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SاFl+w>?>3NXfztghDI*%K%.Hn|Saɑ349 xx-pݯ^܍ٿK;)ɱ󟊂^G 贵 '0+"*JjˢoC5|%E4lEo/B#/wb;u$"*'9izBLLo ;5x!.I0̈́@)3{ͺ /1@LClgpV&0Uq!Dill:pc{-.*M#q#U^Y AKۥ,G``u:ٵ  O,lz Qv!ƚ '%0@`v& p"lB`ك,x EJO!cb͝2W93;Fڳh{xIbR0Cye{~_ !+G~bVXtјhtd=#^g]BCX6$vˎ`mIAG]{5KRJ\q8_,/ltM%0EȅvR*%0olL ++~IFJН^RT1v$6釀=ZepR @{e?%0a XƇv xmaFF0>4䡀ff?WV 6f=EID.n (nEP q ݡ >Dތ[4y5;M bE# < L\!GpA>do'/ Ul][!.+3od4›f%lJK [d${0aKF$k\(Co06$ETҍKw$- zf A%0hA:/c"u$wWq_s'w$Mևi~.bpw!M_LWmYBSua4YAG2pcCř96;q*y@x)¾zP@OReAG[䁤 Y^(DFL#TDSystemnicode MS0z[ 0F.  @n?" dd@  @@`` **H S:'$5 EG ^ ?h *dK)}l cxT EFCG 3+ :  =  YRA . ?XE @'By /S :x H 7'47HH %L ' (1Az bl G? < Z o: !0 -O=u5 ByBCDFEFG;J H- <  x 'O^_tr?!!|4}J x H0 < W -;8 ;  -0OD:9L ( *3A{cmG%K=   z<=b l;kR)!!**..   ,,HH,,HH>>  HG   //() HGHGHG,, -5=uA 8F  rZye+$b$6avfd +ir=$b$}|_1VV*&:`t/QW=r$JJDT&+>LBi'$$$b$֘̔~9S)ؓOK$$2$j'p|V3Z$$$$$$$$$2$ K} ͷ9SRGy"$rgMn-_M[|L2$Rc=Wϱ'0AS2$J^_"Z^ޗf $"$^i?٨y $2$HtkŝRǤRg2$S5Qm AKu2$C [ߍ[O2$ Jef{Zz+$"$(S5y5 x!$$$$$2$S]A96p+]0"2$Sh\LM3F #$$b$d^O6~% 0e0e     A@  A5% 8c8c     ?1 d0u0@Ty2 NP'p<'pA)BCD|E||s " 0e@        @ABC DEEFGHIJK5%LMNOPQRSTUWYZ[ \]^_ `abN E5%  N E5%  N F   5%    !"?N@ABC DEFFGHIJK5%LMNOPQRSTUWYZ[ \]^_ `ab 3@89m:ng48d8d z[ 0ppp@  <4!d!dL< 0<42d2dL< 0g4IdId z[ 0p/ pVuN ʚ;x:ʚ;<4ddddL|- 08<___PPT10 2___PPT9x/ 0\A? %O =$8LGenealogies of time structured data, an application on cave bear ancient DNA(ML3(L Frantz DepaulisOutline of the presentationnIntroduction: Gene genealogies Results .1 Simulation exploratory results .2 Cave bear application Conclusions 1 2;2 3; Wright Fisher Neutral model(  Assumptions Selective neutrality (Ne s <<1) Demography - Isolated panmictic Population, - Constant size N - Poisson Distribution of offspring P (1) - Same sampling time Mutational, sequence data: infinite site model (ISM) - No recombination - Independent mutations - Constant mutation rate Along the sequence Across time - Each mutation affects a new nucleotide sitej %.$~%7$%  H&JY@#  "   C Genealogy of a gene sample     F Coalescent   Constructing coalescents, &     )Constructing-deconstructing coalescents  *(&    GcM  rO2Frequency spectrum of mutations & neutrality tests33 3  IA KMitochondria, correlation LD/distance recombination or mutational effects?V'8(%8'% L d"Time structured data & genealogies##( #  ]J(Algorithm for time structured coalescent)) ) D>)Age structure effect on gene genealogies **( *  OB1Effect of subset size on statistical tests : mean"2- 2  PC?Effect of subset size on statistical tests : significance rate $@- @  QD7Effect of a half subset age on statistical tests: mean "82 8  XFDEffect of a half subset age on statistical tests: significance rates"E2 E  E@%Cave bear: Ursus spelaeus (12-300kYA)0& ((  4  v  v F?Sampling sites   ;94Alignment of polymorphic sites: D-loop of cave bear>5 c$c$c$ 5  REF TTGTCAACTT TCGAATTGAA GT #NOASC3500_40-45 ..A....T.C ..A....... .. #NOASC3800_40-45 ..A....T.C ..A....... .. #NOASC85F16_40-45 .......... .......... .. #NOASC95456_40-45 ..A....T.C ..A....... .. #NOASC92386_40-45 ..A....T.C ..A....... .. #NOASC92413_40-45 C.A....T.C ..A....... .. #NOASC92152_40-45 C.A....T.C ..A....... A. #NOASC5300_50-60 ..A....T.C ..A....... .. #NOASC11600_80 .......... .......... .. #NOASC12500_80 .......... .......... .. #NOASC13800_80 .......... .......... .. #NOASC100801_80 .......... .......... .. #NOASC12400_80 ..A....T.C ..A....... .. #NOASC11800_80 .CA....T.C ..A.G..... .. #NOASC11700_80 C.A....T.C ..A....... A. #NOASC84E16_90-130 C.A....T.C ..A....... .. #NOASC84G19_90-130 C.A....T.C ..A....... .. #NOASCbrC5-02_90-130 C.A....T.C ..A....... .. #NOASC15400_90-130 C.A....T.C ..A......G .. #NOASC15700_90-130 ....T.G.C. .TA..C..G. .. #NOATAB2_40 .......... .......... .. #NOAGrotteMerve_? .......... .T........ .. #NOAAZE_80-130 .......... .......... .C #NOAGigny189F3_? ..A....T.C ..A....... .. #NOAJAL104_? C.A....T.C ..A....... .. #NOATAB15_25-35 ..A......C ..A....... .. #NOAGailenreuth_? ..A......C ..A....... .. #NOA47910_30 ..A....T.C ..A....A.. .. #NOAHohleFels_? ..A....T.C ..A..C.... .. #NOACLA_35 ..A....T.C C.A....... .. #NOACLB_35 ..A....T.C C.A....... .. #NOAChiemsee_35 ..A..G.... ..A...C... .. #NOARamesch1_? ..A..G.... ..A...C... .. #NOARamesch2_? ..A..G.... ..A...C... .. #NOAGeissenklt1_? ...CT..... .T.G.C.... .. #NOAGeissenklt2_? ...CT..... .T.G.C.... .. #NOANixloch_? ...CT..... .T...C.... .. --------------------------------------------- Alp barrier #SOAPoto_? ...CT..... .T...C.... .. #SOAVind1_? ...CT..... .T...C.... .. #SOAVind2_? ...CT..... .T...C.... .. #SOAConturi_? .......T.. .......... .. Fc gc     N   d   d     $ @;Neutrality tests, Belgium cave   A<!Neutrality tests, dated subsample""&     C=Neutrality tests, total sample   \ILD as a function of distance   Time structure , Conclusion   $Acknowledgements  CNRS Nick Barton  /4k $$3/G0J1_2d5}6~789:;<=>?@AP*j* ` +*[[I߾Cqn̙f` R>&tcT̙߯fy4` )Rf{[u333f` c4'aZEEd` ff3f^[^V` X|לk5f` 3f3f3f>?" dd@,?uFd@  d uF@ d`uF n?" dd@   @@``PT $   @ ` `9p>>K0 %%,t K%(  t T  '  t  "' > t   B CDEFy@o5%      v S 6; Z5 `5 f) l x o ] E  zbJD>>82,&    , ,,,2288>>DDJJPPVVVPPPyPgJaJ[JUJUJOPOPIVIVC\C\=b7b+h%hhhVVVVVVVV\\bnzx`NN3N9N?NENKNQNWN]NiTZ``ffff```ZT<$ uW}E_35KpR4c-thVD882,,,& )/555555~5r5f5`5O;I;GMSSYYYYSS}SwYqYkYkYe_e_eeek_k_qYqSwMw<}600    @" ' . t   B CDE F(y @o5%     @" t   BCDEF y @o5%  @" t   B0CDEFy @o5%&&** **0 n`G|dDeM9 0 *$$$$****MP@"< l t   B:CDExFy @o5% *066n6R+(lxcoG(:.("=@@"6 p t   BCoDEtF|y @o5% u:+6 oNo\+;oO    ;<@"\qoV  t   BCcDEFy @o5%-- $<nSVwDDDDDDDDDD JP1V7V7\=\C\C#2crch+xy'_ [\@"}  t   hBCDE\Fdy @o5%<wY ;6F.p4,`/^hDzZ<<</0@"`k  t   HBC,DELFTy @o5% dN@F~.:& ,,b^< '(@"v  t   8BCDEDFLy @o5%2\xR<F#$@"F3n  t   0BCDE@FHy @o5%6qSP>:d6$d6666!$@"Zf t   (BCDE<FDy @o5%&x^>Z& @" R t   pBCDE`Fhy @o5%# 6dX.14@"b \H > t   BYCfDE(F0y @o5% GB0HfBfX8YGGG@" z=  t   pBCDE`Fhy @o5%  S Y`O214@"j'  t HS ?" `} S _'Cliquez pour modifier le style du titre((  t BhS ?#" `] `} S f* 0    t B( S ?#" ``  S `*"0     t BS ?#" `] `} S `*"0    X t <S ?" ` S vCliquez pour modifier les styles du texte du masque Deuxime niveau Troisime niveau Quatrime niveau Cinquime niveau4w b t <zd޽h ?"` ff3f^[^V___PPT10i. 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GLB @ c $D<LB  c $D  L to # toTB B c $D:t6Z"  s *-oLB  c $D w  H# ?  q -Coalescence-4  3 33  B  s *޽h ? ̙33f^___PPT10>+D"' = @B D' = @BA?%,( < +O%,( < +D' =%(D' =%(D' =4@BBBB%(D' =1:Bhidden*o3>+B#style.visibility<*%(D' =4@BBBB%(D' =1:Bhidden*o3>+B#style.visibility<*%(D' =4@BBBB%(D' =1:Bhidden*o3>+B#style.visibility<*%(D' =4@BBBB%(D' =1:Bhidden*o3>+B#style.visibility<*%(D' =4@BBBB%(D' =1:Bhidden*o3>+B#style.visibility<*%(D' =4@BBBB%(D' =1:Bhidden*o3>+B#style.visibility<*%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*%(+' m4K0 &&`>BL&(  ~  s *=t P}   j"  BG0f1?  d2  <f1?, $ l d2  <f1?. j o d2  <f1?. k n d2  <f1?) i j   T@gֳgֳ?p 4 W iaf  hh88   TJgֳgֳ?p 2 W ibf  hh88   TSgֳgֳ?k - R icf  hh88   T8egֳgֳ?k u R idf  hh88   TDggֳgֳ?k  R ief  hh88   Tqgֳgֳ?p YW iff  hh88d2  <f1?- W k d2  <f1?- j pB  Hf1?bF   N{gֳgֳ?x  0Most recent common ancestor of the sample (MRCA)11 1 hh88(  Ngֳgֳ?x l+ Dsample of  genes / of individuals## # hh88  NPgֳgֳ?  {Common ancestor (CA)  hh88^  6? ^  6?l m w ^  6?  Xn^  6?( l 8 ^  6?S d |d  <? B &   Tgֳgֳ? P  xneutral mutations   hh88vB @ Nf1? c:   ZЩgֳgֳ?  iTf  hh88  Zܫgֳgֳ?a r H  iCf  hh88  Zgֳgֳ?   iCf  hh88   Zlgֳgֳ?  iGf  hh88 ! Z4gֳgֳ? l  iCf  hh88 " Zgֳgֳ?" s  iGf  hh88 # ZPgֳgֳ?Mh ; 4 iAf  hh88 $ Zgֳgֳ? X  iAf  hh88vB % Nf1?E E vB & Nf1? vB ' Nf1? vB ( Nf1?^ ^ vB ) Nf1?K : l K vB * Nf1?vvB + Nf1?Q gS vB , Nf1?U pB - Hf1?  pB . Hf1?   pB / Hf1?   pB 0 Hf1? y pB 1 Hf1? | | pB 2 Hf1? l m pB 3 Hf1? pB 4 Hf1? pB 5@ Hf1? pB 6 Hf1? pB 7 Hf1?  i pB 8 Hf1?N N  pB 9 Hf1?z { pB : Hf1?z pB ;@ Hf1?   pB < Hf1?J pB = Hf1? pB > Hf1?f  B H ?  q -Coalescence-4  3 33  H  0޽h ? ff|ffy___PPT10Y+D=' = @B +< mK0  ++@BE8*(  8~ 8 s *t ]D !   pB 8 Hf1?pB 8 Hf1?pB 8 Hf1?TpB 8 Hf1?yypB 8 Hf1?pB 8 Hf1?TTpB  8 Hf1?y78pB  8 Hf1?pB  8 Hf1?xpB  8 Hf1?pB  8 Hf1?xLLpB 8 Hf1?yL8ypB 8@ Hf1?pB 8 Hf1?pB 8 Hf1?^ 8 6?{l%^ 8 6?s J^ 8 6?D^ 8 6?>P L    8#   r" 8 BGf1? v f2 8 61? ; f2 8 61? ?; f2 8 6ff1? ; f2 8 61?j ; f2 8 61? A; f2 8 61?> _;  8 T(gֳgֳ?    iaf  hh88 8 T1gֳgֳ?i -  ibf  hh88  8 T;gֳgֳ?   icf  hh88 !8 TDgֳgֳ?c '  idf  hh88 "8 T$Ugֳgֳ?.   ief  hh88 #8 TH_gֳgֳ?   iff  hh88     $8#   ,$@ 0 %8 Tagֳgֳ?  #  e1)  hh88 &8 Tjgֳgֳ?   vAges of the nodes  hh88  jd '8# j d,$@ 0~B (8 Nf?  ~B )8 Nf1?  *8 T}gֳgֳ? j d |t3,ff  hh88< #  +8# G ,$D 0r" ,8 BG1?M   -8 TPgֳgֳ?  p=1/2N:  hh88 .8 Tgֳgֳ?#   Exp( p ).   hh88*   /8#  ,$@ 0 08 TLgֳgֳ?   |t1,ff  hh88~B 18 Nf1? ~B 28 Nf1? ~B 38 Nf?   0  48# 0 ,$@ 0~B 58 Nf1?  68 T0gֳgֳ?0   |t2,ff  hh88~B 78 Nf?   f 88# f,$@ 0~B 98 Nf1?f :8 TDgֳgֳ?   |t4,ff  hh88~B ;8 Nf?   ] {B <8# h8,$@ 0~B =8 Nf1?] {B| >8 Tgֳgֳ?` >  t5:<fff  hh88~B ?8 Nf?b c C @8 0(T ,$  0 additional assumption: n << N L    0xx,, D8 H ?"6@ NNN?N ,$  0 p = (n (n -1)/2) /2N ( uF2B&     E8 HH ?  