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 domesticus
–
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.
594
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.
© 2005 The Linnean Society of London,
Biological Journal of the Linnean Society,
2005,
84
, 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,
Mus musculus domesticus
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 The Linnean Society of London,
Biological Journal of the Linnean Society,
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
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.
© 2005 The Linnean Society of London,
Biological Journal of the Linnean Society,
2005,
84
, 593–616
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
598 N. RAUFASTE ET AL.
© 2005 The Linnean Society of London, Biological Journal of the Linnean Society, 2005, 84, 593–616
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
SELECTION IN THE HOUSE MOUSE HYBRID ZONE 599
© 2005 The Linnean Society of London, Biological Journal of the Linnean Society, 2005, 84, 593–616
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
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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
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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.
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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/
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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
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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
Locality no. Locality name Year Longitude Latitude Transect coordinate N HI eight loci
APPENDIX 1 Continued
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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
Locality no. Locality name Year Longitude Latitude Transect coordinate N HI eight loci
APPENDIX 1 Continued
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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
Locality no. Locality name Year Longitude Latitude Transect coordinate N HI eight loci
APPENDIX 1 Continued
612 N. RAUFASTE ET AL.
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AP
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2122
;
7376
,85,
1291
2212
22;
7379
,85,
1292
2222
11;
7380
,85,
2291
0012
12;
7381
,85,
2091
2112
21;
7382
,85,
2091
2112
21;
7383
,85,
2291
2222
22;
7384
,85,
2192
2222
02;
7385
,85,
9192
1212
02;
7386
,128
,009
0000
100;
73
87,1
28,0
0900
0010
0;
7388
,128
,909
0000
000;
73
89,1
28,1
0900
0000
0;
7390
,128
,009
0000
100;
739
1,12
8,10
9000
1100
; 73
92,1
28,2
0900
0020
0; 7
393,
128,
1090
0012
00;
7394
,128
,009
0000
000;
739
5,12
8,10
9000
1100
; 73
96,1
28,0
0900
0000
0; 7
397,
128,
0090
0002
00;
7398
,128
,009
0000
200;
739
9,12
8,00
9000
0000
; 74
00,1
28,0
0910
0000
0; 7
974,
129,
9100
0001
00;
7975
,129
,000
0000
200;
797
6,12
9,01
0000
0000
; 79
77,1
29,0
1000
0010
0; 7
978,
129,
0000
0001
00;
7979
,129
,100
0000
100;
798
0,12
9,21
0000
0000
; 79
81,1
29,0
0001
0010
0; 7
982,
129,
0000
0001
00;
7983
,129
,100
0029
000;
798
4,12
9,00
0000
0000
; 79
85,1
29,1
1000
0000
0; 7
986,
129,
0100
0000
00;
7987
,129
,100
0029
000;
798
8,12
9,00
0002
9100
; 79
89,1
29,0
0000
2910
0; 7
990,
129,
1000
1001
00;
7991
,129
,000
0029
100;
799
2,12
9,01
0000
0000
; 79
93,1
29,0
1001
0020
0; 7
994,
129,
0000
0290
00;
7995
,146
,222
2101
001;
799
6,14
6,22
2221
1002
; 79
97,2
0,22
9299
9221
; 79
98,2
0,21
9199
9002
;
7999
,146
,112
1999
222;
80
00,2
0,21
9199
9001
; 80
03,8
6,22
2102
2222
; 80
04,8
6,22
2222
2222
; 80
05,8
6,22
2221
2222
; 80
06,8
6,22
0222
2222
; 80
08,8
6,11
1120
2211
; 80
31,5
5,22
2222
2222
; 80
37,5
5,22
2222
2222
; 80
38,5
5,22
2222
2222
; 80
40,5
5,21
2222
2222
; 80
41,5
5,22
2222
2222
;
8042
,55,
2222
2222
22;
8071
,16,
2212
0122
11;
8072
,16,
1222
9991
22;
8073
,16,
2222
0222
22;
8074
,16,
0221
2122
20;
8075
,120
,211
0112
200;
80
76,1
20,1
2121
1120
2;
8077
,120
,022
0999
100;
80
78,1
20,0
1020
1120
2;
8079
,120
,022
0999
100;
80
80,1
20,1
1020
2200
1;
8081
,120
,122
2999
101;
8082
,120
,112
1999
100;
80
83,1
46,1
0229
9912
2;
8084
,146
,221
1002
222;
80
85,1
46,1
2212
0221
0;
8086
,146
,212
2102
200;
80
87,1
46,2
1920
2222
2;
8091
,86,
2292
2222
22;
8244
,113
,000
0000
000;
82
45,1
13,1
0000
1000
0;
8246
,75,
1222
2222
22;
8247
,75,
1222
2222
22;
8248
,75,
2222
2221
22;
8250
,75,
2222
2221
22;
8251
,75,
2222
2220
22;
8252
,75,
2222
2221
22;
8253
,75,
2222
2222
22;
8254
,75,
2222
2220
22;
8255
,75,
2222
2220
22;
8256
,75,
2222
2221
21;
8257
,86,
2091
1022
01;
8258
,86,
1122
2222
12;
8259
,86,
1222
9992
02;
8260
,86,
2122
1222
12;
8261
,86,
1222
9992
02;
8262
,86,
2222
9992
12;
8263
,86,
2222
2222
20;
8264
,86,
2222
9992
12;
8265
,20,
1021
2121
10;
8266
,20,
0010
1110
10;
8268
,93,
0000
0010
00;
8269
,93,
2000
0210
00;
8270
,120
,102
1101
222;
82
71,1
20,1
0201
2102
0;
9345
,13,
2011
1111
00;
9346
,73,
2019
2220
10;
9348
,21,
0012
2100
12;
9349
,21,
9201
2111
10;
9350
,36,
2221
2122
20;
9351
,36,
2221
2122
21;
9352
,36,
2220
2121
21;
9353
,36,
2211
2120
21;
9354
,36,
2220
2222
20;
9355
,36,
2211
2122
21;
9356
,125
,101
1111
112;
93
57,1
25,0
2199
9911
9;
9358
,125
,912
1001
001;
93
59,1
25,0
2009
9902
9;
9360
,67,
2222
2222
12;
9361
,67,
1212
2222
12;
9362
,67,
2222
2222
02;
9363
,125
,211
0900
101;
93
64,1
25,1
0201
0000
0;
9365
,125
,191
9999
009;
93
66,1
25,2
0100
0002
0;
9367
,131
,111
9102
012;
93
68,1
47,2
2922
1112
1;
9369
,125
,192
1002
