Polychaetes Species Diversity Applying
J10V`1 7 Q` :]CV >VRV`1HQ ? .88 H.QQC > :` . H1VJHV? H1VJHV :JR J$1JVV`1J$ Q` .V V:
!"# $ ! &$ # ' ( $ & ' #$# & #$# $&)$ 5
! ' #$ 5 & ( $# # ( $
.88 %RVJ % Q` ):`G:`: : Q` : `Q`8 1JHVJ .1 `VV:`H. .: GVVJ H:``1VR Q% 1J HQCC:GQ`: 1QJ 11 . VJ01`QJIVJ :C IQJ1 Q`1J$ ]`Q=VH %]]Q` VR G7 .V V$1QJV :<1Q CQH:C :% .Q`1 7 :JR H:``1VR Q% G7 :C1:J : 1QJ:C J 1 % V `Q` "J01`QJIVJ :C`Q VH 1QJ:JR VV:`H. ^ $_8 &Q` $ I7 :H@JQ1CVR$VIVJ $Q Q %1: 1HQCV 15`VV:`H.V`1JH.:`$VQ` .VV]`Q=VH 5:JR Q I71Q`@$`Q%]5`VV:`H.V`:JR %RVJ 5`Q` .V1`:1 :JHV1J`1VCR:H 101 1V:JR`Q` .V1` 1IV :JR 1J0:C%:GCV RVR1H: 1QJ Q :6QJQI1H 1RVJ 1`1H: 1QJ7 :QC: : :CCV5 :J1VC::$:JVCC15$C``VRQ:<<1J18]VH1:C .:J@$Q QI7HQCCV:$%V :``:VCV`Q1V 15 `Q` .1 VVJ 1:C %]]Q` 1J $VQ : 1 1H5 :JR Q`V : : :J<1 ^,%` :R7 Q` :GCV_8 &%` .V`IQ`V5 1Q%CRC1@V Q .:J@ $;VR1IVJ :GQ`: Q`7`Q`VR1IVJ R: : :J:C7V8 &Q`.H.QQCQ`J10V`1 7Q` :]CV>&VRV`1HQ ?I7:H@JQ1CVR$VIVJ $QV QI7 % Q``Q`81JHVJ 8 ! $& !$ 83QJ V6 R .V"`:Q`<1QR10V`1 7 8<1QR10V`1 7Q`2VR1 V``:JV:JV:7Q` RV:GVR I:H`Q 8 "J01`QJIVJ :C0:`1:GCV7%``:HVVR1IVJ 8QC7H.:V V:VIGC:$V7RVH`1] 10V:JRI%C 10:`1: V:J:C7V 882, $ , 3$ , 88 , $ $ 3$ 88$<$ $ 88 " $3 $ 88 4$" $ 8 : 1 1H:C:J:C7V 8]VH1V"HQCQ$1H:C:J:C71 8 ! ! 8 ! ! ! # $ / & & ! # # # # # # # # # 8 ! ,3 , 8 !QJ V6 T $.V `: Q` )1QR10V`1 7 J 1QRVE:JV1`Q53QJ0VJ 1QJQJ<1QCQ$1H:C10V`1 7^ _:$`VVR .: 1V:`VCV:R1J$ ]VH1V Q V6 1JH 1QJ :JR .:G1 : RV `%H 1QJ5 :JR .: QIV IV:%`V .:R Q GV :@VJ8 $` V` .V 1Q 3QJ0VJ 1QJ .V V6]CQ`: 1QJ Q` G1QR10V`1 7 GVH:IV IQ`V 1I]V`: 10V .:JV0V`51JHV1VH:JJQ RV`VJRQ`I:J:$VQIV .1J$1`1VRQJ; @JQ1 1 ^ 1 I:`1JVVHQ7 VIC:$`:`GV.1JR^ JVC$`Q0V V :C8 5 64`:75 666V1] V :C8 5 _:JR .V`V`Q`V .VJVVR`Q`IQ`V`VV:`H.1J .VI:`1JVVJ01`QJIVJ 1$`V: 8 .V1V:C .Q`G1QR10V`1 7Q`I:`1JVVJ01`QJIVJ 1:HQJV_%VJHVQ`I1CC1QJQ`7V:` Q`V0QC% 1QJ:`7.1 Q`78 .1VJ01`QJIVJ .::0V`7.1$.G1QR10V`1 7GVH:%VQ% Q` .VRVH`1GVR:J1I:C].7C::`V`V]`VVJ VR1J .V`V5:JR ].7C::`VV6HC%10VC7 I:`1JV^ Q`V5 _83QJV_%VJ C75I:`1JVQ`$:J1IR1]C:7:I%H.C:`$V`].7CV 1H R10V`1 7 .:J .QVQJC:JR^ :75 _8 2 `V_%1`1J$ RV1$J: 1QJ Q` ]VH1:C $`V: Q` 3QJV`0: 1 QJ5 ]VH1V 1J JVVR Q` `1H ]`Q VH 1QJ5]VH1V1.QV :@1J$``QI .V11CRH:JGV`V `1H VRG7"%`Q]V:JC:1_8 .V R1`VH 10VCVR Q .VV 1J$%]Q`:JV 1Q`@Q`]VH1:C$`V:Q`3QJV`0: 1QJ51.1H. Q$V .V`11 . .VV61 1J$]VH1:C`Q VH 1QJ$`V:`Q`I:JV 1Q`@Q`]`Q VH VR1 V :H`Q .V"H:CCVR : %`:8$ `V$1QJ:CG:1JCV0VC5QIV7V:`GV`Q`V51J 5 .V<:`HVCQJ:3QJ0VJ 1QJ L L""35:IVJRVR1J 5:JR .V`Q QHQCR`:1J%] 1J C1JV 11 . .1 3QJ0VJ 1QJ5 :1I Q ]`Q VH :JR 1I]`Q0V .V I:`1JV :JR HQ: :C VJ01`QJIVJ 1J .V 2VR1 V``:JV:J5 1.1C ]`QIQ 1J$ `V$1QJ:C :JR J: 1QJ:C ]C:J HQJ `1G% 1J$ Q% :1J:GCVRV0VCQ]IVJ 8 8 )1QR10V`1 7 Q` (VR1 V``:JV:J V:7 Q` RV:GVR I:H`Q .V 2VR1 V``:JV:J V: 1 : I:`1JV G1QR10V`1 7 .Q ]Q 8 3QCC V :C8 ^ _ C1 VR :]]`Q61I: VC7 5I:`1JV]VH1VQHH%``1J$1J .V2VR1 V``:JV:JV::JR <1:JH.1 :JR 2Q``1 ^ _ `V]Q` VR .: 8 Q Q` .V 1Q`CR; Q :C RVH`1GVR 1J0V` VG`: V ]VH1V :`V ]`VVJ 1J .V 2VR1 V``:JV:J :`V:8 61$. ]V`HVJ :$V Q` 2VR1 V``:JV:J I:`1JV]VH1V:`VVJRVI1H^ 3 $IQJ$ I:`1JV ]VH1V5 Q` RV:GVR I:H`Q :C1:J Q` RV:GVR I:H`Q 4 $IQJ$ .VV %R1V5 : J%IGV` Q` `VV:`H.V QJ I:H`Q 5 QHV:J1H: IV:RQ15 :GQ0V .V CQ1V` C1I1 5 HQ: :C RV `1 1H Q` I%RR7 RV `1 1H G1QHQVJQV 1V`V `Q%JR8 ,``.Q`V Q .V V0V`V `10V` IQ% . .V G1QHQVJQ1 Q` HQ: :C V``1$VJQ%I%RQHH%``VR:C`V:R7: IQ`RV] .8$CQJ$ : 1%IHQJ 1JVJ :C .VC` .1HQVJQ11:I:1JC7`Q%JR:]]`Q61I: VC7``QI I51.1CV1J:J%IGV`Q` :`V:5GV 1VVJ:JR IQ`RV] .5:VIGC:$V 7]1H:CQ`I16VRQ`I%RR7RV `1 1H VR1IVJ 1V`V 1RVJ 1`1VR8 .1 `VV:`H.5 H:``1VR Q% : `V$1QJ:C H:CV5 HQJ`1`IVR Q` %]R: VR`V%C Q`]`V01Q% %R1V`V:C1 6 `%H %`V:JR1 ]VH1VHQI]Q1 1QJH:JGV%VGQ . Q:V .V_%:C1 7Q`GVJ .1H .:G1 : :JR:>1JR1H: Q`?Q`I:`1JVVJ01`QJIVJ :CH.:J$V5`V%C 1J$``QI.%I:J :H 101 1V : 1VCC : HC1I: 1H H.:J$V ^ 4`:75 6 3C:`@V :JR $1J1Q` .5 6 6:`@:J `: :JR QR`1$%V<5 6 =`V V :C8 5 _8 JRVVR5 GVJ .1H ]VH1V :`V $VJV`:CC7VRVJ :`7Q`.:0V:CQ1IQ 1C1 7:JR .V`V`Q`VIQ C1@VC7 Q`V]QJR QCQH:C VJ01`QJIVJ :CHQJR1 1QJ5HQ0V`:11RV`:J$VQ``VVR1J$$%1CR5.:0V`VC: 10VC7CQJ$ :JRR1``V`VJ C1`VH7HCV^I:J7GVJ .1HQ`$:J1I.:0V .V1`C:`0:C :$V:]:` Q` .V ]VC:$1HHQI]:` IVJ _:JR .V`V`Q`V$10V:J1J V$`: VR`V]QJVQ0V` 1IV Q0:`1: 1QJ Q` 1: V` :JR VR1IVJ _%:C1 75 V6.1G1 R1``V`VJ RV$`VV Q` QCV`:JHV Q `V HQJR1 1QJ5^ :%V`5 63C:`@V:JR:`11H@5 64`:75 5_82Q`VQ0V`51J I:`1JV VJ01`QJIVJ .V GVJ .1H `:%J: 1 HQJ1RV`VR : @V7 $`Q%] 1J .V %JRV` :JR1J$ Q` .V G1QCQ$1H:C ]`QHVV :JR HQII%J1 7 `%H %`V ^ 4:IG1 V :C8 5 6 JVC$`Q0V V :C8 64`:75 _8 $IQJ$ .VIQ 1I]Q` :J IQRVC5]VH1VR$G%JR:JHVR<1QI:IQRVC^ V:`QJ:JR QVJGV`$5 _1: RV0VCQ]VR Q:J:C7 7 QJ 1 V :C8 5 6 3QCQ1Q V :C8 5 _Q``.Q`V]C: `Q`I:JRR`1CC1J$I%R^ 3`VI: V :C85 5 6 `:G%HHQ V :C8 5 6 V`C1<<1 V :C8 5 6 2:JQ%@1:J V :C8 5 _5 HQ: :C RV`VJHV `%H %`V ^ 1HQCV 1 V :C8 5 6 8 `Q%JR QRQI1J: V: `1` 8 1:CQ1VCC@JQ1J . : I:J7Q]]Q` %J1 1H]QC7H.:V V :`V :HQI]CV6Q`1GC1J$]VH1V^ 4`:CV:JR4`:CV5 _5`V=VH 1J$ .VHC:1I .: I:J7Q`%H.]VH1V:`V::HQIQ]QC1 :J]VH1V;;8 <1:JH.1:JR2Q``1^_ `V]Q` VR .: 8 QQ` .V1Q`CR; Q :CRVH`1GVR]QC7H.:V V ]VH1V:`V]`VVJ 1J .V2VR1 V``:JV:J:`V:6 .1]V`HVJ :$VVVI QGVHCQV Q `V:C1 75 1JHV `V1 2VR1 V``:JV:J ]QC7H.:V V :`V GV1J$ RVH`1GVR .VV R:75 1.1CV .V`V :`V I:J7 IQ`V ]VH1V Q GV RVH`1GVR :`Q%JR .V 1Q`CR8 .V`V .:0V GVVJ J%IV`Q% %R1V1J0V 1$: 1J$ .V7 VI: 1H5 :6QJ QI75IQ`].QCQ$7:JRVHQCQ$7Q` 2VR1 V``:JV:J ]QC7H.:V V G% : $VJV`:C :HHQ%J QJ .V VHQCQ$7 Q` .V 2VR1 V``:JV:J ]VH1V5 H:``1VR Q% : : C:`$V H:CV5 R: V G:H@ Q 9 41:J$`:JRV ^ _ :J:C7 10 8 (:1J 1%V 6Q1I:J7]VH1VC10V1JRV`1JVR:`V:Q`.:G1 : -.1H.1 .V1`]: 1:CR1 `1G% 1QJ- $HHQ`R1J$ Q .V 1JJV:J^ <`Q1J:JR QIQC1JQ _:JR:CC:HV:J^ QIQC1JQ5 _ .Q` `:CC5H1VJ 1`1H`VV:`H. 1CCR1]C:7:C:H@Q` :6QJ QI1H:JR]VH1VR1 `1G% 1QJ R: :8 .V`V`Q`V5 Q0V`:CC %R1V Q` :6QJ QI75 VHQCQ$7 :JR ]: 1:C R1 `1G% 1QJ Q` I:`1JVGVJ .1H]VH1V:`V 1CC1JHQI]CV V^V8$8`Q` 2VR1 V``:JV:J V:7 X`W :JR 1H:`R5 6 $%$1V`5 6 1VRC5 6 2:`$:CV` 5 6 <1:JH.1 V :C8 5 :5G5H6 3`VI: V :C8 5 6=Q%@Q%`: V :C8 5 63Q VCCQ V :C8 56 $`0:J1 1R1 V :C8 5 6 VC1J1 5 3QVJ 1JQ :JR 41:HQGGV _8 J ]:` 1H%C:`5 .V`V 1 QJC7 : $VJV`:C :HHQ%J 5R: 1J$G:H@ Q 11 I:`1JV VJ01`QJIVJ H:%V V0V`V R1 %`G:JHV 1J I:H` Q 12 HQ%]CVQ`1JR1HV0:C%V .VR10V`1 7R1 `1G% 1QJ Q`:JVJ 1`V:`V:CV:R Q:$`V: CQ Q`1J`Q`I: 1QJ8$R10V`1 71JR1HV:`V]: 1:CC7 `%H %`VR5 .Q11J$ :% QHQ``VC: 1QJ ^1I1C:` QIQ G1QCQ$1H:CLVHQCQ$1H:CIV:%`V_5$VQ : 1 1H]`Q01RV:0:C%:GCV QQC `Q`:`%CC7RVH`1] 10V:J:C71Q0V`:CC .V1J0V 1$: VR:`V:8$RV :1CVRI:]Q`R10V`1 7 I:@V1 ]Q1GCV QRVC1I1 :JRR1 1J$%1.:`V:11 .$`V: V`R10V`1 7 .`Q%$.Q% .V ]:HV ^R10V`1 7 .Q ]Q _ ``QI :`V: 11 . : CQ1 ]VH1V R10V`1 78 3QJV_%VJ C75 I:]]1J$ %1J$ @`1$1J$ 1J V`]QC: Q` :CCQ1 Q RV V`I1JV 1.V .V` $CQG:C `VJR :`V QGV`0VRCQH:CC7^ %`1JQ V :C8 5 _8$7V 5$VQ : 1 1H:C:]]C1H: 1QJ QR10V`1 7:`V H:`HVR%V Q .V.V V`Q$VJV1 7Q` Q .V]:%H1 7Q` R: : : 1VCC .V 1JHQJ1 VJH7 GV 1VVJR: ::JRJ: %`:C]: V`J^ 2:7JQ% V :C8 5 6=1VJVC V :C8 561J1 13 QR1]C:7 .V0:`1:G1C1 7Q`]QC7H.:V VR10V`1 7 ]: V`J1J]:HV:JR 1IV5:JR Q 1J0V 1$: V .Q1 .1 0:`1:G1C1 7 1 `VC: VR Q .V .:G1 : H.:`:H V`1 1H :JR .%I:J 1I]:H 5 .1`VV:`H.51:1IVR Q7 _1RVJ 1`7R10V`1 7R1 `1G% 1QJ]: V`JQ`]QC7H.:V V:VIGC:$V:JR]VH1V: : `V$1QJ:C^ : 1%I3QJ 1JVJ :C.VC`_`VC: VR QRV] .5VR1IVJ $`:1J1 14 8 (#$" &# # ( $" 8 %R7 1 V : :HQCCVH 1QJ1:H:``1VRQ% 1J`10V:`V:Q` : 1%IHQJ 1JVJ :C.VC`H:CCVR2:H`QR :`V: ^ &1$8 88 _6 .V V$1QJ Q` :<1Q CQH:C :% .Q`1 7 VCVH VR .VV :`V: GVH:%V 11 .1J .VI `VC1H :JR RV]Q1 5 ]Q VJ 1:CC7 V6]CQ1 :GCV `Q` GV:H.V JQ%`1.IVJ 5 1V`V `Q%JR8 $J HQI]`V.VJ10V `VV:`H. Q 1RVJ 1`7 :JR H.:`:H V`1V `VC1H :JR RV]Q1 :CQJ$ : 1%IHQJ 1JVJ :C.VC`1:H:``1VRQ% G7 3.1QHH1:JR :2QJ1H: 1J 8 1$%`V 88 T %R7:`V::CQJ$ : 1%IHQJ 1JVJ :C.VC`^ 7``.VJ1:JV:_8&`QI .VJQ` .V`J Q .VQ% .V`J:`V:72QJ :C QR13: `Q5 Q`0:1:J1H:5:G:%R1:5 V``:H1J::JR4:V :8 15 8 : :V &`QI %] Q : V`1V Q` V: 1: V` HQC%IJ :JR V:GVR :I]C1J$ 1V`V HQJR%H VR G7 :C1:J : 1QJ:C J 1 % V `Q` "J01`QJIVJ :C `Q VH 1QJ :JR VV:`H. ^ $_ :CQJ$ .V : 1%I HQJ 1JVJ :C .VC` :1IVR : H.:`:H V`1<1J$ :JR IQJ1 Q`1J$ `VC1H :JR RV]Q1 5 ]Q VJ 1:CC7 V6]CQ1 :GCV `Q` GV:H.V JQ%`1.IVJ 8 %`0V7 1V`V ]C:JJVR : I:H`QRH:CV T VJ01`QJIVJ :C H.:`:H V`1<: 1QJ R :JR CQH:CRH:CV T IQJ1 Q`1J$ Q` R`VR$1J$ :`V: ^ 1HQCV 1 V :C8 _8 : : ``QI .V I:H`QR:`V: CQH: VR:CQJ$ .V : 1%IHQJ 1JVJ :C.VC`1V`VV6 `:H VR``QI $; R: :V QJ .V G:V Q` : HQIIQJ ]`QHVR%`V Q` :I]C1J$ :JR R: : HQCCVH 1QJ :HHQ`R1J$ Q IV .QRQCQ$1V]`Q]QVRG7 3: VCC1 V :C8 ^ _8 : :QJ .VR1 `1G% 1QJQ` Q1RQJ1: QHV:J1H: V:$`::CQJ$ .VHQ: Q` : 1%I1V`VV6 `:H VR``QI .V`QCCQ11J$1Q`@7 $`R1< V:1: V`:RR1J$Q3:3, RG%``V`VR`Q`I:C1J^ &1$888 _8 %``:HVVR1IVJ :I]CV1V`VHQCCVH VR: V:H. : 1QJ11 .:GQ6RHQ`V`^&1$%`V00_8 &Q`V:H.:I]CVR : 1QJ5RV] .:JR$VQ$`:].1H:C]Q1 1QJ ^ 2 ]`Q=VH 1QJ5 R: %I 4 5 `Q`R: :Q`2:H`Q:`V:Q`2QJ :C QR13: `Q:JR `Q` .V`VI:1J1J$ I:H`QR:`V:_1V`V`VHQ`RVR8 :I]C1J$RV1$J11 .J%IGV`Q` : 1QJ5 : 1QJHQRV ^:`V: :H`QJ7I5 `:JVH :JR RV] ._Q`V:H.:`V:1V`V`V]Q` VR J &1$%`V88588588 588 588 8 16 1$%`V 88 T $ L$ `V: :JR:JVVJ$`:G R%`1J$::I]C1J$8 1$%`V 88 T 1``V`VJ ].:V Q` I:H`Q 1:1V0VRG7RIIIV.:JR]`VV`0VR1J V:1: V`:RR1J$Q3:3, RG%``V`VR`Q`I:C1J8 17 $:GCV 88 T:I]C1J$RV1$J`Q`V:H. %R7:`V:8 (#!& R (#!& R#& # (:H`Q # :G:%R1: &VG`%:`7:JR2:`H. 2:`H.:JR$]`1C $ $V``:H1J: 2:7 2:7 '$ ':V : E:J%:`7 &VG`%:`7 18 1$%`V 88 T:I]C1J$]C:JQ` .V %R7:`V:Q`2QJ :C QR13: `Q7 :I]C1J$ : 1QJ8 19 1$%`V 88 T:I]C1J$]C:JQ` .V %R7:`V:Q` Q`0:1:J1H:7 :I]C1J$ : 1QJ8 20 1$%`V 88 T:I]C1J$]C:JQ` .V %R7:`V:Q`:G:%R1:7 :I]C1J$ : 1QJ 8 21 1$%`V 88 T:I]C1J$]C:JQ` .V %R7:`V:Q` V``:H1J:7 :I]C1J$ : 1QJ 8 22 1$%`V 88 T:I]C1J$]C:JQ` .V %R7:`V:Q`4:V :7 :I]C1J$ : 1QJ8 23 8 :GQ`: Q`7 :J:C7V 2:H`Q %``:HV VR1IVJ :I]CV 1V`V ]`V01Q%C7 1:.VR 11 . : QC% 1QJ Q` 6 , :JR R1 1CCVR1: V`:JR .VJR`1VR QV]:`: V .V`1JV``:H 1QJ``QI .VHQ:`V`8$J:C7V Q` .VHQ:`V```:H 1QJ^] -I_1V`V]V``Q`IVR 11 .$ 21V0V1JV`1V 11 .: `:J$VQ`8 ].151.1CV5 .V`1JV``:H 1QJ^\ -I_%1J$ :JR`:7VR1$`:`^:` 1HCV1 24 8 : : :J:C7V J01`QIVJ :C 0:`1:GCV VR1IVJ `VCV0:J : 1 1H:C]:`:IV V`1V`VH:CH%C: VR:HHQ`R1J$ Q &QC@:JR:`R ^ _82VR1:J1HQJ1RV`VR:$QQRRVH`1] Q`Q`VJ01`QJIVJ :CH.:`:H V`1 1H:JR1 1 :J 1I]Q` :J ]:`:IV V` Q V6]C:1J .V R1 `1G% 1QJ Q` Q` RV:GVR GVJ .1H :VIGC:$V^ V``:1CC1V V :C8 5 _8 2Q`VQ0V`51JV:H. : 1QJ%``:HVVR1IVJ 1V`V :J:C7VR :JR HC:1`1VR :HHQ`R1J$ Q VR1IVJ V6 %`V ]V`HVJ :$V5 `QCCQ11J$ Q : ^ _7:JR^_^] QHQJ VJ Q`:JR_5I%RR7:JR^I_^ T Q_50V`7:JR7I%R ^02_^ TQ_5:JR7I%R^2_^T Q_:JRI%R^2_^\ Q_8 JQ`RV` QI:J:$V11 . .V :IQ%J Q` R: : :JR Q `:H1C1 : V .V 1J V`]`V : 1QJ Q` `V%C 5 .V R: : 1V`V HC:1`1VR1J Q16G: .7IV `1H$`Q%]8 .% .V`1` $`Q%]Q` : 1QJHQ``V]QJR Q .V %]]V`1J``:C1 Q`:C .:G1 : R10V`1 7 861CCR10V`1 7J%IGV` 5 5 5 1J`1V`VH:CH%C: VR1JV:H.Q` .V:I]CVR : 1QJ:%J10:`1: VIV:%`VQ`:C].:R10V`1 7: V:H. : 1QJ8$V:H. R10V`1 7J%IGV`1Q`:R1``V`VJ Q`RV`:JR$10VR1``V`VJ 1V1$. Q .VCVHQIIQJ ]VH1V5 .V1`%VH:J`V0V:CR1``V`VJ :]VH Q`:HQII%J1 78 1V1$.V_%:CC7:CC ]VH1V `V$:`RCV Q` .V1` :G%JR:JHV :JR .V`V`Q`V V_%:C .V J%IGV` Q` ]VH1V6 51.1H. $10V CV 1V1$. Q `:`V ]VH1V .:J 5 1 V_%10:CVJ Q .V CQ$V V6]QJVJ 1:C `Q`I Q` .V .:JJQJT1VJV` 1JRV66 1 .V `VH1]`QH:C Q` 1I]QJ 1JRV6 :JR ]C:HV IQ`V1V1$. QJ HQIIQJ ]VH1V6 :JR 1J`85 .V `VH1]`QH:C Q` .V ]`Q]Q` 1QJ:C :G%JR:JHV Q` .V HQIIQJV ]VH1V ^ .V `VH1]`QH:C Q` .V 25 IV .QRQCQ$7 ^:%01J V :C8 5 _6`:`V`:H 1QJHQI]VJ: V`Q`R1``V`VJ :I]CV1 26 R10V`1 7 H:J GV G`Q:RC7 RV`1JVR : .V 0:`1:G1C1 7 1J ]VH1V HQI]Q1 1QJ :IQJ$ :I]C1J$ %J1 `Q` : $10VJ :`V: : : $10VJ ]: 1:C H:CV8 " 2 QJ .V G:1 Q` VHQCQ$1H:C IV:%`V Q` HQI]Q1 1QJ:C R11I1C:`1 7 H:J GV %VR Q V R1``V`VJHV :IQJ$$`Q%]1JGV :R10V`1 78 .1:J:C711:]V``Q`IVRQJ .VG:1Q`E:HH:`R IV:%`V 1.1H. 1 R1`VH C7 1J V`]`V :GCV : : ]V`HVJ :$V Q` %J.:`VR ]VH1V6 .%5 ]VH1V HQI]Q1 1QJ ^]`VVJHVL:GVJHV_ 1J`Q`I: 1QJ 1J HQJ=%JH 1QJ 11 . .1 `VVIGC:JHVIV:%`V V .V.QIQ$VJV1 7Q`I%C 10:`1: VR1]V`1QJ::IV:%`VQ` GV :R10V`1 7^ $JRV`QJ V :C8 5 _8 .V `VC: 1QJ.1] GV 1VVJ HQII%J1 7 `%H %`V :JR VJ01`QJIVJ :C 0:`1:GCV 1: V6:I1JVR%1J$ .V< ," `Q% 1JV%1J$ .V<`:7T3%` 11I1C:`1 7HQV``1H1VJ ^ 3C:`@V :JR $1J1Q` .5 _8 < ," %V : ]V:`I:J; `:J@ HQ``VC: 1QJ Q V `Q` 1$J1`1H:J HQ``VC: 1QJ GV 1VVJ 1I1C:`1 7 I: `1HV Q` ]QC7H.:V V:G%JR:JHV :JRQ` VJ01`QJIVJ :C 0:`1:GCV8 : 1 %RV5 CQJ$1 %RV5 RV] . :JR VR1IVJ HQI]Q1 1QJ ^]V`HVJ :$V1C THC:7``:H 1QJ\8 II_1V`V .V:G1Q 1H0:`1:GCV:J:C7VR8 .VV :J:C711:%JRV` :@VJ`Q`:CC`10V:`V: Q$V .V`8 :1`11V]V:`I:J;`:J@HQ``VC: 1QJ1:]V``Q`IVRGV 1VVJVJ01`QJIVJ :C0:`1:GCV ^ : 1 %RV5 CQJ$1 %RV5 RV] . :JR VR1IVJ HQI]Q1 1QJ_ :JR G1Q 1H R10V`1 7 `Q` .V 1.QCV %R1VR:`V:8J10:`1: V:JRI%C 10:`1: V:J:C7V1V`V]V``Q`IVR%1J$ .V Q` 1:`V]:H@:$V`1IV`08 8:JR " 2$ ,$ UQ` 1:`V^ $JRV`QJ V :C8 5 _:JR $CVQJ QCQ$1H:C : 1 1H]:H@:$VT$ 0888 ]VH1V VHQCQ$7 :J:C71 JR1H: Q`]VH1V$J:C71^ $_^ %``XJV:JR V$VJR`V8 _.:GVVJ]V``Q`IVR1J Q`RV` Q 1RVJ 1`7 .V ]VH1V :QH1: VR 11 . Q` 1JR1H: 10V Q` `10V $`Q%] Q` : 1QJ RV`10VR``QI Q : ^ _HC:1`1H: 1QJ5:JR16$`Q%]Q` : 1QJRV`10VR``QIRV] . `:J$VHC:1`1H: 1QJ8 .1:J:C71HQIG1JV .V`VC: 10V:G%JR:JHVQ` .V]VH1V11 . .V1``VC: 10V``V_%VJH7Q`QHH%``VJHV1J .V0:`1Q%$`Q%]:JR]`Q01RV:J1JR1H: Q` 0:C%V%GV_%VJ C7 V VRG7`:JRQI1<: 1QJ8 ]: 1:C R10V`1 7 ]`VR1H 1QJ I:] `VR1H 1QJI:]Q`QIV]QC7H.:V V]VH1VRVJ1 7 :JRHQII%J1 71JR1HV^"5 :JRE;_1V`V`V:C1 27 ``QI]`VC1I1J:`7:J:C71HQJR%H VR11 . .VG:1H : 1 1H.1$.C1$. VR .VJVVR Q ]`QHV .V R: : V]:`: VC7 `Q` V:H. I:H`QR:`V:8 .V : 1 1H:C IV .QR Q` ]: 1:C 1J V`]QC: 1QJ ^$VQ : 1 1H_ :`V %VR Q V 1I: V 0:`1:GCV R1 `1G% 1QJ5 V0VJ 1` .V7 .:0VJQ GVVJIV:%`VR5 .`Q%$. .V%VQ`I: .VI: 1H:C`VC: 1QJ.1]8 28 8 & $ 8 J01`QJIVJ :C 0:`1:GCV7 %``:HV VR1IVJ %``:HVVR1IVJ Q`V:H. : 1QJ1V`VHC:1`1VR:HHQ`R1J$ Q Q : ^ _ :JR .V R1 `1G% 1QJQ`VR1IVJ HC:V:CQJ$ .V : 1%IHQJ 1JVJ :C.VC`1:RVH`1GVR1J &1$%`V88 8$$`:R%:C `VJRQ`VR1IVJ $`:1J1 1$%`V 88 T %``:HV VR1IVJ R1 `1G% 1QJ :CQJ$ : 1%I HQJ 1JVJ :C .VC`8 VR1IVJ HC:1`1H: 1QJ:HHQ`R1J$ Q Q : ^ _7:JR^Y_^] QHQJ VJ Q`:JR_5I%RR7:JR^IY_ ^ T Q_5 0V`7 :JR7 I%R ^02Y_ ^ TQ_5 :JR7 I%R ^2Y_ ^T Q_ :JR I%R ^2Y _ ^\ Q_8 29 $]`VR1H 1QJI:]Q`%``:HVVR1IVJ R1 `1G% 1QJ1J .V`10V1J0V 1$: VR:`V:1: `V:C1 30 1$%`V 88 T`VR1H 1QJI:]Q`%``:HVVR1IVJ R1 `1G% 1QJ1J .V`10V1J0V 1$: VR:`V::CQJ$ : 1%IHQJ 1JVJ :C.VC`5`V:C1 31 8 QC7H.:V V :VIGC:$V7 RVH`1] 10V :JR I%C 10:`1: V :J:C7V .V :6QJ QI1H :J:C71 Q` .V 1.QCV :I]CV HQCCVH VR CVR Q .V 1RVJ 1`1H: 1QJ Q` 1JR101R%:C8GVCQJ$1J$ Q`:I1C1V:JR ]VH1V^ :G888 _8 .V _%:J 1 : 10V `%H %`V Q` ]QC7H.:V V :VIGC:$V Q` V:H. :`V: `VC: VR Q G1QHQVJQ1 :``1C1: 1QJ Q` ]VH1V5 ]`V`V``VR VR1IVJ 7]QCQ$1V :JR RV] . `:J$V R1 `1G% 1QJ.:GVVJRVH`1GVR V]:`: VC7``QI .VJQ` .V`J Q .VQ% .V`J:`V:8 $HHQ`R1J$ Q ,HH.1]1J 1 V :C 8 ^ _ :JR 3:J QJV V :C 8 ^ _5 .`VV :C1VJ ]VH1V 1V`V`Q%JR7 VR1QI: % H:]VJ1 5 Q QI: % :GV`:J :JR V :7H.1 $Q Q1 8 .V:I]C1J$ : 1QJ1.V`VJQ]QC7H.:V V]VH1V1V`V`Q%JR1V`VJQ HQJ1RV`VR 11 .1J .V]`VVJ : 1QJQ``V%C 8 32 $:GCV 88 TQC7H.:V :]VH1V`Q%JR1J .V`10V %R7:`V::CQJ$ .V : 1%IHQJ 1JVJ :C .VC`8 !"# $# QC7QRQJ V I:61CCQ% ^ :J<:J15 _ :I1C7 !:]1 VCC1R:V :I1C7 #].`QR1 1R:V :]1 VCC: H:]1 : : ^&:G`1H1%5 _ !:V IQJ1HV .7 `16 ^:01$J75 _ :]1 VCC: $1:`R1 ^2VJ1C5 _ :I1C7 QC7JQ1R:V :]1 QI: % I1J1I% ^ :J$V`.:J5 _ :`IQ .Q[ ]1J1`V`: ^".CV`5 _ V V`QI: % `1C1`Q`I1 ^3C:]:`WRV5 _ :CI$`VJ1VCC: C=%J$I:J1 ^2:CI$`VJ5 _ !V1QH:]1 VCC: $C:G`: 6:` I:J5 :CI$`VJ1VCC: C%J%C: : ^VCCV3.1:=V5 _ !V1QH.`1RV ]8 :I1C7 1$:C1QJ1R:V VR1QI: % H:]VJ1 :75 VCQ$VJ1: :`VJQ: ^VCCV3.1:=V5 _ VR1QI: % ``:$1C1 :I%VJ5 %1$:C1QJ I: .1CR:V $%RQ%1J21CJVR"R1:`R5 Q QI: % :GV`:J :75 %1$:C1QJ _%:IQ%I ^VCCV3.1:=V5 _ Q QI: % C: V`1HV% 28:`5 % .VJVC:1 GQ: ^EQ.J QJ5 _ Q QI: % C1JV: % 3C:]:`WRV5 % .VJVC:1 C1I1HQC: ^".CV`5 _ V%RQCV1QH:]1 VCC: `:%0VC1 6:`IVC1J5 :I1C7 !.`7Q]V :C1R:V :I1C7 (:CR:J1R:V .`7Q]V :C%I RVG1CV ^4`%GV5 _ #61Q .VCC: HQJ `1H : ^3C:]:`WRV5 _ :I1C7 "V1QJ1R:V C7IVJ%`: HC7]V: : ^:1J REQV].5 _ V1QJV ]CVJR1R: :01$J75 $%HC7IVJV C%IG`1HQ1RV ^_%: `V`:$V5 _ :I1C7 V`V1R1R:V $%HC7IVJV ]:CV`I1 :J: ^4`%GV5 _ %GR&:I1C7 V`V1R1R1J:V `:61CCVCC: :``1J1 ^28:`5 _ $%JV`V1 CQJ$11I: ^EQ.J QJ5 _ `:61CCVCC: $`:H1C1 ^28:`5 _ :I1C7 1C:`$1R:V :CR:JV $CVG1`V6 4`%GV5 #JH1 :`$1 .:I: : ^6:` I:J5 _ :CR:JV :`1 2:CI$`VJ5 #JH7 `Q7CC1 $`QVJC:JR1H: 2H J Q.5 V :7H.1 $Q Q1 ^ <%@:5 _ 1C:`$1 0V``%HQ: ^:1J REQV].5 _ 1HQI:H.V C%IG`1H:C1 ^&:G`1H1%5 _ %1$:IG`: VJ :H%C: : ^ `V:R1VCC5 _ V :CQ]`QH % V``1HQC% _%: `V`:$V5 :I1C7 'C7HV`1R:V :I1C7 !Q%`1R:V -C7HV`: :CG: ^,8&82*CCV`5 _ Q%`: Q7V`1 :%G1V`5 -C7HV`: H:]1 : : b` VR5 :I1C7 ].VC11R:V -C7HV`: %J1HQ`J1 :01$J75 %GR&:I1C7,].VC1J1J:V :I1C7 'QJ1:R1R:V #`I:JR1: H1``.Q: &1C1]]15 -C7H1JRV JQ`RI:JJ1 ^2:CI$`VJ5 _ QC7Q]. .:CI% ]1H % ^%=:`R1J5 _ -QJ1:R: VIV`1 : $%RQ%1J21CJVR"R1:`R5 :I1C7 H:C1G`V$I: 1R:V -QJ1:R: I:H%C: : b` VR5 %H:C1G`V$I: 1J`C: %I & : .@V5 :I1C7 V]. 71R:V %HCV`QH.V1C% I1J% % 4`%GV5 V]. .7 H1``Q: ".CV`5 :I1C7 `G1J11R:V V]. .7 .QIGV`$1 :01$J75 %GR&:I1C7,`G1J11J:V V]. .7 .7 `1H1 2H J Q.5 '`G1J1: H%01V`1 ^$%RQ%1J21CJVR"R1:`R5 _ V]. .7 1JH1: 2:CI$`VJ5 .7CQ `QV 1R: ^3C:]:`WRV5 _ V]. .7 ]8 .7CQ C1$% 1H: ^,`C:JR15 _ :I1C7 .7CCQRQH1R:V .7CQ JQ`0V$1H: ^28:`5 _ %GR&:I1C7" VQJ1J:V %HQC:`1H1: 7]1H: "11$5 $ VQJV CQJ$: ^&:G`1H1%5 _ %HQCQ]CQ ^%HQCQ]CQ _ :`I1$V` ^,8&82*CCV`5 _ $ VQJV ]1H : _%: `V`:$V5 :I1C7 :`:QJ1R:V :I1C7 :`:C:H7RQJ11R:V 1``Q].Q`% G`:JH.1: % ".CV`5 :`:C:H7RQJ1: H` ]:`:RQ6: !V01JVJ1: $`:H1C1 ^ :%GV`5 _ %GR&:I1C7.7CCQRQH1J:V :I1C7 :`:QJ1R:V .7CCQRQHV C:I1JQ: :I:`H@5 :`:QJ1 `%C$VJ ^ V01JVJ5 _ .7CCQRQHV C1JV: : ^3C:]:`WRV5 _ :I1C7 !1``: %C1R:V .7CCQRQHV I:H`Q]. .:CI: H.I:`R:5 #].VCQH.:V : I:`1QJ1 ^:1J REQV].5 _ .7CCQRQHV H` I:H%C: : :%CCV`1VCC: I%C 1G`:JH.11 ^4`%GV5 _ :I1C7 %].`Q1J1R:V .V Q 33 !"# $# :I1C7 #I].:`V 1R:V :I1C7 %J1H1R:V %GR&:I1C72VC1JJ1J:V $%J1HV :J VJJ: : ^:01$J71J :I:`H@5 _ VC1JJ: ]:CI: : 4`%GV5 $%J1HV .:`:11 $%RQ%1J21CJVR"R1:`R5 :I1C7 $`1H.QG`:JH.11R:V $%J1HV QV` VR1 1I]QJ5 1V`VGVCC1RV `QVI1 28:`5 $%J1HV ]VJJ: : ^,8&82*CCV`5 _ :I1C7 $V`VGVCC1R:V $%J1HV 01 : : ^VCCV3.1:=V5 _ %GR&:I1C7$I].1 `1 1J:V !71R1HV H` J1JV : 1 : H`1 : : ^,8&82*CCV`5 _ :`].7: GVCC1 ^$%RQ%1J21CJVR"R1:`R5 _ %GR&:I1C7QC7H1``1J:V :`].7: `:CC:6 2:`1QJ 34 88 ( $# $ !#$& :I]C1J$1J2 752 <52 ::JR2 G : 1QJHQJ`1`IVR .V ]`VVJHV Q` Q1RQJ1: QHV:J1H: IV:RQ1 1.1CV 1J 2 R5 2 V5 2 `5 2 $ :JR 2 $ : 1QJ1V`V`Q%JR.:`R%G `: :QHH:1QJ:CC711 . 8 QHV:J1H: 8$ IQ`RV] .5 :I]CV HQCCVH VR 1J 2 75 2 <5 2 :5 2 G5 2 H5 2 R5 2 V :JR 2 ` : 1QJ1JR1H: VR .VQHH%``VJHVQ` 8 QHV:J1H: >RV:RI: V?8,J .VHQJ `:`75 :CQJ$ `:JVH .5]Q1 1QJVR 1J ``QJ Q` 2:` :`10V`5 V:`CQQ` Q` : 1QJ 2 . 1: I:1JC7 HQI]QVR G7 I%RR7 VR1IVJ 8 3QJV_%VJ C7 .V ]QC7H.:V V :VIGC:$V :J:C7VRHQI1J$``QIQ` : 1QJ]C:JJVR8 $ Q :C Q` 1JR101R%:C GVCQJ$1J$ Q ]VH1V 1V`V `Q%JR : .V 1J0V 1$: VR : 1QJ:`V]Q` VR1J $]]VJR16 $ IV V`RV] . .VIQ :G%JR:J ]VH1V`Q%JR 1V`V V]. 7 .QIGV`$1 ^ 1JRLI _5 7]1H:C ]VH1V Q` :JR :JR I16VR VR1IVJ 5 Q QI: % :GV`:J ^ _5 !%IG`1JV`1 C: `V1CC1 ^ _5 GQ . :QH1: VR Q I%RR7 VR1IVJ VJ`1H.VR Q` Q`$:J1H I: V`:JR $%J1HV 01 : : ^_8:JR7]VH1V: '1VJ1: `%1`Q`I15 V]. 7 .QIGV`$1 5 %HQC:`1H1: 7]1H: ^:``1C1: VR Q1VCCQ` VR`1JV:JRG1QHQVJQ1_:JR 1C:`$1 0V``%HQ: Q$V .V` 11 . ]VH1V H.:`:H V`1 1H Q` VR1IVJ HQI]QVR G7 Q`$:J1H RV `1 % : #]QJ%].1 G`VIVJ 15 $%].`Q1JV :`I:R1CCQ :JR %1$:IG`: VJ :H%C: : 1V`V :CQ `Q%JRVR8 J2 H : 1QJ]VH1V: :`:C:H7RQJ1: H` ]:`:RQ6:5 -C7HV`: %J1HQ`J1 :JR #].VCQH.:V : I:`1QJ1 51JR1H: V .VQHH%``VJHVQ`I%RR7``:H 1QJ1J .VVR1IVJ 8$ IV V` Q QI: % :GV`:J ^ _ %HQCV QI: VI:JR1G%C: :RI:G1 1 ^ _:JR VC1JJ: ]:CI: : ^_5 1J.:G1 1J$ 1J I%RR7 VR1IVJ VJ`1H.VR Q` Q`$:J1H I: V`5 1V`V .V IQ :G%JR:J ]VH1VG% 1V`V:CQ_%1 VJ%IV`Q% ]VH1V : !:QJ1HV H` H1``: :5 VH 1J:`1: :%`1HQI:5 V V`QI: % C: `V1CC15 Q QI: % C: V`1HV%5 -8 %J1HQ`J18 V :7H.1 $Q Q1 ^ _5 #I].:`V V :H% 1``QJ ^ _ ^:``1C1: VR Q 3QJ 1JVJ :C V``1$VJQ%2%RG1QHQVJQ1_5 -8 %J1HQ`J1 ^ _5 !8 C: `V1CC1 ^_:JR 8 :GV`:J ^ _ ^GQ .:QH1: VR QI%RR7VR1IVJ VJ`1H.VRQ`Q`$:J1HI: V`_1V`V .VRQI1J:J ]VH1V Q` :VIGC:$V : IV V` RV] .8 .V Q0V`:CC :VIGC:$V 1: 7]1H:C Q` I%RR7V:GVRG% : 2 :52 R:JR2 V:CQQHH%``VR]VH1V:QH1: VR Q $`:0VC ``:H 1QJ Q` VR1IVJ ^V8$8 7C1JQ:H1V: %G1HQC: 5 '8 `%1`Q`I1 5 7`1QH.VCV QH%C: : 5 VH 1J:`1: :%`1HQI: _8 .V :VIGC:$V Q` : 1QJ : I Q` RV] . 1V`V 35 RQI1J: VRG7 !8 C: `V1CC1 ^ _5 8 `1C1`Q`I1 ^ _5 18 `QVI1 ^_5 #8 :H% 1``QJ ^_ :JR %HQCV QI: VI:JR1G%C: :RI:G1 1 ^ _5]VH1V%%:CC7QHH%``1J$1JI%RR7V:GVR :1VCC:1J .VHQJ 1JVJ :C V``1$VJQ%I%RG1QHQVJQ1 V0VJ 1` .V7 H:J QCV`: V HQ:`V ``:H 1QJ Q` VR1IVJ 8 J QIV : 1QJ ]VH1V : #]QJ%].1 `:%0VC15 8 %G1HQC: 5 .`1CQJV`V1 `1C%I 5 8 :%`1HQI: %$$V VR .V ]`VVJHV Q` RV `1 1H .:G1 : 1.V`V .VV]VH1V:`VIQ HQIIQJ8$ I .VIQ :G%JR:J ]VH1V1V`V -8 %J1HQ`J1 ^ _5 %8 VI:JR1G%C: :RI:G1 1 ^ _:JR #8 I:`1QJ1 ^ _51JR1H: Q`Q`VR1IVJ VJ`1H.VRQ`Q`$:J1HI: V`8 .V:VIGC:$VQHH%``VR: .1RV] .1V`V 7]1H:CC7Q` I%RR7V:GVRV6HV] `Q` .QV`Q%JR1J2 G : 1QJ1.V`VQIV]VH1V:``1C1: VR QI%RR7RV `1 1HG1QHQVJQ11:`VHQ`RVR^ 8 %G1HQC: 5 !V1QH:]1 VCC: $C:G`:_ 8 2V:J ]VH1V `1H.JV 0:C%V ^ 1.1H. 1V1$. V_%:CC7 :CC ]VH1V `V$:`RCV .V1` :G%JR:JHV_1V`V`VC: 10VC7HQJ :J :CQJ$ .VRV] .$`:R1VJ G% :_%1H@C7RVH`V:1J$ 1:QGV`0VR: I^ `VJRHQJ`1`IVRG7"1.1H.V 1I: V .VV6]VH VRJ%IGV`Q` ]VH1V 1J :J .7]Q .V 1H:C :I]CV Q` 1JR101R%:C_6 .V ]VH1V :G%JR:JHV ^ _ $`:R%:CC7 RVH`V:V 11 . RV] .8 2V:J R10V`1 1V ^ 1.1H. $10V CV 1V1$. Q .V `:`V ]VH1V5 :JR 1.1H. ]C:HV IQ`V 1V1$. Q .V HQIIQJ ]VH1V_ .Q1 HQI]:`:GCV `VJR5`V:H.1J$ .V.1$.V 0:C%V: I :JR .V CQ1V : I Q` RV] .81VCQ%V0VJJV^E;_RVH`1GV:.1$.V_%1 :G1C1 7:IQJ$ .V]VH1VQHH%``1J$: R1``V`VJ G: .7IV `1V11 . .V`V1V J%IGV`Q`RQI1J:J ]VH1V: I^ 1J` 7 5 _ ^&1$8888 _^ &1$8888 _8 .V:J:C71Q`IV:J0:C%VQ`HQII%J1 71JR1HV:HHQ`R1J$ QVR1IVJ 0:`1: 1QJ.Q1]VH1V`1H.JV^ _:JR:G%JR:JHV^ _RVH`V:V11 . .V 1JH`V:VQ``1JV``:H 1QJ1J .VVR1IVJ 8$:HQJV_%VJHV .VRVV]V`:JRI%RR7 : 1QJ1V`VH.:`:H V`1 RQI1J:J ]VH1V1:`Q%JR:.Q1VRG7E;:JR 1J` 1JR1HV `VJR^ &1$8888 _8 36 A A 5 5 5 5 5 5 5 5 5 5 5 1J` 1J` 1$%`V 888 TQC7H.:V VR10V`1 7:HHQ`R1J$ QRV] .