q -Coalescence-4  3 33  H 8 0޽h ? ff|ff,$___PPT10.+ Dl' = @B D'' = @BA?%,( < +O%,( < +D| ' =%(D$ ' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*$8%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*/8%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*48%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*'8%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*88%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*<8%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*+8%(D' =%(D' =%(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*D8%(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*@8%(+p+0+@8 ++0+D8 +v| mK0 hP`P`x@O(  @{   S  @# S  ,$D  0 @ Ngֳgֳ?N zneutral mutations F  hh88xB @B Hf1?g @ TDgֳgֳ?| iG  hh88 @ T0gֳgֳ?ia@H iTf  hh88 @ T<gֳgֳ?  iC  hh88 @ Tgֳgֳ?{8Q  iCf  hh88  @ T$gֳgֳ?lSI: iG  hh88  @ T((gֳgֳ?t iC  hh88  @ T/gֳgֳ? 1  iA  hh88xB  @ H1?6xB  @ H1?44xB @ H1?6:6xB @ Hf1? xB @ H1?YxxB @ H1?4xB @ Hf1?7; @ NBgֳgֳ?] c,$ 0 %3) uniform distribution of mutations&& & hh88 @ NEgֳgֳ?w4  q gene sample    hh88  @# ,$D 0xB @ Hf1?xB @ Hf1?xB @B Hf1? f @# ,$D 0xB @ Hf1?xB @ Hf1?iixB @ Hf1?f S @# S,$@ 0xB @ Hf1?SxB @ Hf1?SSxB  @ Hf1? !@ TOgֳgֳ? ~ xTopology of the tree  hh88 "@ TZgֳgֳ? > e2)  hh88^ #@ 6? $ ^ $@ 6?M   ^ %@ 6?# ' ^ &@ 6?# ) ^ '@ 6?K t ^ (@ 6? &{ ^ )@ 6? q ^ *@ 6? 4 ^ +@ 6?# @ ^ ,@ 6? # ^     -@#  ,$D 0 .@ T hgֳgֳ? ,   iCf  hh88xB /@B Hf1? J  0@ Tqgֳgֳ?   iA  hh88 1@ Tl{gֳgֳ?   iC  hh88 2@ Tgֳgֳ?1 l  iG  hh88 3@ T@gֳgֳ?K' ! iC  hh88 4@ T<gֳgֳ?   iG  hh88f 5@ 6?3K   6@ T\gֳgֳ? k  iTf  hh88 7@ Tgֳgֳ?O &  iTf  hh88E 8@ Nȶgֳgֳ? ,$ 0 -neutral distribution of sequence polymorphism.. . hh88B 9@ Hf1? ++ ,$@  0B :@ Hf1?  ,$@  0B ;@ Hf1? HH ,$@  0B <@ Hf1?  ,$@  0B =@ Hf1? xx ,$@ 0B >@ Hf1? << ,$D 0^ ?@ 6?6    @@#   ,$D  0 A@ Tgֳgֳ?   iA  hh88 B@ TLgֳgֳ?f S  iA  hh88 C@ Tdgֳgֳ?   iA  hh88^ D@ 6?> jZ E@# l,$D 0 F@ Tgֳgֳ?lZ iA  hh88xB G@ H1?j??pB H@@ Hf1?y z j" I@ BGf1? v ^2 J@ 61? ; ^2 K@ 61? >; ^2 L@ 6ff1? ; ^2 M@ 61? j; ^2 N@ 61? A; ^2 O@ 61? =_;  P@ Tgֳgֳ?   iaf  hh88 Q@ Tgֳgֳ? i-  ibf  hh88 R@ Tgֳgֳ?   icf  hh88 S@ TX gֳgֳ? c'  idf  hh88 T@ T0gֳgֳ? -  ief  hh88 U@ Tgֳgֳ?   iff  hh88 L  ~@# L ,$D 0T 5x_ W@# Lx8xB X@ Hf1?^y_xB Y@ Hf1?5x5xB Z@ Hf1?5y_y [@ T%gֳgֳ?O  pMRCAF  hh88~B \@B Nf1?%f`O  ]@# y,$D 0xB ^@ Hf1?xB _@ Hf1?xB `@ Hf1? )O} a@# O)},$D 0 b@ N3gֳgֳ?O zcommon ancestor (CA)P  hh88xB c@B Hf1?)2}vB d@ Nf? vB e@ Nf1? vB f@ Nf1? vB g@ Nf1?Z vB h@ Nf1?{S 8| i@ TDgֳgֳ?  |t1,ff  hh88 j@ T|?gֳgֳ?/   |t2,ff  hh88 k@ T+B#style.visibility<*@%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*a@%(D4' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*]@%(D4' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*@%(D4' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*@%(D4' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*~@%(D' =%(Du' =%(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*@%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*@%(D ' =%(D ' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*9@%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*:@%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*;@%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*<@%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*=@%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*>@%(D4' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*E@%(D' =%(D' =%(DA' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<*@@%(D' =-}6B!slide(fromLeft)*<3<*@@D4' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*-@%(D' =%(Du' =%(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*u@%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*v@%(D' =%(Du' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*w@%(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*8@%(++0+@ ++0+8@ ++0+u@ +mK0  5 23! 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H ,$@d 02 b  01? Y u H ,$@e 02 c  61?  H ,$@f 02 d  61? V q H ,$@g 0B e  <1? _ _ ,$@h 0B f  <1? ,$@i 0B g  <1? X X ,$@j 0B h  <1? ,$@k 0B i  <1? v v ,$@l 0B j  <1? ,$@m 0:2 k  3 CUENGlfHZJ~Q1? `TU]TU`TU]TU~]TU~ / J ,$@ 0B l  <1?OO O,$@n 0B m  <1? ,$@o 0B n  <1?M {,$@p 0) o  Zgֳgֳ?M < ,$q 0 kT0   hh88) p  Z gֳgֳ?9 0 ,$r 0 kC0   hh88) q  ZPgֳgֳ?`,$ 0 kT0   hh88) r  Z!gֳgֳ?% & ,$ 0 kA0   hh88) s  Z*gֳgֳ?R ,$ 0 kC0   hh88) t  Z4gֳgֳ?<o) ,$ 0 kC0   hh88) u  ZDEgֳgֳ?`,$ 0 kG0   hh88) v  ZPGgֳgֳ?g,$ 0 kC0   hh88) w  ZQgֳgֳ?X,$ 0 kG0   hh88) x  ZZgֳgֳ?%  ,$  0 kC0   hh88) y  Zdgֳgֳ?j (! ,$  0 kT0   hh88) z  ZDugֳgֳ? ! ,$  0 kG0   hh88) {  ZPwgֳgֳ?q T( ,$  0 kG0   hh88) |  Zgֳgֳ?q =( ,$  0 kG0   hh88) }  Zgֳgֳ?% > ,$ 0 kC0   hh88) ~  Zgֳgֳ?% K ,$ 0 kC0   hh88)   ZDgֳgֳ?  U ,$ 0 kC0   hh88)   ZPgֳgֳ?q  ( ,$ 0 kG0   hh88)   Zgֳgֳ?h,$ 0 kA0   hh88)   Zgֳgֳ?4m% ,$ 0 kA0   hh88B   <1?,$@ 0B   <1?6,$@ 0B   <1?I,$@ 0B   <1?,$@ 0B   <1?6UU,$@ 0B   <1?9W,$@ 0"   BG1? % E ,$@ 0B   <1?JJ,$@ 0B  @ <1?WX,$@ 0B  @ <1?,$@ 0B   <1?7,$@ 0B   <1?7U8,$@ 0B   <1?C,$@  0B   <1?E,$@! 0B   <1?,$@" 0B  @ <1?Y,$@# 0B   <1?,$@$ 0^B   6?tu uB   <1?!,$@% 0B   <f1?_~,$@& 0B   <1?,$@' 02   6PP1? ,$@( 02   61?>X ,$@) 02   61? ,$@* 02   6PP1?E` ,$@+ 02   01?  ,$@, 02   61?B] ,$@- 0B   <1? LL ,$@. 0B   <1?  ,$@/ 0B   <1? CC ,$@0 0B   <1?  ,$@1 0B   <1? bb ,$@2 0B   <1?  ,$@3 0F2   S BTCUENGfHL"ZI3JQ1? ӫTUTUӫTUTU3TU3  ) ,$@ 0B   <1?EE,$@4 0B   <1?,$@5 0@2   C CUENGZHI`TJQ1? "U`TU"U`TU`T`TU`T -* ,$@ 0)   Z0gֳgֳ?  ,$6 0 kA0   hh88)   Zgֳgֳ?% ` ,$7 0 kA0   hh88)   Zgֳgֳ?j `! ,$8 0 kT0   hh88   Z4gֳgֳ?H  Haplotype tests: simulations 0(  hh88   Zgֳgֳ?m/g,$ 0 .parameters! : S =8 n =6h0 J  hh889   Zgֳgֳ?E ?,$ 0 {K = 6(0  hh889   Zgֳgֳ?E ?,$ 0 {K = 5(0  hh889   Z@'gֳgֳ?J D*D,$ 0 {K = 4(0  hh88.   Z0*gֳgֳ?PW,$ 0 p10 0000  hh883   ZP4gֳgֳ? ,$ 0 u simulations 0   hh88^   ZT>gֳgֳ? .d,$ 0 haplotype number K {>0  hh88:   TKgֳgֳ?Ge,$ 0 haplotype diversity  0P  hh88T   Z@Qgֳgֳ?,}L,$ 0 H = 1- S@ 0  hh88)   Z0dgֳgֳ?>08,$ 0 kf0  hh88)   Z,$ 0 k20  hh88<   Z|gֳgֳ?MxG,$ 0 ~H = 0.83( 0  hh88<   Zgֳgֳ?M O G,$ 0 ~H = 0.78( 0  hh88<   Zdgֳgֳ?M-G,$ 0 ~H = 0.72( 0  hh88B   <1?  ,$Ds 0B   <1?d~,$@9 0)   Z gֳgֳ?c > ,$: 0 kC0   hh88)   Zԡgֳgֳ?f Q ,$; 0 kC0   hh88)   Zgֳgֳ?%  ,$< 0 kA0   hh88)   Zgֳgֳ?j ! ,$= 0 kT0   hh88+   Z8gֳgֳ? S,$ 0 m{0  hh88  @ 0cC ,$@ 0E   Zgֳgֳ?  Depaulis and Veuille MBE 199860&     hh88T   Z\gֳgֳ?D ,$ 0 ! Hudson 1993.0J   hh887   Zgֳgֳ?xM,$ 0 y ... &0  hh88N   C B(C DEF1?'' @"<   t ,$@ 0    BCDEF? @  ,$@ 0N   C B(CDEF1?'' @"<  T Sk ,$@ 0N   C B(CfDEF1?''ee @"<    ,$@ 0N   C B(CDEF1?'' @"<    ,$@ 0N   C B(C!DEF1?''   @"<  3  ,$@ 0N   C B(CDEF1?'' @"<    ,$@ 0N   C B(CDEF1?'' @"<  m 2 ,$@ 0N   C B'CDEF1?&& @"<   ;S ,$@ 0N   C B(CDEF1?'' @"<   \u ,$@ 0N   C B'CDEF1?&& @"<   ~ ,$@ 0N   C B)CDEF1?(( @"<    ,$@ 0N   C B)CDEF1?(( @"<    ,$@ 0N   C B(C0DEF1?''// @"<  S  ,$@ 0N   C B(CDEF1?'' @"<    ,$@ 0N   C B(CODEF1?''NN @"<   $< ,$@ 0N   C B(CDEF1?'' @"<  ` E] ,$@ 0N   C B'CDEF1?&& @"<   g ,$@ 0N   C B'CvDEF1?&&uu @"<    ,$@ 0N   C B'CGDEF1?&&FF @"<    ,$@ 0N   C B(C(DEF1?'''' @"<    ,$@ 0N   C B'C DEF1?&& @"<    ,$@ 0N   C B(CDEF1?'' @"<  4 % ,$@ 0N   C B(CDEF1?'' @"<  H .F ,$@ 0N   C B(CDEF1?'' @"<  [ Og ,$@ 0N   C B(CDEF1?'' @"<  n o ,$@ 0N   C B(CDEF1?'' @"<  {  ,$@ 0N   C B(CDEF1?'' @"<    ,$@ 0N   C B(CuDEF1?''tt @"<    ,$@ 0N   C B(CfDEF1?''ee @"<     ,$@ 0N   C B)C^DEF1?((]] @"<   0 ,$@ 0N   C B)CNDEF1?((MM @"<   7Q ,$@ 0N   C B(C?DEF1?''>> @"<   Yq ,$@ 0`   c B3CDEF1?22 @"B ` { 0O ,$@ 0B   <1? } ,$@ 0  Q   #  Ql ,$@ 0lB   <1?= lB   <1? = > lB   <1?   lB   <1? c d lB   <1?   lB   <1?   lB   <1? # # lB   <1? lB   <1? HIlB   <1? lB   <1?* Q B   <1?  ,$@ 0B   <1? +,,$@ 0B   <1? LM,$@ 0B   <1? mn,$@ 0B   <1? ,$@ 0B   <1? ,$@ 0B   <1? ,$@ 0B   <1? ,$@ 0B   <1? ,$@ 0B   <1? 56,$@ 0B ! <1? VW,$@ 0B ! <1? xx,$@ 0B ! <1? ,$@ 0B ! <1? ,$@ 0B ! <1? ,$@ 0B ! <1? ,$@ 0B ! <1? ,$@ 0B ! <1? ??,$@ 0B ! <1? `a,$@ 0B  ! <1? ,$@ 0B  ! <1? ,$@ 0B  ! <1? ,$@ 0B  ! <1? ,$@ 0B  ! <1? ,$@ 0B ! <1? '(,$@ 0B ! <1? HI,$@ 0B ! <1? ij,$@ 0B ! <1? ,$@ 0B ! <1? ,$@ 0B ! <1? ,$@ 0B ! <1? ,$@ 0B ! <1? ,$@ 0B ! <1? 12,$@ 0B ! <1? ST,$@ 0B ! <1? tu,$@  0I ! T gֳgֳ?C"? ,$  0 q0.2 0   hh88/ ! ZLgֳgֳ? c,$  0 q0.3 0   hh88/ ! Z gֳgֳ? ,$  0 q0.4 0   hh88/ ! Z)gֳgֳ? ,$  0 q0.5 0   hh88/ ! Z3gֳgֳ? <,$ 0 q0.6 0   hh88/ ! Zl=gֳgֳ? ,$ 0 q0.7 0   hh88/ ! Z4Ggֳgֳ? D!,$ 0 q0.8 0   hh88/  ! ZPgֳgֳ? ,$ 0 q0.9 0   hh88- !! ZZgֳgֳ?6nl0,$ 0 oH 0  hh88- "! Tdgֳgֳ?  ,$ 0 udensity 0  hh88    #!# R,$@ 0lB $! <1?"   %! C B.C0DEF1? /- @ 3  &! Zhygֳgֳ?zN,$ 0 observed H : P = 0.03 *x0 g  hh88X '! Tgֳgֳ?N ,$! 0 Distribution of simulated H60  hh88N (!@ C B(CfDEF1?''ee @"<   ( ,$@ 0 )!@  B)C^DEFA1?((]] @"<    ,$@ 0 *!@  B)CNDEFA1?((MM @"<    ,$@ 0 +!@  B(C?DEFA1?''>> @"<    ,$@ 0 ,!@  B.C$DEFA1?--## @"<    ,$@ 0B -! <1? ,$@ 0B .! <1? ,$@ 0B /! <1? ,$@ 0B 0! <1? ,$@ 0B 1! <1?  ,$@  0I 2! T@gֳgֳ?C"? Z,$ 0 q0.1 0   hh88 3! H̒ ?  q -Coalescence-4  3 33  H   0޽h ?o`8 k      Ca(___PPT10.hn+a>D' = @B Dd' = @BA?