 5 per cent The three most common surnamesamong wives also represent a large part of the populationbut in contrast with the males the rare surnames thatoccur either once or twice constitute 18 per cent morethan 35 times the proportion of rare surnames in hus-bands In the total sample of husbands and wives sixsurnames represent 69 percent of the total populationstudied The surname distribution therefore reflectsgreater prevalence of most frequent (occurring in 20 to 70cases) and rare surnames (occurring once or twice) asagainst the moderately frequent surnames (that occur 3 to20 times in the sample)
The estimates of unbiased Isonymy also differ betweenhusband and wives samples The combined samples(husband and wives) show higher values of estimatedIsonymy (I = 01025) suggesting higher levels of homozy-gosity as a result of more frequent marriages contracted
among a few selected surnames generally found in suchisolated tribes This is reflected in the Karlin andMacGregor distribution which shows the non-lineartrend and the trend remains the same even after thelogarithmic transformation This also explains the widedifferences between the male and female surname distri-butions and the deviations observed in the Karlin andMacGregor and logarithmic distributions Therefore ifthe truncated distribution (omitting the rare surnames) isconsidered it shows a linear trend in the combinedsample suggesting that the non-linearity is contributedby the occurrence of rare surnames in the sample Therare surnames are contributed more through femalesmigrating from other places However the discretePareto distribution shows a linear trend for both thesamples The linear trend is clearer in the female samplethan among males but the genetic implications of thePareto distributions are not apparent (Fox 1983)
The results of the distribution of surnames also showwide differences with respect to progeny size distributionin case of post-reproductive (completed fertility) andreproductive wives and their husbands The neutralallele model would suggest that the distribution shouldbe independent of the fertility and morality (Manrubia2002) but the present study indicates otherwise andthere appears to be some differential fertility for somesurnames The Adi Panggi sample shows greater progenysize (male female and total surviving children) for themost frequent and rare surnames and that is reflected inthe near U-shaped distribution especially in case of hus-band sample This suggests reproductive advantageespecially greater success for the most frequent (6-910)and rare surnames (1-2) as against the less frequentsurnames (3-6) Among the Adi Panggi the four mostfrequent surnames among husbands and wives (especiallyPanyang Paron Mongku and Taku) show higher proge-ny size among the husband sample whereas the wivessample shows a near linear trend of decreasing progenysize from the rare surnames to the most frequent ones Apaired t-test between (eight) different types of surnamesand their progeny size shows higher significance values incase of the husbands (above 45 years) sample (p =0000433) than among the wives (above 45 years) (p =0003263) The mothers and their husbands were all inpost-reproductive period and had attained their maxi-mum fertility potential and do not use family planningand therefore the possibility of other intervening factorscan be excluded It is possible that it could be also dueto selective mortality of children especially higher mor-tality in case of less frequent surnames than with moreand rare frequent ones and this needs to be verifiedHowever there are no apparent cultural active selectivemechanisms that can influence the progeny size amongsome families of the tribe A detailed study on themarriage pattern and surname or clan distribution mightreveal more details The progeny distribution amongwomen of completed fertility is expected to follow Pois-son and Negative binomial distributions (Cavalli-Sforza
10
1971 Vasulu 1980) In patrilineal societies femalesgenerally show a greater surname diversity than malesand as a result the surname dynamics differ with respectto mitochondria and Y-chromosome
In the Indian context surname analysis has been carriedout in a few populations for investigating the effect onpopulation structure of inbreeding (Bhalla 1976 Kashyap1980 Vasulu 1980) genetic kinship (Kashyap 1980Vasulu 1980) preferential mating (Vasulu 1989 Reddy1995 Maji et al 2007) and social network analysisamong castes and tribes (Rao 1995) Surname neutralityhas also been validated in a caste community of Indiansresiding in Texas (Chakraborty 1990)
The above few studies suggest that isonymy can be ofmuch value in inferring population structure in somecastes and tribes (Kashyap 1980 Maji et al 2007) Thesurname structure in India varies among different regionscastes tribes linguistic and religious groups and providesan opportunity to study microevolution and populationstructure where the assumptions of surname or isonymyanalysis are valid Its applicability in diverse populationsespecially among isolated populations is expected to yieldfurther insights of its validity and inferring populationstructure and microevolutionary history
The present study illustrates the importance of surnamein inferring the genetic structure among a small sub-tribeAdi Panggi an isolated Tibeto-Burman- speaking com-munity in Arunachal Pradesh state in the northeasternregion of India The tribe lives on shifting-hunting-and-gathering subsistence and is a patrilocal and patrilinealsociety where the marriage pattern involves movement ofwives from other villages to the husbandrsquos village whichallows more surname diversity through females As aconsequence of differential surname structure betweenhusbands and wives the surname distribution showsdeviation from the expected neutral allele model and italso exhibits a non-linear trend of progeny size distribu-tion with respect to surname occurrence in the husbandsample
The results of the present study illustrate that surnamedistribution could be of much value in deciphering thepopulation structure and microevolution of a population(Crow 1965 Lasker 1978 Yasuda et al 1974) Thenon-linear trend observed in this study appears to beunique in the Indian context especially for the smallisolated tribes
The results of the study will help in further confirming thevalidity of the findings in several other tribes in India Inthe absence of any biological or genetic data for a tribethe surnames provide insights into the population struc-ture and possible genetic implications which can help inthe design of future studies As a prelude to the moleculargenetic investigations about the past genetic history asurname analysis can provide considerable insight into
the expected genetic structure of the population It alsohelps us to frame specific hypothesis related to male andfemale migration history which we will be testing withthe molecular genetic studies The two approachestogether can help to unravel the differential paternaland maternal past genetic history of the tribe
We thank the Government of Arunachal Pradesh andall the participants for their active cooperation in ourstudy Our sincere thanks go to Mr Tozing Tagi andMr Oyam Panyang (Katan and Geku circles) for theirassistance in data collection We acknowledge thesupport and intellectual inputs about the Adi tribesfrom Mr R N Koley (ARO Directorate of Research)and Mr Kaling Borang of Pasighat We thank Ms SKrithika Senior Research Fellow Biological Anthro-pology Unit for suggestions and help with the analysisand draft finalization and the Director Indian Statisti-cal Institute Kolkata for the financial assistance for thestudy among the tribes of Arunachal Pradesh
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11
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Maji S Krithika S Vasulu TS (2007) Genetic kinship among anisolated Adi tribe of Arunachal Pradesh isonymy in Adi Panggi
79321-337
Manrubia SC Zanette DH (2002) At the boundary betweenbiological and cultural evolution The origin of surname distri-butions 216461-477
Neel JV Chagnon NA (1968) The demography of two tribes ofprimitive relatively unacculturated American Indians
(USA) 59680-689
Rao AR (1995) Reciprocity in marital and social networksIllustration with Indian data 67887-904
Reddy BM Malhotra KC (1995) Relevance of surname inpopulation structure A study among the Vadde a fishingcommunity of Kolleru lake in Andhra Pradesh India
6161-168
Relethford JH (1988) Estimation of Kinship and Genetic dis-tance from Surnames 60475-492
Rodriguez LA Barrai I Alfonzo JC (1993) Isonymy structureof four Venezuelan states 20131-145
Rodriguez-Larralde A Formica G Scapoli C (1993) Microevo-lution in Perugia Isonymy 1890-1990 20261-274
Rodriguez-Larralde A Gonzales-Martin A Scapoli C Barrai I(2003) The names of Spain A study of the isonymy structureof Spain 121280-292
Roy S (1960) Aspects of Padam Minyong Culture Directorateof Research Govt of Arunachal Pradesh Itanagar
Roy S (1966) Anthropometry of the Adis of the Siang districtNorth-East Frontier Agency Shillong
Salzano FM Neel JV Maybury Lewis D (1967) Further studieson Xavante Indians I Demographic data on two additionalvillages and genetic structure of the tribe19463-489
Sarma J (1960) Puberty Marriage and Childbirth among thePangi and the Minyong Abor women 5597-113
Scapoli C Rodriguez-Larralde A Beretta M (1997) Correlationbetween isonymy parameters 1217-37
Singh IP (1994) Anthropological Survey of India Oxford
University Press New Delhi
Vasulu TS (1980) Anthrop Soc
of India Mysore
Vasulu TS (1989) Genetic structure of a tribal population 1Breeding isolation among the Yanadis 4255-268
Yasuda N Cavalli-Sfroza LL Skolnir M Moroni A (1974) Theevolution of surnames an analysis of their distribution andextinction 5123-142
Zei G Guglielmino CR Siri E Moroni A Cavalli-Sforza LL(1983a) Surnames as neutral alleles Observation in Sardinia
55357-365
Zei G Matessi RG Siri E et al (1983b) Surnames in SardiniaI Fit of frequency distributions for neutral articles and geneticpopulation structure 47329-352
2
The Adi Tribe consists of several sub-tribes inhabitingdifferent altitudes of the southeastern part of Himalayanmountain terrain along the Siang river valley in the cen-tral region of the State of Arunachal Pradesh (Roy 1960Singh 1994 Blackburn 2004 Lego 2005) In general anAdi tribal village consists of a group of families belongingto a few specific clans living together at different locationsof the valley Adi Panggi is one of the smallest isolatedsub-tribes They reside in 7 villages situated in differentvalleys or hill slopes over an area of about 50 square kmin Geku Circle Upper Siang district and number aboutfour thousand individuals (Koley 2005) The sub-tribespeaks the Adi language a member of the Tibeto-Burmanlinguistic family The northern part of the region sharesa border with China Although the Adi sub-tribes sharea common historical migration possible common originand linguistic and cultural affiliations each sub-tribeforms a separate group that maintains its identity Eachsub-tribe is geographically isolated practices high endog-amy and has specific clans and surname structure differ-ent from other sub-tribes of the Adi (Roy 1960 Lego2005 Koley 2005) Like other tribes in India they arepatrilocal patrilineal and patriarchal sons tend to stayin the village or nearby villages while daughters migrateto their husbandrsquos village after their marriage and adopttheir husbandrsquos surname (Das 1953)
The clan and surname is indicative of their putativeorigin from their possible common ancestral stock andindicates genetic kinship The surname structure playsan important role in their marriage warfare and inhunting and cultural activities and are stable over gener-ations These factors make surname analysis useful forinvestigating the genetic structure of the populationThere are only a few studies on the Adi Panggi egABO polymorphism (Bhattacharjee 1954 Krithika etal 2006) cultural aspects (Sharma 1960) anthropo-metric variation (Roy 1966) and a recent ethnographicstudy (Koley 2005) have been reported
Demographic data and blood samples of Adi Panggi(Pangi) tribal population were collected from six vil-lages in Geku circle ( ) for a molecular popula-tion genetics study among the Adi tribe of ArunachalPradesh India The study was approved by the ethicalcommittee of Indian Statistical Institute Kolkata Thegenetic analysis will be the subject of an upcomingarticle
For the surname analysis the surname of the husbandand the lsquomaiden surnamersquo of the wife were collectedthrough pedigree data from a field survey in 2006 (Majiet al 2007) The surname data were collected from 154husbands and 130 wives Of the seven villages sixvillages were studied for surname distribution Sumsing
GekuGeku
Map showing the location of different Panggi villages ( bull ) in the Upper Siang Districtalong the Siang River Valley in Arunachal Pradesh India
Maji Selective Neutrality of Surname Distribution in Adi Panggi 3
for the total sample was considered separately in thepresent study This distribution is expected to show alinear trend under the assumptions of the neutral allelemodel (Kimura 1983) We also have considered thePareto distribution (Fox 1983) where the relationshipbetween the logarithms of and is expected to be
Sur-Name
ID
Sur-Name
Hus-bands
WivesPanggiOnly
WivesPanggi
and Non-Panggi
N=154 N=130 N=147
1 Paron 1558 1692 1497
2 Panyang 2208 2923 2585
3 Taku 584 1000 884
4 Panggeng 844 385 340
5 Tagi 779 385 340
6 Mongku 1039 308 272
7 Tosang 260 231 204
8 Jopir 065 000 000
9 Taying 195 231 204
10 Tatung 130 154 136
11 Ejing 844 385 340
12 Paloh 390 308 272
13 Padun 130 538 476
14 Tateh 390 308 272
15 Tayom 130 154 136
16 Aje 195 000 000
17 Tarang 195 231 204
18 Taruk 065 385 340
19 Teksin - 077 068
20 Kirom - 077 068
21 Gete - 154 136
22 Tangu - 077 068
23 Non-Panggi
(15)
- - 1156
2ipI or 2
iqI and ii qpI
(SS) and Sibum (SB) are remotely located around 15-30km away from Geku Town (GT) whereas the remainingthree villages Ramku (RK) Kumku (KK) and Peram(PR) are located close to GT
Random isonymy (I) was calculated from the frequencydistribution of the abundance of each surname separatelyin males ( p i ) and females ( q i ) and for both in each( i t h ) village If p i and q i are the frequencies of aparticular surname in males (husbands) and females(wives) respectively then the random isonymy estimateamong males females and both males and females is
Since the random isonymy is biased for the samplesrsquo sizevariations an unbiased isonymy and its variance wasestimated (Dyke et al 1983 Relethford 1988 Barrai etal 1989 Barrai et al 1991) If there are Ni males and Nj
females in the samples the unbiased random isonomy I i jis defined as
I i j = ( n i s bull njs) Ni bull Nj
where nis and njs are the numbers of individuals withsurname in populations i and j respectively and Ni andNj are the total number of surnames in populations i andj and summation is over all s surnames
Since the surnames mimic a genetic trait on the Y-chro-mosome and the surname distribution in a large popula-tion also correlates with genetic diversity (Barrai et al1996) it is expected to conform under certain assump-tions to the selection neutrality of an infinite allele modeldescribed by Karlin and MacGregor (1967) which can beaccounted for by the logarithmic distribution first givenby Fisher That is if we let S represent the number oftimes that a surname occurs in a population and let Krepresent the frequency of S (ie K is the number ofsurnames occurring this same number of S times) thenwe may characterize the distribution by graphing log Kversus log S Karlin and MacGregorrsquos
proposed by Zei et al (1983a 1983b) and Barrai et al(1992) respectively where
I i j ( N i )
and I is random isonymy N i is the total number ofindividuals and
Further the l o g 2 S - l o g 2 K distribution of surnameoccurrence (S) and its frequency K for husband wife and
4
linear where is the number of individuals and is thenumber of surnames but the results are presented for thelog-log distribution only
Occurrence of surname with respect to variation in prog-eny size distribution (separately for male female andtotal children) among the post-reproductive women andtheir husbands were analyzed to investigate the pattern ofsurname distribution in a system of patrilocal marriagewhere males (kin group especially brothers) reside in thesame village and females move out of the village aftertheir marriage Similar analysis was carried out for thecase of reproductive wives and their husbandsrsquo sample
The occurrence of different surnames among husbandsand wives of the Adi Panggi tribe are shown in There are 18 different surnames among 154 husbandsand 22 surnames among 130 wives among the Panggi
GT PR RK KM SS SB
Husband01144 01606 00966 00560 01083 02750
42612 44005 12832 10085 49796 106207
Wife(Panggi)
00967 03444 06965 01467 01334 03131
25678 94574 252379 25790 53877 117380
Wife(Panggi andNon-Panggi)
01679 04706 07450 01827 01607 03444
56482 213309 408934 35767 70828 147090
tribe distributed over six villages There are 15 sur-names of non-Panggi origin that have filtered into thepopulation through marriages with non-Panggi wivesHusbands represent a smaller number of surnameswhereas wives have more diversity of surnames Onaverage a single surname is shared by about 85 ofhusbands while in the case of wives it is about 6 Thethree most common surnames among husbands includeParon Panyang and Mongku which represent about47 of the husbands In case of wives the surnamesPanyang Paron and Taku are the three most commonand occur among 57 of Panggi wives These sevensurnames (which have a frequency above 5) represent785 of the husbands and 615 of thei wives Themost frequent surnames occur among both husbandsand wives with similar percentages and the seven mostfrequent (above 5 occurrence) constitute 707 ofthe sample The sample contains four singly occurringPanggi surnames representing 13 percent of the totalindividuals Considering both Panggi and non-Panggisurnames there are 18 singletons that constitute 6 of
Sample Size F F
Husband 154 00344 54830 00513 83278
Wife(Panggi) 130 00555 76390 00483 66020
Wife(Panggi andNon-Panggi)
147 09010 147215 00559 87113
F and F are distributions based upon random isonymy
Maji Selective Neutrality of Surname Distribution in Adi Panggi 5
Log-log distribution of surnames in Adi Panggi husbands and wives
A) Distribution of surnames among Adi Panggi Husbands
the sample The unbiased random isonymy I i j for theAdi Panggi samples is 01025
shows the estimates the two parametersof Karlin-MacGregor and Fishersrsquo distributions that de-scribe the differential migration rates and surname diver-sity among six villages of Adi Panggi Both the husbandsand wives show wide variation in the rate of migrationand surname diversity between the villages The twovillages KM and RK show the least migration rates andsurname diversity whereas SB shows the highest values of
in case of the husband samples
Panggi wives show higher surname diversity and migra-tion rates than husbands except in case of GT Panggiand non-Panggi wives show higher values than husbands( ) The estimates of F F based on randomisonymy show lesser values than from the Karlin-MacGregor and Fishersrsquo distributions The results showa similar pattern among husbands (n F = 00513 and a F
= 83278) and Panggi plus non-Panggi wives (n F = 00559and a F = 87113)
The logarithmic distribution of surname occurrenceamong husbands and wives and in the total sample for thePanggi is shown in
Both the figures show a non-linear trend suggesting devi-ation from the expected neutral allele model The shapeof the distribution is different in husbands and wives it is
an inverted parabolic curve in males and a slightlyconcave curve in females The best non-linear fit of thecurve with polynomial degree four show a variance ofabout 60 (Rsup2 = 059) in the case of the husbandssample while for wives a polynomial degree threeshows a good fit to the distribution (Rsup2 = 080) whichis better than the linear fit (Rsup2 = 074) In the case ofall Adi Panggi (both husbands and wives) the distribu-tions show a nonlinear trend which is different fromthe trend observed in the case of husband and wifesamples ( ) The best fit to the curve is apolynomial of degree four which accounts for about96 of the variance The non-linear distribution is dueto the higher frequency of occurrence of rare surnames(especially those that occur only once and are contrib-uted by wives) However the truncated log-log distri-bution (with the rare surnames excluded) shows alinear trend
The surname distribution observed among husbandsand wives has also been considered with respect to theirprogeny size to investigate variation of fertility differ-entials with respect to surnames among the post-repro-ductive wives and their husbands separately for malefemale and total progeny size The results of thedistribution show a non-linear trend suggesting a dif-ferential reproductive fitness for different surnamesamong husbands and wives ( ) In the case ofhusbands the rare surnames (occurring once or twice)and common surnames (occurring more than 6-9
Maji Selective Neutrality of Surname Distribution in Adi Panggi 7
Occurrence of surname distribution (s) total number of male ( ) female ( ) andtotal children (times) among Adi Panggi husbands and wives (above 45 Years) and progeny sizedistribution (--)
times) show a trend toward higher progeny size (malesfemales and total children) than those surnames thatoccur less frequently (between 3-5 times) The best fitcurve in case of total children (polynomial degree three)accounts for about 945 percent of the variance (Rsup2 =0945) In case of females the trend shows a negativeassociation between occurrence of surnames and theprogeny size and the best fit curve for the total childrenaccounts (polynomial degree three) about 99 percent ofthe variance (Rsup2 = 0989) Progeny size shows a decreas-
ing trend with the rare surnames that occur once ortwice and a trend of lesser progeny in case of morecommon surnames (except in case the most commonsurnames that occur ten times among wives) Theseresults are a reflection of the population structure of thetribe especially the patrilocal system of marriage whichtends to increase the prevalence of some specific sur-names as a result of preferential marriages among a fewspecific clans or surnames
Husband ( gt 4 5 )
05
101520253035
0 2 4 6 8 10
Occurrence (s)
FrequencyW i f e ( gt 4 5 )
0
10
20
30
40
0 2 4 6 8 10
Occurrence (s)
Frequency
8
imilar distribution of progeny size with respect to sur-name distribution was also considered in case of repro-ductive wives and their husbandsrsquo sample ( )The pattern of distributions is quite varied compared tothe case of post-reproductive samples In the case of thewives the progeny size shows an increasing trend fromthe rare surnames to the most frequent surnamesthough there is some perturbance at the initial case ofrarer ones Among the husband samples a similar initialperturbance of progeny size is noticed with the rarersurnames but the progeny size shows stability with themore commonly occurring surnames of the tribe
Occurrence of surname distribution(s) total number of male ( ) female ( ) and totalchildren (times) among Adi Panggi husbands and wives (below 45 years) and progeny size distribution (--)
Husband (lt45)
0
10
20
30
40
50
60
0 5 10 15 20 25 30
Occurrence (s)
Frequency
W i f e ( lt 4 5 )
0
20
40
60
80
100
0 5 10 15 20 25 30
Occurrence (s)
Frequency
Though selective neutrality of surname distribution hasbeen validated in a number of populations in Europe andLatin American countries a few studies have showndeviation from neutrality implicating the role of migra-tion and operation of selection with respect to certainsurnames The results of the present study indicate thatsuch trends are also observed in small isolated popula-tions Apart from the expected influences of migrationand drift there could be other related factors of popula-tion structure that could cause the deviation from neutral-
Maji Selective Neutrality of Surname Distribution in Adi Panggi 9
