Genetic Structure of Mennonite Populations 
of Kansas and Nebraska 
M.H. CRAWFORD, 1 D.D. DYKES, 2 AND H.F. POLESKY 2 

Abstract We describe the gene frequency distributions for 29 
different blood group, serum, a n d erythrocytic proteins for three 
M e n n o n i t e c o m m u n i t i e s f r o m Kansas a n d Nebraska and c o m p a r e 
their gene frequencies with those of Amish, Hutterite, a n d M e n n o n i t e 
p o p u l a t i o n s using the topological m e t h o d of H a r p e n d i n g a n d Jenkins 
(1973). Subdivision of these c o m m u n i t i e s into congregations reveals 
that the "fission-fusion" m o d e l best characterizes the relationship 
between the genetic patterns a n d historical events. These M e n n o n i t e 
populations, although reproductively isolated at the t u r n of this 
century, are presently entering the m a i n s t r e a m of US rural culture. 

D u r i n g the 1950s a n d 1960s geneticists, anthropologists, a n d physicians 
o f t e n focused o n the genetics of h u m a n isolates. Small, geographically or 
socially defined p o p u l a t i o n s t h a t were reproductively isolated a n d highly 
inbred were being studied t h r o u g h o u t the world ( G o l d s c h m i d t 1963). 
T h e p r i m a r y p u r p o s e s of those investigations were to d o c u m e n t t h e ef-
fect of genetic d r i f t a n d to u n d e r s t a n d t h e m o d e of t r a n s m i s s i o n of r a r e 
genetic diseases. T h i s interest gave i m p e t u s to the study of such h u m a n 
p o p u l a t i o n s as t h e H a b b a n i t e Jews, t h e Samaritans, Tristan da C u n h a , 
P a r m a Valley, a n d St. Bartholemew. D u r i n g this t i m e the genetics of 
various A n a b a p t i s t groups, such as t h e Amish, Dunkers, a n d Hutterites, 
was investigated. T h e s e early genetic isolate studies yielded a b o u n t y 
of i n f o r m a t i o n o n t h e genetics of rare m u t a n t genes, for example, t h e 
Ellis-van Creveld s y n d r o m e , limb-girdle m u s c u l a r dystrophy, h e m o p h i l i a 
B, a n d congenital d e a f n e s s (McKusick et al. 1964). T h e d o c u m e n t a t i o n 
of genetic drift was less d r a m a t i c , even t h o u g h D . F . R o b e r t s ' s study 
of Tristan da C u n h a succeeded in d e m o n s t r a t i n g t h e effects of u n i q u e 

1 Department of Anthropology, University of Kansas, Lawrence, Kansas 66045. 
2 War Memorial Blood Bank, Minneapolis, Minnesota 55455. 
Human Biology, August 1989, Vol. 61, No. 4, pp. 493-514 
© Wayne State University Press, 1989 
KEY WORDS: MENNONITES, POPULATION GENETICS, BLOOD GROUPS 



4 9 4 / CRAWFORD ET AL. 

historical events and stochastic processes on small p o p u l a t i o n s (Roberts 
1968). 

Although the Amish and Hutterites were intensively studied, the 
M e n n o n i t e s were by a n d large ignored by the early geneticists and anthro-
pologists. W i t h the exception of Allen and R e d e k o p ' s (1967) prospectus 
for f u t u r e research a m o n g the O l d Colony M e n n o n i t e s of Mexico and a 
publication of gene frequencies for 51 M e n n o n i t e s encountered by Brown 
et al. (1974) in the G r a n Chaco region of Paraguay, little is known about 
the genetics of the Mennonites. 

T h e genetic structure of the M e n n o n i t e p o p u l a t i o n s of Kansas 
a n d Nebraska is one p o r t i o n of a long-term research program initiated 
by the University of Kansas during the mid-1970s on the genetics 
of differential biological aging. To date, these p o p u l a t i o n s have been 
described demographically (Stevenson et al. 1989; Lin a n d Crawford 
1983), anthropometrically (Devor et al. 1986a,b; Sirijaraya a n d Crawford 
1989), physiologically (Devor a n d Crawford 1984a,b; Koertvelyessy et al. 
1982), and genetically (Crawford a n d Rogers 1982; Rogers 1987). 

T h e primary purposes of this article are (1) t o describe the gene 
frequency distributions for various blood m a r k e r s in t h e Mennonite 
populations of Kansas and Nebraska, (2) t o e x a m i n e the genetic structure 
of these populations and its relationship to the e t h n o h i s t o r y of the 
Anabaptists, and (3) to d e t e r m i n e how systematic pressures, such as 
migration, affect the genetic structure of the M e n n o n i t e s . 

Populations 
T h e M e n n o n i t e c o m m u n i t i e s of K a n s a s a n d Nebraska trace their 

origins to sixteenth-century Europe. These diverse groups, constituting 
the Anabaptist or left wing of the Protestant R e f o r m a t i o n , hold in 
c o m m o n beliefs in the separation of church a n d state, adult baptism, 
and pacifism. Different Anabaptist d e n o m i n a t i o n s c a m e to b e identified 
in Europe by their p r o m i n e n t local leaders. For example, t h e followers 
of Jacob Hutter are known as Hutterites, the followers of Jacob A m a n n 
became the Amish, a n d the followers of M e n n o n S i m o n s call themselves 
the Mennonites. 

Persecution of the Anabaptists in Western Europe, particularly in 
the Netherlands and Austria, forced t h e m to settle in t h e underdeveloped 
agricultural regions of Eastern Europe. In 1669 fifteen M e n n o n i t e refugee 
couples settled in the swampy lowlands south of Danzig in West Prus-
sia and thus founded the Przechowka Church. T h e congregation increased 



Genetic Structure of Mennonites / 495 

in n u m b e r a n d m a i n t a i n e d m e t i c u l o u s records. In 1821 all but seven 
families of this congregation emigrated to Russia a n d settled in the 
U k r a i n e near t h e Molotschna River. T h i s congregation a d o p t e d the n a m e 
Alexanderwohl, in h o n o r of the R u s s i a n czar. 