%,( < +O%,( < +D^' =%(De^' =%(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*q %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*r %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*s %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*t %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*u %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*v %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*w %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*x %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*y %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*z %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*{ %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*| %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*} %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*~ %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<* %(DU' =%(DT' =%(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*; %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*< %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*= %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*> %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*? %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*@ %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*A %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*B %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*C %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*D %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*E %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*F %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*G %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*H %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*I %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*J %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*K %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*L %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*M %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*N %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*O %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*P %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*Q %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*R %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*S %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*T %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*U %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*V %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*W %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*X %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*Y %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*Z %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*[ %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*\ %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*] %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*^ %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*_ %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*` %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*a %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*b %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*c %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*d %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*e %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*f %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*g %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*h %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*i %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*j %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*l %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*m %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*n %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*o %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*p %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(DU' =%(DT' =%(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*! %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*" %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*# %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*$ %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*% %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*& %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*' %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*( %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*) %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<** %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*+ %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*, %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*- %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*. %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*/ %(D' =4@BBBB%(D' 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 ++0+F  ++0+G  ++0+H  ++0+I  ++0+J  ++0+o  ++0+p  ++0+q  ++0+r  ++0+s  ++0+t  ++0+u  ++0+v  ++0+w  ++0+x  ++0+y  ++0+z  ++0+{  ++0+|  ++0+}  ++0+~  ++0+  ++0+  ++0+  ++0+  ++0+  ++0+  ++0+  ++0+  ++0+  ++0+  ++0+  ++0+  ++0+  ++0+  ++0+  ++0+  ++0+  ++0+  ++0+  ++0+  ++0+  ++0+  ++0+  ++0+  ++0+  ++0+  ++0+  ++0+  ++0+  ++0+! ++0+! ++0+! ++0+! ++0+! ++0+! ++0+! ++0+ ! ++0+!! ++0+"! ++0+&! ++0+'! ++0+2! +e m5K0 ,,.4d ,(   R  ZH`gֳgֳ? g. ,$ 0  GCGCGCGAACCCATT outgroup*0   hh88{  Zrgֳgֳ? ET,$ 0  121531416121423 frequenciesR03 3  hh88  6xv < 8Alignment of polymorphic sites: frequencies of mutations"9U $ 9 / L E   # -{ M   ZH~gֳgֳ?E = GCCCGCGAATCCATT^033  hh88m   Zgֳgֳ?E  GCGTGCGATCCGATT~0333  hh88   Zgֳgֳ?E~  GCGTACAATCCCGTC033333  hh88   Zgֳgֳ?EB  -GTGTACAATCTCGAC0333333  hh88   Z|gֳgֳ?E M  -GTGTACAATCTCGAC0333333  hh88   ZPgֳgֳ?E   GCGTGGAATCCCGTT03333  hh88?   Zgֳgֳ?E  CCGCGCGGTCCCATTP033  hh88XB = 0DjJBM ^B > 6DjJOB  ? BlM5<  Tn =7    @ B0h US =15   z H A  Kl,$@ 03 B  Zgֳgֳ? C TP003  hh88 C   `Zgֳgֳ?H !40g$3$  hh88/ D Zgֳgֳ?r,$ 0 qT"03  hh88+ E Zgֳgֳ?<:4,$ 0 mC0  hh88B F NfjJ?~ Z~ ,$@ 0B G NfjJ?^ ^,$@ 0B H @ NfjJ?[} ,$@ 0B I NZfjJ?" #,$@  0B J NfjJ?T < T ,$@  0B K @ NZfjJ? X ,$@  0B L NZfjJ?< < ,$@ 0B M NZfjJ? 9 ,$@ 0B N NZfjJ? ; ,$@ 0B O NZfjJ?F G ,$@ 0B P NZfjJ?ff,$@ 0B Q NZfjJ?fK ,$@ 0B R NfjJ? R,$@  0B S NZfjJ?x Nx ,$@ 0B T NZfjJ?OSx ,$@  0B U NZfjJ? ' ,$@ 0B V NZfjJ?!! ,$@ 0B W NfjJ? ,$@ 0B X N3o?m,$@ 0B Y NZfjJ?+l+,$@ 0B Z NZfjJ?,,,$@ 0B [ NZfjJ? \ ,$@  0B \ NZfjJ?  ,$@ 0B ] @ NZfjJ?+ ,$@ 0 ^ BZjJH2,$@ 0l _ 6 "` 0Cl ` 6 "` :+ a Z"gֳgֳ?g qk ,$! 0 mC0  hh88B b 0D3o9bb$,$D 0B c 6D3>>`,$D" 0 d H, ?  q -Coalescence-4  3 33  H  0޽h ? ff|ff8w8___PPT10W8.Q i+c3D7' = @B D6' = @BA?%,( < +O%,( < +DA' =%(D' =%(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<* %(DA' =%(D' =%(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<* %(D4' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*b %(D7/' =%(D.' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*M %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*A %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*L %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*N %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*J %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*K %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*I %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*R %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*T %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*F %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*S %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*G %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*H %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*P %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*X %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*D %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*Q %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*O %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*U %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*V %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*W %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*Y %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*E %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*^ %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*Z %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*] %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*\ %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*[ %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*a %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*c %(++0+  ++0+  ++0+D  ++0+E  ++0+a  +` m6K0 .