ity In small tribal populations the structure of thesurname distribution might vary widely depending on thehistory of settlement prevailing marriage patterns andother socio-cultural and environmental reasons Furtherthe demographic parameters can vary drastically owing totheir subsistence pattern epidemiological factors andsocial organization such as warfare history Also internaltribal factions which are formed based on kinship or clanor lineages involving several closely knit families (eg inSouth Amerindian tribes Neel 1968 Salzano et al 1967)may play a role This can significantly influence thepopulation structure and can lead to perturbations in thesurname distribution in short time periods and in suchcases the neutral theory predictions may not hold Someof the above factors are important in explaining thedeviation from neutrality in surname distribution amongthe Adi Panggi
In general the Adi tribe consists of several sub-tribes whooccupy different terrains in the upper Himalayan moun-tain ranges and some in plains in the northeastern part ofthe country (Roy 1960 Lego 2005) The formation of thesub-tribes followed the fission-fusion model of popula-tion structure and was triggered by intertribal factionscompeting for survival necessities in this hunting-gather-ing population (Neel 1968 Salzano et al 1967) Theformation of Adi Panggi sub-tribes occurred a couple ofgenerations ago and was a result of conflict between amajor subgroup of the tribe and some other close-kinfactions over hunting rights and other resources (Roy1960 Lego 2005 Lego 2006) The sub-tribes now occu-py a recognized territory and constitute a few close affinaland consanguineous lineages and their affiliated clansSince they have a patrilocal society a majority of theirmen stay in the home village whereas females come fromdifferent villages and from other Adi sub-tribes Thisresults in more female marital migration and greatersurname diversity among wives We therefore observe ahigher proportion of homonymy of surname diversityamong males For example the five most frequent sur-names represent about 65 of total husbands while raresurnames (that occur either once or twice) constituteabout 5 per cent The three most common surnamesamong wives also represent a large part of the populationbut in contrast with the males the rare surnames thatoccur either once or twice constitute 18 per cent morethan 35 times the proportion of rare surnames in hus-bands In the total sample of husbands and wives sixsurnames represent 69 percent of the total populationstudied The surname distribution therefore reflectsgreater prevalence of most frequent (occurring in 20 to 70cases) and rare surnames (occurring once or twice) asagainst the moderately frequent surnames (that occur 3 to20 times in the sample)
The estimates of unbiased Isonymy also differ betweenhusband and wives samples The combined samples(husband and wives) show higher values of estimatedIsonymy (I = 01025) suggesting higher levels of homozy-gosity as a result of more frequent marriages contracted
among a few selected surnames generally found in suchisolated tribes This is reflected in the Karlin andMacGregor distribution which shows the non-lineartrend and the trend remains the same even after thelogarithmic transformation This also explains the widedifferences between the male and female surname distri-butions and the deviations observed in the Karlin andMacGregor and logarithmic distributions Therefore ifthe truncated distribution (omitting the rare surnames) isconsidered it shows a linear trend in the combinedsample suggesting that the non-linearity is contributedby the occurrence of rare surnames in the sample Therare surnames are contributed more through femalesmigrating from other places However the discretePareto distribution shows a linear trend for both thesamples The linear trend is clearer in the female samplethan among males but the genetic implications of thePareto distributions are not apparent (Fox 1983)
The results of the distribution of surnames also showwide differences with respect to progeny size distributionin case of post-reproductive (completed fertility) andreproductive wives and their husbands The neutralallele model would suggest that the distribution shouldbe independent of the fertility and morality (Manrubia2002) but the present study indicates otherwise andthere appears to be some differential fertility for somesurnames The Adi Panggi sample shows greater progenysize (male female and total surviving children) for themost frequent and rare surnames and that is reflected inthe near U-shaped distribution especially in case of hus-band sample This suggests reproductive advantageespecially greater success for the most frequent (6-910)and rare surnames (1-2) as against the less frequentsurnames (3-6) Among the Adi Panggi the four mostfrequent surnames among husbands and wives (especiallyPanyang Paron Mongku and Taku) show higher proge-ny size among the husband sample whereas the wivessample shows a near linear trend of decreasing progenysize from the rare surnames to the most frequent ones Apaired t-test between (eight) different types of surnamesand their progeny size shows higher significance values incase of the husbands (above 45 years) sample (p =0000433) than among the wives (above 45 years) (p =0003263) The mothers and their husbands were all inpost-reproductive period and had attained their maxi-mum fertility potential and do not use family planningand therefore the possibility of other intervening factorscan be excluded It is possible that it could be also dueto selective mortality of children especially higher mor-tality in case of less frequent surnames than with moreand rare frequent ones and this needs to be verifiedHowever there are no apparent cultural active selectivemechanisms that can influence the progeny size amongsome families of the tribe A detailed study on themarriage pattern and surname or clan distribution mightreveal more details The progeny distribution amongwomen of completed fertility is expected to follow Pois-son and Negative binomial distributions (Cavalli-Sforza
10
1971 Vasulu 1980) In patrilineal societies femalesgenerally show a greater surname diversity than malesand as a result the surname dynamics differ with respectto mitochondria and Y-chromosome
In the Indian context surname analysis has been carriedout in a few populations for investigating the effect onpopulation structure of inbreeding (Bhalla 1976 Kashyap1980 Vasulu 1980) genetic kinship (Kashyap 1980Vasulu 1980) preferential mating (Vasulu 1989 Reddy1995 Maji et al 2007) and social network analysisamong castes and tribes (Rao 1995) Surname neutralityhas also been validated in a caste community of Indiansresiding in Texas (Chakraborty 1990)
The above few studies suggest that isonymy can be ofmuch value in inferring population structure in somecastes and tribes (Kashyap 1980 Maji et al 2007) Thesurname structure in India varies among different regionscastes tribes linguistic and religious groups and providesan opportunity to study microevolution and populationstructure where the assumptions of surname or isonymyanalysis are valid Its applicability in diverse populationsespecially among isolated populations is expected to yieldfurther insights of its validity and inferring populationstructure and microevolutionary history
The present study illustrates the importance of surnamein inferring the genetic structure among a small sub-tribeAdi Panggi an isolated Tibeto-Burman- speaking com-munity in Arunachal Pradesh state in the northeasternregion of India The tribe lives on shifting-hunting-and-gathering subsistence and is a patrilocal and patrilinealsociety where the marriage pattern involves movement ofwives from other villages to the husbandrsquos village whichallows more surname diversity through females As aconsequence of differential surname structure betweenhusbands and wives the surname distribution showsdeviation from the expected neutral allele model and italso exhibits a non-linear trend of progeny size distribu-tion with respect to surname occurrence in the husbandsample
The results of the present study illustrate that surnamedistribution could be of much value in deciphering thepopulation structure and microevolution of a population(Crow 1965 Lasker 1978 Yasuda et al 1974) Thenon-linear trend observed in this study appears to beunique in the Indian context especially for the smallisolated tribes
The results of the study will help in further confirming thevalidity of the findings in several other tribes in India Inthe absence of any biological or genetic data for a tribethe surnames provide insights into the population struc-ture and possible genetic implications which can help inthe design of future studies As a prelude to the moleculargenetic investigations about the past genetic history asurname analysis can provide considerable insight into
the expected genetic structure of the population It alsohelps us to frame specific hypothesis related to male andfemale migration history which we will be testing withthe molecular genetic studies The two approachestogether can help to unravel the differential paternaland maternal past genetic history of the tribe
We thank the Government of Arunachal Pradesh andall the participants for their active cooperation in ourstudy Our sincere thanks go to Mr Tozing Tagi andMr Oyam Panyang (Katan and Geku circles) for theirassistance in data collection We acknowledge thesupport and intellectual inputs about the Adi tribesfrom Mr R N Koley (ARO Directorate of Research)and Mr Kaling Borang of Pasighat We thank Ms SKrithika Senior Research Fellow Biological Anthro-pology Unit for suggestions and help with the analysisand draft finalization and the Director Indian Statisti-cal Institute Kolkata for the financial assistance for thestudy among the tribes of Arunachal Pradesh
Barrai I Barbujani G Beretta M Maestri I Russo A Formi-ca G Pinto-Cisternas (1987) Surnames in Ferrara distribu-tion isonymy and levels of inbreeding14415-423
Barrai I Formica G Barale R Beretta M (1989) Isonymy andmigration distance 53249-262
Barrai I Formica G Barale R Scapoli C Canella R BerettaM (1990) Isonymy in emigrants from Ferrara in 1981-1988Ann Hum Biol 177-18
Barrai I Scapoli C Formica G et al (1991) Isonymy inrecords of births and deaths in Ferrara8395-404
Barrai I Formica G Scapoli C et al (1992) Microevolutionin Ferrara Isonymy 1890-1990 19371-385
Barrai I Scapoli C Bereta M et al (1996) Isonymy and thegenetic structure of Switzerland I The distributions of sur-names 23431-455
Barrai I Rodriguez-Larralde A Mamolini E et al (2000)Elements of surname structure of Austria27607-622
Barrai I Rodriguez-Larralde A Manni F et al (2002) Isony-my and isolation by distance in the Netherlands 74263-283
Barrai I Rodriguez-Larralde A Manni F Ruggiero V TartariD Scapoli C (2003) Isolation by language and distance inBeligium 681-16
Bhalla V Bhatia K (1976) Isonymy in a Bhatia leut 39479-500
Bhattacharjee PN (1954) Blood group Investigations in theAbor Tribe 351-54
11
Blackburn S (2004) Memories of migration Notes on legendsand beads in Arunachal Pradesh India
252615ndash60
Boldsen J (1992) Geographical distribution of some Danishsurnames reflection of social and natural selection 24505-513
Cavelli Sfroza LL Bodmer WF (1971) W H Freeman and Co San Francisco
Chakraborty R Weiss KM Rossmann DL Norton SL (1981)Distribution of last names a stochastic model for lilkelihooddetermination in record linking In Morrill W Dyke B (ed)
Academic Press New York
Chakraborty R Schwartz RJ (1990) Selective neutrality ofsurname distribution in an immigrant Indian community ofHouston Texas 21-13
Colantonio SE Lasker GW Kaplan BA Fuster V (2003) Useof surname models in human population biology A review ofrecent developments 75785-807
Crow JF Manage AP (1965) Measurements of Inbreeding ofthe frequencies of marriages between persons of the samesurnames 12199-203
Das TC (1953) Social Organization of the tribal peoples XIV245-262
Dyke B James AV Morrill WT (1983) Estimation of randomisonymy 10295-298
Fox WR Lasker GW (1983) The distribution of surnamefrequencies 5181-87
Karlin S MacGregor J (1967) The number of mutant formsmaintained in a population
4415-438
Kashyap LK (1980) Trends of Isonymy and inbreeding amongthe Ahmadiyyas of Kashmir 1219-25
Kashyap LK Tiwari SC (1980) Kinetics of genetic kinship asinferred by isonymy among the Ahmadiyyas of Kashmir valley
52311-324
Kimura M (1983) Cambridge University Press Cambridge
Koley RN (2005) Socio-cultural aspects of Adi Panggi Direc-torate of Research Government of Arunachal Pradesh Itanagar(in press)
Krithika S Maji S Vasulu TS (2006) Genetic heterogeneityamong three Adi tribes of Arunachal Pradesh India78221-227
Lasker GW (1978) Increments through migration to the coeffi-cient of relationship between communities by isonymy
50235-240
Lego N (2005) History of the Adis of Arunachal Pradesh MrsPonung Lego Itanagar
Lego N (2006) Modern History of Arunachal Pradesh (1825 ndash2006) Mrs Ponung Lego Itanagar
Maji S Krithika S Vasulu TS (2007) Genetic kinship among anisolated Adi tribe of Arunachal Pradesh isonymy in Adi Panggi
79321-337
Manrubia SC Zanette DH (2002) At the boundary betweenbiological and cultural evolution The origin of surname distri-butions 216461-477
Neel JV Chagnon NA (1968) The demography of two tribes ofprimitive relatively unacculturated American Indians
(USA) 59680-689
Rao AR (1995) Reciprocity in marital and social networksIllustration with Indian data 67887-904
Reddy BM Malhotra KC (1995) Relevance of surname inpopulation structure A study among the Vadde a fishingcommunity of Kolleru lake in Andhra Pradesh India
6161-168
Relethford JH (1988) Estimation of Kinship and Genetic dis-tance from Surnames 60475-492
Rodriguez LA Barrai I Alfonzo JC (1993) Isonymy structureof four Venezuelan states 20131-145
Rodriguez-Larralde A Formica G Scapoli C (1993) Microevo-lution in Perugia Isonymy 1890-1990 20261-274
Rodriguez-Larralde A Gonzales-Martin A Scapoli C Barrai I(2003) The names of Spain A study of the isonymy structureof Spain 121280-292
Roy S (1960) Aspects of Padam Minyong Culture Directorateof Research Govt of Arunachal Pradesh Itanagar
Roy S (1966) Anthropometry of the Adis of the Siang districtNorth-East Frontier Agency Shillong
Salzano FM Neel JV Maybury Lewis D (1967) Further studieson Xavante Indians I Demographic data on two additionalvillages and genetic structure of the tribe19463-489
Sarma J (1960) Puberty Marriage and Childbirth among thePangi and the Minyong Abor women 5597-113
Scapoli C Rodriguez-Larralde A Beretta M (1997) Correlationbetween isonymy parameters 1217-37
Singh IP (1994) Anthropological Survey of India Oxford
University Press New Delhi
Vasulu TS (1980) Anthrop Soc
of India Mysore
Vasulu TS (1989) Genetic structure of a tribal population 1Breeding isolation among the Yanadis 4255-268
Yasuda N Cavalli-Sfroza LL Skolnir M Moroni A (1974) Theevolution of surnames an analysis of their distribution andextinction 5123-142
Zei G Guglielmino CR Siri E Moroni A Cavalli-Sforza LL(1983a) Surnames as neutral alleles Observation in Sardinia
55357-365
Zei G Matessi RG Siri E et al (1983b) Surnames in SardiniaI Fit of frequency distributions for neutral articles and geneticpopulation structure 47329-352
Maji Selective Neutrality of Surname Distribution in Adi Panggi 3
for the total sample was considered separately in thepresent study This distribution is expected to show alinear trend under the assumptions of the neutral allelemodel (Kimura 1983) We also have considered thePareto distribution (Fox 1983) where the relationshipbetween the logarithms of and is expected to be
Sur-Name
ID
Sur-Name
Hus-bands
WivesPanggiOnly
WivesPanggi
and Non-Panggi
N=154 N=130 N=147
1 Paron 1558 1692 1497
2 Panyang 2208 2923 2585
3 Taku 584 1000 884
4 Panggeng 844 385 340
5 Tagi 779 385 340
6 Mongku 1039 308 272
7 Tosang 260 231 204
8 Jopir 065 000 000
9 Taying 195 231 204
10 Tatung 130 154 136
11 Ejing 844 385 340
12 Paloh 390 308 272
13 Padun 130 538 476
14 Tateh 390 308 272
15 Tayom 130 154 136
16 Aje 195 000 000
17 Tarang 195 231 204
18 Taruk 065 385 340
19 Teksin - 077 068
20 Kirom - 077 068
21 Gete - 154 136
22 Tangu - 077 068
23 Non-Panggi
(15)
- - 1156
2ipI or 2
iqI and ii qpI
(SS) and Sibum (SB) are remotely located around 15-30km away from Geku Town (GT) whereas the remainingthree villages Ramku (RK) Kumku (KK) and Peram(PR) are located close to GT
Random isonymy (I) was calculated from the frequencydistribution of the abundance of each surname separatelyin males ( p i ) and females ( q i ) and for both in each( i t h ) village If p i and q i are the frequencies of aparticular surname in males (husbands) and females(wives) respectively then the random isonymy estimateamong males females and both males and females is
Since the random isonymy is biased for the samplesrsquo sizevariations an unbiased isonymy and its variance wasestimated (Dyke et al 1983 Relethford 1988 Barrai etal 1989 Barrai et al 1991) If there are Ni males and Nj
females in the samples the unbiased random isonomy I i jis defined as
I i j = ( n i s bull njs) Ni bull Nj
where nis and njs are the numbers of individuals withsurname in populations i and j respectively and Ni andNj are the total number of surnames in populations i andj and summation is over all s surnames
Since the surnames mimic a genetic trait on the Y-chro-mosome and the surname distribution in a large popula-tion also correlates with genetic diversity (Barrai et al1996) it is expected to conform under certain assump-tions to the selection neutrality of an infinite allele modeldescribed by Karlin and MacGregor (1967) which can beaccounted for by the logarithmic distribution first givenby Fisher That is if we let S represent the number oftimes that a surname occurs in a population and let Krepresent the frequency of S (ie K is the number ofsurnames occurring this same number of S times) thenwe may characterize the distribution by graphing log Kversus log S Karlin and MacGregorrsquos
proposed by Zei et al (1983a 1983b) and Barrai et al(1992) respectively where
I i j ( N i )
and I is random isonymy N i is the total number ofindividuals and
Further the l o g 2 S - l o g 2 K distribution of surnameoccurrence (S) and its frequency K for husband wife and
4
linear where is the number of individuals and is thenumber of surnames but the results are presented for thelog-log distribution only
Occurrence of surname with respect to variation in prog-eny size distribution (separately for male female andtotal children) among the post-reproductive women andtheir husbands were analyzed to investigate the pattern ofsurname distribution in a system of patrilocal marriagewhere males (kin group especially brothers) reside in thesame village and females move out of the village aftertheir marriage Similar analysis was carried out for thecase of reproductive wives and their husbandsrsquo sample
The occurrence of different surnames among husbandsand wives of the Adi Panggi tribe are shown in There are 18 different surnames among 154 husbandsand 22 surnames among 130 wives among the Panggi
GT PR RK KM SS SB
Husband01144 01606 00966 00560 01083 02750
42612 44005 12832 10085 49796 106207
Wife(Panggi)
00967 03444 06965 01467 01334 03131
25678 94574 252379 25790 53877 117380
Wife(Panggi andNon-Panggi)
01679 04706 07450 01827 01607 03444
56482 213309 408934 35767 70828 147090
tribe distributed over six villages There are 15 sur-names of non-Panggi origin that have filtered into thepopulation through marriages with non-Panggi wivesHusbands represent a smaller number of surnameswhereas wives have more diversity of surnames Onaverage a single surname is shared by about 85 ofhusbands while in the case of wives it is about 6 Thethree most common surnames among husbands includeParon Panyang and Mongku which represent about47 of the husbands In case of wives the surnamesPanyang Paron and Taku are the three most commonand occur among 57 of Panggi wives These sevensurnames (which have a frequency above 5) represent785 of the husbands and 615 of thei wives Themost frequent surnames occur among both husbandsand wives with similar percentages and the seven mostfrequent (above 5 occurrence) constitute 707 ofthe sample The sample contains four singly occurringPanggi surnames representing 13 percent of the totalindividuals Considering both Panggi and non-Panggisurnames there are 18 singletons that constitute 6 of
Sample Size F F
Husband 154 00344 54830 00513 83278
Wife(Panggi) 130 00555 76390 00483 66020
Wife(Panggi andNon-Panggi)
147 09010 147215 00559 87113
F and F are distributions based upon random isonymy
Maji Selective Neutrality of Surname Distribution in Adi Panggi 5
Log-log distribution of surnames in Adi Panggi husbands and wives
A) Distribution of surnames among Adi Panggi Husbands
the sample The unbiased random isonymy I i j for theAdi Panggi samples is 01025
shows the estimates the two parametersof Karlin-MacGregor and Fishersrsquo distributions that de-scribe the differential migration rates and surname diver-sity among six villages of Adi Panggi Both the husbandsand wives show wide variation in the rate of migrationand surname diversity between the villages The twovillages KM and RK show the least migration rates andsurname diversity whereas SB shows the highest values of
in case of the husband samples
Panggi wives show higher surname diversity and migra-tion rates than husbands except in case of GT Panggiand non-Panggi wives show higher values than husbands( ) The estimates of F F based on randomisonymy show lesser values than from the Karlin-MacGregor and Fishersrsquo distributions The results showa similar pattern among husbands (n F = 00513 and a F
= 83278) and Panggi plus non-Panggi wives (n F = 00559and a F = 87113)
The logarithmic distribution of surname occurrenceamong husbands and wives and in the total sample for thePanggi is shown in
Both the figures show a non-linear trend suggesting devi-ation from the expected neutral allele model The shapeof the distribution is different in husbands and wives it is
an inverted parabolic curve in males and a slightlyconcave curve in females The best non-linear fit of thecurve with polynomial degree four show a variance ofabout 60 (Rsup2 = 059) in the case of the husbandssample while for wives a polynomial degree threeshows a good fit to the distribution (Rsup2 = 080) whichis better than the linear fit (Rsup2 = 074) In the case ofall Adi Panggi (both husbands and wives) the distribu-tions show a nonlinear trend which is different fromthe trend observed in the case of husband and wifesamples ( ) The best fit to the curve is apolynomial of degree four which accounts for about96 of the variance The non-linear distribution is dueto the higher frequency of occurrence of rare surnames(especially those that occur only once and are contrib-uted by wives) However the truncated log-log distri-bution (with the rare surnames excluded) shows alinear trend
The surname distribution observed among husbandsand wives has also been considered with respect to theirprogeny size to investigate variation of fertility differ-entials with respect to surnames among the post-repro-ductive wives and their husbands separately for malefemale and total progeny size The results of thedistribution show a non-linear trend suggesting a dif-ferential reproductive fitness for different surnamesamong husbands and wives ( ) In the case ofhusbands the rare surnames (occurring once or twice)and common surnames (occurring more than 6-9
Maji Selective Neutrality of Surname Distribution in Adi Panggi 7
Occurrence of surname distribution (s) total number of male ( ) female ( ) andtotal children (times) among Adi Panggi husbands and wives (above 45 Years) and progeny sizedistribution (--)
times) show a trend toward higher progeny size (malesfemales and total children) than those surnames thatoccur less frequently (between 3-5 times) The best fitcurve in case of total children (polynomial degree three)accounts for about 945 percent of the variance (Rsup2 =0945) In case of females the trend shows a negativeassociation between occurrence of surnames and theprogeny size and the best fit curve for the total childrenaccounts (polynomial degree three) about 99 percent ofthe variance (Rsup2 = 0989) Progeny size shows a decreas-
ing trend with the rare surnames that occur once ortwice and a trend of lesser progeny in case of morecommon surnames (except in case the most commonsurnames that occur ten times among wives) Theseresults are a reflection of the population structure of thetribe especially the patrilocal system of marriage whichtends to increase the prevalence of some specific sur-names as a result of preferential marriages among a fewspecific clans or surnames
