J409 25d 507....44444444444444444444444444444444444444444444444444444444444444444!!!!! +*2 F@>>. –1 0

!"#$%& ' ()* +,- ' ./ 0"!$"%1 .,,2 ' 333456789:4;<= !"#

$%&&%'( &) *+&,'%

44444444444444444444444444444444444444444444444444444444444444444!"#$%&%' ()*+,%,#& +, -$&*&.)/&'%0+#1),&)2-$&3%#+4#5#&%,6%"78. 9: ;%",&">< =(>%"( ?: @&A),0? B @%C+( 6: ;%"'>

>!"#$%&'#() *+ ,-.-#/#0 1%2-&(3%"( *+ 45%-"*6&-27)0 8999 :*2% ;*-<0,*"*=>=>0 ,-.-#/# ?@ABB0 !CD?E*"(%&%) F-) DG>-&#>3 ;%'%-&57 H"'(#(>(%0 II99 C-"<7*=<( ;*-<0E*'' J-"<#"60 K-=#+*&"#- ?L9M?0 !CD!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!

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

@:A7:=K:9 2--L 7< M:;:=K:9 2--N 67 7G: O636CPC );:65 #C=:QE:9C:E E767C<5 "*)O"L R..! +/" !S 2/N! ,,"TU C5 7G: !<97G V6;CW;E8K79<AC;6B DH9:4 T: X8657CW:F ;:BBE K:B<5DC5D C5 7G: F<=6C5Y6;7:9C6S 65F 73< EA:;CW; AGHB<D:5:7C; D9<8AE C5 7G: F<=6C5"9;G6:6S 3GC;G 69: A9:I6B:57 C5 =69C5: AB65Z7<5[ \D9<8A 2 69;G6:6PR9:J:99:F 7< G:9: 6E A:B6DC; ;9:569;G6:<76U] 65F \D9<8A . 69;G6:6P

20

30

40

50

60

70

80

Dep

th (m

)

% o

f D

AP

I co

unt

5,000

1,000

100

10

5

5

10

15

20

25

30

35

40

Dep

th (m

)

S O N D J F M A M J J A S O N D

1998

% o

f D

AP

I co

unt

5,000

1,000

100

10

5

Bacteria

Crenarchaeota

!"#$%& ' !"#$"%& '("$) "* &+(,$-.+ ,/%#0,#1+) 2-$3 0+'$3 "* /,1$+&-, ,#0 '+(,4-1

1&+#,&13,+"$, 0%&-#4 , 567& ),8'(-#4 +**"&$ ,$ $3+ 9,2,-:- ;1+,# <-8+6)+&-+) )$,$-"#=

>?;9>= -# $3+ @"&$3 A,1-B1 )%/$&"'-1,( 47&+C D3-$+ 0"$) -#0-1,$+ 0,$+) ,#0 0+'$3) 23+&+

),8'(+) 2+&+ 1"((+1$+0C !"#$"%& (-#+) ,&+ '+&1+#$,4+) "* /,1$+&-, ,#0 '+(,4-1

1&+#,&13,+"$, ,) 1"8',&+0 2-$3 $"$,( 8-1&"/-,( ,/%#0,#1+ ,$ +,13 0+'$3C <"$,( 1+((

,/%#0,#1+ 2,) ,))+))+0 %)-#4 $3+ E>AF #%1(+-1 ,1-0 )$,-#C G,1$+&-, ,#0 ,&13,+, 2+&+

+#%8+&,$+0 %)-#4 23"(+61+(( &H@> $,&4+$+0 I%"&+)1+#$ !" #!$% 37/&-0-J,$-"# 2-$3

I%"&+)1+-#6(,/+((+0 '"(7#%1(+"$-0+ '&"/+)C K++ ,()" K%''(+8+#$,&7 F#*"&8,$-"#C

© 2001 Macmillan Magazines Ltd

R9:J:99:F 7< G:9: 6E A:B6DC; :89H69;G6:<76U.4 %CK<E<=6B %!"Q769D:7:F A<BH58;B:<7CF: A9<K:E^ EA:;CW; J<9 :C7G:9 A:B6DC; ;9:569;GQ6:<76S :89H69;G6:<76 <9 K6;7:9C6 3:9: GHK9CFC_:F 3C7G 6BB E6=AB:E;<BB:;7:F J9<= @:A7:=K:9 2--L 857CB M:;:=K:9 2--N R<5: F::AE6=AB: ;<BB:;7:F C5 );7<K:9 2--N G6F E79<5D EA89C<8E K6;ZD9<85F`8<9:E;:5;: 65F 36E <=C77:F J9<= 7G: F676 E:7U4 V9<K:QKC5FC5D;:BBE 3:9: :58=:967:F 9:B67CI: 7< 7<76B ;:BBE E76C5:F KH +"S^QFCQ6=CFC5<Q.QAG:5HBC5F<B: RM"VaUN F<8KB: E76C5C5D-4 V9<K:Q;<5QJ:99:F `8<9:E;:5;: 36E =<E7 C57:5E: C5 E6=AB:E J9<= EG6BB<3:9F:A7GE R,b/,, =US 65F 36E 3:6Z:9 J<9 K<7G 69;G6:6 65F K6;7:9C6 67F:A7GE D9:67:9 7G65 /,, =4 !:D67CI: ;<579<B ;<857E RGHK9CFC_67C<53C7G 6 5<5EA:;CW; A9<K: 65F 687<`8<9:E;:5;:U 3:9: K:B<3 /c <J7<76B ;:BBE C5 -^c <J 6BB E6=AB:ES G:5;: 6 GCDG F:D9:: <J ;<5WF:5;: C57GCE =:7G<F 36E 6;GC:I:F4 R@:: 6BE< @8AAB:=:5769H a5J<9=67C<54U#G: 73< 69;G6:6B D9<8AE E89I:H:FS 6E 3:BB 6E K6;7:9C6 ;<BB:;7CI:BHS

