Inferences of selection and migration in the Danish house ...webpages.icav.up.pt/PTDC/BIA-BEC/103440/2008... · between the house mouse subspecies Mus musculus domesticus and M. m

Biological Journal of the Linnean Society

, 2005,

84

, 593–616. With 5 figures

© 2005 The Linnean Society of London,

Biological Journal of the Linnean Society,

2005,

84

, 593–616

593

Blackwell Science, LtdOxford, UKBIJBiological Journal of the Linnean Society0024-4066The Linnean Society of London, 2005? 2005843593616Original Article

SELECTION IN THE HOUSE MOUSE HYBRID ZONEN. RAUFASTE

Et al.

*Corresponding author. E-mail: [email protected]†Present address: INRA CBGP, Campus de Baillarguet, 34988 Montferrier sur Lez, France

The genus Mus as a model for evolutionary studies

Edited by J. Britton-Davidian and J. B. Searle

Inferences of selection and migration in the Danish house mouse hybrid zone

NATHALIE RAUFASTE

1

, ANNIE ORTH

1

, KHALID BELKHIR

1

, DAVID SENET

1

, CAROLE SMADJA

2

, STUART J. E. BAIRD

1†

, FRANÇOIS BONHOMME

1

, BARBARA DOD

1

and PIERRE BOURSOT

1

*

1

Laboratoire Génome Populations Interactions Adaptation (UMR 5171 IFREMER-CNRS-UMII), and

2

Institut des Sciences de l’Evolution (UMR 5554 CNRS-UMII), Université Montpellier II, France

Received 30 October 2003; accepted for publication 7 October 2004

We analysed the patterns of allele frequency change for ten diagnostic autosomal allozyme loci in the hybrid zonebetween the house mouse subspecies

Mus musculus domesticus

and

M. m. musculus

in central Jutland. After deter-mining the general orientation of the clines of allele frequencies, we analysed the cline shapes along the direction ofmaximum gradient. Eight of the ten clines are best described by steep central steps with coincident positions and anaverage width of 8.9 km (support limits 7.6–12.4) flanked by tails of introgression, indicating the existence of a bar-rier to gene flow and only weak selection on the loci studied. We derived estimates of migration from linkage dise-quilibrium in the centre of the zone, and by applying isolation by distance methods to microsatellite data from someof these populations. These give concordant estimates of

s

=

0.5–0.8 km generation

–

. The barrier to gene flow is ofthe order of 20 km (support limits 14–28), and could be explained by selection of a few per cent at 43–120 under-dominant loci that reduces the mean fitness in the central populations to 0.45. Some of the clines appear symmet-rical, whereas others are strongly asymmetrical, and two loci appear to have escaped the central barrier to gene flow,reflecting the differential action of selection on different parts of the genome. Asymmetry is always in the directionof more introgression into

musculus

, indicating either a general progression of

domesticus

into the

musculus

terri-tory, possibly mediated by differential behaviour, or past movement of the hybrid zone in the opposite direction,impeded by potential geographical barriers to migration in

domesticus

territory. © 2005 The Linnean Society ofLondon,

Biological Journal of the Linnean Society

, 2005,

84

, 593–616.

ADDITIONAL KEYWORDS:

gene flow – genetic barrier – hybridization –


–

Mus

musculus musculus

– speciation.

1 2/

INTRODUCTION

Hybrid zones have been referred to as ‘windows onthe evolutionary process’ (Harrison, 1990) becausethey allow us to study the interplay between migra-tion and selection on the evolution of genetic differen-tiation and adaptation. When parapatric taxa meet

and hybridize, selection against unfit hybrids cancounteract the homogenizing effect of migration andlead to the establishment of frequency clines of diag-nostic characters at the boundary between their dis-tribution areas. A detailed population genetics theoryhas been developed to model the expected patterns ofallele frequency changes in such situations (e.g. forreviews see Barton & Hewitt, 1985; Barton & Gale,1993), showing that they can constitute barriers togene flow between the taxa, of increasing intensitywith the number of loci involved in hybrid unfitness,and with the spread of these loci in the genome.

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2005,

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, 593–616

Briefly, this is because migration brings parental gen-otypes to the centre of the hybrid zone where neutralloci are thus in linkage disequilibrium with loci underselection in hybrids, which impedes free introgressionof these neutral loci and causes an abrupt change oftheir allele frequency in the centre. However, theseloci can eventually extricate themselves from thisnegative genetic background by recombination, andform long tails of introgression into the foreign terri-tory. The resulting clines (described by the shapes ofthe central step and of the tails of introgression) arerelatively independent of the type of selection againsthybrids (which is usually unknown), and can be usedto quantify the intensity of the selection maintainingthe zone and of the resulting barrier to gene flow.Their estimation can also provide some informationabout the number of loci involved in the selection ofhybrids, an important characteristic of the mecha-nisms leading to incompatibilities between differenti-ating genomes.

Since the pioneering work of Hunt & Selander(1973), several authors have studied the genetics ofthe hybrid zone between the two European subspeciesof the house mouse,


and

M. m. musculus

, that are thought to have come intosecondary contact in Europe after a period of indepen-dent geographical expansion from the Middle East,with

M. m. domesticus

colonizing the Mediterraneanbasin and Western Europe while

M. m. musculus

wasexpanding across central Europe (e.g. see Boursot

et al

., 1993, for a review). In previous studies, clinewidths were roughly quantified by visually inspectingthe variations of synthetic morphological or genetichybrid indexes: 90% of the genetic transition occursover 20 km in Denmark (Hunt & Selander, 1973), 75%over 20 km in southern Germany (Sage, Whitney &Wilson, 1986b), 80% over 36 km in Bulgaria (Vanler-berghe

et al

., 1988), and 60% of genetic and morpho-logical variation in 20–40 km in East Holstein (Prager

et al.

, 1993). The first attempt to estimate cline widthsinvolved the application of a simple sigmoid model onthe south German transect (Tucker

et al

., 1992). Theyfound narrower cline widths for the sex chromosomemarkers (Y chromosome, 4 km; X chromosome mark-ers from 4 to 10 km) compared with autosomal alloz-ymes (from 6.4 to 21.2 km). A similar contrastbetween the sex chromosomes and autosomal loci wasalso found in Denmark and Bulgaria (Vanlerberghe

et al

., 1986; Dod

et al

., 1993). However, none of thesestudies had enough samples both in the centre and inthe tails of introgression for a detailed analysis of thecline shape to be realistic. In addition, sampling wasoften carried out in a linear fashion in an arbitrarydirection across the transect, allowing comparisonsbetween markers, but not the calculation of clineparameters along the line of maximum slope. Further-

more, none of these studies included estimations ofmigration, nor of linkage disequilibrium that could becombined with cline widths to estimate the intensityof selection against hybrid mice. Here we analyse alarge dataset on the Danish hybrid zone characterizedfor ten diagnostic allozyme loci, and derive indepen-dent estimates of migration using microsatellite loci.

MATERIAL AND METHODS

M

ICE

Mice were live trapped inside buildings using multi-capture wire traps, during several field trips from1984 to 2000. The location of the sampling sites in theJutland peninsula is indicated on Figure 1 and the listof localities with their Universal Transverse Mercator(UTM) coordinates are given in Appendix 1.

P

ROTEIN

ELECTROPHORESIS

Mice were killed and dissected in the field, and liver,kidney, heart, plasma and blood cells were kept in liq-uid nitrogen for further preparations. Protein extrac-tions, separation by starch gel electrophoresis (oracrylamide gels in the case of Amylase) and detectionof enzyme activity in the gels followed standard pro-tocols, such as described in Pasteur

et al

. (1987). Theloci were chosen for their ability to distinguishbetween the two subspecies in previous studies onhouse mice in the Jutland peninsula and on a broadergeographical scale (Hunt & Selander, 1973; Bonho-mme

et al

., 1984; Britton-Davidian, 1990; Din

et al

.,1996). The alleles were identified by comparison withstandards obtained from mice of known genotypes,and each locality was characterized by the frequencyof

M. m. musculus

alleles.

O

RIENTATION

OF

THE

CLINES

The general orientation of the maximum gradient ofallele frequency across the hybrid zone was deter-mined by fitting the allele frequency data to a simplesigmoid model, where the logit transform of the allelefrequencies is a linear function of the two-dimen-sional (2D) geographical coordinates. The model wasfitted by maximum likelihood, assuming a binomialerror on the estimations of allele frequencies, usingthe computer package GLIM4 (the Numerical Algo-rithm Group). This orientation procedure determinedthe direction of maximum gradient of allele fre-quency, assuming the centre of the hybrid zone is astraight line, and that the frequency change is sig-moid. The coordinate of each locality was then calcu-lated by projection on this direction of maximumgradient.

SELECTION IN THE HOUSE MOUSE HYBRID ZONE

595



2005,

84

, 593–616

F

ITTING

CLINE

SHAPE

The

musculus

allele frequencies for each locus in thedifferent localities along the 1D transect were fitted tovarious models of cline shapes by maximum likelihoodestimation, using the computer package Analyse(by N. Barton & S. Baird, http://helios.bto.ed.ac.uk/evolgen/Mac/Analyse/).

Sample sizes were corrected according to:

(1)

(adapted from Szymura & Barton, 1986, 1991), where

N

is the number of individuals sampled in the locality.

F

IS

is the deficit of heterozygotes (set to zero if not pos-itive) and is used to correct for the non-independencebetween sampling of alleles when there is inbreeding.

F

ST

represents the fluctuations of allele frequenciesbetween loci that are not accounted for by differencesin their cline shapes. It represents the residual varia-tion around the regression of allele frequencies atindividual loci in each locality against the average ofall loci, and was estimated using the ‘concordance’procedure in the Analyse package. The above correc-tion is designed for a single locus. When data for sev-

NeN

N Fst Fis=

+ +2

2 1*

eral loci were pooled, the effective sample size wastaken as the sum of effective sample sizes for the dif-ferent loci.

The Analyse package is then used to compare thelikelihood of the allele frequency data under three dif-ferent models of cline shape: a sigmoid cline, andclines in three parts, with a central sigmoid part andtwo exponential tails of introgression, either identicalon both sides or different. The first model has twoparameters,

w

, the width of the cline (inverse of themaximum slope), and

c

, the geographical position ofthe centre. The second model has two additionalparameters (four in total) describing the shape of theexponential tails, and the last model has two suchparameters for each tail (six parameters in total). Asthe models are nested, a likelihood ratio test can beapplied to choose the model that best explains the datawith a minimum of parameters, by assuming thattwice the difference of log-likelihood between two mod-els follows a chi-squared distribution with the numberof degrees of freedom equal to the difference in thenumber of parameters between the two models.

The Analyse program uses a Metropolis randomexploration algorithm to find the maximum likelihood

Figure 1.

Location of the sampling sites are shown by open triangles on the map, and the axes give their UTM coordinates.Inset: the location of the study area in the Jutland Peninsula. The grey line running across the map is the position of thecentre of the hybrid zone that ends in the east at the head of the Vejle Fjord. The rivers are drawn on this map, and someare highlighted by thicker lines (see text).

6120

6140

6160

6180

6200

6220

475 495 515 535 555 575

10 km

N

http://helios.bto.ed.ac.uk/

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estimate of the parameters, so it was run many timeson each dataset, with different starting conditions anddifferent settings of the parameters controlling theexploration algorithm, in order to explore the param-eter space as thoroughly as possible. Two log-likelihood support limits of parameter estimates weredetermined by inspecting the results of 10 000–50 000explorations of the parameter space with the algo-rithm. The likelihood profiles (Hilborn & Mangel,1997) for position and width were explored using the‘crossection’ option of Analyse, as described in Phil-lips, Baird & Moritz (2004): the parameter of interest(

c

or

w

) is set to a fixed value, and the other parame-ters are searched until the maximum likelihood of thedata is found for this value of

c

(

w

). The procedure isrepeated for a range of values of

c

(

w

) until the rangeof relevant values is covered, and the likelihood profilecan thus be generated for the parameter of interest.

E

STIMATING

SELECTION

PARAMETERS

The parameters of the fitted cline shapes can be usedto infer some population parameters of interest, usingthe existing theory of tension zones, as detailed in anumber of papers (Barton & Hewitt, 1985; Szymura &Barton, 1986, 1991; Barton & Gale, 1993; Kruuk

et al

.,1999; Barton & Shpak, 2000). We will summarize herethe part of this complex theory that was used.

As we will see, a cline shape in three parts, with acentral sigmoid part and two exponential tails, bestdescribes most loci studied. This is the shape expectedfor a locus under weak selection (creating the expo-nential tails of introgression) submitted to the influ-ence of several loci under stronger selection, creatingthe central barrier to gene flow, and the central step inthe clines. Although some of the theory summarizedbelow is derived for neutral loci, the hypothesis ofweak selection on the studied loci needs to be intro-duced to derive equilibrium cline shapes (the onlyequilibrium for neutral loci would be even frequency,e.g. Szymura & Barton, 1986, 1991).

The central sigmoid step can be described by twoparameters,

c

, the position of the centre, and

w

, thewidth of the cline (inverse of the maximum slope). Thefollowing relation relates the total selection acting onthe locus to the width of the cline:

(2)

where

s

2

is the migration parameter (variance of dis-tance from parent to offspring). This is a nuisanceparameter, but it is possible to estimate it by compar-ing the clines for different loci and their linkage dise-quilibria. It is expected to be proportional to thelinkage disequilibrium and the rate of recombinationbetween the loci, and to the gradients of allele fre-

sw

* = 8 2

2

s

quencies for these loci. Standardized linkage disequi-libria (linkage disequilibrium standardized by themaximum possible value given allele frequencies)between pairs of loci were estimated by maximumlikelihood with the program Analyse, in localities fromthe centre of the hybrid zone, where linkage disequi-librium is expected to be maximum, and where thegradient of allele frequencies is estimated as theinverse of cline width. The migration parameter wasestimated for each locality and each pair of loci. Theestimates obtained are then averaged over pairs of lociand localities to obtain a single estimate of the migra-tion parameter. Because the statistical properties ofsuch an average are not known, we also used the vari-ance of the individual hybrid index to estimate theaverage linkage disequilibrium between loci (Barton& Gale, 1993),

D

—.The equations of the exponential tails of introgres-

sion allow the inference of other important parame-ters. The equation of the left tail of the cline is:

(3)

and that of the right tail

(4)

It can be seen that parameters

q

represent thesquare of the ratio between the expected rate of decayin the tails without a barrier and the actual rate ofdecay. They can thus be used to estimate the ratiobetween the selection on the locus under study itselfand the total selection experienced by this locus,including the influence of other loci:

(5)

The program Analyse does not give estimates of the

a

parameters but of parameters

B

/

w

, the ratio of thebarrier to gene flow to the width of the clines. The rela-tionship between these parameters is:

(6)

and similarly for

B

1

.By making the hypothesis that selection on the loci

studied is weak, that the number of loci under selec-tion is not too small (1/

n

<<

1) and that selection actsagainst heterozygotes, one can derive approximateestimates of the number of loci under selection creat-ing the barrier and the intensity of the selection oneach locus using the two following relationships (e.g.Barton & Shpak, 2000):

(7)

p xx cw

( ) = -( )ÊË

ˆ¯a q0 0

2exp ,

p xx cw

( ) = - - -( )ÊË

ˆ¯1

21 1a qexp .

q = sslocus

*.

Bw

00

0 1

021= - -

qa aa

. ,

sB

Bw u

= ÊË

ˆ¯

8 2

22s

ln andD

SELECTION IN THE HOUSE MOUSE HYBRID ZONE 597

© 2005 The Linnean Society of London, Biological Journal of the Linnean Society, 2005, 84, 593–616

(8)

where Du is the height of the central step of allele fre-quency, and is estimated as the difference in the fre-quencies of the two exponential tails taken at thecentre (x = c). We thus have Du = 1 - a0 - a1, whichbecomes:

(9)

Note that in the literature, Du is sometimes omitted,supposing that selection is strong and that it is close to1 (Szymura & Barton, 1986, 1991).There remains onenuisance parameter in the equations above, which isr̄, the average recombination rate between the neutral(or quasi-neutral in that case) locus and the loci underselection. Several methods have been suggested toestimate r̄ (Barton & Hewitt, 1985; Barton & Bengts-son, 1986). Here we take the conservative value of 0.5that gives the upper limit of n, the number of lociunder selection. In the framework of this underdomi-nance model, the average fitness of the central popu-lations can be estimated (taking the fitness of theparental populations arbitrarily at 1):

(10)

MICROSATELLITE TYPING

Genomic DNA was extracted from the spleen usingstandard proteinase K/phenol-chloroform methods.Polymorphism was studied at six microsatellite locithat are described in Table 1. Five of them are com-mon with the previous study of Dallas et al. (1995).The loci were amplified by PCR. One of the amplifica-tion primers was fluorescently labelled (Cy5) andallele sizes were measured after migration in a dena-turing acrylamide gel on an automated sequencer

nr

BB

w u

=ÊË

ˆ¯2s ln

D

Du wB B

B Bw

B B=

+ +

2

2

0 0 1 1

0 0 1 1 0 0 1 1

q q

q q q q.

Wns

H = -ÊË

ˆ¯exp

2.

(Pharmacia). Allele sizes for all loci were measuredrelative to the corresponding allele of the Balb/cmouse laboratory strain, which was arbitrarily attrib-uted size 50. We intentionally selected loci with 3–5-bprepeats (except D17Mit41, 2 bp, Table 1), becauseallele size determination is more reliable than for 2-bprepeats.