022;
93
70,3
0,22
9002
2290
; 93
76,9
,222
2912
010;
93
77,9
,222
2212
091;
9378
,9,2
2229
1209
0;
9379
,9,0
2099
9902
9;
9380
,9,2
2929
2202
1;
9381
,9,2
2222
0219
2;
9382
,9,0
1222
0211
2;
9383
,9,1
2912
2222
2;
9384
,9,2
2122
0211
1;
9385
,9,0
2220
2221
1;
9386
,9,1
2211
1291
2;
9387
,9,2
2222
0201
1;
9388
,9,0
2212
2221
2;
9389
,9,0
2921
2219
2;
9390
,9,0
2222
2012
0;
9391
,9,0
2229
9911
9; 9
392,
9,02
2290
2229
; 93
93,9
,112
1222
120;
939
4,9,
1212
2022
21;
9395
,9,0
2222
2212
1; 9
400,
56,2
2222
2222
2; 9
401,
56,2
2222
2222
2; 9
402,
56,2
2222
2222
2; 9
403,
56,1
2292
2222
2; 9
404,
56,2
2292
2212
2; 9
405,
56,2
2221
2222
2; 9
406,
56,1
2222
2222
2; 9
407,
56,2
2222
2212
2;
9408
,56,
2222
2221
22;
9409
,56,
2222
2222
22;
9410
,56,
2222
2222
22;
9411
,56,
2222
2222
22;
9412
,56,
2222
2221
22;
9428
,56,
1222
2222
22;
9429
,56,
1292
2222
92;
9430
,56,
1292
2229
92;
9431
,56,
9292
2222
91;
9432
,56,
9222
1222
22;
1060
0,1,
1222
2121
12;
1060
1,1,
2221
1122
12;
1060
2,1,
0222
2012
22;
1060
3,89
,101
1112
120;
106
04,8
9,11
2022
0100
; 10
605,
50,1
2211
2200
2; 1
0606
,50,
2211
2222
02;
1060
7,50
,121
2122
101;
106
08,5
0,12
1022
2201
; 10
609,
50,2
2122
2211
2; 1
0610
,94,
0001
0002
00;
1061
1,94
,000
2000
000;
106
14,1
11,2
0122
0202
1; 1
0615
,111
,201
1212
120;
1061
6,11
1,21
1221
2121
; 106
17,1
11,1
2221
1222
1; 1
0618
,111
,221
2102
121;
106
19,1
11,1
2222
1122
1; 1
0620
,111
,202
2202
121;
106
21,7
,200
0110
200;
106
22,7
,200
0001
110;
106
23,7
,200
0111
000;
106
24,5
0,22
1211
2202
; 106
25,5
0,12
1202
2111
; 106
26,5
0,22
1202
2102
; 106
27,6
3,22
1221
2200
;
1062
8,66
,211
2202
202;
106
29,6
6,21
2201
2222
; 106
30,8
4,12
1221
2222
; 106
31,8
4,12
1022
2220
; 106
32,8
4,12
1222
1220
; 106
33,8
4,21
1222
2220
; 106
35,2
4,10
0001
0200
; 106
45,7
,200
0020
000;
106
50,6
6,99
9222
2990
; 106
76,1
14,0
0000
0000
0; 1
0677
,114
,100
0000
000;
106
78,1
14,0
0000
0000
0;
1067
9,11
4,00
0000
0000
; 10
680,
114,
9000
0000
09;
1068
1,11
4,00
0001
0009
; 10
682,
114,
0000
0000
00;
1068
3,11
4,00
0000
0000
; 10
684,
114,
0000
0000
00;
1068
6,11
4,00
0000
0000
; 10
687,
114,
0000
0100
00;
1068
8,11
4,00
0000
0000
; 10
689,
114,
0000
0000
00;
1069
0,11
5,00
0001
0000
;
1069
1,11
5,00
0001
0000
; 10
693,
115,
0000
0000
00;
1069
4,11
5,00
0000
0000
; 10
695,
114,
0000
0000
00;
1081
2,40
,222
1222
222;
10
813,
40,2
2212
1222
2;
1081
4,40
,222
2222
222;
10
815,
40,2
2222
2222
2;
1081
6,40
,122
2222
222;
10
820,
39,1
2222
2222
2;
1082
1,39
,222
2212
221;
1082
2,39
,222
2222
122;
108
23,3
9,22
2222
2222
; 108
24,3
9,22
2222
2122
; 108
25,3
9,22
2222
2222
; 108
26,3
9,22
2222
2222
; 108
27,1
49,2
2002
1222
0; 1
0828
,149
,220
1212
120;
108
29,1
49,2
1022
0122
0; 1
0830
,51,
2121
2221
01; 1
0831
,51,
2210
2222
01; 1
0832
,51,
2200
2222
11; 1
0833
,51,
2210
2222
12;
1083
4,51
,222
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
,220
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
,222
2002
121;
10
857,
37,2
0201
1212
1;
1085
8,35
,222
2102
211;
1085
9,35
,102
2102
222;
108
61,4
1,10
2222
2222
; 10
862,
41,2
2222
2222
1; 1
0863
,41,
2222
2222
22;
1086
4,41
,222
2222
221;
108
65,4
1,22
2222
2222
; 10
866,
41,2
2222
2222
2; 1
0867
,41,
2221
2222
22;
1086
8,41
,122
2222
222;
108
69,4
1,12
2222
2222
; 10
870,
41,1
2229
9922
9; 1
0871
,41,
2222
9992
29;
1087
2,41
,122
2999
229;
108
73,4
1,12
2222
2222
; 10
874,
41,0
2212
2222
2; 1
0875
,41,
1221
2222
22;
1087
6,41
,122
1222
222;
108
77,4
1,02
2222
2222
; 10
878,
37,2
2121
2222
2; 1
0879
,41,
2221
2222
22;
1088
0,37
,220
1202
212;
108
81,3
7,22
2010
2222
; 10
882,
41,2
2222
2222
2; 1
0883
,41,
2222
2222
21;
1088
4,41
,222
2222
222;
108
87,4
,012
2211
210;
108
88,4
,022
2221
221;
108
89,1
30,2
1000
0120
0; 1
0890
,130
,000
0001
200;
108
91,3
7,22
2110
2221
; 10
892,
37,2
1121
0221
2; 1
0893
,37,
2221
2222
22;
1089
4,51
,221
1122
220;
108
95,1
08,2
2100
2222
2; 1
0896
,95,
0201
0222
00;
1089
7,35
,212
2222
222;
1089
8,35
,212
2102
211;
108
99,3
7,22
2020
2212
; 109
00,3
7,22
2110
2222
; 109
01,3
7,21
2022
1210
; 109
02,3
7,22
2022
1200
; 109
03,3
7,22
2112
2211
; 109
05,3
7,12
2020
1120
; 109
06,3
7,12
1222
2220
; 109
07,1
48,2
2120
0110
1; 1
0908
,112
,210
2001
221;
109
09,1
48,2
2211
1212
2; 1
0910
,2,0
1200
1220
1;
1091
1,14
9,21
1020
2221
; 10
912,
149,
2110
2022
21;
1091
4,14
9,22
2121
1220
; 10
916,
149,
2110
2022
20;
1091
7,14
9,01
0120
2220
; 10
919,
17,2
2212
1222
2;
1092
0,17
,222
1212
221;
10
921,
17,1
2222
2221
2;
1092
2,17
,222
2221
211;
10
923,
17,2
2222
2222
0;
1092
4,17
,222
1212
220;
1092
5,17
,222
2222
219;
109
26,1
7,21
2221
1220
; 10
927,
57,2
2222
2222
2; 1
0928
,57,
2222
2222
22;
1092
9,10
8,22
2221
1212
; 10
930,
4,22
2211
0222
; 10
931,
4,22
2211
1220
; 10
932,
17,2
2212
2122
2; 1
0933
,17,
2221
2222
10;
1093
4,51
,220
0222
211;
109
35,5
1,22
0122
2210
; 10
936,
51,2
2012
2221
1;
1093
7,51
,220
1222
220;
109
38,1
7,21
2120
2221
; 10
939,
17,2
1112
2221
1; 1
0940
,17,
2122
2122
21;
1094
1,17
,222
2221
221;
109
62,1
7,22
1212
1121
; 10
963,
17,1
2120
2122
2; 1
0964
,17,
2222
1221
22;
1096
5,17
,212
0212
221;
109
66,1
7,21
1122
2222
; 10
967,
17,2
2102
1221
0; 1
0968
,17,
2220
2112
12;
1096
9,17
,220
2211
222;
109
70,1
7,12
1220
0221
; 10
972,
17,2
1112
1122
1; 1
0973
,17,
2121
2022
11;
1097
4,17
,222
2222
220;
109
76,1
7,22
2202
2221
; 10
977,
17,2
2220
2212
2; 1
0978
,37,
2000
0000
00;
1104
9,51
,920
1122
211;
110
50,5
1,92
0112
2212
; 11
051,
51,9
2012
2220
0; 1
1052
,51,
9201
2222
11;
1105
3,51
,921
0222
201;
110
54,5
1,92
0022
2210
; 110
55,5
1,92
0022
2211
; 110
56,5