B6;Z:F ;B:69 E:6E<56B 79:5FE C5 9:B67CI: ;:BB8B69 6K85F65;: RdCD4 2U4#G:9:J<9:S 6I:96D:E 3:9: ;6B;8B67:F J<9 6BB E6=AB:E J<9 :6;G F:A7GB6H:9S HC:BFC5D 7G: =:65 65586B F:A7G A9<WB: J<9 :6;G AGHB<7HA:RdCD4 .6U4 e:BBE GHK9CFC_C5D 3C7G 7G: K6;7:9C6B A9<K: F<=C567:F 7G:A<A8B67C<5 C5 7G: 8AA:9 2/,= <J 7G: 367:9 ;<B8=5S 9:A9:E:57C5D 8A7< -,c <J 6BB ;:BBE4 Y6;7:9C6 F:;9:6E:F C5 9:B67CI: 6K85F65;: 3C7GC5;9:6EC5D F:A7G RdCDE 2S .6US 65F K:B<3 2S,,,= 7G:H 9:A9:E:57:F<5BH f/b+,c <J 7<76B ;:BBE4 YH ;<=A69CE<5S A:B6DC; ;9:569;G6:<76C5;9:6E:F EG69ABH C5 9:B67CI: 6K85F65;: 67 7G: ./,Q= F:A7G B6H:9S65F K:B<3 2S,,,=3:9: 6E ;<==<5 6E K6;7:9C6 RdCDE 2S .6U4 @6=AB:EJ9<= 2/,= RC5;B8F:F C5 + <J 7G: 2+ E6=ABC5D F67:EU C5FC;67:F 7G677G: 9:B67CI: C5;9:6E: C5 A:B6DC; ;9:569;G6:<76 <;;899:F K:73::5 2,,65F 2/, =S 5:69 7G: F:A7G <J 7G: 2c CE<B8=: RdCDE 2S .6U4 a5 E89J6;:

B6H:9ES A:B6DC; ;9:569;G6:<76 3:9: <5BH A9:E:57 EA<96FC;6BBHS 65F5:I:9 6K85F657 58=:9C;6BBH4 V:B6DC; :89H69;G6:<76 <;;6EC<56BBH6AA:69:F C5 7G: 5:69 E89J6;: B6H:9S K87 D:5:96BBH 9:=6C5:F 67 6 J:3A:9 ;:57 <J 7G: 7<76B ;<857 <I:9 7G: :57C9: 367:9 ;<B8=5S 7<< ;B<E: 7<5:D67CI: ;<857E 7< :E76KBCEG 6 E767CE7C;6BBH 9:BC6KB: :E7C=67: RdCD4 .6U4V:B6DC; ;9:569;G6:<76 65F :89H69;G6:<76 7G8E EG<3:F FCJJ:9:57A677:95E <J 6K85F65;: C5 7G: <A:5 E:6 RdCD4 .6US 3C7G B<3 58=K:9E<J :89H69;G6:<76 C5 E89J6;: 367:9ES C5 ;<5796E7 7< A9:IC<8E <KE:9I6Q7C<5E C5 ;<6E76B 367:9E^S2,S224#G: E8= <J 9:B67CI: 6K85F65;:E <J K6;7:9C6S A:B6DC; :89H69;G6:<76

65F A:B6DC; ;9:569;G6:<76 RE8K796;7C5D 5:D67CI: ;<579<BEU EG<3E 7G:7<76B J96;7C<5 <J A9<K:QKC5FC5D ;:BBE <I:9 ;:BBE E76C5:F KH M"VaRdCD4 .KU4 "B7G<8DG <89 F::AQE:6 F676 <5 9:B67CI: 6K85F65;: <JA:B6DC; ;9:569;G6:<76 RdCD4 .6U 69: DB<K6BBH GCDG:9 7G65 7G<E:9:;:57BH 9:A<97:F C5 ;<6E76B 367:9E^S <89 ;8=8B67CI: 9:B67CI: 6K85QF65;: F676 <5 7<76B A9<K:QA<EC7CI: ;:BBE RdCD4 .KU 69: B<3:94 #GCE =6HK: F8: 7< 7G: <BCD<79<AGC; 56789: <J 7G: E78FH EC7:4 #<76B ;:BB6K85F65;: A677:95E 9:`:;7:F 7G: GCDG:9 7<76B ;:BB 58=K:9E C5E89J6;: 367:9E RdCD4 fU4 #<76B ;:BB 6K85F65;: F:;BC5:F KH 65 <9F:9<J =6D5C78F: K:73::5 2/, 65F 2S,,, =4 #G8ES 7G: 7<76B 58=K:9 <JA:B6DC; ;9:569;G6:<76B ;:BBE A:6Z:F K:73::5 2/, 65F /,, =S K879:=6C5:F 67 GCDG 9:B67CI: 58=K:9E K:B<3 7GCE F:A7G 7< 7G: <;:65`<<94a5 9:;:57 H:69E ;8B7CI67C<5QC5F:A:5F:57 9%!" D:5: E89I:HE G6I:

9:I:6B:F 5:3 7HA:E <J 69;G6:6 C5 IC9786BBH :I:9H :;<EHE7:=:g6=C5:F2.S2f4 1:7 =69C5: AB65Z7<5C; 69;G6:6B 7HA:E G6I: :B8F:F;8B7CI67C<5 7< F67:S 65F J:3 X8657C767CI: F676 :gCE7 <5 7G:C9 ;:BB8B69FCE79CK87C<5 65F 6K85F65;:4 h<E7 A9:IC<8E E78FC:E G6I: 9:BC:F <57G: 9:79C:I6B <J 9%!"QD:5:Q;<576C5C5D ;B<5:E 7< X86BC767CI:BHF:E;9CK: 567896BBH <;;899C5D AC;<AB65Z7<5 A<A8B67C<5E.b+4 "58=K:9 <J E78FC:E 8EC5D X8657C767CI: 9%!" GHK9CFC_67C<57:;G5CX8:E.S2,S22S2+S2/ G6I: E8DD:E7:F 6 3CF:EA9:6F <;;899:5;: 65FGCDG 9:B67CI: ;:BB 6K85F65;: <J 69;G6:6 C5 7G: 8AA:9 367:9 ;<B8=54$5J<978567:BHS 7G:E: :69BC:9 E78FC:E F< 5<7 6BB<3 J<9 X8657CW;67C<5<J 6;786B ;:BB 58=K:9E4 i8657C767CI: =C;9<E;<AC; E89I:HE 8EC5DEC5DB:Q;:BBS `8<9:E;:57 #" '#(> GHK9CFC_67C<5 3C7G 9%!"Q769D:7:F<BCD<58;B:<7CF: A9<K:ES KH ;<5796E7S 6BB<3 FC9:;7 ;:BB :58Q=:967C<52^S2L4 " J:3 C5I:E7CD67C<5E 8EC5D <BCD<58;B:<7CF: A9<K:E 7<:58=:967: =69C5: AC;<AB65Z7<5/S2Nb., G6I: E8;;::F:F C5 F:7:;7C5D

$%&&%'( &) *+&,'%

!"- !"#$%& ' ()* +,- ' ./ 0"!$"%1 .,,2 ' 333456789:4;<=

60 80 100

14

1414

14

4

4

9

9

14

14

9

All probes

0 20 40 60 80

10

100

1,000

5,000

Microbial domains

Crenarchaeota Euryarchaeota Bacteria

Negative controlSum of probes

Percentage of DAPI count

Depth (m)

a b

!"#$%& (L+,# ,##%,( 0+'$3 '&"B(+) "* 8-1&"/-,( 0"8,-#) -# $3+ @"&$3 A,1-B1 )%/$&"'-1,(47&+C @%8/+&) ,&+ '+&1+#$,4+) "* /,1$+&-, ,#0 ,&13,+, ,) 1"8',&+0 $" $"$,( 8-1&"/-,(