ESTIMATION OF MIGRATION FROM MICROSATELLITE ALLELE FREQUENCIES

We estimated the migration parameter s under isola-tion by distance models, using three different methodsapplied to the microsatellite data. The first methodcalculates the regression between FST /(1 - FST) andthe logarithm of geographical distance between popu-lations, r (Rousset, 1997). It applies to pairs of popu-lations in a 2D habitat, and the slope of the regressiongives an estimate of 1/4pDs2, where D is the local pop-ulation density, and s2 is the dispersion parameter. Wechose FST rather than RST (Slatkin, 1995) because it isthought to be more conservative when sample sizesare small, and only a few loci are used (Gaggiotti et al.,1999), but also because of its small variance and of theuncertainties about the mutation model underlyingthe justification of RST. The second method calculatesthe regression of statistic over the logarithm of geo-graphical distance, r (Rousset, 2000b), the slope ofwhich also provides an estimate of 1/4pDs2. Here,however, pairs of individuals rather than of popula-tions are compared, and statistic is given by (Qw - Qr)/(1 - Qw), where Qr is the probability of identity bystate of two alleles separated by a distance r, and Qw isthe probability of identity by state of the two allelesfrom the same individual. The third method used herecalculates the regression of a spatial autocorrelationstatistic, Moran’s I, over the logarithm of geographicaldistance r (Hardy & Vekemans, 1999). It also appliesto pairs of individuals, and the slope of the regressioncan be used as an estimate of (1 - FIT)/2pDs2(1 + FIT).Computer software GENETIX (http://www.univ-montp2.fr/~genetix/genetix/genetix.htm) was used to

)ar

)ar

Table 1. Description of the microsatellite loci studied

LocusName of locus in Dallas et al. (1995)

Balb/c allele in Dallas et al. (1995)

Motif size (bp)

No. ofalleles Chromosome

Position fromcentromere (cM)

Cypla2 34 4 4 11 9 3.1Ckmm 105 1 5 7 7 4.5Ktr2 126 4 4 5 15 58.7Gfap 150 4 3 16 11 62D15Mit16 D15Mit16 3 4 11 15 61.7D17Mit41 – – 2 10 17 53

http://www.univmontp2.fr/~genetix/genetix/genetix.htm

598 N. RAUFASTE ET AL.


calculate FST and convert data files between differentformats, while GENEPOP (ver. 3.3, Raymond & Rous-set, 1995) allowed the calculation of . A programwas written to calculate Moran’s I, according to theformula in Hardy & Vekemans (1999).

RESULTS

ALLOZYME ANALYSES

A total of 170 localities, listed in Appendix 1 with theirgeographical coordinates, were studied here and theirlocation is indicated in Figure 1. Most yielded fivemice or fewer (see Appendix 1 and the distribution ofsample sizes in Table 2). The results of the geneticanalysis at the ten enzymatic autosomal loci (Amy,Es1, Es10, Es2, Gpd, Idh, Mpi, Np, Pgm and Sod) aregiven in Appendix 2. A total of 1233 mice were studied,but only 996–1187 were successfully typed, dependingon the locus.

Samples from the same locality but different collec-tion years were treated separately. An independentstudy (to be published elsewhere) failed to find anyevidence that this hybrid zone has moved since it wassampled in the late 1960s by Hunt & Selander (1973)and our sampling period which extended from 1984 to2000. We therefore considered here that the zone wasstable enough for us to analyse all sampling yearstogether, in order to increase the quality of the sam-pling along the transect.

ORIENTATION OF THE TRANSECT

When fitting a sigmoid cline to the data in two dimen-sions, the centre of the hybrid zone was found to runwest-north-west to east-south-east, at a 7∞ angle clock-wise from the west–east direction (as drawn on the mapof Fig. 1). We will see below that most loci do not fit thesimple sigmoid cline model used here, but rather morecomplex models with a central step of allele frequency.

)ar

We were thus concerned that the use of this simplemodel could lead to an erroneous determination of thedirection of the clines. Using the more complex models,we tested several orientations with angles from 0 to 20∞and found in all cases that the 7∞ orientation gave thebest likelihood (data not shown). The coordinates of thelocalities along the transect were thus calculated byprojecting them onto an axis perpendicular to this cen-tral line. However, as can be seen on the map, the centreof the hybrid zone reaches at its eastern end the headof a deep fjord (the Vejle Fjord). For localities furthereast of this point our standard procedure would cer-tainly underestimate their distance to the centre of thehybrid zone. Thus, the straight line distance betweenthe locality considered and the head of the fjord wasadded to (north of the hybrid zone centre) or subtractedfrom (south of the hybrid zone centre) the transect coor-dinate of the head of the fjord. The amended transectcoordinates are listed in Appendix 1, and are used inthe following analyses.

CLINE SHAPES

The 1D coordinates of the localities calculated asdescribed above were used to analyse the shape of thecline of allele frequency along the transect. The threemodels implemented by the computer program Anal-yse were fitted successively: sigmoid cline (two param-eters), symmetric stepped cline (four parameters) andasymmetric stepped cline (six parameters). The likeli-hood ratio test was used to compare the models andthe choice was made at the 5% significance level.Table 3 shows the type of model retained for eachlocus using this criterion. The sigmoid model (twoparameters) could not be rejected for Idh and Sod,while a stepped symmetric model (four parameters)was retained for Es1 and Mpi, but the stepped asym-metric model (six parameters) provided the best fit forthe six remaining loci. The best positions of the cen-tres for each locus and their 2 Log-Likelihood (LL)support limits are given in Table 3. We then testedwhether an acceptable common centre for all loci couldbe found. To do this, we calculated the best LL of eachlocus dataset for a range of centres spanning the con-fidence intervals of all loci, leaving all the otherparameters free to vary. We then summed the best LLsover loci for each of the centres tested, and determinedthe position that provides the best summed LL(method described in Phillips et al., 2004). This model,with a common centre at position 6191.8 (Table 3), issignificantly worse than that where the centre is freeto vary at each locus (P = 1.4 ¥ 10-12). Inspection of theindividual tests for each locus between the free andconstrained centre models identifies two clear outliers(Table 3): Idh and Sod, the two loci for which sigmoidclines were retained. We reiterated the above proce-

Table 2. The distribution of sample sizes

No. of mice No. of localities

1–5 1036–10 26

11–15 1916–20 1021–25 426–30 531–35 136–40 141–45 046–50 051–55 1



Tab

le 3

.M

axim

um

lik

elih

ood

posi

tion

of

the

cen

tre

for

each

loc

us,

an

d th

e li

keli

hoo

d se

arch

of

a co

mm

on c

entr

e

Loc

us

Bes

t ce

ntr

e fo

r ea

ch l

ocu

sB

est

com

mon

cen

tre

all

loci

Bes

t co

mm

on c

entr

e 8

loci

Mod

elC

entr

e2L

L l

imit

sL

LC

entr

eL

L2D

LL

d.f.

PC

entr

eL

L2D

LL

d.f.

P

Am

y6

6189

.2(6

188.

0–61

91.3

)-8

3.55

6191

.8-8

6.75

6.39

10.

011

6191

.2-8

5.13

3.16

10.

076

Es1

461

92.0

(619

0.8–

6193

.3)

-62.

0761

91.8

-62.

150.

151

0.70

061

91.2

-62.

911.

681

0.19

5E

s10

661

91.6

(618

9.3–

6193

.2)

-70.

6261

91.8

-70.

750.

261

0.60

761

91.2

-70.

890.

541

0.46

2E

s26

6191

.1(6

190.

1–61

96.7

)-9

9.70

6191

.8-9

9.87

0.33

10.

564

6191

.2-9

9.60

0.20

10.

656

Gpd

661

91.2

(618

5.5–

6194

.8)

-85.

4361

91.8

-86.

051.

241

0.26

661

91.2

-85.

430.

001

1.00

0Id

h2

6198

.9(6

196.

1–62

01.7

)-8

2.75

6191

.8-9

4.27

23.0

51

1.6

¥ 10

-6

Mpi

461

90.4

(618

9.3–

6191

.5)

-76.

0461

91.8

-79.

076.

071

0.01

461

91.2

-77.

112.

141

0.14

3N

p6

6191

.7(6

190.

7–61

94.6

)-7

5.53

6191

.8-7

5.55

0.03

10.

862

6191

.2-7

6.03

0.99

10.

320

Pgm

661

93.8

(619

1.6–

6196

.3)

-67.

3061

91.8

-69.

243.

881

0.04

961

91.2

-69.

544.

481

0.03

4S

od2

6199

.0(6

196.

6–62

01.4

)-8

1.30

6191

.8-9

8.21

33.8

41

6.0

¥ 10

-9

All

loc

ibe

st f

or

each

loc

us

-784

.29

6191

.8-8

21.9

075

.23

91.

4 ¥

10-1

2

All

loc

i ex

cept

Idh

an

d S

odbe

st f

or

each

loc

us

-620

.24

6191

.2-6

26.6

412

.79

70.

077 dure for all loci except these two, and found an accept-

able common centre for these loci (at position 6191.2,P = 0.077, Table 3).

Using this common centre we determined the bestcline shapes for these eight loci. The data points andthe fitted cline shapes are plotted on Figure 2, and thecline parameters are given in Table 4. A variety ofshapes are observed among the clines with a centralstep. The height of this step in allele frequency varies,the highest being for Amy. Although two loci, Es1 andMpi, have relatively symmetric introgression pat-terns, all the other loci show an asymmetry that isalways in the same direction. They are characterizedby a steep central change in allele frequency thatoccurs mostly on the domesticus side, where introgres-sion past this barrier is less extensive than on themusculus side (this is particularly pronounced forEs10, Gpd and Pgm). The most pronounced asymme-try of introgression is seen at the Idh and Sod loci,whose centre is displaced almost 8 km into the mus-culus territory as compared with the common centre ofthe other loci. For these two loci, no model with a cen-tral step (around frequency 0.5) fitted the data betterthan the simpler sigmoid model (see Fig. 3).

It can also be seen in Figure 2 that there is consid-erable dispersion of the data points around the model.This is reflected in the poor precision of the parameterestimates, given in Table 4 with their two LL supportlimits. The variations between loci and the supportlimits of the parameters are particularly large for theleft side, with the left barrier parameter absurdlylarge for several loci. The parameters defining theright side of the cline are generally better estimatedand more consistent across loci, but the support limitsstill remain rather wide. When the average of theeight loci with a common centre is used (averagehybrid indices reported in Appendix 1), more reliableparameter estimates with reasonably narrow supportintervals are obtained (Table 4). The parametersdefining the right and left sides are similar to eachother, and the average cline for these eight loci isshown graphically in Figure 4. The Y chromosomedata for the same samples (data from Dod et al., 2005,this issue) are also reported in this figure, showingthat the centre of the allozyme clines corresponds tothe major and abrupt change for this chromosome.

ESTIMATING SELECTION PARAMETERS FROM CLINE SHAPES

Given the estimates of cline widths derived above forthe different loci, it is theoretically possible to esti-mate the migration parameter s using linkage dise-quilibria between loci in the centre of the hybrid zone.We estimated the standardized linkage disequilibriabetween pairs of loci in each locality for the eight loci



that have coincident centres (Amy, Es1, Es2, Es10,Gpd, Mpi, Np and Pgm) using the maximum likeli-hood procedure in the Analyse package. With the esti-mated cline width for each locus given in Table 4, we

then obtained estimates of the migration parameter.All pairs of loci are unlinked on the genetic map,except Es1 and Es2, at 9 cM on chromosome 8, andEs10 and Np, at 22 cM on chromosome 14, and these

Figure 2. Variations of Mus musculus musculus allele frequencies along the transect across the hybrid zone, for eightautosomal loci. Shaded circles represent the real data, and the area of the circles is proportional to the effective numberof alleles sampled (after correction for FIS and FST, see text). The filled circles represent the best fit with a common centrefor the eight loci, the position of which is shown by the thin vertical line.

Amy Es1

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values of recombination were used to estimate s. Theresults were then averaged over pairs of loci and local-ities. We chose localities from the centre of the hybridzone with more than ten mice in the sample (localitiesnumbers 102, 106, 120 and 142: Appendix 1). This ledto an average estimate of the migration parameter s of0.75 km generation– . We also applied the othermethod based on the variance of the hybrid index(Barton & Gale, 1993) to 145 mice from the 24 mostcentral populations. The variance of hybrid index was0.021, and the part due to heterozygosity 0.014. Theexcess variance is attributed to linkage disequilib-

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rium, and with an average cline width of 8.9 km, weget D = 0.015 and s = 0.77 and 0.63 km generation–

(whether sampling occurred before or after migration,Barton & Gale, 1993), in good agreement with the pre-vious estimate.

In order to calculate the intensity of selectionneeded to balance dispersal against selection andrecombination, cline shape has to be estimated. To dothis we used the parameters of the average cline for

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Table 4. Maximum likelihood estimates of cline parameters, and their two log-likelihood support limits, for eight loci witha common centre at position 6191.2

Locus w B0/w q0 B1/w q1

Amy 20.6 (15.2–26.4) 123.51 (2.27–inf) 0.002 (0.000–0.284) 9.67 (2.74–108.45) 0.037 (0.001–0.157)Es1 11.0 (2.2–25.2) 0.67 (0.03–5.57) 0.172 (0.010–0.999)Es10 6.4 (4.9–11.7) 5.15 (1.16–332.67) 0.085 (0.004–0.457) 3.95 (1.81–7.54) 0.016 (0.008–0.055)Es2 3.8 (1.3–9.2) 14.35 (1.91–123.26) 0.004 (0.000–0.058) 6.50 (1.66–23.25) 0.003 (0.000–0.019)Gpd 6.3 (3.4–13.0) 2.59 (0.92–27.71) 0.029 (0.002–0.138) 1.09 (0.68–3.05) 0.032 (0.009–0.135)Mpi 6.2 (2.2–18.9) 2.16 (0.35–9.01) 0.041 (0.004–0.453)Np 5.9 (3.8–44.6) 0.91 (0.15–inf) 0.045 (0.000–1.000) 2.39 (0.39–7.05) 0.009 (0.002–0.715)Pgm 6.8 (4.0–9.1) 146.49 (9.98–inf) 0.016 (0.001–0.098) 2.19 (1.91–5.04) 0.035 (0.010–0.061)Eight loci 8.9 (7.7–12.4) 2.25 (1.09–3.76) 0.072 (0.043–0.163) 2.22 (1.31–3.05) 0.033 (0.023–0.063)

Figure 3. Allele frequency variations along the transectfor two autosomal loci that do not fit the general clineposition and shape of the eight loci in Fig. 2. Symbols areas in Fig. 2.

Idh

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Figure 4. Allele frequency variations along the transect,for the average of eight autosomal loci (Amy, Es1, Es2,Es10, Gpd, Mpi, Np and Pgm), and the Y chromosome (datafrom Dod et al., 2005, this issue). Symbols are as in Fig. 2.

Eight allozymes

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the eight coincident loci. We have seen above that,individually, most of the clines are asymmetrical, butthat this asymmetry is tempered when the averageintrogression over the eight loci is considered. Usingeither the right or the left parameters of this cline fitgives consistent estimates of selection parameters.The barrier to gene flow on both sides is of the order ofB ~ 20 km (2LL support limits 12–33 km on the leftside and 14–28 on the right side), and the height of thecentral step in allele frequency equals 0.45. The totalselection acting on the loci studied is estimated ass* = 0.040–0.059 (eqn 2, with s = 0.63–0.77, seeabove). The selection acting on each of the loci causingthe genetic barrier is estimated as s = 0.021–0.030(eqn 7). The number of loci under selection creatingthe central barrier is estimated to be n ~ 52–78 byassuming that the average recombination between theloci studied and the selected loci is r = 0.5 (eqn 8). Thisis thus an overestimate of the number of loci. Theaverage fitness of hybrid central populations is esti-mated to be WH = 0.45 (eqn 10). Finally, the selectionacting on the loci under study is estimated usingeqn (5) to be on average of the order of slocus = 0.003–0.004, or 0.001–0.002 depending on whether the left orright cline parameters, respectively, are considered.These low values are in agreement with the hypothe-sis that was made of weak selection on these loci inorder to apply the approximations needed for theabove parameter estimations.

ESTIMATING MIGRATION FROM MICROSATELLITE DATA

In order to estimate migration independently of thecline analyses, we typed some of the populations at sixmicrosatellite loci and applied various methods of esti-mation of migration under isolation by distance. Weselected the two sampling years with the largest sam-ples (1992 and 1998) and analysed the results for eachyear separately. Here we are interested in the part ofthe genetic differentiation between populations andindividuals that results from migration and driftalone, so we want to remove the effect of selection inthe hybrid zone. Ideally, this could be done by analys-ing the residual variation of allele frequencies aroundthe cline fits for polymorphic, yet diagnostic, markers.However, most microsatellites tested did not meetthese criteria. We thus chose to study allele frequencyvariation among groups of populations more or lessaligned in a direction perpendicular to that of the cli-nal gradient of the hybrid zone, by restricting the com-parisons to pairs of populations whose average indexof hybridization for the allozymes differed by less than10% (Appendix 1).