1,92
0022
2212
; 112
04,5
8,22
0102
2922
; 112
05,5
8,22
2202
2202
; 112
07,1
1,21
2000
2001
; 112
08,3
1,22
2221
1101
; 112
09,1
07,2
2100
2222
2; 1
1210
,102
,211
0001
001;
112
11,1
35,0
2110
0110
2; 1
1212
,11,
2222
0121
10;
1121
3,11
,221
1002
110;
11
214,
135,
2002
0120
01;
1121
6,15
0,22
2101
0210
; 11
217,
150,
2211
0112
02;
1122
0,13
5,22
2001
2112
; 11
221,
150,
2221
0011
00;
1122
2,15
0,10
0001
2200
; 11
224,
106,
2120
0102
02;
1122
6,10
6,22
2202
2221
; 11
227,
96,2
2000
0110
0;
1122
8,10
6,22
2001
1102
;
1122
9,10
6,21
1101
0102
; 11
243,
106,
2020
0220
02;
1124
4,82
,202
2012
002;
11
245,
82,0
0110
0202
2;
1124
6,82
,202
2022
020;
11
248,
150,
0200
0102
00;
1124
9,14
5,02
0010
0000
; 11
250,
11,2
0110
1222
2;
1125
1,10
6,22
0100
2101
; 11
252,
98,0
1000
1020
1;
1125
3,15
0,20
1000
2210
;
1125
4,98
,110
0000
102;
11
255,
98,0
1000
2000
2;
1125
6,13
4,20
1100
0200
; 11
257,
117,
2000
0000
00;
1125
8,11
7,21
0100
0000
; 11
259,
117,
0000
0002
00;
1126
0,11
7,21
0001
0100
; 11
261,
117,
0200
0100
00;
1126
3,15
0,12
2200
1210
; 11
264,
72,0
2000
0010
0;
1126
5,72
,000
0000
200;
1126
6,72
,090
0000
000;
11
267,
152,
0902
0000
01;
1126
8,98
,191
0000
202;
11
269,
134,
1900
0000
00;
1127
0,72
,090
0000
100;
11
271,
88,0
9000
0000
0;
1127
6,82
,292
2112
121;
11
277,
152,
0922
0000
01;
1128
1,15
2,09
0909
2000
; 11
282,
152,
0920
0001
01;
1128
3,15
2,09
0200
0101
;
1128
4,15
2,09
0290
0000
; 11
285,
152,
0902
0010
01;
1128
7,82
,291
1201
111;
11
288,
11,1
9202
0100
1;
1129
0,10
2,29
1001
2111
; 11
291,
72,0
9000
0010
0;
1129
2,82
,292
1002
111;
11
293,
82,2
9220
0201
2;
1129
4,82
,291
2102
022;
11
295,
98,0
1002
0100
1;
1148
0,10
6,20
2111
0222
;
1148
1,14
5,20
1112
2122
; 114
82,3
4,22
0212
2221
; 114
83,7
2,00
0000
0200
; 114
84,9
6,00
2000
1000
; 114
85,1
1,22
0100
2210
; 114
86,8
3,01
2001
1121
; 114
87,3
4,22
2222
2221
; 114
88,9
6,00
2000
2011
; 114
89,1
50,2
2012
1022
2; 1
1494
,34,
2122
2221
21; 1
1495
,8,1
0012
0000
1; 1
1496
,106
,010
1121
122;
1149
7,10
6,02
1112
2112
; 114
98,1
1,02
0201
2210
; 114
99,3
4,12
2221
2220
; 115
00,3
4,02
2222
2221
; 115
01,7
2,00
0000
1000
; 115
02,9
6,00
0000
2000
; 115
03,9
6,01
0000
0100
; 115
04,1
8,02
0222
2212
; 115
05,1
21,2
2001
0111
0; 1
1506
,90,
0220
2122
21; 1
1507
,14,
0100
0001
00; 1
1508
,106
,211
0021
122;
1150
9,10
6,01
2002
1022
; 11
510,
145,
0000
0000
00;
1151
1,34
,022
2012
222;
11
512,
96,0
0000
0101
1;
1151
3,96
,000
0001
101;
11
514,
152,
2101
0001
02;
1151
5,98
,200
0000
112;
11
516,
152,
0002
0000
01;
1151
7,15
2,00
1200
0001
; 11
518,
119,
0111
1012
11;
1151
9,96
,000
0001
021;
1152
0,87
,010
0001
001;
11
521,
96,0
0000
0200
1;
1152
3,87
,201
0000
001;
11
524,
117,
1100
0000
00;
1152
5,98
,110
1000
202;
11
526,
117,
2000
0000
02;
1152
7,11
7,10
0000
0211
; 11
528,
87,0
1000
2000
1;
1153
0,96
,000
0201
000;
11
531,
152,
0102
1001
10;
1153
2,96
,000
0001
100;
1153
3,14
,012
2001
001;
115
34,8
7,00
0000
0001
; 11
535,
14,0
1020
0100
1; 1
1536
,49,
2222
2222
22;
1153
7,49
,222
1222
222;
115
38,4
9,22
2222
2222
; 11
539,
49,2
2212
2222
2; 1
1540
,49,
2222
2221
22;
1154
1,49
,222
1222
222;
115
42,4
9,22
2222
2122
; 11
543,
49,2
2222
2222
2; 1
1544
,49,
2222
2222
22;
1154
5,49
,222
2222
122;
11
546,
49,2
2022
2222
2;
1155
9,14
,000
0201
000;
11
560,
134,
0001
1200
00;
1156
1,10
2,10
0200
0201
; 11
562,
134,
2200
0000
01;
1156
3,13
4,20
0000
1001
; 11
564,
134,
2000
0010
01;
1156
5,98
,000
0100
102;
11
566,
14,0
0002
0100
0;
1156
7,11
7,12
0010
0201
;
1156
8,10
2,11
0111
2201
; 11
569,
102,
1010
0221
00;
1157
0,10
2,21
1000
2202
; 11
571,
102,
2020
0111
02;
1157
2,10
2,22
1111
1001
; 11
573,
102,
2020
1112
00;
1157
4,13
4,20
0020
0100
; 11
575,
134,
2200
0010
00;
1157
6,13
4,22
0000
1000
; 11
577,
134,
2100
0010
00;
1157
8,13
4,01
0020
0000
;
1157
9,13
4,21
0000
1000
; 11
580,
134,
2000
0222
00;
1158
1,15
0,22
2021
1221
; 11
582,
150,
2201
2212
21;
1158
3,15
0,22
2022
0221
; 11
584,
150,
2220
2112
21;
1158
5,99
,000
0000
102;
11
586,
99,1
1001
0010
1;
1158
7,99
,200
2101
100;
11
588,
152,
0202
0000
10;
1158
9,72
,000
1000
002;
1159
0,15
2,02
0010
0001
; 11
591,
99,1
0000
0020
2; 1
1593
,99,
2102
0121
01;
1159
4,99
,200
0112
101;
115
95,9
8,20
1020
0101
; 11
596,
98,2
1002
0000
0; 1
1597
,98,
2000
0010
21;
1159
8,98
,200
0010
012;
115
99,9
8,21
0001
0112
; 11
600,
98,2
1011
1010
2; 1
1601
,91,
2222
0012
22;
1160
2,28
,222
2122
212;
1160
3,91
,221
0001
110;
116
04,1
50,2
2122
1022
1; 1
1605
,28,
2212
2000
22; 1
1606
,150
,221
1110
220;
116
07,8
7,10
0000
0101
; 116
08,2
9,22
2022
2122
; 116
09,8
8,00
0000
0000
; 116
10,8
7,01
0010
0101
; 116
11,8
7,00
0010
0001
; 116
12,1
02,0
1020
1100
1; 1
1613
,98,
2010
0102
21; 1
1614
,99,
2100
0020
00;
1161
5,98
,200
0000
111;
11
616,
99,2
0000
1010
1;
1161
7,99
,010
0010
102;
11
618,
99,2
0001
0210
2;
1161
9,15
2,01
0220
0001
; 11
620,
152,
0202
2000
00;
1162
1,15
2,01
0200
1001
; 11
622,
152,
0202
0020
01;
1162
3,98
,200
0101
000;
11
624,
152,
0002
1011
01;
1162
5,11
9,21
1210
2222
;
1162
6,11
9,22
1221
2221
; 11
627,
119,
2110
1012
00;
1162
8,11
9,22
1021
2202
; 11
629,
96,0
0000
0101
1;
1163
0,11
,212
2112
002;
11
631,
11,2
1112
0222
2;
1163
2,99
,200
0012
201;
11
633,
88,2
1000
0210
0;
1163
4,99
,010
0002
100;
11
635,
11,2
2222
1221
0;
1163
6,87
,200
0000
100;
1163
7,11
7,12
0010
0101
; 116
38,9
6,11
1010
1110
; 116
39,1
1,01
1111
1121
; 116
40,1
1,11
1100
1212
; 116
41,1
1,21
1010
2220
; 116
42,9
6,20
0000
2000
; 116
43,3
1,21
2011
1221
; 116
44,3
1,02
0001
2100
; 116
45,8
2,22
0021
2022
; 116
46,8
2,22
1001
2022
; 116
47,1
50,2