^QJ .V`1$. _:JRVR1IVJ HC:V ^QJ .VCV` _:CQJ$ .VHQJ 1JVJ :C.VC`Q`2QJ :C QR13: `Q7 ]VH1VRVJ1 7^1JR8LI _5E; 1VCQ%V0VJJV65 5 5 5 1J`61CCR10V`1 7J%IGV`8VR1IVJ HC:V:HHQ`R1J$ Q Q : ^ _7 :JR ^Y_ ^] Q HQJ VJ Q` :JR_5 I%RR7 :JR ^IY_ ^ T Q_5 0V`7 :JR7 I%R ^02Y_^ TQ_5:JR7I%R^2Y_^T Q_:JRI%R^2Y _^\ Q_8 37 1$%`V 888 T`VR1H 1QJI:]Q`"51.1H.V 1I: V .VV6]VH VRJ%IGV`Q`]VH1V1J:J.7]Q .V 1H:C:I]CVQ`1JR101R%:C51J .V:`V:Q`2QJ :C QR1 3: `Q8 38 1$%`V 888 T`VR1H 1QJI:]Q`%``:HVVR1IVJ R1 `1G% 1QJ:CQJ$2QJ :C QR13: `QHQJ 1JVJ :C.VC`8 39 88 $ &## !# ,0V`: Q :CQ` : 1QJ5 ]VH1V:JR 1JR101R%:C1V`VHQCCVH VR^ $]]VJR16 _8 J .V.:CCQ1V : 1QJ^ I_ .VRQI1J:J ]VH1VHQCCVH VR '1VJ1: `%1`QI1 ^ 1JR8LI _5 'J%].7 V`VI1 : ^ _5 %1$:C1QJ I: .1CR:V ^_ :JR VR1QI: % ``:$1C1 ^_:`V 7]1H:CC7`Q%JRQ`:JR7VR1IVJ :JR:``1C1: VR Q .V1VCCQ` VR`1JV:JR G1QHQVJQ18 .V:VIGC:$V1:HQI]QVRV6HC%10VC7G7:G%C1HQC%]VH1V:1VCC :1:QGV`0VR: IQ`RV] .1.V`V .VIQ :G%JR:J ]VH1V1V`V '8 `%1`Q`I1 ^ _5 Q .`1: HQJH71CV$: ^_ ^GQ . V6HC%10VC7 `Q%JR 1J V :JR $_5 %8 I: .1CR:V ^ _ :JR :$VCQJ: ]8 $ I5 ]VH1V .: $VJV`:CC7 ]`V`V` HQ:`V VR1IVJ ^:JRQ`Q`$:J1HRV `1 %_%H.: #]QJ%].1 G1C1JV: : ^ _5 %]1Q RVHQ`: % ^ _5 '8 `%1`Q`I1 ^_HQV61 VR11 .]VH1V5%%:CC71J.:G1 I16VRVR1IVJ :JR :`V QCV`:J QR1``V`VJ :IQ%J Q`I%R:JRQ`$:J1HI: V`5%H.: VC1JJ: ]:CI: : ^ _5 :]1 VCC: H:]1 : : ^ ]VH1IVJV6HC%10VC71J ._:JR Q QI: % :GV`:J ^ _5 .VIQ :G%JR:J 5:JR:CQ C7IVJ%`: CV1Q]7$Q ^ _5 #I].:`V V :H% 1``QJ ^_5 -C7HV`: %J1HQ`J1 ^ _:JR V V`QI: % `1C1`Q`I1 ^ _8 JH`V:1J$ .VRV] .^ I_ .V J%IGV` Q` ]VH1V 7]1H:C Q` I%RR7 VR1IVJ 1JH`V:VR5 18V8 !%IG`1JV`1 C: `V1CC1 ^ _5 2`:R: 01CCQ: ^ _5 %HQCV QI: VI:JR1G%C: :RI:G1 1 ^ _5 -8 %J1HQ`J1 ^_ 8 :GV`:J ^_5 VC1JJ: ]:CI: : ^ _:JR .`1CQJV`V1 `1C%I ^_V0VJ1`:`V1]VH1V5 `Q`V6:I]CV '8 `%1`Q`I1 ^ _5 #8 `:%0VC1 ^ _5 %8 RVHQ`: % ^ _5 7]1H:CC7:QH1: VR Q :JR7V:GVR5 1CCQHH%``VR1JQIV : 1QJ^V8$ $_11 .CQ1V`:G%JR:JHV .:J QGV`0VR1J .V.:CCQ1V`QJV8 40 1 `1` C7 QHH%``VR ``QI I Q` RV] .8 : 1J V`V 1J$ Q 1$J:C .: QIV ]VH1V JQ`I:CC7`Q%JR1J.:CCQ1V`:JR7V:GVR1V`VHQCCVH VR1JQIVRVV] : 1QJ`Q` 1J :JHV #8 `:%0VC1 : R I8 ]VH1V`1H.JV^ :JR"_`V:H.VR .V.1$.V 0:C%V: :JRIRVH`V:1J$11 . RV] .6CQ10:C%V1V`V:CQ`VHQ`RVR1J .V.:CCQ1V : 1QJ^ :JRI_1.V`VV `V1]QC7H.:V VQHH%``VRQHH:1QJ:CC711 .:.1$.J%IGV`Q`]VH1IVJ8 .%: I .V]V:@Q`]VH1V:G%JR:JHV^ _1:I:1JC7R%V Q .VRQI1J:JHVQ` '8 `%1`Q`I1 :JR 8 HQJH.1CV$: 83QI]:`:GCVIV:J0:C%VQ` 1:QGV`0VR: I1.V`V:I:CCV` :IQ%J Q`RQI1J:J ]VH1V1:`Q%JR8 RVH`V:VR8 10V`1 7 1JR1HV ^ :JR _ .Q1 1I1C:` `VJR :JR .V.1$.V 0:C%V 1V`V`VHQ`RVR: :JRIQ`RV] .8 .VQ0V`:CC :VIGC:$VQ` .V:`V:1V`V H.:`:H V`1 `Q` .VRVV]V`QJV^ 1J` 75 : I_^ &1$8888 _^ &1$8888 _8 .V1JR1HV `VJR1J `VC: 1QJ Q VR1IVJ HC:V .Q1 .: ]VH1V `1H.JV ^ _ :JR :G%JR:JHV ^ _ `V:H.VR .V.1$.V 0:CV1JI%RR7R:JR : 1QJ^I_:JRRVH`V:VR]`Q]Q` 1QJ:CC7 Q .V1JH`V:VQ`]V`HVJ :$VQ`GQ .:JR:JRI%R 1J .V VR1IVJ 8 .V .1$.V R10V`1 7^ :JR _1:QGV`0VR1JVR1IVJ `:J$1J$``QII%RR7R:JR Q0V`7R :JR7I%R1.1CV .V]QQ`V :VIGC:$VI:1JC7H.:`:H V`1 RQI1J:J ]VH1VQHH%``VR1J:JR7 : 1QJ^E;75 :JR 1J` 75_^ &1$8888 _^ &1$8 888 _8 41 A A 5 5 5 5 5 5 5 5 5 5 5 5 1J` 1J` 1$%`V 888 TQC7H.:V VR10V`1 7:HHQ`R1J$ QRV] .^QJ .V`1$. _:JRVR1IVJ HC:V ^QJ .VCV` _:CQJ$ .VHQJ 1JVJ :C.VC`Q` Q`0:1:J1H:7 ]VH1VRVJ1 7^1JR8LI _5E;1VCQ% V0VJJV65 5 5 5 1J`61CCR10V`1 7J%IGV`8VR1IVJ HC:V:HHQ`R1J$ Q Q :^ _7 :JR ^Y_^] Q HQJ VJ Q` :JR_5I%RR7 :JR ^IY_ ^ T Q_5 0V`7:JR7I%R ^02Y_ ^ TQ_5:JR7I%R^2Y_^T Q_:JRI%R^2Y _^\ Q_ 42 1$%`V 888 T`VR1H 1QJI:]Q`"51.1H.V 1I: V .VV6]VH VRJ%IGV`Q`]VH1V1J:J.7]Q .V 1H:C:I]CVQ`1JR101R%:C51J .V:`V:Q` Q`0:1:J1H:8 43 1$%`V 888 T`VR1H 1QJI:]Q`%``:HVVR1IVJ R1 `1G% 1QJ:CQJ$ Q`0:1:J1H:HQJ 1JVJ :C.VC`8 44 88 #)## ,0V`:CC ]VH1V11 .: Q :CQ` 1JR101R%:C1V`V:I]CVR1J .V : 1QJQ` :G:%R1: ^ $]]VJR16 _8 Q1RQJ1: QHV:J1H: IV:RQ1 :`V ]`VVJ 1J ``QJ Q` Q``V $ %`:]`QIQJ Q`7:JR .VHQ: :CC:@VQ`Q% .V`J : 1%I5`:J$1J$``QI:GQ% ^%]]V`C1I1 _ QIQ`RV] .8 `:JVH ``QI>H? Q>.?:H`QQ`1V`VCQH: VR1J ]`Q61I1 7Q` .VIV:RQ18 $ IRV] . .VIQ :G%JR:J :JR11RVR1 `1G% VR ]VH1V 1V`V :$VCQJ: ] ^ 1JRLI_5 %HQCQ]CQ :`I1$V` ^ _5 !%IG`1JV`1 V `:%`: ^_5 %1$:C1QJ I: .1CR:V ^ _5 7]1H:C ]VH1V Q` 1VCC Q` VR `1JV :JR G1QHQVJQ15 :JR %]1Q RVHQ`: % ^_8 .VV]VH1VGVH:IVIQ`V:G%JR:J : I1.V`VQ .V`:G%C1HQC%]VH1V%H.: `1QJQ]1Q H:]V`1 ^ _5 V]. 7 .QIGV`$1 ^_ :JR '1VJ1: `%1`Q`I1 ^ _ 1V`V HQCCVH VR8$ IQ`RV] .:VIGC:$V1V`VI:1JC7HQI]QVRG7:JR7]VH1V5V8$8 #]QJ%].1 G`VIVJ 1 ^_5 %8 RVHQ`: % ^ _5 %]1Q].:JV GQIG76 ^ _5 %8 :`I1$V` ^_ :JR ]VH1V Q` I%RR7 V:GVR : !%IG`1JV`1 C: `V1CC1 ^ _5 VC1JJ: ]:CI: : ^ _5 !:QJ1HV H` H1``: : ^ _ ^:``1C1: VR Q HQJ 1JVJ :C V``1$VJQ% I%R G1QHQVJQ1_5 Q QI: % :GV`:J ^ _5 QVH1CQH.V % V`]VJ ^ _8 2Q`VQ0V`5 : $R : 1QJ5 CQH: VR 1J ]`Q61I1 7 Q` .V Q1RQJ1: QHV:J1H: 5 IV:RQ1 1V`V 1$J:CVR ]VH1V :QH1: VR QQ`$:J1HRV `1 1HV:GVR: V]. 7 1JH1: :JR 2 7`1QH.VCV QH%C: : :JR #`:GVCC: 1`1HQCQ`5 ``V_%VJ 1J :C$:C G1Q Q]V8 QIV Q` .V :GQ0V ]VH1V 1V`V :CQ HQCCVH VR: I1.V`V:J1JH`V:VQ` .V1`:G%JR:JHV1:QGV`0VR5V8$8 !8 C: `V1CC1 ^ _5 %8 RVHQ`: % ^_5 %8 :`I1$V` ^ _5 8 V`]VJ ^ _5 #8 G`VIVJ 1 ^_ 8 61$. RVJ1 1V 1V`V 1$J:CVR : 1VCC `Q` Q .V` ]VH1V : V]. 7 .QIGV`$1 ^_5 %]1Q I%C 1QH%C: : ^ _5 :`:C:H7RQJ1: H` ]:`:RQ6: ^ _ % V`J:]1 H% : : ^ _ .: QHH%``VR: .V:IVRV] .8$ `:J1 1QJ ]VH1V`1H.JV^ :JR"_1JH`V:VR%] QIRV] .1.V`V`V:H.VR .V.1$.V 0:C%V6 GVCQ1 .1 RV] . .V J%IGV` Q` ]VH1V RVH`V:VR :JR .V : 1QJ ]C:HVR GV 1VVJ :JRI.Q1VR .VCQ1V 0:C%V8$ I1::CQ`VHQ`RVR:]V:@Q` ]VH1V :G%JR:JHV ^ _ R%V Q .1$. RVJ1 1V Q` : J%IGV` Q` ]VH1V :I]CVR8 .V `1H.V J%IGV`Q`RQI1J:J ]VH1V1:1$J:CVR: I^ 1J` 75 _^ &1$8888 _^ &1$8888 _8 ]VH1V `1H.JV ^ _:JR:G%JR:JHV^ _1JH`V:VR]`Q]Q` 1QJ:CC7 Q .V 1JH`V:V Q` ]V`HVJ :$VQ`I%R1J .VVR1IVJ G% 1.V`V .1``:H 1QJ1:] Q:HQJV_%VJ 1:C V0V`VRVH`V:V1:QGV`0VR8V`7R:JR7I%R:JR:JR7RI%RVR1IVJ .Q VR .V .1$.V R10V`1 7 ^ :JR _5 :JR JQ RQI1J:J ]VH1V ^E; :JR 1J` _ VVIVR Q GV :QH1: VR Q RV`1J1 V $`:J%CQIV `1H H.:`:H V`1 1H Q` VR1IVJ ^ &1$8 888 _ ^ &1$8 888 _8 46 A A 5 5 5 5 5 5 5 5 5 5 5 5 1J` 1J` 1$%`V 888 TQC7H.:V VR10V`1 7:HHQ`R1J$ QRV] .^QJ .V`1$. _:JRVR1IVJ HC:V ^QJ .V CV` _ :CQJ$ .V HQJ 1JVJ :C .VC` Q` :G:%R1:7 ]VH1V RVJ1 7 ^1JR8LI _5 E; 1VCQ% V0VJJV65 5 5 5 1J`61CCR10V`1 7J%IGV`8VR1IVJ HC:V:HHQ`R1J$ Q Q :^ _7 :JR ^Y_^] Q HQJ VJ Q` :JR_5I%RR7 :JR ^IY_ ^ T Q_5 0V`7:JR7I%R ^02Y_ ^ TQ_5:JR7I%R^2Y_^T Q_:JRI%R^2Y _^\ Q_ 47 1$%`V 888 T`VR1H 1QJI:]Q`"51.1H.V 1I: V .VV6]VH VRJ%IGV`Q`]VH1V1J:J.7]Q .V 1H:C:I]CVQ`1JR101R%:C51J .V:`V:Q`:G:%R1: 8 48 1$%`V 888 T`VR1H 1QJI:]Q`%``:HVVR1IVJ R1 `1G% 1QJ:CQJ$:G:%R1:HQJ 1JVJ :C.VC`8 49 88 $ &&#! # ]VH1V :JR 1JR101R%:C 1V`V 1RVJ 1`1VR Q0V`: Q :CQ` Q` .V : 1QJ ]C:HVR 1J V``:H1:J: ^ $]]VJR16 _8 $ 11RV Q1RQJ1: QHV:J1H: IV:RQ1 .:0V GVVJ 1$J:CVR:CQ1J .1:`V:5``QI:GQ% IQ``.Q`V3:]V31`HVQ Q .V]`Q61I1 7Q` ]V`CQJ$: 01CC:$V8 .V CQ1V` C1I1 `V:H. :GQ% I Q` RV] . :JR `QCCQ1 .V G: .7IV `7 `VJR8 .V:VIGC:$V1J0V 1$: VR: IRV] .1:RQI1J: VRG7 :$VCQJ: ] ^ _5 `1QJQ]1Q H:]V`1 ^ _5 :``1C1: VR Q .V 1VCC Q` VR `1JV :JR G1QHQVJQ1 5 !%IG`1JV`1 `%JH.:CVJ1 ^_5:J.:`RV:GVR]VH1V .: %$$V .V]`Q61I1 7Q` `QH@7 .Q:C5 '1VJ1: `%1`Q`I1 ^_5 VR1QI: % ``:$1C1 ^_8 %HQCQ]CQ :`I1$V` ^_5 %]1Q RVHQ`: % ^ _5 %1$:C1QJ I: .1CR:V ^ _8 ,JC7 QIV Q` .V IVJ 1QJVR ]VH1V1V`V`Q%JR: : 1QJ VJ :JR "] 1.V`V .V:VIGC:$V1V`V]QQ`1J V`IQ`]VH1VJ%IGV`:JR:G%JR:JHV8$1I1C:`]VH1VHQI]Q1 1QJ1:QGV`0VR : I1.V`V:`VR%H 1QJQ` .VRVJ1 7Q`QIVQ` .VV]VH1V1:QGV`0VR1J Q]]Q1 1QJ 11 . .V 1JH`V:V `VHQ`RVR `Q` QIV Q .V`5 V8$8 8 H:]V`1 ^ _5 :$VCQJ: ] ^ _5 %]1Q RVHQ`: % ^ _ V %8 :`I1$V` ^ _8 .V.1$.V 0:C%VQ`]VH1V`1H.JV^ :JR"_1:`V:H.VR: I8 Q10:C%VQ` :JR"1V`V:CQ`VHQ`RVR: :JRI1.V`VV`V1]VH1VQHH%``VR11 ..1$. :G%JR:JHV8 J ]:` 1H%C:` : I .V .1$.V ]VH1V :G%JR:JHV ^ _ 1: QG :1JVR I:1JC7`Q` .VRQI1J:JHVQ` 8 H:]V`1 :JR :$VCQJ: ]83QI]:`:GCVIV:J0:C%VQ` 1:QGV`0VR``QI QIQ`RV] .:JRRVH`V:VRGVCQ1 .1`:J$V8 .VIQ R10V`V:VIGC:$V^ :JR 1JR1HV_1V`V`Q%JR``QI Q I8$ I:1VCC :: :JRIV_%1 :G1C1 7^E;_`V:H.VR .VCQ1V 0:C%V:JR: I :J%IGV`Q`RQI1J:J ]VH1V1::I]CVR^ 1J` 75 _V6HV] `Q` : 1QJ VJ :JR "] ^ &1$8 888 _ ^ &1$8 888 _8 .VI:61I%IJ%IGV`Q`]VH1V^ _1: `Q%JR1J I%RR7R:JR : 1QJ :JR .V .1$.V :G%JR:JHV ^ _ 1J :JR6 RVJ1 7 RVH`V:VR ]`Q]Q` 1QJ:CC7 Q .V1JH`V:VQ`]V`HVJ :$VQ`I%R1J .VVR1IVJ 8 .V:VIGC:$V H.:`:H V`1 51 ``QI I%RR7R:JR Q 0V`7 :JR7RI%R 1.1CV .V ]QQ`V :VIGC:$V I:1JC7 H.:`:H V`1 52 A A 5 5 5 5 5 5 5 5 5 5 5 5 1J` 1J` 1$%`V 888 TQC7H.:V VR10V`1 7:HHQ`R1J$ QRV] .^QJ .V`1$. _:JRVR1IVJ HC:V ^QJ .V CV` _ :CQJ$ .V HQJ 1JVJ :C .VC` Q` V``:H1J:7 ]VH1V RVJ1 7 ^1JR8LI _5 E; 1VCQ% V0VJJV65 5 5 5 1J`61CCR10V`1 7J%IGV`8VR1IVJ HC:V:HHQ`R1J$ Q Q :^ _7 :JR ^Y_^] Q HQJ VJ Q` :JR_5I%RR7 :JR ^IY_ ^ T Q_5 0V`7:JR7I%R ^02Y_ ^ TQ_5:JR7I%R^2Y_^T Q_:JRI%R^2Y _^\ Q_ 53 1$%`V 888 T`VR1H 1QJI:]Q`"51.1H.V 1I: V .VV6]VH VRJ%IGV`Q`]VH1V1J:J.7]Q .V 1H:C:I]CVQ`1JR101R%:C51J .V:`V:Q` V``:H1J:8 54 1$%`V 888 T`VR1H 1QJI:]Q`%``:HVVR1IVJ R1 `1G% 1QJ:CQJ$ V``:H1J:HQJ 1JVJ :C.VC`8 55 88 '# $# J : 1QJ ]C:HVR : 4:V : :`V: 1JR101R%:C GVCQJ$1J$ Q ]VH1V 1V`V HQCCVH VR^ $]]VJR16 _8 $ IQ`RV] . .V:VIGC:$V1V`VHQI]QVRG7]VH1VJQ`I:CC7`Q%JR1JHQ:`V VR1IVJ ^:JR:JRQ`$:J1HRV `1 %_: #]QJ%].1 G1C1JV: : ^ _5 V]. 7 .QIGV`$11 ^_ 5 '1VJ1: `%1`Q`I1 ^_:1VCC]VH1VQ`I%RR7VR1IVJ Q QI: % :GV`:J ^_ 5 !%IG`1JV`1 C: `V1CC1 ^ _5 VC1JJ: ]:CI: : ^_8 $ I .V ]VH1V HQI]Q1 1QJ1::CIQ V6HC%10VC7H.:`:H V`1 1HQ`I%RR7%G `: ::1JR1H: VRG7 .V .1$. :G%JR:JHV Q` I%RR7 ]VH1V : !8 C: `V1CC1 ^ _5 % V`J:]1 H% : : ^ _ 5 Q QI: % :GV`:J ^_ 5!:QJ1HV H` H1``: : ^ _8 .1]: V`JQ`]VH1VR1 `1G% 1QJ 1::CQQGV`0VR: IQ`RV] .8&`QI IQ`RV] . Q .VRVV]V` .V.1$.V ]VH1V`1H.JV0:C%V^ :JR"_1V`V`V:H.VRGV 1VVJ:JRI: 1VCC:`Q` .V]VH1V:G%JR:JHV^ _8$:HQJV_%VJHV5 .V :VIGC:$V 11 . .V .1$.V ]VH1VR10V`1 7^ :JR _1V`V .QV:J:C7VR: .1RV] .`:J$V81VCQ% 1JRV6^E;_RVH`1GV.1$.V_%1 :G1C1 7:IQJ$ .V]VH1VQHH%``1J$: .V:VIGC:$VQ` 55:JR I:JRI1JQ`0:C%V1V`V51J V:R5`VHQ`RVR: 5 :JR IQ` RV] .8 V0V` .VCV5 JQ RQI1J:J ]VH1V 1V`V 1JR101R%: VR 1J :CC .V 1J0V 1$: VR G: .7IV `1V : V6]C:1JVR G7 1J` 1JRV6 ^ &1$ 88 8 _ ^ &1$ 88 8 _8 .V `1H.V :VIGC:$V 1J V`I Q` J%IGV` Q` ]VH1V ^ _ 1V`V `Q%JR 1J 0V`7R:JR7 I%R VR1IVJ 1.1CV .V.1$.V RVJ1 1VQ`1JR101R%:C^ _1:QGV`0VR1JI%RR7R:JR VR1IVJ 8 Q $`V: R1``V`VJHV:IQJ$]VH1VR10V`1 1V1V`V]Q1J VRQ% :IQJ$ .VR1``V`VJ VR1IVJ 7]V^GQ .`Q` :JR 1JRV6_:1VCC:1:`VHQ`RVR`Q` .VV_%1 :G1C1 7:JRRQI1J:JHV1JR1HV^ &1$88 8 _^ &1$88 8 _8 56 A A 5 5 5 5 5 5 5 5 5 5 5 5 1J` 1J` 1$%`V 88 8 TQC7H.:V VR10V`1 7:HHQ`R1J$ QRV] .^QJ .V`1$. _:JRVR1IVJ HC:V ^QJ .V CV` _ :CQJ$ .V HQJ 1JVJ :C .VC` Q` 4:V :7 ]VH1V RVJ1 7 ^1JR8LI _5 E; 1VCQ% V0VJJV65 5 5 5 1J`61CCR10V`1 7J%IGV`8VR1IVJ HC:V:HHQ`R1J$ Q Q :^ _7 :JR ^Y_^] Q HQJ VJ Q` :JR_5I%RR7 :JR ^IY_ ^ T Q_5 0V`7:JR7I%R ^02Y_ ^ TQ_5:JR7I%R^2Y_^T Q_:JRI%R^2Y _^\ Q_8 57 1$%`V 88 8 T`VR1H 1QJI:]Q`"51.1H.V 1I: V .VV6]VH VRJ%IGV`Q`]VH1V1J:J.7]Q .V 1H:C:I]CVQ`1JR101R%:C51J .V:`V:Q`4:V :8 58 1$%`V 88 8 T`VR1H 1QJI:]Q`%``:HVVR1IVJ R1 `1G% 1QJ:CQJ$4:V :HQJ 1JVJ :C.VC`8 59 8 : 1 1H:C :J:C71 QC7H.:V V :VIGC:$V7 I%C 10:`1: V :J:C7V 2%C 10:`1: V :J:C7V 1V`V ]V``Q`IVR Q 1J0V 1$: V R1 `1G% 1QJ ]: V`J Q` ]QC7H.:V V :VIGC:$V `Q%JR 1J .V `10V I:H`QR:`V:8 .V J2 Q`R1J: 1QJ Q` ]QC7H.:V VR: :5HQJHV`J1J$:G%JR:JHVQ`]VH1V51RVJ 1`1V]: V`JQ`R1 `1G% 1QJ I:1JC7`VC: VR Q .VRV] .$`:R1VJ ^ &1$8 88 _8 V0V` .VCV R1``V`VJHV 1J ]VH1V R1 `1G% 1QJ1::CQQGV`0VR:IQJ$ .V1JR101R%:C1J0V 1$: VR:`V:^ &1$888 _8 U hs )!Th r! Srriyhpr)!T &!7 h !8 v!vvyh v !9!T r)!!! qru T6i' ! " # T6v' $ T6q $ % T6p' T6s T6r' U@r' T6t% HUu! U@p' U@t' U@' T6u' U@' U@t$ T6! HU% U@i# T6' U@"U@! UWs( U@q# U@$ T6q U@s%U@v' HUr#HUt$ U@t" T6" U@q% U@s# U@y# U@! U@ U@ U@u$ U@q$ T6r$ T6y$ U@s$U@r T6t' U@%U@% T6$ U@#U@h% U@"T6u U@$ U@' T6q% U@r$ T6v! U@h T6p# T6q#U@i% U@# T6$ T6v$ U@h' T6v% U@$ T6% U@p$ U@u# T6 T6p%U@u% U@v$ T6y" T6r U@i$ T6r" T6# T6y' U@%T6q' T6v# T6i"T6u" T6t$U@ U@p U@$ T6s%T6h%T6i$ T6t# T6p HU$ UWp$T6h# T6q" HUi% T6u$ T6q! UWs$ U@y% BUp$ HU#T6s# UWq$ UWi# T6y U@t HUp$UWr$ HUu$HUi$UWs# UWr& HU% UWq#UWi%HUp# T6h! T6s UWs% HUr$ HUi# BUh% HUt#UWu#UWp"HU# UWq' HUq#UWp& UWt$ BUh! U@h$ UWq! UWi$ UWhUWt BUp& BUr% HUh$HUu# HUu"BUr" UWh( UWh$ BUp"BUi"BUq" UWr" UWs! T6h$ UWr BUq&HU$HUs# BUq# UWu! UWt%UWq% HUu%HUq$ BUp! T6i BUi%UWu%BUr& HUr% BUq$ UWh"BUp# HUp! UWi! HUs% BUi& HUh% HUh# BUr! UWu$ BUr$ UWt( T6v BUq% UWi'HUq%BUi# UWt" UWs' BUi! BUi$ HUs$ BUr# BUq! UWi UWh& BUp% BUh# T6s! UWp UWp( HUp% UWr( HUp"UWt& UWu'UWs T6t HUt% T6hUWq UWu UWr 1$%`V 88 TJ2Q`R1J: 1QJ]CQ Q`]QC7H.:V V:VIGC:$V:HHQ`R1J$ QRV] .`:J$V7 Y RI5YRI5YR I5Y R I5 Y R I:JR YR 8IQ`RV] .8 U hs )!Th r! Srriyhpr)!T &!7 h!8 v!vvyh v !9!T r)!!! h rh T6 T6 BU T6 T6 U@ T6 T6 T6 UW U@ U@ T6 HU U@ U@ HU T6 U@ T6 T6 HU U@ HU UW T6 U@ U@ T6 U@ U@ HU U@ BU T6 HU U@ T6 U@ T6 U@ U@ U@ T6 U@ U@ U@ U@ T6 U@ U@T6 T6 T6 U@ T6 T6 T6U@ U@ U@ T6 U@ T6 U@ T6 U@ U@ T6 U@ T6T6 U@ U@ U@ BU U@ HU HU U@ T6 UW T6 U@ U@ HU BU HU T6 T6T6 T6 U@ UW HU UW T6 T6 T6 T6 U@ U@ U@ UW HU UW UW HUUW T6 T6 T6 T6 BU BU HUHUHUUW UW HU T6 U@ U@ HU BUUWHU HU T6 T6U@ T6 T6 BUBUUW UW HUUW UWUW HU T6 UW BU HU HU HU U@ HU BUUW HU BUBUBU BU T6 BU UW BU BU UWBU HU UW BU UW U@ UW T6T6 HUBU UW U@ UW UW UW BU UW HU HU UW UW BU T6 UW HU BU UW UWUW UW UW HU BU BU BU UW T6 T6 UW BU UW UW HU UW UW UW T6 UW UW HU UW 1$%`V 88 TJ2Q`R1J: 1QJ]CQ Q`]QC7H.:V V:VIGC:$V:HHQ`R1J$ Q:`V:72 5 5 $5 ":JR4 8 60 " 2$ ,$ :J:C71 ]V``Q`IVR QJ ]`VVJHVL:GVJHV ^G: VR QJ E:HH:`R 1I1C:`1 7_ :JR ]VH1V :G%JR:JHV ^<`:7 3% 1 1I1C:`1 7_ R: : ]`Q0VR .: R1``V`VJHV :IQJ$ :VIGC:$VR1 `1G% 1QJ1V`V1$J1`1H:J C7`VC: V Q .V`:H Q`RV] .`:J$V5:`V: :JR Q .V `VC: 1QJ.1] GV 1VVJ .VV 1Q `:H Q` ^RV] .:`V:_ ^ $]]VJR16 _8 J RV :1C5 .V V ]V``Q`IVR11 .1JV:H.:`V:`Q`RV] . `:J$V .Q1VR 1$J1`1H:J C7 R1``V`VJHV GV 1VVJ :VIGC:$V :I]CVR : R I :JR : I Q` RV] . : 2QJ :C Q R1 3: `Q 1.1CV : Q`0:1:J1H:5 :VIGC:$V `Q%JR : :CC RV] . `:J$V R1``V`VR QV:H.Q .V`8QC7H.:V VHQII%J1 1V1J.:G1 1J$:G:%R1:V:`CQQ```QI I Q` RV] . Q .V RVV]V` 61 .VJ2]CQ QJ .VG:1Q`E:HH:`RIV:%`V:JR " 2 :J:C71.Q1]: V`J Q` R1``V`VJHV 1J :VIGC:$V HQI]Q1 1QJ ^]`VVJHVL:GVJHV_:IQJ$ .V`10V:`V: ^&1$8 88 _ ^ &1$8 88 _ ^ $]]VJR16 _8 .V _%1 V C:`$V ]`V:R Q` :I]CV ]Q1J HQ``V]QJR Q .V:`V:Q`:G:%R1:1.1CV .V_%1 V .1JHC% V`Q`]Q1J HQ``V]QJR Q 4:V :HQI]:`VR11 . .VQ .V`:`V:^ $]]VJR16 _8 .V V `Q`.QIQ$VJV1 7`V0V:C R1``V`VJHV:IQJ$ .V:`V:1J V`IQ` .V1`0:`1:G1C1 71J]VH1VHQI]Q1 1QJV6HV] GV 1VVJ V``:H1J: 0 :G:%R1::JR Q`0:1:J1H::JR2QJ :C Q 0 4:V :8 .V:0V`:$V E:HH:`RR1 :JHVR QRHVJ `Q1R1:GQ% Q`Q`4:V :^ .VCQ1V`]V`HVJ :$V_:JRC:`$V` `Q` V``:H1J:^:GQ% Q_:JR:G:%R1:^:GQ% Q_ ^ $]]VJR16 _8 .V ]: V`J Q` .V V`Q$VJV1 7Q` .VVC: 1Q:`V:1:I1``Q`VRG71I1C:`]: V`JQ`0:`1:G1C1 71J .VVJ01`QJIVJ :CH.:`:H V`1 1H^V8$.V V`Q$VJV1 7Q`VR1IVJ V6 %`V5]`VVJHVQ` 11RV Q1RQJ1: QHV:J1H: IV:RQ1_8 " 2 :J:C71 ]V``Q`IVR Q IV:%`V R10V`1 7 0:`1:G1C1 7 :CQJ$ .V RV] . $`:R1VJ .Q1 HQIIQJ ]: V`J Q` ]VH1V HQI]Q1 1QJGV 1VVJ .V1J``:C1 Q`:C Srriyhpr)!T&!Ehpph q !9!T r)!! h rh T6 BU U@ T6 T6 T6 T6 UW HU T6 U@ U@ T6 T6 T6 U@ U@ U@ HU T6 HU U@ HU T6 U@ T6 UW T6 U@ U@ U@ T6 T6 T6 T6 U@ T6 U@ T6 U@ T6 T6 U@ HU U@ T6U@ T6 T6 U@ U@U@ U@ U@ U@ U@ T6 T6U@ U@ T6 U@ U@ T6U@ U@ T6U@ U@ T6 T6 T6U@U@ T6 T6 HU T6 U@ U@ U@ U@ U@ T6 T6UWT6 T6T6U@ T6 T6 U@T6 U@ BU BU T6 T6 T6 UW BUHU HUHUUW HU HU T6 T6T6 U@ U@ U@ T6 U@ UW UW UWHUUWUWHUUWHU U@ UW UW UWBU HUUWHUUW HU BUHU T6 T6 UW BUHU HU HUUW BU UW U@ UW UW UW BUUWBU UWHUHUBUBUBU UW U@ UW U@ UW UWUW BU BU UW T6 HU HU BUHU BUHU T6 BU BU BU UWHUBU HU BU UW UW BU HUUW HU BUUW UW BU UW UWT6 UW HUBU BUBU UW UW HUHU T6 UW UW UW T6 UW UW UW HU UW 1$%`V 88 TJ2Q`R1J: 1QJ]CQ Q`]QC7H.:V V:VIGC:$V:HHQ`R1J$ Q:`V:72 5 5 $5 ":JR4 8 62 Srriyhpr)!T&!Ehpph q !9!T r)!! qruf htr ! " # $ % 1$%`V 88 TJ2Q`R1J: 1QJ]CQ Q`]QC7H.:V V:VIGC:$V:HHQ`R1J$ QRV] .`:J$V7 Y RI5YRI5YR I5Y R I5 Y R I:JR YR 8IQ`RV] .8 V%C :J:C7 63 U hs )!Th r! Srriyhpr)!T &!7 h!8 v!vvyh v !9!T r)!!! rqvr.pyh T H H H T 1$%`V 88 T J2 Q`R1J: 1QJ ]CQ Q` ]QC7H.:V V :VIGC:$V :HHQ`R1J$ Q VR1IVJ HC:V^ Q :5 _7:JR^Y_^] QHQJ VJ Q`:JR_5I%RR7:JR^IY_^ T Q_50V`7 :JR7I%R^02Y_^ TQ_5:JR7I%R^2Y_^T Q_:JRI%R^2Y _^\ Q_ QV0:C%: V1.1H.$`:R1VJ 1JVJ01`QJIVJ :C0:`1:GCV^C: 1 %RV5CQJ$1 %RV5RV] .:JR IVR1:J $`:1J 1 64 2 TEn80 1,5 TEe80 TEg80 SAd100 TEd40 SAd115 SAh80 1 TEe50 TEp20 TEo30 SAb80 TEa80 TEl40 SAi80 TEo80 TEn20 TEm30 SAn30 SAn50 SAn80 SAm40 TEf40 TEb40TEb60TEd60 TEm80 TEc80 TEn5 Long TEg30 GTa40 TEf60 SAi100 0,5 TEi80 SAg80 SAl30 TEn40 MTz60 TEe10 TEo10TEc10SAm5TEm10 SAe80 GTc60GTe60 TEh5TEd5 TEa10TEg10 TVb110TEl60 TEf5 SAh10 SAm20 GTd20 TEa50 TEa60TEi50TEc50GTa60TEg50 TEo60TEp60 SAi5 TEp5 SAe30TVg30 SAi60TVa90GTc70 TEb5SAc10 SAf10 SAg5SAe10 SAi20 SAl50 TEp40SAh50SAc40TVb80SAl80TEo100TVg90 SAl10 SAn10 SAd20SAd30SAa20SAh30 GTc30GTd30SAd40 GTc50 SAm60SAc60TVf90 SAa5 GTe40GTa20 GTd60SAe50SAd60GTb70SAf60TEm50 SAb10 Med. SAb30GTb30SAi40TEh40 SAa40 Depth SAa60SAf100 TEh60 SAa100 SAf20 GTc20 SAg40SAf40GTe30 GTb40SAg100 -2 -1,5 -1 -0,5 GTd40 0,5 SAb501 GTd70 TVf80 1,5 2 2,5 SAg60 GTc40TVb60 GTd50TVe50TVg60 GTb20 TVf40TVh40 TVc50TVa30 TEo50 TVc90 GTe20 TVc30 TVg50GTe70TVf60MTc60TVe70SAd80TVd80 TVh80 TVb40 TVg70 GTe50TVh60GTb60 TVd20 MTc20 TVd40 MTd40 MTh60TEn60 -0,5 MTy40 TVe30 MTz40 GTb50TVc70TVa50TVd60 TVa70 TVb20 6v!! TVh20 MTh30MTh40 MTa60 TVb5 TVf20 Lat MTd50 MTb60TVh110MTy60 TVf5 MTh20 TVe90 TVc10 MTg50 TVd5 TVh5 MTh50 MTe50MTe40TVf110 -1 MTf40 MTd60TVd110MTc50 TVa10 MTe60 MTc40 MTb50 TVg10 MTg40MTa50MTy50 TVe10 MTb40 MTf50 TVe110 SAc80 -1,5 MTz50 MTc30MTa40 MTf60 -2 MTg60 -2,5 -3 Axis 1 1$%`V 88 T33$Q`R1J: 1QJ]CQ Q`]QC7H.:V V:VIGC:$V`Q%JR1JV:H.:I]C1J$ : 1QJ ^ : 1QJ]Q1J _84`VVJ:``Q1`V]`VVJ .V`Q%`VJ01`QJIVJ :C`:H Q`^C: 1 %RV5CQJ$1 %RV5 RV] .:JRIVR1:J$`:1J1 4 Sccil 3 Nlong MfxScinFd Preh Lsp 2 Sboc PslcfAxc Lh Ssp Eo Lfr Csp1 Af EufElumHpanHlPhybPlamSechSclm Pmax Phymo Schr Da Schc EaPxa Cd Ocu Phf AbrMoc Epen DrLhy Mf Nar Scfo Site Sccant 1 Prste Aox Sdec Pos Goe Lpx Mk Spkk Scsq Phl Eh Ai Hlu Phyli Msp1CsoyNehy Mdb Ld GcOsp Prca Ab Long Pp Ssc McpMsp2PscCgPepu Neh Nein Cs Lfu Schn Pkf Agra Vinf Pvr Smuo Nlat Poe Ah Scpa MtPhn EvitNlin Htub Ccir Mfr Ltet Php Acu Aa LYn Med. Prci Llat DepthDf Ppp Gom -1,6 Sctr -0,8 PsarSpbx Nab 0,8 1,6 2,4 3,2 4 4,8 Sigm Ap Epal PhmLc Neu Mpc Ac SctcstNel Hps Pcr Tstr Oq SfilStbo Amu Ssp Hf Sigsq Ccly Lat Lem AmGu Pxg Per Nlum Cc -1 Mocu Stlim Mpl 6v!! Prml Agro Dg Ptr Owf Ncch Prsxo Mbl Oer Cm Phyl Pau Mgt Pko Bv EUaPpctStyer Psp -2 Pslcc Hsp Lgr Scty -3 Lcc Necr -4 Ag Psp Msa CocchiGaGn Cb Agr Lg Pf -5 Mgl ElPolya -6 Axis 1 1$%`V 88 T 33$ Q`R1J: 1QJ ]CQ Q` ]QC7H.:V V ]VH1V HQCCVH VR ^]VH1V ]Q1J _8 4`VVJ :``Q1`V]`VVJ .V`Q%`VJ01`QJIVJ :C`:H Q`^C: 1 %RV5CQJ$1 %RV5RV] .:JRIVR1:J$`:1J 1 65 $:GCV 88 T 3Q``VC: 1QJGV 1VVJVJ01`QJIVJ :C0:`1:GCV:JR1 VHQ`V``QI33$ :J:C71 3Q``VC: 1QJGV 1VVJVJ080:`1:GCV:JR1 VHQ`V "J018$61 "J018$61 "J018$61 V] . 5 R5 R5 2VR8 R5 R5 5 : 5 5 5 QJ$ R5 R5 R5 J RV :1C5:VIGC:$V:I]CVR: :JRIQ`RV] .QJ:JR7%G `: :^QJ .VCV` 1J .V]CQ _:`VV]:`: VR1J Q 1Q$`Q%].1$.C1$. 1J$I:1JR1``V`VJHVGV 1VVJ .V ]VH1VHQI]Q1 1QJ`Q%JR1J Q`0:1:J1H::JR .QV`Q%JR1J:G:%R1::JR V``:H1J: ^&1$8 88 _8 1 . .V 1JH`V:V 1J RV] . :JR .V RVH`V:V 1J IVR1:J $`:1J 1 I:1JC7`Q%JR:IQJ$]VH1V`1H.JV5J%IGV`Q`1JR101R%:C5V_%1 :G1C1 71JRV6^ 5"5 :JRE;_:JRRV] .:JRIVR1:J$`:1J1 66 $:GCV 88 T:1`11V]V:`I:J`:J@HQ``VC: 1QJGV 1VVJVJ01`QIVJ :C0:`1:GCV:JRG1Q 1H R10V`1 7`Q` .V1.QCV %R7:`V:8 JGQCR:`V`Q` : 1 1H:CC71$J1`1H:J HQ``VC: 1QJ8 A 1J` V] . (VR8 : QJ$ 5 5 5 A R5 R5 R5 5 5 5 R5 1J` 5 5 5 5 5 V] . R5 R5 R5 5 R5 5 (VR8 5 5 5 R5 5 R5 R5 : 5 5 5 5 5 5 R5 R5 QJ$ R5 R5 5 R5 R5 R5 R5 5 R5 $CCVJ01`QJIVJ :C0:`1:GCV1V`V%VR1J .V< ,R" :J:C716 .V`V%C 1J$I:61I%I ]V:`IVJ HQ``VC: 1QJ ^` _ 1V`V `Q%JR `Q` RV] . :JR IVR1:J $`:1J 1 `:%J:C]: V`J8 V%C 1J$0:C%V:`V1V1$. VRG7]V:`I:J`:J@HQ``VC: 1QJHQV``1H1VJ ^` _8 . !"#$"%&'( ) 34! 5033+67$8&901 ! . ! !"#$"%&'( ) ! *+,,- ) ! ! !! *+,., ) 0! ! "!#$%&'! !! "%! ! 0# ! (!)$*+! "! "#( ! 0!0# ! ,!-.&! !! "# ! 0" ! /!-012 ! "! ""! ! " ! ! ! "! ! !! "! !%$ ! 0"0# ! #! !$' ! 6yy ! !! (" ! !0" ! 67 ]: 1:C R10V`1 7 ]`VR1H 1QJ I:] `VR1H 1QJ I:] Q` QIV HQII%J1 7 1JR1HV ^"5 :JR E;_5 `V:C1 68 1$%`V 88 T`VR1H 1QJI:]Q`]VH1V`1H.JV^"_1J .V`10V %R7:`V:72 ^1J .VI:CCV ]1H %`V_5 5$5 ":JR4 8 69 1$%`V 88 T`VR1H 1QJI:]Q`R10V`1 7^ _1J .V`10V %R7:`V: 72 ^1J .VI:CCV ]1H %`V_5 5$5 ":JR4 8 70 1$%`V 88 T`VR1H 1QJI:]Q`1VCQ%V0VJJV^E;_1J .V`10V %R7:`V: 72 ^1J .VI:CCV ]1H %`V_5 5$5 ":JR4 8 71 8 ]VH1V HQCQ$1H:C :J:C71 V%C .1$.C1$. .: .V ]QC7H.:V V :VIGC:$V `%H %`V :JR .V1` R1 `1G% 1QJ ]: V`J:CQJ$ .V : 1%IHQJ 1JVJ :C.VC`I:1JC7`V]QJRVR Q 1QVJ01`QJIVJ :C `:H Q`5RV] .$`:R1VJ :JRVR1IVJ V6 %`V5:1VCC: .V]: 1:C0:`1:G1C1 71::CQ 1J`C%VJHVRG7C: 1 %RV:JRCQJ$1 %RV8 JQ`RV` Q1RVJ 1`7 .V]VH1V:QH1: VR11 .Q`1JR1H: 10VQ`$`Q%]Q` : 1QJ5`10V $`Q%] RV`10VR ``QI Q : HC:1`1H: 1QJ Q` VR1IVJ :JR 16 ``QI RV] . `:J$V HC:1`1H: 1QJ5 JR1H: Q` ]VH1V $J:C71 ^ $_ .: GVVJ ]V``Q`IVR8 V%C RVIQJ `: V .: QIV ]VH1V R1 `1G% 1QJ RV]VJR QJ RV`1J1 V $`:J%CQIV `1H H.:`:H V`1 1H Q` .V VR1IVJ : 1VCC : RV`1J1 V RV] . `:J$V :JR .V 1JR1H: Q` 0:C%VQ` $ : 1 1H]VH1`7 .V$`Q%]Q` : 1QJ .: :`V:QH1: VR11 . .V]VH1V 11 .R1``V`VJ CV0VCQ`1$J1`1H:JHV^ $]]VJR16 _ ]QC7H.:V V .:0V : 1 1H:CC7 1$J1`1H:J :QH1: 1QJ 1.1 RV`1J1 V $`:J%CQIV `1H H.:`:H V`1 1H1.1CV 11 .RV] .`:J$VVCVH VR8"6HC%10VC7]VH1VR1]C:7:J :QH1: 1QJGQ .11 .VR1IVJ :JRRV] .`:J$V8 .V$`V: V J%IGV`Q` .V]VH1V `Q%JRVVIJQ QGV `QJ$C7:QH1: VR Q]VH1`1HVJ01`QJIVJ :C0:`1:GCV8 JRV :1C5 ]VH1VVVIJQ Q.:0V `QJ$`VC: 1QJ.1]11 .RV`1J1 VVR1IVJ V6 %`V:JR 11 .RV] .`:J$V8 J &1$%`V 88R .V R1 `1G% 1QJ ]: V`J Q` QIV ]VH1V :HHQ`R1J$ Q `V%C QG :1JVR1J .1 %R71V`V1CC% `: VR61 11J V`V 1J$ Q]Q1J VRQ% .VHQJ1 VJH7 GV 1VVJ`V%C Q` $:J:C7V:JR .V]`VR1H 1QJR1 `1G% 1QJI:]Q`QIV]VH1V8 72 1$%`V 88 T`VR1H 1QJI:]Q` #]QJ%].1 G1C1JV: : R1 `1G% 1QJ:CQJ$ .V`10V %R7:`V:7 5$5 ":JR4 8 73 1$%`V 88 T`VR1H 1QJI:]Q` #]QJ%].1 `:%0VC1 R1 `1G% 1QJ:CQJ$ .V`10V %R7:`V:7 5$5 ":JR4 8 74 1$%`V 88 T`VR1H 1QJI:]Q` .:V Q 75 1$%`V 88 T`VR1H 1QJI:]Q` C7IVJ%`: CV1Q]7$Q R1 `1G% 1QJ:CQJ$ .V`10V %R7:`V:2 8 76 1$%`V 88 T`VR1H 1QJI:]Q` .1 `%]: :`1V 1J: R1 `1G% 1QJ:CQJ$ .V`10V %R7:`V:7 5$5 ":JR4 8 77 1$%`V 88 T`VR1H 1QJI:]Q` .