The 1992 sample consisted of 185 individuals from14 different localities. For 1998 we used 436 mice from64 localities. Only 113 mice could be typed for locus

D17Mit14 in the 1992 sample (381 in the 1998 sam-ple), but from 171 to 184 were typed at the five otherloci (from 421 to 436 in 1998). The number of differentalleles found at each locus is indicated in Table 1. Thegenotypes of these 621 mice at the six loci are given inAppendix 3. They include some 1992 data alreadyreported in Dallas et al. (1995). We retyped a fractionof the mice from the earlier study to establish the cor-respondence of allele sizes between the two studies.

For the isolation by distance methods, we not onlyrestricted the comparisons to pairs of populations withsimilar hybridization indexes, as argued above, butalso to pairs not too far from each other, because it isrecommended that pairs be generally separated by nomore than 10s (Rousset, 2000b), and in no case bymore than 20s (Rousset, 2000a). We chose a cut-off at15 km. By using the FST method on pairs of popula-tions selected in this way, we derived an estimate ofDs2 of 3.7 individuals on the 1992 dataset, and of 2.7individuals for 1998. This method relies on estimatesof population frequencies, and in the regression, datapoints are not weighted according to sample sizes, sothese values must be considered with caution. The and Moran I methods do not have this drawback,because they are based on comparisons between indi-viduals, but they appear to be less adapted to the hab-itat structure of mice, which is fragmented ratherthan continuous. They gave consistent estimates ofDs2, but showed differences between 1992 (Ds2 = 1.4and 1.7 for the two methods, respectively) and 1998(Ds2 = 3.7 and 3.9). The 1998 estimate is presumablymore reliable because many more localities were sam-pled, so that the range of pairwise distances betweenindividuals is covered much better.

In order to derive estimates of s, we must nowattempt to estimate the density of mice, which we cando by using our trapping data. In 1998, the year withmost intense trapping, we underwent a random pros-pecting of farms, and found that 25% of them had mice.The average number of mice trapped per farm withmice was 7.2 (with a large variance, see Table 2).Because we intensively trapped in all cases, andstopped the effort only after several unsuccessful trap-ping nights, we believe this is a reasonable estimate ofthe mouse population present. It necessarily providesa lower limit of the number of mice existing at the timeof trapping, but it includes at least two generations (wecounted both adults and young), and so probably over-estimates population densities per generation. Thedensity of farms in the prospected areas was accu-rately determined by counting them on a map, leadingto an average of 3.1 farms km-2. This gives an esti-mated density D of 5.8 mice km-2. The average Ds2

estimated using the three different isolation bydistance methods ranged from 1.4 to 3.9, which givess = 0.51–0.82 km generation– . If our density esti-

)ar

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mates are really lower limits these estimates of s areupper limits, but they are compatible with the esti-mates derived from the cline analyses. With the lowestvalue, s = 0.51 km generation– , we estimate thatthe barrier would be created by a larger number(n = 120) of loci under weaker selection (s = 0.013) thanwhat is found when estimating migration from linkagedisequilibrium. The highest value of s = 0.82 givesn = 46 loci with s = 0.035. The results of the twoapproaches are comparable, given the numeroussources of uncertainty associated with these estimates.

DISCUSSION

For eight of the ten allozyme loci studied, the changesin allele frequencies can be best described by steepcentral steps of allele frequency that are coincidentand flanked by smooth tails of introgression on eitherside. This indicates the presence of a barrier to geneflow in the centre of the zone. We assumed that thisbarrier is caused by selection against hybrids, creatinga tension zone, and applied the existing theory to esti-mate the selection and migration parameters shapingthis hybrid zone.

As quantifying migration is essential to infer selec-tion, we also estimated migration independently fromthe cline analysis, using microsatellites and compar-ing populations of similar positions in the overall gra-dient of the hybrid zone. The results we obtained usingthe different approaches are in good agreement (s =0.5–0.8 km generation– ). There are few relevantdata in the literature with which to compare this esti-mate. Many capture–recapture experiments give dis-cordant results (Lidicker & Patton, 1987), and oftensuch studies concern small study areas and wouldmiss long-distance migrants (Baker, 1981; see Pocock,Hauffe & Searle, 2005, this issue). Myers (1974) wit-nessed colonization events between grids separated by92 m. According to Berry & Jakobson (1974) 25% ofthe mice on an island mate within 50–100 m fromtheir birthplace. Carlsen (1993) captured mice infarms and surrounding fields and reports movementsof up to 130 m. A few studies were able to detect longerdistance migrants. Walkowa, Adamczyk &Chelkowska (1989) trapped 6.5% of the mice living inan enclosure in surrounding fields, up to 300 m away.Cassaing & Croset (1985) recaptured most micewithin 50–480 m in outdoor populations. Auffray et al.(1990a) report on mice that had covered distances ofup to 700 m, and detected immigration from popula-tions up to 820 m apart. Berry (1968) describes morethan 30 mice moving 500 m and 14 moving 1500 m.Migrations over 1000 m are also reported by otherauthors (Pearson, 1963; Tomich, 1970). Although mostmice probably reproduce close to their birthplace, theycan apparently easily reach regions several hundreds

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of metres away. In addition, humans can passivelytransport them over much longer distances. InFigure 5 it can be seen that substantial linkage dis-equilibrium is sometimes found rather far from thecentre of the hybrid zone, and it could be due to suchlong-distance migrations. However, we found nomouse heterozygous at all loci in our dataset, and thusno F1 mice that could result from such long-distancemigrations. Potential first-generation backcrosses,heterozygous at half of their loci, were rare: only onewas found on the domesticus side of the zone (among151 mice from localities with a coordinate lower than6182 and with data available at eight loci or more). Sixsuch mice were found on the musculus side (among359 mice from localities with a coordinate above 6200),but only from the localities closest to the central stepof the hybrid zone (coordinates from 6200–6205).

The average barrier to gene flow appears relativelymoderate (20 km) compared with migration (0.5–0.8 km generation– ), but varies considerablybetween loci in our estimates, as well as the level ofintrogression and degree of asymmetry. Two of the lociwe studied (Idh and Sod) did not show the typical cen-tral step, but rather wide clines with extensive intro-gression into musculus, which could be an indicationthat they have escaped the central barrier. The delayto introgression of neutral alleles across such a barrieris expected to be of the order of 500–1600 generations(B2/s2; Barton & Hewitt, 1985, and referencestherein). There are presumably two reproduction peri-ods per year, one indoors in autumn that we observedduring our trapping campaigns and a second one inspring, mostly in the fields. This would, however, rep-resent only one generation a year in terms of migra-tion. According to what is believed about the patternsof expansion of these subspecies (reviewed in Boursot

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Figure 5. Average standardized linkage disequilibrium(R) between the eight loci of Fig. 2 in localities along thetransect. Only localities with eight mice or more wereselected. Crosses indicate the position of the centre of thehybrid zone.

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et al., 1993), being at the most northern end of thehybrid zone the transect in Jutland should be young-est. The few available archaeological records point toan earliest occurrence of house mice in north-westernEurope around 2000 years BP (Auffray, Vanlerberghe& Britton-Davidian, 1990b; Auffray & Britton-Davidian, 1992; Cucchi, Vigne & Auffray, 2005, thisissue). Unless the hybrid zone in Jutland is muchyounger than this, which cannot be formally excluded,there would have been time for neutral genes toescape the barrier. Our results also imply that the bar-rier created by this hybrid zone would be very weakagainst selectively favourable alleles [T = log(B2ps/2s2)/2s only about 100 years for a selective advantages of 1%]. It will be interesting to determine whethersuch selective introgression has occurred. Because theloci we studied were not chosen at random, but fortheir diagnosticity in this part of the hybrid zone, theyare not suited to address this issue.

With a number of approximations, our analyses pro-vide a rather low estimate of fitness for the centralpopulations (WH = 0.45). Indirect evidence of the lowerfitness of hybrid populations comes from theirobserved higher load of intestinal parasites (Sageet al., 1986a; Moulia et al., 1991) and the confirmationof the probable genetic origin of such differences(Moulia et al., 1993). More direct evidence comes fromthe observation of a slightly reduced testis size in cen-tral populations, as well as very high sterility rates ofmales and substantial reduction of female fertility inF1 crosses between Danish strains of the two subspe-cies (Britton-Davidian et al., 2005, this issue). How-ever, it has also been shown that the natural andartificial hybrids display a better general bilateralsymmetry of skull morphology, which was taken asevidence of heterosis on this character (Alibert et al.,1994, 1997; Auffray et al., 1996).

This relatively narrow hybrid zone is expected to becreated by a moderate number of loci, and if their num-ber was low enough, they could be searched by scan-ning the genome. The range of values of n that wederived (46–120) might appear to be rather high forthis to be feasible. However, it is based on the assump-tion of selection against heterozygotes, neglectingpotentially strong epistasis between loci. Further-more, in deriving our estimates of the number of lociunder selection, we used the parameters of the averagecline for eight loci. Averaging over loci whose clines dif-fer in width and symmetry (although asymmetry isalways in the same direction) probably tends to over-estimate the width, and thus the number of loci underselection (as does the approximation of r = 0.5 betweenthe studied and selected loci). The very limited intro-gression of Y and X chromosome markers in severaltransects (Tucker et al., 1992; Dod et al., 1993, 2005(this issue); Prager, Boursot & Sage, 1997) clearly

points to the possibility that at least some loci areunder strong enough selection to be detected using thisapproach. With progress in the knowledge of the mousegenome (Waterston et al., 2002), and the discovery ofnumerous nucleotide variations between the genomesof the house mouse subspecies (Wade et al., 2002), itshould be possible to identify the regions of the genomemost involved in selection in this hybrid zone (see Pay-seur & Nachman, 2005, this issue). Given the difficul-ties in fitting cline models to these data however, itmay prove difficult to identify the loci under selection.

An intriguing aspect of our cline shape analyses isthat the central step hardly overlaps frequency 0.5,but rather occurs almost entirely below this value, onthe domesticus side (particularly for Es10, Es2, Gpdand Pgm, Fig. 2). We wondered whether this couldresult from the presence of geographical barriers tomigration. The steep edged river valley just south ofthe inferred centre that is highlighted on the map(Fig. 1) could act as such. So could the large area ofheathland that occupies the area between the east-and westward-flowing rivers. There were only a fewscattered farms in this area and in spite of intensivetrapping efforts we were unable to find mice there,which is reflected by the absence of samples in thisarea (Fig. 1). These geographical accidents could com-bine to restrict gene flow in most of the region slightlysouth of the inferred centre, creating an abruptchange in allele frequencies. Depending on the fre-quencies reached south and north of this geographicalbarrier, these steps could either be confounded into asingle step by the inference method, producing a sym-metrical cline (Amy, Es1, Mpi and perhaps Np), or thegeographical rather than the genetic step could becaptured by the method, leading to asymmetric clineswith a step on the domesticus side (Es10, Gpd, Pgm).This would imply that the hybrid zone might be mov-ing south towards the geographical barrier. Loci nottoo tightly linked to selected loci would be laggingbehind the selection front, leaving long traces on themusculus side. Loci more tightly linked to selected lociwould tend to follow the front, and leave less trace. Asthe latter loci will reach higher musculus frequenciesnorth of the geographical barrier, they should tend toshow more introgression into domesticus, a counterin-tuitive conclusion because they are submitted tostronger selection. As the Idh and Sod loci appear tohave escaped the barrier to gene flow, the centre oftheir cline could represent the position of the originalcontact between the subspecies, about 8 km north ofthe present centre. If these loci are not subject toselection, the width of their clines depends only on thetime t elapsed since the initial contact between thesubspecies (w = 2.51s÷t). With an average width of54 km for these two loci, migration between 0.5 and0.8 km generation– , and considering one migration1 2/



episode a year (see above), we find that the initial con-tact would have occurred between 700 and 1800 yearsago, which is compatible with the archaeological data.This would represent an imperceptible movementduring the period that has elapsed since the Hunt &Selander (1973) study. However, the Y chromosomedata appear difficult to reconcile with this scenario.Because it appears to be the locus under strongestselection in this transect, its cline should have beenthe first to be trapped in the presumed density trough,which does not seem to be the case. Barriers to migra-tion such as the one we suspect here could occur else-where in Jutland. The complex interaction betweensuch barriers, genetic drift between the barriers andselection could uncouple the dynamics of epistaticallyinteracting loci, producing staggered clines, explain-ing the differences we see between loci. In such a sit-uation it is not necessary that the major transition ofallele frequency of neutral loci occur at 50%. ThreeRobertsonian (Rb) chromosomal fusions segregate inthe domesticus populations just south of the centre inthe transect we studied, and Fel-Clair et al. (1996)found that their frequencies decreased in a series ofstaggered clines before they could reach the centre ofthe hybrid zone. Experimental crosses have suggesteda complex negative interaction between the cen-tromeres of the two subspecies (Lenormand et al.,1997), and a centromeric marker on a chromosome notinvolved in a Rb fusion seemed, on the basis of thelimited sample available, to show no introgressioninto domesticus (Fel-Clair et al., 1998). It could thusbe that asymmetrical centromeric interactionscontribute to impede introgression into domesticus inthis transect, in addition to potential geographicalbarriers.

However, greater introgression into musculus forseveral diagnostic loci was noted in the broader studyof the Jutland hybrid zone by Hunt & Selander (1973),but also in other geographical regions, despite lessintense sampling (Bulgaria, Vanlerberghe et al., 1988;Bavaria, Tucker et al., 1992; East Holstein, Prageret al., 1993). This apparently general pattern calls fora general explanation. Fisher (1937) predicted that inthe wave of advance of advantageous alleles, the lead-ing edge should be longer than the trailing edge. Theasymmetry observed in the mouse hybrid zone couldthus witness a general movement of the zone in adirection opposite to that predicted above, and reflectan overall advantage of domesticus over musculus.Asymmetric behavioural selection would best accountfor such a situation. There is laboratory evidence thatmice have the capacity to discriminate urine odours oftheir own subspecies from those of the other (Smadja& Ganem, 2002; Ganem et al., 2005, this issue). Odourpreference tests performed on wild-derived mice fromthe tails of the Danish hybrid zone have revealed a

clear assortative choice in M. m. musculus, but nopreference in domesticus (Smadja, Catalan & Ganem,2004). The consequences of this behavioural differenceon gene flow are not straightforward, but one couldthink that it should prevent introgression into muscu-lus, rather than promote it. However, other behav-ioural asymmetries, such as a greater aggressivenessof domesticus males, could act in the opposite direc-tion. It has also been suggested that the choosiness ofthe musculus females could result from reinforcementin the hybrid zone (Smadja et al., 2004), in which casethis behaviour could be a consequence of the asymme-try of introgression, rather than its cause.

A comparison with patterns observed in other partsof this hybrid zone would be very informative, as itwould allow attribution of the confirmed patterns todeterministic forces, and the others to local historical,geographical or stochastic accidents.

ACKNOWLEDGEMENTS

Tonnes Nielsen provided invaluable support to thefieldwork. Patrick Makoundou performed most micro-satellite analyses. Many thanks to Guila Ganem forsharing some samples, and to the Danish farmers fortheir kind co-operation.

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APPENDIX 1

List of the 170 localities studied, with their number, name, year of collection, longitude, latitude (in UTM coordinates),coordinate on the transect (see text), number of mice studied and average hybrid index (% musculus alleles) for the eightloci with coinciding cline centres (Amy, Es1, Es10, Es2, Gpd, Mpi, Np and Pgm).