E c o n o m i c conditions, changes in Russian governmental policies, 
a n d possibly internal strife i n d u c e d the Alexanderwohl M e n n o n i t e s t o 
emigrate to the U n i t e d States in 1874. Virtually the entire congregation 
emigrated, followed by m a n y of their relatives and descendants, w h o 
continued the e x o d u s until the Soviet Revolution e n d e d m o s t of this 
flow. O n their arrival in the U n i t e d States, c o m p e t i t i o n between railroad 
agents for l a n d sales a n d additional strife within the group resulted in t h e 
original Alexanderwohl c o m m u n i t y splitting into three m a j o r divisions. 
O n e group settled west of Lincoln, Nebraska, in t o d a y ' s H e n d e r s o n . T h e 
other two groups settled in Kansas, 40 miles n o r t h of Wichita. 

As these congregations prospered a n d grew in size, offshoots or 
subdivisions f o r m e d . Today there are ten congregations derived f r o m or 
related to the original Przechowka Church. In 1874 the K a n s a s g r o u p 
was subdivided into two c o m m u n i t i e s , HofFnungsau a n d Alexander-
wohl. Shortly after the t u r n of this century (1909), T a b o r split off the 
Alexanderwohl congregation. In 1920 the Goessel church separated f r o m 
the Alexanderwohl congregation. In a d d i t i o n , t h e Nebraska M e n n o n -
ite Bethesda congregation also underwent a fission f r o m t h e Evangelical 
M e n n o n i t e s . 

For p u r p o s e s of c o m p a r i s o n , we include a m o r e conservative M e n -
nonite group f r o m M e r i d i a n , Kansas, in this study. T h i s c o m m u n i t y , 
k n o w n locally as the H o l d e m a n s of the Church of G o d in Christ M e n n o , 
is a heterogeneous p o p u l a t i o n f o u n d e d by J o h n H o l d e m a n in 1859 
in Ohio. T h e m e m b e r s h i p is largely Pennsylvania G e r m a n a n d D u t c h 
(Ostroger M e n n o n i t e s w h o migrated in 1875 f r o m Volhynia in central 
Poland) mixed with the Kleine G e m e i n d e M e n n o n i t e s , w h o migrated to 
M a n i t o b a f r o m southern Russia in 1874. 

Methods 
We collected 1251 b l o o d specimens f r o m three c o m m u n i t i e s of 

K a n s a s a n d Nebraska (Table 1), packed them in ice, a n d shipped t h e m by 
air to the M i n n e a p o l i s War M e m o r i a l Blood Bank for analysis. Although 
the sample size for M e r i d i a n appears to be small, it represents 54% of 
the total c o m m u n i t y a n d almost the entire adult population. T h e Goessel 
sample represents 47% of the entire Alexanderwohl church m e m b e r s h i p . 
T h e H e n d e r s o n sample includes m o r e than 50% of the town residents. 



4 9 6 / CRAWFORD ET AL. 

Table 1. Size of Sample Used for Blood Analyses 
Population Sample Size (N) 
Goessel 
Meridian 
Henderson 
Total 

616 
87 

549 
1252 

We performed red cell typings for the following systems: ABO (anti-
A, Ai, B), Rhesus (anti-C, c, D, E, e), M N S (anti-M, N , S, s), Kell (anti-
K, k), ( K p a , K p b ) , Kidd (anti-Jk a , J k b ) , Duffy ( a n t i - F y a , F y b ) , P (anti-
Pi), Vel (anti-V+), Lutheran (anti-Lu a , L u b ) , a n d several low-frequency 
antigens, including M t a , F r 3 , and B u a . We p e r f o r m e d the red cell typings 
on microtiter (U) plates using 2% suspensions of washed erythrocytes 
and a modification of the m e t h o d of Crawford et al. (1970). We diluted 
antisera to obtain optimal reactivity a n d incubated typings for 1 hr at 
r o o m temperature (20° C). Before reading, we spun the typing plates for 
2 min at 500 rpm. We also carried out tests t h a t required an antiglobulin 
phase. For these tests we used microtiter plates, washing t h e m three times 
with saline and rinsing in a C o o m b s rinser; we spun the plates before 
recording the results. 

We phenotyped serum proteins either electrophoretically or through 
isoelectric focusing (IEF). Transferrin (Tf), haptoglobin (Hp), a n d ceru-
loplasmin (Cp) phenotypes were identified f r o m acrylamide slabs with 
a m i d o black stain (Polesky et al. 1975). T h e group-specific component 
(Gc) and the properdin factor (Bf) systems were simultaneously pheno-
typed using the agarose electrophoretic m e t h o d described by Dykes and 
Polesky (1980). We employed the isoelectric focusing m e t h o d of Dykes 
et al. (1981) for phenotyping Gel. 

We obtained stroma-free hemolysates for d e t e r m i n i n g erythrocytic 
enzyme phenotypes by washing cells three times in saline, diluting 1:1 in 
distilled water, and centrifuging at high speed. Specimens were stored at 
- 7 0 ° C until tested. 

We phenotyped eight red cell protein markers. Adenylate ki-
nase (AK), 6-phosphogluconate dehydrogenase (6-PGD), a n d acid phos-
phatase (AcP) were simultaneously electrophoresed on a single horizontal 
starch gel (Dykes a n d Polesky 1976). T h e esterase D (EsD), isocitrate de-
hydrogenase (ICD), and malate dehydrogenase ( M D H ) phenotypes were 
identified on a starch gel using a citrate-phosphate buffer system, p H = 
5.9, of Karp and Sutton (1967). Locus 1 of the enzyme phosphogluco-
mutase (PGM) was phenotyped using the original technique of Spencer 



Genetic Structure of Mennonites / 497 

et al. (1964). In a d d i t i o n , glyoxalase (GLo) was phenotyped using the 
m e t h o d s of H a r r i s a n d H o p k i n s o n (1976). 

Allelic frequencies for the b l o o d group systems are m a x i m u m 
likelihood estimates c o m p u t e d by a modified MAXLIK program of Reed 
a n d Schull (1968). T h e s e gene frequencies are not corrected for the 
biological relationships t h a t are observed a m o n g the individuals included 
in t h e sample. 