`1CQJV`V1 `1C%I R1 `1G% 1QJ:CQJ$ .V`10V %R7:`V:72 ^1J .VI:CCV ]1H %`V_5 5$5 ":JR4 8 78 1$%`V 88 T`VR1H 1QJI:]Q` $%J1HV 01 : : R1 `1G% 1QJ:CQJ$ .V`10V %R7:`V:2 8 79 1$%`V 88 T`VR1H 1QJI:]Q` V V`QI: % `1C1`Q`I1 R1 `1G% 1QJ:CQJ$ .V`10V %R7:`V:72 ^1J .VI:CCV ]1H %`V_5 5$5 ":JR4 8 80 1$%`V 88 T`VR1H 1QJI:]Q` VC1JJ: ]:CI: : R1 `1G% 1QJ:CQJ$ .V`10V %R7:`V:7 5$5 ":JR4 8 81 1$%`V 88 T`VR1H 1QJI:]Q` V :7H.1 $Q Q1 R1 `1G% 1QJ:CQJ$ .V`10V %R7:`V:72 ^1J .VI:CCV ]1H %`V_5 5$5 ":JR4 8 82 1$%`V 88 T`VR1H 1QJI:]Q` VI: QJV`V1 %J1HQ`J1 R1 `1G% 1QJ:CQJ$ .V`10V %R7:`V:72 ^1J .VI:CCV ]1H %`V_5 5$5 ":JR4 8 83 1$%`V 88 T`VR1H 1QJI:]Q` Q .`1: HQJH.7CV$: R1 `1G% 1QJ:CQJ$ .V`10V %R7:`V:7 5$5 ":JR4 8 84 1$%`V 88 T`VR1H 1QJI:]Q` 'J%].1 V`VI1 : R1 `1G% 1QJ:CQJ$ .V`10V %R7:`V:7 5$5 ":JR4 8 85 1$%`V 88 T`VR1H 1QJI:]Q` '1VJ1: `%1`Q`I1 R1 `1G% 1QJ:CQJ$ .V`10V %R7:`V:7 5$5 ":JR4 8 86 1$%`V 88 T`VR1H 1QJI:]Q` VH 1J:`1: :%`1HQI: R1 `1G% 1QJ:CQJ$ .V`10V %R7:`V:72 ^1J .VI:CCV ]1H %`V_5 5$5 ":JR4 8 87 1$%`V 88 T `VR1H 1QJ I:] Q` 1V`VGCC1RV `QVI1 R1 `1G% 1QJ :CQJ$ .V `10V %R7 :`V:7 2 ^1J .V I:CCV ]1H %`V_5 5 $5 " :JR 4 8 88 1$%`V 88 T `VR1H 1QJ I:] Q` % V`J:]1 H% : : R1 `1G% 1QJ :CQJ$ .V `10V %R7 :`V:7 2 ^1J .V I:CCV ]1H %`V_5 5 $5 " :JR 4 8 89 : : Q` Q` RV:GVR ]VH1V 1J.:G1 1J$ .V `10V 1J0V 1$: V :`V: :CQJ$ .V : 1%I HQJ 1JVJ :C .VC` 1V`V `%JH 1QJ:C Q H:``7 Q% : 11RV` %R7 :1IVR : :J:C7 90 $:GCV 888 %II:`7Q``V%C Q` JR1H: Q`]VH1V$J:C71^ $_:J:C7V``QI .V]`VVJ %R7:JR .QV1CC% `: VR1J :Q` : V :C8 ^ _8QC7H.:V V]VH1V:`V:QH1: VR11 . $`Q%] Q` : 1QJ5 :HHQ`R1J$ Q VR1IVJ HC:V :JR RV] . `:J$V5 11 . R1``V`VJ CV0VC Q` 1$J1`1H:JHV: 1QH:CVQ`1J0V 1$: 1QJ5``QI`V$1QJ:C QJ: 1QJ:CH:CV8 ! #R& ' # !# #R #$ # !# ]QJ%].1 G1C1JV: : IY02^RI_ I ]02^ R I_ ]QJ%].1 G`VIVJ 1 I^RI_ I \02^R I_ ]QJ%].1 `:%0VC1 I I\ 0( ^R I_ :%CCV`1VCC: H:]% VQH1 I 02^R I_ C1IVJ%`: CV7Q]1$Q ^RI_ ^RI_ 1 `%]: :`1V 1J: 2^RI_ 02^R I_ %J1HV 01 : : I 0( ]2^R I_ 7:C1JQVH1: %G1HQC: R 0252^5] _ VC1JJ: ]:CI: : I ]02R2^RI_ 0( \2]2^R] I_ V :1H.1 $Q Q1 ^ R I_ 0( ]2^ RI_ Q .`1: HQJH.7CV$: I ]^ R 6RI_ I^R5RI_ J%].1 V`VI1 : RI^ RI_ ^RI_ 1VJ1: `%1`Q`I1 RI^ RI_ ]I^RI_ VH 1J:`1: :%`1HQI: 0( ]2 I ]02^ R R I_ V`J:]1 H% : : 0( ]252 0252\ (^ R 5\ _ V%C QG :1JVR ``QI .V :J:C71 Q` R10V`1 7 R1 `1G% 1QJ ]: V`J Q` ]QC7H.:V V :VIGC:$V :JR ]VH1V : : `V$1QJ:C ^ : 1%I 3QJ 1JVJ :C .VC`_5 `VC: VR Q QIV VJ01`QJIVJ :C`:H Q`^RV] .5VR1IVJ $`:1J1 91 vqvvqhy /:: ,;: ,:: (;: (:: ";: ":: ;: : HU! HU& HU! HU# HU& HU" HU& HU! HU" HU$ HU! HU( HU! HU" HU$ HU! HU! HU# HU& HU! HU! HU" HU$ HU( HU! 7 7HU" 7HU# 7HU$ 7 s9 s9HU" s9 s9HU$ s9 9HU! 9 9HU# 9 9HU& 9HU( 9 9HU! 6!HU ! 6!HU 6! # 6!HU $ 6!HU 6! 6( 6( # 6(HU 6( & 6(HU 6 # 6 #HU" 6 #HU# 6 #HU$ 6 #HU& 6 # 6 #HU! 6 # 6 6!! " 6!!HU 6!! $ 6!!HU 6!! ( 6!!HU 6!! 6!! ! 6!&HU 6!& # 6!&HU 6!& & 6!&HU 6!& 6!&HU! 6!& hh ":: =: <: /: (: : HU! HU$ HU! HU# HU& HU! HU# HU& HU" HU# HU& " HU $ HU & HU HU! HU! HU# HU( HU! " HU & HU HU! ! HU $ HU ( HU 7 7HU" 7HU# 7 7HU& s9 s9HU" s9 s9HU$ s9 9 9HU" 9 9HU$ 9HU& 9 9HU! 9 6! " 6!HU 6! $ 6!HU 6! ! 6(HU 6( 6( $ 6(HU 6( 6 #HU ! #HU 6 # 6 # #HU 6 # 6 # 6 ( #HU 6 # 6 #HU! 6 ! 6!!HU 6!! # 6!!HU $ 6!!HU 6!! ( 6!!HU 6!! 6!!HU! 6!& " 6!&HU # 6!&HU 6!& & 6!&HU 6!& 6!&HU! 6!&HU! rpvr ,:: (;: (:: ";: ":: ;: : HU! HU$ HU! HU# HU! HU# HU& HU! HU# HU$ HU! HU# HU& HU( HU! HU# HU& HU! HU" HU$ HU! HU! HU$ HU( HU! 7 " 7HU # 7HU 7 & 7HU s9 s9HU" s9 s9 s9HU& 9HU! 9HU" 9 9HU$ 9 9HU( 9 9HU! 6! " 6!HU 6! $ 6!HU & 6!HU 6( " 6(HU 6( $ 6(HU 6( 6 # 6 #HU" 6 # 6 #HU$ 6 # 6 # 6 #HU! 6 # 6 6!! # 6!!HU 6!! & 6!!HU ( 6!!HU 6!! 6!!HU! 6!& " 6!&HU # 6!&HU 6!& & 6!&HU 6!& 6!&HU! 6!& 6!!HU ! 6!!HU QPG`8C6@U6 HPGGVT86 8SVTU68@6 @8CDIP9@SH@U6 1$%`V 88 T:`1: 1QJQ`QC7H.:V :^GC%V_52QCC%H:^$`VVJ_53`% :HV:^7VCCQ1_:JR"H.1JQRV`I: : ^`VR_ :6: Q0V` 1IV51J V`IQ`J%IGV`Q`1JR101R%:C5 :6: :JR]VH1V1J .V : 1QJCQH: VR1J1RV :JRQ% 1RV:R`VR$1J$:`V:: 2QJ :C QR13: `Q5GV`Q`V5R%`1J$:JR:` V`R`VR$1J$:H 101 1V8 92 8 ! .V:J:C7 ]VH1V`1H.JV^ :JR"_1JR1H: VRC1 CVR1``V`VJHVGV 1VVJCQ1V`1J``:C1 Q`:C:JR %]]V`H1`H:C1 Q`:C R1 `1G% 1QJ^RVH`1GVRG7 :JR 1JR1HV_I1``Q`VR]: V`JRVH`1GVR`Q`]VH1V `1H.JV :JR 1JR101R%:C :G%JR:JHV 1J V:H. :`V: ]Q1J 1J$ Q% 5 1J :RR1 1QJ5 QIV RV] . Q` VR1IVJ HC:V 1.V`V .V .1$. ]V`HVJ :$V Q` `VC: 10V ]`VVJHV Q` `:`V ]VH1V HQJ `1G% VR Q 1JH`V:V R10V`1 7 ^V8$8 I : 2QJ :C Q R1 3: `Q5 I : :G:%R1:5 0V`7R:JR7 I%R VR1IVJ : Q`0:1:J1H: :JR V``:H1J:5 :JR7RI%R 1J :G:%R1:_8 :`V]VH1V51J V`IQ``V `1H VR]: 1:C `:J$V :JRLQ` CQ1 :G%JR:JHV5 `V]`VVJ QJV Q` .V `V1 $VJV`:C VHQCQ$1H:C `VC: 1QJ.1] :]]C1VR Q I:`1JV VJ01`QJIVJ 5HQJ `1G% 1J$HQJ1RV`:GC7 Q .V Q :CJ%IGV`Q`]VH1V:JRR10V`1 7Q` :$10VJ:`V:^ V0: Q% V :C8 5 _8 93 0:`1:G1C1 7 1J ]VH1V HQI]Q1 1QJ :IQJ$ :I]C1J$ %J 1 `Q`:$10VJ:`V:: :$10VJ ]: 1:CH:CV8 JRVVRGV :R10V`1 7RV]VJRQJ .VH:CV%VR QRV`1JVCQH:CQ`.:G1 : R10V`1 7^ Q`V:%5 _:JRR1``V`:IQJ$GVJ .1H :6QJ QI1H$`Q%]^ "CC1J$VJ56 "CC1J$VJ:JR4`:75 _82Q GV :R10V`1 7 %R1VHQJHVJ `: VQJC: 1 %R1J:CHC1JV ^_%1`Q 94 :`V: : R1]C:7VR V6HC%10VC7 `Q` :G:%R1: :JR V`` :H1J: ``QI Q I :JR `Q` Q`0:1:J1H:5:G:%R1::JR V``:H1J:5GV 1VVJ:JR I8 .V `VJRQGV`0VR:CQJ$ .VRV] .$`:R1VJ .Q1CQ10:C%VQ`]QC7H.:V V]VH1V `1H.JV:JRR10V`1 71J .V%]]V`1J``:C1 Q`:C 95 RVH`1GVR :J >1J VJ1 7 V``VH ? : :GQ% .V :IV RV ] . `:J$V8 .1 V``VH 1: V6]C:1JVRG7 .V]`VVJHVQ`I16VRVR1IVJ 1.1H.1JH`V:V .V]: 1:CHQI]CV61 7Q` .V%G `: : .: `V%C 1J:C:`$VJ%IGV`J1H.V]:HV:JRG7 .VQ`$:J1HVJ`1H.IVJ R%V Q .V`10V`R1H.:`$V1.1H.RV]Q1 VR1J .VV 96 Q` VR`1JV:JR:VIGC:$V^ 4:IG1 V :C8 5 63.1IVJ< V :C8 5 6H1]1QJV V :C8 5 _8 $CQ : 2QJ :C Q R1 3: `Q 8 QHV:J1H: GVR HQJ `1G% VR Q 1JH`V:V ]VH1V R10V`1 71J .VJV1$.GQ%`1J$ 97 $.:GVVJR1]C:7VR .1 %R71J$VJV`:CHQJ`1`IVR .V]VH1VR1 `1G% 1QJ]: V`J : 1VCC : .V :VIGC:$V HQI]Q1 1QJ QGV`0VR Q0V` 11RV R1``V`VJ :`V: Q` : 7``.VJ1:J V: 1J ]`V01Q% %R1V8 .% : HQJ 1J%%I Q` ]VH1V R1 `1G% 1QJ :CQJ$ 7``.VJ1:J HQJ 1JVJ :C .VC` H:J GV V6]VH VR8 ]VH1V R10V`1 7 I:] H:J `V]`VVJ HQI]`V.VJ10V]VH1VR10V`1 7R1 `1G% 1QJ]: V`J51JHC%R1J$`V%C QG :1JVR`Q` .V : 1%IHQJ 1JVJ :C.VC`:JR .QV`VC: VR Q .VJV1$.GQ%`1J$ 98 %G `: :1J:.Q` VI]Q`:C`:J$V8 .1].VJQIVJQJ1JQ`I:CC7QGV`0VR1JR`VR$VR %G `: : 1.V`V .V RV`:%J: 1QJ :CCQ1 .V Q]]Q` %J1 1H ]VH1V Q `Q`I RVJV ]Q]%C: 1QJ1J .V`1` ].:VQ` .V`VHQCQJ1: 1QJ]`QHV5`QCCQ1VRG7:J1JH`V:V1J .V J%IGV` Q` ]VH1V :JR 1JR101R%:C 11 . R1``V`VJ VHQCQ$1H:C .:G1 ^ 99 8 ! ! # $ / .V]`VVJ `VV:`H.5H:``1VRQ% 1J .V 7``.VJ1:JV:5:CQJ$ : 1%IHQJ 1JVJ :C.VC`5 .:]Q1J VRQ% .: 7 ! .V .`VVI:1JH: V$Q`1VV6]C:1J1J$ .VR`10V`Q`G1QR10V`1 7]: V`J1J .V 7``.VJ1:J HQJ 1JVJ :C .VC` VJ01`QJIVJ :`V7 G: .7IV `1H $`:R1VJ 5 $VQ$`:].1H:C `V: %`V5 `V]QJ1GCV `Q` .V JQ` .RQ% . $`:R1VJ 1J VJ01`QJIVJ :C HQJR1 1QJ5 :JR VJ01`QJIVJ :C .V V`Q$VJV1 7 ^18V8 $`:1J 1 100 .1 %R7.:Q% C1JVR .V1I]Q` :JHVQ` .VVJ01`Q JIVJ :CIQJ1 Q`1J$]`Q$`:IIV : : Q%`HV Q` 0:C%:GCV H1VJ 1`1H R: :5 %H. : `VCV0:J 1J`Q`I: 1QJ QJ ]QC7H.:V V ]VH1V HQI]Q1 1QJ Q` I:`1JV GVJ .1H HQII%J1 1V5 .V1` VHQCQ$7 :JR R10V`1 7 R1 `1G% 1QJ8 V]1 V .V2VR1 V``:JV:JV:1:`V$1QJQ`.1$.G1QR10V`1 7 .: `:J@ .1 G:1J :IQJ$ .VGV @JQ1JI:`1JV:`V:Q` .V1Q`CR5I%H.1Q`@`VI:1J QGVRQJV8$ `V$1QJ:C H:CV I:`1JV `VV:`H. 1J .V 2VR1 V``:JV:J 1 :H %:CC7 ]:` 1:C5 `V`CVH 1J$ ]:`VV``Q` :CQJ$ .VR1``V`VJ G:1J8 JH`V:VR: VI] :`V`V_%1`VR1J :6QJQI7 :JR:I]C1J$Q`]QQ`C7@JQ1JVHQ7 VI:1VCC:CQJ$R V`IIQJ1 Q`1J$]`Q$`:I :1IVR Q:J:C7V]VH1V:JR.:G1 : VHQCQ$78$ .V:IV 1IV5 .V1J0:1QJQ`:C1VJ ]VH1V 1.1H. HQJ :J C7 H.:J$V .V G1QR10V`1 7 Q` .V2VR1 V``:JV:J V: `V_%1`V HQJ 1J%Q%IQJ1 Q`1J$8 .::CQGVVJRVIQJ `: V .: .V]VH1V`1H.JV:JRR10V`1 7Q`QC7H.:V :^:: %``Q$: VQ` .VVJ 1`VI:H`Q 101 _%1 VHQI]CV6 QRV`1JV8$ .V:IV 1IV5 .V]`VVJ `V%C .:0V]Q1J VRQ% .: : $QQRIQJ1 Q`1J$]`Q=VH Q`.%I:J:H 101 1V]`Q01RVR: :%V`%C Q]C:J:J:RV_%: V % :1J:GCV %V Q` I:`1JV VJ01`QJIVJ `VQ%`HV :JR Q 1I]`Q0V `% %`V IQJ1 Q`1J$ :H 1QJ8 `Q VH 1QJ5 HQJV`0: 1QJ :JR % :1J:GCV %V Q` I:`1JV VJ01`QJIVJ `VQ%`HV :JR 1 :QH1: VR G1QR10V`1 7 JVVR :J 1J V$`: VR :]]`Q:H. GV 1VVJ H1VJ 1`1H`VV:`H.:JRI:J:$VIVJ 8$ `V$1QJ:CH:CVIQ J: 1QJ.:0V=%`1R1H 1QJ Q0V` .V1``VQ%`HV:JRH:J5 .V`V`Q`V5GV ]`QR%HVV``1H1VJ HQJV`0: 1QJIV:%`V ^3QJJQCC7 :JR Q%$.$:`RVJ5 _8 JRVVR5 G7 %R71J$ G1QR10V`1 7]: V`J : 1VCC .V1J`C%VJHVQ`QIVVJ01`QJIVJ :C`:H Q`^V8$8RV] .:JR$VQ$`:].1H0:`1:GCV_QJ R10V`1 7 : `V$1QJ:C H:CV5 1 1 V:1V` Q V :GC1. J: %`:C ``QJ 1V` Q GV %VR 1J I:J:$VIVJ `: V$1V ^ V :]V V :C8 5 _5 V :GC1.1J$5 `Q` 1J :JHV5 RV] . 1J V`0:C .: :`V]`1Q`1 1V`Q`HQJV`0: 1QJ8 : :``QIIQJ1 Q`1J$]`Q=VH R: :V H:JGV:Q%`HVQ`1I]Q` :J 1J`Q`I: 1QJ`Q` .V 1I]CVIVJ : 1QJ Q` "%`Q]V:J R1`VH 10V G7 : 1QJ:C 4Q0V`JIVJ ^V8$8 2:`1JV `: V$75 : V` &`:IV 1`VH 10V5 6:G1 : 1`VH 10V_ : 1VCC : QH:C $% .Q`1 1V ^<1QR10V`1 7 `: V$1V :JR $H 1QJ C:J_8 J :RR1 1QJ Q `%` .V` :I]C1J$ :JR :6QJ QI1H V``Q` 5 I%H. Q` 1.: `VI:1J Q GV RQJV `V_%1`V ``VV R1 `1G% 1QJ Q` :0:1C:GCVR: :``QIJ: 1QJ:C:JR`V$1QJ:C`VV:`H.1J1 1: 10V Q .V]%GC1H8 .111CC `:H1C1 : VR: :G:V%]R: V:JRVJ:GCVH1VJ 1`1HR1H%1QJ8$R1]C:7VRG7 3QCC V :C8 ^ _ I:`1JV %`0V7 :`V JQ :C1:7 :HHV1GCV : .V `V$1QJ:C CV0VC :JR5 1.VJ :0:1C:GCV5 R: : HQ0V`:$V 1 Q` VJ 1JHQI]CV V8 JR101R%:C `VV:`H. V``Q` 1J HQCC:GQ`: 1QJ 11 .1J 1 % 1QJ5 ^1JHC%R1J$ I%V%I5 H1VJ 1`1H 1J 1 % V5 $Q0V`JIVJ :JR 1J V`$Q0V`JIVJ :C :$VJH1V5 %J10V`1 1V_ : .V 1J V`J: 1QJ:C5 J: 1QJ:C5 :JR `V$1QJ:CCV0VCI% HQJ 1J%V Q:R0:JHVQ%`@JQ1CVR$VQ`I:`1JVG1QR10V`1 71J .V 2VR1 V``:JV:JV::JRJ:``Q1RQ1J .V%J@JQ1J8 102 & & ! $CGV` VCC1 485 3: :JVQ 285 VCC: 3`QHV 85 `:$Q 8 8 2:H`QGVJ .Q RVCCV QCV R1 3:]`:1:5 1:JQ:5 41$C1Q5 2QJ VH`1 Q541:JJ% `15"CG:^$`H1]VC:$Q QH:JQ_8J10V`1 ;RV$C1 %R1R14VJQ0:5 1 % QR1H1VJ 103 106 4:IG1 28385 41:J$`:JRV $85 &`:H.V 1 8 8 QC1H.V 1 RV1 `QJR1 IQG1C1 RVC 4QC`Q R1 QC1H: `Q ^2:` 1``VJQ 2V`1R1QJ:CV_7R1 `1G%<1QJVV`%QCQR1:CH%JV]VH1V8$ 1 3QJ$`VQRVCC:8 8"85 :]QC15RV VIG`V 8 1` 107 EQV`QJ $8<85 6:JVJ E8 88 8 ]VH1V `1H.JV Q` GVJ .1H I:H`Q`:%J: 1J :J1. V %:`1V :JR HQ: :C :`V:8 ^I:JH:JQ1`1`V`1IVJ 1G1GC1Q_ EQ 8 8"J `Q]7:JRR10V`1 78 '1@Q 5 7 T 8 &:%H.:CR=85E%I:`8$8 8 .VR1V Q`1Q`I7: %R7Q`]QC7H.:V V`VVR1J$$%1CR8 'HV:JQ$`8 :`8 21QC8#JJ%8 &V08 7 R 8`$CC:J6:JHQH@&Q%JR: 1QJ5J10V`1 7Q`Q% .V`J3:C1`Q`J1:5 Q$J$VCV53$:JRV]:` IVJ Q` ,HV:JQ$`:].75J10V`1 7Q`:.1J$ QJ5V: CV5$a8 =: 108 V$VJR`V85 V$VJR`V 8 8 %IV`1H:C"HQCQ$75JR"J$C1.VR8"CV01V`7 ]]8 V01J 8$85" V` 8E85 V628$854QQR:7$8E88"J01`QJIVJ :C1J`C%VJHVQ``V$1QJ:CRVV]RV:]VH1VR10V`1 78 1.V #JJ%:C &V01V1 Q` $HQCQ$75 $0QC% 1QJ5 :JR %7 VI: 1HI 5 7 R 8 Q1:285 ::CCV85 : :J<1 85 :Q` :<85 :`$:1285 1HQCV 1 8 8"``V 1RVCR`:$:$$1QR1:GG1V`VC1 V%C ]Q]QC:IVJ Q:]QC1H.V 11J%J;:`V::CC:`$QR1$J<1Q^ 1``VJQHVJ `:CV_8 21QC8 :`8 VR1 85 ^_7 R QIQC1JQ 288 8 "HQCQ$7; IQ $VJV`:C5 7V ]`Q V:J ]: V`J7 .V ]VH1VR:`V: `VC: 1QJ.1]8 :Q%`J:C Q` 21Q$VQ$`:].7 5 7 R 8 Q`V:%288$`VHQII%J1 1V: %`: VR-,J .V`VC: 1QJ.1]GV 1VVJ:C].:5GV ::JR$:II:R10V`1 78 $HQC8 !V 85 7 R 8 2:`$:CV` 8 8=V7"J01`QJIVJ 7V V`J2VR1 V``:JV:J8 V1Q`@7V`$:IQJ`V8 ]8 2:`<1:CV 1854:GVCC1J1285 :Q` :<85 : :J<1 85 ::CCV85:$:JVCC185:J`1C1285 :`$%1285 1HQCV 1 8 8 `VR$1J$ :H 101 1V `Q` GV:H. JQ%`1.IVJ 1J :J :`V: Q``.Q`V 2QJ :C Q R1 3: `Q ^ _7 V``VH QJ QC7H.:V V HQII%J1 78 21QC8 :`8 VR1 85 ^_7 R 8R 2:`<1:CV 185 1HQCV 185$`R1< 109 Q : 8E848 8 VR1IVJ Q` .V 1V V`J 4%1:J: .VC`8 V]Q` Q` .V ,`1JQHQ .VC` "6]VR1 1QJ5 8 VJRVRVC1J$VJ 0:J RV !:JRGQ%1.Q$VH.QQC V :$VJ1J$VJ 5 7 ]]8 7G:@@VJE88 8 82:`1JV<1QCQ$77$J"HQCQ$1H:C$]]`Q:H.8 :`]V` :JR &Q1 5 V1Q`@7 ]]8 ,HH.1]1J 1R$IG`Q$1$852:`H.1J1$853:J QJV4853: VCC1$853.1IVJ<3853Q`I:H1285&`Q$C1:385&%`J:`14854:IG1 28385 41:HHQJV 485 41:J$`:JRV $85 4`:01C1 385 2: `Q Q :`Q &85 2:<<1Q 1 385 ,`1R VC1J1 85 1`:1JQ 8 8 $C1VJ ]VH1V:CQJ$ .V :C1:JHQ: 7:JQ0V`01V18 21QC8 @J0:1QJ 5 7 T 8 ,C$:`R &85QIV``1VCR8E88%``Q$: V1J GVJ .1H1J0V 1$: 1QJ8.1H. :6QJQI1H%J1 - :Q%`J:C Q` #_%: 1H $HQ7 VI % `V :JR &VHQ0V`7 5 7 R8 ,C$:`R &85 <`: V$:`R 85 6QC .V 8 8 QC7H.:V V : %``Q$: V `Q` I:`1JV G1QR10V`1 77 CQ1V` :6QJQI1H `VQC% 1QJ:JR1JR1H: Q`$`Q%]8 21QR10V`8 QJV`0 85 7R 8 ,%$"85 E=85 7$$<85 8 VC: 1QJ.1]GV 1VVJQ` GQ QII:H`Q`:%J::JR]QC7H7HC1H:`QI: 1H.7R`QH:`GQJ ^$6_``QIIVC V`R1H.:`$V1J Q`1V$1:J`=Q`R:JRHQ: :C1: V`8 :`8 $HQC8 `Q$`8 %V` 85 7 R 8 :JHV`18 83: :CQ$QRV$C1$JVCC1R154V1`1`V1V %`GVCC:`1VR :C1:8$ 1QH8H18J: 821C:JQ5 7R 8 V:`QJ 868 8 &%JH 1QJ:C $`Q%] VHQCQ$7 1J Q` VR1IVJ I:`1JV GVJ .Q7 .V `QCV Q` G1Q %`G: 1QJ8 'HV:JQ$`:].7 :JR :`1JV 21QCQ$77 :J #JJ%:C &V01V1 5 7R 8 V:`QJ 8685 QVJGV`$ 8 82:H`QGVJ .1H%HHV1QJ1J`VC: 1QJ QQ`$:J1HVJ`1H.IVJ :JR]QCC% 1QJQ` .V I:`1JVVJ01`QJIVJ 8 'HV:JQ$`8 :`8 21QC8 #JJ8 &V0 85 7 R8 X`W E28 8 QJ: 1QJ8 4VJV`:C `V: %`V Q` Q`$:J1I1H :VIGC:$V 1J ]VC:$1:C :JR GVJ .:C8 `%H %`V :JR R7J:I1H Q` :VIGC:$V8 2:=Q` GVJ .1H :VIGC:$V8 ]VH1`1H GVJ .1H :VIGC:$V8 J7 =1JJV ,5 VR1 Q`8 2:`1JV "HQCQ$78 QJRQJ7EQ.J1CV7:JRQJ8]]8 T 5 T 5 T 8 X`WE82851H:`RE8 8 Q%0V:%I:J%VCRVG1QJQI1VGVJ .1_%VRVC:2V`2XR1 X``:JXV8 &VH8 1`:08 % J8 :`8 $JR8 5 ^ _7 8 1H:`RE8 8 VH.V`H.V_%:C1 : 10V%`CV G1QHQVJQ1I:`1JVRV%G `: IV%GCVR`:$:GCVRVC:`X$1QJ I:`V1CC:1V8 &VH%V1C RV 1`:0:%6 RV C: % : 1QJ :`1JV R;$JRQ%IV 5 7R 8 QH@C1J$ QJ 85 VCC 848 8 QC7H.:V V =V7 :6: `Q` I:`1JV VJ01`QJIVJ :C _%:C1 7 IQJ1 Q`1J$8 :`8 QCC% 8 2%CC8 5 7 R 8 QJ 1285$GG1: 1288_%:C1 7:VIVJ Q` `:J1 1QJ:C1: V`%1J$:GVJ .1HG1Q 1H1JRV67 .VH:V %R7Q` .V1:C::<:1QJ:^JQ` .V`J$R`1: 1HV:_8 #_%: 1H QJV`0: 1QJ Q` :`1JV :JR )`V.1: V` $HQ7 VI 5 7R 8 Q<:`RQI:H$8 83: :CQ$%VQ` .V]QC7H.:V Q%:JVCC1RQ` .V$R`1: 1HV:88 Q` .V`J:JR3VJ `:C$R`1: 1H8 #H : #R`1: 1H: 5 8^JJH1QJQCV]$_ _%1`Q 110 V6 28$8 8 3QII%J1 7 `%H %`V 1J .V RVV]RV: GVJ .Q8 1.V #JJ%:C &V01V1 Q` $HQCQ$75 $0QC% 1QJ5 :JR %7 VI: 1H 5 7R 8 1VRC 8 8&:%J::JR&CQ`:Q` .V2VR1 V``:JV:J7$7 VI: 1H2:`1JV4%1RV`Q`<1QCQ$1 :JR : %`V Q0V`8 6:IG%`$7V`C:$:%C:`V78 ]8 QVJGV`$ 85 111 QI:V 185V`1:"852V`H: :C1 85:J1852:`%Q85Q``VCCQ8 8"``VH Q`I:`1H%C %`VQJI:H`QGVJ .1H :VIGC:$V1J:1V V`J2VR1 V``:JV:J1 V8 :`1JV QCC% 1QJ 2%CCV 1J 5 7 R 8 QI:V 1 85 <:HH1 85 3VC1: 2:$JQ 285 V0Q 1 85 4VJJ:`Q 85 VQJ1 485 2:J<%V Q 85 2:`%Q 85 2V`H: :C1 85 2QJ :$J: 85 :`C:$`VHQ 85 V`1: "85 1V```:JHVH.1 485 Q``VCCQ 85 QI:JQ "85 %Q 85 1CVJ<1 85 1JVCC1 $85 Q`J:IGW$85 `:G%HHQ<85:J18VJ 1&8531HV`Q$828 8"HQR10V`1 <VR1 `1G%<1QJV]:<1QR VI]Q`:CVRVCCV HQI%J1 < I:H`QGVJ QJ1H.V R1 `QJRQ IQCCV RVCC;:`V: I:`1J: RVCC: VJ% : R1 3: VC]Q`<1:JQ8 J7 $HH:RVI1: RVCCV H1VJ 112 # 7 ]VH1V HQCCVH VR 1J .V :`V: Q` (QJ :C Q R1 !: `Q #( ' $#$ 6FG5HG6! I5JKEL) )>?<: )>@<: )>.<: )>A<: )>8<: )>*<: )>$<: )>B<: )>2<: )>'<: )>?;: )>@;: )>.;: )>A;: )>8;: )>*;: )>$;: )>B;: )>2;: )>';: )>?/: )>@/: )>./: )>A/: )>8/: )>*/: )>$/: )>B/: )>2/: )>'/: )>8,: )>',: )>8(: )>'(: Q 6htr.hqr h 6h $ ; 6uh rr.hpvs . 6p $ $ $ ! $ " $ $ !$ ! !$ $ " $ $ (;; 6uvprv.tr v 6t $$ $$$$ $ ;; 6p yyv.t ryhqvph 6t $ ; 6urypuhrh.h vv 6 $$ $ $ $ $ $ $ ! $ $ ",: 8hyyr vryyh.yvi hpuvv 6 $ ; 6uv.ivyvrhh. 6i !$ (; 6uv.i rrv. 6i !$ (; 6uv.shryv. 6s. $ ; 7 hqh.vyyh 7 $ $!$$ C: 8urr. ps! rh 8 $ ; 8urr. 8ppuv $$$$ $$$$$$ C: 8v u .i hpuvh 8i $ ; 8yr h.pyrhh 8py $$$$$ $$ /; 9vypv .tyhp 9t $ $ $ $ $ $ !$ $! "(: 9 vyr rv.svy 9s $$$!$$$ $$$$ $ D; @rr.yth @y $ ; @vpr.vhh @v # $ $ $"$ " "/: @u vr.h hqvyy @Vh ": Bypr h.hyih Bh $$$ "; Bypr h.vp v B $ " $ $ "$ $ $ $ !$ $ $ $ $ $ $ ! ,:: Bypvqr. qhv B $$ ": Hhyt rvryyh.yyhh Cy $$ ": Ch ur.vvsr h. C $ ; Crr h.svyvs v Cs $ !$ $ $ $ $ $ $ # $ $ ! $ ! (;; Chyvrpvh.ivpyh Ci $ ,; Cq vqr.rqpvh C $ ; Ghvpr. ps .pv hh Gp $ $ $ $!$ $ $ !$ "(: Grvphvryyh.tyhi h Gq $$ (: Grvrvh.t hpvyv Gt $$$ "; Gi vr v.pppvrh Gpp ": Tpyrh.rhqviyhhhivv Gr $ !$ $ $ $ " $ $ $ $ $ $ $ $ $ $ $ ! #$ $ (C: Tpyrh.spuhyrv Gs $ ;: Gi vr v.t hpvyv Gt $$ $$ $ $ <: Gi vr v.yh rvyyv Gyh $ $ ! ! $ $ $ " # ! $ $ ! " $ (=: Gvqvpr. ps .vrh G` $ ; Hhtryyh. . H $$$$ (: Hhyqhr.tyrivsr Hty $!$!$$! =: Hhyqhr.h v Hh $ $$$ ,: Hh uh.iryyv Hiy $$ $$ $ $ $ $$!$ ":: Hh uh.xvir tv Hx $$ ,: Hryvh.hyhh Hy $ $ $ $ ! $ $ ! $ $ $ $ "<: Hrhpuv.tv Ht " $ # $ "$ $ ! ,:: Hvpryyvh.q i hpuvhyv Hqi $$$$$$$ ,; H vpuryr.pyhh. Hp $$$ ! $$ <; @rr.vph Hp $$$ "; Irhr rv.vp v Ir $ % $! $ ""; Iruu.pv h Irp "$! ;; Iruu.uir tv Iru %$ <; Iruu.u vpv Iru $$$$ $ $ <: Ivr.h vphh Ih $$$ (; Iu vh.ppuvyrth. Ippu $ "; Ih.hir h Ihi # $ $ $ !$ $ $ $ $ %$ "$ ,;: Ih.yhr vpr Iyh $ $ $ $ $ $ $! ! ""; Prvh.svs v Ps (: Qh hyhpqvh.h hqh Q $ $ $ $! $ $ $ $$ $ "": Qh hv.sytr Qs $$ ": Qh h vv. ps .vhh Q $$$$ $$ $$ $$$$ D: Qrpvh vh.h vph Qh $$$$$$ $ <; Qrpvh vh.x rv Qx $$ $ (; Qrhy p.r vpy Q ": Quyyqpr.yhvh Quy $$ ": Qvyh tv.r ph Q $$$ (; Qv v.r ph. Qr $ "; Qvh.p vhh Qp $ $ $ $ $$ $ =; Qrpvypuhr.r r Q $$ $$$ $ $ <; Qypv .h hvhp Qyh $ ; Qyuhy.vp Qp $ ; Q hvyyryyh.t hpvyv Qt $ $ $ $ $ $ $ ! $ "(: Hvv.pv vsr h. Q pv $ "; Q vv.ruyr v Q ru $ ; Q vv.shyyh Q y $$ ": Q vv.rr v Q r $ (; Q yh. Q $ ; Tpyh vpvh.vph Tp ! (: Tvthi h.rhpyhh Tvr $ ; Tv.svyvp v Tsvy $ ; Tv.yvpyhh T $ $ (: Tvpuhrr .ph Tpp $ $ $ ,; Tr hv.phh Tp $$$!$$$!$ C; Ur riryyvqr. rv U $ $ $ !"$ # $ $ $ $ "=: Wr vyvv.vsqviy Wvs $ "; E!&0&! =; (: "(: ""; /: D; /: C: (; C: ",; ":; "<; /,: =: =; (/: C: (C; (:; "=; "<: (:; "C; (D; ":: ,": "/: "=: ""; <; ,": ,D; "C: ;(C; 113 # 7 ]VH1V HQCCVH VR 1J .V :`V: Q` $Q`0:1:J1H: #( ' $#$ 6FG5HG6! I5JKEL) >MA; >M*; >MB; >M'; >M.": >M8": >M$": >M2": >MA(: >M*(: >MB(: >M'(: >M.,: >M8,: >M$,: >M2,: >MA/: >M*/: >MB/: >M'/: >M.;: >M8;: >M$;: >M2;: >MA<: >M*<: >MB<: >M'<: >M2<: >M.C: >M8C: >M$C: >M2C: >MA=: >M*=: >MB=: >M'=: >M.D: >M8D: >M$D: >MBD: >M2D: >MA"": >M*"": >M$"": >MB"": >M2"": >M'"": >MB";; >M2";; E&0& 6htr.hqr h 6h $ ; 6uh rr.hpvs . 6p $ $"#$ $$$ ",: 6p yyv.t ryhqvph 6t !$$$$" $$$ D; 6vqr.pruhyh. 6 $ ; 6urypuhrh.h vv 6 $ $$ $#$$$$ "": 8hyyr vryyh.yvi hpuvv 6 $ $$$$!$ <: 6uv.ivyvrhh. 6i $$'$ $ ";; 6uv.shryv. 6s.Csh $$ $$$$$!$ ""; 7 hqh.vyyh 7 " $ #$#$ &!$$ $ $ $ $ ,,: 8hvryyh.phvhh. 8p ""$$$$ ,<: 8hvh.vv. 8 ": 8v hy.pv h. 8pv $ ; 8yr h.pyrhh 8py !$#$$#$$ $$ "/; 9vypv .tyhp 9t $ ; 9 vyr rv.svy 9s ! $$ $$ <: Bypr h.phvhh. Bp $ ; Bypr h.vp v B $ $ $ $ $ $ $ $ $ " $ ! ! $ $ $ $ $ $ $ ,(: Bvhqh.hpyhh. B $ ; Hhyt rvryyh.yyhh Cy $ ; Crr h.svyvs v Cs & $ !$ ! !! $ $ $ $ !$ ! $ $ $ $ $" ,/: Chyvrpvh.ivpyh Ci $ $$ "; Ghvpr. ps .pv hh Gp $!!$$#$ "/; Grvphvryyh.tyhi h Gq $$ ,: Gi vr v.pppvrh Gpp ": Tpyrh.rhqviyhhhivv Gr $ $ $ $ $ !$ ! $ $ $ $ # $ $ $ ! $ $ $ $$$ (=; Tpyrh.spuhyrv Gs $$ $ (; Gi vr v.yh rvyyv Gyh $ #$ ! $ "$ &$ !$ $$ % ! $ $ $$ ==; Tpyrh.r h h. Gr $"!! =; Hhtryyh. . H $$$$$ (; Hhtryh. !! H! $!$"$$$$ $$ "/; Hh uh.iryyv Hiy !$#$$ $$$$$$$$$$ $ $ ";; Hrqvh.s htvyv. Hs %$ "$ $ $ $ ";; Hryvh.hyhh Hy ($ " $ $ $ $ $ $ $ ;(: Hrhpuv.tv Ht $ ; Irhr rv.vp v Ir $ $ (: Iruu.u vpv Iru $$ ": Iu vh.ppuvyrth. Ippu $ $ !%"$$"$$'$ ;:: Ih.hir h Ihi $ "$ "!$$ $ ! $!$ $ $ $ /,; Ih.yhr vpr Iyh !$ $ $$ !$"!!$ $ $ $ $ $ $ $ $ $ $ (,: Puv.r rvh. Pr #($"$!$$$ (:; Puv. ! P $ ; Prvh.svs v Ps "$# $ % " $ $ "$ $$ $ $ (<": Qh hyhpqvh.h hqh Q !$$ /: Qh hv!! Q ! '$ $ !%$" $$ ;/; Qh h vv. ps .vhh Q $ " !$ $$ ! # " $ %$ $ %$ ' $ $ # $ $ ;;; Qrpvh vh.h vph Qh (: Qrpvh vh.x rv Qx $!$ ,: Qvyh tv.r ph Q $$ $ (; Qvh.p vhh Qp $ ; Qrpvypuhr.r r Q !$! $$$$$ D: Q hvyyryyh.t hpvyv Qt !"$ $!!$ $ $ ";; Q vv.ruyr v Q ru $ ; Q vv.shyyh Q y "$ $ !"$ !$$ $ "D: 6vyhv.rpyhh. Q $ ; Q vv.rr v Q r $ ; Q q vyyrh.xrsr rvv. Qxs ": Tpyryrv.phhi h. Tpph $ ; Tpyryrv. vqrhh. Tp $ ; Tvthyv.huvyqhr. Tvt $$"$ $ $"$!$ $ $ (": Tv.qrp h. Tqrp $ $$!#"$#$ "<; Tv.svyvp v Tsvy $$ $# $ "(: Tv.yvpyhh T $$$$$$$$$ /; Tvvqhr!! T $$$$ ,: Tvuhr.ii. Ti $ !$#$$ ":: Tr hv.phh Tp $ " $$! ! $ $ $ (:; Turryhv.ih. Ti $ ; Turryhv.yvvpyh. Tyv $ ; Ur riryyvqr. rv U ! !$ $ $ $ $!$ $ !$ $ $ " !$ $ $ (/; E!&0&! (=; (=; (;; (/: "D: ;; ";(: "/;; /,; ;<: ;(; "/<; =; /(: (,; C: /=; (D: //: ,(: "": (<: ":: "D; (/: ""; /; "C; C; ;; "=: ,: /; ,: ": ,; (; (; ,; (: <: "": "; ": ; (; : ; ": ; ""<<; 114 # 7 ]VH1V HQCCVH VR 1J .V :`V: Q` :G:%R1: #( ' $#$ 6FG5HG6! I5JKEL) 6I.; 6I2; 6I9; 6IN; 6IA": 6I8": 6I$": 6IB": 6I'": 6I7": 6I1": 6I.(: 6I*(: 6IB(: 6I9(: 6IN(: 6IA,: 6I*,: 6I$,: 6I',: 6I7,: 6I1,: 6I./: 6I8/: 6I*/: 6IB/: 6I2/: 6I9/: 6IN/: 6IA;: 6I$;: 6I';: 6I7;: 6I1;: 6I.<: 6I8<: 6I*<: 6IB<: 6I2<: 6I9<: 6IN<: 6IA=: 6I8=: 6I*=: 6I$=: 6I2=: 6I'=: 6I9=: 6I7=: 6I1=: 6I.":: 6I*":: 6IB":: 6I2":: 6I9":: 6I*""; E&0& 6uh rr.hpvs . 6p $ "; 6pvh tv.uhhh. 6u ": 6p yyv.t ryhqvph 6t !$ (; 6iu .t hpvyv. 6t h #$ ;; 6vqr.hpvi hpuvhh. 6 $$$ "; 6urypuhrh.h vv 6 $$ "$ $$ $ $$$ ":; 8hyyr vryyh.yvi hpuvv 6 $$ ": 6uv.ivyvrhh. 6i $ "; 6uv.i rrv. 6i Ci $ & !$$$ $ $ $ &$ $ $ "$ $ /(; 6uv.shryv. 6s.Csh $$ (: 6 hiryyh.v vpy . 6v $$ $ $ /: 6 hqvh.pv uh. 6p $ ; 7 hqh.vyyh 7 $ "; 8hvryyh.phvhh. 8p $ ; 8uhrr ! ! 8 $"($ ",: 8urr. ps! rh 8 $$ ": 8u rhy.qrivyr. 8q $ ; 8yr h.pyrhh 8py $$ $ $$$ $$$! $ $$ "(: 8 h.r v. 8 !$ "$ $ " $ $ "/; 9vypv .tyhp 9t $ (; 9v h.h vrvh. 9h $"$ ;: 9 vyyrh.9 vyyrh. i vhh. 9 $ ; 9 vyr rv.svy 9s $ $$ $$$ $ $ $$ $ $ $ ""; @pyrr.hyr vhh. @hy $$ ": @vpr.uh hvv @u $ "; @vpr.rhh. @r $ $ ;: Bypr h.vp v B $ $ $$ $$ $$!$ $ $ $$ $ $ $ $ ";; Bvhqh.rr vh. Br $$$ "; Hhyt rvryyh.yyhh Cy $$$ $$ $ ;: Ch ur.vvsr h. C $ ; Crr h.svyvs v Cs $ $$ ! ! $$ $ $ $ ",: Chyvrpvh.ivpyh Ci $" $ ;: Cq vqr.rqpvh C $ ; Ghrvpr.u v. Gu $ ; Ghvpr. ps .pv hh Gp #$! $#$ $ $$ $ "<; Grvphvryyh.tyhi h Gq $$$ $ $ ,; Grvpu vqr !! G ": Gi vr vv.h hqh. G $ $ ! $ $ " $ $! $! $ $ $ $$ $ $ (/: Tpyrh.rhqviyhhhivv Gr $ #$ $$ D: Tpyrh.s htvyv. Gs $ ; Gi vr v.t hpvyv Gt $$$ "; Gi vr v.yh rvyyv Gyh $ ! !$ " #$ "$ $ !$ " &$ #! %$ $ ! % $ ' " $ ! $'! $ $ "C": Tpyrh.r h h. Gr $%&$"$ !$%$$$ $ /(; Hhtryyh. . H $$ (: Hhtryh. !! H! !$ $# !$ ($ $$ # # " $ $ ! $ $ $ 115 # 7 ]VH1V HQCCVH VR 1J .V :`V: Q` $V``:H1J: #( ' $#$ 6FG5HG6! I5JKEL) >GA; >G*; >GB; >G'; >G1; >G%; >G.": >G8": >G$": >G2": >GN": >G0": >G1(: >G%(: >G2,: >GN,: >G0,: >GA/: >G*/: >GB/: >G'/: >G7/: >G1/: >G%/: >G.;: >G8;: >G$;: >G2;: >G9;: >GN;: >G0;: >G.<: >GA<: >G*<: >GB<: >G'<: >G7<: >G1<: >G0<: >G%<: >G.=: >G8=: >G$=: >G2=: >G9=: >GN=: >G1=: >G0=: >G1":: >G0":: E&0& 6htr.hqr h 6h $ "; 6uh rr.hpvs . 6p $$$ $ $$ ;: 6uvprv.tr v 6t $ ; 6iu .t hpvyv. 6t h $$ ,: 6vqr.pruhyh. 6 (: 6urypuhrh.h vv 6 $$$$ (: 8hyyr vryyh.yvi hpuvv 6 $$ (: 6uv.ivyvrhh. 