&* (   `   c $A"  ??6b     c $A  ??6b8  $ 0  B_ t {V     6_% 3 &fi : number of occurrences in a sample8' $ %   68f  eNumber of polymorphic sites  B  s *Do Z^( ,$@ 0B  s *DoJN2 ,$@ 0B  s *Dou 9 = ,$@ 0B  6ZDo%Y),$@ 0B  6ZDo N,$@ 0B  6ZDo  D ,$@  0B  0D1 ] ,$@  0B  0D1 r  ,$@  0B  0D1H,$D  02  <jJ# ,$@ 0B  6DjJ ^ c ,$@ 0  6+B#style.visibility<* %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D4' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(Dq' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<* %(D ' =%(D ' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bhidden*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bhidden*o3>+B#style.visibility<* %(DH' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bhidden*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bhidden*o3>+B#style.visibility<* %(DA' =%(D' =%(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =%(Du' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*& %(D' =%(Dh' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*) %(Dq' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*% %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*$ %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*# %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*( %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*' %(DA' =%(D' =%(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*! %(+x+(+  0++(+  0++0+  ++0+  ++0+!  ++0+#  ++0+&  + m1K0, Introduction - $(      6P t "6 ?N  J6 @`P Ss77B^  6^  6\\p   c &A   ??"?f     Tgֳgֳ?p p Q ~ distance d& 0   hh88  6S  2r 2 = !(d ) Pearson s statistic tested by permutations of sites A 3 A   <\ Awadalla et al. (Science 1999)U   sH  0޽h ? 3̙33& TIMING |17.3y___PPT10Y+D=' = @B +H  mK0 G ? P (  ~  s *Pt [   pB  HfjJ?SS pB  HfjJ?22 pB @ HfjJ?6SL f # /@ xB  HfjJ?xB  HfjJ?iixB   HfjJ?fpB   HfjJ?  pB   HfjJ? 6 pB   HfjJ?  pB   HfjJ?1 3 I pB  HfjJ?/pB  HfjJ?3 pB  HfjJ? K pB  HfjJ? w pB  HfjJ?  h  BD    - Parasites during disease evolution (virus& ) - Microbial experimental evolution - Ancient DNA_p_<  B   ` Issue: To what extent the analyses are affected by time structure? How to correct for this? N pqXp- X  H0 ?  q -Coalescence-4  3 33  H  0޽h ? ff|ff___PPT10i.V0 +D=' = @B + m2K0 660.HY `6(   L * Z,gֳgֳ?ec _ ,$ 0 n =2<fff  hh88L 6 Z &gֳgֳ?k + e ,$# 0 n =5<fff  hh88~  s *t0t q5   B  Hf1?. ,$@( 0B  Hf1?- ,$@) 0B  Hf1? T ,$D 0B  @ Hf1?n~~,$@2 0B  Hf1?-  ,$@ 0B  @ Hf1? TT ,$@ 0B  @ Hf1?n23,$D4 0B  @ Hf1? k,$@9 0B  Hf1?n}n,$@3 0B  Hf1?q,$@8 0B  Hf1?jLL,$@? 0B  Hf1?kH4k,$@@ 0B  @ Hf1? ,$@' 0B  Hf1?,$@: 0B  Hf1?  ,$@ 0^  6?ml ^  6?e <^  6? D ^  6?>2  <1? M ,$@ 0d2  <1? + d2  <ff1?  2  <1? nM ,$@ 02  <1? !CM ,$@ 0d2  <1? De   ZpEgֳgֳ? ~  iaf  hh88  Z8Ogֳgֳ? g+  ibf  hh88  ZYgֳgֳ?   icf  hh88' ! Zbgֳgֳ?! c' ,$ 0 idf  hh88' " Zlgֳgֳ?! . ,$ 0 ief  hh88' # ZXvgֳgֳ?!  ,$ 0 iff  hh88B ) Tf1?V ,$@% 0L 0 Zgֳgֳ? ,$  0 n =3<fff  hh88B 1 Tf1? #  ,$@  0B 2 Tf1?  ,$@  0B 3 Tf? ,$@  0B 5 Tf1?s y,$@/ 0B 7 Tf?'  ,$@ 0B 9 Tf1?f,$@6 0L : Z<gֳgֳ?B ,$5 0 n =3<fff  hh88B ; Tf? n,$@7 0B = Tf1?wh8x,$@> 0: > Zgֳgֳ?Y,$< 0 |n =2*ff  hh88B ? Tf?no,$@= 0 B Tgֳgֳ?   }t 1,ff  hh88vB C Nf1?. L2 vB D Nf1? Z vB E Nf?0  B F Nf1?0  5 ,$@ 0B G Nf?/ / * ,$@$ 0B H Nf?ue e ,$@. 0F I T,gֳgֳ?zg t,$- 0 n =4<fff  hh88B J Nf1?# #,$D 0l (   M ! v ,$D 0B K B BDo?"0@NNN?N(  B L  BDo?"0@NNN?N(  B N Hf1?  ,$@  0B O Hf1?4 ,$@& 0B P Hf1?-  ,$@ 0B Q Hf1?n,$@1 0B R Hf1?~~" ,$@* 0B S Hf1?23: ,$@+ 0B T @ Hf1?23q,$D; 0B U @ Hf1?i23,$DA 0B V Hf1?n,$@0 0g W H ?"6@ NNN?N; 0,$,  0 }#The exponential law is memoryless !$$ i X T<gֳgֳ? p ,$ 0 n1 =3^fffff  hh88 Y H ?  r- Simulations-4  333  H  0޽h ? ff|ffss___PPT10s.+9pD$p' = @B Do' = @BA?%,( < +O%,( < +D; ' =%(D ' =%(D' =4@BBBB%(D' =1:Bhidden*o3>+B#style.visibility<* %(D' =A@BBBB0B%(D' =1:Bhidden*o3>+B#style.visibility<*X %(D' =4@BBBB%(D' =1:Bhidden*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bhidden*o3>+B#style.visibility<* %(D' =A@BBBB0B%(D' =1:Bhidden*o3>+B#style.visibility<*" %(D' =A@BBBB0B%(D' =1:Bhidden*o3>+B#style.visibility<*# %(D' =A@BBBB0B%(D' =1:Bhidden*o3>+B#style.visibility<*! %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*0 %(D| ' =%(D$ ' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*2 %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*3 %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*1 %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*N %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(DY' =%(D' =%(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<** %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*7 %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*J %(D4' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*M %(D' =%(D1' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*P %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*X %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*F %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*! %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*" %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*# %(D' =4@BBBB%(D' =1:Bhidden*o3>+B#style.visibility<*M %(D' =A@BBBB0B%(D' =1:Bhidden*o3>+B#style.visibility<** %(D' =4@BBBB%(D' =1:Bhidden*o3>+B#style.visibility<*J %(D' =4@BBBB%(D' =1:Bhidden*o3>+B#style.visibility<*7 %(Dn' =%(D' =%(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*6 %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*G %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*) %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*O %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*R %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*S %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*W %(D' =+4 8?RCBBCB#ppt_wB*Y3>B ppt_w<*W D' =+4 8?RCBBCB#ppt_hB*Y3>B ppt_h<*W D' =+K4 8?CBB#ppt_x+(cos(-2*pi*(1-$))*-#ppt_x-sin(-2*pi*(1-$))*(1-#ppt_y))*(1-$)CB?B*Y3>B ppt_x<*W D' =+K4 8?CBB#ppt_y+(sin(-2*pi*(1-$))*-#ppt_x+cos(-2*pi*(1-$))*(1-#ppt_y))*(1-$)CB?B*Y3>B ppt_y<*W D ' =%(D ' =%(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*I %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*H %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*5 %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*V %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*Q %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D ' =%(D1 ' =%(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*: %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*9 %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*; %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*T %(D ' =%(D ' =%(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*> %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*? %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*= %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*U %(++0+* / ++0+* / ++0+6 / ++0+! / ++0+! / ++0+" / ++0+" / ++0+# / ++0+# / ++0+0 / ++0+: / ++0+> / ++0+I / ++0+W / ++0+X / ++0+X / +# m7K0 ""P-<@  "(   r  S i/t  `}  /   B ?"6@ NNN?NL hFp  # $lB   <jJ?  plB  <jJ?lB   <jJ?~ rB   BjJ?hrB   BjJ?prB   BjJ?uprB   BjJ?FrB   BjJ? FGrB  BjJ?#prB   BjJ?prB   BjJ? rB  BjJ? K .8   ?  rB   BjJ? D rB  BjJ? C rB   BjJ? B CxB   HjJ? xB  B HjJ?  xB   HjJ?xB   HjJ? ! xB !  HjJ? ! xB " HjJ?  xB #  HjJ?  xB $  HjJ?  xB % HjJ?  G 8  F 8 w9rB  B BjJ? FrB  BjJ? FrB   BjJ?  xB  HjJ?= CxB   HjJ?< CxB   HjJ?< = xB  HjJ?{ |= rB &  BjJ?8rB ' BjJ? 7 rB (  BjJ? 67xB )  HjJ? E xB *  HjJ?E E xB +  HjJ?  xB ,  HjJ?xB -  HjJ? xB . HjJ?orxB /  HjJ?qjxB 0 HjJ?  ;xB 1  HjJ?kn xB 2  HjJ?| Y xB 3 B HjJ?U W / 5 Hx/ ?"6@ NNN?N  yContemporaneous sample *  6 H,~/ ?"6@ NNN?NA   `Limited time structure  ; 7 H/ ?"6@ NNN?NlT  #Two subsets with large time spacing*$## 1 : HX/ ?"6@ NNN?N A {'Excess of rare variants Deficit of LD( (' A ; H/ ?"6@ NNN?Nb 7Deficit of rare variants Excess of LD Differentiation 8 87  < Tt/gֳgֳ?  }t 1,ff  hh88|B = Tfo?;5 > T̥/gֳgֳ? n1 =4^fffff  hh88 @ H/ ?  r- Simulations-4  333  H  0޽h ? ff3f^[^V___PPT10i.9[A+D=' = @B +^ m8K0  -7 @(   r + <X99 ?"`Ms l |4   ' 4 | ,$D  0 ?  BC8DEDFL5% 0P!o(/5>871-&M]m} #$@L m @ |4   % |4   @ |4  $ |4 BB J 3 4 L BB K  3 L c BB L B 3 |L M BB M  3 L M BB O 3 i  BB P  3   BB Q B 3   BB R  3   BB T 3 M N BB U  3 M N BB V B 3 6 M BB W  3 M e BB Y 3   BB Z  3   BB [ B 3   BB \  3   BB ^ 3 e f BB _  3 e f# BB ` B 3 N e BB a  3 e | BB c 3  4 BB d  3 4 L BB e B 3 4 5 BB f  3 4 15 BB h 3 1F 2] BB i  3 1] 2t BB j B 3 ] 1^ BB k  3 1] H^ BB m 3  i  BB n  3   BB o B 3   BB p  3   BB r 3  z BB s  3  BB t B 3  BB u  3  BB w 3  t BB x  3  BB y B 3  BB z  3  BB | 3 rQ si BB }  3 ri s BB ~ B 3 [i rj BB   3 ri j BB  3 > ?) 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H2   C h = H2   C ph  H2   C 5h W H2   C h  H2   C h  H2   C m  H2   C em  H2   C = m _ H2   C $ r E H2   C  w " H2   C  |  H2   C  | H2   C  H2   C   H2   C h  H2   C J k H2   C +w M H2   C h * H2   C P r    xB&C&DEF̙&&& @` B    xB'C&DEF̙'&& @`p     xB'C&DEF̙'&& @`5 \    xB'C&DEF̙'&& @`     xB'C&DEF̙'&& @`     xB&C&DEF̙&&& @`     xB'C&DEF̙'&& @`e     xB'C&DEF̙'&& @`= d    xB&C&DEF̙&&& @`$ J    xB'C&DEF̙'&& @` '    xB&C&DEF̙&&& @`    xB&C&DEF̙&&& @`    xB'C&DEF̙'&& @`    xB&C&DEF̙&&& @`     xB'C&DEF̙'&& @`h     xB&C&DEF̙&&& @`J p    xB'C&DEF̙'&& @`+ R    xB'C&DEF̙'&& @` /    xB&C&DEF̙&&& @`  H   C  B H   C p  H   C 5 \ H   C   H   C   H   C   H   C e  H   C = d H   C $ J H   C  ' H   C  H   C  H   C  H   C   H   C h  H   C J p H   C + R H   C  / H   C   N2   S  8 N2   S u N2   S :R N2   S  N2   S  N2   S  N2 !  