Analytical Methods 
P o p u l a t i o n structure of the Anabaptist groups is represented by 

the m e t h o d described by H a r p e n d i n g a n d Jenkins (1973). Sample allelic 
frequencies are converted to a relationship matrix R of dimension (L x 
L), where L is the n u m b e r of sample groups. T h e ijlh element of R is 

where k is t h e n u m b e r of alleles. T h e diagonal elements of /?, rir describe 
the overall d e v i a t i o n of the allelic frequencies of the array ( H a r p e n d i n g 
a n d J e n k i n s 1973; W o r k m a n et al. 1976). T h e weighted mean of the 
diagonal of t h e relationship matrix (Rst), that is, the mean genetic 
heterogeneity of all p o p u l a t i o n s , is equivalent to Wright's FSt-

Relative genetic relationships a m o n g the Anabaptist groups are 
graphically represented by a least-squares approximation of the R matrix. 
Reduced-space eigenvectorial representations provide two-dimensional 
"genetic m a p s " of allelic frequency distributions. T h e eigenvector axes 
of the m a p s are scaled by the root of the corresponding eigenvalues and 
thus equalize t h e scale of projection (Lalouel 1973). 

T h e relative c o n t r i b u t i o n s of systematic versus nonsystematic pres-
sures on the microdifferentiation of subdivided populations are explored 
through the m e t h o d of H a r p e n d i n g and Ward (1982). T h i s m e t h o d is 
based on the a s s u m p t i o n that u n d e r u n i f o r m systematic pressure het-
erozygosity should decrease in p r o p o r t i o n to the increasing genetic dis-
tance f r o m t h e centroid of the array. Thus a u n i f o r m negative slope 
should result f r o m the regression of heterozygosity on a relative ge-
netic distance. T h i s relationship is represented by a two-dimensional plot 
whose o r d i n a t e is genetic distance ( r / 7 ) and whose abscissa is the mean 
per locus heterozygosity (HO)' 

(PL - PL)(PJ - P L ) 
Pl{\-PL) (1) 

(2) 



4 9 8 / CRAWFORD ET AL. 

where PT is the frequency of the zth allele and / is the n u m b e r of loci. 
To c o m p a r e the genetic relationship of the various Mennonite 

congregations with a historically derived d e n d o g r a m , we p e r f o r m e d 
a cluster analysis on the gene frequency data. We used a B M D P 2 M 
c o m p u t e r program (Dixon 1985), which is based on the Euclidean 
distance between two p o p u l a t i o n s (j and k) a n d is defined as 

/ 

where xtj is the value of the ith variable in the jxh p o p u l a t i o n . T h e 
algorithm employed by this cluster program uses the distance between 
centroid clusters as a criterion for a m a l g a m a t i n g clusters. Eight blood 
group systems and 25 alleles were utilized in this cluster analysis. 

Results 
Tables 2 and 3 s u m m a r i z e the phenotypic counts a n d gene frequen-

cies for blood groups, s e r u m proteins, and erythrocytic proteins in three 
M e n n o n i t e communities. 

Of the 31 b l o o d systems tested by the M i n n e a p o l i s Blood Bank (29 
systems reported in this publication), 19 loci exhibit variant f o r m s at 
an incidence of 1% or m o r e . We call these " p o l y m o r p h i c " (Table 4). We 
tested a n u m b e r of rare familial variants, such as Froese (Fr 2 ), Scianna 
2 (Bu a ), Lw, Gregory, Wright, and Miltenberger, for variation in these 
M e n n o n i t e populations. Although these antigens are rare in the general 
U S population, the presence of some of t h e m , namely F r 2 a n d B u a , has 
been reported in M e n n o n i t e groups (Lewis et al. 1978). We did not detect 
any F r 2 * individuals in this study, although 3 out of 547 individuals of 
O b l o o d group were B u a positive. Although the L u t h e r a n b l o o d group 
system is usually p o l y m o r p h i c in northern Europe, L u a ranges f r o m 1% 
to 6%; it is m o n o m o r p h i c in the three M e n n o n i t e c o m m u n i t i e s . 

O n the basis of the b l o o d group frequencies, the Goessel a n d Hen-
derson populations show genetic similarity a n d b o t h groups differ slightly 
f r o m the Meridian population. These gene frequency patterns reflect the 
ethnohistory of these communities, with Goessel and H e n d e r s o n hav-
ing separated reproductively in 1874. The Meridian M e n n o n i t e s are a 
recent mixture of Pennsylvania G e r m a n s , Swiss, and Dutch. Because of 
the small size of the founding population, the gene frequencies consis-
tently fall outside the ranges observed in Germany, Switzerland, and the 
Netherlands. For example, in the M N S system b o t h Goessel a n d Merid-
ian Mennonites differ f r o m the gene frequencies observed in Western 



Genetic Structure of Mennonites / 499 
Table 2. Phenotypes and Gene Frequencies for Blood Group Antigens 

System 
and Goessel Meridian Henderson 

Phenotypes Counts Gene Frequency Counts Gene Frequency Counts Gene Frequency 
ABO 

A, 201 A, = 0.207 25 A! = 0.170 195 A, = 0 . 2 1 5 
A 2 74 A 2 = 0.089 3 A 2 = 0.022 46 A 2 = 0.064 B 83 B = 0.104 9 B = 0.061 78 B = 0.096 
A , B 32 O = 0.600 1 O = 0.747 14 O = 0.625 
A 2 B 7 0 8 
0 219 48 208 

MNSs 
MS 21 8 29 
MSs 75 11 82 
MSs 63 MS = 0.209 3 MS = 0.351 37 MS = 0.236 
MNS 15 Ms = 0.316 1 Ms = 0.23 19 Ms = 0.270 
MNSs 139 N S = 0.031 34 NS = 0.032 119 NS = 0.068 
MNs 175 N s = 0.444 19 Ns = 0.404 120 Ns = 0.426 
NS 0 1 3 
NSs 10 1 33 
Ns 116 8 104 

Rhesus 
CDe 98 15 103 
CcDEe 78 CDe = 0.403 14 CDe = 0.372 69 CDe = 0.428 
CcDe 223 cDE = 0.151 20 cDE = 0.238 205 cDE = 0.145 
cDE 16 cDe = 0.019 0 cDe = 0.000 10 cDe = 0.054 
cDEe 79 Cde = 0.002 5 Cde = 0.000 68 Cde = 0.012 
cDe 11 cdE = 0.002 17 cdE = 0.000 21 cdE = 0.002 
Cede 1 cde = 0.423 0 cde = 0.390 4 cde = 0.359 
cdEe 1 0 1 
cde 109 0 66 