Locality no. Locality name Year Longitude Latitude Transect coordinate N HI eight loci

186 Ringtved 2000 522.1 6125.5 6143.5 1 0.000113 Simmersted 1985 525.0 6128.5 6146.8 2 0.031114 Simmersted 1990 525.0 6128.5 6146.8 14 0.005115 Simmersted A 1990 525.0 6128.5 6146.8 4 0.000181 Kastvra 2000 521.4 6129.1 6146.9 2 0.000182 Kastvra A 2000 522.2 6129.2 6147.2 14 0.006187 Sillerup 2000 535.7 6128.4 6147.3 7 0.057189 Slusen 2000 538.2 6133.7 6152.3 1 0.00092 Oedis 1984 524.4 6140.5 6158.6 13 0.11588 Maltbaek Mark 1992 505.1 6144.8 6160.5 4 0.12993 Oedis F 1985 526.3 6142.3 6160.7 2 0.12587 Maltbaek 1992 504.1 6145.3 6160.9 10 0.113184 Lauritzminde 2000 541.4 6144.1 6162.0 17 0.000175 Binderup 2000 535.1 6143.3 6162.2 3 0.000188 Skartved 2000 534.2 6145.9 6164.8 2 0.042190 Soender Stenderup 2000 538.7 6146.5 6164.9 1 0.000174 Baeklund 2000 516.2 6149.3 6166.4 33 0.047179 Gelballe 2000 522.6 6149.3 6167.1 5 0.000154 Vranderup 1998 524.8 6149.8 6167.9 3 0.18833 Gesten 1998 511.2 6154.7 6171.1 2 0.281133 Trelde 1998 549.1 6162.3 6172.9 1 0.143140 Vejlby 1998 545.4 6163.0 6176.2 2 0.14310 Baekke 1998 509.7 6160.6 6176.8 12 0.213151 Viuf 1998 531.7 6158.9 6177.7 9 0.136153 Vorbasse sk 1992 507.1 6162.2 6178.1 1 0.063101 Rands A 1998 543.7 6164.2 6178.4 1 0.000152 Vorbasse 1992 503.5 6163.2 6178.6 23 0.251100 Rands 1998 543.4 6164.8 6178.9 4 0.234124 Starup 1998 487.1 6166.8 6180.2 1 0.18824 Egtved 1 1990 519.5 6163.3 6180.7 1 0.1887 Ammistbol 1990 525.6 6165.5 6183.6 4 0.250128 Toerskind 1984 519.9 6166.9 6184.3 18 0.104129 Toerskind 1985 519.9 6166.9 6184.3 21 0.099130 Toerskind 1991 519.9 6166.9 6184.3 4 0.2038 Ammistbol A 1992 527.3 6166.5 6184.8 1 0.250137 Tufkaer Mark 1998 531.6 6166.1 6184.9 1 0.21494 Oedsted 1990 523.9 6167.3 6185.1 2 0.15695 Oedsted 1991 523.9 6167.3 6185.1 1 0.43896 Oedsted 1992 523.9 6167.3 6185.1 14 0.20114 Bindeballe 1992 517.6 6168.3 6185.4 5 0.225117 Skoldbjerg 1992 505.9 6171.9 6187.5 12 0.19852 Hojen Kirke 1998 531.0 6168.9 6187.6 13 0.17872 Jerlev V 1992 527.5 6170.2 6188.5 8 0.09071 Jerlev A 1998 528.0 6170.3 6188.6 5 0.118118 Smakaer 1998 520.4 6171.2 6188.6 2 0.344134 Troellund 1992 502.5 6173.8 6189.0 14 0.257102 Ravning 1992 521.9 6171.6 6189.1 10 0.405125 Store Lihme 1987 519.7 6172.3 6189.6 9 0.37599 Plougslund A 1992 505.9 6174.0 6189.7 12 0.28698 Plougslund 1992 506.0 6174.1 6189.7 17 0.26370 Jerlev 1998 528.0 6171.7 6190.0 1 0.643



73 Kaerbolling 1987 525.4 6172.7 6190.7 1 0.571145 Vilstrup 1992 529.4 6172.3 6190.8 3 0.27120 Bredsten 1985 523.2 6173.1 6190.8 5 0.51621 Bredsten 1987 523.2 6173.1 6190.8 2 0.46726 Englesholm Mark 1998 521.1 6173.6 6191.0 1 0.56313 Balle A 1987 523.7 6174.1 6191.9 1 0.43844 Grene Kirke 1998 504.4 6176.5 6191.9 6 0.33011 Balle 1992 522.8 6174.7 6192.3 17 0.60412 Balle 1998 522.8 6174.7 6192.3 2 0.643120 Soedover 1985 521.6 6175.0 6192.5 10 0.500121 Soedover 1992 521.6 6175.0 6192.5 1 0.50080 Lildfrost 1998 525.9 6174.8 6192.9 2 0.56315 Braendgaarde 1998 512.2 6176.7 6193.1 4 0.67245 Grinsted 1998 497.9 6178.9 6193.5 1 0.875142 Vester Hornstrup 1998 526.2 6175.9 6194.0 39 0.473138 Uhre 1998 531.6 6175.6 6194.4 2 0.625143 Vester Hornstrup A 1998 526.8 6176.2 6194.4 1 0.50031 Gammelby 1992 520.8 6177.4 6194.8 3 0.58332 Gammelby 1998 520.8 6177.4 6194.8 1 0.68825 Elkaer 1998 504.8 6179.5 6195.0 1 0.813106 Rugballe 1992 527.8 6176.8 6195.1 14 0.527107 Rugballe A 1992 527.7 6177.0 6195.3 1 0.68822 Brunbjerg 1998 502.8 6180.1 6195.3 1 0.68861 Hovertoft 1998 532.5 6176.5 6195.4 3 0.750108 Rugballe B 1991 528.0 6177.1 6195.4 3 0.708109 Rugballe B 1992 528.0 6177.1 6195.4 3 0.604110 Rugballe B 1998 528.0 6177.1 6195.4 3 0.58360 Hover Kirke A 1998 531.0 6176.7 6195.4 1 0.62581 Lille Hoegsholt 1998 531.0 6176.8 6195.4 1 0.87559 Hover Kirke 1998 529.9 6177.1 6195.6 2 0.68897 Oelgod 1998 478.9 6184.0 6196.3 1 0.313135 Trollerup 1992 522.7 6178.7 6196.3 3 0.458136 Trollerup 1998 522.7 6178.7 6196.3 2 0.71982 Lindeballe 1992 512.3 6180.3 6196.7 10 0.61367 Jelling bis 1987 526.1 6178.8 6196.9 3 0.87583 Lindeballe A 1992 513.0 6180.7 6197.1 1 0.43874 Kiddelund 1998 522.9 6179.5 6197.2 3 0.80489 Moelvang 1990 522.9 6179.6 6197.3 2 0.50090 Moelvang 1992 522.9 6179.6 6197.3 1 0.75066 Jelling 1990 526.5 6179.3 6197.3 3 0.81650 Hoerup 1990 530.2 6179.5 6198.0 8 0.67251 Hoerup 1991 530.2 6179.5 6198.0 25 0.71148 Hoegelund 1998 509.8 6182.1 6198.1 14 0.66847 Hjortlund 1998 504.2 6182.8 6198.2 10 0.63930 Gadbjerg 1987 520.5 6181.2 6198.6 1 0.66791 Norskov 1992 515.3 6182.1 6198.8 2 0.65678 Langelund S 1998 508.3 6183.2 6199.0 13 0.561131 Tofthoj 1987 521.2 6181.7 6199.1 1 0.500119 Smidstrup 1992 516.9 6182.4 6199.3 5 0.68829 Filskov A 1992 500.1 6184.5 6199.3 1 0.81316 Brandbjerg 1985 529.3 6181.2 6199.6 4 0.81717 Brandbjerg 1991 529.3 6181.2 6199.6 28 0.85318 Brandbjerg 1992 529.3 6181.2 6199.6 1 0.68863 Hygum 1990 527.4 6181.4 6199.6 1 0.813


APPENDIX 1 Continued



64 Hygum A 1998 526.0 6181.6 6199.6 2 0.90628 Filskov 1992 499.2 6185.1 6199.8 2 0.78119 Bredal 1998 538.0 6179.3 6199.9 5 0.68877 Langelund F 1998 504.3 6184.8 6200.2 7 0.73665 Hygum B 1998 527.7 6182.6 6200.8 17 0.857146 Vindelev M 1985 530.3 6182.6 6201.1 8 0.729147 Vindelev M 1987 530.3 6182.6 6201.1 1 0.857148 Vindelev M 1991 530.3 6182.6 6201.1 2 0.688149 Vindelev S 1991 530.1 6182.7 6201.2 16 0.740150 Vindelev S 1992 530.1 6182.7 6201.2 17 0.658111 Sandvad 1990 528.0 6183.5 6201.7 7 0.795112 Sandvad 1991 528.0 6183.5 6201.7 1 0.62584 Lindved 1990 536.1 6183.0 6202.4 4 0.828103 Riis A 1998 520.1 6185.5 6202.8 4 0.68836 Givskud Z 1987 522.0 6185.4 6202.9 6 0.86537 Givskud Z 1991 522.0 6185.4 6202.9 15 0.74638 Givskud Z 1998 522.0 6185.4 6202.9 26 0.80135 Givskud E 1991 522.0 6186.0 6203.5 5 0.8382 Aadel A 1991 528.7 6185.4 6203.7 1 0.4383 Aadel A 1998 528.7 6185.4 6203.7 4 0.71262 Hvejsel 1998 526.5 6185.7 6203.8 4 0.8139 Baastrup 1987 540.5 6182.6 6204.0 20 0.76934 Give Mark 1992 514.4 6187.5 6204.0 6 0.875116 Skaerhoved 1998 517.2 6187.4 6204.3 1 0.7501 Aadel 1990 529.0 6186.0 6204.4 3 0.813122 Soendersthoved 1998 510.5 6188.3 6204.4 4 0.810104 Riis 1998 519.9 6187.4 6204.6 4 0.813144 Vibjerg 1998 517.7 6188.1 6205.1 1 0.87546 Hedegaard 1998 510.8 6189.2 6205.3 1 0.8754 Aalsted Moelle 1991 529.2 6187.2 6205.6 4 0.7976 Aalsted Moelle 1998 529.2 6187.2 6205.6 9 0.762105 Riis Mark 1998 521.3 6188.4 6205.8 13 0.77785 Loesning 1984 544.5 6184.5 6208.1 15 0.78286 Loesning 1985 544.5 6184.5 6208.1 14 0.853127 Toerring Mark 1998 532.8 6192.2 6211.0 6 0.814126 Thyregod 1998 520.4 6195.0 6212.2 2 0.62553 Honum 1998 539.7 6193.5 6213.4 54 0.81376 Klovborg 1998 532.6 6198.2 6216.9 1 0.85727 Estrupholm 1998 518.4 6203.9 6220.8 2 0.938139 Underup 1998 543.0 6200.6 6221.2 30 0.869132 Traeden 1998 542.0 6202.7 6222.9 3 0.800141 Vestbirk 1998 545.5 6202.5 6223.9 28 0.797178 Gammel Hampen 2000 523.3 6207.2 6224.7 7 0.923196 Vester Gludsted 2000 518.0 6209.3 6226.2 1 0.750192 Store Noerlund 2000 514.9 6210.5 6227.0 5 0.95849 Hoejby 1992 575.4 6197.4 6240.7 11 0.95539 Gosmer F 1991 574.5 6199.1 6240.8 7 0.97341 Gosmer N 1991 574.7 6199.8 6241.3 21 0.94142 Gosmer N 1992 574.7 6199.8 6241.3 15 0.89643 Gosmer N 1998 574.7 6199.8 6241.3 16 0.871180 Gosmer N 2000 574.7 6199.8 6241.3 7 0.95740 Gosmer J 1991 575.7 6198.8 6241.6 5 0.96355 Hov 1985 576.6 6197.9 6241.9 6 0.99056 Hov 1987 576.6 6197.9 6241.9 18 0.959





57 Hov 1991 576.6 6197.9 6241.9 2 1.00058 Hov 1992 576.6 6197.9 6241.9 5 0.833123 Spoettrup 1998 577.5 6199.2 6243.4 14 0.931191 Spoettrup 2000 577.5 6199.2 6243.4 9 0.958177 Elleskovhuse 2000 552.3 6223.2 6245.7 28 0.94975 Klank 1985 556.8 6223.2 6247.4 10 0.925194 Tovstrup 2000 550.5 6226.0 6247.7 1 1.000197 Voel Oestermark 2000 545.5 6228.3 6248.7 2 1.000173 Abo 2000 564.5 6220.5 6248.8 10 0.965185 Ormslev 2000 566.7 6220.5 6250.1 7 0.974176 Borum Oestergaard 2000 563.9 6227.3 6254.3 4 0.957





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2012

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24,5

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25,5

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26,5

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27,6

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8,66

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30,8

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31,8

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32,8

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33,8

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35,2

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0200

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45,7

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50,6

6,99

9222

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76,1

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106

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0;

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9,11

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114,

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0100

00;

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00;

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00;

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10

813,

40,2

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2;

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222;

10

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2;

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10

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2;

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24,3

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2222

2122

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25,3

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; 108

26,3

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2222

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; 108

27,1

49,2

2002

1222

0; 1

0828

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,220

1212

120;

108

29,1

49,2

1022

0122

0; 1

0830

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2210

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2200

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11; 1

0833

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2210

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12;

1083

4,51

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0222

211;

10

835,

51,2

2002

2220

1;

1083

6,51

,220

0222

222;

10

837,

51,2

2002

2221

2;

1083

9,51

,221

0222

220;

10

840,

51,2

2102

2221

0;

1084

1,51

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0212

220;

10

842,

51,2

2002

0221

0;

1084

4,14

9,22

1121

1220

; 10

845,

149,

2201

2112

20;

1084

6,14

9,22

0120

2221

;

1084

7,14

9,22

1121

2220

; 10

848,

149,

2111

2011

20;

1084

9,14

9,21

0021

2220

; 10

850,

149,

2210

2121

20;

1085

1,14

9,99

9001

0990

; 10

852,

130,

2000

0100

00;

1085

4,13

0,02

0000

0000

; 10

855,

108,

2020

0021

21;

1085

6,35

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2002

121;

10

857,

37,2

0201

1212

1;

1085

8,35

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2102

211;

1085

9,35

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2102

222;

108

61,4

1,10

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41,2

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221;

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69,4

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41,1

2229

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29;

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2,41

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2999

229;

108

73,4

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874,

41,0

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1221

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22;

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22;

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1202

212;

108

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2010

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222;

108

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108

88,4

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108

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1000

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0001

200;

108

91,3

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22;

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108

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2100

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00;

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108

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2020

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2110

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2022

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2020

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1222

2220

; 109

07,1

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0110

1; 1

0908

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2001

221;

109

09,1

48,2

2211

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2; 1

0910

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1200

1220

1;

1091

1,14

9,21

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149,

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2022

21;

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149,

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2022

20;

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7,14

9,01

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2220

; 10

919,

17,2

2212

1222

2;

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0,17

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1212

221;

10

921,

17,1

2222

2221

2;

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2,17

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2221

211;

10

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17,2

2222

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220;

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57,2

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22;

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; 10

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; 10

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10;

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4,51

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211;

109

35,5

1,22

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; 10

936,

51,2

2012

2221

1;

1093

7,51

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1222

220;

109

38,1

7,21

2120

2221

; 10

939,

17,2

1112

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1; 1

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21;

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1,17

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221;

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2; 1

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; 10

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12;

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222;

109

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220;

109

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7,22

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; 10

977,

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2; 1

0978

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2000

0000

00;

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9,51

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1122

211;

110

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1,92

0112

2212

; 11

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51,9

2012

2220

0; 1

1052

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9201

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11;

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201;

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2000

2001

; 112

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2221

1101

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2100

2222

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10;

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1002

110;

11

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135,

2002

0120

01;

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0,22

2101

0210

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217,

150,

2211

0112

02;

1122

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2001

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221,

150,

2221

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00;

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0,10

0001

2200

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224,

106,

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96,2

2000

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2001

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;

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9,10

6,21

1101

0102

; 11

243,

106,

2020

0220

02;

1124

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2012

002;

11

245,

82,0

0110

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2;

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2022

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11

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0200

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00;

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98,0

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1000

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;

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4,98

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0000

102;

11

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98,0

1000

2000

2;

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6,13

4,20

1100

0200

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117,

2000

0000

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0002

00;

1126

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261,

117,

0200

0100

00;

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2200

1210

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264,

72,0

2000

0010

0;

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5,72

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0000

200;

1126

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0000

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11

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152,

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0000

202;

11

269,

134,

1900

0000

00;

1127

0,72

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100;

11

271,

88,0

9000

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1127

6,82

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2112

121;

11

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152,

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2,09

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2000

; 11

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142,

2000

1192

01;

1637

3,14

2,20

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9101

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374,

132,

2292

2199

92;

1637

5,14

2,10

1211

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376,

105,

1211

2012

21;

1637

7,10

5,21

2210

1122

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1637

8,80

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2121

100;

16

379,

142,

1012

2112

00;

1638

0,3,

2211

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12;

1638

1,38

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121;

16

382,

38,2

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3,10

4,22

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384,

105,

2222

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1638

5,77

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1012

221;

16

386,

46,2

2211

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2;

1638

7,13

9,90

9220

2992

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388,

154,

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0211

00;

1638

9,15

4,00

0102

1101

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390,

142,

2011

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01;

1639

1,14

4,22

1121

2220

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392,

26,2

0101

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0;

1639

3,14

2,20

1292

2102

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394,

142,

2121

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1639

5,10

5,22

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396,

105,

1221

2022

10;

1639

7,13

9,22

2220

2922

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398,

15,1

2122

2020

0;

1639

9,15

4,00

0100

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1640

0,14

2,11

2211

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401,

142,

1220

0111

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1640

2,6,

9991

2129

91;

1640

3,47

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202;

16

404,

60,2

1201

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1640

5,14

2,10

1122

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406,

80,2

1122

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1640

7,14

2,20

0200

2100

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408,

104,

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1640

9,14

2,10

1222

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410,

142,

2001

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1641

1,14

2,20

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412,

103,

2222

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1641

3,14

2,10

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414,

151,

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1641

5,15

1,00

1000

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416,

142,

2022

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00;

1641

7,10

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418,

104,

2102

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20;

1641

9,13

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420,

105,

2202

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21;

1642

1,12

3,22

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1642

2,10

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16

423,

126,

2111

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20;

1642

4,64

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16

425,

126,

1121

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11;

1642

6,10

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16

427,

105,

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1642

8,12

3,12

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429,

123,

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1643

0,12

3,22

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431,

123,

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1643

2,12

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1643

3,12

3,22

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434,

123,

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12;

1643

5,12

3,92

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436,

123,

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7,12

3,92

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438,

123,

9920

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1643

9,27

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16

440,

105,

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21;

1644

1,10

4,21

1212

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442,

81,2

2122

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2;

1644

3,11

0,02

2010

2000

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1644

4,10

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16

445,

10,0

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0;

1644

6,13

7,01

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447,

143,

9901

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1644

8,65

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2202

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16

449,

38,1

2212

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2;

1645

0,38

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16

451,

142,

2191

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1645

2,14

2,11

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453,

38,2

2922

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1645

4,38

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1645

5,38

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16

456,

142,

1012

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1645

7,97

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16

458,

142,

1001

1010

00;