,/%#0,#1+ ,$ +,13 0+'$3C <"$,( 1+(( ,/%#0,#1+ 2,) ,))+))+0 %)-#4 $3+ E>AF #%1(+-1 ,1-0

)$,-#C G,1$+&-, ,#0 ,&13,+, 2+&+ +#%8+&,$+0 %)-#4 23"(+61+(( &H@> $,&4+$+0 I%"&+)1+#$

!" #!$% 37/&-0-J,$-"# 2-$3 I%"&+)1+-#6(,/+((+0 '"(7#%1(+"$-0+ '&"/+)C E,$, ,&+ ,.+&,4+) "*

%' $" 5M &"%43(7 8"#$3(7 ),8'(-#4) ".+& , 567& '+&-"0 ,$ $3+ 9,2,-:- ;1+,# <-8+6)+&-+)

)$,$-"#= >?;9>C N&&"& /,&) )3"2 )$,#0,&0 +&&"& "* 8+,#O #"$+ 1"(%8# *"& $"$,( ),8'(+ )-J+

,$ +,13 0+'$3C K++ ,()" K%''(+8+#$,&7 F#*"&8,$-"#C )= E+'$3 '&"B(+) *"& /,1$+&-, P)"(-0)Q%,&+)R= '+(,4-1 1&+#,&13,+"$, P"'+# )Q%,&+)R= '+(,4-1 +%&7,&13,+"$, P"'+# 1-&1(+)R=

,#0 , #"#6)'+1-B1 1"#$&"( '&"/+ PS#+4,$-.+:= )"(-0 1-&1(+)RC *= E+'$3 '&"B(+ "* $3+ )%8 "*

&+(,$-.+ ,/%#0,#1+) "* /,1$+&-,= '+(,4-1 1&+#,&13,+"$, ,#0 '+(,4-1 +%&7,&13,+"$, P"'+#

0-,8"#0)RC H+(,$-.+ ,/%#0,#1+) "* /,1$+&-, ,#0 /"$3 ,&13,+,( 4&"%') 2+&+ )%88,$+0 ,$

+,13 0+'$3 ,#0 #+4,$-.+ 1"#$&"( 0,$, 2+&+ )%/$&,1$+0C

103 104 105 106

10

100

1,000

Sum of probes5,000

Total microbial cells per ml

Depth (m)

ArchaeaBacteria

!"#$%& +L+,# ,##%,( 0+'$3 '&"B(+) "* 8-1&"/-,( 0"8,-#) -# $3+ @"&$3 A,1-B1 )%/$&"'-1,(47&+C @%8/+&) ,&+ $"$,( 1+(( ,/%#0,#1+) "* /,1$+&-, ,#0 ,&13,+, P'+(,4-1 1&+#,&13,+"$,

,#0 +%&7,&13,+"$, 1"8/-#+0RC G,1$+&-, ,#0 ,&13,+, 2+&+ +#%8+&,$+0 %)-#4 23"(+61+((

&H@> $,&4+$+0 I%"&+)1+#$ !" #!$% 37/&-0-J,$-"# 2-$3 I%"&+)1+-#6(,/+((+0 '"(7#%1(+"$-0+

'&"/+)C E,$, ,&+ ,.+&,4+) "* %' $" 5M &"%43(7 8"#$3(7 ),8'(-#4) ".+& , 567& '+&-"0 ,$ $3+

9,2,-:- ;1+,# <-8+6)+&-+) )$,$-"#= >?;9>C K++ ,()" K%''(+8+#$,&7 F#*"&8,$-"#C


567896BBH <;;899C5D K6;7:9C6B 65F 69;G6:6B ;:BBE4 )5: E78FH/ 8EC5DC=6D: C57:5ECW;67C<5 7< :5G65;: 7G: `8<9:E;:5;: ECD56B 9:A<97:FGCDG A:9;:576D:E <J "9;G6:6 R8A 7< ^,cU C5 ;<6E76B 7:=A:967:367:9E 65F 7G: h:FC7:9965:65 @:64 @:I:96B E78FC:ES 8EC5D 65 <BCD<Q58;B:<7CF: A9<K: 769D:7C5D 7G: F<=6C5 "9;G6:6S F:7:;7:F J:3 <9 5<69;G6:6B ;:BBE C5 7G: 8AA:9 367:9 ;<B8=52-S.,4!<5: <J 7G:E: :69BC:9 E78FC:ES G<3:I:9S :g6=C5:F F:A7GE K:B<3

^,, =S 65F <5BH <5:2N CF:57CW:F EA:;CW; 69;G6:6B ;:BB 7HA:E K:H<5F7G: F<=6C5 B:I:B4 d897G:9S EC5DB:Q;:BB GHK9CFC_67C<5 E78FC:E G6I:K::5 G6=A:9:F KH 7G: E=6BB EC_: 65F 6AA69:57 B<3 9%!" ;<57:57 <J=69C5: AC;<AB65Z7<5/S^S2N4 h8B7CABH B6K:BB:FS 9%!"Q769D:7:F A<BHQ58;B:<7CF: A9<K:E^ 6BB<3 =<9: EHE7:=67C; :58=:967C<5 <J EA:;CW;D9<8AE <J AB65Z7<5C; =C;9<K:E4 Y87 7GCE 6AA9<6;G G6E E< J69 K::5F:=<5E7967:F <5BH 67 6 ;<6E76B <;:65 EC7:^S A9<=A7C5D <89 E78FH 677G: O636CC );:65 #C=:QE:9C:E E767C<54a5 7GCE 9:A<97S 3: ;<5;B8ECI:BH EG<3 7G67 A:B6DC; ;9:569;G6:<76