S j N2 "  S B Z N2 #  S ( @ N2 $  S   N2 %  S   N2 &  S   N2 '  S   N2 (  S N2 )  S mYqN2 *  S OgN2 +  S 0wHN2 ,  S  %N2 -  S  .  BCDEF @` =  /  BCDEF @`u  0  BCDEF @`:W  1  BCDEF @`  2  BCDEF @`  3  BCDEF @` 4  BCDEF @`j 5  BCDEF @`B _  6  BCDEF @`( E  7  BCDEF @` "  8  BCDEF @`   9  BCDEF @` z  :  BCDEF @` T q ;  BCDEF @`)E <  BCDEF @`m =  BCDEF @`Ok >  BCDEF @`0wM ?  BCDEF @` !*> @  BCDEF @`  A  67}   d-12 G    B  6,7?   f-0.52 G    C  67   c02 G    D  6P(7` /  e0.52 G    E  6(7D  c12 G    F  6P?7`T e1.52 G    G  6T7d c22 G    H  6P^7`y e2.52 G    I  6\7  c32 G    J  6|r7   g0.0012 G    K  6x77   f0.012 G    L  67  i  e0.12 G    M  6Б7E   c12 G    N  67   d102 G    O  67_   gt1 4 G   P  6X7 ( hin 24 G   Q  607[ - tNe4 G   R  6 ( p generations4   G   S  60O# hMean4 G  BB T  3 7< U  # BB V 3 BB W  3 BB X B 3 BB Y  3  Z  60Od rDt2 G   BB [  3 ̙< \  # 5jBB ] 3 ̙:MBB ^  3 ̙M`BB _ B 3 ̙:MBB ` 3 ̙M` a  60d| fD*fl2 G   BB b  3 @< c  #  BB d 3 BB e  3 BB f B 3 BB g 3 BB h 3 BB i  3  j  60Xd& sHfw2 G   BB k  3 MH l  C  m  6`0 dn  cK2 G   BB n  3  2  o  B'C&DEF'& @`   p  60J d  cH2 G   BB q  3  x H2 r  C  5  s  6)0 dm  sZnS2 G   BB t  3 ̙ (  u  xB&C&DEF̙&&& @`   v  6@0@ d iPearson2 G   <B w  # H x  C *P y  6M0dY eFst2 G   BB z  3 N2 {  S  |  6U05d k Pi/Theta02   G   BB }  3  ~  BCDEF @`Xu   6e0d fS/S02 G   H  0޽h ? ff3f^[^V**___PPT10*.?$\+-1D)' = @B D)' = @BA?%,( < +O%,( < +D(' =%(%(Do(' =%(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*= %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*K %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*L %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*M %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*N %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*O %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*P %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*Q %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*R %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*S %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*T %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*U %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*V %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*W %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*X %(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*Y %(Dc' =4@BBBB%(E5' =1B B`BPB,54*3>B ppt_c='`B@BPB<* D' =?B70, 0; .2, .5; .8, .5; 1, 0-g6B fade*<3<* D' =0l9 BBBB*<3<* )?Dp' =A@BBBB0B%(E5' =1B B`BPB,54*3>B ppt_c='`B@BPB<*" D' =?B70, 0; .2, .5; .8, .5; 1, 0-g6B fade*<3<*" D' =0l9 BBBB*<3<*" )?Dc' =4@BBBB%(E5' =1B B`BPB,54*3>B ppt_c='`B@BPB<*# D' =?B70, 0; .2, .5; .8, .5; 1, 0-g6B fade*<3<*# D' =0l9 BBBB*<3<*# )?Dp' =A@BBBB0B%(E5' =1B B`BPB,54*3>B ppt_c='`B@BPB<*< D' =?B70, 0; .2, .5; .8, .5; 1, 0-g6B fade*<3<*< D' =0l9 BBBB*<3<*< )?++0+ 6 ++0+" 6 ++0+< 6 +q m;K0 p p- o(   x  c $4<t M  < N  H|< ?"6@ NNN?N  Empty symbols: deficit of the statistics; Filled symbols: excess of the statistics. Dt: Tajima's (1989) D; D*fl Fu and Li (1993)'s D*; Hfl Fay and Wu's (2000) H; ZnS Kelly (1997)'s ZnS; K and H Depaulis and Veuille (1998)'s haplotype tests; Pearson: Pearson correlation coefficient between pairwise allelic correlation and distance between mutation tested by permutations according to Awaddala et al. (1999); Fst Hudson et al's (1992) Fst tested by permutations hT  1        L  W  D , F  F   ~8lB  B <jJ? FlB  <jJ? FlB   <jJ?  rB  BjJ?= CrB   BjJ?< CrB   BjJ?< = rB  BjJ?{ |= lB   <jJ?8lB  <jJ? 7 lB   <jJ? 67rB   BjJ? E rB   BjJ?E E rB   BjJ?  rB   BjJ?rB   BjJ? rB  BjJ?orrB   BjJ?qjrB  BjJ?  ;rB   BjJ?kn rB !  BjJ?| Y rB " B BjJ?U W  # T8<gֳgֳ?i` }t 1,ff  hh88|B $ Tfo?gh@" % N;< ?"6@ NNN?ND n1=n/2N    L hFp & #   lB '  <jJ?  plB ( <jJ?lB )  <jJ?~ rB *  BjJ?hrB +  BjJ?prB ,  BjJ?uprB -  BjJ?FrB .  BjJ? FGrB / BjJ?#prB 0  BjJ?prB 1  BjJ? rB 2 BjJ? K dB 3 <jJ?s s wdB 4 <jJ?e e hdB 5 <jJ?c s jB 6 BjJ?h- hjB 7 @ BjJ?g fjB 8 BjJ?f0 0 fjB 9 BjJ?LLEjB : BjJ? LjB ; BjJ?JjB < BjJ?GfjB = BjJ?GGjB > BjJ? jB ? BjJ?uF G jB @ BjJ?x jB A BjJ?wL x B HM< ?  r- Simulations-4  333  OT S3 D  "S3T C  c $X99?S3BB E  3  BB F  3   BB G  3 y z BB H  3 g h BB I  3 U V BB J  3 CDBB K  3 23BB L  3  !BB M  3 BB N  3   BB O 3   BB P 3   BB Q 3 % & BB R 3 S T BB S 3    T  BfC9DE$F, 7,1Y'  9f9@2 D BB U 3 :  V  B CDEF$̙- Z @u 5 BB W 3 ̙   X  BCcDE(F0 [,OY@0 9f0c@ S\  Y  B9CeDE F(e-PZ5 '9W@u m SR  Z  B9CiDE F(i-UY7 99g@M G +?  [  BeCDE$F, , Y 9_e@% " < \  #   : BB ] 3   BB ^  3   1 BB _ B 3    BB `  3    < a  # ] BB b 3 u v BB c  3 u v  BB d B 3 b u BB e  3 u < f  # 5 j BB g 3 M N BB h  3 M N BB i B 3 : M BB j  3 M ` < k  #  Y B BB l 3 % ] & q BB m  3 % q & BB n B 3  q % r BB o  3 % q 8 r < p  #  $  Y BB q 3  ) < BB r  3  < O BB s B 3  < = BB t  3  < = < u  #   O BB v 3   2 BB w  3  2 E BB x B 3  2 3 BB y  3  2 3 < z  # q  BB { 3 u  BB |  3   BB } B 3   BB ~  3   <   # h, a BB  3 1 D BB   3 D W BB  B 3 mD E BB   3 D E B   3 "WBB  3 3f':BB   3 3f:MBB  B 3 3f:;BB   3 3f:;<   # 5 j  BB  3 ̙: M BB   3 ̙M `  BB  B 3 ̙: M  BB  3 ̙M ` <   #  B BB  3 ̙ % BB   3 ̙% 8 BB  B 3 ̙ % BB  3 ̙% 8 <   #   BB  3 ̙ BB   3 ̙ BB  B 3 ̙ BB  3 ̙ <   #  BB  3 ̙ BB   3 ̙ BB  B 3 ̙ BB  3 ̙ <   #   BB  3 ̙  BB   3 ̙  BB  B 3 ̙  BB  3 ̙  <   # h R BB  3 ̙m" 5 BB   3 ̙5 I BB  B 3 ̙m5 I BB  3 ̙" 5 B   3 ̙ 5 BB  3 3f  BB   3 3f , BB  B 3 3f , BB  3 3f  H2   C  H2   C b H2   C : T [ u H2   C  3  H2   C   H2   C  m H2   C   H2   C mT u H2   C @I aj N   S : [ N   S  Y 3 z N   S   N   S  Z | N   S w  N   S m 7 N   S @ a    B&C&DEFff&& @` 8    B&C&DEFff&& @` < b    B&C'DEFff&' @`    B&C&DEFff&& @`4 Z    B&C&DEFff&& @`m|     B&C'DEFff&' @`@E fl  ¸  B&C'DEFff&' @`" >I H ø  C   H ĸ  C  H Ÿ  C   H Ƹ  C m  H Ǹ  C @h fm H ȸ  C >H ɸ  C H ʸ  C  ˸  64MBV   c02 G    ̸  6MD   f0.052 G    ͸  6H,M2   e0.12 G    θ  63M   f0.152 G    ϸ  60=M e0.22 G    и  6GMz f0.252 G    Ѹ  6TMh e0.32 G    Ҹ  6BMV f0.352 G    Ӹ  6,\MF aY  g0.0012 G    Ը  60jM |Y  f0.012 G    ո  61 Y  e0.12 G    ָ  61; Y  c12 G    ׸  61Q Y  d102 G    ظ  6d1 ?g  gt1 4 G   ٸ  61Y   hin 24 G   ڸ  61^ F  tNe4 G   ۸  621)Y   p generations4   G   ܸ  6@1Cr p Significance4   G   ݸ  6J1c i rate4 G  BB ޸  3 E< ߸  # BB  3 BB   3 BB  B 3 BB   3    6L1]+ vDt_inf2 G   BB   3 ^B   3 *BB  3 3f BB   3 3f BB  B 3 3f BB   3 3f     61v+ vDt_sup2 G   BB   3 ̙<   # CxBB  3 ̙H[BB   3 ̙[nBB  B 3 ̙H[BB  3 ̙[n   61( + D*fl_inf2  G &   BB   3 ̙b  B   3 ̙  BB  3 3f  BB   3 3f  BB  B 3 3f  BB  3 3f     6j1  + D*fl_sup2  G &   BB   3   H2   C # E    6 a1 + wZnS_inf2 G   BB   3 KN   S    6@]19+ uK_sup2 G   BB   3 x   B'C&DEFff'& @`   6p1.p+ uH_sup2 G   <B   # CH   C    6t[ + eFst2 G   H  0޽h ? ff3f^[^V___PPT10i.?$\+D=' = @B + m0K0 p- ^(   ~  s *m<t Gl  < T  C ,A,cavebear2 W R  C *A cavebear W   Hp< ?  r- Application-4  333  H  0޽h ? ff|ff___PPT10i.V0 +D=' = @B +y m<K0 xp`- (   r  S u<t >  < d  C <A$cvbreuropsampling-  H3/ ?  r- Application-4  333  H  0޽h ? ff3f^[^V___PPT10i.;Qu+D=' = @B +  m=K0 -| (  | r | S ́<t    < r | S <t E < : | N< ?"6@ NNN?Nr 6x  ~ n =41 S =22 B uF   | NĊ< ?"6@ NNN?Nr R Z(Loreille et al. 2001) (Orlando et al. 2002) (Hofreiter et al. 2002) (Khn et al. 2001) d[+                       | TGH؞IjJ?"0@NNN?N r|I | N< ?"6@ NNN?N 9y   Ne= 13 000 J     | H؅< ?Z r  r- Application-4  333  H | 0޽h ? |  ff3f^[^V___PPT10i.? +D=' = @B + ? m>K0  >> --4 0 =(   x  c $<t C `  < @  N< ?"6@ NNN?NyM a permutation test 8   H< ?  r- Application-4  333  R  s *X99?{  <TY-~. c Statistic*   B     <Y-~0 [ * B     <(H-~h [D* B     <0e-Zl [t* B    <-~r [ * B     <-~2  [D* B     <-|$ @  [** B    <(-< ^  \fl* B    <7-~b {  [ * B     <g-~ S  [H* B     <$Z,C   jfw* B    <Z,~  [ * B     <b,~H  [K* B     <t,~  [ * B     <~,~j [H* B     <,~ [ * B     <8,~ [Z* B     <8, jnS* B    <,~ [ * B     <`,~& aPearson* B     <,~' [ * B   @  C {Z~@B ! C o{Z|@ " C {Z\~@B # C o{Z\|@B $ C o{Z[~@ % C {\~@B & C o{\|@ ' C {~@B ( C o{|@B ) C o{~@ * C { ~@B + C o{ |@ , C {  ~@B - C o{  |@B . C o{  ~@ / C { Q ~@B 0 C o{ Q |@ 1 C {Q S ~@B 2 C o{Q S |@B 3 C o{Q R ~@ 4 C {S ^~@B 5 C o{S ^|@ 6 C {^`~@B 7 C o{^`|@B 8 C o{^_~@ 9 C {`~@B : C o{`|@ ; C {~@B < C o{|@B = C o{~@ > C {~@B ? C o{|@ @ C {~@B A C o{|@B B C o{~@ C C {~@B D C o{| E <6,g^ bObserved*  B    F <>,g [ * B    G <0@,g [-* B    H <@N,g  ^0.82* B    I <T,g [ * B    J <T6,g [-* B    K <L,g  ^1.55* B    L <,gx  [ * B    M <,g   [-* B    N <<,g  ^1.32* B    O <,g  [ * B    P <,gU  [7* B    Q <=g~  [ * B    R <t!gB ^0.79* B    S <THg [ * B    T <xRgs ^0.24* B    U <DZg [ * B    V <^g [-* B    W <\g7 m 0.39 (2.8*)2   F    X <}b [a* B   Y <g [ * B   @ Z C dg@B [ C dg@ \ C dg@B ] C de@B ^ C dgF _ S d@g@B ` C d@e@ a C d@Bg@B b C d@Be@B c C d@AgF d S dBZg@B e C dBZe@ f C dZ\g@B g C dZ\e@B h C dZ[gF i S d\g@B j C d\e@ k C dg@B l C de@B m C dgF n S d g@B o C d e@ p C d  g@B q C d  e@B r C d  gF s S d Q g@B t C d Q e@ u C dQ S g@B v C dQ S e@B w C dQ R gF x S dS ^g@B y C dS ^eF z S d^`g@B { C d^`e@B | C d^_gF } S d`g@B ~ C d`e@  C dg@B  C de@B  C dgF  S dg@B  C de@  C dg@B  C de@B  C dgF  S dg@B  C de  <HE [(* B     <= \P * B     <hEB bvalue %)*  B     <`E*B [ * B     <T> `(21.0)* B     <> [ * B     <>  _(5.3)* B     <>w  [ * B     <>  `(18.4)* B     <>  [ * B     <>  `(16.4)* B     <$ >  [ * B     <@ >! `(37.7)* B     <>  [ * B     <'>R6 `(43.7)* B     <)>$< [ * B     <x6>s h(2.8*)2 F     <?