6i $$"$$$$ C: 6uv.i rrv. 6i Ci $ $ & # %$ $$ & ' ! $ # #$ $ ="; 6uv.shryv. 6s.Csh $ $ $ &$ !!$ $$! !$ !! $ $ " $ $ D(; 6 hiryyh.v vpy . 6v $$$$ ,: 6vuryyh.p vph. 6p $ ; 7 hqh.vyyh 7 $ ; 8hvryyh.phvhh. 8p $ ; 8hvryyh.tvh qv. 8t $ ; 8uhrr ! ! 8 $ $"!$! $$ $ $$ $ "/; 8urr. ps! rh 8 $$$$ (: 8u rhy.qrivyr. 8q $$ ": 8yr h.pyrhh 8py $$$ "; 8 h.r v. 8 $$ (: 9vypv .tyhp 9t $ ; 9v h.h vrvh. 9h $$ ,: 9 vyr rv.svy 9s $ $$ $ $$ =: @pyrr.yi vpvqr. @y $ "; @vpr.hrhh. @h $ ; @vpr.r rqv. @ $$ ": @vpr.rhh. @r $$ $ (; @vpr.vhh @v $ #$"$" $ $ ! (=; @u vr.syvh. @s $ "; Ayhiryyvtr h.qvypuhv. Aq $ ; Bypr h.vp v B $ "$$$$ C; Bvhqh.rr vh. Br ": Hhyt rvryyh.ywthv. Cy ! (: Hhyt rvryyh.yyhh Cy !$$ $$ ;: Crvr.yrqvqh. Ch $$ ;; Crr h.svyvs v Cs $ $ "!$$ $ $$ $ $ "/; Chyvrpvh.ivpyh Ci $$ $$$$$ ;; Cq vqr.rqpvh C ": Ghrvpr.u v. Gu $ ; Ghvpr. ps .pv hh Gp $$!$$ $$ $$ =: Grvphvryyh.tyhi h Gq $ $$ $$ /; Gi vr vv.h hqh. G $ $$$$$$$ $ $ C: Tpyrh.rhqviyhhhivv Gr $ "; Tpyrh.s htvyv. Gs ": Tpyrh.spuhyrv Gs "!' $ /"; Gi vr v.t hpvyv Gt $ ; Gi vr v.yh rvyyv Gyh "$ $ ! $ $ $ $$ ! $ $ ! $ ! $ $ $ /:: Tpyrh.r h h. Gr &$#$!$ !$#$ % $ $ $ ,": Gvqvpr.vrh G` $ ; Hhtryyh. . H $ ; Hhtryh. !! H! $$"$ #&" % & "$ # & % ! $ $ "=/: Hhyhppr .syvtv. Hs $ ; Hh uh.iryyv Hiy $ $ $$ ;: Hh uh.shyyh. Hs $ ; Hh uh.xvir tv Hx $ $$ $ <: Hrqvh.phrv. Hp "$$ $$ <: Hrqvh.s htvyv. Hs !##$($"#$# ! "$ $ $ ;:: Hryvh.hyhh Hy $$ $ (; Hrhpuv.tv Ht !! $$ $ $ $ $ $ $ $ $ $ $ "D: Hvpryyvh.q i hpuvhyv Hqi $$ $$!$ $ C: H vpuryr.pyhh. Hp $ $$$ $ <; Irhr rv.vp v Ir $ %!$$ $ # $ "<; Iruu.uir tv Iru $ $ $ $ $" !$ $!$ "<: Iruu.u vpv Iru $!$$$ $ ;; Iruu.vpvh. Irv ": Ivr.h vphh Ih $$$$ $$ $ <; Iu vh.ppuvyrth. Ippu $ "; Ih.hir h Ihi $ $ $ $ $%$ $#$ $ $ $ $ $ ,D; Ih.yhr vpr Iyh $ !$ $ & $ !$ "$ $ " $ !$ #$ !$ $ $ ,=; P ivvh.pvr v. Pp $$ (: Prvh.svs v Ps " "$"!$ $ $ $ $ ,C: Qhyyh.vpvyvrv. Qp !%$ (<; Qh hyhpqvh.h hqh Q ! ( $ #$##'$ $ $ # !$ $#$ $ $ $$ /D: Qh h vv. ps .vhh Q $#$ $ $ !$ $ $ $ $ $ "$ (/; Qrpvh vh.h vph Qh (: Qrhy p.r vpy Q $$ ": Qrh.vyyh. Qr $ ; Qur h.yh. Qu $ ; Quyyqpr ps! hpyhh Qui ! (: Quyyqpr.yhvh Quy $ ; Quyyqpr.yvrhh. Quyv ": Quyyqpr.hp uuhyh. Qu $$ ": Quy.yvtvph. Quy !$ (; Qvyh tv.r ph Q $ (; Qvh.p vhh Qp $ $ !$$ $ $$ $ "": Qrpvypuhr.r r Q $ $ $ "$ $$ & $ $ & $ #$ $ &$ !$ $ $ " $ $ $ ;/; Qyqr.hvyy. Qh $$$ $ ,: Qhpr .yhh pxvv. Qyh ": Q hvyyryyh.hssvv. Qh ": Q hvyyryyh.t hpvyv Qt $"$ $$$$$ $ =: Q vv.phr v. Q ph "' % $ '$ !$ &$ " ! !$ '$ $ $ $$ "DD; Hvv.pv vsr h. Q pv $ ; Q vv.ruyr v Q ru $ $ $ $ $ $ $ ! $"$ ! $ "D: Q vv.shyyh Q y $$$ ,; Q vv.rr v Q r $ $!$$$$ =: Q q vyyrh.xrsr rvv. Qxs $ "; Qrqyrvphvryyh.shryv. Qyps $ $$$ /: Thiryyvqr.ppv hh. Tip $ "; Tphyvi rth.vsyh. Tpv $ ; 9 vyyrh.Tpuvr vt.rtyrph. Tpu $ $$$ ! <: 9 vyyrh.Tpuvr vt. qyuvv Tpu #$ ;; Tpuyr purvy.v. Tpy ": Tpyryrv.phhi h. Tpph $ "; Tpyryrv.syvh. Tps $$ ": Tpyryrv.hhh. Tp $$ (: Tpyryrv. vqrhh. Tp $ ; Tpyy.Tpyy.h vtr . Tph !"$!$!"# ! # ! "$ $ /:: Tr yh.ppuh . Trpu $ "; Tvthyv.huvyqhr. Tvt $$ ! $# $ $ !$ $ $ $ $ $ (,: Tvthi h.rhpyhh Tvr $$$ !$$ $ ":: Tv.qrp h. Tqrp $ $$$ & ' $ ( " "$ !$ ($ ( =(; Tv.yvpyhh T !$$ /: Tvuhr.ii. Ti $$$ $$$$ /; Tvuhr.x r v.x r v. Txx $$ $$$ ,; Tvuhr. T $ ; Tr hv.phh Tp $$ $$$ /; Ur riryyvqr. rv U "$ $$$$$$$$ $ $ $$! ",; Wr vyvv.vsqviy Wvs % <: E!&0& D<: ";<: /"; ",=; C; ;; /D; ;": ;(: ",; /D: ,"; /C: (C: C;; ;D: /C: /(: ;:: C"; ,C: (<; ,:; "/; ",: "=: <(: D: (=: /; D; ,:; ,/: "=; "": <: <; =; /: C; /; ":: D: /: ;: =; /: ,; : /; ";/(; 116 # 7 ]VH1V HQCCVH VR 1J .V :`V: Q` ':V : #( ' $#$ 6FG5HG6! I5JKEL) O>.(: O>A(: O>8(: O>*(: O>$(: O>A,: O>8,: O>*,: O>$,: O>A/: O>./: O>8/: O>*/: O>$/: O>A;: O>8;: O>*;: O>$;: O>.<: O>A<: O>*<: O>8<: O>$<: O>AC: O>8C: O>*C: O>$C: Q 6uh rr.hpvs . 6p $$$ 6urypuhrh.h vv 6 $ $ $ $ $ $ $ $ $ 6uv.ivyvrhh. 6i $$$!$ 7 hqh.vyyh 7 $$ 8uhrr ! ! 8 $$ $ 8yr h.pyrhh 8py !$ 9vypv .tyhp 9t !$! $ $ $ $ 9v h.h vrvh. 9h 9 vyr rv.svy 9s !!$ $$ $ $ $ @vpr.vhh @v "$ Bypr h.vp v B $ $ $ ! $ $ $ $ $ $ $ $ $ $ $ $ Crr h.svyvs v Cs $ ! $ $ $ $ $ Ghvpr.pv hh Gp "$!$ $ $ $ !$ $ Tpyrh.rhqviyhhhivv Gr # $ $ $ ! $ $ $ " $ ! $ $ $ Tpyrh.spuhyrv Gs Gi vr v.t hpvyv Gt $$$ Gi vr v.yh rvyyv Gyh ! ' $ $ % "$ !$ $ $ ! ! $ Tpyrh.r h h. Gr $ Hhtryyh. . H $ $ $ Hh uh.xvir tv Hx $$ $ Hryvh.hyhh Hy $$$$ "$ $ $ $ $ $ $ Hvpryyvh.q i hpuvhyv Hqi $$ H vpuryr.pyhh. Hp $$ @rr.vph Hp $ Iruu.uir tv Iru !$%"$&$!$ $ Iruu.u vpv Iru !$ $ $ $ $ !$ $ $ $ $ Iruu.vpvh. Irv $ @r rv.ytvvh. Iyt $ Ih.hir h Ihi ($ $ $ #$ $ "$ #$ " $ $ $ $ $ Ih.yhr vpr Iyh $$$$ Prvh.svs v Ps Qh hyhpqvh.h hqh Q $ $ Qh h vv.vhh Q $ $$ $ $ $ Qrpvh vh.h vph Qh $$ Qrpvh vh.x rv Qx $$$ Quy.srvqh. Qus $ Qvyh tv.r ph Q $ ! Qvh.p vhh Qp $$ Qrpvypuhr.r r Q $$$$$ Q hvyyryyh.t hpvyv Qt $$$$ Q vv.rr v Q r Qrqyrvphvryyh.shryv. Qyps #$ $ !$ $ # $ Tpyh vpvh.vph Tp $ Qyq h.pvyvhh. Tppvy $$ $$$ Tpyy.Tpyy.h vtr . Tph $ Tv.svyvp v Tsvy " Tvpuhrr .ph Tpp $ $ Tr hv.phh Tp !$ $ $"$ $$ $ $ $ $ $ $ $ Ur riryyvqr. rv U $$$$$$$$$$ E!&0& ;,: (D; ,": "C; ("; (": (=: ",: (;: <; D: C: ":: <; /: C; <: /: (: C: ,: (; "": ;; /: C: /; ,/<; 117 # 7 : 1 1H Q% ]% &(# # V " 2$ ,$ V`I% : 1QJ:C2$ ,$ &VVIGC:JHV 1Q`@.VV :IV7 VVI : : 7]V71I1C:`1 7 VCVH 1QJ7$CC `:J`Q`I7_%:`V`QQ VVIGC:JHV7 <`:73%` 11I1C:`1 7 %IQ`_%:`V 7]V7 7]V ^]:` 1:C_ V`I% : 1QJIV .QR7V`I% : 1QJQ``V1R%:C%JRV` .V`%CCIQRVC %IGV`Q`]V`I% : 1QJ7 )# 1'&% :IV 7]V V0VC RV] . &16VR :`V: &16VR J1_%V Q%`HV R` 2 V%RQR& ^]V`I_ ]V`I ^23_ RV] . 5 5 5 :`V: 5 5 5 RV] .6:`V: 5 5 5 5 V 5 " Q :C 5 " V`I.:QJVQ`IQ`VVI] 7HVCC .V :1C Q` .V V6]VH VR IV:J _%:`V ^$ %_ `Q` .V IQRVC Q%`HV "2 RV] . VU5 RV] . :`V: VU 5 :`V: RV] .6:`V: VU 5 RV] .6:`V: %IV`: Q` V`I Q%`HV V`I1JHC%RVR1J .VJ%IV`: Q` RV] . RV] . :`V: :`V: RV] .6:`V: RV] .6:`V: .VJQI1J: Q` V`I Q%`HV V`I1JHC%RVR1J .VRVJQI1J: Q` RV] . V :`V: V RV] .6:`V: V ' ' $ 1% ') #&@#1@' Q%`HV " 1I: V _8`QQ RV] . 5 5 :`V: 5 5 RV] .6:`V: 5 5 118 " 2$ ,$ V`I% : 1QJ:C2$ ,$ &VVIGC:JHV 1Q`@.VV :IV7 VVI : : 7]V71I1C:`1 7 VCVH 1QJ7$CC `:J`Q`I7_%:`V`QQ VVIGC:JHV7 <`:73%` 11I1C:`1 7 %IQ`_%:`V 7]V7 7]V ^]:` 1:C_ V`I% : 1QJIV .QR7V`I% : 1QJQ``V1R%:C%JRV` .V`%CCIQRVC %IGV`Q`]V`I% : 1QJ7 )# 1'&% :IV 7]V V0VC RV] . &16VR :`V: &16VR RV] .$`Q%]7 Y RI5YRI5YR I5Y R I5 Y R I:JR YR I #@&@%$ 1$%1% V`IRV] .6:`V:`Q`]:1`Q`CV0VCQ``:H Q`RV] . 1 .1JCV0VC4 Q``:H Q`:`V: J1_%V 4`Q%] ^]V`I_ ]V`I ^23_ 5 5 5 5 5 5 5 5 5 5 5 5 5 Q V 5R`Y 5 Q V 5R`Y 5 Q V 5R`Y $0V`:$V1I1C:`1 7GV 1VVJL11 .1J$`Q%] 5 5 5 5 5 5 1 .1JCV0VC "Q``:H Q`:`V: J1_%V 4`Q%] ^]V`I_ ]V`I ^23_ 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 119 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 $0V`:$V1I1C:`1 7GV 1VVJL11 .1J$`Q%] 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 1 .1JCV0VC$Q``:H Q`:`V: J1_%V 4`Q%] ^]V`I_ ]V`I ^23_ 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 $0V`:$V1I1C:`1 7GV 1VVJL11 .1J$`Q%] 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 1 .1JCV0VC Q``:H Q`:`V: J1_%V 4`Q%] ^]V`I_ ]V`I ^23_ 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 120 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 $0V`:$V1I1C:`1 7GV 1VVJL11 .1J$`Q%] 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 1 .1JCV0VC2 Q``:H Q`:`V: J1_%V 4`Q%] ^]V`I_ ]V`I ^23_ 5 5 5 5 5 5 5 5 5 5 5 5 5 Q V 5R`Y 5 Q V 5R`Y 5 Q V 5R`Y $0V`:$V1I1C:`1 7GV 1VVJL11 .1J$`Q%] 5 5 5 5 5 5 121 " 2$ ,$ V`I% : 1QJ:C2$ ,$ &VVIGC:JHV 1Q`@.VV :IV7 VVI : : 7]V71I1C:`1 7 VCVH 1QJ7$CC `:J`Q`I7_%:`V`QQ VVIGC:JHV7 <`:73%` 11I1C:`1 7 %IQ`_%:`V 7]V7 7]V ^]:` 1:C_ V`I% : 1QJIV .QR7V`I% : 1QJQ``V1R%:C%JRV` .V`%CCIQRVC %IGV`Q`]V`I% : 1QJ7 )# 1'&% :IV 7]V V0VC RV] . &16VR :`V: &16VR RV] .$`Q%]7 Y RI5YRI5YR I5Y R I5 Y R I:JR YR I #@&@%$ 1$%1% V`IRV] .6:`V:`Q`]:1`Q`CV0VCQ``:H Q`:`V: 1 .1JCV0VCQ``:H Q`RV] . J1_%V 4`Q%] ^]V`I_ ]V`I ^23_ 4 5 " 5 5 5 4 5$ 5 5 5 4 5 5 5 5 4 52 5 5 5 "5$ 5 5 5 "5 5 5 5 "52 5 5 5 $5 5 5 5 $52 5 5 5 52 5 5 5 $0V`:$V1I1C:`1 7GV 1VVJL11 .1J$`Q%] 4 " $ 2 4 5 " 5 5 $ 5 5 5 5 5 5 5 2 5 5 5 5 5 1 .1JCV0VCQ``:H Q`RV] . J1_%V 4`Q%] ^]V`I_ ]V`I ^23_ 4 5 " 5 5 5 4 5$ 5 5 5 4 5 5 5 5 4 52 5 5 5 "5$ 5 5 5 "5 5 5 5 "52 5 5 5 $5 5 5 5 122 $52 5 5 5 52 5 5 5 $0V`:$V1I1C:`1 7GV 1VVJL11 .1J$`Q%] 4 " $ 2 4 5 " 5 5 $ 5 5 5 5 5 5 5 2 5 5 5 5 5 1 .1JCV0VCQ``:H Q`RV] . J1_%V 4`Q%] ^]V`I_ ]V`I ^23_ 4 5 " 5 5 5 4 5$ 5 5 5 4 5 5 5 5 4 52 5 5 5 "5$ 5 5 5 "5 5 5 5 "52 5 5 5 $5 5 5 5 $52 5 5 5 52 5 5 5 $0V`:$V1I1C:`1 7GV 1VVJL11 .1J$`Q%] 4 " $ 2 4 5 " 5 5 $ 5 5 5 5 5 5 5 2 5 5 5 5 5 1 .1JCV0VCQ``:H Q`RV] . J1_%V 4`Q%] ^]V`I_ ]V`I ^23_ 4 52 Q V 5R`Y "5$ 5 5 5 "5 5 5 5 $5 5 5 5 $0V`:$V1I1C:`1 7GV 1VVJL11 .1J$`Q%] 4 " $ 2 4 " 5 $ 5 5 5 5 5 2 1 .1JCV0VC Q``:H Q`RV] . J1_%V 4`Q%] ^]V`I_ ]V`I ^23_ 4 52 Q V 5R`Y "5$ 5 5 5 "5 5 5 5 $5 5 5 5 123 $0V`:$V1I1C:`1 7GV 1VVJL11 .1J$`Q%] 4 " $ 2 4 " 5 $ 5 5 5 5 5 2 1 .1JCV0VC Q``:H Q`RV] . J1_%V 4`Q%] ^]V`I_ ]V`I ^23_ 4 52 Q V 5R`Y "5$ 5 5 5 "5 5 5 5 $5 5 5 5 $0V`:$V1I1C:`1 7GV 1VVJL11 .1J$`Q%] 4 " $ 2 4 " $ 5 5 5 5 5 2 124 " 2$ ,$ V`I% : 1QJ:C2$ ,$ &VVIGC:JHV 1Q`@.VV :IV7 VVI : : 7]V71I1C:`1 7 VCVH 1QJ7$CC VVIGC:JHV7 E:HH:`R %IQ`_%:`V 7]V7 7]V ^]:` 1:C_ V`I% : 1QJIV .QR7V`I% : 1QJQ``V1R%:C%JRV` .V`%CCIQRVC %IGV`Q`]V`I% : 1QJ7 )# 1'&% :IV 7]V V0VC RV] .Z`:J$V &16VR :`V: &16VR J1_%V Q%`HV R` 2 V%RQR& ^]V`I_ ]V`I ^23_ RV] .Z`:J$V 5 5 5 :`V: 5 5 5 RV] .Z`:J$V6:`V: 5 5 5 5 V 5 " 5 Q :C 5 " V`I.:QJVQ`IQ`VVI] 7HVCC .V :1C Q` .V V6]VH VR IV:J _%:`V ^$ %_ `Q` .V IQRVC Q%`HV "2 RV] .Z`:J$V VU5 RV] .Z`:J$V :`V: VU 5 :`V: RV] .Z`:J$V6:`V: VU 5 RV] .Z`:J$V6:`V: %IV`: Q` V`I Q%`HV V`I1JHC%RVR1J .VJ%IV`: Q` RV] .Z`:J$V RV] .Z`:J$V :`V: :`V: RV] .Z`:J$V6:`V: RV] .Z`:J$V6:`V: .VJQI1J: Q` V`I Q%`HV V`I1JHC%RVR1J .VRVJQI1J: Q` RV] .Z`:J$V V :`V: V RV] .Z`:J$V6:`V: V ' ' $ 1% ') #&@#1@' Q%`HV " 1I: V _8`QQ RV] .Z`:J$V 5 5 :`V: 5 5 RV] .Z`:J$V6:`V: 5 5 125 " 2$ ,$ V`I% : 1QJ:C2$ ,$ &VVIGC:JHV 1Q`@.VV :IV7 VVI : : 7]V71I1C:`1 7 VCVH 1QJ7$CC VVIGC:JHV7 E:HH:`R %IQ`_%:`V 7]V7 7]V ^]:` 1:C_ V`I% : 1QJIV .QR7V`I% : 1QJQ``V1R%:C%JRV` .V`%CCIQRVC %IGV`Q`]V`I% : 1QJ7 )# 1'&% :IV 7]V V0VC RV] .Z`:J$V &16VR :`V: &16VR RV] .$`Q%]7 Y RI5YRI5YR I5Y R I5 Y R I:JR YR I #@&@%$ 1$%1% V`IRV] .Z`:J$V6:`V:`Q`]:1`Q`CV0VCQ``:H Q`RV] .Z`:J$V 1 .1JCV0VCQ``:H Q`:`V: J1_%V 4`Q%] ^]V`I_ ]V`I ^23_ 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 $0V`:$V1I1C:`1 7GV 1VVJL11 .1J$`Q%] 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 1 .1JCV0VCQ``:H Q`:`V: J1_%V 4`Q%] ^]V`I_ ]V`I ^23_ 5 5 5 5 5 5 5 5 5 5 5 5 126 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 $0V`:$V1I1C:`1 7GV 1VVJL11 .1J$`Q%] 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 1 .1JCV0VCQ``:H Q`:`V: J1_%V 4`Q%] ^]V`I_ ]V`I ^23_ 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 $0V`:$V1I1C:`1 7GV 1VVJL11 .1J$`Q%] 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 1 .1JCV0VC Q``:H Q`:`V: J1_%V 4`Q%] ^]V`I_ ]V`I ^23_ 5 Q V 5R`Y 5 Q V 5R`Y 5 5 5 5 5 5 5 5 127 5 5 5 5 5 Q V 5R`Y $0V`:$V1I1C:`1 7GV 1VVJL11 .1J$`Q%] 5 5 5 5 5 5 1 .1JCV0VCQ``:H Q`:`V: J1_%V 4`Q%] ^]V`I_ ]V`I ^23_ 5 Q V 5R`Y 5 Q V 5R`Y 5 5 5 5 5 5 5 5 5 5 5 5 5 Q V 5R`Y $0V`:$V1I1C:`1 7GV 1VVJL11 .1J$`Q%] 5 5 5 5 5 5 128 " 2$ ,$ V`I% : 1QJ:C2$ ,$ &VVIGC:JHV 1Q`@.VV :IV7 VVI : : 7]V71I1C:`1 7 VCVH 1QJ7$CC VVIGC:JHV7 E:HH:`R %IQ`_%:`V 7]V7 7]V ^]:` 1:C_ V`I% : 1QJIV .QR7V`I% : 1QJQ``V1R%:C%JRV` .V`%CCIQRVC %IGV`Q`]V`I% : 1QJ7 )# 1'&% :IV 7]V V0VC RV] .Z`:J$V &16VR :`V: &16VR RV] .$`Q%]7 Y RI5YRI5YR I5Y R I5 Y R I:JR YR I #@&@%$ 1$%1% V`IRV] .Z`:J$V6:`V:`Q`]:1`Q`CV0VCQ``:H Q`:`V: 1 .1JCV0VCQ``:H Q`RV] .Z`:J$V J1_%V 4`Q%] ^]V`I_ ]V`I ^23_ 5 5 5 5 5 5 5 5 5 5 5 5 5 Q V 5R`Y $0V`:$V1I1C:`1 7GV 1VVJL11 .1J$`Q%] 5 5 5 5 5 5 1 .1JCV0VCQ``:H Q`RV] .Z`:J$V J1_%V 4`Q%] ^]V`I_ ]V`I ^23_ 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 $0V`:$V1I1C:`1 7GV 1VVJL11 .1J$`Q%] 5 129 5 5 5 5 5 5 5 5 5 5 5 5 5 5 1 .1JCV0VCQ``:H Q`RV] .Z`:J$V J1_%V 4`Q%] ^]V`I_ ]V`I ^23_ 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 $0V`:$V1I1C:`1 7GV 1VVJL11 .1J$`Q%] 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 1 .1JCV0VCQ``:H Q`RV] .Z`:J$V J1_%V 4`Q%] ^]V`I_ ]V`I ^23_ 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 $0V`:$V1I1C:`1 7GV 1VVJL11 .1J$`Q%] 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 1 .1JCV0VC Q``:H Q`RV] .Z`:J$V J1_%V 4`Q%] ^]V`I_ ]V`I ^23_ 5 5 5 5 5 5 5 5 5 5 5 5 5 Q V 5R`Y 130 $0V`:$V1I1C:`1 7GV 1VVJL11 .1J$`Q%] 5 5 5 5 5 5 1 .1JCV0VC Q``:H Q`RV] .Z`:J$V J1_%V 4`Q%] ^]V`I_ ]V`I ^23_ 5 5 5 5 5 5 5 5 5 5 5 5 5 Q V 5R`Y $0V`:$V1I1C:`1 7GV 1VVJL11 .1J$`Q%] 5 5 5 5 131 # 7 : 1 1H Q% ]% Q` &( V " 2 1 :JHVRG:VR V `Q`.QIQ$VJV1 7Q`I%C 10:`1: VR1]V`1QJ &VVIGC:JHV 1Q`@.VV :IV7 VVI : : 7]V71I1C:`1 7 VCVH 1QJ7$CC VVIGC:JHV7 E:HH:`R 4`Q%]`:H Q`7:`V: %IGV`Q`$`Q%]7 : 1QJ %IGV`Q`:I]CV7 .$@#1@' % )&' $ 1&'@. &75 ^]V`I_75 #@&@%$ ' #&@%' % 4`Q%] ^]V`I_ ^5_ 5 5 ^5_ 5 5 ^5 _ 5 "R ^5_ 5 "R ^5_ 5 5 "R ^5 _ 5 "R ^5_ 5 "R ^5 _ 5 5 "R ^5_ 5 5 "R ^ 5_ 5 5 $# % # . %1# .#&. $&&'&% 4`Q%] $0V`:$V " 5 5 5 5 5 5 5 5 5 5 132 " 2 1 :JHVRG:VR V `Q`.QIQ$VJV1 7Q`I%C 10:`1: VR1]V`1QJ VVIGC:JHV1Q`@.VV :IV7 VVI : : 7]V71I1C:`1 7 VCVH 1QJ7$CC VVIGC:JHV7 E:HH:`R 4`Q%]`:H Q`7RV] .Z`:J$V %IGV`Q`$`Q%]7 %IGV`Q`:I]CV7 " $ , & ,23" , &75 ^]V`I_75 $ "3,2$ , 4`Q%] ^]V`I_ ^5_ 5 5 "R ^5_ 5 5 ^5_ 5 "R ^5 _ 5 "R ^5 _ 5 5"R ^5_ 5 5 ^5_ 5 5 ^5 _ 5 "R ^5 _ 5 5 "R ^5_ 5 5 "R ^5 _ 5 "R ^5 _ 5 5 "R ^5 _ 5 5 "R ^5 _ 5 5 ^ 5 _ 5 5 2"$ $ $ $ " , 4`Q%] $0V`:$V " 5 5 5 5 5 5 5 5 5 5 5 5 133 #]]VJR16 7 .V ]VH1VR$`Q%] HQJ `1G% 1QJ QG :1JVR ``QI JR1H: Q` ]VH1V :J:C71 ^ $_8 .V1JR1H: Q`0:C%VQ` $]VH1`7 .V$`Q%]Q` : 1QJ .: :`V:QH1: VR11 . .V ]VH1V11 .R1``V`VJ CV0VCQ`1$J1`1H:JHV^1JGQCR_8VR1IVJ HC:V^ Q :5 _7:JR^Y_ ^] Q HQJ VJ Q` :JR_5 I%RR7 :JR ^IY_ ^ T Q_5 0V`7 :JR7 I%R ^02Y_ ^ TQ_5 :JR7I%R^2Y_^T Q_:JRI%R^2Y _^\ Q_8 ]VH1V I 0( ( ( ]RCV0VC I:$V :R]V`: 8 8 8 8 8 8 I].:`V V :H% 1``QJ 8 8 8 8 8 8 #I].1H V1 $%JJV`1 8 8 8 8 8 8 #JH1 :`$1 .:I: : 8 8 8 8 8 8 #JH7 `Q7CC1 $`QVJC:JR1H: 8 8 8 8 8 8 JQGQ .`% $`:H1C1 8 8 8 8 8 8 #QJ1RV Q67HV].:C: 8 8 8 8 8 8 #QJ1RV ]:%H1G`:JH.1: : 8 8 8 8 8 8 #].VCQH.:V : I:`1QJ1 8 8 8 8 8 8 :%CCV`1VCC: I%C 1G`:JH.11 8 8 8 8 8 8 ]QJ%].1 G1C1JV: : 8 8 8 8 8 8 ]QJ%].1 G`VIVJ 1 8 8 8 8 8 8 ]QJ%].1 `:%0VC1 8 8 8 8 8 8 #`:GVCC: 1`1HQCQ` 8 8 8 8 8 8 #`I:JR1: H1``.Q: 8 8 8 8 8 8 61Q .VCC: HQJ `1H : 8 8 8 8 8 8 2`:R: 01CCQ: 8 8 8 8 8 8 :]1 VCC: H:]1 : : 8 8 8 8 8 8 :]1 VCC: $1:`R1 8 8 8 8 8 8 :]1 QI: % I1J1I% 8 8 8 8 8 8 .:V Q $%J1HV :J VJJ: : 8 8 8 8 8 8 $%J1HV .:`:11 8 8 8 8 8 8 $%J1HV QV` VR1 8 8 8 8 8 8 $%J1HV ]VJJ: : 8 8 8 8 8 8 %J1HV 01 : : 8 8 8 8 8 8 $%].`Q1JV :`I:R1CCQ 8 8 8 8 8 8 %].`Q1JV `QC1Q: 8 8 8 8 8 8 #C:GVCC1$V`: R1]CQH.:1 % 8 8 8 8 8 8 -C7HV`: :CG: 8 8 8 8 8 8 -C7HV`: H:]1 : : 8 8 8 8 8 8 134 -C7HV`: %J1HQ`J1 8 8 8 8 8 8 -C7H1JRV JQ`RI:JJ1 8 8 8 8 8 8 -QJ1:R: VIV`1 : 8 8 8 8 8 8 -QJ1:R: I:H%C: : 8 8 8 8 8 8 :CI$`VJ1VCC: C=%J$I:J1 8 8 8 8 8 8 :CI$`VJ1VCC: C%J%C: : 8 8 8 8 8 8 :`IQ .QV ]1J1`V`: 8 8 8 8 8 8 V1QJV ]CVJR1R: 8 8 8 8 8 8 V V`QI: % `1C1`Q`I1 8 8 8 8 8 8 7:C1JQVH1: %G1HQC: 8 8 8 8 8 8 7R`Q1RV ]V%RQ%JH1J: % 8 8 8 8 8 8 !:V IQJ1HV .7 `16 8 8 8 8 8 8 %:QJ1HV H` H1``: : 8 8 8 8 8 8 !V1QH:]1 VCC: $C:G`: 8 8 8 8 8 8 !V1QH.`1RV ]8 8 8 8 8 8 8 !V01JVJ1: $`:H1C1 8 8 8 8 8 8 !%IG`1JV`1Q]1 ]:`:RQ6: 8 8 8 8 8 8 %%IG`1JV`1 HQHH1JV: 8 8 8 8 8 8 %HQCV QI: VI:JR1G%C: :RI:G1 1 8 8 8 8 8 8 %HQCV QI: ``:$1C1 8 8 8 8 8 8 HQCV QI: `%JH.:CVJ1 8 8 8 8 8 8 !%IG`1JV`1 $`:H1C1 8 8 8 8 8 8 %%IG`1JV`1 C: `V1CC1 8 8 8 8 8 8 HQCV QI: V `:%`: 8 8 8 8 8 8 !71R1HV H` J1JV : 8 8 8 8 8 8 :$VCQJ: :CCVJ1 8 8 8 8 8 8 :$VCQJ: ] 8 8 8 8 8 8 :C:HQHV`Q `%C1$1JQ% 8 8 8 8 8 8 :C:HQHV`Q V `:HV`Q 8 8 8 8 8 8 :CR:JV $CVG1`V6 8 8 8 8 8 8 :CR:JV :`1 8 8 8 8 8 8 :`].7: GVCC1 8 8 8 8 8 8 :`].7: `:CC:6 8 8 8 8 8 8 :`].7: @1JGV`$1 8 8 8 8 8 8 VR1QI: % H:]VJ1 8 8 8 8 8 8 VR1QI: % ``:$1C1 8 8 8 8 8 8 VC1JJ: ]:CI: : 8 8 8 8 8 8 V :7H.1 $Q Q1 8 8 8 8 8 8 QJ 1HVCC1J: RQ`QG`:JH.1:C1 8 8 8 8 8 8 7`1QH.VCV QH%C: : 8 8 8 8 8 8 $ VQJV ]1H : 8 8 8 8 8 8 VI: QJV`V1 %J1HQ`J1 8 8 8 8 8 8 V]. .7 H1``Q: 8 8 8 8 8 8 V]. .7 .QIGV`$1 8 8 8 8 8 8 V]. .7 .7 `1H1 8 8 8 8 8 8 V]. .7 1JH1: 8 8 8 8 8 8 V]. .7 ]8 8 8 8 8 8 8 $%JV`V1 CQJ$11I: 8 8 8 8 8 8 1H.QI:H.V C%IG`1H:C1 8 8 8 8 8 8 1JQV :`IQ`1H:J: 8 8 8 8 8 8 Q .`1: HQJH.1CV$: 8 8 8 8 8 8 Q QI: % :GV`:J 8 8 8 8 8 8 Q QI: % C: V`1HV% 8 8 8 8 8 8 Q QI: % C1JV: % 8 8 8 8 8 8 J%].1 V`VI1 : 8 8 8 8 8 8 'J%].1 ] 8 8 8 8 8 8 135 'J%].1 ]8 8 8 8 8 8 8 '`G1J1: H%01V`1 8 8 8 8 8 8 1VJ1: `%1`Q`I1 8 8 8 8 8 8 :CQC: 1H1C1VJ1 8 8 8 8 8 8 :J .:C1 QV` VR1 8 8 8 8 8 8 :`:C:H7RQJ1: ]:`:RQ6: 8 8 8 8 8 8 :`:QJ1 `%C$VJ 8 8 8 8 8 8 :`:]`1QJQ]1Q H` ]1JJ: : 8 8 8 8 8 8 VH 1J:`1: :%`1HQI: 8 8 8 8 8 8 VH 1J:`1: @Q`VJ1 8 8 8 8 8 8 V :CQ]`QH % V``1HQC% 8 8 8 8 8 8 V : ]%1CC: 8 8 8 8 8 8 .V`%: IQJ1C1`V`: 8 8 8 8 8 8 .V`%: ]C%IQ: 8 8 8 8 8 8 .7CCQRQHV H` I:H%C: : 8 8 8 8 8 8 .7CCQRQHV C:I1JQ: 8 8 8 8 8 8 .7CCQRQHV C1JV: : 8 8 8 8 8 8 .7CCQRQHV I:H`Q]. .:CI: 8 8 8 8 8 8 .7CQ `QV 1R: 8 8 8 8 8 8 .7CQ C1$% 1H: 8 8 8 8 8 8 .7CQ JQ`0V$1H: 8 8 8 8 8 8 1C:`$1 0V``%HQ: 8 8 8 8 8 8 1`QI1 V`%H: 8 8 8 8 8 8 1 : H`1 : : 8 8 8 8 8 8 QVH1CQH.:V % V`]VJ 8 8 8 8 8 8 QC7H1``% :%`:J 1:H% 8 8 8 8 8 8 QC7QRQJ V I:61CCQ% 8 8 8 8 8 8 QC7Q]. :CI% ]1H % 8 8 8 8 8 8 QI: QHV`Q C:I:`H@11 8 8 8 8 8 8 `:61CCVCC: :``1J1 8 8 8 8 8 8 `:61CCVCC: $`:H1C1 8 8 8 8 8 8 `1QJQ]1Q H:]V`1 8 8 8 8 8 8 1J%]1Q H1``1`V`: 8 8 8 8 8 8 `1QJQ]1Q V.CV`1 8 8 8 8 8 8 `1QJQ]1Q `:CC:6 8 8 8 8 8 8 _%1C:]1Q V6QH%C: : 8 8 8 8 8 8 `1QJQ]1Q VVJ `%]1 8 8 8 8 8 8 `Q QRQ`01CCV: @V`V` V1J1 8 8 8 8 8 8 `Q %C: ]8 8 8 8 8 8 8 :IIQC7HV :`VJQ: 8 8 8 8 8 8 V%RQCV1QH:]1 VCC: `:%0VC1 8 8 8 8 8 8 :GVCC1RV QH QH1``: : 8 8 8 8 8 8 H:C1G`V$I: 1J`C: %I 8 8 8 8 8 8 Q`01CCV: ^ H.1 QIV`1J$Q_ JV$CVH : 8 8 8 8 8 8 .Q`01CCV: ^%H.1 QIV`1J$Q_ `%RQC].11 8 8 8 8 8 8 H.CV`QH.V1C% I1J% % 8 8 8 8 8 8 %HQC:`1H1: 7]1H: 8 8 8 8 8 8 HQCVCV]1 H:J :G`: 8 8 8 8 8 8 QC7RQ`: H1C1: : 8 8 8 8 8 8 HQCVCV]1 `QC1Q: 8 8 8 8 8 8 HQCVCV]1 _%:I: : 8 8 8 8 8 8 HQCVCV]1 `1RVJ : : 8 8 8 8 8 8 HQCQ]CQ ^ HQCQ]CQ_ :`I1$V` 8 8 8 8 8 8 V`]%C: HQJH.:`%I 8 8 8 8 8 8 1$:C1QJ I: .1CR:V 8 8 8 8 8 8 136 1$:C1QJ _%:IQ%I 8 8 8 8 8 8 1$:IG`: VJ :H%C: : 8 8 8 8 8 8 ]1Q RVHQ`: % 8 8 8 8 8 8 ]1Q `1C1HQ`J1 8 8 8 8 8 8 %]1Q I%C 1QH%C: : 8 8 8 8 8 8 %]1QH.:V Q] V`% HQ :`%I 8 8 8 8 8 8 ]1QJ1R:V]8 8 8 8 8 8 8 ]1Q].:JV GQIG76 8 8 8 8 8 8 ]1Q].:JV @`Q7V`1 @`Q7V`1 8 8 8 8 8 8 %]1Q].:JV ]8 8 8 8 8 8 8 V`J:]1 H% : : 8 8 8 8 8 8 .VJVC:1 GQ: 8 8 8 8 8 8 .VJVC:1 C1I1HQC: 8 8 8 8 8 8 *V`VGVCC1RV `QVI1 8 8 8 8 8 8 V`I1C1Q]1 1J`%JR1G%C%I 8 8 8 8 8 8 137 #]]VJR16 7 .V ]VH1VR$`Q%] HQJ `1G% 1QJ QG :1JVR ``QI JR1H: Q` ]VH1V :J:C71 ^ $_8 .V1JR1H: Q`0:C%VQ` $]VH1`7 .V$`Q%]Q` : 1QJ .: :`V:QH1: VR11 . .V ]VH1V 11 . R1``V`VJ CV0VC Q` 1$J1`1H:JHV ^1J GQCR_8 V] . $`Q%]7 Y R I5 YR I5 YR I5Y R I5 Y R I:JR YR I ]VH1V R R R R R R ]RCV0VC #I:$V :R]V`: 8 8 8 8 8 8 8 #I].:`V V :H% 1``QJ 8 8 8 8 8 8 8 I].1H V1 $%JJV`1 8 8 8 8 8 8 8 #JH1 :`$1 .:I: : 8 8 8 8 8 8 8 #JH7 `Q7CC1 $`QVJC:JR1H: 8 8 8 8 8 8 8 JQGQ .`% $`:H1C1 8 8 8 8 8 8 8 #QJ1RV Q67HV].:C: 8 8 8 8 8 8 8 #QJ1RV ]:%H1G`:JH.1: : 8 8 8 8 8 8 8 ].VCQH.:V : I:`1QJ1 8 8 8 8 8 8 8 :%CCV`1VCC: I%C 1G`:JH.11 8 8 8 8 8 8 8 ]QJ%].1 G1C1JV: : 8 8 8 8 8 8 8 ]QJ%].1 G`VIVJ 1 8 8 8 8 8 8 8 #]QJ%].1 `:%0VC1 8 8 8 8 8 8 8 `:GVCC: 1`1HQCQ` 8 8 8 8 8 8 8 #`I:JR1: H1``.Q: 8 8 8 8 8 8 8 #61Q .VCC: HQJ `1H : 8 8 8 8 8 8 8 +`:R: 01CCQ: 8 8 8 8 8 8 8 :]1 VCC: H:]1 : : 8 8 8 8 8 8 8 :]1 VCC: $1:`R1 8 8 8 8 8 8 8 :]1 QI: % I1J1I% 8 8 8 8 8 8 8 .:V Q 7:C1JQVH1: %G1HQC: 8 8 8 8 8 8 8 7R`Q1RV ]V%RQ%JH1J: % 8 8 8 8 8 8 8 !:V IQJ1HV .7 `16 8 8 8 8 8 8 8 %:QJ1HV H` H1``: : 8 8 8 8 8 8 8 %V1QH:]1 VCC: $C:G`: 8 8 8 8 8 8 8 %V1QH.`1RV ]8 8 8 8 8 8 8 8 !V01JVJ1: $`:H1C1 8 8 8 8 8 8 8 !%IG`1JV`1Q]1 ]:`:RQ6: 8 8 8 8 8 8 8 !%IG`1JV`1 HQHH1JV: 8 8 8 8 8 8 8 HQCV QI: VI:JR1G%C: :RI:G1 1 8 8 8 8 8 8 8 %HQCV QI: ``:$1C1 8 8 8 8 8 8 8 %HQCV QI: `%JH.:CVJ1 8 8 8 8 8 8 8 !%IG`1JV`1 $`:H1C1 8 8 8 8 8 8 8 %%IG`1JV`1 C: `V1CC1 8 8 8 8 8 8 8 HQCV QI: V `:%`: 8 8 8 8 8 8 8 !71R1HV H` J1JV : 8 8 8 8 8 8 8 :$VCQJ: :CCVJ1 8 8 8 8 8 8 8 :$VCQJ: ] 8 8 8 8 8 8 8 :C:HQHV`Q `%C1$1JQ% 8 8 8 8 8 8 8 :C:HQHV`Q V `:HV`Q 8 8 8 8 8 8 8 :CR:JV $CVG1`V6 8 8 8 8 8 8 8 :CR:JV :`1 8 8 8 8 8 8 8 :`].7: GVCC1 8 8 8 8 8 8 8 :`].7: `:CC:6 8 8 8 8 8 8 8 :`].7: @1JGV`$1 8 8 8 8 8 8 8 VR1QI: % H:]VJ1 8 8 8 8 8 8 8 VR1QI: % ``:$1C1 8 8 8 8 8 8 8 VC1JJ: ]:CI: : 8 8 8 8 8 8 8 V :7H.1 $Q Q1 8 8 8 8 8 8 8 QJ 1HVCC1J: RQ`QG`:JH.1:C1 8 8 8 8 8 8 8 7`1QH.VCV QH%C: : 8 8 8 8 8 8 8 $ VQJV ]1H : 8 8 8 8 8 8 8 VI: QJV`V1 %J1HQ`J1 8 8 8 8 8 8 8 V]. .7 H1``Q: 8 8 8 8 8 8 8 V]. .7 .QIGV`$1 8 8 8 8 8 8 8 V]. .7 .7 `1H1 8 8 8 8 8 8 8 V]. .7 1JH1: 8 8 8 8 8 8 8 V]. .7 ]8 8 8 8 8 8 8 8 $%JV`V1 CQJ$11I: 8 8 8 8 8 8 8 1H.QI:H.V C%IG`1H:C1 8 8 8 8 8 8 8 1JQV :`IQ`1H:J: 8 8 8 8 8 8 8 Q .