3:9: :X8CI6B:57 C5 58=K:9E 7< K6;7:9C6 7G9<8DG<87 7G: :57C9: =:E<Q65F K67GHA:B6DC; _<5:E C5 6=6j<9 <;:65 DH9:S <I:9 6 2QH9 7C=:QE:9C:EE89I:H4 #3< =6C5 A<C57E 69CE: J9<= <89 <KE:9I67C<5E4 dC9E7S <56I:96D:S 7G: J96;7C<5 <J A:B6DC; ;9:569;G6:<76 9:B67CI: 7< M!"Q;<576C5C5D =C;9<<9D65CE=E :X86BE <9 :g;::FE 7G: K6;7:9C6B J96;7C<5K:B<3 2S,,,= RdCD4 .6U4 #GCE 9:E8B7 E8DD:E7E 7G67 A:B6DC; ;9:569;GQ6:<76 69: 6 ;<5ECE7:57 65F ECD5CW;657 ;<=A<5:57 <J F::A E:6=C;9<KC<76S 65F 7G67 7G:H =6H 9CI6B K6;7:9C6B 6K85F65;:E C5 7G:=:E<Q 65F K67GHA:B6DC; _<5:E4 d897G:9S 7G: E765FC5D E7<;ZE <JK6;7:9C6 65F A:B6DC; ;9:5<9;G6:<76 3:9: C5I:9E:BH A9<A<97C<56B4@:;<5FS 7G: 7<76B A9<A<97C<5 <J ;:BBE RK6;7:9C6B 65F 69;G6:6BUF:7:;7:F 3C7G A<BH58;B:<7CF: A9<K:E RdCD4 .KU 9:=6C5:F J6C9BH;<5E7657 C5 E89J6;: 367:9E 7< F:A7GE <J 2S,,, =S 9<8DGBH N,c <J7G: 7<76B M!"Q;<576C5C5D ;:BBEN4 "7 F:A7GE K:73::5 .S,,, 65F fS,,,=S 7GCE 58=K:9 F:;9:6E:F 7< 6K<87 ^,c K:J<9: C5;9:6EC5D 6D6C5 7<=<9: 7G65 L,c ;B<E: 7< 7G: E:6 `<<9 RdCD4 .KUS EC=CB69 7< A9:IC<8EBH9:A<97:F =C;9<KC6B ;:BB 58=K:9E 65F =:76K<BC; 6;7CIC7H A9<WB:E C57G: F::A V6;CW;.24 M!"Q;<576C5C5D ;:BBE 85B6K:BB:F KH A<BH58;B:<Q7CF: A9<K:E =CDG7 C5;B8F: ;:BBE 3C7G B<3 9%!" ;<57:57S ;:BBE 5<79:;<D5C_:F KH 7G: A9<K:E RJ<9 :g6=AB:S 858E86BBH E=6BB N>5-&)- C57G: 2Q"= 965D:..U <9 F:6F ;:BBE RDG<E7EU.f4 )89 9:E8B7E AB6;: 658AA:9 BC=C7 <5 7G: E8=<J 7G: 6K<I: ;67:D<9C:E4T: ;<5;B8F: 7G67 7G:8E: <J A<BH58;B:<7CF: A9<K:E A9<ICF:E 6 9<K8E7 6AA9<6;G J<9CF:57CW;67C<5 <J F::AQE:6 =C;9<K:ES 65F 7G67 7G: J96;7C<5 <J 9:=6C5QC5D 85B6K:BB:F ;:BBE 9:=6C5E K:B<3 .,bf,c <J 7G: 7<76B ;:BBE4" A9:IC<8E E78FH.+ E8DD:E7:F 7G67 7G: 58=K:9 <J ;:BBE A<EC7CI: J<9

9%!" GHK9CFC_67C<5 7:;G5CX8:E =6H 9:`:;7 7G: 58=K:9 <J =:76K<QBC;6BBH 6;7CI: ;:BBE4 aJ 798:S 7G:5 7G: 7<76B 58=K:9 <J A9<K:QKC5FC5D;:BBE EG<8BF 9:A9:E:57 7G: A9<A<97C<5 <J =:76K<BC_C5D ;:BBE C5 7G:367:9 ;<B8=5 R^,bN,c <J 7<76B ;:BBE A9:E:57U4 #G: F:;9:6E: C5A9<K:QA<EC7CI: ;:BBE K:73::5 .S,,, 65F fS,,,= =CDG7 9:`:;7F:;9:6EC5D 6I6CB6KCBC7H <9 X86BC7H <J D9<37G E8KE7967:S 9:E8B7C5D C56 =:76K<BC;6BBH B:EE 6;7CI: A<A8B67C<54 eB<E: 7< 7G: E:6 `<<9SD96IC767C<56BBH F:A<EC7:F <9D65C; =677:9 =CDG7 79CDD:9 7G: <KE:9I:FC5;9:6E: <J 7G: A9<A<97C<5 <J 6BB A9<K:QA<EC7CI: ;:BBE RdCD4 .KU4 #GCE9:E8B7 CE ;<5ECE7:57 3C7G <KE:9I:F C5;9:6E:E C5 =:76K<BC; 6;7CIC7H5:69 7G: E:6 `<<9 A9:IC<8EBH J<85F C5 V6;CW; K67GHA:B6DC; =C;9<KC6BA<A8B67C<5E.24 )89 F676 C5FC;67: 7G67 =<E7 F::AQE:6 =C;9<K:ER69;G6:6 65F K6;7:9C6U ;<576C5 6AA9:;C6KB: 6=<857E <J 9%!"S 65FE< =6H K: 6;7CI: ;<579CK87<9E 7< 7G: :;<EHE7:=4 d897G:9=<9:S DCI:57G67 7G: 7<76B 58=K:9 <J =C;9<K:E E79<5DBH F:;9:6E:E 3C7G F:A7G C57G: <;:65.2S./S <89 F676 B:6F 7< 65 :;<B<DC;6BBH ECD5CW;657 ;<5;B8EC<5[D9<37G ;<5FC7C<5E J<9 A:B6DC; =C;9<K:E F< 6AA:69 7< K: 6;;8967:BH9:`:;7:F KH =C;9<KC6B E765FC5D E7<;Z4 #G:9: =6H 6BE< K: 6 A<A8B6Q7C<5 EGCJ7 3C7G C5;9:6EC5D F:A7G 7< =C;9<KC6B D9<8AE K:77:9 6F6A7:F7< F::A E:6 ;<5FC7C<5ES 6 B69D: J96;7C<5 <J 3GC;G 6AA:69E 7< K:A:B6DC; ;9:569;G6:<764e<=KC5C5D <89 A9:E:57 F676 E:7 3C7G ;<57C5:576B EG:BJ F676^S 3:

;65 :g796A<B67: 7<76B ;:BB 6K85F65;:E J<9 69;G6:6 7< 7G: DB<K6B<;:65.^4 @8;G 65 :g796A<B67C<5 E8DD:E7E 7G67 7G: 3<9BF <;:65;<576C5E 6AA9<gC=67:BH 24f ! 2,.N 69;G6:6B ;:BBES 65F f42 ! 2,.N

K6;7:9C6B ;:BBE4 "K<87 24, ! 2,.N ;:BBES 7G67 CES .,c <J 6BB 7G: AC;<QAB65Z7<5 ;:BBE C5 7G: 3<9BF <;:65 6AA:69 7< K: 9:A9:E:57:F KH <5:EA:;CW; ;B6F:S 7G: A:B6DC; ;9:569;G6:<764 #G: G6KC767 965D: J<9 7GCEEC5DB: =C;9<KC6B D9<8AS EA655C5D J9<==:E<A:B6DC; 7< K67GHA:B6DC;F:A7GES CE 858E86BBH K9<6F4 "E 6 F<=C5657 ;<=A<5:57 <J 7G: F::A<;:65S &697GPE B69D:E7 KC<=:S 69;G6:6 69: 7G8E J69 J9<= ;<5W5:F 7<:g79:=: 5C;G: G6KC767E4 %67G:9S 7G: FCE79CK87C<5 <J 7G:E: 69;G6:6E8DD:E7E 7G67 6 ;<==<5 6F6A7CI: E7967:DH G6E 6BB<3:F 7G:= 7<96FC67: 7G9<8DG<87 5:69BH 7G: :57C9: <;:65C; 367:9 ;<B8=54 !