>ay [ * B   @  C ;@5=@B  C o;@5<@  C ;58=@B  C o;58<@B  C o;56=@  C ;8Z=@B  C o;8Z<@  C ;Z\=@B  C o;Z\<@B  C o;Z[=@  C ;\=@B  C o;\<@  C ;=@B  C o;<@B  C o;=@  C ; =@B  C o; <@  C ;  =@B  C o;  <@B  C o;  =@  C ; Q =@B  C o; Q <@  C ;Q S =@B  C o;Q S <@B  C o;Q R =@  C ;S ^=@B  C o;S ^<@  C ;^`=@B  C o;^`<@B  C o;^_=@  C ;`=@B  C o;`<@  C ;=@B  C o;<@B  C o;=@  C ;=@B  C o;<@  C ;=@B  C o;<@B „ C o;=@ Ä C ;=@B Ą C o;< ń <a# ^Mean* B    Ƅ <6 bNo time *  B     <4 c structure*   B     <x4 [ * B     <D46> d % rejected*   B     <p4-> [ * B     <ؼ4> c (4.9;5.5)*   B     <4> [ * B     <4 > c (5.2;2.8)*   B     <@ > [ * B     <( > c (5.4;4.8)*   B     < > [ * B     < > c (1.7;3.9)*   B     <H > [ * B     <$6> c (4.9;4.6)*   B     <5> [ * B     <h3'c> c (5.5;5.1)*   B     <P4Kc> [ * B     <lL> a(5.0;/)* B     <pQo> [ * B     <a  [(* B     <n  [P* B     <tD  c value %)*   B     <w)A  [ * B     <l  `(30.0)* B     <  [ * B     <4   _(8.8)* B     <0w   [ * B     <   `(17.2)* B     <Ľ   [ * B     <    _(8.6)* B     <ѷ   [ * B     <T%  `(31.2)* B     <d  [ * B     <(U2  `(31.7)* B     < !9  [ * B     <t s  h(2.7*)2 F     < ay  [ * B   @  C @5@B  C o@5@  C 58@B  C o58@B  C o56@  C 8Z@B  C o8Z@  C Z\@B  C oZ\@B  C oZ[@  C \@B  C o\@ ! C @B " C o@B # C o@ $ C  @B % C o @ & C   @B ' C o  @B ( C o  @ ) C  Q @B * C o Q @ + C Q S @B , C oQ S @B - C oQ R @ . C S ^@B / C oS ^@ 0 C ^`@B 1 C o^`@B 2 C o^_@ 3 C `@B 4 C o`@ 5 C @B 6 C o@B 7 C o@ 8 C @B 9 C o@ : C @B ; C o@B < C o@ = C @B > C o ? <,-    ^Mean* B    @ <,    [ * B    A <l@    [-* B    B <H    ^0.30* B    C <(W    [ * B    D <\    [-* B    E <g    ^0.38* B    F <q  {  [ * B    G <l|  n  ]0.3* B    H <  c  [9* B    I <   [ * B    J <|  6  ]9.1* B    K <   [ * B    L <  B  ^0.80* B    M <    [ * B    N <  s  ^0.22* B    O <    [ * B    P <(  P  ^0.00* B    Q <  C[  [ * B    R <4 [ (  \CI* B    S < [   [ * B    T <[   [[* B    U < [   [-* B    V <` [ $  d 1.56;1.26]*   B    W <[ !  [ * B    X <<[ c  [[* B    Y <8[   [-* B    Z <\B[   d 1.89;0.84]*   B    [ <P[   [ * B    \ <[[   [[* B    ] <e[  [-* B    ^ <o[ '  d 4.04;2.56]*   B    _ <@p[ #  [ * B    ` <$[  `[6;12]* B    a <[  [ * B    b <8[ f  e [0.66;0.89]*   B    c <L[ F^  [ * B    d <4[   e [0.08;0.47]*   B    e <ܰ[ w  [ * B    f <h[ 6Y  [[* B    g <[ Vv  [-* B    h <[ b  d 0.29;0.23]*   B    i <[   [ * B    j <L   bAverage *  B    k <Pu   _time * B    l <LI   c structure*   B    m <I   [ * B    n </ 6  d % rejected*   B    o <\/ -  [ * B    p <+/   c (7.8;3.0)*   B    q <2/   [ * B    r <@/   c (8.2;1.0)*   B    s <8/  [ * B    t <M/  `(4.2;3* B    u <`/ x  [.* B    v <f/  \7)* B    w <k/  [ * B    x <v/   c (0.8;9.5)*   B    y <̉/   [ * B    z <ؐ/ 6  c (3.3;7.8)*   B    { <#/ 5  [ * B    | <4/ v  d (11.5;2.9)*   B    } <p/ [s  [ * B    ~ <д/   a(4.9;/)* B     <L/ o  [ * B     <   [(* B     < y  [P* B     < D  c value %)*   B     < )A  [ * B     <X y  `(30.0)* B     < y  [ * B     <   y  _(8.6)* B     < w y  [ * B     < ' y  _(17.4* B     < y  [)* B     <$9 y  [ * B     <,=  y  _(7.9)* B     <P y  [ * B     <P !y  `(30.9)* B     <a  y  [ * B     <$f U2y  `(31.9)* B     <{ !9y  [ * B     <l{ sy  h(2.8*)2 F     < ayy  [ * B   F  S  @5 @B  C  @5 @  C  58 @B  C  58 @B  C  56 F  S  8Z @B  C  8Z @  C  Z\ @B  C  Z\ @B  C  Z[ F  S  \ @B  C  \ @  C   @B  C   @B  C   F  S    @B  C    @  C     @B  C     @B  C     F  S   Q  @B  C   Q  @  C  Q S  @B  C  Q S  @B  C  Q R  F  S  S ^ @B  C  S ^ F  S  ^` @B  C  ^` @B  C  ^_ F  S  ` @B  C  ` @  C   @B  C   @B  C   F  S   @B  C   @  C   @B  C   @B  C   F  S   @B  C     < ^  ^Mean* B     <x ^  [ * B     <\ ^  [-* B     < ^  ^0.33* B    … < ^  [ * B    Å <D ^  [-* B    ą <  ^  ^0.42* B    Ņ < { ^  [ * B    ƅ <\ ^  ^0.37* B    Dž < ^  [ * B    ȅ <  6 ^  ]9.1* B    Ʌ < ^  [ * B    ʅ < $ B^  ^0.80* B    ˅ <. ^  [ * B    ̅ <d9 s^  ^0.22* B    ͅ <A ^  [ * B    ΅ <M P^  ^0.00* B    υ <tN C[^  [ * B    Ѕ <XY (2 \CI* B    х <$d 2 [ * B    ҅ <r 2 [[* B    Ӆ < 2 [-* B    ԅ < 2 d 1.59;1.18]*   B    Յ <c 2 [ * B    օ <  c2 [[* B    ׅ < 2 [-* B    ؅ <  2 d 1.89;0.84]*   B    م <̰  2 [ * B    څ <  2 [[* B    ۅ <lE 2 [-* B    ܅ < ' 2 d 4.20;2.54]*   B    ݅ < # 2 [ * B    ޅ < 2 `[6;12]* B    ߅ <D 2 [ * B     <, f2 e [0.66;0.89]*   B     < F^2 [ * B     < 2 e [0.08;0.48]*   B     <@) w2 [ * B     </ 6Y2 [[* B     <= Vv2 [-* B     <C b2 d 0.29;0.24]*   B     <tP 2 [ * B     <0Yg1 pScladina*  B     <(gg0 [ * B     <\LDy [n* B     <,sTo  ]=20* B     <\L [ * B     <2%U [S* B     <:J ]=15* B     < 2 [ * B     < Mey  [ * B     < c[M  f Uncertainty *   B     <0   bin time *  B     < d structure *   B     <$ [ * B     <6 d % rejected*   B     <- [ * B     < c (9.3;2.8)*   B     <+ [ * B     <+  c (9.3;0.8)*   B     <+  [ * B     <"+  c (4.5;3.6)*   B     <)+  [ * B     <D+  c (0.7;9.8)*   B     <@+  [ * B     <K+6 c (3.7;7.5)*   B     <U+5 [ * B     <^+v d (11.6;2.8)*   B     <(c+[s [ * B     <|v+{ a(4.8;/)* B     <}+) [ * B   @  C g@i@B  C og@h@  C gCFi@B  C ogCFh@B  C ogCDi@  C gF5i@B  C ogF5h@  C g9;i@B  C og9;h@B  C og9:i@  C g;Zi@B  C og;Zh@  C g]`i@B  C og]`h@B  C og]^i@  C g`i@B  C og`h@  C gi@B  C ogh@B  C ogi@  C g i@B  C og h@  C g  i@B  C og  h@B  C og  i@  C g Q i@B  C og Q h@  C gU W i@B ! C ogU W h@B " C ogU V i@ # C gW ^i@B $ C ogW ^h@ % C gbdi@B & C ogbdh@B ' C ogbci@ ( C gdi@B ) C ogdh@ * C gi@B + C ogh@B , C ogi@ - C gi@B . C ogh@ / C gi@B 0 C ogh@B 1 C ogi@ 2 C gi@B 3 C ogh 4 <+jPh [ * B   H  0޽h ? ff3f^[^V___PPT10i.Aep+D=' = @B +;B m?K0 :A2A0-7< 0 @(      <0e0e ?t #" 0eK  < <  N< ?"6@ NNN?NyM a permutation test 4    H< ?  r- Application-4  333  R  s *X99?L  < =b c Statistic*   B     < 6Nb [ * B     <t  W b [D* B     < N ` W [t* B    < i b [ * B     << K b [D* B     Ho9 g B [** B    <4b^ W \fl* B    <y b [ * B     < # q b [H* B     <tk W jfw* B    <h b [ * B     <4yb [K* B     <xb [ * B     <]b [H* B     <[tb [ * B     <i1b [Z* B     <,gW jnS* B    <mb [ * B     <Rb aPearson* B     <8b [ * B   @  C +@B  C o+@  C +.@B ! C o+.@B " C o+,@ # C . @B $ C o. @ % C   @B & C o  @B ' C o  @ ( C  < @B ) C o < @ * C < ? @B + C o< ? @B , C o< = @ - C ? `@B . C o? `@ / C `c@B 0 C o`c@B 1 C o`a@ 2 C c@B 3 C oc@ 4 C @B 5 C o@B 6 C o@ 7 C @B 8 C o@ 9 C "@B : C o"@B ; C o @ < C " @B = C o" @ > C  @B ? C o @B @ C o  @ A C 0@B B C o0 C <dS bObserved*  B    D < ZrS [ * B    E <@oS [-* B    F <o S ^1.21* B    G <\ո S [ * B    H <ո  S [-* B    I <` S ^2.28* B    J < S [ * B    K <a & S [-* B    L <}, S ^0.69* B    M <t S [ * B    N <5,S \12* B    O <,~S [ * B    P <FS ^0.86* B    Q <ptS [ * B    R <S ^0.14* B    S <Hr|S [ * B    T <yIiS [-* B    U <yS e 0.27 (11.4)*   B    V < [ * B   W <` [a* B   X <Hɱ&>S [ * B   @ Y C @B Z C o@ [ C @B \ C o@B ] C o@ ^ C @B _ C o@ ` C @B a C o@B b C o@ c C )@B d C o)@ e C ),@B f C o),@B g C o)*@ h C +.@B i C o+.@B j C o+,@ k C . @B l C o. @ m C   @B n C o  @B o C o  @ p C  < @B q C o < @ r C < ? @B s C o< ? @B t C o< = @ u C ? `@B v C o? `@ w C `c@B x C o`c@B y C o`a@ z C c@B { C oc@ | C @B } C o@B ~ C o@  C @B  C o@  C "@B  C o"@B  C o @  C " @B  C o" @  C  @B  C o @B  C o  @  C 0@B  C o0  <h9 [(* B     <<O1 [P* B     <ث^9 c value %)*   B     <9 [ * B     < 1 `(10.5)* B     < 1 [ * B     <|& 1 i(0.6**)2 F     <@1 1 c 2 F     <$ 1 `(25.7)* B     <  1 [ * B     <|R1 `(16.5)* B     <O1 [ * B     <1 `(32.1)* B     <Ġ1 [ * B     <X1 `(24.3)* B     <H1 [ * B     <1 `(11.5)* B     <1 [ * B   F  S a@B  C a@  C ad@B  C ad@B  C abF  S d)@B  C d)F  S ),@B  C ),@B  C )*F  S , @B  C , @  C   @B  C   @B  C   F  S  $ @B  C  $ F  S $ ' @B  C $ ' @B  C $ % F  S ' < @B  C ' < @  C < ? @B  C < ? @B  C < = F  S ? `@B  C ? `@  C `c@B  C `c@B  C `aF  S c@B  C c@  C @B  C @B  C F  S  @B  C  @ ˆ C  #@B È C  #@B Ĉ C  !F ň S #@B ƈ C #@ Lj C "@B Ȉ C "@B Ɉ C  F ʈ S " @B ˈ C " F ̈ S  @B ͈ C  @B Έ C   F ψ S 0@B Ј C 0 ҈ <`֭pX ^Mean* B    ӈ <$խSk [ * B    Ԉ << [-* B    Ո <x  ^0.09* B    ֈ <7  [ * B    ׈ <   [-* B    ؈ <|  ^0.08* B    و <Z $ c $V"`=ad@@B % C V=ad>@B & C V=ab@Z ' c $V"`=d)@@B ( C V=d)>@ ) C =),@@B * C V=),>@B + C V=)*@Z , c $V"`=,$ @@B - C V=,$ >@ . C =$ ' @@B / C V=$ ' >@B 0 C V=$ % @Z 1 c $V"`=' < @@B 2 C V=' < >Z 3 c $V"`=< ? @@B 4 C V=< ? >@B 5 C V=< = @Z 6 c $V"`=? `@@B 7 C V=? `>@ 8 C =`c@@B 9 C V=`c>@B : C V=`a@Z ; c $V"`=c @@B < C V=c >Z = c $V"`= #@@B > C V= #>@B ? C V= !