1645

9,65

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2192

211;

16

461,

139,

2221

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10;

1646

2,11

8,21

2002

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463,

65,2

1212

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1;

1646

4,65

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2212

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16

465,

65,2

2221

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0;

1646

6,65

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220;

1646

7,13

2,22

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468,

124,

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1000

00;

1646

9,27

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16

470,

62,2

2122

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1647

1,62

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220;

16

472,

101,

9990

0009

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1647

3,15

1,01

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474,

151,

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1647

6,10

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16

478,

65,2

2212

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1;

1647

9,14

0,01

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9100

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1648

0,10

0,00

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481,

100,

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1648

2,10

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483,

100,

0100

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1648

4,10

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100;

16

485,

65,1

1221

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1648

6,65

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16

487,

61,1

1222

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2;

1648

8,65

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220;

16

489,

127,

9911

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19;

1649

0,13

3,00

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9000

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1649

1,10

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164

92,1

0,00

0029

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93,1

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94,1

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95,1

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96,1

0,01

0000

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97,7

4,22

1110

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98,1

18,1

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6499

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6500

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2021

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22; 1

6502

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165

03,3

3,01

0020

2000

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1650

4,33

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16

506,

151,

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00;

1650

7,15

1,00

0000

1200

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508,

151,

0000

1091

00;

1650

9,15

1,00

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9100

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510,

151,

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1011

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1651

1,13

8,12

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512,

61,1

1022

1222

2;

1651

3,61

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2212

220;

16

514,

139,

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2221

20;

1651

5,13

9,22

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1651

6,13

9,22

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517,

139,

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1651

8,13

9,22

2922

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519,

139,

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1652

0,13

9,22

2121

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521,

139,

2221

2120

22;

1652

2,13

9,21

2122

2120

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523,

139,

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22;

1652

4,13

9,22

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525,

139,

2122

2122

12;

1652

6,13

9,22

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9029

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1652

7,13

9,20

2099

9029

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528,

139,

2129

9991

19;

1652

9,14

1,22

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530,

139,

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2221

12;

1653

1,14

1,22

1299

9119

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532,

141,

2102

9991

19;

1653

3,14

1,22

0222

2202

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534,

141,

2212

2221

12;

1653

5,14

1,22

0222

2221

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536,

141,

2202

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19;

1653

7,14

1,22

0299

9219

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1653

8,14

1,22

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539,

141,

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2221

10;

1654

0,14

1,22

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541,

141,

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2221

11;

1654

2,14

1,22

0299

9109

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543,

141,

2212

2221

10;

1654

4,14

1,12

1222

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545,

141,

2212

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1654

6,14

1,22

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547,

141,

2202

2221

11;

1654

8,14

1,22

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1654

9,52

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0999

009;

165

50,7

1,00

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9009

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51,5

2,00

0099

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52,5

2,00

0099

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53,5

2,01

0099

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54,5

2,00

0099

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55,5

2,01

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56,5

2,00

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57,5

2,01

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58,1

27,2

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165

60,1

27,2

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9;

1656

1,12

7,12

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562,

127,

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1656

3,14

2,21

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564,

142,

2021

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01;

1656

5,14

2,21

1000

2200

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566,

142,

2101

0021

01;

1656

7,14

2,20

0200

2100

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568,

142,

2002

0221

00;

1656

9,14

2,20

1220

2210

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570,

142,

2002

0020

00;

1657

1,14

2,20

0299

9119

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1657

2,12

3,22

2299

9129

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73,1

23,2

2229

9922

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6574

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2221

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6575

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29; 1

6576

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2221

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29; 1

6577

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29; 1

6578

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29; 1

6579

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29; 1

6580

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29; 1

6581

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2929

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29; 1

6582

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2929

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29; 1

6583

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0929

9991

19;

1658

4,70

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1201

111;

165

85,5

2,09

2999

9209

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586,

52,0

9099

9911

9; 1

6587

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1929

9992

29;

1658

8,43

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229;

165

89,4

3,22

0999

9229

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590,

43,1

2009

9992

9; 1

6591

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1202

9999

29;

1659

2,43

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1999

929;

165

93,4

3,22

0222

2922

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594,

38,1

2029

9992

9; 1

6595

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2212

2022

12;

1659

6,38

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222;

165

97,3

8,20

2211

1212

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598,

38,2

2022

2192

2; 1

6599

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2202

2229

10;

1660

0,38

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929;

166

01,3

8,92

0222

2922

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602,

38,2

2022

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2; 1

6603

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0202

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20;

1660

4,38

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929;

166

05,3

8,22

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606,

38,2

2022

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0; 1

6607

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2202

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21;

1660

8,38

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2999

929;

166

09,1

42,1

0219

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9; 1

6610

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1002

901;

166

11,1

42,0

1119

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9; 1

6612

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1020

901;

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13,1

42,2

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6614

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166

15,1

5,19

0122

2210

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16,1

5,29

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17,3

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221;

166

18,3

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219;

166

19,6

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1221

121;

1662

0,6,

2229

9990

19;

1662

4,32

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119;

166

25,6

4,12

2210

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626,

65,2

1222

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6627

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2212

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10;

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8,65

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120;

166

29,6

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630,

65,2

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6631

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1663

2,65

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9909

210;

166

33,1

42,9

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5,17

3,22

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886,

174,

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00;

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7,17

8,92

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2992

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888,

184,

0009

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1688

9,18

5,22

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2992

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890,

185,

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22;

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1,17

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174,

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00;

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3,17

4,00

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9000

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894,

174,

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90;

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5,19

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1689

6,18

7,00

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897,

187,

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00;

1689

8,19

1,22

2199

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899,

191,

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1690

0,17

5,00

0999

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901,

184,

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2,19

2,92

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2992

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903,

174,

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00;

1690

4,17

4,00

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905,

182,

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1690

8,17

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1690

9,19

1,22

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910,

187,

2009

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09;

1691

1,18

7,00

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9009

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912,

191,

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29;

1691

3,18

7,00

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9009

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914,

189,

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09;

1691

5,17

7,22

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9999

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916,

177,

2229

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09;

1691

7,18

7,00

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918,

184,

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9900

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1691

9,18

4,00

0999

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1692

0,18

0,22

2999

9229

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921,

180,

2299

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99;

1692

2,19

1,22

2999

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923,

184,

0009

9900

00;

1692

4,18

4,00

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925,

184,

0099

9900

90;

1692

6,18

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927,

180,

2299

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1692

8,17

7,22

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929,

177,

2229

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22;

1693

0,17

7,22

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9229

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1693

1,17

7,22

2999

2229

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933,

184,

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9900

00;

1693

4,18

5,22

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935,

177,

2229

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29;

1693

6,17

7,22

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937,

174,

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9920

09;

1693

8,18

0,22

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9999

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939,

191,

2229

9922

22;

1694

1,17

3,22

2999

2922

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943,

192,

2229

9922

29;

1694

6,18

4,00

0999

0000

;

1694

9,18

4,00

0999

0000

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950,

179,

0009

9900

09;

1695

2,18

4,00

0999

0090

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953,

178,

2229

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29;

1695

4,19

0,00

0999

0000

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955,

188,

1009

9900

00;

1695

6,17

9,00

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9000

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958,

177,

2229

9922

22;

1695

9,18

2,00

0999

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960,

177,

2229

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29;

1696

1,17

7,22

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2222

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1696

2,19

7,92

9999

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963,

177,

2229

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29;

1696

4,18

5,92

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965,

185,

9299

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6,17

7,22

2999

9929

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967,

177,

2229

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22;

1696

8,18

0,22

2999

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969,

191,

2229

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29;

1697

0,18

2,00

0999

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971,

182,

0009

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00;

1697

2,18

2,00

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0000

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1697

3,18

2,00

0999

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974,

182,

1009

9900

00;

1697

5,18

1,00

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977,

184,

0009

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00;

1697

8,18

2,00

0999

0000

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979,

182,

0009

9900

00;

1698

0,18

2,00

0999

0000

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981,

182,

0009

9900

00;

1698

2,17

7,22

2999

2122

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983,

194,

9299

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99;

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4,17

5,00

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;

1698

5,18

4,00

0999

0000

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986,

184,

0009

9900

00;

1698

7,18

4,00

0999

0000

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988,

179,

0009

9990

09;

1698

9,17

4,02

0999

9009

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990,

174,

0009

9990

09;

1699

1,17

4,00

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9009

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993,

174,

0009

9990

09;

1699

4,17

4,00

0999

9000

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995,

174,

0009

9921

09;

1699

6,17

9,90

9999

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1699

7,17

4,90

9999

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998,

174,

9099

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99;

1699

9,18

2,00

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001,

177,

9299

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1700

2,17

7,22

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177,

2229

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29;

1700

4,17

7,92

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177,

2229

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29;

1700

6,17

7,22

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007,

178,

2229

9922

21;

1700

8,17

8,22

1999

2129

;

1700

9,17

6,22

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9222

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010,

173,

2229

9922

22;

1701

1,19

7,22

2999

2229

; 17

012,

185,

2299

9999

99;

1701

3,17

3,22

2999

2229

; 17

014,

185,

2299

9999

99;

1701

5,17

3,22

2999

9222

; 17

016,

173,

2229

9922

22;

1701

7,17

3,22

2999

2222

; 17

019,

182,

0009

0000

00;

1702

0,18

2,00

0000

0000

;

1702

1,17

4,90

0090

9000

; 17

022,

174,

9000

9090

00;

1702

6,17

4,90

0090

9000

; 17

027,

174,

9009

9999

09;

1702

8,17

4,00

0090

9000

; 17

029,

174,

0000

9090

00;

1703

0,17

4,20

0090

9000

; 17

031,

176,

9222

9292

22;

1703

2,17

4,20

0090

9000

; 17

033,

174,

0000

9090

00;

1703

4,17

4,00

0090

9000

;

1703

5,17

4,00

0090

9000

; 17

036,

174,

2000

9090

00;

1703

7,17

7,02

2292

9222

; 17

038,

177,

0222

9292

22;

1703

9,17

7,02

2292

9222

; 17

040,

177,

2222

9292

22;

1704

1,17

7,02

2292

9222

; 17

042,

177,

0222

9292

22;

1704

3,17

7,02

2292

9222

; 17

044,

186,

0000

9000

00;

1704

5,18

7,90

9000

9990

;

1704

9,18

8,00

0999

0000

; 17

050,

173,

9222

2299

22;

1705

1,17

4,90

0000

9900

; 17

052,

174,

9000

0099

00;

1705

3,17

4,90

0000

9900

; 17

054,

174,

2000

0090

00;

1705

5,17

4,90

0000

9900

; 17

056,

174,

9000

0099

00;

1705

7,17

8,22

1229

2220

; 17

058,

196,

2221

2299

02;

1705

9,19

2,99

2999

9229

;

1706

0,19

2,92

2222

9920

; 17

061,

175,

0009

9900

00;

1706

2,18

0,92

2092

9922

; 17

063,

191,

2222

2222

22;

1706

4,17

8,92

9291

2992

; 17

065,

178,

2201

9022

22;

1706

6,17

3,22

2199

2222

; 17

067,

179,

0000

9090

00;

1706

8,18

1,00

0999

0000

; 17

069,

184,

0009

9900

00;

1707

0,18

4,00

0999

0000

;

1707

5,17

6,02

2999

2229

;



AP

PE

ND

IX 3

Th

e ge

not

ypes

of t

he

621

mic

e te

sted

for

six

mic

rosa

tell

ite

loci

. For

eac

h m

ouse

(mic

e se

para

ted

by s

emic

olon

s) w

e gi

ve s

ucc

essi

vely

(sep

arat

ed b

y co

m-

mas

): m

ouse

nu

mbe

r, lo

cali

ty n

um

ber

(as

in A

ppen

dix

1), s

ampl

ing

year

, an

d th

e ge

not

ypes

at

the

six

mic

rosa

tell

ite

loci

in t

he

orde

r li

sted

in t

he

firs

tre

cord

(an

d re

pres

ente

d fo

r ea

ch lo

cus

by t

he

con

cate

nat

ion

of

the

size

s of

th

e tw

o al

lele

s, w

ith

th

ree

digi

ts f

or e

ach

all

ele,

e.g

. 050

058

mea

nin

g a

het

-er

ozyg

ote

for

alle

les

of s

izes

50

and

58).

Un

dete

rmin

ed g

enot

ypes

are

cod

ed 0

0000

0

mou

sen

o.,L

ocn

o.,y

ear,

Ktr

2,G

fap,

D15

Mit

16,C

kmm

,Cyp

la2,

D17

Mit

41;

1166

1,42

,199

2,03

8038

,071

077,

0620

66,0

6506

5,06

6066

,000

000;

11

662,

42,1

992,

0380

38,0

7107

1,05

4070

,075

075,

0500

66,0

0000

0;

1166

8,42

,199

2,03

8038

,071

071,

0620

66,0

6006

5,05

0050

,000

000;

1166

9,42

,199

2,03

8038

,071

071,

0540

66,0

6006

5,06

6066

,000

000;

11

670,

42,1

992,

0380

38,0

7107

1,05

4066

,060

065,

0660

74,0

0000

0;

1167

1,42

,199

2,03

8038

,071

071,

0660

66,0

6506

5,05

0058

,046

046;

11

672,

42,1

992,

0380

38,0

7707

7,05

8066

,065

075,

0660

66,0

0000

0;

1167

3,42

,199

2,03

8038

,071

077,

0620

70,0

7507

5,05

0074

,000

000;

11

674,

42,1

992,

0380

38,0

7107

7,06

6066

,065

075,

0660

70,0

0000

0;

1167

7,42

,199

2,03

8038

,071

071,

0660

66,0

6006

5,05

0066

,000

000;

11

681,

42,1

992,

0380

38,0

7107

1,05

4078

,065

065,

0540

66,0

0000

0;

1168

3,42

,199

2,03

8038

,071

077,

0540

66,0

7507

5,05

0066

,000

000;

11

697,

42,1

992,

0380

38,0

7107

7,05

4054

,075

075,

0620

66,0

0000

0;

1169

8,42

,199

2,03

8038

,071

071,

0580

66,0

6506

5,05

0066

,000

000;

11

699,

42,1

992,

0380

38,0

7107

7,06

6066

,065

065,

0660

70,0

4604

6;

1170

7,42

,199

2,03

8038

,071

071,

0620

66,0

6006

5,05

0066

,000

000;

11

536,

49,1

992,

0380

38,0

7107

7,06

2062

,060

065,

0500

54,0

0000

0;

1153

7,49

,199

2,03

8038

,068

071,

0580

62,0

6506

5,05

4066

,000

000;

11

538,

49,1

992,

0380

38,0

7107

1,05

8070

,065

075,

0540

54,0

0000

0;

1153

9,49

,199

2,03

8038

,068

071,

0540

58,0

6507

0,05

0058

,046

046;

11

540,

49,1

992,

0380

38,0

7107

7,07

0070

,065

070,

0540

54,0

0000

0;

1154

1,49

,199

2,03

8038

,068

077,

0580

62,0

6507

0,05

8058

,000

000;

11

542,

49,1

992,

0380

38,0

7107

1,05

4078

,065

070,

0540

62,0

0000

0;

1154

3,49

,199

2,03

8038

,071

077,

0660

66,0

6506

5,05

0062

,000

000;

11

544,

49,1

992,

0380

46,0

7107

1,05

8062

,065

065,

0540

66,0

0000

0;

1154

5,49

,199

2,03

8038

,071

077,

0700

70,0

7507

5,05

0058

,000

000;

11

546,

49,1

992,

0380

38,0

6807

1,05

8070

,065

065,

0540

54,0

0000

0;

1127

8,69

,199

2,04

6046

,071

071,

0580

58,0

5506

5,04

2054

,050

050;

11

490,

69,1

992,

0460

46,0

4407

1,05

8058

,055

055,

0420

54,0

5005

0;

1149

1,69

,199

2,04

6046

,044

071,

0580

58,0

5505

5,04

2054

,000

000;

11

492,

69,1

992,

0420

46,0

4404

4,05

8058

,055

055,

0540

54,0

5005

0;

1149

3,69

,199

2,04

6046

,044

044,

0580

58,0

5505

5,04

2042

,048

050;

11

547,

69,1

992,

0460

50,0

7107

1,05

8058

,055

055,

0700

70,0

4805

0;

1154

8,69

,199

2,04

6050

,044

044,

0580

58,0

5505

5,04

2054

,050

050;

11

551,

69,1

992,

0460

46,0

4407

1,05

8062

,055

055,

0420

70,0

0000

0;

1155

2,69

,199

2,04

6046

,044

071,

0580

58,0

5506

5,04

2070

,048

050;

11

554,

69,1

992,

0460

46,0

7107

1,05

8058

,055

055,

0420

42,0

5005

0;

1155

5,69

,199

2,04

6046

,044

044,

0580

58,0

5505

5,04

2054

,000

000;

11

556,

69,1

992,

0460

46,0

4407

1,05

8058

,055

055,

0420

70,0

5005

0;

1155

7,69

,199

2,04

6046

,044

071,

0580

58,0

5505

5,05

4070

,000

000;

11

558,

69,1

992,

0460

46,0

4407

1,05

8058

,055

055,

0540

70,0

0000

0;