2221
1212
0; 1
1648
,11,
2120
1010
10;
1164
9,15
0,22
0021
1222
; 11
650,
11,2
1202
0100
1;
1165
1,10
6,21
0122
1122
; 11
652,
106,
2220
2112
22;
1165
3,10
9,21
1100
1210
; 11
654,
58,2
2222
2222
2;
1165
5,11
,212
0201
000;
11
656,
58,2
2220
0222
2;
1165
7,11
7,20
0002
0001
; 11
659,
117,
0012
0000
00;
1166
0,10
9,20
0202
2221
;
SELECTION IN THE HOUSE MOUSE HYBRID ZONE 613
© 2005 The Linnean Society of London, Biological Journal of the Linnean Society, 2005, 84, 593–616
1166
1,42
,222
1202
222;
116
62,4
2,02
2102
2222
; 11
667,
106,
0211
0211
00;
1166
8,42
,222
2222
222;
116
69,4
2,22
2219
2222
; 11
670,
42,0
2211
2222
2; 1
1671
,42,
2222
2222
21;
1167
2,42
,022
2222
221;
116
73,4
2,22
2222
2222
; 11
674,
42,0
2222
2222
2; 1
1677
,42,
0222
2222
22;
1167
8,58
,222
2022
221;
1168
1,42
,022
2022
222;
11
682,
88,0
0002
0000
0;
1168
3,42
,222
2022
222;
11
685,
134,
2100
2210
00;
1168
6,15
2,01
0200
0101
; 11
687,
152,
0102
0000
00;
1168
9,15
2,01
0100
0101
; 11
692,
153,
0100
0100
00;
1169
4,10
9,21
1100
2122
; 11
695,
11,1
2020
1221
0;
1169
6,15
2,01
0200
0101
;
1169
7,42
,222
2222
222;
116
98,4
2,22
2122
2222
; 11
699,
42,2
2222
2212
2; 1
1700
,87,
0000
0010
00;
1170
2,15
2,02
0200
0101
; 11
703,
87,0
0000
0110
0; 1
1710
,150
,121
1212
220;
161
88,1
2,20
2022
1912
; 16
189,
12,0
2222
1290
0; 1
6190
,136
,122
2212
101;
161
91,6
,222
2222
120;
161
92,6
,202
2222
911;
1619
3,53
,222
0222
901;
161
94,5
3,22
2022
2922
; 16
195,
53,2
2202
2292
2; 1
6196
,53,
2220
2229
21;
1619
7,53
,222
9999
929;
161
98,5
3,22
2022
2920
; 16
199,
53,2
9202
2290
1; 1
6200
,53,
2220
2229
22;
1620
1,53
,292
0222
902;
162
02,6
,222
2292
912;
162
03,5
3,22
2022
2921
; 16
204,
53,2
2202
2292
2;
1620
5,53
,292
0222
902;
162
06,5
3,22
0022
2221
; 16
207,
136,
2220
2111
12;
1620
8,53
,220
0221
021;
162
09,5
3,29
2122
2911
; 16
210,
53,2
2202
2222
1; 1
6211
,53,
2221
2229
21;
1621
2,53
,290
0222
902;
162
13,5
3,22
0022
2902
; 16
214,
53,2
2102
2290
2; 1
6215
,53,
2220
2222
21;
1621
6,53
,292
0222
902;
1621
7,53
,222
0222
922;
162
18,5
3,22
2022
2922
; 16
219,
53,2
2202
2292
1; 1
6220
,53,
2220
2229
21;
1622
1,53
,222
0222
121;
162
22,5
3,22
2022
2922
; 16
223,
53,2
2202
2292
1; 1
6224
,53,
2220
2229
22;
1622
5,76
,222
2222
901;
162
26,5
3,22
2022
2121
; 16
227,
53,2
2202
2212
1; 1
6228
,6,2
1211
1092
2;
1622
9,53
,222
0222
920;
162
30,4
8,22
2029
2920
; 16
231,
48,1
2202
2192
0; 1
6232
,48,
1229
2219
20;
1623
3,48
,999
9221
990;
162
34,7
8,20
2209
1900
; 16
235,
77,2
2221
1292
0; 1
6236
,78,
1029
9999
29;
1623
7,25
,220
2112
222;
162
38,7
8,22
2002
1001
; 16
239,
48,9
1001
2100
1; 1
6240
,22,
2020
2121
20;
1624
1,48
,220
0220
220;
162
42,4
8,11
2112
2000
; 162
43,4
8,12
2121
2120
; 162
44,1
22,2
1222
1299
0; 1
6245
,48,
2210
1111
22; 1
6251
,122
,102
1212
290;
162
52,1
22,2
2221
0221
0; 1
6253
,47,
2120
0122
11; 1
6254
,44,
1110
0021
20; 1
6255
,44,
0901
0001
00; 1
6256
,44,
0101
0001
00; 1
6257
,44,
2100
0102
00;
1625
8,47
,221
2212
111;
162
59,7
8,12
1202
2021
; 16
260,
47,2
2211
9212
1; 1
6261
,47,
2100
2110
22;
1626
2,47
,910
1110
022;
162
63,7
7,22
0011
2202
; 16
264,
78,2
1211
2112
2; 1
6265
,122
,112
2102
122;
162
66,5
3,22
2022
2222
; 16
267,
19,2
2112
0112
2; 1
6268
,19,
2222
0012
22;
1627
0,53
,222
0222
121;
1627
1,6,
1212
2220
12;
1627
2,47
,101
0021
111;
162
73,1
9,21
1200
2122
; 16
274,
103,
0221
2020
21;
1627
5,19
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0212
222;
162
76,1
05,9
2920
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2; 1
6277
,78,
2221
1910
01;
1627
8,78
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2011
122;
162
79,7
8,22
2000
0121
; 16
280,
78,2
1200
2102
2; 1
6281
,78,
1220
1201
22;
1628
2,78
,111
0120
022;
1628
3,78
,122
0121
020;
162
84,5
3,22
2022
2221
; 162
85,5
3,22
2022
2022
; 162
86,5
3,22
2022
2122
; 162
87,5
3,22
2022
2121
; 162
88,5
3,22
2022
2222
; 162
89,5
3,22
2022
2121
; 162
90,5
3,22
2022
2220
; 162
91,5
3,22
2922
2020
; 162
92,5
3,22
2222
2120
; 162
93,1
05,0
2220
2221
0; 1
6294
,105
,222
2211
119;
1629
5,10
5,22
2212
2010
; 162
96,1
16,2
2102
1212
2; 1
6297
,47,
2022
1222
12; 1
6298
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2021
2212
09; 1
6299
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1222
1022
12; 1
6300
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1212
2120
21; 1
6301
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2212
2121
22; 1
6302
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1202
1011
00; 1
6303
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1221
2211
1; 1
6304
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1202
0011
10; 1
6305
,141
,221
2222
110;
163
06,3
8,02
2221
2220
;
1630
7,6,
2112
1221
19;
1630
8,53
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2122
122;
163
09,4
5,22
2022
2221
; 16
311,
38,0
1222
0112
2; 1
6312
,53,
2220
2222
21;
1631
3,53
,122
0222
022;
163
14,5
3,22
2022
2021
; 16
315,
44,0
2020
0010
0; 1
6316
,38,
2212
2222
20;
1631
7,14
1,12
2222
2222
; 16
318,
53,2
2222
2202
2; 1
6319
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2222
102;
1632
0,14
1,22
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2102
; 16
321,
141,
2202
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22;
1632
2,14
1,22
1222
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; 16
323,
141,
2202
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11;
1632
4,38
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2202
121;
16
325,
103,
0121
0220
20;
1632
6,10
3,02
2202
2021
; 16
327,
110,
1220
2121
00;
1632
8,53