,&-./012)345& 4%&4)%)-"/6

@6=AB:E 3:9: Wg:F 3C7G ,4.Q"= WB7:9:F J<9=6BC5 R.c W56B ;<5;:57967C<5U 65F WB7:9:F<57< ,4.Q"= A<BH;69K<567: WB7:9E4 "J7:9 WB7967C<5 ;:BBE 3:9: 79:67:F KH <I:9B6HC5D 7G:WB7:9 J<9 .=C5 3C7G 6 ,4./ h !6eB E<B87C<5 =6F: 8A C5 /,c RIkIU :7G65<B4 @6=AB:E3:9: E7<9:F J9<_:5 :C7G:9 K:J<9: <9 6J7:9 WB7967C<5S 3C7G :X86B A9:E:9I67C<5 <J A9<KC5DA<7:57C6B4

7./5&89&55 :$/%&19&6- !" #!$% .;*%"0"<)-"/6 9/$6-1

%CK<E<=6B %!" 769D:7:F GHK9CFC_67C<5 36E ;699C:F <87 <5 7G: WB7:9QK<85F ;:BBE 8EC5D`8<9:E;:C5 CE<7GC<;H6567: Rda#eU =8B7CABH B6K:BB:F A9<K:E^ <J =<9: 7G65 2,, K6E: A6C9E C5B:5D7GS 769D:7C5D ;9:569;G6:<76 R=69C5: D9<8A 2 69;G6:6U <9 :89H69;G6:<76 R=69C5: D9<8A. 69;G6:6U2,S <9 K6;7:9C64 " 5<5QKC5FC5D A9<K: R;<5ECE7C5D <J 7G: ;<=AB:=:5769H E7965F <J7G: A:B6DC; ;9:569;G6:<76B A9<K:U E:9I:F 6E 7G: 5:D67CI: ;<579<BS 3GCB: 7G: E8==67C<5 <J69;G6:6B 65F K6;7:9C6B J96;7C<5E R=C58E 5:D67CI: ;<579<BU A9<ICF:F 6 ;G:;Z <5 G<3 7<76BA9<K:QA<EC7CI: ;:BBE ;<=A69:F 3C7G 7<76B ;:BB ;<857E 8EC5D 7G: M"Va E76C5N4 V9<K: 36E6FF:F 7< 6 GHK9CFC_67C<5 K8JJ:9 67 6 W56B ;<5;:57967C<5 <J . 5D"B"24 #G: GHK9CFC_67C<5K8JJ:9 <J 2,c R3kIU F:g7965 E8BAG67:S ,4,2c A<BHR"U 65F ,42c E<FC8= F<F:;HB E8BAG67:R@M@U C5 /! @&#36E 8E:F 3C7G :C7G:9 /,c J<9=6=CF: RK6;7:9C6B 65F 5:D67CI: ;<579<BA9<K:EU <9 L,c J<9=6=CF: RJ<9 K<7G 69;G6:6B A9<K:ES 2! @&# CE 2/,=h !6eBS 2=h:7GHB:5:FC6=C5:7:7966;:7C; 6;CF R&M#"US .,=h #9CES AO N4,U4T: FCF 5<7 687<=67C;6BBH E8K796;7 7G: 5:D67CI: ;<857 ;<99:EA<5FC5D 7< :6;G E6=AB:

K:;68E: ;:BBE EG<3C5D 6 \5:D67CI:P C=6D:S J<9 :g6=AB:S <3C5D 7< 687<`8<9:E;:5;:S ;<8BF K:<J 65H D9<8A[ ;9:569;G6:<76S :89H69;G6:<76 <9 K6;7:9C64 TG:5 9:A<97C5D 7G: E8= <J9:B67CI: ;:BB 58=K:9E J9<= 6BB A9<K:ES G<3:I:9S 3: FCF E8K796;7 7G: 5:D67CI: ;<857 6E G:9:3: ;65 9:B67: 7G: 5:D67CI: ;<857 7< 6BB ;:BBES HC:BFC5D 6 ;<5E:9I67CI: :E7C=67: J<9 7<76B A9<K:A<EC7CI: ;:BBE4 M89C5D "A9CB 2--NS GCDG 5:D67CI: I6B8:E <J ./c 65F f^c 3:9: <KE:9I:F J<97G: /Q= 65F 7G: ./Q= B6H:9ES 9:EA:;7CI:BH] J<9 7G: <7G:9 2., E6=AB:E 7G: 5:D67CI: ;<579<B6I:96D:F .4,# 24/c R=:65# E4F4U4 OHK9CFC_67C<5 7:=A:96789:E 3:9: ^/ !e J<9 7G: 73<69;G6:6B A9<K:ES 65F // !e J<9 K6;7:9C6B 65F 5:D67CI: ;<579<B A9<K:E4 OHK9CFC_67C<5 7C=:36E 2. G4 V<E7QGHK9CFC_67C<5 36EG R. GU 36E 67 /, !e J<9 69;G6:6B A9<K:ES 65F +/ !e J<9K6;7:9C6Bk5:D67CI: ;<579<B A9<K:E C5 6 ,4.! @&# E<B87C<5 =6F: 8A C5 /,c J<9=6=CF:4dCB7:9E 3:9: =<857:F C5 eC7C`8<9 R#:F V:BB6US 65F da#eQA<EC7CI: ;:BBE 3:9: ;<857:F C59:B67C<5 7< M"VaQA<EC7CI: ;:BBE <5 7G: E6=: WB7:9-4 e<857C5D 36E A:9J<9=:F 8EC5D 6 l:CEE:AC`8<9:E;:5;: =C;9<E;<A: :X8CAA:F 3C7G 6 !2,, !:<`869 <Kj:;7CI: 65F 2,,QT=:9;89HB6=A CBB8=C567C<5S 6E 3:BB 6E 7G: 6AA9<A9C67: WB7:9 E:7E J<9 M"Va 65F da#e ReG9<=6#:;G5<B<DHU4