`1: HQJH.1CV$: 8 8 8 8 8 8 8 Q QI: % :GV`:J 8 8 8 8 8 8 8 Q QI: % C: V`1HV% 8 8 8 8 8 8 8 Q QI: % C1JV: % 8 8 8 8 8 8 8 J%].1 V`VI1 : 8 8 8 8 8 8 8 'J%].1 ] 8 8 8 8 8 8 8 J%].1 ]8 8 8 8 8 8 8 8 '`G1J1: H%01V`1 8 8 8 8 8 8 8 1VJ1: `%1`Q`I1 8 8 8 8 8 8 8 :CQC: 1H1C1VJ1 8 8 8 8 8 8 8 :J .:C1 QV` VR1 8 8 8 8 8 8 8 139 :`:C:H7RQJ1: ]:`:RQ6: 8 8 8 8 8 8 8 :`:QJ1 `%C$VJ 8 8 8 8 8 8 8 :`:]`1QJQ]1Q H` ]1JJ: : 8 8 8 8 8 8 8 VH 1J:`1: :%`1HQI: 8 8 8 8 8 8 8 VH 1J:`1: @Q`VJ1 8 8 8 8 8 8 8 V :CQ]`QH % V``1HQC% 8 8 8 8 8 8 8 V : ]%1CC: 8 8 8 8 8 8 8 .V`%: IQJ1C1`V`: 8 8 8 8 8 8 8 .V`%: ]C%IQ: 8 8 8 8 8 8 8 .7CCQRQHV H` I:H%C: : 8 8 8 8 8 8 8 .7CCQRQHV C:I1JQ: 8 8 8 8 8 8 8 .7CCQRQHV C1JV: : 8 8 8 8 8 8 8 .7CCQRQHV I:H`Q]. .:CI: 8 8 8 8 8 8 8 .7CQ `QV 1R: 8 8 8 8 8 8 8 .7CQ C1$% 1H: 8 8 8 8 8 8 8 .7CQ JQ`0V$1H: 8 8 8 8 8 8 8 1C:`$1 0V``%HQ: 8 8 8 8 8 8 8 1`QI1 V`%H: 8 8 8 8 8 8 8 1 : H`1 : : 8 8 8 8 8 8 8 QVH1CQH.:V % V`]VJ 8 8 8 8 8 8 8 QC7H1``% :%`:J 1:H% 8 8 8 8 8 8 8 QC7QRQJ V I:61CCQ% 8 8 8 8 8 8 8 QC7Q]. :CI% ]1H % 8 8 8 8 8 8 8 QI: QHV`Q C:I:`H@11 8 8 8 8 8 8 8 `:61CCVCC: :``1J1 8 8 8 8 8 8 8 `:61CCVCC: $`:H1C1 8 8 8 8 8 8 8 `1QJQ]1Q H:]V`1 8 8 8 8 8 8 8 1J%]1Q H1``1`V`: 8 8 8 8 8 8 8 `1QJQ]1Q V.CV`1 8 8 8 8 8 8 8 `1QJQ]1Q `:CC:6 8 8 8 8 8 8 8 #_%1C:]1Q V6QH%C: : 8 8 8 8 8 8 8 `1QJQ]1Q VVJ `%]1 8 8 8 8 8 8 8 `Q QRQ`01CCV: @V`V` V1J1 8 8 8 8 8 8 8 `Q %C: ]8 8 8 8 8 8 8 8 :IIQC7HV :`VJQ: 8 8 8 8 8 8 8 V%RQCV1QH:]1 VCC: `:%0VC1 8 8 8 8 8 8 8 %:GVCC1RV QH QH1``: : 8 8 8 8 8 8 8 %H:C1G`V$I: 1J`C: %I 8 8 8 8 8 8 8 Q`01CCV: ^ H.1 QIV`1J$Q_ JV$CVH : 8 8 8 8 8 8 8 .Q`01CCV: ^%H.1 QIV`1J$Q_ `%RQC].11 8 8 8 8 8 8 8 %H.CV`QH.V1C% I1J% % 8 8 8 8 8 8 8 HQC:`1H1: 7]1H: 8 8 8 8 8 8 8 %HQCVCV]1 H:J :G`: 8 8 8 8 8 8 8 QC7RQ`: H1C1: : 8 8 8 8 8 8 8 HQCVCV]1 `QC1Q: 8 8 8 8 8 8 8 %HQCVCV]1 _%:I: : 8 8 8 8 8 8 8 %HQCVCV]1 `1RVJ : : 8 8 8 8 8 8 8 HQCQ]CQ ^ HQCQ]CQ_ :`I1$V` 8 8 8 8 8 8 8 %V`]%C: HQJH.:`%I 8 8 8 8 8 8 8 1$:C1QJ I: .1CR:V 8 8 8 8 8 8 8 %1$:C1QJ _%:IQ%I 8 8 8 8 8 8 8 1$:IG`: VJ :H%C: : 8 8 8 8 8 8 8 ]1Q RVHQ`: % 8 8 8 8 8 8 8 ]1Q `1C1HQ`J1 8 8 8 8 8 8 8 %]1Q I%C 1QH%C: : 8 8 8 8 8 8 8 ]1QH.:V Q] V`% HQ :`%I 8 8 8 8 8 8 8 ]1QJ1R:V]8 8 8 8 8 8 8 8 140 ]1Q].:JV GQIG76 8 8 8 8 8 8 8 %]1Q].:JV @`Q7V`1 @`Q7V`1 8 8 8 8 8 8 8 %]1Q].:JV ]8 8 8 8 8 8 8 8 % V`J:]1 H% : : 8 8 8 8 8 8 8 % .VJVC:1 GQ: 8 8 8 8 8 8 8 % .VJVC:1 C1I1HQC: 8 8 8 8 8 8 8 *V`VGVCC1RV `QVI1 8 8 8 8 8 8 8 V`I1C1Q]1 1J`%JR1G%C%I 8 8 8 8 8 8 8 141 #]]VJR16 7 : Q` : <85 :`$%1 285 : :J<1 85 : :CCV 85 :$:JVCC185 1HQCV 1 8 8 VC1H :JRR`VR$1J$`Q`GV:H.JQ%`1.IVJ 1J .VHVJ `:C 7``.VJ1:JV:^ :C7_7V``VH QJ GVJ .1H:VIGC:$V8 :`1JV $HQCQ$7 5 ^%]]C8_7 R8 :Q` :<85 QI:V 185 QI1`1852:`<1:CV 185:J185VJJ:285 :JV`:8:JR 1HQCV 1 8R$J%]R: VQ` .VVHQCQ$7:JR]: 1:CR1 `1G% 1QJQ`QIV]QC7H.:V V ]VH1V``QI .V :C1:JHQJ 1JVJ :C.VC`8 @ :C1:J :Q%`J:C Q` QQCQ$7 5%]]C87 R 8 142 Italian Journal of Zoology , 2011, i First , 1–14 Ecology and spatial distribution of selected polychaete species from the Italian continental shelf B. LA PORTA 1* , P. TOMASSETTI 1, S. LOMIRI 1, S. MARZIALETTI 2, D. VANI 1, M. PENNA 1, P. LANERA 1, & L. NICOLETTI 1 1ISPRA – Institute for Environmental Protection and Research, Rome, Italy and 2Marine Ecological Surveys Ltd, Bath, UK (Received 26 June 2010; accepted 12 May 2011) Abstract Data from several research and monitoring projects carried out between 1999 and 2009, for a total of 26 study areas located along the Italian continental shelf (Mediterranean Sea), were extracted from ISPRA’s data set, to revise and update the existing information on the ecology and spatial distribution of 20 selected soft-sediment polychaete species. The species were selected taking into account their spatial distribution and ecological role in the benthic assemblages and the existence of voucher specimens deposited in ISPRA’s reference collection. Samplings were taken at 872 stations on soft sediments, at depths ranging from 1 to 155 m. Surface sediment composition data were available at each site. The number of specimens from the selected species was extracted at each site, and relative abundance (%)calculated. The spatial distribution of each species was investigated according to the biogeographical zones identified in the Italian Seas. The distribution of five species (Aponuphis bilineata , A. brementi , A. fauveli , Nothria conchylega , and Onuphis eremita ) was updated. Several species that were previously considered to be characteristic of a specific biocenosis, sensu Pérès & Picard (1964), e.g. Diopatra neapolitana , Ditrupa arietina , Notria conchylega and Sternaspis scutata , were found to be distributed over a wider bathymetric and granulo- metric range of surface sediments. Indicator Species Analysis highlights that the distribution of 17 selected species depends on definite granulometric characteristics of the sediment. This new relevant information outlines the important contribution of environmental monitoring programmes to scientific knowledge. Keywords: Polychaete assemblages , biocoenosis , soft bottoms , sediment texture , Mediterranean Sea Introduction Information on species distribution is incomplete, and many species are frequently recorded outside Polychaetes are amongst the most frequent and Downloaded by [B. La Porta] at 06:00 23 August 2011 their known bathymetric and granulometric range abundant organisms characterizing marine benthic and of their characteristic ecological boundaries, communities, accounting for up to more than a which define specific biocoenosis sensu Pérès & third of the total number of macrobenthic species in Picard (1964). There is only a general account, soft substrata (Day 1967; Knox 1977). Polychaetes dating back to Bellan (1964), on the ecology of have proven to be excellent indicators of environ- Mediterranean polychaetes. Further studies only mental conditions due to their distribution over a analysed the distribution of polychaetes at local scales broad range of environments and their ample display (e.g. Gravina 1986; Castelli et al. 1992; Bianchi of ecological requirements. Because of their eco- et al. 1993a,b,c; Crema et al. 1993; Simboura et al. logical variability, they are widely used in applied 2000; Çinar 2005; Moreira et al. 2006; Cosentino & environmental research (Giangrande et al. 2005). Giacobbe 2008; Zaâbi et al. 2009). The information Although there have been numerous studies inves- on the ecological and spatial distribution of species tigating the systematics, taxonomy, morphology and should therefore be revised and updated on the base ecology of Mediterranean polychaetes, there are still on newly collected information. significant gaps in the understanding of their ecology. *Correspondence: B. La Porta, ISPRA, Institute for Environmental Protection and Research, Rome, Italy. Email: [email protected] ISSN 1125-0003 print/ISSN 1748-5851 online © 2011 Unione Zoologica Italiana DOI: 10.1080/11250003.2011.588443 2 B. La Porta et al. Environmental monitoring programmes can be a In this study we present a first update on the spa- source of valuable scientific data to increase the tial distribution and ecology (related to sediment scientific knowledge of benthic communities and composition and depth range) of 20 soft-sediment species, as these programmes allow the integration polychaete species along the Italian continental shelf. of multidisciplinary information. Moreover, frequent These species were selected taking into account their monitoring surveys and common sampling method- spatial distribution and their ecological role in ben- ologies allow the collection of numerous and compa- thic assemblages. For each species, voucher speci- rable data from different study areas. mens were available in ISPRA’s reference collection, This study proves the relevance of multidisci- which is based on the material collected in numerous plinary environmental characterization and monitor- monitoring surveys. ing programmes for scientific research collating data extracted from a number of surveys (e.g. sand dredg- Material and methods ing, offshore platforms developments, aquaculture, beach nourishments) carried out in Italy over the Data from 26 study areas located along the Italian last 10 years by the Italian National Institute for continental shelf were extracted from ISPRA’s data Environmental Protection and Research (ISPRA). set (Figure 1). These locations were selected with These projects have provided data on sediment com- base on the common procedures of data collection position, associated benthic communities and depth and analyses at each sampling period from 1999 to range of a number of species in different areas of the 2009. Samples had been taken in two replicates at Italian continental shelf. 872 stations along transects perpendicular to the Downloaded by [B. La Porta] at 06:00 23 August 2011 Figure 1. Location of the 26 study areas from which data was extracted. Stations are numbered from the Ligurian Sea to the northern Adriatic Sea. The dimension of the circle is proportional to the number of stations investigated in each area. 1, Rosignano; 2, Elba; 3, Porto Ercole; 4, Montalto; 5, Civitavecchia; 6, Ostia; 7, Torpaterno; 8, Torvaianica; 9, Anzio; 10, Sabaudia; 11, Terracina; 12, Gaeta; 13, Ponza; 14, Porto Torres; 15, Olbia; 16, Baia; 17, Bagnoli; 18, Castellammare; 19, Bisceglie; 20, Molise; 21, Ortona; 22, Giulianova; 23, Marche; 24, Civitanova Marche; 25, Ravenna; 26, Chioggia. The nine biogeographical zones described in the Italian Seas are as according to Bianchi (2004) and followed in the checklist of flora and fauna of the Italian Seas (Relini 2008) Ecology of selected polychaete species from the Italian coast 3 Table I. The 26 study areas located in the nine biogeographical zones identified in the Italian Seas (Bianchi 2004), the number of sampling stations for each area, divided into depth ranges ( C = Central; S = South; N =North). Study Areas Sea Biogeographic Total number of Depth ranges (m) Zone Stations 0-10 10,1-20 20,1-30 30,1-50 >50 1. Rosignano Ligurian Sea 1 35 7 15 7 4 2 2. Elba N Tyrrhenian Sea 1 −2 41 2 2 15 22 3. Porto Ercole N Tyrrhenian Sea 2 11 11 4. Montalto C Tyrrhenian Sea 2 34 2 12 20 5. Civitavecchia C Tyrrhenian Sea 2 70 1 5 21 43 6. Ostia C Tyrrhenian Sea 2 68 4 27 14 23 7. Torpaterno C Tyrrhenian Sea 2 26 18 8 8. Torvaianica C Tyrrhenian Sea 2 50 4 4 4 12 26 9. Anzio C Tyrrhenian Sea 2 31 4 7 4 16 10. Sabaudia C Tyrrhenian Sea 2 57 4 7 5 13 28 11. Terracina C Tyrrhenian Sea 2 50 7 5 2 10 26 12. Gaeta C Tyrrhenian Sea 2 27 5 4 9 9 13. Ponza C Tyrrhenian Sea 2 19 3 16 14. Porto Torres Sardinian Sea 2 17 5 12 15. Olbia C Tyrrhenian Sea 2 10 10 16. Baia C Tyrrhenian Sea 3 17 4 5 4 4 17. Bagnoli C Tyrrhenian Sea 3 33 33 18. Castellammare S Tyrrhenian Sea 3 21 3 17 1 19. Bisceglie S Adriatic Sea 7 11 11 20.Molise CAdriaticSea 8 52 20 11 11 7 3 21. Ortona C Adriatic Sea 8 5 5 22. Giulianova C Adriatic Sea 8 28 10 7 4 5 2 23. Marche C Adriatic Sea 8 75 75 24. Civitanova Marche C Adriatic Sea 8 23 23 25. Ravenna N Adriatic Sea 9 9 9 26. Chioggia N Adriatic Sea 9 52 7 4 41 coast, at depths ranging from 1 to 155 m (Table I), ranges (1–10 m; 10.1–20 m; 20.1–30 m; 30.1–50 m; using a Van Veen grab (0.1 m 2 covering area). The >50 m). sediments sampled were sieved through a 1-mm The spatial distribution of each species was fur- mesh and the retained material was preserved in ther analysed according to the division of the Italian seawater adding 4% CaCO 3-buffered formalin. Seas into nine biogeographical zones, proposed by Downloaded by [B. La Porta] at 06:00 23 August 2011 Macrozoobenthic samples were further sorted Bianchi (2004) (Figure 1) and applied in the Italian into major taxonomical groups and the collected Checklist of marine flora and fauna (Relini 2008). polychaetes counted and classified to the lowest No study areas were available for this study from possible taxonomic level. Surface sediments were biogeographical zones 4, 5 and 6. collected at each station with a box-corer, analysed Where available, information on the selected and classified according to sand percentage, follow- species on biocoenosis affiliation (Bellan 1964; Pérès ing Nota (1958): sand (S) ( >95% content of sand), & Picard 1964; Picard 1965; Augier 1982), preferred muddy sand (mS) (95–70%), very sandy mud (vsM) sediment typologies and depth range distribution was (70–30%), sandy mud (sM) (30–5%) and mud (M) also extracted from existing literature. (<5%). Indicator Species Analysis (ISA) (Dufréne & Twenty species, listed in Table II, were selected Legendre 1997) has been performed in order to according to their spatial distribution, their ecolog- identify the species associated with or indicative ical role and the existence of voucher specimens of five groups of stations, derived from Nota deposited in ISPRA’s reference collection. For each (1958) classification. This analysis combines the rel- species number of specimens and relative abun- ative abundance of the species with their relative dance (%) were calculated. Box-and-whisker plots frequency of occurrence in the various groups and (Tukey 1977) were performed to analyse the depth provides an indicator value subsequently tested by distribution of species. Depths were grouped in five randomization. 4 B. La Porta et al. Table II. Distribution of the 20 selected species by the nine biogeographical zones of the Italian Seas. In white, the distribution of the selected species as reported in the Checklist of Flora and Fauna of the Italian Seas (Relini 2008); in grey, records confirming the species distribution; in black, new records distribution of five species ( Aponuphis bilineata , A. brementi , A. fauveli , Nothria conchylega , and Onuphis eremita ). Species Biogeographical Zones of Italian Seas 123456789 Aponuphis bilineata + + + + + + + + + (Baird, 1870) Aponuphis brementi + ++++++++ (Fauvel, 1916) Aponuphis fauveli + + + + + + + + + (Rioja, 1918) Chaetozone caputesocis +++++++++ (Saint-Joseph, 1894) Chaetozone gibber + ++ +++ Woodham & Chambers, 1994 Clymenura leiopygos +++ +++ (Grube, 1860) Diopatra neapolitana + +++ ++++ Delle Chiaje, 1841 Ditrupa arietina +++ +++ + (O.F. Muller, 1776) Drilonereis filum + + + + + + + + + (Claparède, 1868) Eunice vittata +++++++++ (Delle Chiaje, 1828) Heteromastus filiformis (Claparède, + + + + + + + + + 1864) Hyalinoecia tubicola +++++++++ (O.F. Müller, 1776) Melinna palmata +++ ++ + + + Grube, 1870 Metasychis gotoi +++ + + + + (Izuka, 1902) Nematonereis unicornis +++++++++ (Grube, 1840) Nothria conchylega + + + + + + + + (M. Sars, 1835) Onuphis eremita + + + + + + Audouin & Milne-Edwards, 1833 Owenia fusiformis +++ ++ + + + Downloaded by [B. La Porta] at 06:00 23 August 2011 Delle Chiaje, 1841 Pectinaria auricoma +++++ ++ + (O.F. Müller, 1776) Sternaspis scutata + ++ ++ + + + (Ranzani, 1817) Results and discussion of five of these species (in black). Aponuphis bilin- eata , A. brementi and A. fauveli , were recorded along A total of 55,000 individuals from 200 species of the northern coast of the Island of Elba (Tuscan polychaetes distributed across the Italian biogeo- Archipelagos) in zone 1, which is considered to graphical zones 1, 2, 3, 7, 8, and 9 (Figure 1) were be a climate transitional area between zones 1 and identified. 2 (Relini 2008). Nothria conchylega was found in The allocation of the 20 selected species by bio- zone 2, in the southern coast of the Island of Elba geographical zones, as reported in the Checklist of and along the coast of Latium (Central Tyrrhenian flora and fauna of the Italian seas (Relini 2008), is Sea) and Onuphis eremita was collected in zone 7, presented in white in Table II. The results confirmed along the eastern coast of Apulia (Southern Adriatic the presence of these species in some biogegraph- Sea). ical zones (in grey) and updated the distribution Ecology of selected polychaete species from the Italian coast 5 In Figure 2, box-and-whisker plots describe the exclusively from shallow waters to 50 m depth, depths distribution of the selected species. The anal- Diopatra neapolitana was mostly distributed in shal- ysis of the distribution revealed that most of the low areas (1–10 m depth) and absent below 30 m, 20 species occurred at all depth ranges (Figure 3). Hyalinoecia tubicola was sampled deeper than 20 m, Amongst the exceptions, Chaetozone gibber occurred Metasychis gotoi and Sternaspis scutata were only A. bilineata A. brementi A. fauveli C. caputesocis 90 112 128 90 80 96 80 112 70 70 80 96 60 60 50 64 80 50 48 64 40 40 30 Depth (m) 32 48 30 32 20 20 16 16 10 10 0 0 0 0 C. gibber C. leiopygos D. neapolitana D. arietina 45 72 27 54 40 64 24 48 35 56 21 42 30 48 18 36 25 40 15 30 20 32 12 24 Depth (m) 15 24 9 18 10 16 6 12 5 8 3 6 0 0 0 0 D. filum E. vittata H. filiformis H. tubicola 180 64 90 108 160 80 56 96 140 70 84 48 120 60 72 40 50 60 100 32 40 48 80 Depth (m) 30 24 36 60 20 16 24 40 10 8 12 20 0 0 0 0 Downloaded by [B. La Porta] at 06:00 23 August 2011 M. palmata M. gotoi N. unicornis N. conchylega 108 128 108 112 96 112 96 96 84 84 96 72 80 72 80 60 64 60 64 48 48 48 48 36 36 32 Depth (m) 24 32 24 16 12 12 16 0 0 0 0 O. eremita O. fusiformis P. auricoma S. scutata 80 72 108 96 96 70 64 84 56 84 60 72 48 72 50 60 40 60 40 48 32 48 36 30 24 36 20 Depth (m) 24 16 24 12 10 8 12 0 0 0 0 Figure 2. Box-and-whisker plots describe the distribution of depths where species were found. 6 B. La Porta et al. Figure 3. Distribution of the 20 selected species along the different depth ranges (in meters), taking into account the percentage of analysed stations, for each depth range, where the species were found. The maximum found percentage was 90%. found deeper than 10 m, and Onuphis eremita , was granulometric characteristics of the sediment and the absent in the 30–50 m depth range. indicator values of ISA statistic specify the groups The species distributions show that all species were of stations that are associated with the species with found in all sediment classes (Nota’s sediment classi- different levels of significance. It is interesting to fication) except Diopatra neapolitana , which was not highlight that 17 of the selected species are associ- found in sandy-muds (sM) and Metasychis gotoi and ated with specific groups with a high level of sig- Pectinaria auricoma , which were absent from sands nificance ( p≤0.01 and p≤0.001) while Hyalinoecia (S) (Figure 4). tubicola , Melinna palmata and Nothria conchylega are The species selected were present in the range associated with a low level of significance ( p≤0.05). of 0–20% relative abundance within the associated Chaetozone gibber , Clymenura leiopygos and Diopatra polychaete assemblages in all sediment typologies; neapolitana have no significant relationship with higher values of relative abundance were recorded the groups identified, suggesting a wide ecological only for some species and in some classes of sediment tolerance against the sediment fractions. Moreover, (Table III). Drilonereis filum and Eunice vittata have values of ISA The species-group contributions obtained from that suggest a high tolerance to mixed sediment (vsM ISA are reported in Table IV. Results demonstrate and sM) and Sternaspis scutata show a significant that the species distribution depends on definite preference for muddy sediment (sM and M). Downloaded by [B. La Porta] at 06:00 23 August 2011 Figure 4. Distribution of the 20 selected species amongst different sediment classes according to Nota’s classification (1958): sand (S) (>95% content of sand), muddy sand (mS) (70–95% content of sand), very sandy mud (vsM) (70–30% content of sand), sandy mud (sM) (30–5% content of sand) and mud (M) ( <5% content of sand). The distributions take into account the percentage of analysed stations, for each sediment class, where the species were found. The maximum found percentage was 63%. Ecology of selected polychaete species from the Italian coast 7 Table III. Frequency of station in each Nota’s sediment class where each species was found. Species occurrence is also clustered into classes of relative abundances (%) of each species within the whole polychaete assemblages (S = sand; mS =muddy sand; vsM =very sandy mud; sM =sandy mud; M =mud). 