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221;
16
329,
53,2
2202
2212
1;
1633
0,53
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0222
222;
1633
1,53
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0222
121;
16
332,
53,2
2202
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1;
1633
4,13
9,22
2221
2112
; 16
335,
139,
2222
2222
22;
1633
6,11
0,12
2211
2101
; 16
337,
141,
2202
2221
21;
1633
8,14
1,22
0222
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; 16
339,
71,0
0002
1000
0;
1634
0,71
,100
0101
000;
16
342,
71,1
0000
1001
0;
1634
3,52
,000
0000
101;
1634
4,78
,222
1112
020;
163
45,7
7,22
2111
2220
; 16
346,
77,2
2012
1222
2; 1
6347
,77,
2201
1122
20;
1634
8,77
,221
0010
220;
163
49,5
2,00
1122
2001
; 16
350,
48,2
1222
0222
1; 1
6351
,59,
1222
2111
12;
1635
2,48
,212
0111
112;
163
53,4
8,12
1120
2110
; 16
354,
48,1
2110
1202
9; 1
6355
,48,
1112
2222
12;
1635
6,13
9,22
1020
9220
; 16
357,
139,
2222
2221
11;
1635
8,13
9,22
1222
2121
; 16
359,
132,
1122
2212
12;
1636
0,71
,000
0110
010;
16
361,
139,
1222
2222
22;
1636
2,13
9,21
2121
2022
; 16
363,
59,1
1012
1122
2;
1636
4,14
2,10
0101
0200
; 16
365,
142,
2011
0111
00;
1636
6,15
,211
1229
201;
1636
7,13
9,22
2222
9220
; 16
368,
139,
2222
2291
12;
1636
9,13
9,22
2222
9121
; 16
370,
139,
2222
2291
01;
1637
1,14
2,10
2100
9101
; 16
372,
142,
2000
1192
01;
1637
3,14
2,20
0011
9101
; 16
374,
132,
2292
2199
92;
1637
5,14
2,10
1211
9201
; 16
376,
105,
1211
2012
21;
1637
7,10
5,21
2210
1122
;
1637
8,80
,101
2121
100;
16
379,
142,
1012
2112
00;
1638
0,3,
2211
1201
12;
1638
1,38
,212
1202
121;
16
382,
38,2
2122
0222
2;
1638
3,10
4,22
2120
2121
; 16
384,
105,
2222
1121
21;
1638
5,77
,220
1012
221;
16
386,
46,2
2211
1222
2;
1638
7,13
9,90
9220
2992
; 16
388,
154,
0011
0211
00;
1638
9,15
4,00
0102
1101
; 16
390,
142,
2011
0012
01;
1639
1,14
4,22
1121
2220
; 16
392,
26,2
0101
2212
0;
1639
3,14
2,20
1292
2102
; 16
394,
142,
2121
9111
00;
1639
5,10
5,22
0221
2111
; 16
396,
105,
1221
2022
10;
1639
7,13
9,22
2220
2922
; 16
398,
15,1
2122
2020
0;
1639
9,15
4,00
0100
0100
;
1640
0,14
2,11
2211
2211
; 16
401,
142,
1220
0111
01;
1640
2,6,
9991
2129
91;
1640
3,47
,222
2022
202;
16
404,
60,2
1201
0202
2;
1640
5,14
2,10
1122
0202
; 16
406,
80,2
1122
1012
0;
1640
7,14
2,20
0200
2100
; 16
408,
104,
2222
1212
21;
1640
9,14
2,10
1222
1201
; 16
410,
142,
2001
1021
02;
1641
1,14
2,20
9110
2202
; 16
412,
103,
2222
2121
22;
1641
3,14
2,10
2122
1201
; 16
414,
151,
0010
1001
02;
1641
5,15
1,00
1000
0000
; 16
416,
142,
2022
1012
00;
1641
7,10
5,21
0022
2222
; 16
418,
104,
2102
1222
20;
1641
9,13
8,02
0222
1010
; 16
420,
105,
2202
2022
21;
1642
1,12
3,22
2222
2222
;
1642
2,10
,002
0101
000;
16
423,
126,
2111
1211
20;
1642
4,64
,222
2222
212;
16
425,
126,
1121
1212
11;
1642
6,10
,001
0001
200;
16
427,
105,
1222
9022
21;
1642
8,12
3,12
0222
2222
; 16
429,
123,
2202
2222
22;
1643
0,12
3,22
2222
2222
; 16
431,
123,
2222
2222
22;
1643
2,12
3,22
2222
2222
;
1643
3,12
3,22
2222
2222
; 16
434,
123,
9222
2221
12;
1643
5,12
3,92
2212
2222
; 16
436,
123,
9212
2222
12;
1643
7,12
3,92
2222
2212
; 16
438,
123,
9920
2122
21;
1643
9,27
,222
2222
202;
16
440,
105,
2222
1221
21;
1644
1,10
4,21
1212
2122
; 16
442,
81,2
2122
2212
2;
1644
3,11
0,02
2010
2000
;
1644
4,10
,010
1101
100;
16
445,
10,0
0012
0120
0;
1644
6,13
7,01
9020
0000
; 16
447,
143,
9901
1021
10;
1644
8,65
,212
2202
221;
16
449,
38,1
2212
1202
2;
1645
0,38
,929
2222
992;
16
451,
142,
2191
2112
01;
1645
2,14
2,11
9211
1201
; 16
453,
38,2
2922
2220
2;
1645
4,38
,211
1121
212;
1645
5,38
,919
1222
992;
16
456,
142,
1012
1022
01;
1645
7,97
,020
0011
201;
16
458,
142,
1001
1010
00;
1645
9,65
,221
2192
211;
16
461,
139,
2221
2221
10;
1646
2,11
8,21
2002
0010
; 16
463,
65,2
1212
1222
1;
1646
4,65
,212
2212
212;
16
465,
65,2
2221
0220
0;
1646
6,65
,212
2202
220;
1646
7,13
2,22
1021
2122
; 16
468,
124,
0200
1000
00;
1646
9,27
,222
2212
222;
16
470,
62,2
2122
0222
0;
1647
1,62
,120
2111
220;
16
472,
101,
9990
0009
90;
1647
3,15
1,01
0000
0002
; 16
474,
151,
0000
0001
01;
1647
6,10
,010
1101
101;
16
478,
65,2
2212
1222
1;
1647
9,14
0,01
0299
9100
;
1648
0,10
0,00
0200
1000
; 16
481,
100,
0100
0021
00;
1648
2,10
0,01
0200
1100
; 16
483,
100,
0100
0011
00;
1648
4,10
,000
0091
100;
16
485,
65,1
1221
9222
2;
1648
6,65
,202
2192
022;
16
487,
61,1
1222
9201
2;
1648
8,65
,222
2292
220;
16
489,
127,
9911
2920
19;
1649
0,13
3,00
0209
9000
;
1649
1,10
,000
0199
000;
164
92,1
0,00
0029
9200
; 164
93,1
0,00
0010
0100
; 164
94,1
0,00
0010
0200
; 164
95,1
0,00
0000
0000
; 164
96,1
0,01
0000
1200
; 164
97,7
4,22
1110
1220
; 164
98,1
18,1
0011
0101
1; 1
6499
,74,
2221
2022
22; 1
6500
,74,
2021
2091
22; 1
6502
,140
,000
0000
000;
165
03,3
3,01
0020
2000
;
1650
4,33
,010
0201
000;
16
506,
151,
0000
1001
00;
1650
7,15
1,00
0000
1200
; 16
508,
151,
0000
1091
00;
1650
9,15
1,00
0020
9100
; 16
510,
151,
0000
1011
01;
1651
1,13
8,12
2222
1110
; 16
512,
61,1
1022
1222
2;
1651
3,61
,102
2212
220;
16
514,
139,
2222
2221
20;
1651
5,13
9,22
2222
2011
;
1651
6,13
9,22
2222
2220
; 16
517,
139,
2222
2222
11;
1651
8,13
9,22
2922
2221
; 16
519,
139,
2222
2221
21;
1652
0,13
9,22
2121
2211
; 16
521,
139,
2221
2120
22;
1652
2,13
9,21
2122
2120
; 16
523,
139,
2222
2222
22;
1652
4,13
9,22
2221
2110
; 16
525,
139,
2122
2122
12;
1652
6,13
9,22
2099
9029
;
1652
7,13
9,20
2099
9029
; 16
528,
139,
2129
9991
19;
1652
9,14
1,22
2222
2112
; 16
530,
139,
2222
2221
12;
1653
1,14
1,22
1299
9119
; 16
532,
141,
2102
9991
19;
1653
3,14
1,22
0222
2202
; 16
534,
141,
2212
2221
12;
1653
5,14
1,22
0222
2221
; 16
536,
141,
2202
9991
19;