Duffy 
Fy a + b - 115 Fy a = 0.456 23 Fy a = 0.500 131 Fy a = 0.499 
a + b + 311 F y b = 0.544 39 F y b = 0.500 284 F y b = 0.501 
a - b + 150 23 135 

Kidd 
J k a + b - 166 Jk a = 0.521 20 Jk a = 0.477 65 Jka = 0.433 
a + b + 276 J k b = 0.479 42 J k b = 0.523 337 Jk b = 0.567 
a - b + 134 24 147 

p i 
+ 460 P, = 0.495 70 Pj = 0.555 439 P! = 0.554 
- 156 P 2 = 0.505 16 P 2 = 0.445 110 P 2 = 0.446 

Kell 
KK 4 K = 0.066 1 K = 0.076 4 K = 0.093 
Kk 71 k = 0.934 11 k = 0.924 94 k = 0.907 
kk 525 74 449 
K p a + b - 2 K p a = 0.042 0 K p a = 0.020 0 K p a = 0.065 
a + b + 11 K p b = 0.958 2 K p b = 0.980 27 K p b = 0.935 
a - b + 197 47 197 



5 0 0 / CRAWFORD ET AL. 

(Table 2 cont.) 
System 

and Goessel Meridian Henderson 
Phenotypes Counts Gene Frequency Counts Gene Frequency Counts Gene Frequency 
Lewis 

Le a + b - 73 L e ( a + b - ) = 0.126 11 L e ( a + b - ) = 0.129 102 L e ( a + b - ) = 0.187 
a + b + 0 L e ( a - b + ) = 0.766 0 L e ( a - b + ) = 0.742 0 L e ( a - b + ) = 0.754 
a - b + 446 L e ( a - b - ) = 0.108 63 L e ( a - b - ) = 0.129 412 L e ( a - b - ) = 0.059 
a - b - 63 11 32 

Europe, whereas Henderson M e n n o n i t e s c o n f o r m m o r e closely to the ob-
served European pattern ( M o u r a n t et al. 1976). These d a t a suggest that 
the founding Mennonites do not represent a r a n d o m sample of either 
G e r m a n y or the Netherlands but constitute a u n i q u e genetic amalgam. 

There is little comparative i n f o r m a t i o n on the b l o o d genetics of 
Mennonite populations. Brown et al. (1974) described t h e blood group 
frequencies of 51 M e n n o n i t e settlers w h o were encountered by the 
researchers in the G r a n Chaco region of Paraguay. T h i s C h a c o colony 
emigrated f r o m Canada in 1926. T h e r e is considerable genetic difference 
between this small Paraguayan sample a n d the M e n n o n i t e s of Kansas and 
Nebraska. Specifically, the Chaco M e n n o n i t e s exhibit high frequencies of 
O blood group (79%), M s (42%), and C D e (59%) a n d low frequencies of B 
(0), MS (16%), and cde (23%). T h e differences between these M e n n o n i t e 
groups are probably a result of a c o m b i n a t i o n of factors, namely, the 
genetic composition of t h e f o u n d e r s a n d the small size of the sample 
f r o m Paraguay. 

Most of the serum protein gene frequencies observed in M e n n o n i t e 
populations fall within or j u s t outside t h e published E u r o p e a n gene 
frequency ranges. For example, in Europe a n p o p u l a t i o n s the group-
specific c o m p o n e n t (also t e r m e d the v i t a m i n D b i n d i n g c o m p o n e n t or 
DBP) varies for the G c 2 allele between 20% a n d 30% (Constans et al. 
1985). T h e frequency of this allele in the three M e n n o n i t e c o m m u n i t i e s 
is slightly elevated and beyond the published E u r o p e a n ranges. Although 
the G c i s allele has a frequency of 55-60% in Europe, t h e M e n n o n i t e 
populations range between 54% a n d 57%. A sample of M e n n o n i t e s 
(combined Goessel a n d Meridian c o m m u n i t i e s ) has been described for 
Gc a n d P G M isoelectric focusing subtypes (Dykes et al. 1983). As 
expected, these reported frequencies are intermediate between values for 
each of the two communities. 

One interesting finding in one of the M e n n o n i t e c o m m u n i t i e s is 
the presence of the G c 1 A 1 gene. T h i s gene is rarely observed in European 
populations unless there is a history of African or Australian aborigine 
admixture. Although G c 1 A 1 was first detected in Australian aborigines 
and was initially n a m e d G c A b , its highest incidence is in central Africa 



Genetic Structure of Mennonites / 501 

Table 3. Phenotypes and Gene Frequencies for Serum and Red Blood Cell 
Proteins 

Systems 
and Goessel Meridian Henderson 

Phenotypes Counts Gene Frequency Counts Gene Frequency Counts Gene Frequency 
Haptoglobin 

Hp 1-1 80 Hp 1 = 0 . 3 5 3 3 Hp 1 = 0.235 58 Hp 1 = 0.345 
2-1 278 H p 2 = 0.642 34 H p 2 = 0.765 260 H p 2 = 0.650 
2 - 2 259 H p C a r l = 0.005 48 H p C a r , = 0.000 224 H p C a r l = 0.005 
2-Carl 6 0 4 

Ceruloplasmin 
CpAB 4 C p A = 0.003 0 C p A = 0 001 10 C p A = 0.009 

BB 585 C p B = 0.997 86 C p B = 1.000 536 C p B = 0.990 
BC 0 C p c = 0.000 0 C p c = 0.000 1 C p c = 0.001 

Bf 
F 22 6 36 
FS 153 F = 0.167 22 F = 0.224 188 F = 0.247 
S 391 S = 0.813 40 S = 0.724 274 S = 0.705 
FF1 1 SO.7 = 0.006 0 SO.7 = 0.019 4 SO.7 = 0.024 
FSO.7 2 F, = 0.014 0 F, = 0.033 7 Fj = 0.024 
F,S 16 5 21 
SSO.7 6 3 16 
SO.7 0 0 1 
S0.7F1 0 0 1 