%:;:CI:F + 08BH] 6;;:A7:F .2 !<I:=K:9 .,,,4

24 T<:E:S e4 %4S m65FB:9S )4 n TG::BCES h4 *4 #<369FE 6 567896B EHE7:= <J <9D65CE=E[ V9<A<E6B J<9 7G:

F<=6C5E "9;G6:6S Y6;7:9C6S 65F &8;69H64 :&*5O P-(= D5-<O C5#O !CD XM5 +/L^b+/L- R2--,U4

.4 M:*<5DS &4 d4 "9;G6:6 C5 ;<6E76B =69C5: :5IC9<5=:57E4 :&*5O P-(= D5-<O C5#O !CD XR5 /^N/b/^N-

R2--.U4

f4 d8G9=65S 04 "4S h;e6BB8=S m4 n M6ICES "4 "4 !<I:B =6j<9 69;G6:K6;7:9C6B D9<8A J9<= =69C5:

AB65Z7<54 P-(>&% QFG5 2+Nb2+- R2--.U4

+4 d8G9=65S 04 "4 n M6ICES "4 "4 TCF:EA9:6F "9;G6:6 65F 5<I:B Y6;7:9C6 J9<= 7G: F::A E:6 6E EG<35 KH

2^@ 9%!" D:5: E:X8:5;:E4 E-&O N5*=O :&*6O C%&O >FN5 .L/b.N/ R2--LU4

/4 d8G9=65S 04 "4 n )8I:95:HS e4 e4 h69C5: =C;9<KC6B FCI:9EC7H E78FC:E IC6 2^@ 9%!" E:X8:5;:E[ ;B<5C5D

9:E8B7E J9<= ;<6E76B 367:9E 65F ;<857C5D <J 567CI: 69;G6:6 3C7G `8<9:E;:57 EC5DB: A9<K:E4 DG>-(O N5*=O

Q?5 fb2/ R2--NU4

^4 M:*<5DS &4 d4S #6HB<9S *4 #4S h69EGS #4 *4 n V9:E7<5S e4 h4 (CE86BC_67C<5 65F :58=:967C<5 <J =69C5:

AB65Z7<5C; D&57-%- 65F F-5(%&#- KH 8EC5D A<BH9CK<58;B:<7CF: A9<K:E 65F `8<9:E;:57 #" '#(>

GHK9CFC_67C<54 D22=O N"$#&*"O E#5&*Q#*=O GF5 ///+b//^f R2---U4

L4 m69BS M4 h4 n *8Z6ES %4 #G: O636CC );:65 #C=:QE:9C:E RO)#U A9<D96=[ Y6;ZD9<85FS 967C<56B: 65F

W:BF C=AB:=:5767C<54 1%%2RC%- ;%'O AQ5 2.-b2/^ R2--Û4

N4 V<97:9S m4 n d:CDS 14 @4 #G: 8E: <J M"Va J<9 CF:57CJHC5D 65F ;<857C5D 6X867C; =C;9<`<964 J#3"*=O

45%-"*6&O ?F5 -+fb-+N R2-N,U4

-4 OC;ZES %4 &4S "=655S %4 a4 n @76GBS M4 "4 M86B E76C5C5D <J 567896B K6;7:9C<AB65Z7<5 3C7G +"S ^QFC6=CFC5<Q.QAG:5HBC5F<B: 65F `8<9:E;:57 <BCD<58;B:<7CF: A9<K:E 769D:7C5D ZC5DF<=QB:I:B 2^@ 9%!"

E:X8:5;:E4 D22=O N"$#&*"O E#5&*Q#*=O FX5 .2/Nb.2^f R2--.U4

2,4 h6EE656S %4S h8996HS "4 &4S V9:E7<5S e4 h4 n M:*<5DS &4 d4 (:97C;6B FCE79CK87C<5 65F AGHB<D:5:7C;

;G696;7:9C_67C<5 <J=69C5: AB65Z7<5C;D&57-%- C5 7G: @6576 Y69K696 eG655:B4D22=O N"$#&*"OE#5&*Q#*=O

GQ5 /,b/^ R2--LU4

224 h8996HS "4 &4 %( -=O " 7C=: E:9C:E 6EE:EE=:57 <J AB65Z7<5C; 69;G6:6B I69C6KCBC7H C5 7G: @6576 Y69K696

eG655:B4 DG>-(O E#5&*QO N5*=O ?N5 2.-b2+/ R2---U4

2.4 )BE:5S o4 04 "9;G6:6S 69;G6:6 :I:9H3G:9:4 P-(>&% QM>5 ^/Lb^/N R2--+U4

2f4 @7:C5S 04 *4 n @C=<5S h4 a4 "9;G6:6B 8KCX8C7H4 :&*5O P-(= D5-<O C5#O !CD RQ5 ^..Nb^.f, R2--Û4

2+4 M:*<5DS &4ST8S m4 14S V9:p_BC5S Y4 Y4 n 0<IC5:S %4 (4h4OCDG 6K85F65;: <J "9;G6:6 C5 "5769;7C;=69C5:

AC;<AB65Z7<54 P-(>&% QM>5 ^-/b^-L R2--+U4

$%&&%'( &) *+&,'%

!"#$%& ' ()* +,- ' ./ 0"!$"%1 .,,2 ' 333456789:4;<= !".© 2001 Macmillan Magazines Ltd

2/4 h6EE656S %4 %( -=O (:97C;6B FCE79CK87C<5 65F 7:=A<96B I69C67C<5 <J =69C5: AB65Z7<5C; "9;G6:6 C5 7G:

o:9B6;G: E796C7S "5769;7C;6S F89C5D :69BH EA9C5D4 J#3"*=O 45%-"*6&O AQ5 ^,Lb^2L R2--NU4

2^4 "=655S %4 a4S m98=G<B_S *4 n @76GBS M4 "4 dB8<9:E;:57Q<BCD<58;B:<7CF: A9<KC5D <J 3G<B: ;:BBE J<9

F:7:9=C567CI:S AGHB<D:5:7C;S 65F :5IC9<5=:576B E78FC:E C5 =C;9<KC<B<DH4 SO F-5(%&#*=O >M?5 L^.bLL,

R2--,U4

2L4 *::S @4 n m:=AS V4 d4 @C5DB:Q;:BB %!" ;<57:57 <J 567896B =69C5: AB65Z7<5C; K6;7:9C6 =:6E89:F KH

GHK9CFC_67C<5 3C7G =8B7CAB: 2^@ 9%!"Q769D:7:F `8<9:E;:57 A9<K:E4 J#3"*=O 45%-"*6&O QR5 N^-bNL-

R2--+U4

2N4 h8996HS "4 &4 %( -=O @:6E<56B 65F EA67C6B I69C6KCBC7H <J K6;7:9C6B 65F 69;G6:6B 6EE:=KB6D:E C5 7G: ;<6E76B

367:9E 5:69 "5I:9E CEB65FS "5769;7C;64 D22=O N"$#&*"O E#5&*Q#*=O GA5 ./N/b./-/ R2--NU4

2-4 oB<q;Z5:9S d4 )4S d8;GES Y4 h4 n "=655S %4 Y6;7:9C<AB65Z7<5 ;<=A<EC7C<5E <J B6Z:E 65F <;:65E[ 6 W9E7

;<=A69CE<5 K6E:F <5 `8<9:E;:5;: #" '#(> GHK9CFC_67C<54 D22=O N"$#&*"O E#5&*Q#*=O GF5 fL.2bfL.^

R2---U4

.,4 @C=<5S h4S oB<q;Z5:9S d4 )4 n "=655S %4 MCJJ:9:57 ;<==85C7H E798;789: 65F 7:=A:96789: <A7C=6 <J