1166

3,69

,199

2,04

6046

,044

065,

0580

58,0

5505

5,04

2054

,000

000;

11

664,

69,1

992,

0460

46,0

4406

5,05

8058

,055

055,

0420

54,0

0000

0;

1166

6,69

,199

2,04

6046

,071

071,

0580

58,0

5505

5,05

4054

,000

000;

11

675,

69,1

992,

0420

46,0

4407

1,05

8058

,055

055,

0420

54,0

0000

0;

1167

6,69

,199

2,04

6050

,044

071,

0500

58,0

5505

5,05

4070

,000

000;

11

679,

69,1

992,

0420

46,0

4407

1,05

8058

,055

055,

0420

54,0

0000

0;

1168

0,69

,199

2,04

2046

,044

071,

0580

58,0

5506

5,05

4054

,000

000;

11

684,

69,1

992,

0460

46,0

4407

1,05

8058

,055

055,

0540

54,0

0000

0;

1169

0,69

,199

2,04

6046

,044

071,

0580

58,0

5505

5,04

2054

,050

050;

11

691,

69,1

992,

0420

46,0

4407

1,05

8058

,055

055,

0420

54,0

5005

0;

1169

3,69

,199

2,04

2046

,044

071,

0580

58,0

5505

5,05

4054

,000

000;

11

701,

69,1

992,

0460

46,0

4404

4,05

8058

,055

055,

0420

42,0

0000

0;

1170

4,69

,199

2,04

6050

,044

044,

0580

58,0

5505

5,05

4070

,000

000;

11

705,

69,1

992,

0460

46,0

4404

4,05

8058

,055

055,

0420

54,0

0000

0;

1170

6,69

,199

2,04

6046

,044

071,

0580

58,0

5505

5,04

2054

,000

000;

11

708,

69,1

992,

0460

46,0

4407

1,05

8058

,055

065,

0420

54,0

0000

0;

1170

9,69

,199

2,04

2046

,071

071,

0580

58,0

5505

5,04

2042

,000

000;

11

711,

69,1

992,

0460

46,0

4407

1,05

8058

,055

055,

0420

70,0

0000

0;

1124

2,72

,199

2,05

0050

,044

053,

0500

50,0

6506

5,04

2046

,048

048;

11

264,

72,1

992,

0500

50,0

5305

6,05

0066

,055

065,

0460

46,0

4805

0;

1126

5,72

,199

2,05

0050

,053

053,

0500

50,0

5506

5,04

2042

,000

000;

11

266,

72,1

992,

0500

50,0

5305

6,05

0066

,065

065,

0420

46,0

5005

0;

1127

0,72

,199

2,05

0050

,053

056,

0500

66,0

5506

5,04

6046

,048

050;

11

291,

72,1

992,

0500

50,0

5605

6,05

0066

,065

065,

0420

42,0

4805

0;

1148

3,72

,199

2,05

0050

,053

056,

0500

50,0

5506

5,04

2046

,048

048;

11

501,

72,1

992,

0500

50,0

4405

6,05

0050

,055

065,

0460

46,0

4805

0;

1158

9,72

,199

2,04

2050

,044

065,

0620

62,0

5505

5,05

4054

,000

000;

11

520,

87,1

992,

0500

50,0

5005

0,05

4062

,055

060,

0000

00,0

5005

0;

1152

8,87

,199

2,05

0050

,050

056,

0500

50,0

0000

0,05

0050

,048

050;

11

534,

87,1

992,

0500

50,0

5005

3,05

8058

,050

055,

0500

50,0

5005

0;

1161

0,87

,199

2,03

8038

,050

056,

0620

62,0

5005

5,03

8058

,050

052;

11

611,

87,1

992,

0500

50,0

5005

0,05

0058

,055

055,

0380

58,0

4604

6;

1163

6,87

,199

2,05

0050

,044

044,

0460

46,0

5505

5,04

6050

,050

050;

11

700,

87,1

992,

0500

50,0

4405

3,05

0050

,070

075,

0420

46,0

5005

0;

1170

3,87

,199

2,05

0054

,044

053,

0460

50,0

5005

5,04

2046

,050

050;

11

484,

96,1

992,

0500

50,0

6807

1,00

0000

,050

055,

0000

00,0

5005

2;

1148

8,96

,199

2,05

0050

,053

071,

0000

00,0

5505

5,03

8038

,052

052;

11

502,

96,1

992,

0420

42,0

6807

1,05

8058

,050

060,

0000

00,0

5005

2;

1150

3,96

,199

2,04

6054

,050

053,

0580

58,0

5505

5,00

0000

,050

050;

11

512,

96,1

992,

0420

42,0

5305

3,00

0000

,060

070,

0000

00,0

5005

2;

1151

3,96

,199

2,05

0050

,053

053,

0460

46,0

5505

5,05

0050

,052

052;

11

519,

96,1

992,

0500

50,0

6806

8,05

4058

,055

060,

0500

74,0

5005

0;

1152

1,96

,199

2,05

0050

,053

053,

0580

58,0

5505

5,03

8038

,050

052;

11

530,

96,1

992,

0500

54,0

7107

1,05

4058

,060

060,

0380

50,0

5205

2;

1153

2,96

,199

2,04

6050

,053

068,

0580

58,0

5505

5,03

8050

,050

050;

11

629,

96,1

992,

0380

46,0

5605

6,05

4058

,065

065,

0460

54,0

5005

0;

1163

8,96

,199

2,04

2050

,044

068,

0540

58,0

5005

5,05

0050

,052

052;

11

642,

96,1

992,

0420

50,0

6807

1,05

4058

,055

070,

0380

42,0

5005

0;

1125

2,98

,199

2,05

4054

,053

053,

0620

62,0

5506

0,05

0054

,050

050;

11

254,

98,1

992,

0500

50,0

5305

6,05

8066

,055

055,

0420

54,0

5005

0;

1125

5,98

,199

2,05

4054

,050

050,

0500

58,0

5505

5,05

0054

,050

052;

11

268,

98,1

992,

0540

54,0

5005

6,05

0066

,055

060,

0380

50,0

5005

0;

1129

5,98

,199

2,04

6046

,056

056,

0580

62,0

5505

5,05

0058

,050

050;

11

515,

98,1

992,

0540

54,0

5607

1,05

0066

,055

055,

0500

54,0

4605

8;

1152

5,98

,199

2,05

0054

,053

053,

0500

58,0

5506

0,04

2050

,046

058;

11

565,

98,1

992,

0540

54,0

5005

9,04

6050

,050

060,

0500

54,0

0000

0;

1159

5,98

,199

2,03

8042

,050

056,

0500

66,0

5506

0,04

2042

,000

000;

11

596,

98,1

992,

0500

50,0

5305

3,05

0058

,055

060,

0500

50,0

0000

0;

1159

7,98

,199

2,04

6054

,050

053,

0500

58,0

5505

5,05

4058

,000

000;

11

598,

98,1

992,

0500

54,0

5607

1,05

8062

,055

055,

0380

54,0

0000

0;

1159

9,98

,199

2,05

0054

,053

071,

0500

58,0

5506

0,04

2042

,000

000;

11

600,

98,1

992,

0500

54,0

5307

1,05

0058

,055

055,

0420

42,0

0000

0;

1161

3,98

,199

2,05

4054

,050

071,

0580

66,0

5505

5,05

0054

,000

000;

11

615,

98,1

992,

0540

54,0

5007

1,05

0062

,060

060,

0500

50,0

0000

0;

1162

3,98

,199

2,05

0050

,062

071,

0460

70,0

5505

5,04

6050

,000

000;

11

585,

99,1

992,

0540

54,0

5005

6,05

0050

,050

060,

0460

50,0

0000

0;

1158

6,99

,199

2,05

4054

,050

056,

0460

50,0

5006

0,04

6050

,050

050;

11

587,

99,1

992,

0460

54,0

7107

4,05

0054

,055

055,

0380

54,0

5005

0;

1159

1,99

,199

2,05

4054

,050

056,

0460

66,0

5505

5,05

0054

,050

050;

11

593,

99,1

992,

0380

38,0

7108

0,04

6058

,055

055,

0500

58,0

0000

0;

1159

4,99

,199

2,05

4054

,056

071,

0500

50,0

5505

5,04

6050

,050

050;

11

614,

99,1

992,

0540

54,0

5605

9,04

6050

,050

055,

0500

50,0

5005

0;

1161

6,99

,199

2,03

8038

,050

059,

0500

66,0

5505

5,05

0054

,050

050;

11

617,

99,1

992,

0540

54,0

5907

1,04

6050

,050

060,

0460

50,0

5005

0;

1161

8,99

,199

2,03

8038

,050

059,

0500

66,0

5505

5,05

0054

,050

050;

11

632,

99,1

992,

0540

54,0

5005

9,04

6066

,055

055,

0500

54,0

5005

0;

1163

4,99

,199

2,05

4054

,050

071,

0500

66,0

5506

0,04

6054

,050

050;

11

568,

102,

1992

,050

050,

0500

50,0

5405

8,05

5065

,054

054,

0000

00;

1125

7,11

7,19

92,0

5405

4,05

0050

,050

062,

0550

60,0

5005

0,05

0050

; 11

258,

117,

1992

,050

050,

0500

53,0

5005

0,05

0055

,038

050,

0500

56;

1125

9,11

7,19

92,0

4205

4,05

3053

,050

054,

0550

55,0

4604

6,05

2052

; 11

260,

117,

1992

,050

054,

0530

53,0

5405

8,05

0060

,038

046,

0480

52;

1126

1,11

7,19

92,0

5005

4,05

3053

,050

058,

0500

55,0

3805

4,04

8052

; 11

262,

117,

1992

,050

054,

0530

53,0

5406

2,05

0055

,038

050,

0480

50;

1152

4,11

7,19

92,0

5005

0,05

3053

,058

062,

0500

55,0

3804

2,04

8052

; 11

526,

117,

1992

,038

038,

0000

00,0

0000

0,05

5055

,000

000,

0000

00;

1152

7,11

7,19

92,0

5005

4,05

3053

,054

054,

0550

60,0

3804

2,00

0000

; 11

567,

117,

1992

,050

050,

0000

00,0

5005

0,05

0055

,038

042,

0500

50;

1163

7,11

7,19

92,0

5005

4,05

3053

,054

062,

0550

60,0

4204

2,05

2052

; 11

657,

117,

1992

,050

054,

0530

53,0

5005

0,05

5055

,046

050,

0500

52;

1165

9,11

7,19

92,0

5005

4,04

4050

,050

058,

0500

55,0

4605

0,05

0052

; 11

269,

134,

1992

,050

050,

0500

50,0

5005

0,05

5060

,050

050,

0000

00;

1156

2,13

4,19

92,0

5005

4,05

3053

,046

050,

0550

60,0

4604

6,05

0050

; 11

563,

134,

1992

,050

050,

0500

50,0

5005

0,05

5060

,042

046,

0000

00;

1156

4,13

4,19

92,0

5005

0,05

0050

,050

050,

0550

60,0

4204

6,04

8048

; 11

574,

134,

1992

,046

050,

0500

56,0

5005

0,05

5065

,046

046,

0500

50;

1157

5,13

4,19

92,0

5005

0,05

0050

,054

054,

0550

60,0

4605

0,05

2052

; 11

576,

134,

1992

,050

050,

0500

50,0

6206

2,05

5055

,000

000,

0500

50;

1157

7,13

4,19

92,0

5005

0,05

0050

,000

000,

0550

60,0

0000

0,05

0050

; 11

578,

134,

1992

,050

050,

0500

50,0

0000

0,05

5055

,042

054,

0500

50;

1157

9,13

4,19

92,0

5005

0,05

0050

,050

050,

0550

60,0

4605

0,05

0050

; 11

580,

134,

1992

,038

050,

0500

53,0

5805

8,05

5055

,058

058,

0500

50;

1126

3,15

0,19

92,0

3803

8,05

3071

,062

062,

0550

70,0

0000

0,05

0050

; 11

489,

150,

1992

,038

038,

0710

77,0

0000

0,05

5055

,000

000,

0500

50;

1158

1,15

0,19

92,0

3803

8,05

3053

,062

062,

0650

70,0

5805

8,05

0056

; 11

582,

150,

1992

,050

050,

0470

47,0

0000

0,05

5070

,058

062,

0500

56;

1158

3,15

0,19

92,0

0000

0,00

0000

,000

000,

0650

70,0

5805

8,04

6056

; 11

584,

150,

1992

,038

038,

0530

53,0

5405

4,06

5070

,058

070,

0460

52;

1160

4,15

0,19

92,0

5005

0,06

8071

,054

054,

0550

70,0

5805

8,04

6058

; 11

606,

150,

1992

,038

038,

0710

71,0

5405

4,05

5070

,058

058,

0500

56;

1164

9,15

0,19

92,0

3805

4,07

1071

,050

062,

0550

70,0

5805

8,05

0056

; 11

267,

152,

1992

,042

042,

0530

53,0

5805

8,05

5055

,000

000,

0500

50;

1127

7,15

2,19

92,0

5005

4,05

3053

,046

050,

0550

55,0

5405

4,05

0050

; 11

279,

152,

1992

,042

054,

0530

53,0

5005

8,05

0050

,046

050,

0500

50;

1128

0,15

2,19

92,0

5405

4,05

3056

,054

062,

0600

60,0

4605

0,00

0000

; 11

281,

152,

1992

,050

054,

0530

53,0

5005

0,05

0055

,046

050,

0500

50;

1128

2,15

2,19

92,0

5405

4,05

3056

,054

062,

0600

60,0

5005

0,05

0050

; 11

283,

152,

1992

,050

050,

0530

53,0

0000

0,05

0055

,046

050,

0500

50;

1128

4,15

2,19

92,0

5405

4,00

0000

,000

000,

0500

55,0

0000

0,05

0050

; 11

285,

152,

1992

,050

054,

0530

53,0

0000

0,05

0060

,046

054,

0500

50;

1151

4,15

2,19

92,0

5005

0,05

3053

,050

050,

0500

60,0

5005

4,05

0050

; 11

516,

152,

1992

,050

050,

0530

53,0

5005

8,05

0055

,050

054,

0500

50;

1151

7,15

2,19

92,0

5405

4,05

3053

,046

058,

0500

55,0

5005

4,05

0050

; 11

531,

152,

1992

,054

054,

0530

53,0

4605

8,06

5065

,000

000,

0000

00;

1158

8,15

2,19

92,0

5005

0,05

3053

,050

050,

0550

55,0

4605

4,05

0050

; 11

590,

152,

1992

,050

054,

0530

53,0

5005

0,05

5055

,000

000,

0500

50;

1161

9,15

2,19

92,0

4604

6,05

3053

,054

058,

0550

55,0

4605

0,05

0050

; 11

620,

152,

1992

,050

050,

0530

53,0

5005

0,05

0055

,054

054,

0500

50;

1162

1,15

2,19

92,0

4604

6,05

0053

,054

062,

0550

60,0

4206

2,04

8050

; 11

624,

152,

1992

,050

050,

0500

53,0

5006

2,05

5055

,042

046,

0480

50;

1168

6,15

2,19

92,0

5005

0,05

3053

,050

058,

0550

60,0

5005

0,05

0050

; 11

687,

152,

1992

,050

050,

0530

53,0

4605

0,06

5070

,046

054,

0500

50;

1168

9,15

2,19

92,0

5005

4,05

3056

,046

050,

0650

65,0

5005

0,05

2052

; 11

696,

152,

1992

,050

050,

0530

53,0

5005

0,06

5070

,046

050,

0500

50;

1170

2,15

2,19

92,0

5005

0,05

3053

,050

050,

0650

65,0

5005

4,05

0050

; 11

230,

157,

1992

,038

038,

0710

77,0

6206

6,07

5075

,066

070,

0460

46;

1123

1,15

7,19

92,0

3803

8,07

1071

,058

066,

0650

65,0

5405

4,05

8058

; 11

232,

157,

1992

,038

038,

0680

71,0

5807

8,06

5070

,066

066,

0580

58;

1123

3,15

7,19

92,0

3803

8,07

1071

,054

066,

0600

60,0

6607

4,00

0000

; 11

234,

157,

1992

,038

038,

0710

71,0

6607

8,06

5065

,066

070,

0000

00;

1123

6,15

7,19

92,0

3803

8,07

1071

,066

066,

0650

70,0

6607

0,00

0000

; 11

237,

157,

1992

,038

038,

0710

77,0

6207

8,06

5075

,066

070,

0000

00;

1123

8,15

7,19

92,0

3803

8,07

1077

,054

070,

0600

65,0

5406

6,00

0000

; 11

239,

157,

1992

,038

038,

0770

77,0

7007

8,06

5075

,066

070,

0000

00;

16

303,

3,19

98,0

3803

8,07

4074

,066

074,

0650

65,0

6606

6,04

6058

; 16

380,

3,19

98,0

3803

8,07

1080

,066

074,

0650

65,0

5005

8,00

0000

;

1661

7,3,

1998

,038

038,

0680

80,0

6206

2,05

0065

,058

062,

0460

56;

16

618,

3,19

98,0

3803

8,05

9083

,050

070,

0650

65,0

6206

2,04

6046

; 16

191,

6,19

98,0

3803

8,05

6056

,074

074,

0500

65,0

6206

6,04

6058

; 16

192,

6,19

98,0

3803

8,07

1074

,070

074,

0650

70,0

7007

4,04

6046

;

