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<5472616E7363726962656420> Tj 1 0 0 1 361.44 570.00 Tm <53706163657220> Tj 1 0 0 1 387.60 570.24 Tm <284954532D4929> Tj 0.72 Tc 1 0 0 1 275.76 559.68 Tm <44697363757373696F6E> Tj 0.48 Tc 1 0 0 1 275.76 549.12 Tm <41636B6E6F776C656467656D656E7473> Tj 0.48 Tc 1 0 0 1 275.76 538.56 Tm <5265666572656E636573> Tj 0.48 Tc /F3 10.0 Tf 1 0 0 1 275.76 501.60 Tm <496E74726F64756374696F6E> Tj 0.24 Tc /F3 8.0 Tf 1 0 0 1 326.16 802.56 Tm <436F6E747269627574696F6E7320> Tj 1 0 0 1 376.08 802.32 Tm <746F20> Tj 1 0 0 1 384.72 802.56 Tm <5A6F6F6C6F67792C20> Tj 1 0 0 1 417.84 802.32 Tm <363920> Tj 1 0 0 1 428.88 802.56 Tm <286C2F322920> Tj 1 0 0 1 448.32 802.32 Tm <38392D393820> Tj 1 0 0 1 470.16 803.04 Tm <283230303029> Tj 0.48 Tc 1 0 0 1 350.64 792.24 Tm <53504220> Tj 1 0 0 1 366.00 792.24 Tm <41636164656D696320> Tj 1 0 0 1 402.48 792.24 Tm <5075626C697368696E6720> Tj 1 0 0 1 442.08 792.00 Tm <62762C20> Tj 1 0 0 1 455.04 792.00 Tm <54686520> Tj 1 0 0 1 469.44 792.00 Tm <4861677565> Tj 0.48 Tc /F3 9.0 Tf 1 0 0 1 130.32 632.64 Tm <4C6570656F70687468656972757320> Tj 1 0 0 1 199.92 632.40 Tm <736F6C6D6F6E69732C20> Tj 1 0 0 1 243.84 632.40 Tm <636F7065706F642C20> Tj 1 0 0 1 285.60 632.16 Tm <4372757374616365612C20> Tj 1 0 0 1 333.36 632.16 Tm <41746C616E74696320> Tj 1 0 0 1 371.04 631.92 Tm <73616C6D6F6E2C20> Tj 1 0 0 1 407.52 631.92 Tm <73657175656E63696E672C20> Tj 1 0 0 1 462.00 631.92 Tm <313853> Tj 0.48 Tc 1 0 0 1 129.84 619.92 Tm <444E412C20> Tj 1 0 0 1 158.64 620.16 Tm <696E7465726E616C20> Tj 1 0 0 1 194.16 619.92 Tm <7472616E7363726962656420> Tj 1 0 0 1 244.80 619.68 Tm <7370616365722C20> Tj 1 0 0 1 277.44 620.16 Tm <706F70756C6174696F6E20> Tj 1 0 0 1 326.16 619.92 Tm <67656E6574696373> Tj 0.72 Tc 1 0 0 1 483.36 590.64 Tm <3932> Tj 0.72 Tc 1 0 0 1 483.36 580.32 Tm <3932> Tj 0.96 Tc 1 0 0 1 483.60 569.76 Tm <3932> Tj 0.96 Tc 1 0 0 1 483.36 559.44 Tm <3933> Tj 0.96 Tc 1 0 0 1 483.36 548.64 Tm <3937> Tj 0.96 Tc 1 0 0 1 483.36 538.32 Tm <3937> Tj 0.96 Tc /F3 8.0 Tf 1 0 0 1 241.68 264.72 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Bx`H'K.WPŴ J,T lDIU ?q?Ie>stream Utility of 18S rDNA and ITS sequences as population markers for Lepeophtheirus sulmonis (Copepoda: Caligidae) parasitising Atlantic salmon (Sulmo sular) in ScotlandrA.P. Shinn, rB.A. Banks,2N. Tange, 1J.E. Bron, rC. Sommerville, 2T. Aoki & rR.Woottenllnstitute of Aquaculture, 鶹ýAV, 鶹ýAV FKg 4LA, Scotland; 2Laboratory of Genetics andBiochemistry, Department of Aquatic Bioscience, Tolqto University of Fisheries, Konan 4-5-7, Minqto-ku,Tolq,o l0B, Japan Keywords: Sea louse,ribosomalAbstract Genetic differentiation within the salmon louse Zepeophtheirussalmonis (Kroyer, I 837), was investigated by the sequencingofspecific nucleotide regions. Partial sequences ofthe l8Sribosomal RNA gene and the ribosomal internal transcribedspacer (ITS-1) region from single sea lice were amplified bythe polymerase chain reaction (PCR). Lice were collected fromwild and farmed Atlantic salmon (Salmo salar L., 1758) fromnine selected localities around the Scottish coastline. A 0.gkbfragment of the 18S ribosomal RNA gene was amplified andcompared for several samples of lice which showed noobservable differences between the lice from different collectionsites confirming the absence of cryptic species. The 454nucleotide ITS- 1 sequence showed differences between derivedsequences from 13 sea lice samples from 4 collection siteswhich included 2 farm sites and 2 sites where lice were takenfrom wild fish. Across all samples, there wasa92.14o/o similarityin the ITS-1 sequence. The percentage similarity in the ITS-1sequence in samples of lice from two fish farms were 99.'71%(site A) and 95.'72o/o (site D) but only 86.90% (site B) and86.03% (site C) similarity was shown in lice samples takenfrom sites where wild salmonids were caught. The