1653
7,14
1,22
0299
9219
;
1653
8,14
1,22
0222
2222
; 16
539,
141,
9202
2221
10;
1654
0,14
1,22
2222
2212
; 16
541,
141,
9212
2221
11;
1654
2,14
1,22
0299
9109
; 16
543,
141,
2212
2221
10;
1654
4,14
1,12
1222
2200
; 16
545,
141,
2212
2222
12;
1654
6,14
1,22
1222
2222
; 16
547,
141,
2202
2221
11;
1654
8,14
1,22
0222
2212
;
1654
9,52
,000
0999
009;
165
50,7
1,00
0099
9009
; 165
51,5
2,00
0099
9119
; 165
52,5
2,00
0099
9019
; 165
53,5
2,01
0099
9009
; 165
54,5
2,00
0099
9019
; 165
55,5
2,01
0099
9019
; 165
56,5
2,00
0099
9209
; 165
57,5
2,01
0099
9009
; 165
58,1
27,2
2211
2222
2; 1
6559
,127
,212
2202
121;
165
60,1
27,2
2199
9922
9;
1656
1,12
7,12
0222
2121
; 16
562,
127,
2222
0122
22;
1656
3,14
2,21
1001
1200
; 16
564,
142,
2021
1102
01;
1656
5,14
2,21
1000
2200
; 16
566,
142,
2101
0021
01;
1656
7,14
2,20
0200
2100
; 16
568,
142,
2002
0221
00;
1656
9,14
2,20
1220
2210
; 16
570,
142,
2002
0020
00;
1657
1,14
2,20
0299
9119
;
1657
2,12
3,22
2299
9129
; 165
73,1
23,2
2229
9922
9; 1
6574
,43,
2221
9992
09; 1
6575
,43,
2222
9992
29; 1
6576
,43,
2221
9992
29; 1
6577
,43,
2222
9992
29; 1
6578
,43,
2221
9992
29; 1
6579
,43,
2222
9992
29; 1
6580
,43,
2221
9992
29; 1
6581
,43,
2929
9992
29; 1
6582
,43,
2929
9992
29; 1
6583
,52,
0929
9991
19;
1658
4,70
,192
1201
111;
165
85,5
2,09
2999
9209
; 16
586,
52,0
9099
9911
9; 1
6587
,43,
1929
9992
29;
1658
8,43
,292
9999
229;
165
89,4
3,22
0999
9229
; 16
590,
43,1
2009
9992
9; 1
6591
,43,
1202
9999
29;
1659
2,43
,220
1999
929;
165
93,4
3,22
0222
2922
; 16
594,
38,1
2029
9992
9; 1
6595
,38,
2212
2022
12;
1659
6,38
,220
2222
222;
165
97,3
8,20
2211
1212
; 16
598,
38,2
2022
2192
2; 1
6599
,38,
2202
2229
10;
1660
0,38
,220
2999
929;
166
01,3
8,92
0222
2922
; 16
602,
38,2
2022
2292
2; 1
6603
,38,
0202
1129
20;
1660
4,38
,120
2999
929;
166
05,3
8,22
0222
2922
; 16
606,
38,2
2022
2292
0; 1
6607
,38,
2202
2029
21;
1660
8,38
,020
2999
929;
166
09,1
42,1
0219
9990
9; 1
6610
,142
,202
1002
901;
166
11,1
42,0
1119
9990
9; 1
6612
,142
,211
1020
901;
166
13,1
42,2
9120
0200
1; 1
6614
,142
,190
9999
019;
166
15,1
5,19
0122
2210
; 166
16,1
5,29
2222
0212
; 166
17,3
,091
2222
221;
166
18,3
,992
9999
219;
166
19,6
,211
1221
121;
1662
0,6,
2229
9990
19;
1662
4,32
,022
2112
119;
166
25,6
4,12
2210
2220
; 16
626,
65,2
1222
2221
9; 1
6627
,65,
2212
1121
10;
1662
8,65
,202
2212
120;
166
29,6
5,22
2211
2212
; 16
630,
65,2
2122
0221
2; 1
6631
,65,
2222
1121
21;
1663
2,65
,212
9909
210;
166
33,1
42,9
9999
0999
2; 1
6884
,192
,202
2922
222;
1688
5,17
3,22
2192
9222
; 16
886,
174,
0000
9090
00;
1688
7,17
8,92
9292
2992
; 16
888,
184,
0009
9900
00;
1688
9,18
5,22
9192
2992
; 16
890,
185,
2222
9222
22;
1689
1,17
6,22
2292
2222
; 16
892,
174,
0000
9091
00;
1689
3,17
4,00
0099
9000
; 16
894,
174,
0090
9099
90;
1689
5,19
1,22
2292
2122
;
1689
6,18
7,00
0090
9000
; 16
897,
187,
0002
9090
00;
1689
8,19
1,22
2199
2222
; 16
899,
191,
2221
9922
02;
1690
0,17
5,00
0999
0900
; 16
901,
184,
0009
9900
00;
1690
2,19
2,92
9292
2992
; 16
903,
174,
0000
9090
00;
1690
4,17
4,00
0090
9000
; 16
905,
182,
0009
9900
00;
1690
8,17
3,02
2999
9229
;
1690
9,19
1,22
2999
2222
; 16
910,
187,
2009
9990
09;
1691
1,18
7,00
0999
9009
; 16
912,
191,
2229
9922
29;
1691
3,18
7,00
0999
9009
; 16
914,
189,
0009
9990
09;
1691
5,17
7,22
9999
9999
; 16
916,
177,
2229
9922
09;
1691
7,18
7,00
0999
9009
; 16
918,
184,
0009
9900
00;
1691
9,18
4,00
0999
0000
;
1692
0,18
0,22
2999
9229
; 16
921,
180,
2299
9999
99;
1692
2,19
1,22
2999
2222
; 16
923,
184,
0009
9900
00;
1692
4,18
4,00
9999
0090
; 16
925,
184,
0099
9900
90;
1692
6,18
0,22
9999
9999
; 16
927,
180,
2299
9999
99;
1692
8,17
7,22
2999
9229
; 16
929,
177,
2229
9922
22;
1693
0,17
7,22
2999
9229
;
1693
1,17
7,22
2999
2229
; 16
933,
184,
0009
9900
00;
1693
4,18
5,22
9999
9999
; 16
935,
177,
2229
9922
29;
1693
6,17
7,22
2999
2222
; 16
937,
174,
0009
9920
09;
1693
8,18
0,22
9999
9999
; 16
939,
191,
2229
9922
22;
1694
1,17
3,22
2999
2922
; 16
943,
192,
2229
9922
29;
1694
6,18
4,00
0999
0000
;
1694
9,18
4,00
0999
0000
; 16
950,
179,
0009
9900
09;
1695
2,18
4,00
0999
0090
; 16
953,
178,
2229
9922
29;
1695
4,19
0,00
0999
0000
; 16
955,
188,
1009
9900
00;
1695
6,17
9,00
0999
9000
; 16
958,
177,
2229
9922
22;
1695
9,18
2,00
0999
0000
; 16
960,
177,
2229
9922
29;
1696
1,17
7,22
2999
2222
;
1696
2,19
7,92
9999
9999
; 16
963,
177,
2229
9922
29;
1696
4,18
5,92
9999
9999
; 16
965,
185,
9299
9999
99;
1696
6,17
7,22
2999
9929
; 16
967,
177,
2229
9922
22;
1696
8,18
0,22
2999
2222
; 16
969,
191,
2229
9922
29;
1697
0,18
2,00
0999
0000
; 16
971,
182,
0009
9900
00;
1697
2,18
2,00
0999
0000
;
1697
3,18
2,00
0999
0000
; 16
974,
182,
1009
9900
00;
1697
5,18
1,00
0999
0000
; 16
977,
184,
0009
9900
00;
1697
8,18
2,00
0999
0000
; 16
979,
182,
0009
9900
00;
1698
0,18
2,00
0999
0000
; 16
981,
182,
0009
9900
00;
1698
2,17
7,22
2999
2122
; 16
983,
194,
9299
9999
99;
1698
4,17
5,00
0999
0000
;
1698
5,18
4,00
0999
0000
; 16
986,
184,
0009
9900
00;
1698
7,18
4,00
0999
0000
; 16
988,
179,
0009
9990
09;
1698
9,17
4,02
0999
9009
; 16
990,
174,
0009
9990
09;
1699
1,17
4,00
0999
9009
; 16
993,
174,
0009
9990
09;
1699
4,17
4,00
0999
9000
; 16
995,
174,
0009
9921
09;
1699
6,17
9,90
9999
9999
;
1699
7,17
4,90
9999
9999
; 16
998,
174,
9099
9999
99;
1699
9,18
2,00
0999
0000
; 17
001,
177,
9299
9999
99;
1700
2,17
7,22
2999
2222
; 17
003,
177,
2229
9922
29;
1700
4,17
7,92
9999
9999
; 17
005,
177,
2229
9922
29;
1700
6,17
7,22
2999
9929
; 17
007,
178,
2229
9922
21;
1700
8,17
8,22
1999
2129
;
1700
9,17
6,22
2999
9222
; 17
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
;