Gc 
1-1 284 G c 1 = 0.678 38 G c 1 = 0.698 232 Gc 1 = 0.665 
2 - 1 245 G c 2 = 0.322 32 G c 2 = 0.302 265 G c 2 = 0.334 
2 - 2 70 G c A b = 0.000 6 G c A b = 0.00 50 G c A b = 0.001 
2 - A B 0 0 1 

Gc: (isoelectric focus) 
IS 176 28 40 
IS 1F 67 IS = 0.565 9 IS = 0.572 196 IS = 0.535 
1S2 177 IF = 0.116 30 IF = 0.121 211 IF = 0.130 
IF 12 2 = 0.319 3 2 = 0.307 4 2 = 0.334 
IF2 43 1A1 = 0 . 0 0 0 5 1A1 = 0 . 0 0 0 47 1A1 = 0 . 0 0 1 
2 58 8 49 
l A l ( A b ) 0 0 1 

ADA 
1-1 570 A D A 1 = 0.968 77 A D A 1 = 0.895 498 A D A 1 = 0.958 
2 - 1 39 A D A 2 = 0.032 9 A D A 2 = 0.105 44 A D A 2 0.042 
2 - 2 0 0 1 

Adenylate Kinase 
A K 1 = 0 . 9 7 1 AK1-1 557 A K 1 = 0.954 86 A K 1 = 1.000 512 A K 1 = 0 . 9 7 1 

2 - 2 57 A K 2 = 0.046 0 A K 2 = 0.000 32 A K 2 = 0.029 



5 0 2 / CRAWFORD ET AL. 

(Table 3 cont.) 
Systems 

and 
Phenotypes 

Goessel Meridian Henderson 
Counts Gene Frequency Counts Gene Frequency Counts Gene Frequency 

6-PGD 
AA 605 P d A = 0.993 86 P d A = 1.000 539 P d A = 0.995 
AC 9 P d c = 0.007 0 P d c = 0.000 1 P d B = 0.001 
AB 0 0 5 P d c = 0.004 

AcP 
A 94 14 40 
AB 254 a c P A = 0.377 40 a c P A = 0.401 214 A = 0.291 
B 214 a c P B = 0.594 31 a c P B = 0.593 213 B = 0.639 
BC 25 a c P 0 = 0.029 0 a c P 0 = 0.006 53 C = 0.070 
AC 7 1 21 
C 1 0 1 

Esterase D 
1-1 509 EsD 1 = 0.911 58 E s D 1 = 0.808 433 EsD 1 = 0 . 9 0 1 
2 - 1 93 E s D 2 = 0.089 23 E s D 2 = 0.192 96 E s D 2 = 0.099 
2 - 2 8 5 5 

PGM j 
1-1 428 PGM j 1 = 0.848 61 P G M , 1 = 0.843 P G M 1 = 0.815 
2 - 1 156 P G M , 2 = 0.152 23 P G M , 2 = 0.157 P G M 2 = 0.184 
2 - 2 13 P G M r = 0.000 2 P G M r = 0.000 P G M r = 0.001 
1-r 0 1 

PGMj 
1 + 1 - 315 1 + = 0.764 64 l + = 0.807 246 1+ = 0.704 
1 - 76 1 - = 0.085 3 1" = 0.026 85 1~ = 0.112 
1+2+ 1 2 + = 0.141 0 2 + = 0.146 5 2+ = 0.173 
1 + 2 - 136 2~ = 0.010 21 2" = 0.021 153 2" = 0.010 
1 - 2 + 1 IT = 0.001 3 8 lr = 0.001 
1 - 2 - 11 2 14 
2 + 1 0 1 
2 + 2 - 9 2 13 
lr 1 1 1 

GLO 
1 - 1 25 GLO 1 = 0.456 N T 21 G L O 1 = 0.353 
2-1 80 G L O 2 = 0.564 64 G L O 2 = 0.647 
2 - 2 44 65 

ICD 
1-1 610 ICD 1 = 1.000 N T 527 I C D 1 = 0.998 2-1 0 I C D 2 = 0.000 2 I C D 2 = 0.002 

M D H 
1-1 610 M D H 1 = 1.000 N T 529 M D H 1 = 1.000 



Genetic Structure of Mennonites / 503 

Table 4. Summary of Polymorphic and Monomorphic Loci among at Least One 
of Three Populations Reported in This Study 
Polymorphic Loci Monomorphic Loci 
ABO Lutheran 
Rhesus Ceruloplasmin 
MNS 6-Phosphogluconate dehydrogenase 
Kidd Isocitrate dehydrogenase 
Duffy Malate dehydrogenase 
P Gregory 
Kell Froese (Fr3) 
Lewis Scianna 2(Bu a ) 
Gm* Lw 
Glyoxalase I Vel 
Km* Wright 
Haptoglobin Miltenberger 
Properdin Factor 
Group specific component 
Adenosine deaminase 
Adenylate kinase 
Acid phosphatase 
Esterase D 
Phosphoglucomutase 
* Reported in another publication. 

( C o n s t a n s et al. 1985). Considering t h e presence of an African marker 
PGM'j gene in t h e s a m e person with the G c 1 A 1 , it is highly likely that 
b o t h these genes were introduced through American black gene flow into 
the p o p u l a t i o n . 

T h e f r e q u e n c y of the H p 1 allele in the three M e n n o n i t e c o m m u n i -
ties is lower t h a n its incidence in Germany, Switzerland, or the Nether-
lands. In particular, H p 1 occurs in Meridian at 23.5% versus the 36-40% 
in those regions of E u r o p e f r o m which the Mennonites originated. 

A rare haptoglobin variant, Carlberg, was detected in two of the 
M e n n o n i t e c o m m u n i t i e s . H p Carlberg ( H p Ca) was originally described 
by Galatius-Jensen (1958). T h i s phenotype resembles a mixture of H p 2 -
2 a n d H p 2 - 1 in variable proportions, p r o m p t i n g Sutton (1965) to suggest 
that this variant may be the result of genetic mosaicism. More recently, 
the H p Ca p h e n o t y p e has been explained on the basis of a possible 
m u t a t i o n of t h e H p 1 a chain followed by a decrease in the synthesis of 
the p o l y p e p t i d e (Bowman a n d Kurosky 1982). 