greatersimilarity between the ITS-1 sequence within farm sites maybe attributed to a restricted gene flow within lice populationsin Atlantic salmon cage sites.Contents IntroductionMaterial and methodsSample collection and preparationGenomic DNA extractionPCR amplification18S ribosomal DNAInternal Transcribed Spacer (ITS)Cloning into pMOS Blue T-vectorDideoxynucleotide sequencing and analysis Resultsl8S ribosomal DNAInternal Transcribed Spacer (ITS-I)DiscussionAcknowledgementsReferencesIntroduction Contributions to Zoology, 69 (l/2) 89-98 (2000)SPB Academic Publishing bv, The Hague Lepeophtheirus solmonis, copepod, Crustacea, Atlantic salmon, sequencing, 18SDNA, internal transcribed spacer, population genetics 929292939797 8990909l91919l9192 The characterization of salmon louse (Lepeo-phtheirus salmonis (Kroyer, 1837)) populations hasrecently become the subject of considerable inter-est as a tool to investigate louse population ori-gins and interactions. Nordhagen, Heuch & Schram(1998) investigated experimentally the usefulnessof morphometric parameters to identify the ori-gins of the salmon lice populations in Norway. Theyconcluded that it was not possible to discriminatepopulations of lice on the basis of size, as lice ofdifferent origins raised at the same temperature ona common host showed no significant differencesin dimensions and similar life-cycle features suchas egg production and growth rate. Similarly, inan unpublished experiment, the present authorsapplying multivariate analyses to morphometricdata ofseveral populations oflice taken from natu-rally infected wild and farmed salmonids sampledthroughout a year, concluded that the discrimina-tion of lice on the basis of size was not sufficientlyclear to be useful.Other studies have investigated the allelic fre-quencies of polymorphic enzymes (allozymes) ofpopulations ofthe genus Lepeophtheiras. For ex-ample, Shinn et al. 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Tm <77697468696E20> Tj 1 0 0 1 244.08 333.84 Tm <6365727461696E> Tj 0.48 Tc 1 0 0 1 57.36 322.32 Tm <696E646976696475616C20> Tj 1 0 0 1 103.20 322.08 Tm <77696C6420> Tj 1 0 0 1 124.56 322.08 Tm <6C69636520> Tj 1 0 0 1 142.56 321.84 Tm <73756767657374656420> Tj 1 0 0 1 186.48 321.60 Tm <7468657920> Tj 1 0 0 1 207.60 321.36 Tm <6F726967696E6174656420> Tj 1 0 0 1 253.44 321.36 Tm <66726F6D> Tj 0.48 Tc 1 0 0 1 57.36 309.84 Tm <6661726D20> Tj 1 0 0 1 81.60 309.60 Tm <73697465732E20> Tj 1 0 0 1 107.28 309.60 Tm <5768696C6520> Tj 1 0 0 1 137.28 309.60 Tm <524150447320> Tj 1 0 0 1 174.00 309.12 Tm <616C6C6F7720> Tj 1 0 0 1 201.84 309.12 Tm <666F7220> Tj 1 0 0 1 217.68 309.12 Tm <74686520> Tj 1 0 0 1 234.96 308.88 Tm <636F6D706172692D> Tj 0.72 Tc 1 0 0 1 57.60 297.36 Tm <736F6E20> Tj 1 0 0 1 75.36 297.12 Tm <6F6620> Tj 1 0 0 1 87.12 297.12 Tm <616D706C696669656420> Tj 1 0 0 1 131.52 297.12 Tm <444E4120> Tj 1 0 0 1 157.44 296.88 Tm <62616E647320> Tj 1 0 0 1 185.76 296.64 Tm <6F6620> Tj 1 0 0 1 197.76 296.64 Tm <73696D696C617220> Tj 1 0 0 1 230.40 296.64 Tm <6D6F6C6563756C6172> Tj 0.72 Tc 1 0 0 1 57.12 284.88 Tm <7765696768742C20> Tj 1 0 0 1 95.04 284.64 Tm <776974686F757420> Tj 1 0 0 1 133.92 284.40 Tm <64697265637420> Tj 1 0 0 1 164.40 284.40 Tm <73616D706C696E6720> Tj 1 0 0 1 209.76 284.16 Tm <697420> Tj 1 0 0 1 220.80 284.16 Tm <697320> Tj 1 0 0 1 233.04 283.92 Tm <756E6B6E6F776E> Tj 0.48 Tc 1 0 0 1 57.12 272.40 Tm <7768657468657220> Tj 1 0 0 1 96.24 272.40 Tm <746865736520> Tj 1 0 0 1 123.12 272.16 Tm <73696D696C617220> Tj 1 0 0 1 157.44 271.92 Tm <73697A656420> Tj 1 0 0 1 183.36 271.68 Tm <62616E647320> Tj 1 0 0 1 213.12 271.68 Tm <636F72726573706F6E6420> Tj 1 0 0 1 265.44 271.44 Tm <746F> Tj 0.48 Tc 1 0 0 1 57.12 259.92 Tm <74686520> Tj 1 0 0 1 74.88 259.92 Tm <73616D6520> Tj 1 0 0 1 100.80 260.16 Tm <706F7274696F6E20> Tj 1 0 0 1 136.80 259.44 Tm <6F6620> Tj 1 0 0 1 150.00 259.68 