1620

2,6,

1998

,038

038,

0590

71,0

6207

4,06

5070

,050

066,

0460

46;

16

228,

6,19

98,0

3805

0,07

1074

,058

074,

0600

65,0

5007

4,05

2056

; 16

271,

6,19

98,0

3803

8,04

4044

,070

074,

0650

65,0

6607

0,04

6050

; 16

307,

6,19

98,0

3805

0,07

4077

,058

074,

0650

65,0

5807

0,05

6056

;

1640

2,6,

1998

,038

038,

0770

83,0

5406

6,06

5065

,058

070,

0000

00;

16

619,

6,19

98,0

3803

8,07

4077

,058

062,

0650

65,0

7007

4,05

6056

;

1662

0,6,

1998

,038

038,

0710

83,0

5806

6,05

5065

,066

070,

0460

58;

16

422,

10,1

998,

0500

50,0

5305

3,05

8058

,055

055,

0380

38,0

5005

2;

1642

6,10

,199

8,03

8054

,053

053,

0540

62,0

5505

5,03

8038

,050

050;

1644

4,10

,199

8,05

0050

,053

053,

0580

66,0

5505

5,03

8038

,050

050;

1644

5,10

,199

8,05

0054

,053

053,

0580

66,0

5505

5,03

8054

,050

050;

1647

6,10

,199

8,05

0050

,053

053,

0580

66,0

5505

5,03

8038

,050

052;



1648

4,10

,199

8,05

0050

,053

053,

0580

66,0

5505

5,04

2054

,050

052;

16

491,

10,1

998,

0500

50,0

5305

3,05

8066

,055

055,

0380

54,0

5005

0;

1649

2,10

,199

8,05

0054

,053

053,

0580

66,0

5505

5,03

8038

,050

052;

16

493,

10,1

998,

0380

54,0

5305

3,05

4066

,050

055,

0380

54,0

5205

2;

1649

4,10

,199

8,04

2050

,053

053,

0580

58,0

5005

5,03

8038

,050

052;

16

495,

10,1

998,

0500

50,0

4405

3,05

8066

,055

055,

0500

54,0

5005

2;

1649

6,10

,199

8,03

8050

,053

053,

0540

58,0

5006

0,03

8050

,050

052;

16

188,

12,1

998,

0500

50,0

5005

0,06

6066

,050

050,

0500

50,0

5005

2;

1618

9,12

,199

8,05

0050

,050

050,

0540

66,0

5505

5,05

8058

,000

000;

16

366,

15,1

998,

0380

38,0

7407

4,06

6078

,065

065,

0540

54,0

5005

0;

1639

8,15

,199

8,03

8038

,050

050,

0700

78,0

6506

5,05

4054

,000

000;

16

615,

15,1

998,

0380

38,0

5007

4,06

6066

,065

065,

0540

54,0

5005

8;

1661

6,15

,199

8,03

8038

,050

050,

0700

78,0

6506

5,05

8074

,050

050;

16

267,

19,1

998,

0380

38,0

4404

4,06

2062

,065

065,

0740

74,0

4605

6;

1626

8,19

,199

8,03

8038

,050

071,

0620

62,0

6507

0,07

0070

,046

046;

16

273,

19,1

998,

0460

46,0

4404

4,00

0000

,050

050,

0000

00,0

5005

0;

1627

5,19

,199

8,00

0000

,071

071,

0660

66,0

6507

0,07

0074

,046

046;

16

302,

19,1

998,

0380

38,0

4407

1,05

4062

,065

065,

0580

62,0

4604

6;

1624

0,22

,199

8,04

2046

,071

071,

0540

58,0

6006

5,06

6074

,000

000;

16

237,

25,1

998,

0380

38,0

7107

1,06

2062

,050

065,

0580

74,0

4605

0;

1639

2,26

,199

8,03

8050

,056

080,

0580

66,0

5005

0,00

0000

,000

000;

16

439,

27,1

998,

0380

38,0

6807

1,06

6066

,060

060,

0620

62,0

4605

8;

1646

9,27

,199

8,03

8038

,056

077,

0660

70,0

6506

5,06

2070

,046

046;

16

624,

32,1

998,

0380

50,0

6806

8,07

4078

,050

065,

0500

58,0

4604

6;

1650

3,33

,199

8,04

2042

,059

065,

0580

58,0

5005

5,05

0058

,050

050;

16

504,

33,1

998,

0460

50,0

5305

3,04

6050

,050

055,

0500

54,0

5005

0;

1630

6,38

,199

8,03

8038

,071

071,

0580

66,0

5507

5,05

4066

,046

058;

16

311,

38,1

998,

0420

50,0

5008

0,05

0074

,060

065,

0500

50,0

4605

6;

1631

6,38

,199

8,03

8038

,071

071,

0580

66,0

6507

5,05

4054

,050

050;

16

324,

38,1

998,

0380

38,0

7107

1,05

8074

,065

075,

0660

66,0

4605

2;

1638

1,38

,199

8,03

8038

,071

077,

0540

62,0

5506

5,00

0000

,000

000;

16

382,

38,1

998,

0380

38,0

6807

1,06

2062

,065

075,

0540

54,0

0000

0;

1644

9,38

,199

8,03

8038

,071

083,

0620

74,0

6506

5,06

2066

,050

056;

16

450,

38,1

998,

0380

38,0

7107

1,06

2062

,055

075,

0700

70,0

4605

0;

1645

3,38

,199

8,03

8038

,071

071,

0620

62,0

5506

5,05

4070

,050

056;

16

454,

38,1

998,

0380

38,0

7107

1,05

4058

,065

075,

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16

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16

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16

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16

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16

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53,1

998,

0380

38,0

6807

4,05

4062

,065

065,

0580

70,0

5605

8;

1633

0,53

,199

8,03

8038

,068

068,

0500

58,0

6506

5,05

8070

,046

056;

16

332,

53,1

998,

0380

38,0

6806

8,05

4062

,065

065,

0580

58,0

5605

8;

1634

1,53

,199

8,03

8038

,068

068,

0620

66,0

6006

5,05

8070

,058

058;

16

351,

59,1

998,

0500

50,0

7107

1,05

8058

,060

060,

0000

00,0

4605

0;

1636

3,59

,199

8,05

0054

,053

071,

0620

70,0

6006

5,06

6074

,046

058;

16

404,

60,1

998,

0380

54,0

5307

1,05

8062

,060

060,

0000

00,0

0000

0;

1648

7,61

,199

8,03

8054

,071

071,

0580

66,0

6006

5,05

4054

,050

050;

16

512,

61,1

998,

0380

50,0

4407

1,06

6066

,065

065,

0500

50,0

4605

0;

1630

0,62

,199

8,03

8038

,062

062,

0580

74,0

6506

5,05

0066

,046

056;

16

301,

62,1

998,

0380

38,0

5907

7,06

6066

,065

070,

0420

50,0

5405

4;

1647

0,62

,199

8,03

8050

,068

074,

0700

70,0

5506

0,05

0050

,046

052;

16

471,

62,1

998,

0000

00,0

4408

6,07

0078

,065

065,

0500

50,0

5605

8;

1642

4,64

,199

8,03

8050

,053

071,

0460

62,0

5505

5,05

8058

,052

056;

16

625,

64,1

998,

0500

50,0

7107

1,06

6066

,065

065,

0660

66,0

0000

0;

1644

8,65

,199

8,03

8050

,071

071,

0580

70,0

5506

5,05

4058

,046

046;

16

459,

65,1

998,

0380

38,0

7107

1,05

0050

,055

070,

0580

58,0

4604

6;

1646

3,65

,199

8,03

8038

,053

071,

0580

74,0

6506

5,05

8062

,056

058;

16

464,

65,1

998,

0380

38,0

6807

1,00

0000

,050

065,

0000

00,0

5005

0;

1646

5,65

,199

8,05

0054

,053

053,

0500

58,0

5005

5,05

0058

,050

050;

16

466,

65,1

998,

0380

38,0

7107

7,06

6070

,065

070,

0500

66,0

4604

6;

1647

8,65

,199

8,03

8050

,071

071,

0700

70,0

5005

5,05

4066

,046

046;

16

485,

65,1

998,

0380

38,0

6806

8,07

4074

,055

055,

0580

62,0

5205

8;

1648

6,65

,199

8,03

8038

,071

071,

0500

50,0

5506

5,06

2062

,052

058;

16

488,

65,1

998,

0500

50,0

7107

1,07

0070

,055

065,

0620

66,0

4605

8;

1662

6,65

,199

8,03

8050

,068

068,

0000

00,0

5005

0,05

8058

,000

000;

16

627,

65,1

998,

0380

38,0

6807

4,07

0070

,050

065,

0580

66,0

0000

0;

1662

8,65

,199

8,03

8038

,071

071,

0500

62,0

6506

5,06

6066

,000

000;

16

629,

65,1

998,

0420

50,0

6807

1,07

0070

,055

065,

0580

58,0

0000

0;

1663

0,65

,199

8,03

8050

,071

071,

0620

70,0

6506

5,05

8058

,000

000;

16

631,

65,1

998,

0380

50,0

7107

4,06

2070

,055

065,

0540

58,0

0000

0;

1658

4,70

,199

8,05

0054

,059

071,

0580

58,0

5505

5,04

6046

,048

052;

16

339,

71,1

998,

0500

50,0

4404

4,06

2066

,045

045,

0380

46,0

0000

0;

1634

0,71

,199

8,03

8050

,044

053,

0580

62,0

5506

0,04

6046

,050

056;

16

342,

71,1

998,

0380

50,0

4405

3,05

8062

,055

065,

0380

46,0

5605

6;

1636

0,71

,199

8,03

8050

,044

053,

0620

62,0

5506

0,04

6046

,056

056;

16

550,

71,1

998,

0380

50,0

5307

1,04

6062

,050

070,

0460

46,0

5005

0;

1649

7,74

,199

8,03

8050

,050

071,

0580

74,0

5505

5,05

0074

,046

046;

16

499,

74,1

998,

0380

38,0

7107

1,05

8070

,060

065,

0620

74,0

4605

2;

1650

0,74

,199

8,04

2050

,053

071,

0460

50,0

6506

5,04

6070

,050

050;

16

225,

76,1

998,

0380

38,0

7108

0,05

8066

,065

065,

0500

50,0

4605

0;

1623

5,77

,199

8,03

8046

,056

071,

0460

66,0

6506

5,04

6074

,000

000;

16

263,

77,1

998,

0380

50,0

5607

1,06

2062

,065

065,

0420

74,0

4604

6;

1634

5,77

,199

8,03

8038

,056

071,

0620

66,0

6506

5,05

4074

,046

050;

16

346,

77,1

998,

0380

46,0

5607

1,04

6066

,065

065,

0420

62,0

4604

6;

1634

7,77

,199

8,03

8046

,056

071,

0460

66,0

6506

5,04

2062

,046

050;

16

348,

77,1

998,

0380

50,0

0000

0,06

2066

,060

065,

0540

62,0

4604

6;

1638

5,77

,199

8,03

8038

,056

071,

0620

66,0

6506

5,05

4074

,046

050;

16

234,

78,1

998,

0500

50,0

5008

0,05

8058

,055

065,

0500

58,0

0000

0;

1623

6,78

,199

8,03

8050

,050

080,

0580

74,0

6506

5,05

0050

,000

000;

16

238,

78,1

998,

0380

50,0

5008

0,05

8074

,065

065,

0500

70,0

4604

6;

1625

9,78

,199

8,03

8050

,000

000,

0580

58,0

5006

5,05

0074

,000

000;

16

264,

78,1

998,

0380

50,0

8008

0,05

8062

,055

065,

0500

58,0

4604

6;

1627

7,78

,199

8,03

8038

,050

050,

0620

62,0

6506

5,05

0070

,046

046;

16

278,

78,1

998,

0380

50,0

5008

0,05

8070

,055

065,

0500

70,0

4604

6;

1627

9,78

,199

8,03

8050

,050

080,

0580

62,0

5506

5,05

0050

,046

056;

16

280,

78,1

998,

0380

50,0

5008

0,05

8062

,055

065,

0500

70,0

4606

0;

1628

1,78

,199

8,03

8038

,050

080,

0620

74,0

5505

5,05

0070

,056

060;

16

282,

78,1

998,

0380

50,0

5005

0,05

8062

,055

065,

0500

50,0

4606

0;

1628

3,78

,199

8,03

8050

,050

080,

0580

62,0

6506

5,05

0058

,046

060;

16

344,

78,1

998,

0380

50,0

8008

0,05

8062

,065

065,

0500

50,0

4605

6;

1647

5,79

,199

8,03

8038

,074

080,

0620

70,0

6506

5,05

4066

,046

056;

16

477,

79,1

998,

0380

38,0

7408

0,06

2062

,065

065,

0540

66,0

4605

6;

1650

1,79

,199

8,03

8038

,074

080,

0620

70,0

6506

5,05

4066

,046

056;

16

378,

80,1

998,

0380

50,0

7108

0,06

2070

,055

055,

0460

66,0

0000

0;

1640

6,80

,199

8,03

8038

,080

080,

0580

62,0

5005

5,04

6050

,000

000;

16

442,

81,1

998,

0380

54,0

8008

0,05

8074

,065

065,

0740

74,0

5605

6;

1645

7,97

,199

8,04

2042

,050

050,

0500

58,0

5505

5,05

4066

,050

052;

16

480,

100,

1998

,042

050,

0440

44,0

4605

0,05

0050

,046

058,

0500

50;

1648

1,10

0,19

98,0

5005

0,04

4050

,046

046,

0500

50,0

4604

6,05

0050

; 16

482,

100,

1998

,050

050,

0440

44,0

4605

0,05

0050

,046

058,

0500

54;



1648

3,10

0,19

98,0

5005

0,04

4053

,046

046,

0500

50,0

4604

6,05

0050

; 16

472,

101,

1998

,050

050,

0440

44,0

5005

8,05

0050

,038

046,

0500

50;

1627

4,10

3,19

98,0

3805

0,07

4080

,050

054,

0650

65,0

5406

2,04

6046

; 16

325,

103,

1998

,038

050,

0740

74,0

5005

8,05

5065

,054

058,

0460

56;

1632

6,10

3,19

98,0

3803

8,07

4074

,050

054,

0650

65,0

5806

6,04

6056

; 16

412,

103,

1998

,038

054,

0590

71,0

5406

2,05

5065

,000

000,

0000

00;

1638

3,10

4,19

98,0

5005

0,07

1074

,058

058,

0550

55,0

6206

2,00

0000

; 16

408,

104,

1998

,038

038,

0710

74,0

6607

4,06

5065

,046

050,

0000

00;

1641

8,10

4,19

98,0

3803

8,07

1074

,000

000,

0550

65,0

5005

4,04

6058

; 16

441,

104,

1998

,038

038,

0500

50,0

6207

4,06

5065

,050

054,

0460

46;

1627

6,10

5,19

98,0

4204

2,00

0000

,000

000,

0450

50,0

0000

0,04

6046

; 16

293,

105,

1998

,038

050,

0500

50,0

5806

2,05

5065

,070

070,

0460

56;

1629

4,10

5,19

98,0

3805

0,05

6077

,054

054,

0650

65,0

5405

8,04

6046

; 16

295,

105,

1998

,038

050,

0710

71,0

5806

2,06

5070

,054

070,

0500

52;

1637

6,10

5,19

98,0

3803

8,06

8071

,050

050,

0550

65,0

5005

4,04

6046

; 16

377,

105,

1998

,038

038,

0680

71,0

5806

2,06

5065

,050

058,

0440

58;

1638

4,10

5,19

98,0

3805

0,07

1074

,050

058,

0550

60,0

6206

2,00

0000

; 16

395,

105,

1998

,038

054,

0740

74,0

5005

0,06

5065

,054

054,

0000

00;

1639

6,10

5,19

98,0

5005

4,07

4074

,050

058,

0550

65,0

5406

2,00

0000

; 16

417,

105,

1998

,038

050,

0710

74,0

5806

6,06

0065

,062

070,

0460

50;

1642

0,10

5,19

98,0

3803

8,07

1074

,050

050,

0550

65,0

5406

2,04

6056

; 16

427,

105,

1998

,038

038,

0710

74,0

5005

0,05

5065

,050

066,

0500

56;

1644

0,10

5,19

98,0

3805

0,05

6071

,050

058,

0550

70,0

6207

0,04

6052

; 16

327,

110,

1998

,042

054,

0710

74,0

5005

8,05

5055

,050

054,

0500

50;

1633

6,11

0,19

98,0

5405

4,07

1071

,062

062,

0600

65,0

5005

8,04

8050

; 16

443,

110,

1998

,038

050,

0710

77,0

0000

0,05

0050

,000

000,

0000

00;

1629

6,11

6,19

98,0

3804

2,07

1071

,050

054,

0550

65,0

5805

8,04

6056

; 16

462,

118,

1998

,046

050,

0680

68,0

5807

4,05

0050

,000

000,

0460

56;

1649

8,11

8,19

98,0

3803

8,05

3053

,046

046,

0600

65,0

4205

8,05

0050

; 16

244,

122,

1998

,038

038,

0680

71,0

5005

8,05

0065

,050

066,

0460

58;

1625

1,12

2,19

98,0

3805

4,05

0071

,050

050,

0650

70,0

5405

4,04

6060

; 16

252,

122,

1998

,038

038,

0530

71,0

5005

4,06

5065

,050

062,

0460

58;