Husband ( gt 4 5 )
05
101520253035
0 2 4 6 8 10
Occurrence (s)
FrequencyW i f e ( gt 4 5 )
0
10
20
30
40
0 2 4 6 8 10
Occurrence (s)
Frequency
8
imilar distribution of progeny size with respect to sur-name distribution was also considered in case of repro-ductive wives and their husbandsrsquo sample ( )The pattern of distributions is quite varied compared tothe case of post-reproductive samples In the case of thewives the progeny size shows an increasing trend fromthe rare surnames to the most frequent surnamesthough there is some perturbance at the initial case ofrarer ones Among the husband samples a similar initialperturbance of progeny size is noticed with the rarersurnames but the progeny size shows stability with themore commonly occurring surnames of the tribe
Occurrence of surname distribution(s) total number of male ( ) female ( ) and totalchildren (times) among Adi Panggi husbands and wives (below 45 years) and progeny size distribution (--)
Husband (lt45)
0
10
20
30
40
50
60
0 5 10 15 20 25 30
Occurrence (s)
Frequency
W i f e ( lt 4 5 )
0
20
40
60
80
100
0 5 10 15 20 25 30
Occurrence (s)
Frequency
Though selective neutrality of surname distribution hasbeen validated in a number of populations in Europe andLatin American countries a few studies have showndeviation from neutrality implicating the role of migra-tion and operation of selection with respect to certainsurnames The results of the present study indicate thatsuch trends are also observed in small isolated popula-tions Apart from the expected influences of migrationand drift there could be other related factors of popula-tion structure that could cause the deviation from neutral-
Maji Selective Neutrality of Surname Distribution in Adi Panggi 9
ity In small tribal populations the structure of thesurname distribution might vary widely depending on thehistory of settlement prevailing marriage patterns andother socio-cultural and environmental reasons Furtherthe demographic parameters can vary drastically owing totheir subsistence pattern epidemiological factors andsocial organization such as warfare history Also internaltribal factions which are formed based on kinship or clanor lineages involving several closely knit families (eg inSouth Amerindian tribes Neel 1968 Salzano et al 1967)may play a role This can significantly influence thepopulation structure and can lead to perturbations in thesurname distribution in short time periods and in suchcases the neutral theory predictions may not hold Someof the above factors are important in explaining thedeviation from neutrality in surname distribution amongthe Adi Panggi
In general the Adi tribe consists of several sub-tribes whooccupy different terrains in the upper Himalayan moun-tain ranges and some in plains in the northeastern part ofthe country (Roy 1960 Lego 2005) The formation of thesub-tribes followed the fission-fusion model of popula-tion structure and was triggered by intertribal factionscompeting for survival necessities in this hunting-gather-ing population (Neel 1968 Salzano et al 1967) Theformation of Adi Panggi sub-tribes occurred a couple ofgenerations ago and was a result of conflict between amajor subgroup of the tribe and some other close-kinfactions over hunting rights and other resources (Roy1960 Lego 2005 Lego 2006) The sub-tribes now occu-py a recognized territory and constitute a few close affinaland consanguineous lineages and their affiliated clansSince they have a patrilocal society a majority of theirmen stay in the home village whereas females come fromdifferent villages and from other Adi sub-tribes Thisresults in more female marital migration and greatersurname diversity among wives We therefore observe ahigher proportion of homonymy of surname diversityamong males For example the five most frequent sur-names represent about 65 of total husbands while raresurnames (that occur either once or twice) constituteabout 5 per cent The three most common surnamesamong wives also represent a large part of the populationbut in contrast with the males the rare surnames thatoccur either once or twice constitute 18 per cent morethan 35 times the proportion of rare surnames in hus-bands In the total sample of husbands and wives sixsurnames represent 69 percent of the total populationstudied The surname distribution therefore reflectsgreater prevalence of most frequent (occurring in 20 to 70cases) and rare surnames (occurring once or twice) asagainst the moderately frequent surnames (that occur 3 to20 times in the sample)
The estimates of unbiased Isonymy also differ betweenhusband and wives samples The combined samples(husband and wives) show higher values of estimatedIsonymy (I = 01025) suggesting higher levels of homozy-gosity as a result of more frequent marriages contracted
among a few selected surnames generally found in suchisolated tribes This is reflected in the Karlin andMacGregor distribution which shows the non-lineartrend and the trend remains the same even after thelogarithmic transformation This also explains the widedifferences between the male and female surname distri-butions and the deviations observed in the Karlin andMacGregor and logarithmic distributions Therefore ifthe truncated distribution (omitting the rare surnames) isconsidered it shows a linear trend in the combinedsample suggesting that the non-linearity is contributedby the occurrence of rare surnames in the sample Therare surnames are contributed more through femalesmigrating from other places However the discretePareto distribution shows a linear trend for both thesamples The linear trend is clearer in the female samplethan among males but the genetic implications of thePareto distributions are not apparent (Fox 1983)
The results of the distribution of surnames also showwide differences with respect to progeny size distributionin case of post-reproductive (completed fertility) andreproductive wives and their husbands The neutralallele model would suggest that the distribution shouldbe independent of the fertility and morality (Manrubia2002) but the present study indicates otherwise andthere appears to be some differential fertility for somesurnames The Adi Panggi sample shows greater progenysize (male female and total surviving children) for themost frequent and rare surnames and that is reflected inthe near U-shaped distribution especially in case of hus-band sample This suggests reproductive advantageespecially greater success for the most frequent (6-910)and rare surnames (1-2) as against the less frequentsurnames (3-6) Among the Adi Panggi the four mostfrequent surnames among husbands and wives (especiallyPanyang Paron Mongku and Taku) show higher proge-ny size among the husband sample whereas the wivessample shows a near linear trend of decreasing progenysize from the rare surnames to the most frequent ones Apaired t-test between (eight) different types of surnamesand their progeny size shows higher significance values incase of the husbands (above 45 years) sample (p =0000433) than among the wives (above 45 years) (p =0003263) The mothers and their husbands were all inpost-reproductive period and had attained their maxi-mum fertility potential and do not use family planningand therefore the possibility of other intervening factorscan be excluded It is possible that it could be also dueto selective mortality of children especially higher mor-tality in case of less frequent surnames than with moreand rare frequent ones and this needs to be verifiedHowever there are no apparent cultural active selectivemechanisms that can influence the progeny size amongsome families of the tribe A detailed study on themarriage pattern and surname or clan distribution mightreveal more details The progeny distribution amongwomen of completed fertility is expected to follow Pois-son and Negative binomial distributions (Cavalli-Sforza
10
1971 Vasulu 1980) In patrilineal societies femalesgenerally show a greater surname diversity than malesand as a result the surname dynamics differ with respectto mitochondria and Y-chromosome
In the Indian context surname analysis has been carriedout in a few populations for investigating the effect onpopulation structure of inbreeding (Bhalla 1976 Kashyap1980 Vasulu 1980) genetic kinship (Kashyap 1980Vasulu 1980) preferential mating (Vasulu 1989 Reddy1995 Maji et al 2007) and social network analysisamong castes and tribes (Rao 1995) Surname neutralityhas also been validated in a caste community of Indiansresiding in Texas (Chakraborty 1990)
The above few studies suggest that isonymy can be ofmuch value in inferring population structure in somecastes and tribes (Kashyap 1980 Maji et al 2007) Thesurname structure in India varies among different regionscastes tribes linguistic and religious groups and providesan opportunity to study microevolution and populationstructure where the assumptions of surname or isonymyanalysis are valid Its applicability in diverse populationsespecially among isolated populations is expected to yieldfurther insights of its validity and inferring populationstructure and microevolutionary history
The present study illustrates the importance of surnamein inferring the genetic structure among a small sub-tribeAdi Panggi an isolated Tibeto-Burman- speaking com-munity in Arunachal Pradesh state in the northeasternregion of India The tribe lives on shifting-hunting-and-gathering subsistence and is a patrilocal and patrilinealsociety where the marriage pattern involves movement ofwives from other villages to the husbandrsquos village whichallows more surname diversity through females As aconsequence of differential surname structure betweenhusbands and wives the surname distribution showsdeviation from the expected neutral allele model and italso exhibits a non-linear trend of progeny size distribu-tion with respect to surname occurrence in the husbandsample
The results of the present study illustrate that surnamedistribution could be of much value in deciphering thepopulation structure and microevolution of a population(Crow 1965 Lasker 1978 Yasuda et al 1974) Thenon-linear trend observed in this study appears to beunique in the Indian context especially for the smallisolated tribes
The results of the study will help in further confirming thevalidity of the findings in several other tribes in India Inthe absence of any biological or genetic data for a tribethe surnames provide insights into the population struc-ture and possible genetic implications which can help inthe design of future studies As a prelude to the moleculargenetic investigations about the past genetic history asurname analysis can provide considerable insight into
the expected genetic structure of the population It alsohelps us to frame specific hypothesis related to male andfemale migration history which we will be testing withthe molecular genetic studies The two approachestogether can help to unravel the differential paternaland maternal past genetic history of the tribe
We thank the Government of Arunachal Pradesh andall the participants for their active cooperation in ourstudy Our sincere thanks go to Mr Tozing Tagi andMr Oyam Panyang (Katan and Geku circles) for theirassistance in data collection We acknowledge thesupport and intellectual inputs about the Adi tribesfrom Mr R N Koley (ARO Directorate of Research)and Mr Kaling Borang of Pasighat We thank Ms SKrithika Senior Research Fellow Biological Anthro-pology Unit for suggestions and help with the analysisand draft finalization and the Director Indian Statisti-cal Institute Kolkata for the financial assistance for thestudy among the tribes of Arunachal Pradesh
Barrai I Barbujani G Beretta M Maestri I Russo A Formi-ca G Pinto-Cisternas (1987) Surnames in Ferrara distribu-tion isonymy and levels of inbreeding14415-423
Barrai I Formica G Barale R Beretta M (1989) Isonymy andmigration distance 53249-262
Barrai I Formica G Barale R Scapoli C Canella R BerettaM (1990) Isonymy in emigrants from Ferrara in 1981-1988Ann Hum Biol 177-18
Barrai I Scapoli C Formica G et al (1991) Isonymy inrecords of births and deaths in Ferrara8395-404
Barrai I Formica G Scapoli C et al (1992) Microevolutionin Ferrara Isonymy 1890-1990 19371-385
Barrai I Scapoli C Bereta M et al (1996) Isonymy and thegenetic structure of Switzerland I The distributions of sur-names 23431-455
Barrai I Rodriguez-Larralde A Mamolini E et al (2000)Elements of surname structure of Austria27607-622
Barrai I Rodriguez-Larralde A Manni F et al (2002) Isony-my and isolation by distance in the Netherlands 74263-283
Barrai I Rodriguez-Larralde A Manni F Ruggiero V TartariD Scapoli C (2003) Isolation by language and distance inBeligium 681-16
Bhalla V Bhatia K (1976) Isonymy in a Bhatia leut 39479-500
Bhattacharjee PN (1954) Blood group Investigations in theAbor Tribe 351-54
11
Blackburn S (2004) Memories of migration Notes on legendsand beads in Arunachal Pradesh India
252615ndash60
Boldsen J (1992) Geographical distribution of some Danishsurnames reflection of social and natural selection 24505-513
Cavelli Sfroza LL Bodmer WF (1971) W H Freeman and Co San Francisco
Chakraborty R Weiss KM Rossmann DL Norton SL (1981)Distribution of last names a stochastic model for lilkelihooddetermination in record linking In Morrill W Dyke B (ed)
Academic Press New York
Chakraborty R Schwartz RJ (1990) Selective neutrality ofsurname distribution in an immigrant Indian community ofHouston Texas 21-13
Colantonio SE Lasker GW Kaplan BA Fuster V (2003) Useof surname models in human population biology A review ofrecent developments 75785-807
Crow JF Manage AP (1965) Measurements of Inbreeding ofthe frequencies of marriages between persons of the samesurnames 12199-203
Das TC (1953) Social Organization of the tribal peoples XIV245-262
Dyke B James AV Morrill WT (1983) Estimation of randomisonymy 10295-298
Fox WR Lasker GW (1983) The distribution of surnamefrequencies 5181-87
Karlin S MacGregor J (1967) The number of mutant formsmaintained in a population
4415-438
Kashyap LK (1980) Trends of Isonymy and inbreeding amongthe Ahmadiyyas of Kashmir 1219-25
Kashyap LK Tiwari SC (1980) Kinetics of genetic kinship asinferred by isonymy among the Ahmadiyyas of Kashmir valley
52311-324
Kimura M (1983) Cambridge University Press Cambridge
Koley RN (2005) Socio-cultural aspects of Adi Panggi Direc-torate of Research Government of Arunachal Pradesh Itanagar(in press)
Krithika S Maji S Vasulu TS (2006) Genetic heterogeneityamong three Adi tribes of Arunachal Pradesh India78221-227
Lasker GW (1978) Increments through migration to the coeffi-cient of relationship between communities by isonymy
50235-240
Lego N (2005) History of the Adis of Arunachal Pradesh MrsPonung Lego Itanagar
Lego N (2006) Modern History of Arunachal Pradesh (1825 ndash2006) Mrs Ponung Lego Itanagar
Maji S Krithika S Vasulu TS (2007) Genetic kinship among anisolated Adi tribe of Arunachal Pradesh isonymy in Adi Panggi
79321-337
Manrubia SC Zanette DH (2002) At the boundary betweenbiological and cultural evolution The origin of surname distri-butions 216461-477
Neel JV Chagnon NA (1968) The demography of two tribes ofprimitive relatively unacculturated American Indians
(USA) 59680-689
Rao AR (1995) Reciprocity in marital and social networksIllustration with Indian data 67887-904
Reddy BM Malhotra KC (1995) Relevance of surname inpopulation structure A study among the Vadde a fishingcommunity of Kolleru lake in Andhra Pradesh India
6161-168
Relethford JH (1988) Estimation of Kinship and Genetic dis-tance from Surnames 60475-492
Rodriguez LA Barrai I Alfonzo JC (1993) Isonymy structureof four Venezuelan states 20131-145
Rodriguez-Larralde A Formica G Scapoli C (1993) Microevo-lution in Perugia Isonymy 1890-1990 20261-274
Rodriguez-Larralde A Gonzales-Martin A Scapoli C Barrai I(2003) The names of Spain A study of the isonymy structureof Spain 121280-292
Roy S (1960) Aspects of Padam Minyong Culture Directorateof Research Govt of Arunachal Pradesh Itanagar
Roy S (1966) Anthropometry of the Adis of the Siang districtNorth-East Frontier Agency Shillong
Salzano FM Neel JV Maybury Lewis D (1967) Further studieson Xavante Indians I Demographic data on two additionalvillages and genetic structure of the tribe19463-489
Sarma J (1960) Puberty Marriage and Childbirth among thePangi and the Minyong Abor women 5597-113
Scapoli C Rodriguez-Larralde A Beretta M (1997) Correlationbetween isonymy parameters 1217-37
Singh IP (1994) Anthropological Survey of India Oxford
University Press New Delhi
Vasulu TS (1980) Anthrop Soc
of India Mysore
Vasulu TS (1989) Genetic structure of a tribal population 1Breeding isolation among the Yanadis 4255-268
Yasuda N Cavalli-Sfroza LL Skolnir M Moroni A (1974) Theevolution of surnames an analysis of their distribution andextinction 5123-142
Zei G Guglielmino CR Siri E Moroni A Cavalli-Sforza LL(1983a) Surnames as neutral alleles Observation in Sardinia
55357-365
Zei G Matessi RG Siri E et al (1983b) Surnames in SardiniaI Fit of frequency distributions for neutral articles and geneticpopulation structure 47329-352
4
linear where is the number of individuals and is thenumber of surnames but the results are presented for thelog-log distribution only
Occurrence of surname with respect to variation in prog-eny size distribution (separately for male female andtotal children) among the post-reproductive women andtheir husbands were analyzed to investigate the pattern ofsurname distribution in a system of patrilocal marriagewhere males (kin group especially brothers) reside in thesame village and females move out of the village aftertheir marriage Similar analysis was carried out for thecase of reproductive wives and their husbandsrsquo sample
The occurrence of different surnames among husbandsand wives of the Adi Panggi tribe are shown in There are 18 different surnames among 154 husbandsand 22 surnames among 130 wives among the Panggi
GT PR RK KM SS SB
Husband01144 01606 00966 00560 01083 02750
42612 44005 12832 10085 49796 106207
Wife(Panggi)
00967 03444 06965 01467 01334 03131
25678 94574 252379 25790 53877 117380
Wife(Panggi andNon-Panggi)
01679 04706 07450 01827 01607 03444
56482 213309 408934 35767 70828 147090
tribe distributed over six villages There are 15 sur-names of non-Panggi origin that have filtered into thepopulation through marriages with non-Panggi wivesHusbands represent a smaller number of surnameswhereas wives have more diversity of surnames Onaverage a single surname is shared by about 85 ofhusbands while in the case of wives it is about 6 Thethree most common surnames among husbands includeParon Panyang and Mongku which represent about47 of the husbands In case of wives the surnamesPanyang Paron and Taku are the three most commonand occur among 57 of Panggi wives These sevensurnames (which have a frequency above 5) represent785 of the husbands and 615 of thei wives Themost frequent surnames occur among both husbandsand wives with similar percentages and the seven mostfrequent (above 5 occurrence) constitute 707 ofthe sample The sample contains four singly occurringPanggi surnames representing 13 percent of the totalindividuals Considering both Panggi and non-Panggisurnames there are 18 singletons that constitute 6 of
Sample Size F F
Husband 154 00344 54830 00513 83278
Wife(Panggi) 130 00555 76390 00483 66020
Wife(Panggi andNon-Panggi)
147 09010 147215 00559 87113
F and F are distributions based upon random isonymy
Maji Selective Neutrality of Surname Distribution in Adi Panggi 5
Log-log distribution of surnames in Adi Panggi husbands and wives
A) Distribution of surnames among Adi Panggi Husbands
the sample The unbiased random isonymy I i j for theAdi Panggi samples is 01025
shows the estimates the two parametersof Karlin-MacGregor and Fishersrsquo distributions that de-scribe the differential migration rates and surname diver-sity among six villages of Adi Panggi Both the husbandsand wives show wide variation in the rate of migrationand surname diversity between the villages The twovillages KM and RK show the least migration rates andsurname diversity whereas SB shows the highest values of
in case of the husband samples
Panggi wives show higher surname diversity and migra-tion rates than husbands except in case of GT Panggiand non-Panggi wives show higher values than husbands( ) The estimates of F F based on randomisonymy show lesser values than from the Karlin-MacGregor and Fishersrsquo distributions The results showa similar pattern among husbands (n F = 00513 and a F
= 83278) and Panggi plus non-Panggi wives (n F = 00559and a F = 87113)
The logarithmic distribution of surname occurrenceamong husbands and wives and in the total sample for thePanggi is shown in
Both the figures show a non-linear trend suggesting devi-ation from the expected neutral allele model The shapeof the distribution is different in husbands and wives it is
an inverted parabolic curve in males and a slightlyconcave curve in females The best non-linear fit of thecurve with polynomial degree four show a variance ofabout 60 (Rsup2 = 059) in the case of the husbandssample while for wives a polynomial degree threeshows a good fit to the distribution (Rsup2 = 080) whichis better than the linear fit (Rsup2 = 074) In the case ofall Adi Panggi (both husbands and wives) the distribu-tions show a nonlinear trend which is different fromthe trend observed in the case of husband and wifesamples ( ) The best fit to the curve is apolynomial of degree four which accounts for about96 of the variance The non-linear distribution is dueto the higher frequency of occurrence of rare surnames(especially those that occur only once and are contrib-uted by wives) However the truncated log-log distri-bution (with the rare surnames excluded) shows alinear trend
The surname distribution observed among husbandsand wives has also been considered with respect to theirprogeny size to investigate variation of fertility differ-entials with respect to surnames among the post-repro-ductive wives and their husbands separately for malefemale and total progeny size The results of thedistribution show a non-linear trend suggesting a dif-ferential reproductive fitness for different surnamesamong husbands and wives ( ) In the case ofhusbands the rare surnames (occurring once or twice)and common surnames (occurring more than 6-9
Maji Selective Neutrality of Surname Distribution in Adi Panggi 7
Occurrence of surname distribution (s) total number of male ( ) female ( ) andtotal children (times) among Adi Panggi husbands and wives (above 45 Years) and progeny sizedistribution (--)
times) show a trend toward higher progeny size (malesfemales and total children) than those surnames thatoccur less frequently (between 3-5 times) The best fitcurve in case of total children (polynomial degree three)accounts for about 945 percent of the variance (Rsup2 =0945) In case of females the trend shows a negativeassociation between occurrence of surnames and theprogeny size and the best fit curve for the total childrenaccounts (polynomial degree three) about 99 percent ofthe variance (Rsup2 = 0989) Progeny size shows a decreas-
ing trend with the rare surnames that occur once ortwice and a trend of lesser progeny in case of morecommon surnames (except in case the most commonsurnames that occur ten times among wives) Theseresults are a reflection of the population structure of thetribe especially the patrilocal system of marriage whichtends to increase the prevalence of some specific sur-names as a result of preferential marriages among a fewspecific clans or surnames
Husband ( gt 4 5 )
05
101520253035
0 2 4 6 8 10
Occurrence (s)
FrequencyW i f e ( gt 4 5 )
0
10
20
30
40
0 2 4 6 8 10
Occurrence (s)
Frequency
8
imilar distribution of progeny size with respect to sur-name distribution was also considered in case of repro-ductive wives and their husbandsrsquo sample ( )The pattern of distributions is quite varied compared tothe case of post-reproductive samples In the case of thewives the progeny size shows an increasing trend fromthe rare surnames to the most frequent surnamesthough there is some perturbance at the initial case ofrarer ones Among the husband samples a similar initialperturbance of progeny size is noticed with the rarersurnames but the progeny size shows stability with themore commonly occurring surnames of the tribe
Occurrence of surname distribution(s) total number of male ( ) female ( ) and totalchildren (times) among Adi Panggi husbands and wives (below 45 years) and progeny size distribution (--)