@Z @ c $V"`=#@@B A C V=#>@ B C ="@@B C C V=">@B D C V= @Z E c $V"`=" @@B F C V=" >@ G C = @@B H C V= >@B I C V=  @Z J c $V"`=0@@B K C V=0> L </ pX  ^Mean* B    M </ Sk  [ * B    N </   [-* B    O </  u  ]0.4* B    P <0/ t  [2* B    Q <x/  [ * B    R <>/   [-* B    S <X:/   ^0.59* B    T <XW/  [ * B    U <a/   ^0.35* B    V <pk/  [ * B    W < T}  d 5.34;2.98]*   B    l <P Jb}  [ * B    m <Z }  `[8;15]* B    n <[ }  [ * B    o <xe }}  e [0.71;0.91]*   B    p <p }  [ * B    q <Dz m }  e [0.07;0.39]*   B    r < }  [ * B    s <ԙ Ux}  [[* B    t <ܡ y}  [-* B    u < $}  d 0.23;0.20]*   B    v <( 2}  [ * B    w <䬈@ bAverage *  B    x < '`  htime structure* B    y <ӈ Rj  [ * B    z <8 X  d % rejected*   B    { < X  [ * B    | <,  X  c (9.3;2.1)*   B    } <A X  [ * B    ~ <t A X  c (6.9;0.3)*   B     <A  X  [ * B     <A X  c (4.7;2.6)*   B     <!A #X  [ * B     <2A X  d (1.2;11.1)*   B     <  [(* B     <A O6  [P* B     <ܓA ^>  c value %)*   B     <A >  [ * B     <A  6  `(18.5)* B     <ĹA 6  [ * B     <A 6  h(1.9*)2 F     <PA 6  c 2 F     <A 6  `(23.4)* B     <A 6  [ * B     <`A 6  `(39.9)* B     <DA 6  [ * B     < 6  `(43.2)* B     < 6  [ * B     < 6  `(41.1)* B     <% 6  [ * B     <. 6  `(11.9)* B     <+ 6  [ * B   F  S  a @B  C  a @  C  ad @B  C  ad @B  C  ab F  S  d) @B  C  d) F  S  ), @B  C  ), @B  C  )* F  S  ,$ @B  C  ,$ F  S  $ ' @B  C  $ ' @B  C  $ % F  S  ' < @B  C  ' < @  C  < ? @B  C  < ? @B  C  < = F  S  ? ` @B  C  ? ` @  C  `c @B  C  `c @B  C  `a F  S  c  @B  C  c  @  C  # @B  C  # @B  C  ! F  S  # @B  C  # @  C  " @B  C  " @B  C    F  S  "  @B  C  "  F ‰ S   @B É C   @B ĉ C    F ʼn S  0 @B Ɖ C  0  lj <R pX'  ^Mean* B    ȉ <Q Sk'  [ * B    ɉ <e '  [-* B    ʉ <(n  '  ^0.44* B    ˉ <| '  [ * B    ̉ < '  [-* B    ͉ <̎  '  ^0.61* B    Ή <R '  [ * B    ω <0  '  ^0.37* B    Љ <䣇 '  [ * B    щ < '  ^11.8* B    ҉ < '  [ * B    Ӊ <$̇ '  ^0.84* B    ԉ <և '  [ * B    Չ <D '  ^0.18* B    ։ <L |'  [ * B    ׉ < '  ^0.00* B    ؉ < '  [ * B    ى <  \CI* B    ډ <d * [ * B    ۉ <# d [[* B    ܉ < $  [-* B    ݉ <* $  d 1.70;1.09]*   B    މ <T6  3  [ * B    ߉ <0D g  [[* B     <T  [-* B     <c 6  d 2.28;0.72]*   B     <ti , D  [ * B     <x  [[* B     <  [-* B     < T d 5.23;2.99]*   B     <| Jb [ * B     <   `[8;16]* B     <ة  [ * B     < } e [0.71;0.91]*   B     <ཉ  [ * B     < m  e [0.07;0.40]*   B     <̉  [ * B     <։ X{ [[* B     < | [-* B     <   d 0.24;0.19]*   B     < . [ * B     <fS d all dated *   B     < 8. [n* B     < ? 6 _=27, * B     <H'3 [S* B     <0+7$ ]=20* B     <; [ * B     <@IXq  [ * B     <lO QI6  f Uncertainty *   B     <tJ   bin time *  B     <H_z   c structure*   B     <0`z   [ * B     <4~q d % rejected*   B     <샋q [ * B     <$q  c (9.3;2.4)*   B     <|q  [ * B     <q  c (7.0;0.2)*   B     <\q   [ * B     <Tq  c (4.6;2.7)*   B     <Ƌq # [ * B     <t̋q d (1.2;11.7)*   B     <X܋q ! [ * B     <q c (3.5;9.7)*   B     <q [ * B     <q d (14.1;2.5)*   B     <q [ * B     <q ](5.* B     <q! ^4;/)* B     <0#q [ * B   @  C LO@B  C oLM@  C LO@B  C oLM@B  C oLO@  C LaO@B  C oLaM@  C LehO@B  C oLehM@B  C oLefO@  C Lh)O@B  C oLh)M@  C L-0O@B  C oL-0M@B  C oL-.O@  C L0$ O@B  C oL0$ M@  C L) , O@B  C oL) , M@B  C oL) * O@ ! C L, < O@B " C oL, < M@ # C LA C O@B $ C oLA C M@B % C oLA B O@ & C LC `O@B ' C oLC `M@ ( C LehO@B ) C oLehM@B * C oLefO@ + C Lh O@B , C oLh M@ - C L%'O@B . C oL%'M@B / C oL%&O@ 0 C L'O@B 1 C oL'M@ 2 C L#&O@B 3 C oL#&M@B 4 C oL#$O@ 5 C L& O@B 6 C oL& M@ 7 C LO@B 8 C oLM@B 9 C oLO@ : C L0O@B ; C oL0M < <@O c 2 F   H  0޽h ? ff3f^[^V___PPT10i.Aep+D=' = @B +Vh m@K0 UgMg@-ot @f(   R  s *X99? x  c $<t [ `)  < <  N$< ?"6@ NNN?NyM a permutation test 4    H< ?  r- Application-4  333    <lg =6 c Statistic*   B     <xt ":6 [ * B     <q (r6 [D* B     <n ew+ [t* B    < |6 [ * B     <8 ( r 6 [D* B     <^ e  [** B    < ~ + \fl* B    <8`  6 [ * B     < 8 6 [H* B     < x + jfw* B    <  6 [ * B     <+ d 6 [K* B     <`^ w 6 [ * B     <p.6 [H* B     <'@6 [ * B     <p6 [Z* B     <]+ jnS* B    <j66 [ * B     <@GA6 aPearson* B     <tRk6 [ * B     <dZ6 [F* B     <d+ jst* B    <\Y76 [ * B   F  S ;@B ! C ;@ " C ;=@B # C ;=@B $ C ;<F % S =@@B & C =@F ' S @C@B ( C @C@B ) C @AF * S Cc @B + C Cc @ , C c e @B - C c e @B . C c d F / S e c @B 0 C e c F 1 S c f @B 2 C c f @B 3 C c d F 4 S f @B 5 C f @ 6 C  @B 7 C  @B 8 C  F 9 S  @B : C  @ ; C @B < C @B = C F > S @B ? C @ @ C @B A C @B B C F C S @B D C @ E C @B F C @B G C F H S @B I C  J <{& bObserved*  B    K <Ĕ& [ * B    L <& [-* B    M <& ^0.45* B    N <& [ * B    O <= 0 & [-* B    P <(D+ & ^0.88* B    Q < & [ * B    R <1 & ^1.35* B    S < & [ * B    T <lw l & \17* B    U <te } & [ * B    V <vW& ^0.91* B    W <PIa& [ * B    X <8& ^0.10* B    Y < +C& [ * B    Z <H& [-* B    [ <"& e 0.09 (22.0)*   B    \ <! 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BB   3 8c&BB   3 8c&c d BB  3 8c&c ' BB  3 8cc  BB  3 8cic j BB  3 8c c BB  3 8c c BB  3 8cGc H BB  3 8cc  BB  3 8cc  <   # DXBB  3 1L`BB   3 1BB  B 3 1LBB  3 1`<   # 3 X BB  3 1: `q BB   3 1q  BB  B 3 1: q BB  3 1q ` <   #  XQ BB  3 1 ` BB !  3 1 A BB " B 3 1  BB # 3 1 `A < $  #   OBB % 3 1  BB &  3 1  @BB ' B 3 1  @BB ( 3 1  < )  # tBB * 3 1|BB +  3 1BB , B 3 1|BB - 3 1< .  # Z#BB / 3 1BB 0  3 1KBB 1 B 3 1BB 2 3 1K< 3  # i#BB 4 3 1qBB 5  3 1BB 6 B 3 1qBB 7 3 1< 8  # 3  BB 9 3 1: q QBB :  3 1q Q BB ; B 3 1: Qq BB < 3 1q  Q< =  # F  BB > 3 1 N V BB ?  3 1V  BB @ B 3 1  V BB A 3 1VN  < B  # {#BB C 3 1BB D  3 1BB E B 3 1BB F 3 1< G  # DBB H 3 1LQBB I  3 1QBB J B 3 1LQBB K 3 1Q< L  # ) F BB M 3 11 N g BB N  3 1g BB O B 3 11 g BB P 3 1g N < Q  # 5BB R 3 1QBB S  3 1Q%BB T B 3 1QBB U 3 1%Q< V  #  F  BB W 3 1 N BB X  3 1 BB Y B 3 1 BB Z 3 1 N < [  #  Q > BB \ 3 1 BB ]  3 1 A / BB ^ B 3 1 / BB _ 3 1 A  `  B CDELFT8c  $ ' +.269=@DGKORVY]`d!h"k#o$r%v'y(})*+,./012456789;<=>?ABCDEFHIJKLNOPQRSUVW XY[\]^!_%`(b,c/d3e7f:h>iAjEkHlLmPoSpWqZr^saueviwlxpyszw|z}~~ '(@p  a  6PPfJ  cR2 G   b  6ڦ+( c22 G    c  6xJ  k = 0.41742   G   d  6    f0.012 G   e  6P   e0.12 G   f  6 dQ c12 G   g  6   c02 G   h  6 z [ d102 G   i  6   d202 G   j  6r   d302 G   k  6P X 9 d402 G   l  6([   d502 G   m  6D  | d602 G   n  6\ =  d702 G   o  6( = q distance (nt)4  G   p  6  er4 G   q  6 eI e24 G  H  0޽h ? ff3f^[^V___PPT10i.Fj+D=' = @B + mK0  7(  y  0 f -C Can substantially bias the results Even if within 10% of the age of the MRCA bottom of the tree with more branches non random subset of mutations (rare ones) small: long external branches, excess of rare variants (negative D, deficit of LD) great: a long internal branch apparent differentiation excess of intermediate frequency variants (positive D, excess of LD) if equilibratedj#n*nTnn#*T  ~  s *pqt <X   H  0޽h ? ff|ff___PPT10e.+D=' = @B + m/K0  0(  x  c $^t  `}   x  c $`_t  `  H  0޽h ? ff|ff___PPT10i.%=!+D=' = @B +F 0 p@(  d  c $?n     s *X 8r   *& we can draw one of the possible tree from the probabilistic distribution predicted by the model then lay neutral mutations on such trees and deduce their repartition in the sample below The difference is that in this case, there is an explicit probabilistic model that describes how the branching occur and what s the corresponding lengths of the branches. 4n1f_H  0og3 ? ̙33 0 04F(  4d 4 c $)Y$     4 s *,L ^   0 x The focus is on model testing This is our null hypothesis of molecular evolution: the neutral model. This is in particular its simplest and most used version for the polymorphism level: the Wright Fisher model. Based on this model, we can derive how things evolve under mutation/drift dynamic equilibrium . And that s the model we re going to test in those tests called for short neutrality tests , but which really test a full neutral model with all its assumptions : I won t detail But also heavy assumptions about demographic issues in the broad sense of anything acting at the population scale& . Implicit You can apply several mutational models that depend on the type of data. I will focus on nucleotide variation This type of variation shows the best fit with a mutational model, the ISM, which is also the most powerful to be tested, with the corresponding counterpart of heavy assumptions, in its original version, relevant mtDNA specific to this model that is there is no homoplasy So any departure from the model can potentially arise from any of these assumptions. There are ways to distinguish the different forces, which have some different predictions, but I will not really address this difficult issue today. We ll rather focus on the effect of departure from the last infinite site assumption of constant sampling time. . To make inference and test such models a widely used efficient framework relies on coalescent theory. <   ? ;   K  f  -   .  :  d   < ^ j  f g @  -    !<H 4 0og3 ? ̙33 0 P<h(  <d < c $->kO    < s *dO $   ROne can easily simulate such trees One step is to draw& in the continuous time approx They are given by probability laws of parameter p, the proba for 2 genes to have a CA in the previous generation. that is 1 over twice the pop size for 2 genes from a diploid population for n gene the prob of CA of a pair among the sampled is multiplied by the number of pairs slows down for internal branches & which allows to neglect multiple events at the same generation. 