Tm <74686520> Tj 1 0 0 1 167.52 259.92 Tm <67656E6F6D652E20> Tj 1 0 0 1 208.80 259.20 Tm <5468697320> Tj 1 0 0 1 232.56 258.96 Tm <737475647920> Tj 1 0 0 1 260.16 258.96 Tm <64652D> Tj 0.48 Tc 1 0 0 1 57.36 247.44 Tm <7363726962657320> Tj 1 0 0 1 89.76 247.20 Tm <616E20> Tj 1 0 0 1 102.48 247.44 Tm <696E7665737469676174696F6E20> Tj 1 0 0 1 160.80 246.96 Tm <776869636820> Tj 1 0 0 1 190.80 246.72 Tm <636861726163746572697A657320> Tj 1 0 0 1 248.40 246.72 Tm <706F70752D> Tj 0.72 Tc 1 0 0 1 57.12 234.96 Tm <6C6174696F6E7320> Tj 1 0 0 1 91.20 234.96 Tm <7468726F75676820> Tj 1 0 0 1 130.08 234.72 Tm <64657465726D696E6174696F6E20> Tj 1 0 0 1 196.08 234.24 Tm <6F6620> Tj 1 0 0 1 209.52 234.24 Tm <74686520> Tj 1 0 0 1 227.52 234.24 Tm <6E75636C656F74696465> Tj 0.48 Tc 1 0 0 1 57.36 222.48 Tm <73657175656E636520> Tj 1 0 0 1 100.08 222.48 Tm <6F6620> Tj 1 0 0 1 112.80 222.24 Tm <73656C656374656420> Tj 1 0 0 1 150.96 222.24 Tm <617265617320> Tj 1 0 0 1 176.64 222.00 Tm <6F6620> Tj 1 0 0 1 189.60 222.00 Tm <74686520> Tj 1 0 0 1 205.20 222.00 Tm <4C2E20> Tj 1 0 0 1 217.68 221.76 Tm <73616C6D6F6E697320> Tj 1 0 0 1 261.36 222.00 Tm <67652D> Tj 0.72 Tc 1 0 0 1 56.88 210.24 Tm <6E6F6D652C20> Tj 1 0 0 1 87.36 210.24 Tm <6E616D656C7920> Tj 1 0 0 1 122.88 210.00 Tm <74686520> Tj 1 0 0 1 141.12 210.00 Tm <6C385320> Tj 1 0 0 1 160.56 210.00 Tm <7269626F736F6D616C20> Tj 1 0 0 1 207.60 209.52 Tm <524E4120> Tj 1 0 0 1 234.00 209.76 Tm <67656E6520> Tj 1 0 0 1 258.24 209.28 Tm <616E64> Tj 0.48 Tc 1 0 0 1 56.88 197.76 Tm <74686520> Tj 1 0 0 1 73.92 197.76 Tm <696E7465726E616C20> Tj 1 0 0 1 110.40 197.76 Tm <7472616E7363726962656420> Tj 1 0 0 1 162.72 197.28 Tm <73706163657220> Tj 1 0 0 1 193.92 197.52 Tm <28495453292E> Tj 0.48 Tc 1 0 0 1 67.68 185.52 Tm <496E666F726D6174696F6E20> Tj 1 0 0 1 122.88 185.28 Tm <726573756C74696E7320> Tj 1 0 0 1 164.64 185.04 Tm <66726F6D20> Tj 1 0 0 1 189.12 184.80 Tm <616E616C7673697320> Tj 1 0 0 1 227.52 184.56 Tm <6F6620> Tj 1 0 0 1 240.00 184.56 Tm <74686520> Tj 1 0 0 1 258.00 184.56 Tm <6C3835> Tj 0.48 Tc 1 0 0 1 300.96 770.40 Tm <72524E4120> Tj 1 0 0 1 330.24 770.64 Tm <67656E6520> Tj 1 0 0 1 354.00 769.92 Tm <73657175656E636520> Tj 1 0 0 1 396.48 769.92 Tm <68617320> Tj 1 0 0 1 413.76 770.16 Tm <70726576696F75736C7920> Tj 1 0 0 1 462.72 769.44 Tm <6265656E20> Tj 1 0 0 1 486.48 769.20 Tm <7573656420> Tj 1 0 0 1 509.04 769.20 Tm <746F> Tj 0.48 Tc 1 0 0 1 300.96 757.92 Tm <64657465726D696E6520> Tj 1 0 0 1 346.80 757.68 Tm <7768657468657220> Tj 1 0 0 1 384.96 757.44 Tm <6F7220> Tj 1 0 0 1 396.72 757.44 Tm <6E6F7420> Tj 1 0 0 1 413.28 757.44 Tm <74686520> Tj 1 0 0 1 429.84 757.20 Tm <7370656369657320> Tj 1 0 0 1 463.68 756.96 Tm <756E64657220> Tj 1 0 0 1 491.52 756.96 Tm <696E7665732D> Tj 0.72 Tc 1 0 0 1 300.72 745.68 Tm <7469676174696F6E20> Tj 1 0 0 1 336.24 745.20 Tm <636F6E7461696E7320> Tj 1 0 0 1 374.64 744.96 Tm <6372797074696320> Tj 1 0 0 1 407.28 744.96 Tm <7370656369657320> Tj 1 0 0 1 440.64 744.72 Tm <6F7220> Tj 1 0 0 1 451.92 744.96 Tm <67656E65746963616C6C7920> Tj 1 0 0 1 501.84 744.48 Tm <69736F2D> Tj 0.48 Tc 1 0 0 1 300.96 733.20 Tm <6C6174656420> Tj 1 0 0 1 324.96 733.44 Tm <706F70756C6174696F6E732E20> Tj 1 0 0 1 382.56 732.48 Tm <53696D696C61726C792C20> Tj 1 0 0 1 428.64 732.24 Tm <64657465726D696E6174696F6E20> Tj 1 0 0 1 492.24 732.00 Tm <6F6620> Tj 1 0 0 1 504.48 732.00 Tm <746865> Tj 0.72 Tc 1 0 0 1 300.72 720.72 Tm <6E75636C656F7469646520> Tj 1 0 0 1 349.92 720.24 Tm <73657175656E636520> Tj 1 0 0 1 393.12 720.00 Tm <6F6620> Tj 1 0 0 1 405.84 720.00 Tm <696E7465726E616C20> Tj 1 0 0 1 442.80 719.76 Tm <7472616E7363726962656420> Tj 1 0 0 1 495.12 719.28 Tm <737061632D> Tj 0.72 Tc 1 0 0 1 300.72 708.00 Tm <65727320> Tj 1 0 0 1 317.04 708.24 Tm <284954532920> Tj 1 0 0 1 343.92 707.76 Tm <6861766520> Tj 1 0 0 1 367.68 707.52 Tm <6265656E20> Tj 1 0 0 1 392.16 707.52 Tm <7375636365737366756C6C7920> Tj 