G:7:9<79<AGC; AC;<AB65Z7<5 C5 I69C<8E 9:DC<5E <J 7G: @<87G:95 );:654 DG>-(O E#5&*QO N5*=O >X5 .L/b

.N+ R2---U4

.24 !6D676S #4S d8Z8F6S O4S d8Z8F6S %4 n m<CZ:S a4 Y6;7:9C<AB65Z7<5 FCE79CK87C<5 65F A9<F8;7C<5 C5 F::A

V6;CW; 367:9E[ *69D:QE;6B: D:<D96AGC; I69C67C<5E 65F A<EECKB: ;<8ABC5D 3C7G EC5ZC5D A697C;B: `8g:E4

J#3"*=O 45%-"*6&O AF5 +.^b+f/ R.,,,U4

..4 e<897C:ES e4 %( -=O @=6BB:E7 :8Z69H<7C; <9D65CE=4 P-(>&% QMN5 .// R2--+U4

.f4 l3:CJ:BS $4 *4 nO6DE79<q=S "r 4 #<76B ;<857E <J=69C5: K6;7:9C6 C5;B8F: 6 B69D: J96;7C<5 <J 5<5Q58;B:<CFQ

;<576C5C5D K6;7:9C6 RDG<E7EU4 D22=O N"$#&*"O E#5&*Q#*=O G>5 .2N,b.2N/ R2--/U4

.+4 m695:9S h4 n d8G9=65S 04 M:7:9=C567C<5 <J 6;7CI: =69C5: K6;7:9C<AB65Z7<5[ 6 ;<=A69CE<5 <J

85CI:9E6B 2^@ 9%!" A9<K:ES 687<96FC<D96AGHS 65F 58;B:<CF E76C5C5D4 D22=O N"$#&*"O E#5&*Q#*=O GQ5

2.,Nb2.2f R2--LU4

./4 TCBBC6=ES V4 h4S e69B8;;CS "4 d4 n)BE<5S %4 " F::A A9<WB: <J E<=: KC<B<DC;6BBH C=A<97657 A9<A:97C:E C5

7G: ;:5796B !<97G V6;CW; DH9:4 45%-"*=O D5(- Q5 +L2b+L^ R2-N,U4

.^4 h:569FS O4 T4 n @=C7GS @4 h4 OHAE<=:79H <J <;:65 K6EC5 A9<IC5;:E4 SO T%*27)'O ;%'O M>5 +f,/b+f./

R2-^Û4

Y6::/$2$(,'-4 +(=%-2',+%( CE 6I6CB6KB: <5 P-(>&%PE T<9BFQTCF: T:K EC7:RG77A[kk333456789:4;<=U <9 6E A6A:9 ;<AH J9<= 7G: *<5F<5 :FC7<9C6B <JW;: <J P-(>&%4

=9>6/?5&0#&3&6-1

T: 7G65Z 7G: ;9:3 <J 7G: ;U E*-"- V-$%S *4 #8A6E 65F m4 Yj<q9Z=65 J<9 G:BA 3C7GE6=ABC5D4 #4 #6HB<9 G:BA:F E765F69FC_C5D A9<KC5D A9<7<;<BES 65F *4 d8jC:ZC A9<ICF:F;<=A8767C<56B E8AA<974 #GCE E78FH 36E E8AA<97:F KH !)""Q@:6D9657 )JW;: RhYmkMhmUS !@d RMhmUS 65F E8AA<97 J9<= 7G: M6ICF 65F *8;CB: V6;Z69F d<85F67C<5 R&dMU4#GCE CE @)&@# ;<579CK87C<5 58=K:9 /f2f 65F $@ 0o)d@ ;<579CK87C<5 58=K:9 ^+L4

e<99:EA<5F:5;: 65F 9:X8:E7E J<9 =67:9C6BE EG<8BF K: 6FF9:EE:F 7< h4Y4m4R:Q=6CB[ =Z695:9sE<:E74G636CC4:F8U4

$%&&%'( &) *+&,'%

!/" !"#$%& ' ()* +,- ' ./ 0"!$"%1 .,,2 ' 333456789:4;<=

44444444444444444444444444444444444444444444444444444444444444444D&.-+,0 E%*+'-),F. "8'& >+-$#)*/&-+-+), G&->&&, "&'%-+C&.H-8%"- !: I&.->< 6%"-J, K: 68""%J>< L%"'). !: 6%#$%()?<!.$'&+0$ H: K"+24,> B =(>%"( !''&, E&""&t

> H"'(#(>(% *+ K%==0 D"#3-= W :*2>=-(#*" F#*=*6)0 !"#$%&'#() *+ N<#"Q>&670N<#"Q>&67 N,? MSX0 !Y?1%2-&(3%"( *+ T%"%(#5'0 ;>(6%&' !"#$%&'#()0 :#'5-(-.-)0 P%. S%&'%) 9AALZ0 !CDt C3#(7'*"#-" X&*2#5-= ;%'%-&57 H"'(#(>(%0 D2-&(-<* B9IB0 F-=Q*-0;%2>Q=#5 *+ :-"-3-

!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!

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

,#$%-4F@>>[,#$ /$7$/ %= D0#,+(0 1$,I$$( 2'/$* *#%I* (% &%--$/'8,+%( I+,# ,#$ $*,+2',$9 -$/',$9($** %= +(,$-'&,+(0 2'/$*5 16, +*($0',+7$/4 &%--$/',$9 I+,# =6,6-$ 2',+(0 %::%-,6(+,+$*!O6=CB7<5PE 98B:2S. A9<ICF:E 6 7<<B J<9 85F:9E765FC5D 6 965D: <J

E<;C6B C57:96;7C<5ES C5;B8FC5D 6B798CE=S 6DD9:EEC<5S E:BWEG5:EE 65FEAC7:4 a7 E767:E 7G67 6B798CE= R<9 B:EE 6DD9:EEC<5U CE J6I<89:F 3G:5&Q " 5 ! ,S 3G:9: 5 CE 7G: W75:EE ;<E7 7< 7G: 6B798CE7S Q CE 7G: W75:EEK:5:W7 7< 7G: K:5:W;C69H 65F & CE 7G:C9 D:5:7C; 9:B67:F5:EE4 d<9 6DCI:5 K:5:W7 65F ;<E7S 7G: :I<B87C<5 <J 6B798CE= 7G:9:J<9: 9:BC:E8A<5 6 E8JW;C:57BH GCDG 9:B67:F5:EE K:73::5 C57:96;7C5D C5FCICF86BE4O6=CB7<5. <9CDC56BBH E8DD:E7:F 7G67 6 GCDG 9:B67:F5:EE ;<8BF 69CE: C573< 36HE[ R2U K:G6IC<89 K6E:F 8A<5 FC9:;7 ZC5 9:;<D5C7C<5 K:73::5C5FCICF86BES <9 R.U BC=C7:F FCEA:9E6B RA<A8B67C<5 ICE;<EC7HU4O<3:I:9S 7G: C=A<9765;: <J BC=C7:F FCEA:9E6B C5 C5;9:6EC5D 7G:

9:B67:F5:EE 6=<5D C57:96;7C5D C5FCICF86BE 65F J6I<89C5D 6B798CE=G6E K::5 ;<579<I:9EC6Bfb224 O6=CB7<5PE <9CDC56B E8DD:E7C<5 G6E K::5;<57:E7:F K:;68E: BC=C7:F FCEA:9E6B ;65 6BE< C5;9:6E: ;<=A:7C7C<5K:73::5 5:CDGK<89C5D 9:B67CI:ES 3GC;G <AA<E:E 7G: :I<B87C<5 <J6B798CE7C; K:G6IC<89fb224 $5J<978567:BHS :=AC9C;6B 7:E7E <J 7G:<9HS7G67 F:7:9=C5: 7G: 9:B67CI: C=A<9765;: <J C5;9:6E:E C5 K<7G 9:B67:FQ5:EE 65F ;<=A:7C7C<5 K:73::5 9:B67CI:ES G6I: K::5 GC5F:9:F K:;68E:K<7G J6;7<9E 69: C5`8:5;:F KH FCEA:9E6BS 65F E< 7G:C9 :JJ:;7E 69:8E86BBH ;<5J<85F:F+SNb2,4#G: I69C6KB: J<9= <J =67: ;<=A:7C7C<5 65F A<A8B67C<5 E798;789:

6;9<EE WD 36EA EA:;C:E 3C7G3C5DB:EE =6B:E <JJ:9E 65 <AA<9785C7H J<9FCE:5765DBC5D 7G: ;<5J<85F:F :JJ:;7E <J 9:B67:F5:EE 65F ;<=A:7CQ7C<5 K:73::5 9:B67CI:E C5 ICE;<8E A<A8B67C<5E2.b2N4 dCD 36EAE 69:EA:;C:E 7G67 F:I:B<A 3C7GC5 7G: J98C7 <J WD 79::ES 65F C5;B8F:=8786BCE7C; A<BBC567C5D EA:;C:E 6E 3:BB 6E A696EC7C; 5<5QA<BBC567C5DEA:;C:E2/4 a5 =65H EA:;C:E 7G: =6B:E 69: 3C5DB:EES 65F =67: 3C7G 7G:3C5D:F J:=6B:E K:J<9: 7G: J:=6B:E FCEA:9E:4 #G: B:I:B <J 6DD9:EEC<5K:73::5 7G:E: 5<5QFCEA:9EC5D =6B:E I69C:E :5<9=<8EBH 6;9<EEEA:;C:E2.b2/S2N4 "7 <5: :g79:=:S =6B:E <J E<=: 5<5QA<BBC567C5DEA:;C:E 69: GCDGBH =<FCW:F J<9 ;<=K67 3C7G 69=<89:F K<FC:E 65FG8D: =65FCKB:E4 #G:E: =65FCKB:E 69: 8E:F 7< 7:69 E<J7 7CEE8: 65FE:I:9 K<FH A697ES C5;B8FC5D BC=KES G:6F 65F 6KF<=:5S 65F ;659:E8B7 C5 :g79:=:BH GCDG=<976BC7H B:I:BE4 "7 7G: <7G:9 :g79:=:S =6B:E<J <7G:9 5<5QA<BBC567C5D 65F =<E7 A<BBC567C5D EA:;C:E EG<3 5<=<FCW;67C<5E J<9 ;<=K67 <9 6DD9:EEC<54";9<EE 7G:E: EA:;C:ES 7G: 6I:96D: 9:B67:F5:EE <J ;<=A:7C5D =6B:E

I69C:E :5<9=<8EBH <3C5D 7< I69C67C<5 C5 7G: 58=K:9 <J J:=6B:E 7G67B6H :DDE C5 :6;G J98C72.b2^4 d<9 :g6=AB:S CJ <5BH <5: J:=6B: B6HE :DDE C56 J98C7 7G:5 6BB 7G: ;<=A:7C5D =6B:E 3CBB K: K9<7G:9E] C5;9:6EC5D58=K:9E <J J:=6B:E B6HC5D :DDE C5 6 J98C7 3CBB B:6F 7< =6B:E

–2

–1

0

1

2

Mea

n in

jury

leve

l (LE

I) co

ntra

sts

0.00 0.05 0.10 0.15 0.20 0.25 0.30

Relatedness contrasts

!"#$%& ' L+,# -#T%&7 (+.+( 1"#$&,)$) '("$$+0 ,4,-#)$ +)$-8,$+0 &+(,$+0#+)) 1"#$&,)$)C>1&")) )'+1-+)= $3+ 8+,# -#T%&7 (+.+( P(-*+$-8+ +U$+#$ "* -#T%&7= ?NFR ,#0 $3+ '&"'"&$-"# "*

-#0-.-0%,() )+.+&+(7 -#T%&+0 PKFR )3"2+0 #" )-4#-B1,#$ &+(,$-"#)3-' 2-$3 +)$-8,$+0

&+(,$+0#+)) P?NFV ,(( 1"#$&,)$)= & P5=5WR X 5CY5= ' Z X YCY[= ( X YC\\O #"$ -#1(%0-#4

1"#$&,)$) 2-$3-# $3+ '"((-#,$"& (-#+,4+= & P5=55R X YC]Z= ' Z X YCY[= ) X YCMZO KFV ,((

1"#$&,)$)= & P5=5WR X YCYM= ' Z $ YCY5= ( X YC^MO #"$ -#1(%0-#4 1"#$&,)$) 2-$3-# $3+

'"((-#,$"& (-#+,4+= & P5=55R X YCYW= ' Z $ YCY5= ( X YC^\RC !-&1(+)= 1"#$&,)$) /+$2++# $3+

#"#6'"((-#,$-#4 )'+1-+)O )Q%,&+)= 1"#$&,)$) /+$2++# $3+ '"((-#,$"& )'+1-+)O $&-,#4(+= $3+

1"#$&,)$ /+$2++# '"((-#,$"&) ,#0 #"#6'"((-#,$"&)C


Documents

J409 25d 507....44444444444444444444444444444444444444444444444444444444444444444!!!!! +*2 F@>>. –1 0