0 K@     H < 0og3 ? ̙33 0 ~pD(  Dd D c $>n   r D s *Ȥ 's   ^1-7 a 2nd step is to build the topology, by linking nodes randomly by pairs backward in time 8,9 Mutations can then be distributed uniformly on the tree, taking into account the sizes of the branches The distribution of genetic variation in the sample below can then be deduced according to the mutational model and thus to the kind of genetic markers considered. 10 Here, I ve plotted sequence data, with a non orthodox vertical representation & 11,12 for instance, this mutation A affects the 3 individuals descendents of its branch. 13 and so on. d c*r0j 0 H D 0og3 ? ̙33  0  z  (  p  0 ?O ;  h  N` R    Here is a coalescent a gene genealogy which describes the genetic history of the sample of a sample of guys the black dots on the bottom line, the present generation we go back in time along the ancestral lineages of this guys, the thin lines when a common ancestor is reached, we call this event a coalescence until the MRCA of the sample the middle tree is the corresponding coalescent representation the branches are proportional to time nodes represent common ancestors and the root the MRCA of the sample The consequence of a neutral model is that this genealogical part can be entirely uncoupled from the mutational process that s because the number of offsprings is independent of the allelic state an ind carries In this sense, it s quite different from a phylogeny. Instead of starting from the distribution of mutations in the sample to reconstruct the tree. we do just the opposite. dl fU fe, ( B  s *og3 ? ̙33Z 0 $j(  $ $  `Cj"Cj" ?Z    f We simulate thousands such trees with the parameters of the sequence data set: the sample size and number of segregating sites. The tests are conditional on these parameters. So, this is not the traditional way that conditions on theta. It was proposed by D. Hudson, discussed and validated. I ll introduce some of the stat I ll use which describe LD structure with haplotype dist. The number of haplotypes K observed on each simulation is computed Haplotype diversity H is a variant of the former test that takes the haplotype frequencies pi into account. The probability of a dataset is simply the proportions of simulations showing less the observed number of haplotypes. f?F  x $ <A  ??    H $ 0og3 ? a(80___PPT10.h`7* 0 F(  d  c $>n     s * 8r   0 H  0og3 ? ̙33 0   (   ^  S >n     c $ 8r   i viral can be followed during the whole duration of the disease no good dataset ascertainment bias issue  j H  0og3 ? ̙3380___PPT10.: 0 f^ h (  h d h c $>n   R h s *d 9q    talk about& ,that is Coalescent theory show some So let s start with...KJ K H h 0og3 ? ̙338 0 #0 N(  0 d 0 c $>n   a 0 s *la 9q  a 8LNot mathematician diff job acquiuring and checking the ancient DNA data                    H 0 0og3 ? ̙33 0  % ,(   ^  S >n     c $,  8r   " H  0og3 ? ̙3380___PPT10.0ai@ 0  - (   d  c $>n     s * 8r   1Application 12 -200 kY caves throughout europe @     H  0og3 ? ̙3380___PPT10.:A 0 - -(   d  c $)Y$     s *$ ^    Another stat we looked at as to do with the debate about potential rec in human mt DNA the most compelling indirect evidence for ,it as to do with the observation by Awadalla and coWrs of a neg correlation between LD and distance among mutations along the genome which is most simply explained by rec the data points being far from independent the r2 was tested by permutation among sites Our indirect argument is that our data are limited to a small portion of the genome D-loop region so that if such a correlation is found , recombination would need to be absurdly high to explain it and it should be due to something else like and mutational effects known to be frequent in those regions mutational effects can however predict this as suggested by Innan and Nordborg"  O C     U      i                         !<H  0og3 ? ̙33\J 0 @. (   d  c $->kO     s *JO $    simple case one subset of size n1 and time spacing t1 with the most recent subsample start with most recent subset allow coalescence among them if goes back to before the other subsampling then the evt is cancelled process started back from this point t1 with sample size updated by adding the n1 guys because& iow given no CA at this point still same P CA o otM   _      r  - H  0og3 ? ̙33p M 0   .  (   d  c $>n     s *{+ S^   jthat s the kind of data available to test the model, an alignment of polymorphic sites indicated by columns only variable sites are shown there are 15 of them here that s the S parameter with mutated nt indicated in green on a sample of 7 individuals the n parameter on the different rows Another way to sum up this information is simply to look at the frequencies of the mutations on the last row. In general you don t know what is the mutant the derived state and what is the ancestral one what is green and what is white and this affects drastically the frequencies of the mutations one way to orientate mutations is to use an outgroup to root the trees & orientate mutations. This is the kind of tree which could have lead to this data set with the corresponding mutations that s the corresponding tree, with C ancetral and a mutation to T in the intraspecific tree. and not the contrary then you can pool mutations according to their frequencies here for instance, there are & 6R@=     { H  0og3 ? ̙3380___PPT10.'nO 0 xp / (   X  C >n   p  S   8r   And this leads to this that we call the oriented frequency spectrum and in orange the unfolded one where you pool the class i with the n-i one. negative Ds result from an excess of rare variants &  > H  0og3 ? ̙3380___PPT10.p2> 0 *"0/ (   X  C >n   "  S ̔ 8r   zqualitatively pattern: because lineage cannot coalesce before contemporaneous. without rec LD stat reflect freq &_   H  0og3 ? ̙3380___PPT10.@dB 0 SK@/ (   X  C >n   K  S  8r    moderate age / MRCA sim cond theta, tested according to the resulting S left most and rightmost points correspond to the no time struct case effect max for intermediate subset sizes ^theta increases neg freq spect stat HFW= f disym int nd arbre weak effects on topology pw LD ZnS, excess of hap Differentiation gen between subpop mode position irrelevant f (sampling scheme/permutation test) only differenciation due to time struct prop of the largest subpopulation of greater age tested by perm. .C    f  C                !     E     D H  0og3 ? ̙3380___PPT10.IxC 0 4,P/ (   X  C >n     S H 8r   )     Nж g g   "   NH g g S 2J ocorresp p rej of the H0 only those above the chosen error risk >5% curves separated as one way tests depending on the direction of departure empty symbols means deficit of the stat and filled excess up to 15% for Fst neut stat >10% meaning 2 times two many N     % H  0og3 ? ̙3380___PPT10.  D 0 `X`/ (   X  C >n   X  S  8r   fix subset to half of the total f time spacing on the x axes left neutral standard case Fect inc as t1 increases non monotonic intermediate deficit of rare higher excess&[  R H  0og3 ? ̙3380___PPT10. -iF 0 p/$ y(  $ X $ C >n    $ S  8r   {9corresponding rej rates various direction f time spacing &  ) H $ 0og3 ? ̙3380___PPT10.'B? 0 D</( (  ( X ( C >n   < ( S  8r   sampling throughout Europe co Wr Lyon S. Paabo s group big sample from a single cave in belgium with precise statigraphic records Question evidence of bottleneck potentially resp for extinction struct pop alp provide barrier to gene flow in glaciations era      *     6    < H ( 0og3 ? ̙3380___PPT10.\9 0 JB/, (  , X , C >n   B , S  8r   2check contamination # of tests also chemical degeneration of DNA during and after fossilization process lead to errors in the PCR then checks between several clones. small DNA fragments D loop highly var Sums to 80 pb that can be aligned age statigraphic records or carbodated unknown bp cont                 H , 0og3 ? ̙3380___PPT10.PD; 0 /0 T(  0 X 0 C >n    0 S  8r   VD1 cav B statig. local cons discrete columns various tests 1st row obs values standard no time struct sim p, mean =0 for freq spec CI % rejection<=5% correla come back to it take into account t struct inc P() data shift stat inc rej rate timle uncertainty did not change much not general depends on sampling scheme.     7      .  '      J H 0 0og3 ? ̙3380___PPT10.2^x< 0 /4 (  4 X 4 C >n    4 S  8r   Hsame all with age estimates throughout Eur ** D* becomes more likely &(   H 4 0og3 ? ̙3380___PPT10.pZP= 0 /8 `(  8 X 8 C >n    8 S   8r   b:total inc all undated ones Fst ** across the alps turns * ; H 8 0og3 ? ̙3380___PPT10.w;I 0 80/< (  < X < C >n   0 < S   8r   `mutat 80 bp would require higher an absurd rec rate much higher than its nuclear counterpart. N   "  1 H < 0og3 ? ̙3380___PPT10.Nѫ>vx} |3i. .^움+ MiMʢ\Q.7DDdU .眙ӦN0&3sy=k ӷ?);HיD"ǫD=5/!Cb$P_k^ !,#kBID7g=2b}-ȳv{y<<FY}&A_DJ2\l `2\]$ aOH>kTL|d&ǽ)o'ُKkLc:5\TPȭI'*,P#|Ơ&459 j j5Ti6vNlPg.n^ޠ>~6B" :44 t\y5. wPËh\y >/]th,JUq h(AAIBPT*M1oaXk>wuA3 z1wP ǁ[_&P>|p/OB_bD(9?K9z=rgW u.Km1 LAVr )3ԫL:_^Yڧſg䳺^By|oʰ⿠݇x \! 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