Husband (lt45)
0
10
20
30
40
50
60
0 5 10 15 20 25 30
Occurrence (s)
Frequency
W i f e ( lt 4 5 )
0
20
40
60
80
100
0 5 10 15 20 25 30
Occurrence (s)
Frequency
Though selective neutrality of surname distribution hasbeen validated in a number of populations in Europe andLatin American countries a few studies have showndeviation from neutrality implicating the role of migra-tion and operation of selection with respect to certainsurnames The results of the present study indicate thatsuch trends are also observed in small isolated popula-tions Apart from the expected influences of migrationand drift there could be other related factors of popula-tion structure that could cause the deviation from neutral-
Maji Selective Neutrality of Surname Distribution in Adi Panggi 9
ity In small tribal populations the structure of thesurname distribution might vary widely depending on thehistory of settlement prevailing marriage patterns andother socio-cultural and environmental reasons Furtherthe demographic parameters can vary drastically owing totheir subsistence pattern epidemiological factors andsocial organization such as warfare history Also internaltribal factions which are formed based on kinship or clanor lineages involving several closely knit families (eg inSouth Amerindian tribes Neel 1968 Salzano et al 1967)may play a role This can significantly influence thepopulation structure and can lead to perturbations in thesurname distribution in short time periods and in suchcases the neutral theory predictions may not hold Someof the above factors are important in explaining thedeviation from neutrality in surname distribution amongthe Adi Panggi
In general the Adi tribe consists of several sub-tribes whooccupy different terrains in the upper Himalayan moun-tain ranges and some in plains in the northeastern part ofthe country (Roy 1960 Lego 2005) The formation of thesub-tribes followed the fission-fusion model of popula-tion structure and was triggered by intertribal factionscompeting for survival necessities in this hunting-gather-ing population (Neel 1968 Salzano et al 1967) Theformation of Adi Panggi sub-tribes occurred a couple ofgenerations ago and was a result of conflict between amajor subgroup of the tribe and some other close-kinfactions over hunting rights and other resources (Roy1960 Lego 2005 Lego 2006) The sub-tribes now occu-py a recognized territory and constitute a few close affinaland consanguineous lineages and their affiliated clansSince they have a patrilocal society a majority of theirmen stay in the home village whereas females come fromdifferent villages and from other Adi sub-tribes Thisresults in more female marital migration and greatersurname diversity among wives We therefore observe ahigher proportion of homonymy of surname diversityamong males For example the five most frequent sur-names represent about 65 of total husbands while raresurnames (that occur either once or twice) constituteabout 5 per cent The three most common surnamesamong wives also represent a large part of the populationbut in contrast with the males the rare surnames thatoccur either once or twice constitute 18 per cent morethan 35 times the proportion of rare surnames in hus-bands In the total sample of husbands and wives sixsurnames represent 69 percent of the total populationstudied The surname distribution therefore reflectsgreater prevalence of most frequent (occurring in 20 to 70cases) and rare surnames (occurring once or twice) asagainst the moderately frequent surnames (that occur 3 to20 times in the sample)
The estimates of unbiased Isonymy also differ betweenhusband and wives samples The combined samples(husband and wives) show higher values of estimatedIsonymy (I = 01025) suggesting higher levels of homozy-gosity as a result of more frequent marriages contracted
among a few selected surnames generally found in suchisolated tribes This is reflected in the Karlin andMacGregor distribution which shows the non-lineartrend and the trend remains the same even after thelogarithmic transformation This also explains the widedifferences between the male and female surname distri-butions and the deviations observed in the Karlin andMacGregor and logarithmic distributions Therefore ifthe truncated distribution (omitting the rare surnames) isconsidered it shows a linear trend in the combinedsample suggesting that the non-linearity is contributedby the occurrence of rare surnames in the sample Therare surnames are contributed more through femalesmigrating from other places However the discretePareto distribution shows a linear trend for both thesamples The linear trend is clearer in the female samplethan among males but the genetic implications of thePareto distributions are not apparent (Fox 1983)
The results of the distribution of surnames also showwide differences with respect to progeny size distributionin case of post-reproductive (completed fertility) andreproductive wives and their husbands The neutralallele model would suggest that the distribution shouldbe independent of the fertility and morality (Manrubia2002) but the present study indicates otherwise andthere appears to be some differential fertility for somesurnames The Adi Panggi sample shows greater progenysize (male female and total surviving children) for themost frequent and rare surnames and that is reflected inthe near U-shaped distribution especially in case of hus-band sample This suggests reproductive advantageespecially greater success for the most frequent (6-910)and rare surnames (1-2) as against the less frequentsurnames (3-6) Among the Adi Panggi the four mostfrequent surnames among husbands and wives (especiallyPanyang Paron Mongku and Taku) show higher proge-ny size among the husband sample whereas the wivessample shows a near linear trend of decreasing progenysize from the rare surnames to the most frequent ones Apaired t-test between (eight) different types of surnamesand their progeny size shows higher significance values incase of the husbands (above 45 years) sample (p =0000433) than among the wives (above 45 years) (p =0003263) The mothers and their husbands were all inpost-reproductive period and had attained their maxi-mum fertility potential and do not use family planningand therefore the possibility of other intervening factorscan be excluded It is possible that it could be also dueto selective mortality of children especially higher mor-tality in case of less frequent surnames than with moreand rare frequent ones and this needs to be verifiedHowever there are no apparent cultural active selectivemechanisms that can influence the progeny size amongsome families of the tribe A detailed study on themarriage pattern and surname or clan distribution mightreveal more details The progeny distribution amongwomen of completed fertility is expected to follow Pois-son and Negative binomial distributions (Cavalli-Sforza
10
1971 Vasulu 1980) In patrilineal societies femalesgenerally show a greater surname diversity than malesand as a result the surname dynamics differ with respectto mitochondria and Y-chromosome
In the Indian context surname analysis has been carriedout in a few populations for investigating the effect onpopulation structure of inbreeding (Bhalla 1976 Kashyap1980 Vasulu 1980) genetic kinship (Kashyap 1980Vasulu 1980) preferential mating (Vasulu 1989 Reddy1995 Maji et al 2007) and social network analysisamong castes and tribes (Rao 1995) Surname neutralityhas also been validated in a caste community of Indiansresiding in Texas (Chakraborty 1990)
The above few studies suggest that isonymy can be ofmuch value in inferring population structure in somecastes and tribes (Kashyap 1980 Maji et al 2007) Thesurname structure in India varies among different regionscastes tribes linguistic and religious groups and providesan opportunity to study microevolution and populationstructure where the assumptions of surname or isonymyanalysis are valid Its applicability in diverse populationsespecially among isolated populations is expected to yieldfurther insights of its validity and inferring populationstructure and microevolutionary history
The present study illustrates the importance of surnamein inferring the genetic structure among a small sub-tribeAdi Panggi an isolated Tibeto-Burman- speaking com-munity in Arunachal Pradesh state in the northeasternregion of India The tribe lives on shifting-hunting-and-gathering subsistence and is a patrilocal and patrilinealsociety where the marriage pattern involves movement ofwives from other villages to the husbandrsquos village whichallows more surname diversity through females As aconsequence of differential surname structure betweenhusbands and wives the surname distribution showsdeviation from the expected neutral allele model and italso exhibits a non-linear trend of progeny size distribu-tion with respect to surname occurrence in the husbandsample
The results of the present study illustrate that surnamedistribution could be of much value in deciphering thepopulation structure and microevolution of a population(Crow 1965 Lasker 1978 Yasuda et al 1974) Thenon-linear trend observed in this study appears to beunique in the Indian context especially for the smallisolated tribes
The results of the study will help in further confirming thevalidity of the findings in several other tribes in India Inthe absence of any biological or genetic data for a tribethe surnames provide insights into the population struc-ture and possible genetic implications which can help inthe design of future studies As a prelude to the moleculargenetic investigations about the past genetic history asurname analysis can provide considerable insight into
the expected genetic structure of the population It alsohelps us to frame specific hypothesis related to male andfemale migration history which we will be testing withthe molecular genetic studies The two approachestogether can help to unravel the differential paternaland maternal past genetic history of the tribe
We thank the Government of Arunachal Pradesh andall the participants for their active cooperation in ourstudy Our sincere thanks go to Mr Tozing Tagi andMr Oyam Panyang (Katan and Geku circles) for theirassistance in data collection We acknowledge thesupport and intellectual inputs about the Adi tribesfrom Mr R N Koley (ARO Directorate of Research)and Mr Kaling Borang of Pasighat We thank Ms SKrithika Senior Research Fellow Biological Anthro-pology Unit for suggestions and help with the analysisand draft finalization and the Director Indian Statisti-cal Institute Kolkata for the financial assistance for thestudy among the tribes of Arunachal Pradesh
Barrai I Barbujani G Beretta M Maestri I Russo A Formi-ca G Pinto-Cisternas (1987) Surnames in Ferrara distribu-tion isonymy and levels of inbreeding14415-423
Barrai I Formica G Barale R Beretta M (1989) Isonymy andmigration distance 53249-262
Barrai I Formica G Barale R Scapoli C Canella R BerettaM (1990) Isonymy in emigrants from Ferrara in 1981-1988Ann Hum Biol 177-18
Barrai I Scapoli C Formica G et al (1991) Isonymy inrecords of births and deaths in Ferrara8395-404
Barrai I Formica G Scapoli C et al (1992) Microevolutionin Ferrara Isonymy 1890-1990 19371-385
Barrai I Scapoli C Bereta M et al (1996) Isonymy and thegenetic structure of Switzerland I The distributions of sur-names 23431-455
Barrai I Rodriguez-Larralde A Mamolini E et al (2000)Elements of surname structure of Austria27607-622
Barrai I Rodriguez-Larralde A Manni F et al (2002) Isony-my and isolation by distance in the Netherlands 74263-283
Barrai I Rodriguez-Larralde A Manni F Ruggiero V TartariD Scapoli C (2003) Isolation by language and distance inBeligium 681-16
Bhalla V Bhatia K (1976) Isonymy in a Bhatia leut 39479-500
Bhattacharjee PN (1954) Blood group Investigations in theAbor Tribe 351-54
11
Blackburn S (2004) Memories of migration Notes on legendsand beads in Arunachal Pradesh India
252615ndash60
Boldsen J (1992) Geographical distribution of some Danishsurnames reflection of social and natural selection 24505-513
Cavelli Sfroza LL Bodmer WF (1971) W H Freeman and Co San Francisco
Chakraborty R Weiss KM Rossmann DL Norton SL (1981)Distribution of last names a stochastic model for lilkelihooddetermination in record linking In Morrill W Dyke B (ed)
Academic Press New York
Chakraborty R Schwartz RJ (1990) Selective neutrality ofsurname distribution in an immigrant Indian community ofHouston Texas 21-13
Colantonio SE Lasker GW Kaplan BA Fuster V (2003) Useof surname models in human population biology A review ofrecent developments 75785-807
Crow JF Manage AP (1965) Measurements of Inbreeding ofthe frequencies of marriages between persons of the samesurnames 12199-203
Das TC (1953) Social Organization of the tribal peoples XIV245-262
Dyke B James AV Morrill WT (1983) Estimation of randomisonymy 10295-298
Fox WR Lasker GW (1983) The distribution of surnamefrequencies 5181-87
Karlin S MacGregor J (1967) The number of mutant formsmaintained in a population
4415-438
Kashyap LK (1980) Trends of Isonymy and inbreeding amongthe Ahmadiyyas of Kashmir 1219-25
Kashyap LK Tiwari SC (1980) Kinetics of genetic kinship asinferred by isonymy among the Ahmadiyyas of Kashmir valley
52311-324
Kimura M (1983) Cambridge University Press Cambridge
Koley RN (2005) Socio-cultural aspects of Adi Panggi Direc-torate of Research Government of Arunachal Pradesh Itanagar(in press)
Krithika S Maji S Vasulu TS (2006) Genetic heterogeneityamong three Adi tribes of Arunachal Pradesh India78221-227
Lasker GW (1978) Increments through migration to the coeffi-cient of relationship between communities by isonymy
50235-240
Lego N (2005) History of the Adis of Arunachal Pradesh MrsPonung Lego Itanagar
Lego N (2006) Modern History of Arunachal Pradesh (1825 ndash2006) Mrs Ponung Lego Itanagar
Maji S Krithika S Vasulu TS (2007) Genetic kinship among anisolated Adi tribe of Arunachal Pradesh isonymy in Adi Panggi
79321-337
Manrubia SC Zanette DH (2002) At the boundary betweenbiological and cultural evolution The origin of surname distri-butions 216461-477
Neel JV Chagnon NA (1968) The demography of two tribes ofprimitive relatively unacculturated American Indians
(USA) 59680-689
Rao AR (1995) Reciprocity in marital and social networksIllustration with Indian data 67887-904
Reddy BM Malhotra KC (1995) Relevance of surname inpopulation structure A study among the Vadde a fishingcommunity of Kolleru lake in Andhra Pradesh India
6161-168
Relethford JH (1988) Estimation of Kinship and Genetic dis-tance from Surnames 60475-492
Rodriguez LA Barrai I Alfonzo JC (1993) Isonymy structureof four Venezuelan states 20131-145
Rodriguez-Larralde A Formica G Scapoli C (1993) Microevo-lution in Perugia Isonymy 1890-1990 20261-274
Rodriguez-Larralde A Gonzales-Martin A Scapoli C Barrai I(2003) The names of Spain A study of the isonymy structureof Spain 121280-292
Roy S (1960) Aspects of Padam Minyong Culture Directorateof Research Govt of Arunachal Pradesh Itanagar
Roy S (1966) Anthropometry of the Adis of the Siang districtNorth-East Frontier Agency Shillong
Salzano FM Neel JV Maybury Lewis D (1967) Further studieson Xavante Indians I Demographic data on two additionalvillages and genetic structure of the tribe19463-489
Sarma J (1960) Puberty Marriage and Childbirth among thePangi and the Minyong Abor women 5597-113
Scapoli C Rodriguez-Larralde A Beretta M (1997) Correlationbetween isonymy parameters 1217-37
Singh IP (1994) Anthropological Survey of India Oxford
University Press New Delhi
Vasulu TS (1980) Anthrop Soc
of India Mysore
Vasulu TS (1989) Genetic structure of a tribal population 1Breeding isolation among the Yanadis 4255-268
Yasuda N Cavalli-Sfroza LL Skolnir M Moroni A (1974) Theevolution of surnames an analysis of their distribution andextinction 5123-142
Zei G Guglielmino CR Siri E Moroni A Cavalli-Sforza LL(1983a) Surnames as neutral alleles Observation in Sardinia
55357-365
Zei G Matessi RG Siri E et al (1983b) Surnames in SardiniaI Fit of frequency distributions for neutral articles and geneticpopulation structure 47329-352
Maji Selective Neutrality of Surname Distribution in Adi Panggi 5
Log-log distribution of surnames in Adi Panggi husbands and wives
A) Distribution of surnames among Adi Panggi Husbands
the sample The unbiased random isonymy I i j for theAdi Panggi samples is 01025
shows the estimates the two parametersof Karlin-MacGregor and Fishersrsquo distributions that de-scribe the differential migration rates and surname diver-sity among six villages of Adi Panggi Both the husbandsand wives show wide variation in the rate of migrationand surname diversity between the villages The twovillages KM and RK show the least migration rates andsurname diversity whereas SB shows the highest values of
in case of the husband samples
Panggi wives show higher surname diversity and migra-tion rates than husbands except in case of GT Panggiand non-Panggi wives show higher values than husbands( ) The estimates of F F based on randomisonymy show lesser values than from the Karlin-MacGregor and Fishersrsquo distributions The results showa similar pattern among husbands (n F = 00513 and a F
= 83278) and Panggi plus non-Panggi wives (n F = 00559and a F = 87113)
The logarithmic distribution of surname occurrenceamong husbands and wives and in the total sample for thePanggi is shown in
Both the figures show a non-linear trend suggesting devi-ation from the expected neutral allele model The shapeof the distribution is different in husbands and wives it is
an inverted parabolic curve in males and a slightlyconcave curve in females The best non-linear fit of thecurve with polynomial degree four show a variance ofabout 60 (Rsup2 = 059) in the case of the husbandssample while for wives a polynomial degree threeshows a good fit to the distribution (Rsup2 = 080) whichis better than the linear fit (Rsup2 = 074) In the case ofall Adi Panggi (both husbands and wives) the distribu-tions show a nonlinear trend which is different fromthe trend observed in the case of husband and wifesamples ( ) The best fit to the curve is apolynomial of degree four which accounts for about96 of the variance The non-linear distribution is dueto the higher frequency of occurrence of rare surnames(especially those that occur only once and are contrib-uted by wives) However the truncated log-log distri-bution (with the rare surnames excluded) shows alinear trend
The surname distribution observed among husbandsand wives has also been considered with respect to theirprogeny size to investigate variation of fertility differ-entials with respect to surnames among the post-repro-ductive wives and their husbands separately for malefemale and total progeny size The results of thedistribution show a non-linear trend suggesting a dif-ferential reproductive fitness for different surnamesamong husbands and wives ( ) In the case ofhusbands the rare surnames (occurring once or twice)and common surnames (occurring more than 6-9
Maji Selective Neutrality of Surname Distribution in Adi Panggi 7
Occurrence of surname distribution (s) total number of male ( ) female ( ) andtotal children (times) among Adi Panggi husbands and wives (above 45 Years) and progeny sizedistribution (--)
times) show a trend toward higher progeny size (malesfemales and total children) than those surnames thatoccur less frequently (between 3-5 times) The best fitcurve in case of total children (polynomial degree three)accounts for about 945 percent of the variance (Rsup2 =0945) In case of females the trend shows a negativeassociation between occurrence of surnames and theprogeny size and the best fit curve for the total childrenaccounts (polynomial degree three) about 99 percent ofthe variance (Rsup2 = 0989) Progeny size shows a decreas-
ing trend with the rare surnames that occur once ortwice and a trend of lesser progeny in case of morecommon surnames (except in case the most commonsurnames that occur ten times among wives) Theseresults are a reflection of the population structure of thetribe especially the patrilocal system of marriage whichtends to increase the prevalence of some specific sur-names as a result of preferential marriages among a fewspecific clans or surnames
Husband ( gt 4 5 )
05
101520253035
0 2 4 6 8 10
Occurrence (s)
FrequencyW i f e ( gt 4 5 )
0
10
20
30
40
0 2 4 6 8 10
Occurrence (s)
Frequency
8
imilar distribution of progeny size with respect to sur-name distribution was also considered in case of repro-ductive wives and their husbandsrsquo sample ( )The pattern of distributions is quite varied compared tothe case of post-reproductive samples In the case of thewives the progeny size shows an increasing trend fromthe rare surnames to the most frequent surnamesthough there is some perturbance at the initial case ofrarer ones Among the husband samples a similar initialperturbance of progeny size is noticed with the rarersurnames but the progeny size shows stability with themore commonly occurring surnames of the tribe
Occurrence of surname distribution(s) total number of male ( ) female ( ) and totalchildren (times) among Adi Panggi husbands and wives (below 45 years) and progeny size distribution (--)
Husband (lt45)
0
10
20
30
40
50
60
0 5 10 15 20 25 30
Occurrence (s)
Frequency
W i f e ( lt 4 5 )
0
20
40
60
80
100
0 5 10 15 20 25 30
Occurrence (s)
Frequency
Though selective neutrality of surname distribution hasbeen validated in a number of populations in Europe andLatin American countries a few studies have showndeviation from neutrality implicating the role of migra-tion and operation of selection with respect to certainsurnames The results of the present study indicate thatsuch trends are also observed in small isolated popula-tions Apart from the expected influences of migrationand drift there could be other related factors of popula-tion structure that could cause the deviation from neutral-
Maji Selective Neutrality of Surname Distribution in Adi Panggi 9
ity In small tribal populations the structure of thesurname distribution might vary widely depending on thehistory of settlement prevailing marriage patterns andother socio-cultural and environmental reasons Furtherthe demographic parameters can vary drastically owing totheir subsistence pattern epidemiological factors andsocial organization such as warfare history Also internaltribal factions which are formed based on kinship or clanor lineages involving several closely knit families (eg inSouth Amerindian tribes Neel 1968 Salzano et al 1967)may play a role This can significantly influence thepopulation structure and can lead to perturbations in thesurname distribution in short time periods and in suchcases the neutral theory predictions may not hold Someof the above factors are important in explaining thedeviation from neutrality in surname distribution amongthe Adi Panggi
In general the Adi tribe consists of several sub-tribes whooccupy different terrains in the upper Himalayan moun-tain ranges and some in plains in the northeastern part ofthe country (Roy 1960 Lego 2005) The formation of thesub-tribes followed the fission-fusion model of popula-tion structure and was triggered by intertribal factionscompeting for survival necessities in this hunting-gather-ing population (Neel 1968 Salzano et al 1967) Theformation of Adi Panggi sub-tribes occurred a couple ofgenerations ago and was a result of conflict between amajor subgroup of the tribe and some other close-kinfactions over hunting rights and other resources (Roy1960 Lego 2005 Lego 2006) The sub-tribes now occu-py a recognized territory and constitute a few close affinaland consanguineous lineages and their affiliated clansSince they have a patrilocal society a majority of theirmen stay in the home village whereas females come fromdifferent villages and from other Adi sub-tribes Thisresults in more female marital migration and greatersurname diversity among wives We therefore observe ahigher proportion of homonymy of surname diversityamong males For example the five most frequent sur-names represent about 65 of total husbands while raresurnames (that occur either once or twice) constituteabout 5 per cent The three most common surnamesamong wives also represent a large part of the populationbut in contrast with the males the rare surnames thatoccur either once or twice constitute 18 per cent morethan 35 times the proportion of rare surnames in hus-bands In the total sample of husbands and wives sixsurnames represent 69 percent of the total populationstudied The surname distribution therefore reflectsgreater prevalence of most frequent (occurring in 20 to 70cases) and rare surnames (occurring once or twice) asagainst the moderately frequent surnames (that occur 3 to20 times in the sample)