5 0 4 / CRAWFORD ET AL. 

S - l e u t 
n ( 5 ) L - l e u t I n d i a n a P a r a g u a y ^ ^ ^ 

© ® ( z ) 
M e r i d i a n 

West E u r o p e 

( 2 ) ® © G o e s s e l 
© 

H e n d e r s o n 

Figure 1. Least-squares reduction genetic map of the eight Anabaptist populations based 
on allelic frequencies for ten alleles and three loci. Three Kansas and Nebraska 
Mennonite communities are represented by Meridian, Goessel, and Hender-
son. Gene frequencies for the other Anabaptist groups were obtained from 
Juberg et al. (1971), Brown et al. (1974), and Steinberg et al. (1967). 

Population Structure 
Figure 1 shows the population structure in the reduced-space ge-

netic m a p of the three Kansas and Nebraska M e n n o n i t e c o m m u n i t i e s 
compared to other Anabaptist groups a n d Western Europe (represented 
by mean gene frequencies f r o m the regions of origin of t h e M e n n o n -
ites). This plot is based on ten alleles and three loci. T h e n u m b e r of 
loci used in this analysis was necessitated by the availability of data on 
other Anabaptist groups. Samples are arrayed along the first two scaled 
eigenvectors, which together account for 71% of the total variation in 
the sample (eigenvector 1, 51%; eigenvector 2, 20%). T h e addition of the 
third eigenvector in a pseudo-three-dimensional representation accounts 
for 83% of the total variation. T h e genetic m a p in Figure 1 indicates 
little genetic divergence a m o n g the various Anabaptist groups, with the 
cluster populations in close proximity to the centroid of the distribution. 
Of the three Mennonite c o m m u n i t i e s studied in Kansas a n d Nebraska, 
the Meridian population differs f r o m the other two groups. T h i s genetic 
difference is a result of their ethnohistory and genetic f o u n d a t i o n , with 
Goessel and Henderson being a single c o m m u n i t y until 1874. Meridian 
had few founders. Goessel and H e n d e r s o n individuals, w h o are General 
Conference Mennonites (the most m a i n s t r e a m of the Mennonites), have 
the closest genetic affinity to the composite Western E u r o p e popul a t i on. 
The Paraguay Mennonites differ f r o m the other groups probably because 
of their small sample size (N = 51) and may represent a fission of the 
larger Canadian gene pool along familial lines. 



Genetic Structure of Mennonites / 505 

Figure 2. Plot of mean per locus heterozygosity H ( / / 0 is used interchangeably) against 
distance from the centroid rH of the relationship matrix for eight Anabaptist 
communities. The R matrix is based on ten blood group alleles and three loci. 

Figure 2 indicates t h e relationship between mean per locus het-
erozygosity ( / / 0 ) a n d the relative distance f r o m the centroid of the allelic 
d i s t r i b u t i o n ( r / z ) for the seven Anabaptist groups and a composite West-
ern E u r o p e g r o u p . T h i s c o m p a r i s o n reflects the ethnohistory and origins 
of the A n a b a p t i s t groups. T h e Paraguayan Mennonites exhibit low het-
erozygosity a n d high rih suggesting that this population is experiencing 
the action of n o n s y s t e m a t i c pressures because of its geographic repro-
ductive isolation a n d small size. By contrast, the Indiana Amish have 
high heterozygosity a n d high ru and, like the Paraguay Mennonites, are 
located far f r o m expectation, as d e m o n s t r a t e d by their distance f r o m the 
theoretical regression line. T h i s unusually high level of ru and H0 a m o n g 
the A m i s h is probably t h e result of a highly heterogeneous f o u n d i n g pop-
ulation acted on by n o n s y s t e m a t i c pressures. However, the Goessel and 
H e n d e r s o n p o p u l a t i o n s exhibit relatively high levels of heterozygosity 
and low rih suggesting elevated migration rates and exogamy. 



5 0 6 / CRAWFORD ET AL. 

Wright's FST statistics have been widely used to measure the ge-
netic microdifferentiation of subdivided populations. In c o m p a r i s o n with 
other studies of FSt, the Anabaptist p o p u l a t i o n s exhibit a low FST level 
of 0.002. T h i s value indicates t h a t these Anabaptist groups can be consid-
ered genetically homogeneous (based on 10 alleles) and have experienced 
little genetic differentiation. In contrast, subdivided c i r c u m p o l a r popu-
lations attain FST values of 0.12, a n d the m a j o r geographic entities of 
Homo sapiens exhibit values u p to 0.15 (Crawford and Enciso 1982). 

Figure 3 contains a least-squares reduction genetic m a p of the 
three M e n n o n i t e c o m m u n i t i e s subdivided by congregations. We reduced 
44 alleles a n d 15 genetic loci into 2 eigenvectors. T h e first two scaled 
eigenvectors account for almost 62% of the gene frequency variance in 
the array. T h e first eigenvector (e\) has almost twice the discriminating 
power and accounts for 41.5% of the variance, versus 21.5% for e2. The 
first scaled eigenvector separates K a n s a n f r o m Nebraskan Mennonites. 
T h e Goessel c o m m u n i t y is subdivided into three congregations, namely, 
the Alexanderwohl, Tabor, and Goessel churches. Goessel a n d Tabor split 
f r o m the Alexanderwohl church in 1909 a n d 1920, respectively. In Figure 
4 the Alexanderwohl church is the m o s t proximal to t h e centroid of the 
array, whereas the Goessel and Tabor churches have b e e n separated f r o m 
their parental groups by the frequencies of the P~ and esterase D 1 alleles. 

T h e bases for the distribution of the groups along the second axis 
( e ^ 1 ) shown in Figure 4 is less obvious. T h e reduced-space representation 
of the alleles used to c o m p u t e the genetic m a p shows that G c 1 F and cde 
separate the populations along the second axis. 