1 0 0 1 448.80 707.28 Tm <696D706C656D656E74656420> Tj 1 0 0 1 508.56 706.80 Tm <6173> Tj 0.48 Tc 1 0 0 1 300.72 695.52 Tm <6120> Tj 1 0 0 1 307.92 695.52 Tm <7461786F6E6F6D696320> Tj 1 0 0 1 356.16 695.28 Tm <746F6F6C20> Tj 1 0 0 1 375.84 695.04 Tm <746F20> Tj 1 0 0 1 386.88 695.04 Tm <6469736372696D696E61746520> Tj 1 0 0 1 443.28 694.56 Tm <636C6F73656C7920> Tj 1 0 0 1 476.40 694.56 Tm <72656C6174656420> Tj 1 0 0 1 508.80 694.32 Tm <6F72> Tj 0.72 Tc 1 0 0 1 300.48 683.04 Tm <6D6F7270686F6C6F676963616C6C7920> Tj 1 0 0 1 374.64 682.56 Tm <696E64697374696E677569736861626C6520> Tj 1 0 0 1 450.96 682.08 Tm <7370656369657320> Tj 1 0 0 1 484.32 681.84 Tm <6F6620> Tj 1 0 0 1 495.36 682.32 Tm <706172612D> Tj 0.72 Tc 1 0 0 1 300.96 670.32 Tm <7369746520> Tj 1 0 0 1 322.08 670.32 Tm <652E6720> Tj 1 0 0 1 341.04 670.08 Tm <617069636F6D706C6578616E7320> Tj 1 0 0 1 413.52 670.08 Tm <28486F6D616E20> Tj 1 0 0 1 455.28 669.60 Tm <657420> Tj 1 0 0 1 469.20 669.36 Tm <616C2E2C20> Tj 1 0 0 1 490.32 669.60 Tm <31393937293B> Tj 0.48 Tc 1 0 0 1 300.48 658.32 Tm <6779726F64616374796C69647320> Tj 1 0 0 1 361.68 657.84 Tm <284D6F6E6F67656E65612920> Tj 1 0 0 1 421.92 657.60 Tm <2843756E6E696E6768616D2C20> Tj 1 0 0 1 486.00 656.88 Tm <4D6347696C2D> Tj 0.72 Tc 1 0 0 1 300.72 645.60 Tm <6C697672617920> Tj 1 0 0 1 329.76 645.12 Tm <2620> Tj 1 0 0 1 341.76 645.36 Tm <4D61634B656E7A69652C20> Tj 1 0 0 1 399.36 644.88 Tm <313939353B20> Tj 1 0 0 1 426.96 644.64 Tm <43756E6E696E6768616D2C20> Tj 1 0 0 1 490.56 644.40 Tm <31393937293B> Tj 0.72 Tc 1 0 0 1 300.72 632.88 Tm <7363686973746F736F6D657320> Tj 1 0 0 1 361.68 632.88 Tm <28446967656E65612920> Tj 1 0 0 1 408.96 632.64 Tm <284C6974746C65776F6F6420> Tj 1 0 0 1 464.16 631.92 Tm <2620> Tj 1 0 0 1 476.16 631.68 Tm <4A6F686E73746F6E2C> Tj 0.72 Tc 1 0 0 1 301.68 620.40 Tm <313939353B20> Tj 1 0 0 1 328.32 620.16 Tm <56616E20> Tj 1 0 0 1 349.20 620.16 Tm <48657277657264656E2C20> Tj 1 0 0 1 403.20 619.68 Tm <426C61697220> Tj 1 0 0 1 428.16 619.44 Tm <2620> Tj 1 0 0 1 439.20 619.68 Tm <4167617473756D612C20> Tj 1 0 0 1 490.08 619.44 Tm <31393938293B> Tj 0.72 Tc 1 0 0 1 300.48 607.68 Tm <616E6420> Tj 1 0 0 1 321.60 607.92 Tm <6E656D61746F64657320> Tj 1 0 0 1 374.40 607.68 Tm <284368696C746F6E2C20> Tj 1 0 0 1 419.76 607.20 Tm <47617373657220> Tj 1 0 0 1 455.28 606.96 Tm <2620> Tj 1 0 0 1 469.20 606.96 Tm <4265766572696467652C> Tj 0.72 Tc 1 0 0 1 301.44 595.44 Tm <31393935292E20> Tj 1 0 0 1 331.68 595.20 Tm <4E75636C656F7469646520> Tj 1 0 0 1 383.04 594.96 Tm <766172696174696F6E7320> Tj 1 0 0 1 429.84 594.72 Tm <696E20> Tj 1 0 0 1 441.84 594.72 Tm <49545320> Tj 1 0 0 1 461.28 594.48 Tm <726567696F6E7320> Tj 1 0 0 1 496.80 594.48 Tm <68617665> Tj 0.72 Tc 1 0 0 1 300.48 582.96 Tm <616C736F20> Tj 1 0 0 1 321.60 582.72 Tm <6265656E20> Tj 1 0 0 1 346.08 582.72 Tm <7573656420> Tj 1 0 0 1 370.08 582.72 Tm <746F20> Tj 1 0 0 1 383.04 582.48 Tm <63686172616374657269736520> Tj 1 0 0 1 439.20 582.24 Tm <73747261696E7320> Tj 1 0 0 1 471.60 582.00 Tm <6F7220> Tj 1 0 0 1 484.80 582.00 Tm <69736F6C61746573> Tj 0.48 Tc 1 0 0 1 300.48 570.48 Tm <6F6620> Tj 1 0 0 1 312.00 570.48 Tm <437970746F636172796F6E20> Tj 1 0 0 1 369.36 570.24 Tm <6972726974616E7320> Tj 1 0 0 1 404.16 570.48 Tm <28446967676C657320> Tj 1 0 0 1 443.52 569.76 Tm <2620> Tj 1 0 0 1 453.84 569.76 Tm <41646C6172642C20> Tj 1 0 0 1 490.08 569.52 Tm <3139392737293B> Tj 0.48 Tc 1 0 0 1 299.76 558.00 Tm <456E7465726F6379746F7A6F6F6E20> Tj 1 0 0 1 376.08 557.52 Tm <6269656E6575736920> Tj 1 0 0 1 419.52 558.00 Tm <2852696E6465722C20> Tj 1 0 0 1 462.48 557.04 Tm <4B61747A77696E6B656C2D> Tj 0.48 Tc 1 0 0 1 300.24 545.28 Tm <576C6164617273636820> Tj 1 0 0 1 351.12 545.04 Tm <2620> Tj 1 0 0 1 363.60 545.04 Tm <4C6F73636865722C20> Tj 1 0 0 1 407.04 544.80 Tm <3139393761293A20> Tj 1 0 0 1 443.52 544.80 Tm <45696D6572696120> Tj 1 0 0 1 482.88 544.32 Tm <6D6178696D61> Tj 0.72 Tc 1 0 0 1 300.24 533.28 Tm <28426172746120> Tj 1 0 0 1 331.44 532.80 Tm <657420> Tj 1 0 0 1 343.20 532.56 Tm <616C2E2C20> Tj 1 0 0 1 361.68 532.56 Tm <31393938293B20> Tj 1 0 0 1 392.64 532.56 Tm <456368696E6F73746F6D6120> Tj 1 0 0 1 453.60 532.08 Tm <636170726F6E6920> Tj 1 0 0 1 491.52 532.56 Tm <284D6F722D> Tj 0.72 Tc 1 0 0 1 301.20 520.80 Tm <67616E20> Tj 1 0 0 1 320.88 520.08 Tm <2620> Tj 1 0 0 1 334.08 520.32 Tm <426C6169722C20> Tj 1 0 0 1 365.28 520.08 Tm <313939382920> Tj 1 0 0 1 395.76 519.84 Tm <616E6420> Tj 1 0 0 1 414.96 519.84 Tm <48796D656E6F6C6570697320> Tj 1 0 0 1 477.84 519.36 Tm <64696D696E757461> Tj 0.72 Tc 1 0 0 1 300.24 508.32 Tm <284F6B616D6F746F20> Tj 1 0 0 1 346.56 507.60 Tm <657420> Tj 1 0 0 1 357.12 507.36 Tm <616C2E2C20> Tj 1 0 0 1 374.64 507.60 Tm <31393937292E20> Tj 1 0 0 1 404.40 507.12 Tm <53657175656E636520> Tj 1 0 0 1 448.56 507.12 Tm <73696D696C617269747920> Tj 1 0 0 1 492.72 506.88 Tm <696E20> Tj 1 0 0 1 503.76 506.88 Tm <746865> Tj 0.48 Tc 1 0 0 1 300.48 495.60 Tm <49545320> Tj 1 0 0 1 317.76 495.36 Tm <726567696F6E20> Tj 1 0 0 1 346.32 495.36 Tm <68617320> Tj 1 0 0 1 362.40 495.12 Tm <616C736F20> Tj 1 0 0 1 380.88 495.36 Tm <70726F766964656420> Tj 1 0 0 1 420.24 494.64 Tm <65766964656E636520> Tj 1 0 0 1 459.60 494.64 Tm <666F7220> Tj 1 0 0 1 473.76 494.40 Tm <73796E6F6E6F6D79> Tj 0.48 Tc 1 0 0 1 300.96 482.88 Tm <65672E20> Tj 1 0 0 1 315.36 482.88 Tm <546520> Tj 1 0 0 1 325.44 482.88 Tm <6C61646F7220> Tj 1 0 0 1 347.52 482.64 Tm <7320> Tj 1 0 0 1 352.08 482.64 Tm <6120> Tj 1 0 0 1 356.64 482.88 Tm <676920> Tj 1 0 0 1 365.52 482.64 Tm <6120> Tj 1 0 0 1 373.92 482.40 Tm <7370702E20> Tj 1 0 0 1 392.40 482.88 Tm <2854726963686F7374726F6E67796C696461652920> Tj 1 0 0 1 481.44 482.40 Tm <2853746576656E2D> Tj 0.72 Tc 1 0 0 1 300.24 470.40 Tm <736F6E2C20> Tj 1 0 0 1 322.08 470.16 Tm <47617373657220> Tj 1 0 0 1 355.68 470.16 Tm <2620> Tj 1 0 0 1 368.40 469.92 Tm <4368696C746F6E2C20> Tj 1 0 0 1 408.96 469.92 Tm <313939362920> Tj 1 0 0 1 437.52 469.68 Tm <616E6420> Tj 1 0 0 1 456.48 469.68 Tm <46617363696F6C6120> Tj 1 0 0 1 500.16 469.44 Tm <7370702E> Tj 0.48 Tc 1 0 0 1 300.00 458.16 Tm <2848617368696D6F746F20> Tj 1 0 0 1 353.52 457.68 Tm <657420> Tj 1 0 0 1 364.56 457.44 Tm <616C2E2C20> Tj 1 0 0 1 380.88 457.68 Tm <31393937293B20> Tj 1 0 0 1 410.40 457.20 Tm <7375622D73706563696174696F6E20> Tj 1 0 0 1 475.44 456.96 Tm <652E672E20> Tj 1 0 0 1 492.96 456.96 Tm <456368692D> Tj 0.72 Tc 1 0 0 1 300.00 445.44 Tm <6E6F636F6363757320> Tj 1 0 0 1 342.72 445.20 Tm <6D756C74696C6F63756C6172697320> Tj 1 0 0 1 406.32 445.20 Tm <2852696E64657220> Tj 1 0 0 1 441.84 444.72 Tm <657420> Tj 1 0 0 1 452.16 444.72 Tm <616C2E2C20> Tj 1 0 0 1 469.20 444.72 Tm <31393937622920> Tj 1 0 0 1 501.12 444.48 Tm <616E64> Tj 0.72 Tc 1 0 0 1 300.00 432.96 Tm <50617261676F6E696D757320> Tj 1 0 0 1 360.00 432.72 Tm <7765737465726D616E6920> Tj 1 0 0 1 412.08 432.72 Tm <28426C61697220> Tj 1 0 0 1 439.68 432.24 Tm <657420> Tj 1 0 0 1 449.52 432.24 Tm <616C2E2C20> Tj 1 0 0 1 466.32 432.24 Tm <4920> Tj 1 0 0 1 470.64 432.00 Tm <3939372920> Tj 1 0 0 1 492.48 432.00 Tm <6F7220> Tj 1 0 0 1 503.52 432.00 Tm <746865> Tj 0.48 Tc 1 0 0 1 300.00 420.72 Tm <6964656E74696669636174696F6E20> Tj 1 0 0 1 361.68 420.24 Tm <6F6620> Tj 1 0 0 1 374.40 420.24 Tm <6372797074696320> Tj 1 0 0 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Shinn et al. - 185 rDNA qnd ITS sequences as population markers for Lepeophtheirus salmonis polymorphic enzymes, it was possible to obtainsome discrimination of lice from wild fish on theeast coast of Scotland from both wild and farmedfish sampled on the west coast of Scotland. Isdal,Nylund & Nevdal's (1997) allozyme study ofNorwegian populations of lice similarly founddifferences between northern and southern popu-lations of farmed lice. De Meeiis et al. (1992) usedallozymes to