The estimates of unbiased Isonymy also differ betweenhusband and wives samples The combined samples(husband and wives) show higher values of estimatedIsonymy (I = 01025) suggesting higher levels of homozy-gosity as a result of more frequent marriages contracted
among a few selected surnames generally found in suchisolated tribes This is reflected in the Karlin andMacGregor distribution which shows the non-lineartrend and the trend remains the same even after thelogarithmic transformation This also explains the widedifferences between the male and female surname distri-butions and the deviations observed in the Karlin andMacGregor and logarithmic distributions Therefore ifthe truncated distribution (omitting the rare surnames) isconsidered it shows a linear trend in the combinedsample suggesting that the non-linearity is contributedby the occurrence of rare surnames in the sample Therare surnames are contributed more through femalesmigrating from other places However the discretePareto distribution shows a linear trend for both thesamples The linear trend is clearer in the female samplethan among males but the genetic implications of thePareto distributions are not apparent (Fox 1983)
The results of the distribution of surnames also showwide differences with respect to progeny size distributionin case of post-reproductive (completed fertility) andreproductive wives and their husbands The neutralallele model would suggest that the distribution shouldbe independent of the fertility and morality (Manrubia2002) but the present study indicates otherwise andthere appears to be some differential fertility for somesurnames The Adi Panggi sample shows greater progenysize (male female and total surviving children) for themost frequent and rare surnames and that is reflected inthe near U-shaped distribution especially in case of hus-band sample This suggests reproductive advantageespecially greater success for the most frequent (6-910)and rare surnames (1-2) as against the less frequentsurnames (3-6) Among the Adi Panggi the four mostfrequent surnames among husbands and wives (especiallyPanyang Paron Mongku and Taku) show higher proge-ny size among the husband sample whereas the wivessample shows a near linear trend of decreasing progenysize from the rare surnames to the most frequent ones Apaired t-test between (eight) different types of surnamesand their progeny size shows higher significance values incase of the husbands (above 45 years) sample (p =0000433) than among the wives (above 45 years) (p =0003263) The mothers and their husbands were all inpost-reproductive period and had attained their maxi-mum fertility potential and do not use family planningand therefore the possibility of other intervening factorscan be excluded It is possible that it could be also dueto selective mortality of children especially higher mor-tality in case of less frequent surnames than with moreand rare frequent ones and this needs to be verifiedHowever there are no apparent cultural active selectivemechanisms that can influence the progeny size amongsome families of the tribe A detailed study on themarriage pattern and surname or clan distribution mightreveal more details The progeny distribution amongwomen of completed fertility is expected to follow Pois-son and Negative binomial distributions (Cavalli-Sforza
10
1971 Vasulu 1980) In patrilineal societies femalesgenerally show a greater surname diversity than malesand as a result the surname dynamics differ with respectto mitochondria and Y-chromosome
In the Indian context surname analysis has been carriedout in a few populations for investigating the effect onpopulation structure of inbreeding (Bhalla 1976 Kashyap1980 Vasulu 1980) genetic kinship (Kashyap 1980Vasulu 1980) preferential mating (Vasulu 1989 Reddy1995 Maji et al 2007) and social network analysisamong castes and tribes (Rao 1995) Surname neutralityhas also been validated in a caste community of Indiansresiding in Texas (Chakraborty 1990)
The above few studies suggest that isonymy can be ofmuch value in inferring population structure in somecastes and tribes (Kashyap 1980 Maji et al 2007) Thesurname structure in India varies among different regionscastes tribes linguistic and religious groups and providesan opportunity to study microevolution and populationstructure where the assumptions of surname or isonymyanalysis are valid Its applicability in diverse populationsespecially among isolated populations is expected to yieldfurther insights of its validity and inferring populationstructure and microevolutionary history
The present study illustrates the importance of surnamein inferring the genetic structure among a small sub-tribeAdi Panggi an isolated Tibeto-Burman- speaking com-munity in Arunachal Pradesh state in the northeasternregion of India The tribe lives on shifting-hunting-and-gathering subsistence and is a patrilocal and patrilinealsociety where the marriage pattern involves movement ofwives from other villages to the husbandrsquos village whichallows more surname diversity through females As aconsequence of differential surname structure betweenhusbands and wives the surname distribution showsdeviation from the expected neutral allele model and italso exhibits a non-linear trend of progeny size distribu-tion with respect to surname occurrence in the husbandsample
The results of the present study illustrate that surnamedistribution could be of much value in deciphering thepopulation structure and microevolution of a population(Crow 1965 Lasker 1978 Yasuda et al 1974) Thenon-linear trend observed in this study appears to beunique in the Indian context especially for the smallisolated tribes
The results of the study will help in further confirming thevalidity of the findings in several other tribes in India Inthe absence of any biological or genetic data for a tribethe surnames provide insights into the population struc-ture and possible genetic implications which can help inthe design of future studies As a prelude to the moleculargenetic investigations about the past genetic history asurname analysis can provide considerable insight into
the expected genetic structure of the population It alsohelps us to frame specific hypothesis related to male andfemale migration history which we will be testing withthe molecular genetic studies The two approachestogether can help to unravel the differential paternaland maternal past genetic history of the tribe
We thank the Government of Arunachal Pradesh andall the participants for their active cooperation in ourstudy Our sincere thanks go to Mr Tozing Tagi andMr Oyam Panyang (Katan and Geku circles) for theirassistance in data collection We acknowledge thesupport and intellectual inputs about the Adi tribesfrom Mr R N Koley (ARO Directorate of Research)and Mr Kaling Borang of Pasighat We thank Ms SKrithika Senior Research Fellow Biological Anthro-pology Unit for suggestions and help with the analysisand draft finalization and the Director Indian Statisti-cal Institute Kolkata for the financial assistance for thestudy among the tribes of Arunachal Pradesh
Barrai I Barbujani G Beretta M Maestri I Russo A Formi-ca G Pinto-Cisternas (1987) Surnames in Ferrara distribu-tion isonymy and levels of inbreeding14415-423
Barrai I Formica G Barale R Beretta M (1989) Isonymy andmigration distance 53249-262
Barrai I Formica G Barale R Scapoli C Canella R BerettaM (1990) Isonymy in emigrants from Ferrara in 1981-1988Ann Hum Biol 177-18
Barrai I Scapoli C Formica G et al (1991) Isonymy inrecords of births and deaths in Ferrara8395-404
Barrai I Formica G Scapoli C et al (1992) Microevolutionin Ferrara Isonymy 1890-1990 19371-385
Barrai I Scapoli C Bereta M et al (1996) Isonymy and thegenetic structure of Switzerland I The distributions of sur-names 23431-455
Barrai I Rodriguez-Larralde A Mamolini E et al (2000)Elements of surname structure of Austria27607-622
Barrai I Rodriguez-Larralde A Manni F et al (2002) Isony-my and isolation by distance in the Netherlands 74263-283
Barrai I Rodriguez-Larralde A Manni F Ruggiero V TartariD Scapoli C (2003) Isolation by language and distance inBeligium 681-16
Bhalla V Bhatia K (1976) Isonymy in a Bhatia leut 39479-500
Bhattacharjee PN (1954) Blood group Investigations in theAbor Tribe 351-54
11
Blackburn S (2004) Memories of migration Notes on legendsand beads in Arunachal Pradesh India
252615ndash60
Boldsen J (1992) Geographical distribution of some Danishsurnames reflection of social and natural selection 24505-513
Cavelli Sfroza LL Bodmer WF (1971) W H Freeman and Co San Francisco
Chakraborty R Weiss KM Rossmann DL Norton SL (1981)Distribution of last names a stochastic model for lilkelihooddetermination in record linking In Morrill W Dyke B (ed)
Academic Press New York
Chakraborty R Schwartz RJ (1990) Selective neutrality ofsurname distribution in an immigrant Indian community ofHouston Texas 21-13
Colantonio SE Lasker GW Kaplan BA Fuster V (2003) Useof surname models in human population biology A review ofrecent developments 75785-807
Crow JF Manage AP (1965) Measurements of Inbreeding ofthe frequencies of marriages between persons of the samesurnames 12199-203
Das TC (1953) Social Organization of the tribal peoples XIV245-262
Dyke B James AV Morrill WT (1983) Estimation of randomisonymy 10295-298
Fox WR Lasker GW (1983) The distribution of surnamefrequencies 5181-87
Karlin S MacGregor J (1967) The number of mutant formsmaintained in a population
4415-438
Kashyap LK (1980) Trends of Isonymy and inbreeding amongthe Ahmadiyyas of Kashmir 1219-25
Kashyap LK Tiwari SC (1980) Kinetics of genetic kinship asinferred by isonymy among the Ahmadiyyas of Kashmir valley
52311-324
Kimura M (1983) Cambridge University Press Cambridge
Koley RN (2005) Socio-cultural aspects of Adi Panggi Direc-torate of Research Government of Arunachal Pradesh Itanagar(in press)
Krithika S Maji S Vasulu TS (2006) Genetic heterogeneityamong three Adi tribes of Arunachal Pradesh India78221-227
Lasker GW (1978) Increments through migration to the coeffi-cient of relationship between communities by isonymy
50235-240
Lego N (2005) History of the Adis of Arunachal Pradesh MrsPonung Lego Itanagar
Lego N (2006) Modern History of Arunachal Pradesh (1825 ndash2006) Mrs Ponung Lego Itanagar
Maji S Krithika S Vasulu TS (2007) Genetic kinship among anisolated Adi tribe of Arunachal Pradesh isonymy in Adi Panggi
79321-337
Manrubia SC Zanette DH (2002) At the boundary betweenbiological and cultural evolution The origin of surname distri-butions 216461-477
Neel JV Chagnon NA (1968) The demography of two tribes ofprimitive relatively unacculturated American Indians
(USA) 59680-689
Rao AR (1995) Reciprocity in marital and social networksIllustration with Indian data 67887-904
Reddy BM Malhotra KC (1995) Relevance of surname inpopulation structure A study among the Vadde a fishingcommunity of Kolleru lake in Andhra Pradesh India
6161-168
Relethford JH (1988) Estimation of Kinship and Genetic dis-tance from Surnames 60475-492
Rodriguez LA Barrai I Alfonzo JC (1993) Isonymy structureof four Venezuelan states 20131-145
Rodriguez-Larralde A Formica G Scapoli C (1993) Microevo-lution in Perugia Isonymy 1890-1990 20261-274
Rodriguez-Larralde A Gonzales-Martin A Scapoli C Barrai I(2003) The names of Spain A study of the isonymy structureof Spain 121280-292
Roy S (1960) Aspects of Padam Minyong Culture Directorateof Research Govt of Arunachal Pradesh Itanagar
Roy S (1966) Anthropometry of the Adis of the Siang districtNorth-East Frontier Agency Shillong
Salzano FM Neel JV Maybury Lewis D (1967) Further studieson Xavante Indians I Demographic data on two additionalvillages and genetic structure of the tribe19463-489
Sarma J (1960) Puberty Marriage and Childbirth among thePangi and the Minyong Abor women 5597-113
Scapoli C Rodriguez-Larralde A Beretta M (1997) Correlationbetween isonymy parameters 1217-37
Singh IP (1994) Anthropological Survey of India Oxford
University Press New Delhi
Vasulu TS (1980) Anthrop Soc
of India Mysore
Vasulu TS (1989) Genetic structure of a tribal population 1Breeding isolation among the Yanadis 4255-268
Yasuda N Cavalli-Sfroza LL Skolnir M Moroni A (1974) Theevolution of surnames an analysis of their distribution andextinction 5123-142
Zei G Guglielmino CR Siri E Moroni A Cavalli-Sforza LL(1983a) Surnames as neutral alleles Observation in Sardinia
55357-365
Zei G Matessi RG Siri E et al (1983b) Surnames in SardiniaI Fit of frequency distributions for neutral articles and geneticpopulation structure 47329-352
6
Log-log distribution of surnames in the Adi Panggi tribe
Distribution of surnames among the Adi Panggi tribe
the sample The unbiased random isonymy I i j for theAdi Panggi samples is 01025
shows the estimates the two parametersof Karlin-MacGregor and Fishersrsquo distributions that de-scribe the differential migration rates and surname diver-sity among six villages of Adi Panggi Both the husbandsand wives show wide variation in the rate of migrationand surname diversity between the villages The twovillages KM and RK show the least migration rates andsurname diversity whereas SB shows the highest values of
in case of the husband samples
Panggi wives show higher surname diversity and migra-tion rates than husbands except in case of GT Panggiand non-Panggi wives show higher values than husbands( ) The estimates of F F based on randomisonymy show lesser values than from the Karlin-MacGregor and Fishersrsquo distributions The results showa similar pattern among husbands (n F = 00513 and a F
= 83278) and Panggi plus non-Panggi wives (n F = 00559and a F = 87113)
The logarithmic distribution of surname occurrenceamong husbands and wives and in the total sample for thePanggi is shown in
Both the figures show a non-linear trend suggesting devi-ation from the expected neutral allele model The shapeof the distribution is different in husbands and wives it is
an inverted parabolic curve in males and a slightlyconcave curve in females The best non-linear fit of thecurve with polynomial degree four show a variance ofabout 60 (Rsup2 = 059) in the case of the husbandssample while for wives a polynomial degree threeshows a good fit to the distribution (Rsup2 = 080) whichis better than the linear fit (Rsup2 = 074) In the case ofall Adi Panggi (both husbands and wives) the distribu-tions show a nonlinear trend which is different fromthe trend observed in the case of husband and wifesamples ( ) The best fit to the curve is apolynomial of degree four which accounts for about96 of the variance The non-linear distribution is dueto the higher frequency of occurrence of rare surnames(especially those that occur only once and are contrib-uted by wives) However the truncated log-log distri-bution (with the rare surnames excluded) shows alinear trend
The surname distribution observed among husbandsand wives has also been considered with respect to theirprogeny size to investigate variation of fertility differ-entials with respect to surnames among the post-repro-ductive wives and their husbands separately for malefemale and total progeny size The results of thedistribution show a non-linear trend suggesting a dif-ferential reproductive fitness for different surnamesamong husbands and wives ( ) In the case ofhusbands the rare surnames (occurring once or twice)and common surnames (occurring more than 6-9
Maji Selective Neutrality of Surname Distribution in Adi Panggi 7
Occurrence of surname distribution (s) total number of male ( ) female ( ) andtotal children (times) among Adi Panggi husbands and wives (above 45 Years) and progeny sizedistribution (--)
times) show a trend toward higher progeny size (malesfemales and total children) than those surnames thatoccur less frequently (between 3-5 times) The best fitcurve in case of total children (polynomial degree three)accounts for about 945 percent of the variance (Rsup2 =0945) In case of females the trend shows a negativeassociation between occurrence of surnames and theprogeny size and the best fit curve for the total childrenaccounts (polynomial degree three) about 99 percent ofthe variance (Rsup2 = 0989) Progeny size shows a decreas-
ing trend with the rare surnames that occur once ortwice and a trend of lesser progeny in case of morecommon surnames (except in case the most commonsurnames that occur ten times among wives) Theseresults are a reflection of the population structure of thetribe especially the patrilocal system of marriage whichtends to increase the prevalence of some specific sur-names as a result of preferential marriages among a fewspecific clans or surnames
Husband ( gt 4 5 )
05
101520253035
0 2 4 6 8 10
Occurrence (s)
FrequencyW i f e ( gt 4 5 )
0
10
20
30
40
0 2 4 6 8 10
Occurrence (s)
Frequency
8
imilar distribution of progeny size with respect to sur-name distribution was also considered in case of repro-ductive wives and their husbandsrsquo sample ( )The pattern of distributions is quite varied compared tothe case of post-reproductive samples In the case of thewives the progeny size shows an increasing trend fromthe rare surnames to the most frequent surnamesthough there is some perturbance at the initial case ofrarer ones Among the husband samples a similar initialperturbance of progeny size is noticed with the rarersurnames but the progeny size shows stability with themore commonly occurring surnames of the tribe
Occurrence of surname distribution(s) total number of male ( ) female ( ) and totalchildren (times) among Adi Panggi husbands and wives (below 45 years) and progeny size distribution (--)
Husband (lt45)
0
10
20
30
40
50
60
0 5 10 15 20 25 30
Occurrence (s)
Frequency
W i f e ( lt 4 5 )
0
20
40
60
80
100
0 5 10 15 20 25 30
Occurrence (s)
Frequency
Though selective neutrality of surname distribution hasbeen validated in a number of populations in Europe andLatin American countries a few studies have showndeviation from neutrality implicating the role of migra-tion and operation of selection with respect to certainsurnames The results of the present study indicate thatsuch trends are also observed in small isolated popula-tions Apart from the expected influences of migrationand drift there could be other related factors of popula-tion structure that could cause the deviation from neutral-
Maji Selective Neutrality of Surname Distribution in Adi Panggi 9
ity In small tribal populations the structure of thesurname distribution might vary widely depending on thehistory of settlement prevailing marriage patterns andother socio-cultural and environmental reasons Furtherthe demographic parameters can vary drastically owing totheir subsistence pattern epidemiological factors andsocial organization such as warfare history Also internaltribal factions which are formed based on kinship or clanor lineages involving several closely knit families (eg inSouth Amerindian tribes Neel 1968 Salzano et al 1967)may play a role This can significantly influence thepopulation structure and can lead to perturbations in thesurname distribution in short time periods and in suchcases the neutral theory predictions may not hold Someof the above factors are important in explaining thedeviation from neutrality in surname distribution amongthe Adi Panggi
In general the Adi tribe consists of several sub-tribes whooccupy different terrains in the upper Himalayan moun-tain ranges and some in plains in the northeastern part ofthe country (Roy 1960 Lego 2005) The formation of thesub-tribes followed the fission-fusion model of popula-tion structure and was triggered by intertribal factionscompeting for survival necessities in this hunting-gather-ing population (Neel 1968 Salzano et al 1967) Theformation of Adi Panggi sub-tribes occurred a couple ofgenerations ago and was a result of conflict between amajor subgroup of the tribe and some other close-kinfactions over hunting rights and other resources (Roy1960 Lego 2005 Lego 2006) The sub-tribes now occu-py a recognized territory and constitute a few close affinaland consanguineous lineages and their affiliated clansSince they have a patrilocal society a majority of theirmen stay in the home village whereas females come fromdifferent villages and from other Adi sub-tribes Thisresults in more female marital migration and greatersurname diversity among wives We therefore observe ahigher proportion of homonymy of surname diversityamong males For example the five most frequent sur-names represent about 65 of total husbands while raresurnames (that occur either once or twice) constituteabout 5 per cent The three most common surnamesamong wives also represent a large part of the populationbut in contrast with the males the rare surnames thatoccur either once or twice constitute 18 per cent morethan 35 times the proportion of rare surnames in hus-bands In the total sample of husbands and wives sixsurnames represent 69 percent of the total populationstudied The surname distribution therefore reflectsgreater prevalence of most frequent (occurring in 20 to 70cases) and rare surnames (occurring once or twice) asagainst the moderately frequent surnames (that occur 3 to20 times in the sample)
The estimates of unbiased Isonymy also differ betweenhusband and wives samples The combined samples(husband and wives) show higher values of estimatedIsonymy (I = 01025) suggesting higher levels of homozy-gosity as a result of more frequent marriages contracted
among a few selected surnames generally found in suchisolated tribes This is reflected in the Karlin andMacGregor distribution which shows the non-lineartrend and the trend remains the same even after thelogarithmic transformation This also explains the widedifferences between the male and female surname distri-butions and the deviations observed in the Karlin andMacGregor and logarithmic distributions Therefore ifthe truncated distribution (omitting the rare surnames) isconsidered it shows a linear trend in the combinedsample suggesting that the non-linearity is contributedby the occurrence of rare surnames in the sample Therare surnames are contributed more through femalesmigrating from other places However the discretePareto distribution shows a linear trend for both thesamples The linear trend is clearer in the female samplethan among males but the genetic implications of thePareto distributions are not apparent (Fox 1983)
The results of the distribution of surnames also showwide differences with respect to progeny size distributionin case of post-reproductive (completed fertility) andreproductive wives and their husbands The neutralallele model would suggest that the distribution shouldbe independent of the fertility and morality (Manrubia2002) but the present study indicates otherwise andthere appears to be some differential fertility for somesurnames The Adi Panggi sample shows greater progenysize (male female and total surviving children) for themost frequent and rare surnames and that is reflected inthe near U-shaped distribution especially in case of hus-band sample This suggests reproductive advantageespecially greater success for the most frequent (6-910)and rare surnames (1-2) as against the less frequentsurnames (3-6) Among the Adi Panggi the four mostfrequent surnames among husbands and wives (especiallyPanyang Paron Mongku and Taku) show higher proge-ny size among the husband sample whereas the wivessample shows a near linear trend of decreasing progenysize from the rare surnames to the most frequent ones Apaired t-test between (eight) different types of surnamesand their progeny size shows higher significance values incase of the husbands (above 45 years) sample (p =0000433) than among the wives (above 45 years) (p =0003263) The mothers and their husbands were all inpost-reproductive period and had attained their maxi-mum fertility potential and do not use family planningand therefore the possibility of other intervening factorscan be excluded It is possible that it could be also dueto selective mortality of children especially higher mor-tality in case of less frequent surnames than with moreand rare frequent ones and this needs to be verifiedHowever there are no apparent cultural active selectivemechanisms that can influence the progeny size amongsome families of the tribe A detailed study on themarriage pattern and surname or clan distribution mightreveal more details The progeny distribution amongwomen of completed fertility is expected to follow Pois-son and Negative binomial distributions (Cavalli-Sforza