Figure 5 is a plot of the m e a n per locus heterozygosity and ru for 
the various congregations. T h i s m e a n of H a n d ru is based on 44 alleles 
instead of the 10 shown in Figure 2. Alexanderwohl, Meridian, a n d , 
to a lesser extent, Goessel a n d Tabor r e m a i n closest to the theoretical 
regression line (the predicted relationship between the two variables). T h e 
Henderson Mennonite congregation exhibits t h e highest heterozygosity 
level, whereas the Bethesda a n d the Evangelical congregations have the 
lowest H . In this plot the heterozygosity levels of the congregations are 
reduced to some extent because of the a d d i t i o n of a large n u m b e r of 
diallelic loci. 

M a j o r differences are observed in a c o m p a r i s o n of two M e n n o n i t e 
congregation dendograms, one a historical reconstruction of p o p u l a t i o n 
fission and fusion (Figure 6) and the other based on the cluster analysis 
of gene frequencies (Figure 7). Genetically, the Tabor p o p u l a t i o n is sep-
arate from the other M e n n o n i t e c o m m u n i t i e s , even t h o u g h historically 
it split off only two generations ago. T h e Evangelical church in H e n d e r -



Genetic Structure of Mennonites / 507 

Bethesda 

Evangelical 

e X 1 1 1/2 
Goessel 

Tabor 

Meridan 

Alexanderwohl 

Henderson M.B. 

Figure 3. Least-squares reduction genetic map ot the three Mennonite communities 
subdivided into seven congregations. Frequencies from 44 alleles and 15 
genetic loci were used to construct the R matrix. 

0) o < O 
A , 

A C P B K cdE 

0 B F f B M CDE F y b 

A c P c Gc1 G c , s S J k A 

cde ADA 1 B I f ' Ak1 B f 5 0 7 Hp1 B f s Acp* cDe 

\ I 
*= 03 O) _Q J- ~ C 
® > ° (0 © 0) <J) 
CO UJ O I - 2 < I 

1 A c p B S M \ e > l / 2 
CdE Hp1 BfF« / 2 2 

B f 5 0 7 B f F B k Gc 1 I 

cde E s D ' A] 0 ADA1 AcpA J k A A c P c G c ^ J 

Figure 4. Dispersion of the seven Mennonite congregations and alleles along eigenvectors 
associated with the two largest eigenvectors of R and A matrices. 



5 0 8 / CRAWFORD ET AL. 

0.42 

0.40 Henderson 
Meridan 

H 0 0.38 

0.36 

0.34 Evangelical 

0.32 
0.02 0 . 0 3 0.04 0 . 0 5 0.06 0.07 0.08 0.09 0.10 

Figure 5. Plot of mean per locus heterozygosity ( H 0 ) against distance from the centroid 
r / 7 of the R matrix for seven Mennonite congregations. Frequencies from 44 
alleles and 15 loci were used. 

son, Nebraska, and the Goessel congregation c a m e off the genetic tree 
next, even though historically their origins m u s t be traced to Russia. 
These findings support the hypothesis t h a t the fission of the M e n n o n i t e 
gene pools occurred along familial lines a n d does n o t represent a ran-
d o m subset of the p o p u l a t i o n . T h e Bethesda a n d Alexanderwohl com-
munities, both r e m n a n t s of the original division of the Alexanderwohl 
congregation in 1874, cluster together genetically. Yet, despite their het-
erogeneous European origins, b o t h Meridian a n d the H e n d e r s o n Men-
nonite congregations show closer genetic affinities t h a n d o the offshoot 
congregations. 

Discussion 
T h e population history of the M e n n o n i t e s can be characterized in 

t e r m s of a fission-fusion model, which is schematically represented in 
Figure 6. The founders of the various M e n n o n i t e groups c a m e f r o m 
the Netherlands, Switzerland, n o r t h e r n Germany, Moravia, Alsace, and 
Tirol. T h e earliest groups were f o u n d e d by a small n u m b e r of individuals 
coming from different regions of Europe. 

T h e Przechowka Church in West Prussia, which eventually gave rise 
to the Alexanderwohl congregation, was f o u n d e d by 15 refugee couples 
f r o m the Netherlands a n d Switzerland. T h i s congregation, through high 

m 



Genetic Structure of Mennonites / 509 

W e s t e r n E u r o p e R u s s i a U n i t e d S t a t e s 

Tabor 1909 

1874 Alexanderwohl 

A l e x a n d e r w o h l 

A l e x a n d e r w o h l 
(Kansas) G o e s s e l 1920 

A l e x a n d e r w o h l 

N e t h e r l a n d s Evangelical 

B e t h e s d a 

G a l l i c i a n s (Nebraska) Bethesda 

W e s t P r u s s i a n s 
Church of G o d 

in Christ 
Church of G o d 

in Christ Meridian 
O s t o r o g e r s Mennonites 

Mennonite Brethern Henderson M B 

Figure 6. The ethnohistory of the seven Mennonite congregations summarized by a 
dendogram. The dotted lines indicate that various combinations of Dutch 
and German populations contributed to the formation of the Nebraskan and 
Kansan Mennonites. This dendogram is based on an unpublished flow diagram 
constructed by John Janzen. 

fertility a n d t h e a d d i t i o n of other migrants f r o m Prussia a n d Moravia, in-
creased in n u m b e r a n d in 1821 relocated in the Ukraine. The community 
was r e n a m e d Alexanderwohl, a n d when the Russian government revoked 
the military e x e m p t i o n s for Mennonites, the entire c o m m u n i t y emigrated 
t o the U n i t e d States in 1874. O n arrival, Alexanderwohl underwent the 
first fission, with one g r o u p settling in Nebraska a n d founding the town 
of H e n d e r s o n a n d the second group settling in Kansas. T h i s Kansas sub-
division of the Alexanderwohl c o m m u n i t y settled in two communities, 
H o f f n u n g s a u a n d Goessel. Shortly after the t u r n of the twentieth century, 
in 1909, T a b o r split f r o m the Alexanderwohl congregation (located in the 
town of Goessel) a n d established the Goessel church in 1920. In addi-
tion, the Nebraska M e n n o n i t e Bethesda congregation also underwent a 
fission, with t h e separation of the Evangelical Mennonites. 

A similar fission a n d fusion of p o p u l a t i o n s has been described by 
Olsen (1987) for t h e H u t t e r i t e s of N o r t h America. Olsen characterized the 
f o r m a t i o n of H u t t e r i t e colonies on the basis of historic a n d demographic 



5 1 0 / CRAWFORD ET AL. 