demonstrate an adaptive polymorphismto salinity in L. europaensis Zeddam, Berrebi, Re-naud, Raibaut & Gabrion, 1988 parasitic on Scoph-thalmus rhombus L., 1758 and Plqtichthys flesusL., 1758 which they suggested, when coupled witha restricted movement of hosts between environ-ments of differing salinity, could in time give riseto genetically isolated sub-populations.Other approaches in the investigation of thepopulation characteristics of Z. salmonis includethe use of inductively coupled plasma mass spec-trometry (fCPMS) to ascertain the elemental sig-nature of adult female lice (Shinn et al., this vol-ume). In this study it was demonstrated that adultfemale lice from seven localities could bp discrimi-nated from each other on the basis of l6 elementsand lice from farmed fish could be discriminatedfrom lice on wild salmonids within the same lo-cality.Molecular techniques permit finer resolution ofgenetic differences between populations and indi-viduals that cannot be detected using techniquessuch as morphometrics and allozyme analysis. Toddet al. (1997) using RAPDs, stated that genetic dif-ferentiation between populations of lice on farmedsalmon and wild salmonids and between individualfarm sites was evident to the extent that the pres-ence of putative "farm markers" within certainindividual wild lice suggested they originated fromfarm sites. While RAPDs allow for the compari-son of amplified DNA bands of similar molecularweight, without direct sampling it is unknownwhether these similar sized bands correspond tothe same portion of the genome. This study de-scribes an investigation which characterizes popu-lations through determination of the nucleotidesequence of selected areas of the L. salmonis ge-nome, namely the l8S ribosomal RNA gene andthe internal transcribed spacer (ITS).Information resultins from analvsis of the l85 rRNA gene sequence has previously been used todetermine whether or not the species under inves-tigation contains cryptic species or genetically iso-lated populations. Similarly, determination of thenucleotide sequence of internal transcribed spac-ers (ITS) have been successfully implemented asa taxonomic tool to discriminate closely related ormorphologically indistinguishable species of para-site e.g apicomplexans (Homan et al., 1997);gyrodactylids (Monogenea) (Cunningham, McGil-livray & MacKenzie, 1995; Cunningham, 1997);schistosomes (Digenea) (Littlewood & Johnston,1995; Van Herwerden, Blair & Agatsuma, 1998);and nematodes (Chilton, Gasser & Beveridge,1995). Nucleotide variations in ITS regions havealso been used to characterise strains or isolatesof Cyptocaryon irritans (Diggles & Adlard, 199'7);Enterocytozoon bieneusi (Rinder, Katzwinkel-Wladarsch & Loscher, 1997a): Eimeria maxima(Barta et al., 1998); Echinostoma caproni (Mor-gan & Blair, 1998) and Hymenolepis diminuta(Okamoto et al., 1997). Sequence similarity in theITS region has also provided evidence for synonomyeg. Te lador s a gi a spp. (Trichostrongylidae) (Steven-son, Gasser & Chilton, 1996) and Fasciola spp.(Hashimoto et al., 1997); sub-speciation e.g. Echi-nococcus multilocularis (Rinder et al., 1997b) andParagonimus westermani (Blair et al., I 997) or theidentification of cryptic species as has been sug-gested for geographically distanced isolates ofNecator americanus (Romstad et al., 1998). Overall,these studies suggest that the 18S and ITS regionsmay be used to provide a means of population char-acterization and it is with this in mind that thisstudy has been undertaken.Materials and methodsSample collection and preparationSea lice were collected from eight sites in Scot-land. 