10
1971 Vasulu 1980) In patrilineal societies femalesgenerally show a greater surname diversity than malesand as a result the surname dynamics differ with respectto mitochondria and Y-chromosome
In the Indian context surname analysis has been carriedout in a few populations for investigating the effect onpopulation structure of inbreeding (Bhalla 1976 Kashyap1980 Vasulu 1980) genetic kinship (Kashyap 1980Vasulu 1980) preferential mating (Vasulu 1989 Reddy1995 Maji et al 2007) and social network analysisamong castes and tribes (Rao 1995) Surname neutralityhas also been validated in a caste community of Indiansresiding in Texas (Chakraborty 1990)
The above few studies suggest that isonymy can be ofmuch value in inferring population structure in somecastes and tribes (Kashyap 1980 Maji et al 2007) Thesurname structure in India varies among different regionscastes tribes linguistic and religious groups and providesan opportunity to study microevolution and populationstructure where the assumptions of surname or isonymyanalysis are valid Its applicability in diverse populationsespecially among isolated populations is expected to yieldfurther insights of its validity and inferring populationstructure and microevolutionary history
The present study illustrates the importance of surnamein inferring the genetic structure among a small sub-tribeAdi Panggi an isolated Tibeto-Burman- speaking com-munity in Arunachal Pradesh state in the northeasternregion of India The tribe lives on shifting-hunting-and-gathering subsistence and is a patrilocal and patrilinealsociety where the marriage pattern involves movement ofwives from other villages to the husbandrsquos village whichallows more surname diversity through females As aconsequence of differential surname structure betweenhusbands and wives the surname distribution showsdeviation from the expected neutral allele model and italso exhibits a non-linear trend of progeny size distribu-tion with respect to surname occurrence in the husbandsample
The results of the present study illustrate that surnamedistribution could be of much value in deciphering thepopulation structure and microevolution of a population(Crow 1965 Lasker 1978 Yasuda et al 1974) Thenon-linear trend observed in this study appears to beunique in the Indian context especially for the smallisolated tribes
The results of the study will help in further confirming thevalidity of the findings in several other tribes in India Inthe absence of any biological or genetic data for a tribethe surnames provide insights into the population struc-ture and possible genetic implications which can help inthe design of future studies As a prelude to the moleculargenetic investigations about the past genetic history asurname analysis can provide considerable insight into
the expected genetic structure of the population It alsohelps us to frame specific hypothesis related to male andfemale migration history which we will be testing withthe molecular genetic studies The two approachestogether can help to unravel the differential paternaland maternal past genetic history of the tribe
We thank the Government of Arunachal Pradesh andall the participants for their active cooperation in ourstudy Our sincere thanks go to Mr Tozing Tagi andMr Oyam Panyang (Katan and Geku circles) for theirassistance in data collection We acknowledge thesupport and intellectual inputs about the Adi tribesfrom Mr R N Koley (ARO Directorate of Research)and Mr Kaling Borang of Pasighat We thank Ms SKrithika Senior Research Fellow Biological Anthro-pology Unit for suggestions and help with the analysisand draft finalization and the Director Indian Statisti-cal Institute Kolkata for the financial assistance for thestudy among the tribes of Arunachal Pradesh
Barrai I Barbujani G Beretta M Maestri I Russo A Formi-ca G Pinto-Cisternas (1987) Surnames in Ferrara distribu-tion isonymy and levels of inbreeding14415-423
Barrai I Formica G Barale R Beretta M (1989) Isonymy andmigration distance 53249-262
Barrai I Formica G Barale R Scapoli C Canella R BerettaM (1990) Isonymy in emigrants from Ferrara in 1981-1988Ann Hum Biol 177-18
Barrai I Scapoli C Formica G et al (1991) Isonymy inrecords of births and deaths in Ferrara8395-404
Barrai I Formica G Scapoli C et al (1992) Microevolutionin Ferrara Isonymy 1890-1990 19371-385
Barrai I Scapoli C Bereta M et al (1996) Isonymy and thegenetic structure of Switzerland I The distributions of sur-names 23431-455
Barrai I Rodriguez-Larralde A Mamolini E et al (2000)Elements of surname structure of Austria27607-622
Barrai I Rodriguez-Larralde A Manni F et al (2002) Isony-my and isolation by distance in the Netherlands 74263-283
Barrai I Rodriguez-Larralde A Manni F Ruggiero V TartariD Scapoli C (2003) Isolation by language and distance inBeligium 681-16
Bhalla V Bhatia K (1976) Isonymy in a Bhatia leut 39479-500
Bhattacharjee PN (1954) Blood group Investigations in theAbor Tribe 351-54
11
Blackburn S (2004) Memories of migration Notes on legendsand beads in Arunachal Pradesh India
252615ndash60
Boldsen J (1992) Geographical distribution of some Danishsurnames reflection of social and natural selection 24505-513
Cavelli Sfroza LL Bodmer WF (1971) W H Freeman and Co San Francisco
Chakraborty R Weiss KM Rossmann DL Norton SL (1981)Distribution of last names a stochastic model for lilkelihooddetermination in record linking In Morrill W Dyke B (ed)
Academic Press New York
Chakraborty R Schwartz RJ (1990) Selective neutrality ofsurname distribution in an immigrant Indian community ofHouston Texas 21-13
Colantonio SE Lasker GW Kaplan BA Fuster V (2003) Useof surname models in human population biology A review ofrecent developments 75785-807
Crow JF Manage AP (1965) Measurements of Inbreeding ofthe frequencies of marriages between persons of the samesurnames 12199-203
Das TC (1953) Social Organization of the tribal peoples XIV245-262
Dyke B James AV Morrill WT (1983) Estimation of randomisonymy 10295-298
Fox WR Lasker GW (1983) The distribution of surnamefrequencies 5181-87
Karlin S MacGregor J (1967) The number of mutant formsmaintained in a population
4415-438
Kashyap LK (1980) Trends of Isonymy and inbreeding amongthe Ahmadiyyas of Kashmir 1219-25
Kashyap LK Tiwari SC (1980) Kinetics of genetic kinship asinferred by isonymy among the Ahmadiyyas of Kashmir valley
52311-324
Kimura M (1983) Cambridge University Press Cambridge
Koley RN (2005) Socio-cultural aspects of Adi Panggi Direc-torate of Research Government of Arunachal Pradesh Itanagar(in press)
Krithika S Maji S Vasulu TS (2006) Genetic heterogeneityamong three Adi tribes of Arunachal Pradesh India78221-227
Lasker GW (1978) Increments through migration to the coeffi-cient of relationship between communities by isonymy
50235-240
Lego N (2005) History of the Adis of Arunachal Pradesh MrsPonung Lego Itanagar
Lego N (2006) Modern History of Arunachal Pradesh (1825 ndash2006) Mrs Ponung Lego Itanagar
Maji S Krithika S Vasulu TS (2007) Genetic kinship among anisolated Adi tribe of Arunachal Pradesh isonymy in Adi Panggi
79321-337
Manrubia SC Zanette DH (2002) At the boundary betweenbiological and cultural evolution The origin of surname distri-butions 216461-477
Neel JV Chagnon NA (1968) The demography of two tribes ofprimitive relatively unacculturated American Indians
(USA) 59680-689
Rao AR (1995) Reciprocity in marital and social networksIllustration with Indian data 67887-904
Reddy BM Malhotra KC (1995) Relevance of surname inpopulation structure A study among the Vadde a fishingcommunity of Kolleru lake in Andhra Pradesh India
6161-168
Relethford JH (1988) Estimation of Kinship and Genetic dis-tance from Surnames 60475-492
Rodriguez LA Barrai I Alfonzo JC (1993) Isonymy structureof four Venezuelan states 20131-145
Rodriguez-Larralde A Formica G Scapoli C (1993) Microevo-lution in Perugia Isonymy 1890-1990 20261-274
Rodriguez-Larralde A Gonzales-Martin A Scapoli C Barrai I(2003) The names of Spain A study of the isonymy structureof Spain 121280-292
Roy S (1960) Aspects of Padam Minyong Culture Directorateof Research Govt of Arunachal Pradesh Itanagar
Roy S (1966) Anthropometry of the Adis of the Siang districtNorth-East Frontier Agency Shillong
Salzano FM Neel JV Maybury Lewis D (1967) Further studieson Xavante Indians I Demographic data on two additionalvillages and genetic structure of the tribe19463-489
Sarma J (1960) Puberty Marriage and Childbirth among thePangi and the Minyong Abor women 5597-113
Scapoli C Rodriguez-Larralde A Beretta M (1997) Correlationbetween isonymy parameters 1217-37
Singh IP (1994) Anthropological Survey of India Oxford
University Press New Delhi
Vasulu TS (1980) Anthrop Soc
of India Mysore
Vasulu TS (1989) Genetic structure of a tribal population 1Breeding isolation among the Yanadis 4255-268
Yasuda N Cavalli-Sfroza LL Skolnir M Moroni A (1974) Theevolution of surnames an analysis of their distribution andextinction 5123-142
Zei G Guglielmino CR Siri E Moroni A Cavalli-Sforza LL(1983a) Surnames as neutral alleles Observation in Sardinia
55357-365
Zei G Matessi RG Siri E et al (1983b) Surnames in SardiniaI Fit of frequency distributions for neutral articles and geneticpopulation structure 47329-352
Maji Selective Neutrality of Surname Distribution in Adi Panggi 7
Occurrence of surname distribution (s) total number of male ( ) female ( ) andtotal children (times) among Adi Panggi husbands and wives (above 45 Years) and progeny sizedistribution (--)
times) show a trend toward higher progeny size (malesfemales and total children) than those surnames thatoccur less frequently (between 3-5 times) The best fitcurve in case of total children (polynomial degree three)accounts for about 945 percent of the variance (Rsup2 =0945) In case of females the trend shows a negativeassociation between occurrence of surnames and theprogeny size and the best fit curve for the total childrenaccounts (polynomial degree three) about 99 percent ofthe variance (Rsup2 = 0989) Progeny size shows a decreas-
ing trend with the rare surnames that occur once ortwice and a trend of lesser progeny in case of morecommon surnames (except in case the most commonsurnames that occur ten times among wives) Theseresults are a reflection of the population structure of thetribe especially the patrilocal system of marriage whichtends to increase the prevalence of some specific sur-names as a result of preferential marriages among a fewspecific clans or surnames
Husband ( gt 4 5 )
05
101520253035
0 2 4 6 8 10
Occurrence (s)
FrequencyW i f e ( gt 4 5 )
0
10
20
30
40
0 2 4 6 8 10
Occurrence (s)
Frequency
8
imilar distribution of progeny size with respect to sur-name distribution was also considered in case of repro-ductive wives and their husbandsrsquo sample ( )The pattern of distributions is quite varied compared tothe case of post-reproductive samples In the case of thewives the progeny size shows an increasing trend fromthe rare surnames to the most frequent surnamesthough there is some perturbance at the initial case ofrarer ones Among the husband samples a similar initialperturbance of progeny size is noticed with the rarersurnames but the progeny size shows stability with themore commonly occurring surnames of the tribe
Occurrence of surname distribution(s) total number of male ( ) female ( ) and totalchildren (times) among Adi Panggi husbands and wives (below 45 years) and progeny size distribution (--)
Husband (lt45)
0
10
20
30
40
50
60
0 5 10 15 20 25 30
Occurrence (s)
Frequency
W i f e ( lt 4 5 )
0
20
40
60
80
100
0 5 10 15 20 25 30
Occurrence (s)
Frequency
Though selective neutrality of surname distribution hasbeen validated in a number of populations in Europe andLatin American countries a few studies have showndeviation from neutrality implicating the role of migra-tion and operation of selection with respect to certainsurnames The results of the present study indicate thatsuch trends are also observed in small isolated popula-tions Apart from the expected influences of migrationand drift there could be other related factors of popula-tion structure that could cause the deviation from neutral-
Maji Selective Neutrality of Surname Distribution in Adi Panggi 9
ity In small tribal populations the structure of thesurname distribution might vary widely depending on thehistory of settlement prevailing marriage patterns andother socio-cultural and environmental reasons Furtherthe demographic parameters can vary drastically owing totheir subsistence pattern epidemiological factors andsocial organization such as warfare history Also internaltribal factions which are formed based on kinship or clanor lineages involving several closely knit families (eg inSouth Amerindian tribes Neel 1968 Salzano et al 1967)may play a role This can significantly influence thepopulation structure and can lead to perturbations in thesurname distribution in short time periods and in suchcases the neutral theory predictions may not hold Someof the above factors are important in explaining thedeviation from neutrality in surname distribution amongthe Adi Panggi
In general the Adi tribe consists of several sub-tribes whooccupy different terrains in the upper Himalayan moun-tain ranges and some in plains in the northeastern part ofthe country (Roy 1960 Lego 2005) The formation of thesub-tribes followed the fission-fusion model of popula-tion structure and was triggered by intertribal factionscompeting for survival necessities in this hunting-gather-ing population (Neel 1968 Salzano et al 1967) Theformation of Adi Panggi sub-tribes occurred a couple ofgenerations ago and was a result of conflict between amajor subgroup of the tribe and some other close-kinfactions over hunting rights and other resources (Roy1960 Lego 2005 Lego 2006) The sub-tribes now occu-py a recognized territory and constitute a few close affinaland consanguineous lineages and their affiliated clansSince they have a patrilocal society a majority of theirmen stay in the home village whereas females come fromdifferent villages and from other Adi sub-tribes Thisresults in more female marital migration and greatersurname diversity among wives We therefore observe ahigher proportion of homonymy of surname diversityamong males For example the five most frequent sur-names represent about 65 of total husbands while raresurnames (that occur either once or twice) constituteabout 5 per cent The three most common surnamesamong wives also represent a large part of the populationbut in contrast with the males the rare surnames thatoccur either once or twice constitute 18 per cent morethan 35 times the proportion of rare surnames in hus-bands In the total sample of husbands and wives sixsurnames represent 69 percent of the total populationstudied The surname distribution therefore reflectsgreater prevalence of most frequent (occurring in 20 to 70cases) and rare surnames (occurring once or twice) asagainst the moderately frequent surnames (that occur 3 to20 times in the sample)
The estimates of unbiased Isonymy also differ betweenhusband and wives samples The combined samples(husband and wives) show higher values of estimatedIsonymy (I = 01025) suggesting higher levels of homozy-gosity as a result of more frequent marriages contracted
among a few selected surnames generally found in suchisolated tribes This is reflected in the Karlin andMacGregor distribution which shows the non-lineartrend and the trend remains the same even after thelogarithmic transformation This also explains the widedifferences between the male and female surname distri-butions and the deviations observed in the Karlin andMacGregor and logarithmic distributions Therefore ifthe truncated distribution (omitting the rare surnames) isconsidered it shows a linear trend in the combinedsample suggesting that the non-linearity is contributedby the occurrence of rare surnames in the sample Therare surnames are contributed more through femalesmigrating from other places However the discretePareto distribution shows a linear trend for both thesamples The linear trend is clearer in the female samplethan among males but the genetic implications of thePareto distributions are not apparent (Fox 1983)
The results of the distribution of surnames also showwide differences with respect to progeny size distributionin case of post-reproductive (completed fertility) andreproductive wives and their husbands The neutralallele model would suggest that the distribution shouldbe independent of the fertility and morality (Manrubia2002) but the present study indicates otherwise andthere appears to be some differential fertility for somesurnames The Adi Panggi sample shows greater progenysize (male female and total surviving children) for themost frequent and rare surnames and that is reflected inthe near U-shaped distribution especially in case of hus-band sample This suggests reproductive advantageespecially greater success for the most frequent (6-910)and rare surnames (1-2) as against the less frequentsurnames (3-6) Among the Adi Panggi the four mostfrequent surnames among husbands and wives (especiallyPanyang Paron Mongku and Taku) show higher proge-ny size among the husband sample whereas the wivessample shows a near linear trend of decreasing progenysize from the rare surnames to the most frequent ones Apaired t-test between (eight) different types of surnamesand their progeny size shows higher significance values incase of the husbands (above 45 years) sample (p =0000433) than among the wives (above 45 years) (p =0003263) The mothers and their husbands were all inpost-reproductive period and had attained their maxi-mum fertility potential and do not use family planningand therefore the possibility of other intervening factorscan be excluded It is possible that it could be also dueto selective mortality of children especially higher mor-tality in case of less frequent surnames than with moreand rare frequent ones and this needs to be verifiedHowever there are no apparent cultural active selectivemechanisms that can influence the progeny size amongsome families of the tribe A detailed study on themarriage pattern and surname or clan distribution mightreveal more details The progeny distribution amongwomen of completed fertility is expected to follow Pois-son and Negative binomial distributions (Cavalli-Sforza
10
1971 Vasulu 1980) In patrilineal societies femalesgenerally show a greater surname diversity than malesand as a result the surname dynamics differ with respectto mitochondria and Y-chromosome
In the Indian context surname analysis has been carriedout in a few populations for investigating the effect onpopulation structure of inbreeding (Bhalla 1976 Kashyap1980 Vasulu 1980) genetic kinship (Kashyap 1980Vasulu 1980) preferential mating (Vasulu 1989 Reddy1995 Maji et al 2007) and social network analysisamong castes and tribes (Rao 1995) Surname neutralityhas also been validated in a caste community of Indiansresiding in Texas (Chakraborty 1990)
The above few studies suggest that isonymy can be ofmuch value in inferring population structure in somecastes and tribes (Kashyap 1980 Maji et al 2007) Thesurname structure in India varies among different regionscastes tribes linguistic and religious groups and providesan opportunity to study microevolution and populationstructure where the assumptions of surname or isonymyanalysis are valid Its applicability in diverse populationsespecially among isolated populations is expected to yieldfurther insights of its validity and inferring populationstructure and microevolutionary history
The present study illustrates the importance of surnamein inferring the genetic structure among a small sub-tribeAdi Panggi an isolated Tibeto-Burman- speaking com-munity in Arunachal Pradesh state in the northeasternregion of India The tribe lives on shifting-hunting-and-gathering subsistence and is a patrilocal and patrilinealsociety where the marriage pattern involves movement ofwives from other villages to the husbandrsquos village whichallows more surname diversity through females As aconsequence of differential surname structure betweenhusbands and wives the surname distribution showsdeviation from the expected neutral allele model and italso exhibits a non-linear trend of progeny size distribu-tion with respect to surname occurrence in the husbandsample
The results of the present study illustrate that surnamedistribution could be of much value in deciphering thepopulation structure and microevolution of a population(Crow 1965 Lasker 1978 Yasuda et al 1974) Thenon-linear trend observed in this study appears to beunique in the Indian context especially for the smallisolated tribes
The results of the study will help in further confirming thevalidity of the findings in several other tribes in India Inthe absence of any biological or genetic data for a tribethe surnames provide insights into the population struc-ture and possible genetic implications which can help inthe design of future studies As a prelude to the moleculargenetic investigations about the past genetic history asurname analysis can provide considerable insight into
the expected genetic structure of the population It alsohelps us to frame specific hypothesis related to male andfemale migration history which we will be testing withthe molecular genetic studies The two approachestogether can help to unravel the differential paternaland maternal past genetic history of the tribe
We thank the Government of Arunachal Pradesh andall the participants for their active cooperation in ourstudy Our sincere thanks go to Mr Tozing Tagi andMr Oyam Panyang (Katan and Geku circles) for theirassistance in data collection We acknowledge thesupport and intellectual inputs about the Adi tribesfrom Mr R N Koley (ARO Directorate of Research)and Mr Kaling Borang of Pasighat We thank Ms SKrithika Senior Research Fellow Biological Anthro-pology Unit for suggestions and help with the analysisand draft finalization and the Director Indian Statisti-cal Institute Kolkata for the financial assistance for thestudy among the tribes of Arunachal Pradesh
Barrai I Barbujani G Beretta M Maestri I Russo A Formi-ca G Pinto-Cisternas (1987) Surnames in Ferrara distribu-tion isonymy and levels of inbreeding14415-423
Barrai I Formica G Barale R Beretta M (1989) Isonymy andmigration distance 53249-262
Barrai I Formica G Barale R Scapoli C Canella R BerettaM (1990) Isonymy in emigrants from Ferrara in 1981-1988Ann Hum Biol 177-18
Barrai I Scapoli C Formica G et al (1991) Isonymy inrecords of births and deaths in Ferrara8395-404
Barrai I Formica G Scapoli C et al (1992) Microevolutionin Ferrara Isonymy 1890-1990 19371-385
Barrai I Scapoli C Bereta M et al (1996) Isonymy and thegenetic structure of Switzerland I The distributions of sur-names 23431-455
Barrai I Rodriguez-Larralde A Mamolini E et al (2000)Elements of surname structure of Austria27607-622
Barrai I Rodriguez-Larralde A Manni F et al (2002) Isony-my and isolation by distance in the Netherlands 74263-283
Barrai I Rodriguez-Larralde A Manni F Ruggiero V TartariD Scapoli C (2003) Isolation by language and distance inBeligium 681-16
Bhalla V Bhatia K (1976) Isonymy in a Bhatia leut 39479-500
Bhattacharjee PN (1954) Blood group Investigations in theAbor Tribe 351-54
11
Blackburn S (2004) Memories of migration Notes on legendsand beads in Arunachal Pradesh India
252615ndash60
Boldsen J (1992) Geographical distribution of some Danishsurnames reflection of social and natural selection 24505-513
Cavelli Sfroza LL Bodmer WF (1971) W H Freeman and Co San Francisco
Chakraborty R Weiss KM Rossmann DL Norton SL (1981)Distribution of last names a stochastic model for lilkelihooddetermination in record linking In Morrill W Dyke B (ed)
Academic Press New York
Chakraborty R Schwartz RJ (1990) Selective neutrality ofsurname distribution in an immigrant Indian community ofHouston Texas 21-13
Colantonio SE Lasker GW Kaplan BA Fuster V (2003) Useof surname models in human population biology A review ofrecent developments 75785-807
Crow JF Manage AP (1965) Measurements of Inbreeding ofthe frequencies of marriages between persons of the samesurnames 12199-203
Das TC (1953) Social Organization of the tribal peoples XIV245-262
Dyke B James AV Morrill WT (1983) Estimation of randomisonymy 10295-298