0-20 % 21-40 % 41-60 % 61-80 % S mS vsM sM M S mS vsM sM M S mS vsM sM M S mS vsM sM M Aponuphis bilineata 15 123 41 9 31 1 16 2 Aponuphis brementi 10 63 52 10 2 3 7 2 Aponuphis fauveli 940631011 4 1 3 3 1 Chaetozone caputesocis 6 16 27 10 7 1 1 Chaetozone gibber 5 6151119 1 Clymenura leiopygos 1921212232 1 1 Diopatra neapolitana 327 8 Ditrupa arietina 142456 2 Drilonereis filum 128582946 2 1 Eunice vittata 4 32 66 39 41 Heteromastus filiformis 616423754 13 Hyalinoecia tubicola 2 12 24 5 4 Melinna palmata 570764473 7145 1 Metasychis gotoi 15212212 2 2 Nematonereis unicornis 3 49 45 20 21 Nothria conchylega 5251021 1 1 Onuphis eremita 12 9 3 210 1 1 Owenia fusiformis 5616157 531710 1 42 31 Pectinaria auricoma 71 28 12 10 Sternaspis scutata 452575469 111013 212 25 Table IV. The species-group contributions obtained from Indicator Species Analysis (ISA). The indicator values of ISA specify the groups of stations that are associated with the species with different levels of significance. Species S mS vsM sM M P-level Aponuphis bilineata 1.326 25.808 6.499 0.970 - 0.000 Aponuphis brementi 0.449 4.619 16.738 0.796 0.001 0.000 Aponuphis fauveli 0.242 3.816 17.674 0.531 0.231 0.000 Caulleriella caputesocis 1.112 0.998 6.531 1.195 0.131 0.001 Caulleriella gibber 0.645 0.069 1.276 2.794 3.381 0.167 Clymenura leiopygos 4.518 0.911 5.358 3.627 0.823 0.362 Diopatra neapolitana 0.322 0.014 2.411 - 0.983 0.074 Ditrupa arietina 0.008 0.024 8.932 0.349 0.059 0.000 Drilonereis filum 0.006 0.431 11.515 8.242 3.989 0.002 Downloaded by [B. La Porta] at 06:00 23 August 2011 Eunice vittata 0.152 1.194 12.180 8.935 2.700 0.000 Heteromastus filiformis 0.320 0.747 6.284 1.223 0.050 0.000 Hyalinoecia tubicola 0.148 0.157 11.931 11.883 3.457 0.022 Melinna palmata 0.073 3.796 7.793 13.975 8.344 0.023 Metasychis gotoi - 0.401 5.442 8.945 0.362 0.001 Nematonereis unicornis 0.143 2.719 11.495 4.522 0.954 0.000 Nothria conchylega 0.903 5.518 0.944 0.108 0.002 0.021 Onuphis eremita 8.653 0.53 0.101 0.068 0.308 0.000 Owenia fusiformis 64.494 8.374 0.803 0.004 0.001 0.000 Pectinaria auricoma - 14.735 3.054 1.490 0.157 0.000 Sternaspis scutata 0.034 0.229 6.068 16.599 18.048 0.002 For each species, the information on biocoenoses Aponuphis bilineata (Onuphidae) was found from affiliation, ecological significance, sediment 4 to 100 m depth (Figure 2) although it was mostly typologies and depth ranges, as emerging from frequent between 20 and 30 m depth (Figure 3). The previous literature and from the results of this study, species mainly prefers muddy sand (mS) and very is summarized in Table V. These results are also sandy mud (vsM) (Figure 4 and Table IV), where listed below, in alphabetic order. gravels are present, as along the Island of Elba and 8 B. La Porta et al. e results of biocoenoses = species living on = tol. L CHARACTERISTICS biocoenoses of coastal terrigenous mud, mud (vsM), sandy mud (sM), mud (M) mud (vsM), sandy mud (sM), mud (M) mud (vsM), sandy mud (sM), mud (M) mud (vsM), sandy mud (sM), mud (M) mud (vsM), sandy mud (sM), mud (M) mud (vsM), sandy mud (sM), mud (M) mud (vsM), mud (M) sand (S), muddy sand (mS), very sandy sand (S), muddy sand (mS), very sandy Depth range (m) Sediment typology sand (S), muddy sand (mS), very sandy sand (S), muddy sand (mS), very sandy 4–100 m 5–116 m sand (S), muddy sand (mS), very sandy 5–90 m 3,5–80 m 1–43 m sand (S), muddy sand (mS), very sandy 2,5–73 m sand (S), muddy sand (mS), very sandy 3–22,7 m = from ISPRA’ dataset biocoenoses of muddy detritic, DL species living on mixed sediments. = = pth ranges, as emerging from previous literature and from th species living on sands and tolerant to other fractions, mud = biocoenoses of coastal detritic, DE = species with wide ecological distribution, Mixt. = biocoenoses of surface muddy sands in sheltered waters, VTC Chimenz Gusso et al. 2001 1993 Gambi & Giangrande 1986; Bellan1991; & Bianchi Bellan-Santini et al. 1993Cosentino b; & Chimenz Giacobbe Gusso 2008 et al. 2001; Intès & Le Loeuff 1986; Bianchi et al. 1993a, b, c = Crema et al. 1993; Intès & Le Loeuff 1986; Bianchi et al. 1993a, b, c; Cacabelos et al. 2008 ; Zaabi et al. 2009 Massé 1972; Giangrande & Gambi 1985; Crema et al. Bellan 1964; Picard 1965; Giangrande & Gambi 1985; Bellan 1964; Picard 1965; Gambi & Giangrande 1986; preferential species, sand tol. = coralligenous biocoenoses, DC = ies, pref. Downloaded by [B. La Porta] at 06:00 23 August 2011 ation, ecological significance, sediment typologies and de species living on gravely sediments, Lre = muddy sand, mixed sediments mud, very sandy mud. DE, Lre, Grav. Mixt. mud-tol. Bianchi et al. 1993 b DE, Mixt. sand, muddy sand, very fine sand, sand, detritic sand, mud Biocoenoses - ecological significance Depth range (vertical zonation) Sediment typology SFBC excl Circalittoral, Bathyal Circalittoral Infralittoral Sand, tolerant of small amount of biocoenoses of well-sorted fine sand, SVMC = Communities of unstable soft seabeds, excl. exclusive spec = of the shelf-edge detritic, SFBC muds and tolerant to other fractions, Grav. Table V. For each species,this information study. Results on are biocoenoses listed affili in alphabetic order. Abbreviations: C SPECIESAponuphis bilineata ECOLOGICALCHARACTERISTICS REFERENCES ECOLOGICA Aponuphis brementi Aponuphis fauveli Caulleriella caputesocis Chaetozone gibber MI Clymenura leiopygos Diopatra neapolitana Ecology of selected polychaete species from the Italian coast 9 L CHARACTERISTICS mud (vsM), sandy mud (sM), mud (M) mud (vsM), sandy mud (sM), mud (M) mud (vsM), sandy mud (sM), mud (M) mud (vsM), sandy mud (sM), mud (M) mud (vsM), sandy mud (sM), mud (M) Depth range (m) Sediment typology sand (S), muddy sand (mS), very sandy 22–155 m 3–90 m sand (S), muddy sand (mS), very sandy 1,5–99 m sand (S), muddy sand (mS), very sandy 5–51 m sand (S), muddy sand (mS), very sandy 1,5–60 m sand (S), muddy sand (mS), very sandy from ISPRA’ dataset UNEP 1986; / ˇ neda & Safran 1988; 1965; Picard. 1972; Diaz-Casta Bellan & Bellan-Santini 1991 et al. 1993a,b,c Giangrande 1986; Intès & LeBellan-Santini Loeuff 1991; 1986; Bianchi Bellan et & al. 1993a, b, c. Gravina & Somaschini 1990; DewarumezBianchi et et al. al. 1992, 1993a; Carvalho et al. 2005 1965; Augier 1982; Gambi &Grémare Giangrande et 1986; al. 1989; Crema et al. 1993 Gaillande 1968; Picard 1972; FAO Bellan 1961; Bellan 1964; Pérès & Picard 1964; Picard Pérès & Picard 1964; Bellan 1964; Picard 1965; Bianchi Bellan 1964; Picard 1965; Picard 1972; Gambi & Bellan 1961; Bellan 1964; Pérès & Picard 1964; Picard Pérès & Picard 1964; Bellan 1964; Picard 1965; De Downloaded by [B. La Porta] at 06:00 23 August 2011 sediment, sand, mixed sediments, median grain size sand, sedimentorganic with matter (Om) (with high fraction of sand),sand, detritic sediment with organic matter (Om) without specific ecological role, indicator of artificial or natural disturbance, euryhaline muddy enriched sediments DC pref., Mixt. SVMC excl., companion species Lre DC excl., DC, MI C, Lre, opportunist, colonizer Detritic muddy sand Depth range (vertical zonation) sandy mud, mud, mixed sediments Circalittoral, Bathyal Sediment typology mixed sediments Circalittoral, Infralittoral eurybathic Biocoenoses - ecological significance Eunice vittata Hyalinoaecia tubicola Drilonereis filum Heteromastus filiformis Table V. (Continued) SPECIESDitrupa arietina ECOLOGICALCHARACTERISTICS REFERENCES ECOLOGICA 10 B. La Porta et al. L CHARACTERISTICS sandy mud (sM), mud (M) mud (vsM), sandy mud (sM), mud (M) mud (vsM), sandy mud (sM), mud (M) mud (vsM), sandy mud (sM), mud (M) mud (vsM), sandy mud (sM), mud (M) sandy mud (sM), mud (M) mud (vsM), sandy mud (sM), mud (M) mud (vsM), sandy mud (sM), mud (M) sand (S), muddy sand (mS), very sandy Depth range (m) muddy sand (mS), very sandy mud (vsM), Sediment typology sand (S), muddy sand (mS), very sandy 13,5 –100 m muddy sand (mS), very sandy mud (vsM), 4,5 – 60,5 m sand (S), muddy sand (mS), very sandy 1,5 –73 m sand (S), muddy sand (mS), very sandy 5 –100 m 4 –23 m and 90 m sand (S), muddy sand (mS), very sandy 3–110 m sand (S), muddy sand (mS), very sandy 17,5–100 m 17,5–100 m from ISPRA’ dataset 1972; Zavodnik et al. 1985;1986;Crema Gambi et & al. Giangrande 1991; BianchiBen-Eliahu et & al. Fiege 1993a; 1995 Desbruyères et al. 1972-73; IntèsAbbiati & et Le al. Loeuff 1987; 1986; Hily 1987; Bianchi et al. 1993a,b,c Bianchi et al. 1993c,b,c Zavodnik et al. 1985; Bellan1986; 1985; Sardà Intès 1986; & Bianchi Le et Loeuff al. 1993a, b, c Gambi 1985; Gambi & GiangrandeLoeuff 1986; 1986; Intès Hily & 1987; Le Duineveldet et al. al. 1993b; 1991, Dauvin Bianchi 2000;Gonçalves Dos da Santos Silva Brasil 2000 & Picard 1965; Zavodnik et al. 1985, Cantone et al. 2004 Picard 1965; Bellan 1964; Bianchi et al. 1993a Pérès & Picard 1964; Bellan 1964; Picard 1965; Picard Pérès & Picard 1964; Bellan 1964; Picard 1965; Bellan 1964; Bellan 1963; Intès & Le Loeuff 1986 ; Bellan 1964; Picard 1965; Drago & Arbetelli 1978; Bellan 1963; Bellan 1964; Pérès 1982; Giangrande & Downloaded by [B. La Porta] at 06:00 23 August 2011 sheaths. Posidonia oceanica ecotonal zone enriched with organic matter (Om), sediment with organic matter (Om) DE pref., Mixt. VTC excl DE, Mixt. MI SFBC excl. Picard 1965; Bianchi et al. 1993a DC, SFBC DL, VTC, Mixt. Mud DE Mud. tol. sandy mud Eurybathic, Circalittoral, Bathyal Infralittoral, Circalittoral Sand tol., fine sand, sandy mud, sandy mud Circalittoral, Bathyal mud, sediment with organic matter Depth range (vertical zonation) Infralittoral, Circalittoral, Bathyal Sediment typology Mixed sediments, muddy fine sand, Biocoenoses - ecological significance Sternaspis scutata Pectinaria auricoma Onuphys eremita Owenia fusiformis Nothria conchylega Methasychis gotoi Nematonereis unicornis Table V. (Continued) SPECIESMelinna palmata ECOLOGICALCHARACTERISTICS REFERENCES ECOLOGICA Ecology of selected polychaete species from the Italian coast 11 western Sardinia coasts. Moreover, this species seems Ditrupa arietina (Serpulidae) was mostly collected to tolerate a high percentage of mud occurring in between 30 and 50 m depth, although its range spans both sandy mud and muddy stations (sM and M). from 5 to 51 m depth (Figures 2 and 3). According Our results partially agree with the existing informa- to our results, this species seems to tolerate different tion that signalled A. bilineata in detritic and sandy fractions of mud in the sediment, since it was mostly substrata and in mixed sediment (Table V). recorded on very sandy mud (vsM) and in other types Aponuphis brementi and Aponuphis fauveli of sediment, ranging from sand to mud (Figure 4 (Onuphidae) displayed similar sediment prefer- and Table IV). It is also interesting to underline that ences to the congeneric species A. bilineata showing the gravel fraction was very scarce in all the stations a significant association with very sandy mud (vsM) where D. arietina was recorded, although this is a (Figure 4 and Table IV). A. brementi was more species normally affiliated to detritic coastal assem- frequent below 20 m depth and up to 50 m while blages and it has been signalled on mixed sediments A. fauveli occurred below 30 m and up to 90 m (Table V). depth (Figures 2 and 3). Our results add information Drilonereis filum (Oenonidae) was found from 3 to to the existing literature where these species were 90 m depth, mostly below 20 m (Figures 2 and 3), described as ‘misticolous’ and A. fauveli in particular in all sediment types even if it showed a high toler- was affiliated to the muddy detritic biocoenoses ance to mixed sediment (vsM and sM) (Figure 4 and (Table V). Table IV); moreover, it was very rare in sandy (S) and Caulleriella caputesocis (Cirratulidae) was found shallow stations and up to 20 m depth. Our results, from 3.5 to 80 m depth (Figure 2) and more on the whole, confirm the information existing on frequently in deeper stations, below 30 m depth this species (Table V). (Figure 3). It occurred in stations with high per- Eunice vittata (Eunicidae) was found from 1.5 centages of mud (vsM, sM and M) (Figure 4 and to 60 m depth (Figure 2). It is mostly frequent Table IV), in accordance with the existing informa- on muddy substrata, preferably in very sandy mud tion (Table V), but also in few stations where the (vsM), sandy mud (sM) (Table IV) and mud (M) sandy fraction was ≥95–70% of the sediment (S (depth range 30–50m), rather than on sandy sedi- and mS). ments (S and mS) (Figures 3 and 4). The percentage Chaetozone gibber (Cirratulidae) was distributed in of gravel (mainly composed of organic detritic mat- all sediment typologies, with higher frequency in ter) was more abundant in vsM sediments which muddy stations (vsM, sM and M) (Figure 4) as indi- characterized stations located around Island of Elba cated in the literature (Table V). The species was and few stations of Baia (Gulf of Naples). Our mainly found in shallow water and never below 43 m results partially agree with previous scientific litera- depth (Figures 2 and 3). ture (Table V). Clymenura leiopygos (Maldanidae) was collected Heteromastus filiformis (Capitellidae) was recorded from 2.5 to 73 m (Figure 2) especially in stations between 1.5 and 99 m depth (Figures 2 and 3) located between 20 and 30 m depth (Figure 3). The in sediments characterized by a high percentages Downloaded by [B. La Porta] at 06:00 23 August 2011 presence of this species was signalled in sandy sta- of mud (vsM, sM and M) and only in few sandy tions, as well as in all the other sediment typologies, (S) and muddy sand (mS) stations (Figure 4 and with comparable frequencies (Figure 4). Our data Table IV). These results agree with the existing infor- show that C. leiopygos has a higher tolerance to dif- mation, which defined the species as an exclusive ferent fractions of mud than what expressed in the species of upper muddy-sand assemblages living also existing literature (Table V). in mud-rich sediments (Table V). Diopatra neapolitana (Onuphidae) was mostly Hyalinoaecia tubicola (Onuphidae) was sampled found in shallow stations (above 10 m depth) but exclusively below 20 m depth, frequently between 30 is present up to about 23 m depth (Figures 2 and 50 m (Figures 2 and 3). This species inhabited and 3). It was mainly collected in sediments with primarily very sandy mud (vsM), sandy mud (sM) high percentages of mud (vsM and M) and only in and muddy sand (mS) sediments and was found a few sandy stations (S and mS) (Figure 4). These in other sediment typologies in only a few stations results suggest that this species has a good tolerance (S and M) (Figure 4 and Table IV). In some of for mud, in partial agreement with previous litera- the stations where H. tubicola was recorded (mainly ture that considered D. neapolitana a species typical vsM and mS stations located in the Tyrrhenian Sea of fine well-sorted sand assemblages (Picard 1965) along the Island of Elba and the coast of Terracina and characteristic of sandy sediments, but tolerant to and Sabaudia), the percentage of gravel suggested small amounts of mud (Table V). the presence of detritic habitats where, according to 12 B. La Porta et al. the scientific literature, this species is most common (Figures 2 and 3). This species was mainly found in (Table V). stations characterized by a high percentage of sand (S Melinna palmata (Ampharetidae) was collected and mS) (Figure 4 and Table IV), in accordance with from 1 to 99 m depth (Figures 2 and 3), and results authors who described it as typical of fine well-sorted show that the species is strongly linked to the muddy sand assemblages (Table V). Moreover, a significant fraction of the sediment (Figure 4 and Table IV), in percentage of gravel characterized some of the sta- accordance with several authors (Table V). In fact, tions were the species was found at the Island of Elba, the stations where the species mostly occurred were Porto Torres (Western Sardinia), Montalto (Central characterized by an increasing percentages of mud Tyrrhenian Sea) and Marche (Central Adriatic Sea) (from mS to M classes). indicating that O. fusiformis inhabits also detritic sub- Metasychis gotoi has been defined as an alien strata, as stated by Pérès & Picard (1964) (Table V). species for the Mediterranean Sea (Cantone et al. Pectinaria auricoma (Pectinariidae) was mostly 2004; Occhipinti-Ambrogi et al. 2011). The species recorded between 20 and 30 m depth, although it was sampled, in comparable frequencies, in sedi- was collected in the range of 4.5–60.5 m (Figures 2 ments with a variable fraction of mud, with a signif- and 3), on sandy substrata with low fractions of mud icant preference for sandy mud (sM), and at depths (mS) (Figure 4 and Table IV). We noticed that the ranging from about 17 to 100 m (Figures 2–4 and increase of mud in the sediment led to the decrease of Table IV). This species was absent in shallow waters P.auricoma , and the species was absent in exclusively with sand percentages >95%. The data that we col- sandy sediment (S). Our results are partially in accor- lected on this species integrate the information on the dance with the common description of this species as substrata inhabited by M. gotoi (Table V). affiliated to the muddy-detritic assemblages (Picard Nematonereis unicornis distribution (Eunicidae) 1965) and as a mud tolerant species (Table V). highlights the wide bathymetric range of this species, Sternaspis scutata (Sternaspidae) occurred exclu- recorded from 3 to 110 m depth with highest fre- sively below 13 m depth, mainly between 20 and quencies between 30 and 50 m depth (Figures 2 30 m (Figures 2 and 3). Our results indicate that and 3). N. unicornis inhabits sediments with differ- this species inhabits primarily muddy sediments (sM ent percentages of mud (from mS to M) (Figure 4, and M), even if it was also found in several sta- Table IV) and – in some locations – with abun- tions with very sandy mud (vsM) or muddy sand dant fractions of gravel (Elba Island, Baia and Anzio, (mS) (Figure 4 and Table IV). These results partially Tyrrhenian Sea). These results confirmed the infor- confirm the previous information, which described mation about this species, which normally has been this species as typical of the terrigenous mud-shelf affiliated to muddy-detritic assemblages (Table V) assemblages and inhabiting sandy mud (Table V). and classified as one of the few taxa able to burrow inside Posidonia oceanica sheaths (Gambi 2002). Nothria conchylega (Onuphidae) was recorded from Conclusions 5 to 100 m depth with variable frequencies at differ- Downloaded by [B. La Porta] at 06:00 23 August 2011 ent depth ranges (Figures 2 and 3). It was mostly New relevant information emerged from the com- found in sediments with 70–95% sand (mS) and parison of the results of this study with pre-existing it was less frequent either at stations characterized knowledge. In fact, the data analysed allow us to exclusively by sand (S) or by lower percentages of update the distribution in the Italian Biogeographical sand (vsM, sM and M) (Figure 4 and Table IV). Zones of five species ( Aponuphis bilineata , A. bre- These results add information to previous literature, menti and A. fauveli , Nothria conchylega , and Onuphis which described N. conchylega as a species typical eremita ). of muddy sediment, often found in shelf terrigenous These results contribute to update the ecological mud and in shelf edge detritic assemblages (Table V). characteristics of the investigated species in relation Onuphis eremita (Onuphidae) was collected up to to depth range and sediment typologies, highlighting 90 m depth and mostly between 4 and 10 m; no significant associations of a number of species with individuals were found between 30 and 50 m depth definite sediment classes. Moreover, this results out- (Figures 2 and 3). O. eremita was frequent in sandy line that multidisciplinary data sets, as ISPRA’s, are stations (S and mS) (Figure 4 and Table IV), in essential in increasing the scientific knowledge on the accordance with the literature that affiliated this benthic communities and species that populates soft species to the biocoenoses of well-sorted fine sand seabed substrata. (Table V). Nevertheless, we signal that this species This study furthermore outlines the importance of was also frequent in mud (M). environmental monitoring programmes as a source Owenia fusiformis (Oweniidae) was present in the of important scientific data such as relevant infor- depth range 1.5–73 m with variable frequencies mation on marine invertebrate ecology. This work is Ecology of selected polychaete species from the Italian coast 13 meant to be a first contribution in the framework Bianchi CN. 2004. Proposta di suddivisione dei mari italiani in of a more extensive revision of our understanding settori biogeografici. Notiziario SIBM, 46:57–59. of the ecological requirements and the distribution Bianchi CN, Ceppodomo I, Galli C, Sgorbini S, Dell’Amico F, Morri C. 1993a. Benthos dei mari toscani. I: Livorno – Isola of polychaete species in soft sediments, improving d’Elba (crociera Enea 1985). In: Ferretti O, Immordino F, our knowledge on the functioning and evolution of Damiani V, editors. Arcipelago Toscano. Studio oceanografico, marine ecosystems. sedimentologico, geochimico e biologico. Rome: ENEA. pp. 263–290. Bianchi CN, Ceppodomo I, Niccolai I, Aliani S, De Ranieri S, Abbiati M, Dell’Amico F, Morri C. 1993b. Benthos dei mari Acknowledgements toscani. II: Isola d’Elba – Montecristo (crociera Enea 1986). We are grateful to the editor and the two refer- In: Ferretti O, Immordino F, Damiani V, editors. Arcipelago Toscano. Studio oceanografico, sedimentologico, geochimico e ees for their invaluable comments and contribution. biologico. Rome: ENEA. pp. 291–316. Many thanks to Dr Maria Cristina Gambi (Stazione Bianchi CN, Ceppodomo I, Cocito S, Aliani S, Cattaneo Vietti Zoologica Anton Dohrn of Naples) and Prof. R, Morri C. 1993c. Benthos dei mari toscani. III: La Spezia – Michele Scardi (University of Rome ‘Torvergata’), Livorno (crociera Enea 1987). In: Ferretti O, Immordino F, for their invaluable support on data analysis and for Damiani V, editors. Arcipelago Toscano. Studio oceanografico, sedimentologico, geochimico e biologico. Rome: ENEA. pp. the suggestions that improved this work considerably. 