CD CD 
AMALG. S r-
DISTANCE ct> co 

C D CO ^t r -
CT) C D CD f ^ 
NT W C\l CO co co co hv! 

Meridian 

H e n d e r s o n MB 

B e t h e s d a 

Alexanderwohl 

Goessel 1920 

Evangelical 

T a b o r 1909 

Figure 7. A dendogram based on the cluster analysis of the frequencies of 8 blood group 
systems and 25 alleles. This clustering method is based on the distance between 
centroids as a criterion for amalgamation. 

evidence for 1878-1970 and likened their structure in m a n y respects to 
the Y a n o m a m o of Venezuela. Although the M e n n o n i t e s d o not explicitly 
follow a policy of splitting off daughter colonies because of high fertility, 
this population subdivision does occur de facto. Most of the General 
Conference M e n n o n i t e splits appear to be d u e t o personality conflicts, 
doctrinal disagreements, or both. Although e c o n o m i c considerations are 
not voiced, they should n o t be ignored as one of the mot i va t i n g factors 
for population fission. 

This fission-fusion process is reflected in the genetic structure of 
the Mennonites. For example, the c o n t e m p o r a r y Alexanderwohl congre-
gation remains proximal to the center of the allelic array, whereas its 
offshoots, Tabor and Goessel, show marked genetic difference. Given the 
short time since the fission of these p o p u l a t i o n s ( 2 - 3 generations), it is 
unlikely that the Tabor a n d Goessel M e n n o n i t e s would experience this 
magnitude of genetic microdifferentiation. T h e two m o r e likely explana-
tions for this genetic uniqueness are p o p u l a t i o n sampling a n d subdivi-
sion by families. Sampling may play a m a j o r role, because the T a b o r and 



Genetic Structure of Mennonites / 511 

Goessel s a m p l e s are the smallest of all the congregations studied. It is 
also likely t h a t the fission process involving Tabor, Goessel, a n d Alexan-
derwohl t o o k place along familial lines, producing a n o n r a n d o m division 
of the gene pool. In s u p p o r t of this, Rogers (1984) d e m o n s t r a t e d through 
the use of s u r n a m e analysis t h a t the initial fissioning of the M e n n o n i t e 
i m m i g r a n t g r o u p was n o n r a n d o m . T h e r e were 63 surnames a m o n g the 
191 families t h a t i m m i g r a t e d ; however, only 3 of these s u r n a m e s (5%) 
are f o u n d in all three settlements. 

T h e M e r i d i a n c o m m u n i t y (Church of G o d in Christ M e n n o ) is a 
heterogeneous p o p u l a t i o n f o u n d e d in 1859. Although this group is small 
a n d highly conservative with regard to endogamy, it has experienced 
a low level of inbreeding (Sirijaraya 1983). As a result of the genetic 
heterogeneity of its f o u n d e r s , the M e r i d i a n congregation r e m a i n s close 
to the regression line in the ru versus H0 c o m p a r i s o n . Meridian displays 
a relatively high heterozygosity b u t a low distance f r o m the centroid of 
the allelic f r e q u e n c y d i s t r i b u t i o n (see Figure 5). 

T h e K a n s a s M e n n o n i t e p o p u l a t i o n s also experienced various de-
grees of r e p r o d u c t i v e isolation a n d inbreeding t h ro u g h o u t their history. 
According to Rogers (1984), their inbreeding coefficient ( F ) varied f r o m 
2% in 1 8 0 0 - 1 8 1 9 to 0.5% in 1860-1874. At present, the General Confer-
ence M e n n o n i t e c o m m u n i t i e s of K a n s a s a n d Nebraska cannot be char-
acterized as genetic isolates because a high p r o p o r t i o n of marriages are 
with n o n - M e n n o n i t e s . D u r i n g the last 2 0 - 2 5 years, approximately 46% 
of all the marraiges in Hoffnungsau, Kansas, were contracted with non-
M e n n o n i t e s (Kay 1978). However, f r o m 1874-1915 the c o m m u n i t y was 
99% e n d o g a m o u s . Since World War I the reproductive isolation has 
rapidly b r o k e n d o w n with the congregation m e m b e r s becoming m o r e 
integrated into m a i n s t r e a m American life. Before 1936 m o r e than 95% 
of the marriages were M e n n o n i t e e n d o g a m o u s . Marital partners were 
M e n n o n i t e b u t f r o m a different village or town. By 1957 M e n n o n i t e en-
d o g a m y o c c u r r e d in 72.3% of the marriages a n d 27.7% were exogamous 
(Kay 1978). 

T h e b l o o d genetics s u p p o r t the conclusion that the General Con-
ference M e n n o n i t e s described in this study are n o longer m e m b e r s of 
genetic isolates a n d are culturally approaching the m a i n s t r e a m farming 
c o m m u n i t i e s . T h e high levels of heterozygosity reflect considerable sys-
t e m a t i c pressure in the f o r m of migration. T h e m e a n per locus heterozy-
gosity varies f r o m 33% to 41%, based on 29 b l o o d systems. In addition, 
the presence of several African m a r k e r genes suggests gene flow f r o m the 
American black gene p o o l into the M e n n o n i t e groups. 

C o n t e m p o r a r y K a n s a s a n d Nebraska General Conference M e n n o n -
ite genetic s t r u c t u r e can best be explained on the basis of episodes of 



5 1 2 / CRAWFORD ET AL. 

fission and fusion, followed by a rapid breakdown of reproductive isola-
tion. Thus these Mennonite populations can n o longer be characterized 
as highly inbred genetic isolates. 
Acknowledgments This research was supported in part by the National 
Institute of Aging under grant AGO 1646 and the University of Kansas under 
Faculty Research Award 3326-5038. Crawford received support through a Public 
Health Service Research Career Development Award (K04DE0028-05). We 
would like to thank the Mennonite communities of Goessel, Henderson, and 
Meridian for their patience and good will in this project. Special words of thanks 
go to John Janzen, whose reconstruction of the ethnohistory was used in this 
publication, and to Laurine Rogers, who helped organize and administer this 
research program. 

Revision received 3 June 1989. 

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