Fox WR Lasker GW (1983) The distribution of surnamefrequencies 5181-87
Karlin S MacGregor J (1967) The number of mutant formsmaintained in a population
4415-438
Kashyap LK (1980) Trends of Isonymy and inbreeding amongthe Ahmadiyyas of Kashmir 1219-25
Kashyap LK Tiwari SC (1980) Kinetics of genetic kinship asinferred by isonymy among the Ahmadiyyas of Kashmir valley
52311-324
Kimura M (1983) Cambridge University Press Cambridge
Koley RN (2005) Socio-cultural aspects of Adi Panggi Direc-torate of Research Government of Arunachal Pradesh Itanagar(in press)
Krithika S Maji S Vasulu TS (2006) Genetic heterogeneityamong three Adi tribes of Arunachal Pradesh India78221-227
Lasker GW (1978) Increments through migration to the coeffi-cient of relationship between communities by isonymy
50235-240
Lego N (2005) History of the Adis of Arunachal Pradesh MrsPonung Lego Itanagar
Lego N (2006) Modern History of Arunachal Pradesh (1825 ndash2006) Mrs Ponung Lego Itanagar
Maji S Krithika S Vasulu TS (2007) Genetic kinship among anisolated Adi tribe of Arunachal Pradesh isonymy in Adi Panggi
79321-337
Manrubia SC Zanette DH (2002) At the boundary betweenbiological and cultural evolution The origin of surname distri-butions 216461-477
Neel JV Chagnon NA (1968) The demography of two tribes ofprimitive relatively unacculturated American Indians
(USA) 59680-689
Rao AR (1995) Reciprocity in marital and social networksIllustration with Indian data 67887-904
Reddy BM Malhotra KC (1995) Relevance of surname inpopulation structure A study among the Vadde a fishingcommunity of Kolleru lake in Andhra Pradesh India
6161-168
Relethford JH (1988) Estimation of Kinship and Genetic dis-tance from Surnames 60475-492
Rodriguez LA Barrai I Alfonzo JC (1993) Isonymy structureof four Venezuelan states 20131-145
Rodriguez-Larralde A Formica G Scapoli C (1993) Microevo-lution in Perugia Isonymy 1890-1990 20261-274
Rodriguez-Larralde A Gonzales-Martin A Scapoli C Barrai I(2003) The names of Spain A study of the isonymy structureof Spain 121280-292
Roy S (1960) Aspects of Padam Minyong Culture Directorateof Research Govt of Arunachal Pradesh Itanagar
Roy S (1966) Anthropometry of the Adis of the Siang districtNorth-East Frontier Agency Shillong
Salzano FM Neel JV Maybury Lewis D (1967) Further studieson Xavante Indians I Demographic data on two additionalvillages and genetic structure of the tribe19463-489
Sarma J (1960) Puberty Marriage and Childbirth among thePangi and the Minyong Abor women 5597-113
Scapoli C Rodriguez-Larralde A Beretta M (1997) Correlationbetween isonymy parameters 1217-37
Singh IP (1994) Anthropological Survey of India Oxford
University Press New Delhi
Vasulu TS (1980) Anthrop Soc
of India Mysore
Vasulu TS (1989) Genetic structure of a tribal population 1Breeding isolation among the Yanadis 4255-268
Yasuda N Cavalli-Sfroza LL Skolnir M Moroni A (1974) Theevolution of surnames an analysis of their distribution andextinction 5123-142
Zei G Guglielmino CR Siri E Moroni A Cavalli-Sforza LL(1983a) Surnames as neutral alleles Observation in Sardinia
55357-365
Zei G Matessi RG Siri E et al (1983b) Surnames in SardiniaI Fit of frequency distributions for neutral articles and geneticpopulation structure 47329-352
8
imilar distribution of progeny size with respect to sur-name distribution was also considered in case of repro-ductive wives and their husbandsrsquo sample ( )The pattern of distributions is quite varied compared tothe case of post-reproductive samples In the case of thewives the progeny size shows an increasing trend fromthe rare surnames to the most frequent surnamesthough there is some perturbance at the initial case ofrarer ones Among the husband samples a similar initialperturbance of progeny size is noticed with the rarersurnames but the progeny size shows stability with themore commonly occurring surnames of the tribe
Occurrence of surname distribution(s) total number of male ( ) female ( ) and totalchildren (times) among Adi Panggi husbands and wives (below 45 years) and progeny size distribution (--)
Husband (lt45)
0
10
20
30
40
50
60
0 5 10 15 20 25 30
Occurrence (s)
Frequency
W i f e ( lt 4 5 )
0
20
40
60
80
100
0 5 10 15 20 25 30
Occurrence (s)
Frequency
Though selective neutrality of surname distribution hasbeen validated in a number of populations in Europe andLatin American countries a few studies have showndeviation from neutrality implicating the role of migra-tion and operation of selection with respect to certainsurnames The results of the present study indicate thatsuch trends are also observed in small isolated popula-tions Apart from the expected influences of migrationand drift there could be other related factors of popula-tion structure that could cause the deviation from neutral-
Maji Selective Neutrality of Surname Distribution in Adi Panggi 9
ity In small tribal populations the structure of thesurname distribution might vary widely depending on thehistory of settlement prevailing marriage patterns andother socio-cultural and environmental reasons Furtherthe demographic parameters can vary drastically owing totheir subsistence pattern epidemiological factors andsocial organization such as warfare history Also internaltribal factions which are formed based on kinship or clanor lineages involving several closely knit families (eg inSouth Amerindian tribes Neel 1968 Salzano et al 1967)may play a role This can significantly influence thepopulation structure and can lead to perturbations in thesurname distribution in short time periods and in suchcases the neutral theory predictions may not hold Someof the above factors are important in explaining thedeviation from neutrality in surname distribution amongthe Adi Panggi
In general the Adi tribe consists of several sub-tribes whooccupy different terrains in the upper Himalayan moun-tain ranges and some in plains in the northeastern part ofthe country (Roy 1960 Lego 2005) The formation of thesub-tribes followed the fission-fusion model of popula-tion structure and was triggered by intertribal factionscompeting for survival necessities in this hunting-gather-ing population (Neel 1968 Salzano et al 1967) Theformation of Adi Panggi sub-tribes occurred a couple ofgenerations ago and was a result of conflict between amajor subgroup of the tribe and some other close-kinfactions over hunting rights and other resources (Roy1960 Lego 2005 Lego 2006) The sub-tribes now occu-py a recognized territory and constitute a few close affinaland consanguineous lineages and their affiliated clansSince they have a patrilocal society a majority of theirmen stay in the home village whereas females come fromdifferent villages and from other Adi sub-tribes Thisresults in more female marital migration and greatersurname diversity among wives We therefore observe ahigher proportion of homonymy of surname diversityamong males For example the five most frequent sur-names represent about 65 of total husbands while raresurnames (that occur either once or twice) constituteabout 5 per cent The three most common surnamesamong wives also represent a large part of the populationbut in contrast with the males the rare surnames thatoccur either once or twice constitute 18 per cent morethan 35 times the proportion of rare surnames in hus-bands In the total sample of husbands and wives sixsurnames represent 69 percent of the total populationstudied The surname distribution therefore reflectsgreater prevalence of most frequent (occurring in 20 to 70cases) and rare surnames (occurring once or twice) asagainst the moderately frequent surnames (that occur 3 to20 times in the sample)
The estimates of unbiased Isonymy also differ betweenhusband and wives samples The combined samples(husband and wives) show higher values of estimatedIsonymy (I = 01025) suggesting higher levels of homozy-gosity as a result of more frequent marriages contracted
among a few selected surnames generally found in suchisolated tribes This is reflected in the Karlin andMacGregor distribution which shows the non-lineartrend and the trend remains the same even after thelogarithmic transformation This also explains the widedifferences between the male and female surname distri-butions and the deviations observed in the Karlin andMacGregor and logarithmic distributions Therefore ifthe truncated distribution (omitting the rare surnames) isconsidered it shows a linear trend in the combinedsample suggesting that the non-linearity is contributedby the occurrence of rare surnames in the sample Therare surnames are contributed more through femalesmigrating from other places However the discretePareto distribution shows a linear trend for both thesamples The linear trend is clearer in the female samplethan among males but the genetic implications of thePareto distributions are not apparent (Fox 1983)
The results of the distribution of surnames also showwide differences with respect to progeny size distributionin case of post-reproductive (completed fertility) andreproductive wives and their husbands The neutralallele model would suggest that the distribution shouldbe independent of the fertility and morality (Manrubia2002) but the present study indicates otherwise andthere appears to be some differential fertility for somesurnames The Adi Panggi sample shows greater progenysize (male female and total surviving children) for themost frequent and rare surnames and that is reflected inthe near U-shaped distribution especially in case of hus-band sample This suggests reproductive advantageespecially greater success for the most frequent (6-910)and rare surnames (1-2) as against the less frequentsurnames (3-6) Among the Adi Panggi the four mostfrequent surnames among husbands and wives (especiallyPanyang Paron Mongku and Taku) show higher proge-ny size among the husband sample whereas the wivessample shows a near linear trend of decreasing progenysize from the rare surnames to the most frequent ones Apaired t-test between (eight) different types of surnamesand their progeny size shows higher significance values incase of the husbands (above 45 years) sample (p =0000433) than among the wives (above 45 years) (p =0003263) The mothers and their husbands were all inpost-reproductive period and had attained their maxi-mum fertility potential and do not use family planningand therefore the possibility of other intervening factorscan be excluded It is possible that it could be also dueto selective mortality of children especially higher mor-tality in case of less frequent surnames than with moreand rare frequent ones and this needs to be verifiedHowever there are no apparent cultural active selectivemechanisms that can influence the progeny size amongsome families of the tribe A detailed study on themarriage pattern and surname or clan distribution mightreveal more details The progeny distribution amongwomen of completed fertility is expected to follow Pois-son and Negative binomial distributions (Cavalli-Sforza
10
1971 Vasulu 1980) In patrilineal societies femalesgenerally show a greater surname diversity than malesand as a result the surname dynamics differ with respectto mitochondria and Y-chromosome
In the Indian context surname analysis has been carriedout in a few populations for investigating the effect onpopulation structure of inbreeding (Bhalla 1976 Kashyap1980 Vasulu 1980) genetic kinship (Kashyap 1980Vasulu 1980) preferential mating (Vasulu 1989 Reddy1995 Maji et al 2007) and social network analysisamong castes and tribes (Rao 1995) Surname neutralityhas also been validated in a caste community of Indiansresiding in Texas (Chakraborty 1990)
The above few studies suggest that isonymy can be ofmuch value in inferring population structure in somecastes and tribes (Kashyap 1980 Maji et al 2007) Thesurname structure in India varies among different regionscastes tribes linguistic and religious groups and providesan opportunity to study microevolution and populationstructure where the assumptions of surname or isonymyanalysis are valid Its applicability in diverse populationsespecially among isolated populations is expected to yieldfurther insights of its validity and inferring populationstructure and microevolutionary history
The present study illustrates the importance of surnamein inferring the genetic structure among a small sub-tribeAdi Panggi an isolated Tibeto-Burman- speaking com-munity in Arunachal Pradesh state in the northeasternregion of India The tribe lives on shifting-hunting-and-gathering subsistence and is a patrilocal and patrilinealsociety where the marriage pattern involves movement ofwives from other villages to the husbandrsquos village whichallows more surname diversity through females As aconsequence of differential surname structure betweenhusbands and wives the surname distribution showsdeviation from the expected neutral allele model and italso exhibits a non-linear trend of progeny size distribu-tion with respect to surname occurrence in the husbandsample
The results of the present study illustrate that surnamedistribution could be of much value in deciphering thepopulation structure and microevolution of a population(Crow 1965 Lasker 1978 Yasuda et al 1974) Thenon-linear trend observed in this study appears to beunique in the Indian context especially for the smallisolated tribes
The results of the study will help in further confirming thevalidity of the findings in several other tribes in India Inthe absence of any biological or genetic data for a tribethe surnames provide insights into the population struc-ture and possible genetic implications which can help inthe design of future studies As a prelude to the moleculargenetic investigations about the past genetic history asurname analysis can provide considerable insight into
the expected genetic structure of the population It alsohelps us to frame specific hypothesis related to male andfemale migration history which we will be testing withthe molecular genetic studies The two approachestogether can help to unravel the differential paternaland maternal past genetic history of the tribe
We thank the Government of Arunachal Pradesh andall the participants for their active cooperation in ourstudy Our sincere thanks go to Mr Tozing Tagi andMr Oyam Panyang (Katan and Geku circles) for theirassistance in data collection We acknowledge thesupport and intellectual inputs about the Adi tribesfrom Mr R N Koley (ARO Directorate of Research)and Mr Kaling Borang of Pasighat We thank Ms SKrithika Senior Research Fellow Biological Anthro-pology Unit for suggestions and help with the analysisand draft finalization and the Director Indian Statisti-cal Institute Kolkata for the financial assistance for thestudy among the tribes of Arunachal Pradesh
Barrai I Barbujani G Beretta M Maestri I Russo A Formi-ca G Pinto-Cisternas (1987) Surnames in Ferrara distribu-tion isonymy and levels of inbreeding14415-423
Barrai I Formica G Barale R Beretta M (1989) Isonymy andmigration distance 53249-262
Barrai I Formica G Barale R Scapoli C Canella R BerettaM (1990) Isonymy in emigrants from Ferrara in 1981-1988Ann Hum Biol 177-18
Barrai I Scapoli C Formica G et al (1991) Isonymy inrecords of births and deaths in Ferrara8395-404
Barrai I Formica G Scapoli C et al (1992) Microevolutionin Ferrara Isonymy 1890-1990 19371-385
Barrai I Scapoli C Bereta M et al (1996) Isonymy and thegenetic structure of Switzerland I The distributions of sur-names 23431-455
Barrai I Rodriguez-Larralde A Mamolini E et al (2000)Elements of surname structure of Austria27607-622
Barrai I Rodriguez-Larralde A Manni F et al (2002) Isony-my and isolation by distance in the Netherlands 74263-283
Barrai I Rodriguez-Larralde A Manni F Ruggiero V TartariD Scapoli C (2003) Isolation by language and distance inBeligium 681-16
Bhalla V Bhatia K (1976) Isonymy in a Bhatia leut 39479-500
Bhattacharjee PN (1954) Blood group Investigations in theAbor Tribe 351-54
11
Blackburn S (2004) Memories of migration Notes on legendsand beads in Arunachal Pradesh India
252615ndash60
Boldsen J (1992) Geographical distribution of some Danishsurnames reflection of social and natural selection 24505-513
Cavelli Sfroza LL Bodmer WF (1971) W H Freeman and Co San Francisco
Chakraborty R Weiss KM Rossmann DL Norton SL (1981)Distribution of last names a stochastic model for lilkelihooddetermination in record linking In Morrill W Dyke B (ed)
Academic Press New York
Chakraborty R Schwartz RJ (1990) Selective neutrality ofsurname distribution in an immigrant Indian community ofHouston Texas 21-13
Colantonio SE Lasker GW Kaplan BA Fuster V (2003) Useof surname models in human population biology A review ofrecent developments 75785-807
Crow JF Manage AP (1965) Measurements of Inbreeding ofthe frequencies of marriages between persons of the samesurnames 12199-203
Das TC (1953) Social Organization of the tribal peoples XIV245-262
Dyke B James AV Morrill WT (1983) Estimation of randomisonymy 10295-298
Fox WR Lasker GW (1983) The distribution of surnamefrequencies 5181-87
Karlin S MacGregor J (1967) The number of mutant formsmaintained in a population
4415-438
Kashyap LK (1980) Trends of Isonymy and inbreeding amongthe Ahmadiyyas of Kashmir 1219-25
Kashyap LK Tiwari SC (1980) Kinetics of genetic kinship asinferred by isonymy among the Ahmadiyyas of Kashmir valley
52311-324
Kimura M (1983) Cambridge University Press Cambridge
Koley RN (2005) Socio-cultural aspects of Adi Panggi Direc-torate of Research Government of Arunachal Pradesh Itanagar(in press)
Krithika S Maji S Vasulu TS (2006) Genetic heterogeneityamong three Adi tribes of Arunachal Pradesh India78221-227
Lasker GW (1978) Increments through migration to the coeffi-cient of relationship between communities by isonymy
50235-240
Lego N (2005) History of the Adis of Arunachal Pradesh MrsPonung Lego Itanagar
Lego N (2006) Modern History of Arunachal Pradesh (1825 ndash2006) Mrs Ponung Lego Itanagar
Maji S Krithika S Vasulu TS (2007) Genetic kinship among anisolated Adi tribe of Arunachal Pradesh isonymy in Adi Panggi
79321-337
Manrubia SC Zanette DH (2002) At the boundary betweenbiological and cultural evolution The origin of surname distri-butions 216461-477
Neel JV Chagnon NA (1968) The demography of two tribes ofprimitive relatively unacculturated American Indians
(USA) 59680-689
Rao AR (1995) Reciprocity in marital and social networksIllustration with Indian data 67887-904
Reddy BM Malhotra KC (1995) Relevance of surname inpopulation structure A study among the Vadde a fishingcommunity of Kolleru lake in Andhra Pradesh India
6161-168
Relethford JH (1988) Estimation of Kinship and Genetic dis-tance from Surnames 60475-492
Rodriguez LA Barrai I Alfonzo JC (1993) Isonymy structureof four Venezuelan states 20131-145
Rodriguez-Larralde A Formica G Scapoli C (1993) Microevo-lution in Perugia Isonymy 1890-1990 20261-274
Rodriguez-Larralde A Gonzales-Martin A Scapoli C Barrai I(2003) The names of Spain A study of the isonymy structureof Spain 121280-292
Roy S (1960) Aspects of Padam Minyong Culture Directorateof Research Govt of Arunachal Pradesh Itanagar
Roy S (1966) Anthropometry of the Adis of the Siang districtNorth-East Frontier Agency Shillong
Salzano FM Neel JV Maybury Lewis D (1967) Further studieson Xavante Indians I Demographic data on two additionalvillages and genetic structure of the tribe19463-489
Sarma J (1960) Puberty Marriage and Childbirth among thePangi and the Minyong Abor women 5597-113
Scapoli C Rodriguez-Larralde A Beretta M (1997) Correlationbetween isonymy parameters 1217-37
Singh IP (1994) Anthropological Survey of India Oxford
University Press New Delhi
Vasulu TS (1980) Anthrop Soc
of India Mysore
Vasulu TS (1989) Genetic structure of a tribal population 1Breeding isolation among the Yanadis 4255-268
Yasuda N Cavalli-Sfroza LL Skolnir M Moroni A (1974) Theevolution of surnames an analysis of their distribution andextinction 5123-142
Zei G Guglielmino CR Siri E Moroni A Cavalli-Sforza LL(1983a) Surnames as neutral alleles Observation in Sardinia
55357-365
Zei G Matessi RG Siri E et al (1983b) Surnames in SardiniaI Fit of frequency distributions for neutral articles and geneticpopulation structure 47329-352
Maji Selective Neutrality of Surname Distribution in Adi Panggi 9
ity In small tribal populations the structure of thesurname distribution might vary widely depending on thehistory of settlement prevailing marriage patterns andother socio-cultural and environmental reasons Furtherthe demographic parameters can vary drastically owing totheir subsistence pattern epidemiological factors andsocial organization such as warfare history Also internaltribal factions which are formed based on kinship or clanor lineages involving several closely knit families (eg inSouth Amerindian tribes Neel 1968 Salzano et al 1967)may play a role This can significantly influence thepopulation structure and can lead to perturbations in thesurname distribution in short time periods and in suchcases the neutral theory predictions may not hold Someof the above factors are important in explaining thedeviation from neutrality in surname distribution amongthe Adi Panggi
In general the Adi tribe consists of several sub-tribes whooccupy different terrains in the upper Himalayan moun-tain ranges and some in plains in the northeastern part ofthe country (Roy 1960 Lego 2005) The formation of thesub-tribes followed the fission-fusion model of popula-tion structure and was triggered by intertribal factionscompeting for survival necessities in this hunting-gather-ing population (Neel 1968 Salzano et al 1967) Theformation of Adi Panggi sub-tribes occurred a couple ofgenerations ago and was a result of conflict between amajor subgroup of the tribe and some other close-kinfactions over hunting rights and other resources (Roy1960 Lego 2005 Lego 2006) The sub-tribes now occu-py a recognized territory and constitute a few close affinaland consanguineous lineages and their affiliated clansSince they have a patrilocal society a majority of theirmen stay in the home village whereas females come fromdifferent villages and from other Adi sub-tribes Thisresults in more female marital migration and greatersurname diversity among wives We therefore observe ahigher proportion of homonymy of surname diversityamong males For example the five most frequent sur-names represent about 65 of total husbands while raresurnames (that occur either once or twice) constituteabout 5 per cent The three most common surnamesamong wives also represent a large part of the populationbut in contrast with the males the rare surnames thatoccur either once or twice constitute 18 per cent morethan 35 times the proportion of rare surnames in hus-bands In the total sample of husbands and wives sixsurnames represent 69 percent of the total populationstudied The surname distribution therefore reflectsgreater prevalence of most frequent (occurring in 20 to 70cases) and rare surnames (occurring once or twice) asagainst the moderately frequent surnames (that occur 3 to20 times in the sample)
The estimates of unbiased Isonymy also differ betweenhusband and wives samples The combined samples(husband and wives) show higher values of estimatedIsonymy (I = 01025) suggesting higher levels of homozy-gosity as a result of more frequent marriages contracted
among a few selected surnames generally found in suchisolated tribes This is reflected in the Karlin andMacGregor distribution which shows the non-lineartrend and the trend remains the same even after thelogarithmic transformation This also explains the widedifferences between the male and female surname distri-butions and the deviations observed in the Karlin andMacGregor and logarithmic distributions Therefore ifthe truncated distribution (omitting the rare surnames) isconsidered it shows a linear trend in the combinedsample suggesting that the non-linearity is contributedby the occurrence of rare surnames in the sample Therare surnames are contributed more through femalesmigrating from other places However the discretePareto distribution shows a linear trend for both thesamples The linear trend is clearer in the female samplethan among males but the genetic implications of thePareto distributions are not apparent (Fox 1983)