614 N. RAUFASTE ET AL.
© 2005 The Linnean Society of London, Biological Journal of the Linnean Society, 2005, 84, 593–616
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;
SELECTION IN THE HOUSE MOUSE HYBRID ZONE 615
© 2005 The Linnean Society of London, Biological Journal of the Linnean Society, 2005, 84, 593–616
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
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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
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056;
16
454,
38,1
998,
0380
38,0
7107
1,05
4058
,065
075,
0540
70,0
5005
0;
1645
5,38
,199
8,03
8038
,071
071,
0580
62,0
5506
5,05
4070
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050;
16
594,
38,1
998,
0380
38,0
7108
3,05
8062
,065
075,
0620
66,0
4604
6;
1659
5,38
,199
8,03
8050
,083
083,
0620
62,0
5506
0,05
8066
,046
058;
16
596,
38,1
998,
0380
38,0
7107
1,06
2062
,055
075,
0540
70,0
5005
8;
1659
7,38
,199
8,03
8050
,071
071,
0500
70,0
6006
5,07
0070
,046
046;
16
598,
38,1
998,
0380
38,0
7107
1,05
8058
,075
075,
0540
62,0
5805
8;
1659
9,38
,199
8,03
8050
,068
080,
0540
62,0
5506
0,06
2066
,050
056;
16
600,
38,1
998,
0380
50,0
7107
4,05
0058
,055
065,
0540
54,0
4605
6;
1660
1,38
,199
8,03
8054
,071
077,
0500
62,0
5507
5,05
4070
,056
056;
16
602,
38,1
998,
0380
38,0
7107
1,06
2062
,055
075,
0700
70,0
5605
0;
1660
3,38
,199
8,03
8038
,071
071,
0580
62,0
5507
5,05
4054
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058;
16
604,
38,1
998,
0380
38,0
7108
3,06
2066
,065
075,
0540
62,0
4604
6;
1660
5,38
,199
8,03
8038
,068
077,
0580
66,0
6507
5,05
4070
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056;
16
606,
38,1
998,
0380
38,0
7107
1,05
8062
,055
065,
0540
62,0
4605
2;
1660
7,38
,199
8,03
8038
,071
083,
0660
66,0
5506
5,05
4062
,052
058;
16
608,
38,1
998,
0380
38,0
7107
1,05
8062
,065
075,
0540
70,0
5005
6;
1657
4,43
,199
8,04
6050
,068
071,
0580
62,0
6506
5,06
6066
,058
058;
16
575,
43,1
998,
0380
38,0
6807
1,07
0070
,065
070,
0620
62,0
5805
8;
1657
6,43
,199
8,03
8038
,071
077,
0700
70,0
6507
0,05
0050
,058
058;
16
577,
43,1
998,
0380
38,0
6807
1,06
6070
,065
070,
0500
50,0
5805
8;
1657
8,43
,199
8,03
8038
,071
077,
0580
62,0
6506
5,04
6050
,056
056;
16
579,
43,1
998,
0380
38,0
7107
1,05
8062
,050
065,
0500
50,0
4605
6;
1658
0,43
,199
8,03
8038
,068
071,
0700
70,0
6507
0,06
2066
,058
058;
16
581,
43,1
998,
0380
38,0
7107
1,06
2070
,065
065,
0620
66,0
5805
8;
1658
2,43
,199
8,03
8038
,071
077,
0580
70,0
6507
0,04
6050
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058;
16
587,
43,1
998,
0380
38,0
5305
3,06
6066
,070
070,
0620
62,0
5805
8;
1658
8,43
,199
8,03
8038
,068
071,
0000
00,0
6506
5,06
2062
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058;
16
589,
43,1
998,
0380
38,0
7107
7,06
2066
,065
070,
0460
50,0
5605
6;
1659
0,43
,199
8,03
8038
,071
077,
0580
58,0
7007
0,04
6050
,056
056;
16
591,
43,1
998,
0380
38,0
6807
1,00
0000
,070
070,
0620
62,0
5805
8;
1659
2,43
,199
8,03
8038
,071
071,
0000
00,0
6507
0,04
6050
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058;
16
593,
43,1
998,
0380
38,0
7107
1,00
0000
,065
065,
0460
50,0
5805
8;
1625
4,44
,199
8,04
2054
,050
050,
0460
62,0
5505
5,05
4054
,046
046;
16
255,
44,1
998,
0380
38,0
5005
0,06
6066
,060
060,
0540
54,0
4604
8;
1625
6,44
,199
8,03
8050
,053
068,
0000
00,0
5506
0,00
0000
,046
050;
16
257,
44,1
998,
0380
46,0
5006
8,05
4066
,055
060,
0420
50,0
4605
0;
1630
4,44
,199
8,03
8050
,050
068,
0620
66,0
5506
0,05
0074
,046
050;
16
315,
44,1
998,
0420
46,0
5005
0,05
4054
,055
055,
0420
62,0
4805
2;
1638
6,46
,199
8,03
8038
,050
053,
0620
66,0
6506
5,06
2066
,000
000;
16
253,
47,1
998,
0460
46,0
7107
1,04
6046
,050
065,
0500
74,0
4605
0;
1625
8,47
,199
8,05
0050
,065
065,
0500
50,0
6506
5,07
4074
,046
046;
16
260,
47,1
998,
0380
38,0
5006
5,05
8058
,065
065,
0500
74,0
4605
0;
1626
1,47
,199
8,03
8038
,071
074,
0620
62,0
6506
5,03
8038
,050
050;
16
262,
47,1
998,
0500
50,0
5307
4,06
2062
,060
065,
0380
38,0
5005
0;
1626
9,47
,199
8,03
8050
,071
074,
0620
62,0
6506
5,03
8054
,050
050;
16
272,
47,1
998,
0380
50,0
7107
1,06
2066
,060
065,
0380
54,0
5005
0;
1629
7,47
,199
8,03
8050
,071
074,
0580
62,0
6006
5,06
2074
,046
058;
16
298,
47,1
998,
0380
50,0
7107
1,06
2062
,065
065,
0380
74,0
4805
0;
1629
9,47
,199
8,03
8038
,071
071,
0620
62,0
5005
0,04
6050
,050
058;
16
403,
47,1
998,
0380
38,0
5605
6,04
6058
,060
065,
0740
74,0
5605
6;
1623
0,48
,199
8,03
8042
,071
071,
0660
66,0
5506
5,05
4070
,046
046;
16
231,
48,1
998,
0380
42,0
7107
1,06
6066
,055
065,
0580
70,0
5005
6;
1623
2,48
,199
8,03
8038
,071
071,
0580
74,0
6506
5,05
8070
,046
056;
16
233,
48,1
998,
0380
42,0
5607
1,06
6066
,065
065,
0580
70,0
4605
6;
1623
9,48
,199
8,03
8054
,053
074,
0460
74,0
5506
5,05
0058
,046
046;
16
241,
48,1
998,
0380
42,0
5607
1,06
6074
,065
065,
0580
58,0
4606
0;
1624
2,48
,199
8,03
8038
,050
074,
0660
74,0
5506
5,05
4070
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060;
16
243,
48,1
998,
0380
38,0
5607
1,05
8074
,065
065,
0540
70,0
4605
0;
1624
5,48
,199
8,05
0050
,050
071,
0580
58,0
5505
5,07
0070
,050
060;
16
246,
48,1
998,
0380
50,0
5007
1,05
8074
,065
065,
0540
70,0
5005
6;
1624
7,48
,199
8,03
8050
,050
050,
0580
74,0
6506
5,05
4054
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056;
16
248,
48,1
998,
0380
50,0
7107
1,05
8074
,065
065,
0540
54,0
5005
6;
1624
9,48
,199
8,03
8050
,071
071,
0580
74,0
6506
5,05
4070
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056;
16
250,
48,1
998,
0380
50,0
5005
0,05
8074
,065
065,
0460
54,0
4605
6;
1635
0,48
,199
8,05
0050
,050
050,
0580
58,0
5005
0,05
0050
,046
056;
16
352,
48,1
998,
0380
38,0
5005
0,07
4074
,055
065,
0580
58,0
4606
0;
1635
3,48
,199
8,03
8038
,053
071,
0620
62,0
6006
5,03
8054
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060;
16
354,
48,1
998,
0380
42,0
5607
1,06
6074
,065
065,
0540
70,0
5005
6;
1635
5,48
,199
8,03
8038
,074
074,
0620
62,0
6506
5,05
4054
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050;
16
349,
52,1
998,
0420
50,0
5305
6,05
4054
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050,
0460
50,0
4604
6;
1654
9,52
,199
8,03
8050
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056,
0540
62,0
7007
0,04
6046
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050;
16
551,
52,1
998,
0380
50,0
5005
3,05
4058
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055,
0420
46,0
5005
0;
1655
2,52
,199
8,05
0050
,050
056,
0460
54,0
5006
0,04
6050
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050;
16
553,
52,1
998,
0380
50,0
4407
1,04
6062
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070,
0460
50,0
4805
0;
1655
4,52
,199
8,03
8050
,053
056,
0460
62,0
7007
0,04
6046
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050;
16
555,
52,1
998,
0500
50,0
5305
6,04
6046
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070,
0460
50,0
4605
2;
1655
6,52
,199
8,04
6050
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050,
0540
62,0
7007
0,04
2046
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052;