317–337. Thanks to the researchers and the students for their Cacabelos E, Moreira J, Troncoso JS. 2008. Distribution of assistance in field activities and for their time and Polychaeta in soft-bottoms of a Galician Ria (NW Spain). invaluable dedication to taxonomic identification. Scientia Marina 72:655–667. We want to express our fond memory and grati- Cantone G, Castelli A, Gambi MC, Giangrande A, Catalano D, Cigliano M, Como S, Musco L. 2004. Relazione finale tude also to Prof. Eugenio Fresi (University of Rome nell’ambito del programma di ricerca ‘Aspim’, area scientifica ‘Tor Vergata’) for the precious contribution he has della tassonomia dei policheti. ICRAM, Roma. made to marine biology and to the study of benthic Carvalho S, Moura A, Miguel BG, Pereira P, Cancela da Fonseca communities. L, Falcão M, Drago T, Leitão F, Regala J. 2005. Spatial Part of this study was supported by the local and inter-annual variability of the macrobenthic communities within a coastal lagoon (Óbidos lagoon) and its relationship authorities of the Regione Lazio and the Regione with environmental parameters. Acta Oecologica 27:143–159. Marche. Bioservice s.c.r.l. (Napoli) also contributed Castelli A, Crema R, Prevedelli D, Zunarelli Vandini R. 1992. with a set of data. Caratterizzazione bionomica di fondi molli infralitorali dell’ Alto Tirreno Toscano Atti della Società Toscana di Scienze Naturali, Memorie Serie B, 99:39–60. Chimenz Gusso C, Gravina MF, Maggiore FR. 2001. Temporal References variations in soft bottom benthic communities in Central Tyrrhenian Sea (Italy). Archivio di Oceanografia e Limnologia Abbiati M, Bianchi CN, Castelli A. 1987. Polychate vertical 22:175–182. zonation along a littoral cliff in the western Mediterranean. Çinar ME. 2005. Polychaetes from the coast of northern Cyprus P.S.Z.N.: Marine Ecology 8:33–48. (eastern Mediterranean Sea) with two new records for the Augier H. 1982. Inventaire et classification des biocénoses marines Mediterranean Sea. Cahiers de Biologie Marine 46:143–159. benthiques de la Méditerranée. Publication du Conseil de Downloaded by [B. La Porta] at 06:00 23 August 2011 Cosentino A, Giacobbe S. 2008. Distribution and functional l’Europe, Collection Sauvegarde de la Nature. pp. 25–59. response of sublittoral soft bottom assemblages to sedimentary Bellan G. 1961. Résultats scientifiques des campagnes de constraints. Estuarine, Coastal and Shelf Science 79:263–276. la ‘Calypso’. Annélides Polychètes. Campagne de la Crema R, Castelli A, Bonvicini Pagliai AM, Zunarelli Vandini Calypso (Septembre–Octobre 1955). Annales de l’Institut E, Prevedelli D, Alboni L. 1993. Studio delle comunità ben- Océanographique 39:161–178. toniche di fondi molli infralitorali dell’Alto Tirreno Toscano. Bellan G. 1963. Nouvelle contribution à l’étude de la microfaune In: Progetto Mare. Ricerca sullo stato biologico chimico e fisico annélidienne de la region de Marseille. Recueil des Travaux de dell’Alto Tirreno Toscano. Relazioni conclusive delle ricerche. la Station Marine d’Endoume 29:43–56. pp. 445–488. Bellan G. 1964. Contribution il l’étude systématique, bionomique Dauvin JC. 2000. The muddy fine sand Abra alba-Melinna palmata et écologique des Annelides Polychètes de la Méditerranée. community of the Bay of Morlaix twenty years after the Amoco Thèse Doct. Sci. Marseille. No. 135. Cadiz Oil Spill. Marine Pollution Bulletin 40:528–536. Bellan G. 1985. Effects of pollution and man-made modifica- Day JH. 1967. Monograph on the Polychaeta of southern Africa. tions on marine benthic communities in the Mediterranean: Vol. Part 1 Errantia. British Museum (Natural History), A review. In: Moraitou-Apostolopoulou M, Kiortsis V, edi- London. 458 pp. tors. Mediterranean Marine Ecosystems, NATO Conf. Ser. 1, De Gaillande D. 1968. Monographie des peuplements benthiques Ecology. New York, NY: Plenum Press. pp. 163–194. d’une calanque des côtes de Provence: Port-Miou. Recueil des Bellan G, Bellan-Santini D. 1991. Polychaetous annelids (exclud- Travaux de la Station Marine d’Endoume 44:358–401. ing Serpulidae) from artificial reefs in the Marseille area Desbruyères D, Guille A, Ramos JM. 1972–73. Bionomie ben- (French Mediterranean coast). Ophelia 5:433–442. thique du plateau continental de la côte catalane espagnole. Ben-Eliahu MN, Fiege D. 1995. Polychaeta from the continental Vie et Milieu 23:335–363. shelf and slope of Israel collected by the ’Meteor’ 5 expedition Dewarumez JM, Davoult D, Glacon R. 1992. Nouvelles signalisa- (1987). Senckenbergiana Maritima 25:85–105. tions d’espèces macrobenthiques sur les côtes françaises de la 14 B. La Porta et al. Manche orientale et de la Mer du Nord. I. Annélides. Cahiers Knox GA. 1977. The role of polychaetes in benthic soft-bottom de Biologie Marine 33:83–93. communities. In: Reish DJ, Fauchald K, editors. Essays on Diaz-Castañeda V, Safran P. 1988. Dynamique de la colonization Polychaetous Annelids in Memory of Dr. Olga Hartman. Los par les annélides polychètes de sédiments défaunés par la pol- Angeles, CA: Allan Hancock Foundation. pp. 547–604. lution dans des enceintes expérimentales en rade de Toulon Massé H. 1972. Contribution à l’étude de la macrofaune (France). Oceanologica Acta 11:285–297. de peuplements des sables fins infralittoraux des côtes de Dos Santos Brasil AC, Gonçalves da Silva SH. 2000. Spatial dis- Provence. VI. Données sur la biologie des espèces. Téthys 4: tribution of Polychaeta in a soft-bottom community at Saco do 63–84. Céu, Ilha Grande, Rio de Janeiro, Brazil. Bulletin of Marine Moreira J, Quintas P, Troncoso JS. 2006. Spatial distribution of Science 67:103–112. soft-bottom polychaete annelids in the Ensenada de Baiona Drago N, Albertelli G. 1978. Étude faunistique et bionomique du (Ría de Vigo, Galicia, noth-west Spain). Scientia Marina 70S3: littoral de Cogoleto (golfe de Genes). Téthys 8: 203–212. 217–224. Dufrêne M, Legendre P. 1997. Species assemblages and indica- Nota DJG. 1958. Sediments of the western Guiana shelf. Reports tor species: The need for a flexible asymmetrical approach. of the Orinoco Shelf Expedition, 2. Mendedelingen van de Ecological Monographs 67:345–366. Landbouwhogeschool te Wageningen 58: 98 pp. Duineveld GCA, Kunitzer A, Niermann U, De Wilde PAWJ, Gray Occhipinti-Ambrogi A, Marchini A, Cantone G, Castelli A, JS. 1991. The macrobenthos of the North Sea. Netherlands Chimenz C, Cormaci M, Froglia C, Furnari G, Gambi MC, Journal of Sea Research 28:53–65. Giaccone G, et al. 2011. Alien species along the Italian coasts: FAO /UNEP. 1986. Report of the FAO /UNEP meeting on the An overview 2011. Biological Invasions 13:215–237. effects of pollution on marine ecosystems (Blanes, Spain, 7–11 Pérès JM. 1954. Contribution à l’étude des Annélides Polychètes October 1985). FAO fisheries report. 352, pp. 20. de la Méditerranée occidentale. Recueil des Travaux de la Gambi MC. 2002. Spatio-temporal distribution and ecological Station Marine d’Endoume 13:83–155. role of polychaete borers of Posidonia oceanica (L.) Delile Pérès JM. 1982. Zonations. General features of organismic assem- scales. Bulletin of Marine Science 71:1323–1331. blages in pelagial and benthal. Structure and dynamics of Gambi MC, Giangrande A. 1986. Distribution of soft-bottom assemblages. Major benthic assemblages. Specific benthic polychaetes in two coastal areas of the Tyrrhenian Sea (Italy): assemblages. In: Kinne O, editor. Marine Ecology. London: structural analysis. Estuarine, Coastal and Shelf Science John Wiley & Sons. pp. 9–66, 119–186, 375–581. 23:847–862. Pérès JM, Picard J. 1964. Nouveau manuel de bionomie benthique Giangrande A, Gambi MC. 1985. Distribution of soft bot- de la Mer Méditérranée. Recueil des travaux de la Station tom Polychaetes in the Gulf of Salerno (Tyrrhenian Sea). Marine d’Endoume 31:137 pp. Rapport de la Commission Internationale pour l’Exploration Picard C. 1972. Les peuplements de vase au large du Golfe de Scientifique de la Mer Méditerranée 29:233–235. Fos. Téthys 3:603–607. Giangrande A, Licciano M, Musco L. 2005. Polychaetes as Picard J. 1965. Recherches qualitatives sur les biocoenoses environmental indicators revisited. Marine Pollution Bulletin marines des substrats meubles dragables de la région marseil- 50:1153–1162. laise. Recueil des Travaux de la Station Marine d’Endoume Gravina MF. 1986. Analisi della distribuzione dei policheti nei 36:1–160. fondi mobili costieri delle Cinque Terre (Liguria). Bollettino Relini G. 2008. Checklist of flora and fauna of the Italian seas. dei Musei e degli Istituti Biologici dell’Università di Genova 2008. Biologia Marina Mediterranea 15(Suppl. 1): 385 pp. 52(Suppl.):197–212. Sardà R. 1986. Fauna de Anélidos Poliquetos de la Región del Gravina MF, Somaschini A. 1990. Censimento dei policheti Estrecho de Gibraltar, III – Eunicida, Orbiniida, Spionida, dei mari italiani: Capitellidae Grube, 1862. Atti della Magelonida, Chaetopterida, Ctenodrilida, Flabelligerida, Società Toscana di Scienze Naturali. Memorie. Serie B, 97: Opheliida, Oweniida, Capitellida, Terebellida, Sabellida y 259–285. Nerillida. Miscellània Zoològica 10:71–85. Downloaded by [B. La Porta] at 06:00 23 August 2011 Grémare A, Amouroux JM, Vétion G. 1998. Long-term compar- Simboura N, Nicolaidou A, Thessalou-Legaki M. 2000. ison of macrobenthos within the soft bottoms of the Bay of Polychaete community of Greece: An ecological overview. Banyuls-sur-Mer (Northwestern Mediterranean Sea). Journal Marine Ecology 21:129–144. of Sea Research 40:281–302. Tukey JW. 1977. Box-and-whisker plots. Exploratory Data Hily C. 1987. Spatio-temporal variability of Chaetozone setosa Analysis 39–43. (Malmgren) populations on an organic gradient in the Bay Zaâbi S, Gillet P, Afli A, Mboumaiza M. 2009. Biodiversity of Brest, France. Journal of Experimental Marine Biology and of polychaetous annelids from the peninsula of Cap Bon, Ecology 112:201–216. northeast coast of Tunisia. Zoosymposia 2:587–600. Intès A, Le Loeuff P. 1986. Les Annélides Polychètes de Côtes Zavodnik D, Vidakovic J, Amoureux L. 1985. Contribution to sed- d’Ivoire. V. Mise en evidence et description des peuplements. iment macrofauna in the area of Rovinj (North Adriatic Sea). Tropical Oceanography 21:117–142. Cahiers de Biologie Marine 26:431–44. Marine Ecology. ISSN 0173-9565 ORIGINAL ARTICLE Relict sand dredging for beach nourishment in the central Tyrrhenian Sea (Italy): effects on benthic assemblages Barbara La Porta, Monica Targusi, Loretta Lattanzi, Paola La Valle, Daniela Paganelli & Luisa Nicoletti ISPRA formerly ICRAM, Central Institute for Marine Research, Rome, Italy Keywords Abstract Environmental monitoring; macrozoobenthos; Mediterranean Sea; recolonisation process; The aim of this study is to analyse the effects in space and time of relict sand- sand extraction. dredging activities on macrobenthic assemblages, in an area situated offshore Montalto di Castro (central Tyrrhenian Sea, Italy), and to analyse the recoloni- Correspondence sation processes of macrobenthos in the dredged areas. The area in question is B. La Porta, ISPRA formerly ICRAM, Central characterised by relict sand deposits (Holocenic paleo-beaches), used for beach Institute for Marine Research, via di Casolotti nourishment along the Latium coast. The effects of sand extraction on benthic 300, 00166 Rome, Italy. E-mail: [email protected] assemblages were investigated before, during and after three dredging opera- tions. The sites analysed are located within the dredged areas (inside stations) Conflicts of interest and in neighbouring, not dredged, areas (outside stations). The results showed All authors declare no conflicts of interest. that the impact of sand extraction was confined to the dredged stations and to the areas in proximity to the dredged areas. During dredging activities, the doi:10.1111/j.1439-0485.2009.00321.x structure of benthic assemblages within the impacted stations was characterised by low species richness and diversity. Both the direct removal of sediment and the re-suspension and consequent deposition of fine sediment affected benthic assemblages of the impacted stations. A few months after the dredgings, a recolonisation process was still observed at all the impacted stations. A gradual recolonisation process was observed at those stations affected by only one dredging, whereas a different recolonisation was observed at those stations affected by two dredgings over time. This study suggests that differences of re-colonisation processes of benthic assemblages are related to the intensity of dredging operations in terms of dredging frequency. To combat coastal erosion along the Italian coasts, the the development of shallow furrows 1–3 m in width and local governments and the environmental protection sometimes up to 5 m in depth (Desprez 2000). Anchor agencies of several regions have planned nourishment dredging leads to the formation of deep, cup-shaped operations exploiting relict sand deposits, within the depressions, typically up to 8–10 m deep (Boyd & Rees framework of the European project INTERREG IIIC 2003). Both dredging methods can result in significant BEACHMED-e (http://www.beachmed.eu). environmental alterations, which may take place on both Relict sands are non-diagnosed sedimentary deposits physical and biological levels. The main physical effects situated along the continental shelf in a state of disequi- involve variations in morphological and bathymetric fea- librium with the present sedimentary dynamics. The tures, modifications of superficial sediment characteristics, removal of such sediments, occurring offshore at high and an increase in water turbidity caused by the re-sus- depths, does not affect the wave motion regime and, pension of fine sediment in the water column during therefore, coastal dynamics. The relict sand extraction is dredging activities. Concerning the biological effects, both performed through the use of suction trailers or anchor dredging methods cause severe disturbances in macrozoo- dredges. A common consequence of trailer dredging is benthos assemblages in terms of the direct effect on Marine Ecology 30 (Suppl. 1) (2009) 97–104 ª 2009 Blackwell Verlag GmbH 97 Response of macrobenthos to relict sand dredging La Porta, Targusi, Lattanzi, La Valle, Paganelli & Nicoletti sediment removal and the indirect effect associated with macrobenthos assemblages; (ii) the recolonisation pro- the deposition of suspended sediment caused by sand cesses of macrobenthos in the dredged areas; (iii) the extraction (Desprez 2000; Sarda` et al. 2000; Boyd & Rees effects over time of repeated dredging activities on macro- 2003; Szymelfenig et al. 2006; Simonini et al. 2007). Nev- benthos assemblages. ertheless, the type of dredge employed, as well as the nat- ure of the receiving environment, can potentially Material and Methods influence the spatial scale of impact on the benthic fauna, in terms of both direct and indirect effects caused by sand The study area was located 3.5 nautical miles offshore extraction (Boyd & Rees 2003). Boyd & Rees (2003), Ne- from Montalto di Castro (Lazio, Italy) in the central Tyr- well et al. (2004), Robinson et al. (2005) and, more rhenian Sea, on the continental shelf at 50 m of water recently, Cooper et al. (2007) have shown that the impact depth. on benthic assemblages is also related to the process of The relict sand-dredging activities in this area took repeated dredgings within the dredged site. Robinson place in three different periods, July 2004 (first dredging), et al. (2005) and Cooper et al. (2007) also highlighted June 2005 (second dredging), and September 2005 (third that benthic recolonisation processes in repeatedly dredging). Over this period, three changes in the bound- dredged areas are particularly difficult to predict, because aries of the extraction areas were reported (Fig. 1). For of both the different benthic responses to the intensity of the first dredging, an anchor dredge was used, whereas dredging operations in terms of dredging frequency and for the second and third dredging a trailer dredge was the variations in environmental characteristics. used. The monitoring surveys were carried out from May Between July 2004 and September 2005, three relict 2004 to October 2006, before, during and after the dredg- sand-dredging activities were performed in an area ing activities, as indicated in Nicoletti et al. (2006) offshore Montalto di Castro (Lazio, Italy) in the central (Table 1). The sampling plan provided five stations Tyrrhenian Sea, with the final aim of nourishing various (named stations 1, 2, 3, 4 and 5), one of which was beaches along the Lazio coasts. This area was character- located inside the dredged area in order to monitor the ised by the presence of relict sand deposits that were first dredging. The second and the third dredging activi- covered by a muddy layer of recent deposition, with a ties were carried out in proximity (N–NE) to the first thickness that varies between a few centimetres and a few area dredged. Three stations (6, 7 and 8) were added to metres (Chiocci & La Monica 1999). For these operations, the sampling plan to monitor these dredgings, as shown ISPRA, formerly ICRAM (Central Institute for Marine in Fig. 1. Macrobenthos sampling was carried out using a Research), carried out an environmental impact study Van Veen grab with a surface of 0.1 m 2. Two replicates related to marine relict sand extraction for beach nourish- were collected at each station. Samples were sieved ment, funded by the Regione Lazio local authority. This through a 1-mm mesh and the retained material was pre- monitoring program has provided an opportunity to col- served in 4% CaCO 3 buffered formalin in seawater. For lect useful information for the evaluation of the conse- each station, samples of superficial sediments were col- quences of sand extraction over a relatively short time lected through a box-corer to determine grain size distri- period in an offshore area that until now has been poorly bution. Superficial sediments were classified according to investigated. In particular, in this study we analysed: Shepard (1954). The collected organisms were counted (i) the effects of relict sand-dredging activities on the and classified to the lowest possible taxonomic level. In Fig. 1. On the left, the location of relict sand-extraction areas with a map of sampling stations (black point) is represented; on the right, side scan sonar reliefs of the dredged areas (fD = first dredged area; sD = second dredged area; tD = third dredged area) is reported. 98 Marine Ecology 30 (Suppl. 1) (2009) 97–104 ª 2009 Blackwell Verlag GmbH La Porta, Targusi, Lattanzi, La Valle, Paganelli & Nicoletti Response of macrobenthos to relict sand dredging Table 1. Sand-dredging characteristics and sampling plan of the three dredged areas. First dredged area Second dredged area Third dredged area Volume sand extracted (m 3) 600,000 150,000 700,000 Water depth (m) 50 50 50 Type of dredge Anchor dredge Trailer dredge Trailer dredge Dredging period July 2004 June 2005 September 2005 Sampling stations Inside the dredged area 5 6,7 3, 4, 6 Outside the dredged area 1, 2, 3, 4 1, 2, 3, 4, 5, 8 1, 2, 5, 7, 8 Surveys May 2004 – before dredging (B) Ö – – July 2004 – during first dredging (fD) Ö – – September 2004 – 2 months after dredging (A2) Ö – – April 2005 – 9 months after dredging (A9) Ö – – August 2005 – 14 months after dredging (A14) Ö Ö Ö September 2005 – during third dredging (tD) Ö Ö Ö May 2006 – 22 months after dredging (A22) Ö Ö Ö October 2006 – 27 months after dredging (A27) Ö Ö Ö particular, Polychaeta, Crustacea, Mollusca and Echino- The univariate analysis showed that the first dredging dermata were analysed. The main ecological indices caused a drastic reduction of the ecological indices exclu- (abundance, number of species, Margalef species richness sively at station 5 located within the dredged area. Sta- and Shannon–Wiener diversity) were calculated. Multi- tions 1, 2, 3 and 4, located outside the first dredged area, variate analysis was performed with abundance data to seemed not to have been affected by dredging. Fourteen analyse the benthic assemblage variation patterns in terms months after the end of the first dredging activity, of species composition and numerically dominant species. impacted station 5 showed an increase in the ecological The output from the non-metric multidimensional scaling parameters. During the second dredging, no surveys for (nMDS) ordination model of the Bray–Curtis similarity the macrobenthos monitorings were carried out. Never- matrix was obtained for each station and sampling per- theless, the monitoring survey carried out 2 months after iod. Univariate and multivariate analyses were performed the second extraction, showed that only station 6 was using the software package primer v. 6.1.5 (Clarke & characterised by extremely low indices values. During the Gorley 2001). third dredging, all stations except 1, 2 and 8, which were located outside the dredged area, showed a drastic decrease of the ecological indices (Fig. 2 and Table 3). Results In general, data relating to the two monitoring surveys During the study period, 4553 individuals belonging to carried out after the end of the third dredging highlighted 191 species were collected (Table 2). Polychaetes were the that all the impacted stations showed an increase in the most abundant taxon (3371 individuals and 103 species), ecological indices. Stations 5 and 6 were characterised by followed by crustaceans (626 individuals and 48 species), a particularly strong increase in these values (Fig. 2, echinoderms (328 individuals and 10 species), and mol- Table 3), mainly due to the high abundance of a few luscs (228 individuals and 30 species). The most abun- opportunistic species (e.g. C. gibba and T. stroemi ) and to dant species were the polychaetes Paralacydonia paradoxa , the presence and abundance of previously absent species Glycera unicornis , Paraprionospio pinnata , Metasychis gotoi , that colonised the impacted substrata (e.g. S. bairdi, the tanaid Tuberapseudes echinatus, and the ophiuroid N. hombergi, D. glaucus ). Amphiura chiajei . In general, benthic assemblages were The nMDS ordination plot of data relating to each sta- characteristic of muddy bottoms. The species composition tion and to each sampling period shows an overlapping did not show considerable variations over time. Only a of samples (Fig. 3). Station distribution confirms the few taxa showed variation over time; these were the homogeneity of the benthic assemblage observed over opportunistic species Corbula gibba and Terebellides stro- time. Station 5, which was affected during the first and emi, and the sabulicolous polychaetes Streblosoma bairdi , the third dredging, segregated on the left side of the plot. Nephtys hombergi and Diplocirrus glaucus . These latter Furthermore, on the right side we find stations 5 and 6 species were mainly found at the dredged stations. analysed during the last two monitoring surveys and Marine Ecology 30 (Suppl. 