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Samples were then fro-zen in liquid nitrogen and stored at -70"C.Genomic DNA exlractionFrozen lice samples were kept on ice, homogenisedin a cell lysis buffer (1OmM Tris-HCl pH 8.0;150mM NaCl; lOmM EDTA-NaOH pH 8.0; andl% SDS) and then treated with proteinase K (lmg/ml) overnight at 55'C and RNaseA (0.5mg/ml) forone hour at 37"C to digest cellular proteins andRNA. Genomic DNA was then precipitated using0.3M NaOH-Ac and isolated using a standardphenolic-chloroform extraction (Sambrook et al.,1989) and precipitated in ice-cold l5o/o ethanol.The purified DNA pellets were dried at room tem-perature overnight and resuspended in TE buffer(lOmM Tris-HCl pH 8.0, lmM EDTA).PCR ampliJicationUniversal and species specific primers (given be-low) were selected for each gene and amplifiedusing PCR (Saiki et al., 1988).l8S ribosomal DIVAThe nematode Caenorhabditis elegans rRNA prim-ers complementary to the annealing positions 2074-2098 5'-GGGCAAGTCTGGTGCC-3' (18S-l) and2931-2946 5'-GGTCTGTGATGCCCTT-3', (l 8S-2) were used (Ellis, Sulston & Coulson, 1986). A25pl reaction mix was used, containing 25pM ofeach primer, 50-l00ng DNA and a PCR bead(Amersham Pharmacia-Biotech)(1.5 units of Taq,1OmM Tris-HCl, 50mM KCl, 1.5mM MgCl'200uM of each dNTP). The thermocycle programmewas 5 min at 95oC, 30 cycles of 30s at 95oC, 30sat 55'C. 90s at '72"C, then 5min at 72"C.Interual Transcribed Spacer (ITS)The Gyrodactylus salaris 18S rDNA forward primerITS- I 5'-TTTCCGTAGGTGAACCT-3' correspond- 7654 21# 91 Fig. l. Map of Lepeophtheiru,s salmonis collection sites inScotland. The sequence ofITS- I was determined and comparedfor individual lice collected from sites A-D. Lice from thesesites and from sites E-H were used to determine a partial sequenceof the l8S rRNA gene.ing to the annealing position 1,938-1955 (Cun-ningham, McGillivray & MacKenzie, 1995) andthe Scftislosoma 285 rDNA reverse primer ITS-25'-TCCTCCGCTTAGTGATA-3' (Littlewood &Johnston, 1995) were used as primers in the PCR.The PCR reaction mix was as for the l8S rDNAamplification. The thermocycling programme was5min at 95'C, 30 cycles of 60s at 95"C, 90s at 52"C,120s at 72oC, then 5min at l2'C.Cloning into pMOS Blue T:vectorPCR products were purified and ligated into pMOSBlue T-vector (pTTBlue) (Amersham, UK) using endstream endobj 22 0 obj <> endobj 16 0 obj <>/ProcSet[/PDF/Text/ImageB/ImageC/ImageI]/Subtype 21 0 R/Font<>>>/MediaBox[0 0 595.20 841.68]>> endobj 24 0 obj <>stream % CANON_PFINF_TYPE0_TEXTON endstream endobj 25 0 obj <>stream q 595.20 0 0 841.68 0.00 0.00 cm /Obj2 Do Q endstream endobj 26 0 obj <>stream BT 3 Tr 0.48 Tc /F3 9.0 Tf 1 0 0 1 84.72 800.88 Tm <412E502E20> Tj 1 0 0 1 106.56 800.64 Tm <5368696E6E20> Tj 1 0 0 1 133.68 800.64 Tm <657420> Tj 1 0 0 1 144.24 800.64 Tm <616C2E20> Tj 1 0 0 1 157.92 802.80 Tm <2D20> Tj 1 0 0 1 167.28 800.64 Tm <6C20> Tj 1 0 0 1 171.84 800.40 Tm <425320> Tj 1 0 0 1 185.28 800.40 Tm <72444E4120> Tj 1 0 0 1 213.60 800.16 Tm <616E6420> Tj 1 0 0 1 234.00 799.92 Tm <49545320> Tj 1 0 0 1 249.84 799.68 Tm 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<64697374616E636520> Tj 1 0 0 1 96.48 437.76 Tm <6D617472697820> Tj 1 0 0 1 128.16 437.52 Tm <616E6420> Tj 1 0 0 1 146.88 437.52 Tm <74686520> Tj 1 0 0 1 163.44 437.28 Tm <626F6F74737472617020> Tj 1 0 0 1 207.12 437.04 Tm <7472656520> Tj 1 0 0 1 226.56 436.80 Tm <776869636820> Tj 1 0 0 1 256.32 437.04 Tm <70726F2D> Tj 0.48 Tc 1 0 0 1 57.60 425.52 Tm <766964657320> Tj 1 0 0 1 84.00 425.28 Tm <636F6E666964656E636520> Tj 1 0 0 1 134.88 425.04 Tm <657374696D6174657320> Tj 1 0 0 1 178.56 424.80 Tm <666F7220> Tj 1 0 0 1 194.16 424.80 Tm <74686520> Tj 1 0 0 1 210.96 424.80 Tm <67726F7570696E677320> Tj 1 0 0 1 257.76 424.08 Tm <776173> Tj 0.72 Tc 1 0 0 1 57.60 413.04 Tm <647261776E20> Tj 1 0 0 1 87.84 412.80 Tm <7573696E6720> Tj 1 0 0 1 115.20 412.80 Tm <547265657669657720> Tj 1 0 0 1 160.56 412.56 Tm <312E3520> Tj 1 0 0 1 176.64 412.80 Tm <28506167652C20> Tj 1 0 0 1 208.56 412.32 Tm <31393936292E> Tj 0.48 Tc 1 0 0 1 57.36 375.84 Tm <526573756C7473> Tj 0.48 Tc 1 0 0 1 57.36 350.64 