16
557,
52,1
998,
0380
50,0
5305
6,05
4062
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070,
0460
46,0
4805
0;
1658
3,52
,199
8,03
8050
,053
056,
0620
62,0
7007
0,04
6046
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050;
16
585,
52,1
998,
0500
50,0
4404
4,05
4058
,065
065,
0500
50,0
4805
0;
1658
6,52
,199
8,05
0050
,053
053,
0420
46,0
7007
0,04
6050
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050;
16
635,
52,1
998,
0500
50,0
4405
3,04
6054
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070,
0460
46,0
0000
0;
1619
3,53
,199
8,03
8038
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074,
0660
66,0
6006
0,05
8070
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058;
16
194,
53,1
998,
0000
00,0
0000
0,00
0000
,065
065,
0000
00,0
5805
8;
1619
5,53
,199
8,03
8038
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068,
0500
66,0
6506
5,05
8070
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056;
16
196,
53,1
998,
0380
38,0
6806
8,05
4066
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060,
0700
70,0
5605
6;
1619
7,53
,199
8,03
8038
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000,
0540
58,0
6506
5,05
8070
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058;
16
198,
53,1
998,
0380
38,0
6806
8,05
0058
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065,
0580
70,0
5605
6;
1619
9,53
,199
8,03
8038
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074,
0580
62,0
6006
5,05
8058
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058;
16
200,
53,1
998,
0380
38,0
6807
4,05
8062
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065,
0580
62,0
5605
8;
1620
1,53
,199
8,03
8038
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068,
0580
66,0
6006
5,05
8070
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058;
16
203,
53,1
998,
0380
38,0
7407
4,05
4062
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060,
0580
70,0
4605
6;
1620
4,53
,199
8,03
8038
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068,
0500
66,0
6006
5,05
8058
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056;
16
205,
53,1
998,
0380
38,0
6806
8,05
4066
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065,
0700
70,0
5605
8;
1620
6,53
,199
8,03
8038
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068,
0620
66,0
6006
0,05
8070
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058;
16
208,
53,1
998,
0380
38,0
6806
8,05
4066
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065,
0580
70,0
5605
6;
1620
9,53
,199
8,03
8038
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068,
0540
66,0
6506
5,05
8062
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056;
16
210,
53,1
998,
0380
38,0
7407
4,06
2066
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060,
0580
62,0
5605
8;
1621
1,53
,199
8,03
8038
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074,
0620
66,0
6006
0,05
8070
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058;
16
212,
53,1
998,
0380
38,0
6806
8,05
8066
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065,
0580
70,0
5605
8;
1621
3,53
,199
8,03
8038
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068,
0540
62,0
6006
5,05
8070
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058;
16
214,
53,1
998,
0000
00,0
6806
8,00
0000
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065,
0700
70,0
0000
0;
1621
5,53
,199
8,03
8038
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074,
0000
00,0
6006
5,07
0070
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056;
16
216,
53,1
998,
0380
38,0
6806
8,05
0054
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065,
0580
70,0
5605
6;
1621
7,53
,199
8,03
8038
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068,
0540
58,0
6006
5,07
0070
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058;
16
218,
53,1
998,
0380
38,0
6806
8,05
4058
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065,
0620
70,0
4605
8;
1621
9,53
,199
8,03
8038
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074,
0580
62,0
6506
5,05
8070
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056;
16
220,
53,1
998,
0380
38,0
6806
8,05
4062
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060,
0580
62,0
5605
8;
1622
1,53
,199
8,03
8038
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068,
0540
62,0
6506
5,05
8058
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056;
16
222,
53,1
998,
0380
38,0
6806
8,05
0054
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065,
0580
70,0
0000
0;
1622
3,53
,199
8,03
8038
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068,
0500
62,0
6006
5,05
8062
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056;
16
224,
53,1
998,
0380
38,0
6807
4,06
2066
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060,
0620
62,0
5605
8;
1622
6,53
,199
8,03
8038
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071,
0540
58,0
6506
5,05
8070
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058;
16
227,
53,1
998,
0380
38,0
6806
8,05
4066
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065,
0580
58,0
5605
6;
1622
9,53
,199
8,03
8038
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068,
0540
62,0
6006
5,07
0070
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058;
16
266,
53,1
998,
0380
50,0
6806
8,05
8066
,065
065,
0580
70,0
4605
8;
1627
0,53
,199
8,03
8038
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068,
0540
66,0
6006
5,05
8070
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058;
16
284,
53,1
998,
0380
38,0
6806
8,05
8058
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060,
0500
58,0
5605
8;
1628
5,53
,199
8,03
8038
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068,
0500
66,0
6506
5,05
8062
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056;
16
286,
53,1
998,
0380
50,0
5005
0,05
4058
,060
065,
0580
62,0
5605
8;
1628
7,53
,199
8,00
0000
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068,
0620
62,0
5005
0,05
0050
,000
000;
16
288,
53,1
998,
0380
38,0
6806
8,05
4058
,065
065,
0580
70,0
4604
6;
1628
9,53
,199
8,03
8038
,068
068,
0580
62,0
6006
0,05
8070
,056
058;
16
290,
53,1
998,
0380
46,0
6806
8,04
6054
,065
065,
0700
70,0
5605
8;
1629
1,53
,199
8,03
8038
,068
068,
0540
54,0
6506
5,05
8062
,046
056;
16
292,
53,1
998,
0380
38,0
6807
4,05
0058
,060
065,
0580
70,0
5805
8;
1630
8,53
,199
8,03
8038
,068
068,
0580
66,0
6006
5,05
8062
,056
056;
16
310,
53,1
998,
0380
38,0
6807
4,05
4066
,060
060,
0620
70,0
4605
8;
1631
2,53
,199
8,03
8038
,068
074,
0540
62,0
6006
5,05
8058
,046
058;
16
313,
53,1
998,
0380
38,0
6806
8,05
4066
,060
060,
0700
70,0
5805
8;
1631
4,53
,199
8,03
8038
,068
068,
0500
54,0
6006
5,05
8062
,056
056;
16
318,
53,1
998,
0380
38,0
6806
8,05
4062
,065
065,
0580
70,0
5605
8;
1632
8,53
,199
8,03
8038
,068
068,
0540
62,0
6506
5,06
2070
,056
058;
16
329,
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;
616 N. RAUFASTE ET AL.
© 2005 The Linnean Society of London, Biological Journal of the Linnean Society, 2005, 84, 593–616
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