The results of the distribution of surnames also showwide differences with respect to progeny size distributionin case of post-reproductive (completed fertility) andreproductive wives and their husbands The neutralallele model would suggest that the distribution shouldbe independent of the fertility and morality (Manrubia2002) but the present study indicates otherwise andthere appears to be some differential fertility for somesurnames The Adi Panggi sample shows greater progenysize (male female and total surviving children) for themost frequent and rare surnames and that is reflected inthe near U-shaped distribution especially in case of hus-band sample This suggests reproductive advantageespecially greater success for the most frequent (6-910)and rare surnames (1-2) as against the less frequentsurnames (3-6) Among the Adi Panggi the four mostfrequent surnames among husbands and wives (especiallyPanyang Paron Mongku and Taku) show higher proge-ny size among the husband sample whereas the wivessample shows a near linear trend of decreasing progenysize from the rare surnames to the most frequent ones Apaired t-test between (eight) different types of surnamesand their progeny size shows higher significance values incase of the husbands (above 45 years) sample (p =0000433) than among the wives (above 45 years) (p =0003263) The mothers and their husbands were all inpost-reproductive period and had attained their maxi-mum fertility potential and do not use family planningand therefore the possibility of other intervening factorscan be excluded It is possible that it could be also dueto selective mortality of children especially higher mor-tality in case of less frequent surnames than with moreand rare frequent ones and this needs to be verifiedHowever there are no apparent cultural active selectivemechanisms that can influence the progeny size amongsome families of the tribe A detailed study on themarriage pattern and surname or clan distribution mightreveal more details The progeny distribution amongwomen of completed fertility is expected to follow Pois-son and Negative binomial distributions (Cavalli-Sforza
10
1971 Vasulu 1980) In patrilineal societies femalesgenerally show a greater surname diversity than malesand as a result the surname dynamics differ with respectto mitochondria and Y-chromosome
In the Indian context surname analysis has been carriedout in a few populations for investigating the effect onpopulation structure of inbreeding (Bhalla 1976 Kashyap1980 Vasulu 1980) genetic kinship (Kashyap 1980Vasulu 1980) preferential mating (Vasulu 1989 Reddy1995 Maji et al 2007) and social network analysisamong castes and tribes (Rao 1995) Surname neutralityhas also been validated in a caste community of Indiansresiding in Texas (Chakraborty 1990)
The above few studies suggest that isonymy can be ofmuch value in inferring population structure in somecastes and tribes (Kashyap 1980 Maji et al 2007) Thesurname structure in India varies among different regionscastes tribes linguistic and religious groups and providesan opportunity to study microevolution and populationstructure where the assumptions of surname or isonymyanalysis are valid Its applicability in diverse populationsespecially among isolated populations is expected to yieldfurther insights of its validity and inferring populationstructure and microevolutionary history
The present study illustrates the importance of surnamein inferring the genetic structure among a small sub-tribeAdi Panggi an isolated Tibeto-Burman- speaking com-munity in Arunachal Pradesh state in the northeasternregion of India The tribe lives on shifting-hunting-and-gathering subsistence and is a patrilocal and patrilinealsociety where the marriage pattern involves movement ofwives from other villages to the husbandrsquos village whichallows more surname diversity through females As aconsequence of differential surname structure betweenhusbands and wives the surname distribution showsdeviation from the expected neutral allele model and italso exhibits a non-linear trend of progeny size distribu-tion with respect to surname occurrence in the husbandsample
The results of the present study illustrate that surnamedistribution could be of much value in deciphering thepopulation structure and microevolution of a population(Crow 1965 Lasker 1978 Yasuda et al 1974) Thenon-linear trend observed in this study appears to beunique in the Indian context especially for the smallisolated tribes
The results of the study will help in further confirming thevalidity of the findings in several other tribes in India Inthe absence of any biological or genetic data for a tribethe surnames provide insights into the population struc-ture and possible genetic implications which can help inthe design of future studies As a prelude to the moleculargenetic investigations about the past genetic history asurname analysis can provide considerable insight into
the expected genetic structure of the population It alsohelps us to frame specific hypothesis related to male andfemale migration history which we will be testing withthe molecular genetic studies The two approachestogether can help to unravel the differential paternaland maternal past genetic history of the tribe
We thank the Government of Arunachal Pradesh andall the participants for their active cooperation in ourstudy Our sincere thanks go to Mr Tozing Tagi andMr Oyam Panyang (Katan and Geku circles) for theirassistance in data collection We acknowledge thesupport and intellectual inputs about the Adi tribesfrom Mr R N Koley (ARO Directorate of Research)and Mr Kaling Borang of Pasighat We thank Ms SKrithika Senior Research Fellow Biological Anthro-pology Unit for suggestions and help with the analysisand draft finalization and the Director Indian Statisti-cal Institute Kolkata for the financial assistance for thestudy among the tribes of Arunachal Pradesh
Barrai I Barbujani G Beretta M Maestri I Russo A Formi-ca G Pinto-Cisternas (1987) Surnames in Ferrara distribu-tion isonymy and levels of inbreeding14415-423
Barrai I Formica G Barale R Beretta M (1989) Isonymy andmigration distance 53249-262
Barrai I Formica G Barale R Scapoli C Canella R BerettaM (1990) Isonymy in emigrants from Ferrara in 1981-1988Ann Hum Biol 177-18
Barrai I Scapoli C Formica G et al (1991) Isonymy inrecords of births and deaths in Ferrara8395-404
Barrai I Formica G Scapoli C et al (1992) Microevolutionin Ferrara Isonymy 1890-1990 19371-385
Barrai I Scapoli C Bereta M et al (1996) Isonymy and thegenetic structure of Switzerland I The distributions of sur-names 23431-455
Barrai I Rodriguez-Larralde A Mamolini E et al (2000)Elements of surname structure of Austria27607-622
Barrai I Rodriguez-Larralde A Manni F et al (2002) Isony-my and isolation by distance in the Netherlands 74263-283
Barrai I Rodriguez-Larralde A Manni F Ruggiero V TartariD Scapoli C (2003) Isolation by language and distance inBeligium 681-16
Bhalla V Bhatia K (1976) Isonymy in a Bhatia leut 39479-500
Bhattacharjee PN (1954) Blood group Investigations in theAbor Tribe 351-54
11
Blackburn S (2004) Memories of migration Notes on legendsand beads in Arunachal Pradesh India
252615ndash60
Boldsen J (1992) Geographical distribution of some Danishsurnames reflection of social and natural selection 24505-513
Cavelli Sfroza LL Bodmer WF (1971) W H Freeman and Co San Francisco
Chakraborty R Weiss KM Rossmann DL Norton SL (1981)Distribution of last names a stochastic model for lilkelihooddetermination in record linking In Morrill W Dyke B (ed)
Academic Press New York
Chakraborty R Schwartz RJ (1990) Selective neutrality ofsurname distribution in an immigrant Indian community ofHouston Texas 21-13
Colantonio SE Lasker GW Kaplan BA Fuster V (2003) Useof surname models in human population biology A review ofrecent developments 75785-807
Crow JF Manage AP (1965) Measurements of Inbreeding ofthe frequencies of marriages between persons of the samesurnames 12199-203
Das TC (1953) Social Organization of the tribal peoples XIV245-262
Dyke B James AV Morrill WT (1983) Estimation of randomisonymy 10295-298
Fox WR Lasker GW (1983) The distribution of surnamefrequencies 5181-87
Karlin S MacGregor J (1967) The number of mutant formsmaintained in a population
4415-438
Kashyap LK (1980) Trends of Isonymy and inbreeding amongthe Ahmadiyyas of Kashmir 1219-25
Kashyap LK Tiwari SC (1980) Kinetics of genetic kinship asinferred by isonymy among the Ahmadiyyas of Kashmir valley
52311-324
Kimura M (1983) Cambridge University Press Cambridge
Koley RN (2005) Socio-cultural aspects of Adi Panggi Direc-torate of Research Government of Arunachal Pradesh Itanagar(in press)
Krithika S Maji S Vasulu TS (2006) Genetic heterogeneityamong three Adi tribes of Arunachal Pradesh India78221-227
Lasker GW (1978) Increments through migration to the coeffi-cient of relationship between communities by isonymy
50235-240
Lego N (2005) History of the Adis of Arunachal Pradesh MrsPonung Lego Itanagar
Lego N (2006) Modern History of Arunachal Pradesh (1825 ndash2006) Mrs Ponung Lego Itanagar
Maji S Krithika S Vasulu TS (2007) Genetic kinship among anisolated Adi tribe of Arunachal Pradesh isonymy in Adi Panggi
79321-337
Manrubia SC Zanette DH (2002) At the boundary betweenbiological and cultural evolution The origin of surname distri-butions 216461-477
Neel JV Chagnon NA (1968) The demography of two tribes ofprimitive relatively unacculturated American Indians
(USA) 59680-689
Rao AR (1995) Reciprocity in marital and social networksIllustration with Indian data 67887-904
Reddy BM Malhotra KC (1995) Relevance of surname inpopulation structure A study among the Vadde a fishingcommunity of Kolleru lake in Andhra Pradesh India
6161-168
Relethford JH (1988) Estimation of Kinship and Genetic dis-tance from Surnames 60475-492
Rodriguez LA Barrai I Alfonzo JC (1993) Isonymy structureof four Venezuelan states 20131-145
Rodriguez-Larralde A Formica G Scapoli C (1993) Microevo-lution in Perugia Isonymy 1890-1990 20261-274
Rodriguez-Larralde A Gonzales-Martin A Scapoli C Barrai I(2003) The names of Spain A study of the isonymy structureof Spain 121280-292
Roy S (1960) Aspects of Padam Minyong Culture Directorateof Research Govt of Arunachal Pradesh Itanagar
Roy S (1966) Anthropometry of the Adis of the Siang districtNorth-East Frontier Agency Shillong
Salzano FM Neel JV Maybury Lewis D (1967) Further studieson Xavante Indians I Demographic data on two additionalvillages and genetic structure of the tribe19463-489
Sarma J (1960) Puberty Marriage and Childbirth among thePangi and the Minyong Abor women 5597-113
Scapoli C Rodriguez-Larralde A Beretta M (1997) Correlationbetween isonymy parameters 1217-37
Singh IP (1994) Anthropological Survey of India Oxford
University Press New Delhi
Vasulu TS (1980) Anthrop Soc
of India Mysore
Vasulu TS (1989) Genetic structure of a tribal population 1Breeding isolation among the Yanadis 4255-268
Yasuda N Cavalli-Sfroza LL Skolnir M Moroni A (1974) Theevolution of surnames an analysis of their distribution andextinction 5123-142
Zei G Guglielmino CR Siri E Moroni A Cavalli-Sforza LL(1983a) Surnames as neutral alleles Observation in Sardinia
55357-365
Zei G Matessi RG Siri E et al (1983b) Surnames in SardiniaI Fit of frequency distributions for neutral articles and geneticpopulation structure 47329-352
10
1971 Vasulu 1980) In patrilineal societies femalesgenerally show a greater surname diversity than malesand as a result the surname dynamics differ with respectto mitochondria and Y-chromosome
In the Indian context surname analysis has been carriedout in a few populations for investigating the effect onpopulation structure of inbreeding (Bhalla 1976 Kashyap1980 Vasulu 1980) genetic kinship (Kashyap 1980Vasulu 1980) preferential mating (Vasulu 1989 Reddy1995 Maji et al 2007) and social network analysisamong castes and tribes (Rao 1995) Surname neutralityhas also been validated in a caste community of Indiansresiding in Texas (Chakraborty 1990)
The above few studies suggest that isonymy can be ofmuch value in inferring population structure in somecastes and tribes (Kashyap 1980 Maji et al 2007) Thesurname structure in India varies among different regionscastes tribes linguistic and religious groups and providesan opportunity to study microevolution and populationstructure where the assumptions of surname or isonymyanalysis are valid Its applicability in diverse populationsespecially among isolated populations is expected to yieldfurther insights of its validity and inferring populationstructure and microevolutionary history
The present study illustrates the importance of surnamein inferring the genetic structure among a small sub-tribeAdi Panggi an isolated Tibeto-Burman- speaking com-munity in Arunachal Pradesh state in the northeasternregion of India The tribe lives on shifting-hunting-and-gathering subsistence and is a patrilocal and patrilinealsociety where the marriage pattern involves movement ofwives from other villages to the husbandrsquos village whichallows more surname diversity through females As aconsequence of differential surname structure betweenhusbands and wives the surname distribution showsdeviation from the expected neutral allele model and italso exhibits a non-linear trend of progeny size distribu-tion with respect to surname occurrence in the husbandsample
The results of the present study illustrate that surnamedistribution could be of much value in deciphering thepopulation structure and microevolution of a population(Crow 1965 Lasker 1978 Yasuda et al 1974) Thenon-linear trend observed in this study appears to beunique in the Indian context especially for the smallisolated tribes
The results of the study will help in further confirming thevalidity of the findings in several other tribes in India Inthe absence of any biological or genetic data for a tribethe surnames provide insights into the population struc-ture and possible genetic implications which can help inthe design of future studies As a prelude to the moleculargenetic investigations about the past genetic history asurname analysis can provide considerable insight into
the expected genetic structure of the population It alsohelps us to frame specific hypothesis related to male andfemale migration history which we will be testing withthe molecular genetic studies The two approachestogether can help to unravel the differential paternaland maternal past genetic history of the tribe
We thank the Government of Arunachal Pradesh andall the participants for their active cooperation in ourstudy Our sincere thanks go to Mr Tozing Tagi andMr Oyam Panyang (Katan and Geku circles) for theirassistance in data collection We acknowledge thesupport and intellectual inputs about the Adi tribesfrom Mr R N Koley (ARO Directorate of Research)and Mr Kaling Borang of Pasighat We thank Ms SKrithika Senior Research Fellow Biological Anthro-pology Unit for suggestions and help with the analysisand draft finalization and the Director Indian Statisti-cal Institute Kolkata for the financial assistance for thestudy among the tribes of Arunachal Pradesh
Barrai I Barbujani G Beretta M Maestri I Russo A Formi-ca G Pinto-Cisternas (1987) Surnames in Ferrara distribu-tion isonymy and levels of inbreeding14415-423
Barrai I Formica G Barale R Beretta M (1989) Isonymy andmigration distance 53249-262
Barrai I Formica G Barale R Scapoli C Canella R BerettaM (1990) Isonymy in emigrants from Ferrara in 1981-1988Ann Hum Biol 177-18
Barrai I Scapoli C Formica G et al (1991) Isonymy inrecords of births and deaths in Ferrara8395-404
Barrai I Formica G Scapoli C et al (1992) Microevolutionin Ferrara Isonymy 1890-1990 19371-385
Barrai I Scapoli C Bereta M et al (1996) Isonymy and thegenetic structure of Switzerland I The distributions of sur-names 23431-455
Barrai I Rodriguez-Larralde A Mamolini E et al (2000)Elements of surname structure of Austria27607-622
Barrai I Rodriguez-Larralde A Manni F et al (2002) Isony-my and isolation by distance in the Netherlands 74263-283
Barrai I Rodriguez-Larralde A Manni F Ruggiero V TartariD Scapoli C (2003) Isolation by language and distance inBeligium 681-16
Bhalla V Bhatia K (1976) Isonymy in a Bhatia leut 39479-500
Bhattacharjee PN (1954) Blood group Investigations in theAbor Tribe 351-54
11
Blackburn S (2004) Memories of migration Notes on legendsand beads in Arunachal Pradesh India
252615ndash60
Boldsen J (1992) Geographical distribution of some Danishsurnames reflection of social and natural selection 24505-513
Cavelli Sfroza LL Bodmer WF (1971) W H Freeman and Co San Francisco
Chakraborty R Weiss KM Rossmann DL Norton SL (1981)Distribution of last names a stochastic model for lilkelihooddetermination in record linking In Morrill W Dyke B (ed)
Academic Press New York
Chakraborty R Schwartz RJ (1990) Selective neutrality ofsurname distribution in an immigrant Indian community ofHouston Texas 21-13
Colantonio SE Lasker GW Kaplan BA Fuster V (2003) Useof surname models in human population biology A review ofrecent developments 75785-807
Crow JF Manage AP (1965) Measurements of Inbreeding ofthe frequencies of marriages between persons of the samesurnames 12199-203
Das TC (1953) Social Organization of the tribal peoples XIV245-262
Dyke B James AV Morrill WT (1983) Estimation of randomisonymy 10295-298
Fox WR Lasker GW (1983) The distribution of surnamefrequencies 5181-87
Karlin S MacGregor J (1967) The number of mutant formsmaintained in a population
4415-438
Kashyap LK (1980) Trends of Isonymy and inbreeding amongthe Ahmadiyyas of Kashmir 1219-25
Kashyap LK Tiwari SC (1980) Kinetics of genetic kinship asinferred by isonymy among the Ahmadiyyas of Kashmir valley
52311-324
Kimura M (1983) Cambridge University Press Cambridge
Koley RN (2005) Socio-cultural aspects of Adi Panggi Direc-torate of Research Government of Arunachal Pradesh Itanagar(in press)
Krithika S Maji S Vasulu TS (2006) Genetic heterogeneityamong three Adi tribes of Arunachal Pradesh India78221-227
Lasker GW (1978) Increments through migration to the coeffi-cient of relationship between communities by isonymy
50235-240
Lego N (2005) History of the Adis of Arunachal Pradesh MrsPonung Lego Itanagar
Lego N (2006) Modern History of Arunachal Pradesh (1825 ndash2006) Mrs Ponung Lego Itanagar
Maji S Krithika S Vasulu TS (2007) Genetic kinship among anisolated Adi tribe of Arunachal Pradesh isonymy in Adi Panggi
79321-337
Manrubia SC Zanette DH (2002) At the boundary betweenbiological and cultural evolution The origin of surname distri-butions 216461-477
Neel JV Chagnon NA (1968) The demography of two tribes ofprimitive relatively unacculturated American Indians
(USA) 59680-689
Rao AR (1995) Reciprocity in marital and social networksIllustration with Indian data 67887-904
Reddy BM Malhotra KC (1995) Relevance of surname inpopulation structure A study among the Vadde a fishingcommunity of Kolleru lake in Andhra Pradesh India
6161-168
Relethford JH (1988) Estimation of Kinship and Genetic dis-tance from Surnames 60475-492
Rodriguez LA Barrai I Alfonzo JC (1993) Isonymy structureof four Venezuelan states 20131-145
Rodriguez-Larralde A Formica G Scapoli C (1993) Microevo-lution in Perugia Isonymy 1890-1990 20261-274
Rodriguez-Larralde A Gonzales-Martin A Scapoli C Barrai I(2003) The names of Spain A study of the isonymy structureof Spain 121280-292
Roy S (1960) Aspects of Padam Minyong Culture Directorateof Research Govt of Arunachal Pradesh Itanagar
Roy S (1966) Anthropometry of the Adis of the Siang districtNorth-East Frontier Agency Shillong
Salzano FM Neel JV Maybury Lewis D (1967) Further studieson Xavante Indians I Demographic data on two additionalvillages and genetic structure of the tribe19463-489
Sarma J (1960) Puberty Marriage and Childbirth among thePangi and the Minyong Abor women 5597-113
Scapoli C Rodriguez-Larralde A Beretta M (1997) Correlationbetween isonymy parameters 1217-37
Singh IP (1994) Anthropological Survey of India Oxford
University Press New Delhi
Vasulu TS (1980) Anthrop Soc
of India Mysore
Vasulu TS (1989) Genetic structure of a tribal population 1Breeding isolation among the Yanadis 4255-268
Yasuda N Cavalli-Sfroza LL Skolnir M Moroni A (1974) Theevolution of surnames an analysis of their distribution andextinction 5123-142
Zei G Guglielmino CR Siri E Moroni A Cavalli-Sforza LL(1983a) Surnames as neutral alleles Observation in Sardinia
55357-365
Zei G Matessi RG Siri E et al (1983b) Surnames in SardiniaI Fit of frequency distributions for neutral articles and geneticpopulation structure 47329-352
11
Blackburn S (2004) Memories of migration Notes on legendsand beads in Arunachal Pradesh India
252615ndash60
Boldsen J (1992) Geographical distribution of some Danishsurnames reflection of social and natural selection 24505-513
Cavelli Sfroza LL Bodmer WF (1971) W H Freeman and Co San Francisco
Chakraborty R Weiss KM Rossmann DL Norton SL (1981)Distribution of last names a stochastic model for lilkelihooddetermination in record linking In Morrill W Dyke B (ed)
Academic Press New York
Chakraborty R Schwartz RJ (1990) Selective neutrality ofsurname distribution in an immigrant Indian community ofHouston Texas 21-13
Colantonio SE Lasker GW Kaplan BA Fuster V (2003) Useof surname models in human population biology A review ofrecent developments 75785-807
Crow JF Manage AP (1965) Measurements of Inbreeding ofthe frequencies of marriages between persons of the samesurnames 12199-203
Das TC (1953) Social Organization of the tribal peoples XIV245-262
Dyke B James AV Morrill WT (1983) Estimation of randomisonymy 10295-298
Fox WR Lasker GW (1983) The distribution of surnamefrequencies 5181-87
Karlin S MacGregor J (1967) The number of mutant formsmaintained in a population
4415-438
Kashyap LK (1980) Trends of Isonymy and inbreeding amongthe Ahmadiyyas of Kashmir 1219-25
Kashyap LK Tiwari SC (1980) Kinetics of genetic kinship asinferred by isonymy among the Ahmadiyyas of Kashmir valley
52311-324
Kimura M (1983) Cambridge University Press Cambridge
Koley RN (2005) Socio-cultural aspects of Adi Panggi Direc-torate of Research Government of Arunachal Pradesh Itanagar(in press)
Krithika S Maji S Vasulu TS (2006) Genetic heterogeneityamong three Adi tribes of Arunachal Pradesh India78221-227
Lasker GW (1978) Increments through migration to the coeffi-cient of relationship between communities by isonymy
50235-240
Lego N (2005) History of the Adis of Arunachal Pradesh MrsPonung Lego Itanagar
Lego N (2006) Modern History of Arunachal Pradesh (1825 ndash2006) Mrs Ponung Lego Itanagar
Maji S Krithika S Vasulu TS (2007) Genetic kinship among anisolated Adi tribe of Arunachal Pradesh isonymy in Adi Panggi
79321-337
Manrubia SC Zanette DH (2002) At the boundary betweenbiological and cultural evolution The origin of surname distri-butions 216461-477
Neel JV Chagnon NA (1968) The demography of two tribes ofprimitive relatively unacculturated American Indians
(USA) 59680-689
Rao AR (1995) Reciprocity in marital and social networksIllustration with Indian data 67887-904
Reddy BM Malhotra KC (1995) Relevance of surname inpopulation structure A study among the Vadde a fishingcommunity of Kolleru lake in Andhra Pradesh India
6161-168
Relethford JH (1988) Estimation of Kinship and Genetic dis-tance from Surnames 60475-492
Rodriguez LA Barrai I Alfonzo JC (1993) Isonymy structureof four Venezuelan states 20131-145
Rodriguez-Larralde A Formica G Scapoli C (1993) Microevo-lution in Perugia Isonymy 1890-1990 20261-274
Rodriguez-Larralde A Gonzales-Martin A Scapoli C Barrai I(2003) The names of Spain A study of the isonymy structureof Spain 121280-292
Roy S (1960) Aspects of Padam Minyong Culture Directorateof Research Govt of Arunachal Pradesh Itanagar
Roy S (1966) Anthropometry of the Adis of the Siang districtNorth-East Frontier Agency Shillong
Salzano FM Neel JV Maybury Lewis D (1967) Further studieson Xavante Indians I Demographic data on two additionalvillages and genetic structure of the tribe19463-489
Sarma J (1960) Puberty Marriage and Childbirth among thePangi and the Minyong Abor women 5597-113
Scapoli C Rodriguez-Larralde A Beretta M (1997) Correlationbetween isonymy parameters 1217-37
Singh IP (1994) Anthropological Survey of India Oxford
University Press New Delhi
Vasulu TS (1980) Anthrop Soc
of India Mysore
Vasulu TS (1989) Genetic structure of a tribal population 1Breeding isolation among the Yanadis 4255-268
Yasuda N Cavalli-Sfroza LL Skolnir M Moroni A (1974) Theevolution of surnames an analysis of their distribution andextinction 5123-142
Zei G Guglielmino CR Siri E Moroni A Cavalli-Sforza LL(1983a) Surnames as neutral alleles Observation in Sardinia
55357-365
Zei G Matessi RG Siri E et al (1983b) Surnames in SardiniaI Fit of frequency distributions for neutral articles and geneticpopulation structure 47329-352