1) (2009) 97–104 ª 2009 Blackwell Verlag GmbH 99 Response of macrobenthos to relict sand dredging La Porta, Targusi, Lattanzi, La Valle, Paganelli & Nicoletti Table 2. Species collected during the study period. Table 2. (Continued .) Mollusca Othomaera schmidtii (Stephensen, 1915) Pseudotorinia architae (O.G. Costa, 1839) Westwoodilla rectirostris (Delia Valle, 1893) Calliostoma (Ampullotrochus) gramtlatum (Von Born, 1 778) Harpinia agna G Karaman, 1987 Turritella communis Risso, 1826 Harpinia ala G. Karaman, 1987 Hyala vitrea (Montagu, 1803) Harpinia antennaria Meinert, 1890 Calyptraea chinensis (Linnaeus, 1758) Harpinia karamani King, 2004 Polinices macilenta (Philippi, 1844) Harpinia sp. Polinices nitida (Donovan, 1804) Metaphoxus fultoni (Scott, 1890) Eulima glabra (Da Costa, 1778) Phtisica marina Slabber, 1769 Nassarius (Gussonea) cfr. comiculus (Olivi, 1792) Alpheus glaber (Olivi, 1792) Nucula nucleus (Linnaeus, 1758) Athanas nitescens (Leach, 1814) Nucula sulcata (Bronn, 1831) Processa canaliculata Leach, 1815 Saccella commutata (Philippi, 1844) Callianassa subterranea (Montagu, 1808) Thyasira biplicata (Philippi, 1836) Goutretia denticulata (Lutze, 1937) Glans aculeata (Poli, 1795) Jaxea nocturna Nardo, 1847 Astarte sulcata (Da Costa, 1778) Paguristes eremita (Linnaeus, 1767) Plagiocardium papillosum (Poli, 1795) Anapagurus laevis (Bell, 1845) Lutraria sp. Anapagurus serripes (Hope, 1851) Phaxas adriaticus (Coen, 1933) Pagurus cuanensis Bell, 1845 Tellina donacina Linnaeus 1758 Medorippe lanata (Linnaeus, 1767) Tellina serrata Brocchi, 1814 Ebalia deshayesi Lucas, 1845 Gari fervensis (Gmelin, 1791) Liocarcinus maculatus (Risso, 1827) Abra alba (Wood,1802) Goneplax rhomboides (Linnaeus, 1758) Abra prismatica (Montagu, 1808) Polychaeta Abra renierii (Bronn, 1831) Capitella capitata (Fabricius, 1870) Pitar rudis (Poli 1795) Heteromastus filiformis (Claparede, 1864) Timoclea ovata (Pennant, 1777) Leiocapitella glabra Hartman, 1947 Corbula gibba (Olivi,1792) Notomastus aberans Day, 1957 Antalis inaequicostata (Dautzenberg, 1891) Notomastus latericeus Sars, 1850 Crustacea Notomastus lineatus Claparede, 1870 Iphinoe rhodaniensis Ledoyer, 1965 Pseudoleiocapitella fauveli Harmelin, 1964 Iphinoe serrata Norman, 1867 Cossura soyeri Laubier, 1964 Apseudes acutifrons G O. Sars, 1882 Clymenura clypeata (Saint-Joseph, 1894) Apseudes elisae Bacescu, 1961 Praxillella affinis (M. Sars, 1872) Apseudes latreilli (Milne-Edwards, 1828) Praxillella gracilis (M. Sars, 1872) Tuberapseudes echinatus (GO. Sars, 1882) Maldane glebifex Grube, 1860 Leptochelia savignyi (Kroyer, 1842) Maldane sarsi Malmgren, 1865 Arcturella dilatata (GO. Sars, 1883) Nematonereis unicornis (Schmarda, 1861) Gnathia sp. Palola siciliensis (Grube, 1840) Anthura gracilis (Montagu, 1808) Metasychis gotoi (Izuka, 1902) Cirolana borealis Lilljeborg, 1852 Nicomache lumbricalis (Fabricius, 1780) Cirolana sp. Maldanidae gen.sp Ampelisca diadema (A Costa, 1853) Polyophthalmus pictus (Dujardin, 1839) Ampelisca spinifer Reid, 1951 Polyodontes maxillosus (Ranzani, 1817) Ampelisca spinipes Boeck, 1861 Harmothoe longisetis (Grube, 1863) Ampelisca typica (Bate, 1856) Lepidonotus clava (Montagu, 1808) Haploops dellavallei Chevreux, 1900 Lepidonotus squamatus (Linnaeus, 1767) Haploops nirae Kaim Malka, 1976 Malmgreniella lunulata (Delle Chiaje, 1830) Leptocheirus guttatus (Grube, 1864) Sthenelais boa (Johnston, 1833) Leptocheirus mariae G Karaman, 1973 Podarkeopsis arenicola (La Greca, 1947) Medicorophium rotundirostre (Stephensen, 1915) Pilargis verrucosa (Saint-Joseph, 1899) Photis longicaudata (Bate & Westwood, 1862) Sigambra tentaculata (Treadwell, 1941) Leucothoe incisa Robertson, 1892 Glycera alba (O.F. Muller, 1776) Leucothoe lilljeborgi Boeck, 1861 Glycera tesselata Grube, 1863 Leucothoe oboa G. Karaman, 1971 Glycera unicornis Savigny, 1818 Liljeborgia dellavallei Stebbing, 1906 Glycinde nordmanni (Malmgren, 1866) Hippomedon massiliensis Bellan-Santini, 1965 Goniada maculata Oersted, 1843 Maera grossimana (Montagu, 1808) Nephtys hombergi Savigny, 1818 100 Marine Ecology 30 (Suppl. 1) (2009) 97–104 ª 2009 Blackwell Verlag GmbH La Porta, Targusi, Lattanzi, La Valle, Paganelli & Nicoletti Response of macrobenthos to relict sand dredging Table 2. (Continued .) Table 2. (Continued .) Nepthys hystricis Mclntosh, 1900 Prionospio fallax Soderstrom, 1920 Paralacydonia paradoxa Fauvel, 1913 Prionospio steenstrupi Malmgren, 1867 Phyllodoce lineata (Claparede, 1870) Scolelepis bonnieri (Mesnil, 1896) Dorvillea (Schistomeringos) neglecta (Fauvel, 1923) Scolelepis foliosa (Audouin & Milne-Edwards, 1833) Dorvillea (Schistomeringos) rudolphii (Delle Chiaje, 1828) Spio decoratus Bobretzky, 1870 Aglaophamus rubellus (Michaelsen, 1897) Spio filicornis (O. F. Muller, 1776) Eunice pennata (O.F. Muller, 1776) Spio multioculata (Rioja, 1918) Eunice vittata (Delle Chiaje, 1828) Spiophanes bombyx (Claparede, 1870) Lysibranchia paucibranchiata Cantone, 1983 Spiophanes kroyeri Grube, 1860 Marphysa belli (Audouin & Milne-Edwards, 1833) Spiophanes kroyeri reyssi Laubier, 1961 Marphysa kinbergi Mclntosh, 1910 Poecilochaetus serpens Alien, 1904 Lumbrineriopsis paradoxa (Saint-Joseph, 1888) Echinodermata Lumbrineris gracilis (Fillers, 1868) Pseudotrachytyone sp. Lumbrineris latreilli Audouin & Milne Edwars, 1834 Trachythyone elongata (Duben Koren, 1844) Scoletoma emandibulata-mabiti (Ramos, 1976) Trachythyone tergestina (M. Sars, 1857) Scoletoma fragilis (O.F. Muller, 1776) Thyone fusus (O.F. Muller, 1776) Scotetoma tetrawa (Schmarda, 1861) Phyllophorus urna Grube, 1840 Arabella tricolor (Montagu, 1804) Labidoplax digitata (Montagu, 1815) Drilonereis filum (Claparede, 1870) Amphiura chiajei Forbes, 1843 Apomtphis bilineata (Baird, 1870) Amphiura filiformis (O.F. Muller, 1776) Apomtphys brementi (Fauvel, 1916) Ophiopsila aranea Forbes, 1843 Apomtphis fauveli (Rioja, 1918) Ophiura albida Forbes, 1839 Hyalinoecia tubicola (O.F. Muller, 1776) Schizaster canaliferus (Lamarck, 1816) Myriochele oculata Zachs, 1923 Owenia fusiformis Delle Chiaje, 1841 Aphelochaeta marioni (Saint-Joseph, 1894) characterised by high species richness and diversity. Con- Caulleriella mitltibranchiis (Grube, 1863) cerning the grain size distribution of the sediments, some Chaetozone caputesocis (Saint-Joseph, 1894) grain size variations were observed after the dredgings, Chaetozone setosa Malmgren, 1867 both inside and outside the dredged areas. In particular, a Monticellina dorsobranchialis (Kirkegaard, 1959) significant increase in the sandy fraction (from 28% to Brada villosa (Rathke, 1843) 94.3%) was observed after the first dredging in station 5 Diplocirrus glaucus (Malmgren, 1867) (inside the dredged area) and another (from 47% to Flabelligera affinis M. Sars, 1829 Sternaspis scutata (Ranzani, 1817) 88.7% of sand) was recorded after the third dredging in Amage adspersa (Grube, 1863) station 6 (inside the dredged area). No relevant grain size Amage gallasii Marion, 1875 variations were reported in the other stations. Ampharete acutifrons (Grube, 1860) Amphicteis gunneri (M. Sars, 1835) Anobothrus gracilis (Malmgren, 1866) Discussion Eclysippe vanelli (Fauvel, 1936) The results obtained from this study, as expected and in Lysippe labiata Malmgren, 1866 Sabellides octocirrata (M. Sars, 1835) accordance with some authors (Blake et al. 1996; Newell Melinna palmata Grube, 1870 et al. 1998; Sarda` et al. 2000; Van Dalfsen et al. 2000; Pectinaria auricoma (O. F. Muller, 1776) Boyd & Rees 2003; Simonini et al. 2005), highlighted that Pectinaria koreni (Malmgren, 1866) the direct effects of relict sand dredgings on macroben- Pista brevibranchia Caullery, 1915 thos assemblages were limited to the dredged areas. In Pista cnstata (O. F. Muller, 1776) particular, all the stations located inside the dredged areas Streblosoma bairdi (Malmgren, 1866) during the first (station 5) and the third dredging (sta- Terebellides stroemi M. Sars, 1835 Magellona spl tions 3, 4 and 6) showed a strong decrease in ecological Magelona sp2 indices as a consequence of the complete removal of Spiochaetopteus costarum (Claparede, 1868) superficial sediments. Despite the lack of data, both Aonides paucibranchiata Southern, 1914 before and during the second dredging, it is important to Laonice cirrata (M. Sars, 1851) highlight the case of station 6, where both the low values ⁄ Minuspio cirri era Wiren, 1883 of the ecological indices recorded a few months after the Paraprionospio pinnata (Fillers, 1901) second extraction and its position (inside the second Prionospio caspersi Laubier, 1962 Prionospio ehlersi Fauvel, 1928 dredged area) allowed us to hypothesise that this station was dredged during the second extraction. Marine Ecology 30 (Suppl. 1) (2009) 97–104 ª 2009 Blackwell Verlag GmbH 101 Response of macrobenthos to relict sand dredging La Porta, Targusi, Lattanzi, La Valle, Paganelli & Nicoletti Fig. 2. Number of individuals (black line) and species (grey line) collected at each station over time. Table 3. Species richness (d) and Shannon diversity (H’) values calculated for each station over time. Stations B fD A2 A9 A14 tD A22 A27 1 d 5.74 8.03 5.62 5.34 5.86 4.37 5.06 9.37 H’ 4.21 4.86 4.10 3.97 4.14 3.30 3.95 4.67 2 d 6.40 5.35 5.29 6.53 7.92 7.90 5.15 8.62 H’ 4.05 4.02 3.99 4.50 4.77 4.87 3.77 4.96 3 d 5.96 5.94 8.05 5.32 5.23 4.25 5.64 5.93 H’ 4.24 4.36 4.71 3.97 3.88 3.51 4.11 4.07 4 d 6.18 8.68 8.93 5.35 7.34 5.03 1.85 7.77 H’ 3.95 4.93 5.00 4.11 4.64 3.91 2.15 4.64 5 d 7.61 3.75 6.87 6.07 6.51 3.96 6.40 9.99 H’ 4.54 3.28 4.50 4.25 4.47 3.36 4.38 5.11 6 d – – – – 3.97 5.19 5.31 9.60 H’ – – – – 3.45 4.09 3.34 4.07 7 d – – – – 6.75 4.75 3.74 5.30 H’ – – – – 4.47 3.71 3.22 4.03 8 d – – – – 7.19 5.85 7.09 7.31 H’ – – – – 4.63 4.28 4.53 4.35 This study highlighted that the impacts of relict sand quent deposition of fine sediments caused by sand-extrac- dredgings on macrobenthos assemblages were observed tion operations and was mainly evident at stations 5 and in the zones in proximity to the dredged areas. These 7, which were located in proximity to the third dredged indirect impacts were due to the re-suspension and subse- area. The increase in the fine fraction of superficial 102 Marine Ecology 30 (Suppl. 1) (2009) 97–104 ª 2009 Blackwell Verlag GmbH La Porta, Targusi, Lattanzi, La Valle, Paganelli & Nicoletti Response of macrobenthos to relict sand dredging Fig. 3. 2D-nMDS ordination plot of abundance data of each station and each sampling period. sediments observed in station 5 after dredging confirmed tions in the first phase of the recolonisation process, fol- that fine sediment re-deposition had occurred. lowed by an increase in the number of species and These results also highlighted that a stronger sediment individuals (Bonsdorff 1980; Kenny & Rees 1994, 1996; suspension was generated by the trailer dredge (used for Newell et al. 1998; Sarda` et al. 2000; Van Dalfsen et al. the second and the third dredging), whose impact was 2000; Nicoletti et al. 2004). The differences between two greater than that of the anchor dredge (used for the first recolonisation processes at the impacted stations were dredging). probably related to the fact that the first group of stations Concerning the analysis of the recolonisation processes (3, 4 and 7) was influenced exclusively by only one of macrobenthos assemblages, our results showed that a dredging (the third one), whereas the second group few months after the end of dredgings, the recolonisation (5 and 6) was affected by two dredgings (respectively the processes could still be observed at all the impacted sta- first and the third one for station 5 and the second and tions, in accordance with Green (2002), Boyd & Rees the third one for station 6). Moreover, these differences (2003), Simonini et al. (2005). In general, these processes could be related to the intensity of dredging operations in are mainly due to the settlement of new recruits from the terms of dredging frequency, as also observed by Boyd & planktonic larvae and immigration of the adults from the Rees (2003), Newell et al. (2004), Robinson et al. (2005) neighbouring areas (Bonvicini Pagliai et al. 1985; Rees & and Cooper et al. (2007). Dare 1993; Newell et al. 1998; Van Dalfsen et al. 2000), This study has confirmed the observations of some but recolonisation processes are difficult to predict authors (Robinson et al. 2005; Smith et al. 2006) con- because they are strongly influenced by many different cerning the difficulties in evaluating the effects over time factors ( e.g. biological cycles of different species, hydrody- of relict sand dredgings on benthic assemblages, due to namic regime, changes in sediment structure depth). the high number of factors involved. In our specific case, This study also revealed differences in the recolonisa- the analysis of the impact on the assemblages was further tion processes of the impacted stations. The gradual complicated by the use of two different types of dredge, recolonisation process was observed at stations 3, 4 and and by the fact that dredging activities were repeated 7, whereas different processes (with an exponential trend) within a relatively short period of time, as well as in areas were observed at stations 5 and 6. These stations were ini- that are very close to one another. Further, medium-term tially characterised by the abundance of a few opportunis- monitoring surveys will provide a more detailed descrip- tic species ( e.g. Corbula gibba and Terebellides stroemi ) tion of how the recolonisation process of macrobenthos and, subsequently (in the last monitoring), by an increase assemblages affected by sand dredging will occur, as well in abundance and in the number of sabulicolous species as how long this will take. (e.g. Streblosoma bairdi, Nephtys hombergi and Diplocirrus glaucus ) which had not been collected in the previous References investigated periods. This phenomenon is normally observed in dredged substrata where the defaunation Blake N.J., Doyle L.J., Culter J.J. (1996). Impacts and direct allows the opportunistic species to form dense popula- effects of sand dredging for beach renourishment on the Marine Ecology 30 (Suppl. 1) (2009) 97–104 ª 2009 Blackwell Verlag GmbH 103 Response of macrobenthos to relict sand dredging La Porta, Targusi, Lattanzi, La Valle, Paganelli & Nicoletti benthic organisms and geology of the west Florida shelf. una off the South coast of the United Kingdom. Journal of Contract 14-35- 0001-30644. US Dept. of Interior, MMS, Coastal Research , 20 (1), 115–125. Office of International Activities and Marine Minerals: 109 Nicoletti L., La Valle P., Chimenz Gusso C. (2004) Specie indi- pp. catrici: il caso Corbula gibba (Olivi, 1792). Biologia Marina Bonsdorff E. (1980) Macrozoobenthic recolonization of a Mediterranea , 11 (2), 273–277. dredged brackish water bay in SW Finland. Ophelia , Suppl. Nicoletti L., Paganelli D., Gabellini M. (2006) Environmental 1, 145–155. aspects of relict sand dredging for beach nourishment: pro- Bonvicini Pagliai A.M., Cognetti Varriale A.M., Crema R., Cu- posal for a monitoring protocol. Quaderno ICRAM n. 5: rini Galletti M., Vandini Zunarelli R. (1985) Environmental 155 pp. impact of extensive dredging in a coastal marine area. Mar- Rees H.L., Dare P.J. (1993) - Sources of mortality and associ- ine Pollution Bulletin , 16 , 483–488. ated life-cycle traits of selected benthic species: a review. Boyd S.E., Rees H.L. (2003) An examination of the spatial Ministry of Agriculture, Fisheries and Food Directorate of scale of impact on the marine benthos arising from marine Fisheries Research. Fisheries Research Data Report, no. 83, aggregate extraction in the central English Channel. Estua- Lowestoft, 1993: 36 pp. rine, Coastal and Shelf Science , 57 , 1–16. Robinson J.E., Newell R.C., Seiderer L.J., Simpson N.M. (2005) Chiocci F.L., La Monica G.B. (1999). Individuazione e carat- Impacts of aggregate dredging on sediment composition and terizzazione dei depositi sabbiosi presenti sulla piattaforma associated benthic fauna at an offshore dredge site in the continentale della Regione Lazio e valutazione di un loro southern North Sea. Marine Environmental Research , 60 , 51– utilizzo ai fini del ripascimento dei litorali in erosione. Rap- 68. porto della prima fase. Universita` degli Studi di Roma ‘La Sarda` R., Pinedo S., Gremare A., Taboada S. (2000) Changes Sapienza’, Dipartimento di Scienze della Terra – Regione in the dynamic of shallow-bottom assemblages due to sand Lazio, Dipartimento OO.PP. e Servizi per il Territorio – extraction in the Catalan Western Mediterranean Sea. ICES ROMA, settembre 1999: 100 pp. Journal of Marine Science , 57 , 1446–1453. Clarke K.R., Gorley R.N. (2001) PRIMER v5: User man- Shepard F.P. (1954) Nomenclature based on sand, silt, clay ual ⁄ tutorial, PRIMER-E, Plymouth UK, 91pp. ratios. Journal Sedimentary Petrology , 24 , 151–158. Cooper K., Boyd S., Aldridge J., Rees H. (2007) Cumulative Simonini R., Ansaloni I., Bonvicini Pagliai A.M., Cavallini F., impacts of aggregate extraction on seabed macro-inverte- Iotti M., Mauri M., Montanari G., Preti M., Rinaldi A., brate communities in an area off the east coast of the Uni- Prevedelli D. (2005) The effects of sand extraction on the ted Kingdom. Journal of Sea Research , 57 , 288–302. macrobenthos of a relict sands area (northern Adriatic Sea): Desprez M. (2000) Physical and biological impact of marine results 12 months post-extraction. Marine Pollution Bulletin , aggregate extraction along the French coast of the Eastern 50 , 768–777. English Channel: short- and long-term post-dredging resto- Simonini R., Ansaloni I., Bonini P., Grandi V., Graziosi F., ration. ICES Journal of Marine Science , 57 , 1428–1438. Iotti M., Massamba-N’Siala G., Mauri M., Montanari G., Green K. (2002). Beach nourishment: a review of the biological Preti M., De Nigris N., Prevedelli D. (2007) Recolonization and physical impacts. ASMFC (Atlantic States Marine Fish- and recovery dynamics of the macrozoobenthos after sand eries Commission), Habitat Management Series, 7: 174 pp. extraction in relict sand bottoms of the Northern Adriatic Kenny A.J., Rees H.L. (1994) The effects of marine gravel Sea. Marine Environmental Research , 64 , 574–589. extraction on the macrobenthos: early post-dredging recol- Smith R., Boyd S.E., Rees H.L., Dearnaley M.P., Stevenson J.R. onization. Marine Pollution Bulletin , 28 , 442–447. (2006) Effects of dredging activity on epifaunal communities Kenny A.J., Rees H.L. (1996) The effects of marine gravel and surveys following cessation of dredging. Estuarine, extraction on the macrobenthos: results of 2 years post- Coastal and Shelf Science , 70 , 207–223. dredging. Marine Pollution Bulletin , 32 , 615–622. Szymelfenig M., Kotwichi L., Graca B. (2006) Benthic re-colo- Newell R.C., Seiderer L.J., Hitchcock D.R. (1998) The impact nization in post-dredging pits in the Puck Bay (Southern of dredging works in coastal waters: a review of the sensitiv- Baltic Sea). Estuarine, Coastal and Shelf Science , 68 , 489–498. ity to disturbance and subsequent recovery of biological Van Dalfsen J.A., Essink K., Toxvig Madsen H., Birklund J., resources on the sea bed. Oceanography and Marine Biology: Romero J., Manzanera M. (2000) Differential response of an Annual Review , 36 , 127–178. macrozoobenthos to marine sand extraction in the North Newell R.C., Seiderer L.J., Simpson N.M., Robinson J.E. (2004) Sea and the Western Mediterranean. ICES Journal of Marine Impacts of marine aggregate dredging on benthic macrofa- Science , 57 , 1439–1445. 104 Marine Ecology 30 (Suppl. 1) (2009) 97–104 ª 2009 Blackwell Verlag GmbH Q`0V`$: :?_5`Q`.11J0:C%:GCV%]]Q` QJR: ::J:C71:JR`Q` .V%$$V 1QJ .: 1I]`Q0VR .11Q`@HQJ1RV`:GC78]VH1:C .:J@$Q Q`Q`841:J2:`1:&1I1:^ 33R ]:CC:J<:J16Q]1 :CQ` QIV_`Q`.10:C%:GCV%$$V 1QJR%`1J$:CC .V %R7]V`1QR8 27 $`V: V .:J@ $Q Q 1Q ]VH1:C ]VQ]CV7 :0V`1Q V0Q 1 1.Q `% VR :JR %]]Q` VRIV1JHV .VGV$1JJ1J$Q` .V.:JR Q2QJ1H: :`$%15 .VGV ``1VJR :JRHQCCV:$%V`Q`.:`1J$%H.:]`Q`V1QJ:C:JRC1`VV6]V`1VJHV8 ((#& 1JHV `VV:`H. QJ Q` RV:GVR I:H`Q