1626

5,12

2,19

98,0

3803

8,06

8071

,058

066,

0650

65,0

0000

0,05

6058

; 16

421,

123,

1998

,038

038,

0710

71,0

5805

8,06

5065

,050

050,

0460

46;

1642

8,12

3,19

98,0

3803

8,07

1071

,046

066,

0650

65,0

5005

0,04

6046

; 16

429,

123,

1998

,038

038,

0710

71,0

5005

0,06

5065

,054

062,

0460

72;

1643

0,12

3,19

98,0

3803

8,07

1071

,058

066,

0650

65,0

5005

0,04

6046

; 16

431,

123,

1998

,038

038,

0710

71,0

5805

8,06

5065

,050

050,

0560

56;

1643

2,12

3,19

98,0

3803

8,07

1071

,058

062,

0650

65,0

5405

4,04

6046

; 16

433,

123,

1998

,038

038,

0710

71,0

6606

6,06

5065

,050

070,

0460

58;

1643

4,12

3,19

98,0

3803

8,07

1071

,058

058,

0650

65,0

5005

0,04

6046

; 16

435,

123,

1998

,038

038,

0710

71,0

5405

8,06

5065

,054

066,

0460

46;

1643

6,12

3,19

98,0

3803

8,07

1071

,050

058,

0650

65,0

5405

4,04

6046

; 16

437,

123,

1998

,038

038,

0710

71,0

5006

6,06

5065

,054

070,

0460

46;

1643

8,12

3,19

98,0

3803

8,07

1071

,050

058,

0650

65,0

5405

4,04

6046

; 16

572,

123,

1998

,038

038,

0710

71,0

4605

8,06

5065

,050

050,

0460

46;

1657

3,12

3,19

98,0

3803

8,07

1071

,054

058,

0650

65,0

5406

6,05

6058

; 16

468,

124,

1998

,038

038,

0680

71,0

6207

4,05

5065

,062

066,

0460

58;

1642

3,12

6,19

98,0

3804

2,04

7077

,058

062,

0650

70,0

6607

0,04

6046

; 16

425,

126,

1998

,038

042,

0560

71,0

6206

2,06

5070

,066

070,

0460

46;

1648

9,12

7,19

98,0

3804

2,07

1077

,062

062,

0650

70,0

7007

4,04

6046

; 16

558,

127,

1998

,038

038,

0710

71,0

6206

6,06

5075

,058

070,

0460

50;

1655

9,12

7,19

98,0

3805

0,07

1080

,054

074,

0650

70,0

5005

4,04

6050

; 16

560,

127,

1998

,038

038,

0710

71,0

5806

6,06

5075

,054

070,

0460

50;

1656

1,12

7,19

98,0

3803

8,07

1077

,062

066,

0650

75,0

5407

0,04

6050

; 16

562,

127,

1998

,038

046,

0710

77,0

5806

6,06

5070

,070

070,

0460

50;

1635

9,13

2,19

98,0

3803

8,07

1080

,050

070,

0650

65,0

6206

6,04

6046

; 16

374,

132,

1998

,038

038,

0710

71,0

5007

0,06

5070

,050

062,

0460

46;

1646

7,13

2,19

98,0

3803

8,06

8068

,050

074,

0650

65,0

5806

2,05

2058

; 16

490,

133,

1998

,046

050,

0470

53,0

5805

8,05

0065

,050

070,

0500

50;

1619

0,13

6,19

98,0

5005

0,07

1071

,058

070,

0500

65,0

5807

0,04

6046

; 16

207,

136,

1998

,038

038,

0710

80,0

6206

2,05

5060

,050

054,

0460

56;

1644

6,13

7,19

98,0

5005

0,04

4050

,050

050,

0500

70,0

4204

6,05

0058

; 16

419,

138,

1998

,038

050,

0710

71,0

6607

0,05

5065

,046

054,

0460

46;

1651

1,13

8,19

98,0

3805

0,07

1071

,066

070,

0550

65,0

5406

2,04

6046

; 16

334,

139,

1998

,038

038,

0710

71,0

5806

6,06

5065

,046

046,

0460

46;

1633

5,13

9,19

98,0

3803

8,07

1071

,058

066,

0650

65,0

4607

4,04

6046

; 16

356,

139,

1998

,038

042,

0710

71,0

5805

8,06

5065

,046

074,

0460

56;

1635

7,13

9,19

98,0

3804

2,07

1071

,058

058,

0650

65,0

7407

4,04

6056

; 16

358,

139,

1998

,038

038,

0710

71,0

5805

8,06

5065

,074

074,

0460

56;

1636

1,13

9,19

98,0

4204

2,07

1071

,058

058,

0650

65,0

7407

4,04

6056

; 16

362,

139,

1998

,042

042,

0710

71,0

5805

8,06

5065

,046

058,

0460

56;

1636

7,13

9,19

98,0

3803

8,07

1071

,058

066,

0650

65,0

7407

4,04

6056

; 16

368,

139,

1998

,038

042,

0710

71,0

5805

8,06

5065

,046

046,

0460

56;

1636

9,13

9,19

98,0

4204

2,07

1071

,058

066,

0650

65,0

4604

6,04

6046

; 16

370,

139,

1998

,038

038,

0710

71,0

5806

6,06

5065

,046

074,

0460

46;

1638

7,13

9,19

98,0

3804

2,07

1071

,058

058,

0650

65,0

4604

6,04

6046

; 16

397,

139,

1998

,038

042,

0710

71,0

5805

8,06

5065

,074

074,

0460

46;

1646

1,13

9,19

98,0

3803

8,07

1071

,058

058,

0650

65,0

4607

4,04

6046

; 16

514,

139,

1998

,042

042,

0710

71,0

5805

8,06

5065

,074

074,

0560

58;

1651

5,13

9,19

98,0

3804

2,07

1071

,058

058,

0500

65,0

5005

0,04

6056

; 16

516,

139,

1998

,042

042,

0710

71,0

5805

8,06

5065

,074

074,

0460

56;

1651

7,13

9,19

98,0

3803

8,07

1071

,058

058,

0650

65,0

5807

4,04

6056

; 16

518,

139,

1998

,042

042,

0710

71,0

5805

8,06

5065

,046

074,

0460

56;

1651

9,13

9,19

98,0

4204

2,07

1071

,058

058,

0650

65,0

4607

4,04

6046

; 16

520,

139,

1998

,038

042,

0710

71,0

5806

6,06

5070

,046

058,

0460

46;

1652

1,13

9,19

98,0

3804

2,07

1071

,058

058,

0650

65,0

5005

0,04

6046

; 16

522,

139,

1998

,042

042,

0710

71,0

5805

8,06

5065

,046

046,

0460

56;

1652

3,13

9,19

98,0

3803

8,07

1071

,058

058,

0650

65,0

4607

4,04

6056

; 16

524,

139,

1998

,038

038,

0710

71,0

5805

8,06

5065

,046

046,

0460

56;

1652

5,13

9,19

98,0

3803

8,07

1071

,058

058,

0650

65,0

7407

4,04

6056

; 16

526,

139,

1998

,038

042,

0710

71,0

5805

8,06

5065

,046

046,

0460

56;

1652

7,13

9,19

98,0

3804

2,07

1071

,058

066,

0650

65,0

4607

4,04

6056

; 16

528,

139,

1998

,038

038,

0000

00,0

5805

8,06

5065

,074

074,

0460

46;

1653

0,13

9,19

98,0

3803

8,07

1071

,050

058,

0650

65,0

5405

4,04

6056

; 16

479,

140,

1998

,050

050,

0440

53,0

7407

4,06

5065

,046

046,

0500

50;

1650

2,14

0,19

98,0

5005

0,04

4053

,074

074,

0650

65,0

4604

6,05

0050

; 16

305,

141,

1998

,038

038,

0530

71,0

5405

8,06

5065

,046

054,

0520

52;

1631

7,14

1,19

98,0

3803

8,07

1071

,050

058,

0650

65,0

4605

4,05

6056

; 16

319,

141,

1998

,038

038,

0710

71,0

5005

0,06

5065

,046

054,

0460

58;

1632

0,14

1,19

98,0

3803

8,06

8071

,050

050,

0650

65,0

4605

4,04

6058

; 16

321,

141,

1998

,038

038,

0710

71,0

5805

8,06

5065

,054

054,

0460

56;

1632

2,14

1,19

98,0

3803

8,06

8071

,058

058,

0650

65,0

4605

4,05

6056

; 16

323,

141,

1998

,038

038,

0710

71,0

5005

4,06

5065

,054

054,

0460

52;

1633

7,14

1,19

98,0

3803

8,07

1071

,062

062,

0650

65,0

4605

4,04

6046

; 16

338,

141,

1998

,038

038,

0710

71,0

5805

8,06

5065

,046

054,

0520

56;

1652

9,14

1,19

98,0

3803

8,07

1071

,058

066,

0650

70,0

4607

4,04

6056

; 16

531,

141,

1998

,038

038,

0710

71,0

5005

4,06

5065

,054

054,

0460

52;

1653

2,14

1,19

98,0

3803

8,07

1071

,050

058,

0650

65,0

5405

4,05

6058

; 16

533,

141,

1998

,038

038,

0710

71,0

5405

8,06

5065

,054

054,

0520

56;

1653

4,14

1,19

98,0

3803

8,07

1071

,050

050,

0650

65,0

5405

4,04

6056

; 16

535,

141,

1998

,038

038,

0710

71,0

5805

8,06

5065

,054

054,

0560

56;

1653

6,14

1,19

98,0

3803

8,07

1071

,058

058,

0650

65,0

4605

4,04

6056

; 16

537,

141,

1998

,038

038,

0710

71,0

5005

8,06

5065

,046

054,

0560

58;

1653

8,14

1,19

98,0

3803

8,06

8071

,050

058,

0650

65,0

5405

4,04

6056

; 16

539,

141,

1998

,038

038,

0710

71,0

5005

0,06

5065

,046

054,

0520

52;

1654

0,14

1,19

98,0

3803

8,06

8071

,058

058,

0650

65,0

4605

4,05

6058

; 16

541,

141,

1998

,038

038,

0710

71,0

5805

8,06

5065

,046

054,

0560

58;

1654

2,14

1,19

98,0

3803

8,07

1071

,058

058,

0650

65,0

5405

4,05

6058

; 16

543,

141,

1998

,038

038,

0710

71,0

5005

8,06

5065

,054

054,

0460

58;

1654

4,14

1,19

98,0

3803

8,06

8071

,050

058,

0650

65,0

4605

4,05

2058

; 16

545,

141,

1998

,038

038,

0710

71,0

5006

2,06

5065

,054

054,

0460

56;

1654

6,14

1,19

98,0

3803

8,06

8071

,058

058,

0650

65,0

4605

4,05

6056

; 16

547,

141,

1998

,038

038,

0710

71,0

5005

8,06

5065

,046

054,

0460

46;

1636

4,14

2,19

98,0

3803

8,05

3053

,062

062,

0550

55,0

4605

0,05

0056

; 16

365,

142,

1998

,042

050,

0530

53,0

5005

0,06

0065

,046

070,

0500

50;

1637

1,14

2,19

98,0

3805

0,05

0050

,062

066,

0550

65,0

5807

0,04

6050

; 16

372,

142,

1998

,038

054,

0500

50,0

6206

2,05

5065

,046

050,

0460

56;

1637

3,14

2,19

98,0

3803

8,05

0050

,062

062,

0550

65,0

5005

4,05

0050

; 16

375,

142,

1998

,038

054,

0500

50,0

6206

2,06

0065

,050

054,

0480

52;

1637

9,14

2,19

98,0

3803

8,05

0050

,058

062,

0550

60,0

5005

4,05

2052

; 16

390,

142,

1998

,038

054,

0500

50,0

6206

6,05

5065

,050

054,

0000

00;

1639

3,14

2,19

98,0

5005

0,05

0071

,066

066,

0550

55,0

5405

4,00

0000

; 16

394,

142,

1998

,038

038,

0500

50,0

6206

2,06

0065

,050

070,

0000

00;

1640

0,14

2,19

98,0

3804

2,05

0050

,050

062,

0550

65,0

7007

0,00

0000

; 16

401,

142,

1998

,038

042,

0500

50,0

5006

2,06

0060

,050

070,

0000

00;

1640

5,14

2,19

98,0

5405

4,05

0050

,062

062,

0550

60,0

5005

0,00

0000

; 16

407,

142,

1998

,038

054,

0500

71,0

5806

2,05

5065

,070

070,

0000

00;

1640

9,14

2,19

98,0

3805

4,05

0050

,058

062,

0550

55,0

5005

4,00

0000

; 16

410,

142,

1998

,050

050,

0710

71,0

5006

6,05

5065

,058

070,

0000

00;

1641

1,14

2,19

98,0

5005

0,07

1071

,050

066,

0550

65,0

5807

0,00

0000

; 16

413,

142,

1998

,038

038,

0500

50,0

6206

2,06

0065

,050

054,

0000

00;

1641

6,14

2,19

98,0

3805

4,05

0050

,062

062,

0550

65,0

5007

0,00

0000

; 16

451,

142,

1998

,038

054,

0500

50,0

6206

2,05

5065

,050

054,

0460

50;

1645

2,14

2,19

98,0

3805

4,05

0050

,062

062,

0550

60,0

5005

4,04

6050

; 16

456,

142,

1998

,038

038,

0500

50,0

6206

2,05

5055

,054

054,

0500

50;

1645

8,14

2,19

98,0

4204

2,06

5071

,046

046,

0550

55,0

5005

4,04

6058

; 16

563,

142,

1998

,042

054,

0500

50,0

5006

2,05

5065

,050

070,

0460

50;

1656

4,14

2,19

98,0

3805

4,05

0050

,058

062,

0650

65,0

5005

0,05

0050

; 16

565,

142,

1998

,038

038,

0500

50,0

6206

2,06

5065

,070

070,

0500

56;

1656

6,14

2,19

98,0

5005

4,05

0071

,062

066,

0550

65,0

5405

8,04

6056

; 16

567,

142,

1998

,050

050,

0500

50,0

6606

6,05

5065

,070

070,

0460

56;

1656

8,14

2,19

98,0

3805

0,05

0050

,062

066,

0600

65,0

7007

0,05

0056

; 16

569,

142,

1998

,038

054,

0710

77,0

5406

2,05

0055

,066

070,

0460

50;

1657

0,14

2,19

98,0

5005

0,05

0071

,066

066,

0550

60,0

7007

0,05

0056

; 16

571,

142,

1998

,050

050,

0500

71,0

6606

6,05

5065

,070

070,

0460

56;

1660

9,14

2,19

98,0

3805

4,05

0050

,062

062,

0550

65,0

4605

4,04

6056

; 16

610,

142,

1998

,050

050,

0500

71,0

6606

6,05

5065

,058

070,

0500

56;

1661

1,14

2,19

98,0

3805

4,05

0050

,058

058,

0550

60,0

7007

0,05

0050

; 16

612,

142,

1998

,038

042,

0500

50,0

5006

2,05

5065

,050

050,

0560

56;

1661

3,14

2,19

98,0

3805

0,05

0059

,062

066,

0550

55,0

7007

0,04

6056

; 16

614,

142,

1998

,050

054,

0500

59,0

5006

2,05

5065

,054

054,

0560

56;

1644

7,14

3,19

98,0

4205

0,07

7080

,050

070,

0550

60,0

6607

4,05

0052

; 16

391,

144,

1998

,050

050,

0710

71,0

5005

0,06

5065

,062

074,

0000

00;

1641

4,15

1,19

98,0

3805

0,05

0056

,054

066,

0500

60,0

5005

4,00

0000

; 16

415,

151,

1998

,050

050,

0500

53,0

4605

0,06

0070

,046

046,

0000

00;

1647

3,15

1,19

98,0

5005

0,05

0056

,066

066,

0500

70,0

5005

4,04

8050

; 16

474,

151,

1998

,050

050,

0500

53,0

0000

0,05

0050

,046

054,

0500

50;

1650

6,15

1,19

98,0

3803

8,05

0056

,054

058,

0700

70,0

5005

4,04

8050

; 16

507,

151,

1998

,050

050,

0530

53,0

6606

6,06

0070

,054

054,

0500

50;

1650

8,15

1,19

98,0

5005

0,05

3053

,066

066,

0600

60,0

4605

4,05

0050

; 16

509,

151,

1998

,038

050,

0500

50,0

5806

6,05

0070

,046

050,

0480

50;

1651

0,15

1,19

98,0

3805

0,05

3053

,054

066,

0600

70,0

5405

4,05

0050

; 16

388,

154,

1998

,046

050,

0500

56,0

5005

8,05

0055

,046

070,

0000

00;

1638

9,15

4,19

98,0

4605

0,05

0053

,058

058,

0500

60,0

4607

0,00

0000

; 16

399,

154,

1998

,046

050,

0530

56,0

5005

8,06

0060

,046

046,

0000

00;

1663

4,15

9,19

98,0

5005

0,05

0053

,054

062,

0550

60,0

4204

6,00

0000

;

1

6460

,720

,199

8,03

8038

,074

077,

0580

58,0

6506

5,05

4058

,000

000;

AP

PE

ND

IX 3

Con

tin

ued

Documents

Inferences of selection and migration in the Danish house ...webpages.icav.up.pt/PTDC/BIA-BEC/103440/2008... · between the house mouse subspecies Mus musculus domesticus and M. m