Tm <612920> Tj 1 0 0 1 71.04 350.64 Tm <31383520> Tj 1 0 0 1 87.36 350.64 Tm <7269626F736F6D716C20> Tj 1 0 0 1 132.24 350.40 Tm <444E41> Tj 0.48 Tc 1 0 0 1 57.36 325.68 Tm <54686520> Tj 1 0 0 1 77.04 325.92 Tm <7072656369736520> Tj 1 0 0 1 111.60 325.44 Tm <73657175656E636520> Tj 1 0 0 1 154.56 325.44 Tm <666F7220> Tj 1 0 0 1 170.40 325.20 Tm <6120> Tj 1 0 0 1 178.56 325.20 Tm <302E396B6220> Tj 1 0 0 1 206.16 324.96 Tm <667261676D656E7420> Tj 1 0 0 1 248.40 324.72 Tm <6F6620> Tj 1 0 0 1 260.64 324.72 Tm <746865> Tj 0.72 Tc 1 0 0 1 58.32 313.44 Tm <31385320> Tj 1 0 0 1 78.48 313.44 Tm <7269626F736F6D616C20> Tj 1 0 0 1 126.48 313.20 Tm <524E4120> Tj 1 0 0 1 153.60 313.20 Tm <67656E6520> Tj 1 0 0 1 178.32 312.72 Tm <77617320> Tj 1 0 0 1 199.68 312.48 Tm <64657465726D696E656420> Tj 1 0 0 1 252.96 312.48 Tm <66726F6D> Tj 0.48 Tc 1 0 0 1 57.36 300.96 Tm <656967687420> Tj 1 0 0 1 83.76 300.72 Tm <73656120> Tj 1 0 0 1 102.00 300.72 Tm <6C74636520> Tj 1 0 0 1 122.16 301.44 Tm <283220> Tj 1 0 0 1 135.84 300.72 Tm <66726F6D20> Tj 1 0 0 1 161.28 300.48 Tm <6561636820> Tj 1 0 0 1 185.76 300.24 Tm <7369746520> Tj 1 0 0 1 214.80 300.24 Tm <3220> Tj 1 0 0 1 225.36 300.00 Tm <77696C6420> Tj 1 0 0 1 249.60 299.76 Tm <616E6420> Tj 1 0 0 1 268.56 299.76 Tm <32> Tj 0.72 Tc 1 0 0 1 57.36 288.48 Tm <6661726D656420> Tj 1 0 0 1 91.20 288.24 Tm <7369746573292C20> Tj 1 0 0 1 119.76 288.00 Tm <7573696E6720> Tj 1 0 0 1 146.16 288.00 Tm <74776F20> Tj 1 0 0 1 165.36 288.00 Tm <636C6F6E657320> Tj 1 0 0 1 195.84 288.00 Tm <70657220> Tj 1 0 0 1 213.12 287.52 Tm <73706563696D656E20> Tj 1 0 0 1 256.80 287.76 Tm <28692E652E> Tj 0.48 Tc 1 0 0 1 57.12 276.00 Tm <6120> Tj 1 0 0 1 65.76 276.00 Tm <746F74616C20> Tj 1 0 0 1 89.28 275.76 Tm <6F6620> Tj 1 0 0 1 103.44 275.76 Tm <6C3620> Tj 1 0 0 1 117.12 275.76 Tm <636C6F6E65732920> Tj 1 0 0 1 152.40 275.52 Tm <616E6420> Tj 1 0 0 1 172.08 275.52 Tm <66726F6D20> Tj 1 0 0 1 196.80 275.28 Tm <626F746820> Tj 1 0 0 1 220.08 275.28 Tm <666F727761726420> Tj 1 0 0 1 258.24 274.80 Tm <616E64> Tj 0.48 Tc 1 0 0 1 56.88 263.52 Tm <7265766572736520> Tj 1 0 0 1 90.24 263.28 Tm <73657175656E63657320> Tj 1 0 0 1 135.60 263.28 Tm <666F7220> Tj 1 0 0 1 150.00 263.04 Tm <6561636820> Tj 1 0 0 1 171.60 262.80 Tm <636C6F6E652E20> Tj 1 0 0 1 199.44 262.80 Tm <467572746865722C20> Tj 1 0 0 1 235.44 262.56 Tm <74686520> Tj 1 0 0 1 250.80 262.56 Tm <444E41> Tj 0.48 Tc 1 0 0 1 57.36 251.04 Tm <666F7220> Tj 1 0 0 1 72.48 251.04 Tm <7468697320> Tj 1 0 0 1 91.20 250.80 Tm <726567696F6E20> Tj 1 0 0 1 121.68 250.56 Tm <77617320> Tj 1 0 0 1 141.36 250.56 Tm <6578747261637465642C20> Tj 1 0 0 1 186.24 250.56 Tm <70757269666965642C20> Tj 1 0 0 1 226.56 250.08 Tm <636C6F6E656420> Tj 1 0 0 1 258.24 249.84 Tm <616E64> Tj 0.48 Tc 1 0 0 1 57.36 238.56 Tm <73657175656E63656420> Tj 1 0 0 1 106.32 238.56 Tm <666F7220> Tj 1 0 0 1 123.12 238.32 Tm <6C69636520> Tj 1 0 0 1 142.56 238.32 Tm <74616B656E20> Tj 1 0 0 1 170.40 237.84 Tm <66726F6D20> Tj 1 0 0 1 195.12 237.84 Tm <74686520> Tj 1 0 0 1 212.88 237.84 Tm <73616D6520> Tj 1 0 0 1 239.04 237.60 Tm <73616D706C6573> Tj 0.48 Tc 1 0 0 1 57.12 226.08 Tm <6F6E20> Tj 1 0 0 1 71.04 226.08 Tm <74776F20> Tj 1 0 0 1 90.96 226.08 Tm <736570617261746520> Tj 1 0 0 1 129.60 225.84 Tm <6F63636173696F6E7320> Tj 1 0 0 1 174.72 225.84 Tm <706C6163696E6720> Tj 1 0 0 1 210.48 225.60 Tm <696E6372656173656420> Tj 1 0 0 1 254.64 225.12 Tm <636F6E2D> Tj 0.72 Tc 1 0 0 1 57.12 213.84 Tm <666964656E636520> Tj 1 0 0 1 91.68 213.60 Tm <696E20> Tj 1 0 0 1 102.48 213.60 Tm <74686520> Tj 1 0 0 1 118.80 213.36 Tm <73657175656E636520> Tj 1 0 0 1 160.80 213.36 Tm <6964656E746974792E20> Tj 1 0 0 1 199.20 213.12 Tm <4120> Tj 1 0 0 1 210.00 212.88 Tm <39303420> Tj 1 0 0 1 228.48 212.88 Tm <6E75636C656F74696465> Tj 0.48 Tc 1 0 0 1 57.12 201.60 Tm <286E742920> Tj 1 0 0 1 74.40 201.12 Tm <73657175656E636520> Tj 1 0 0 1 114.48 201.12 Tm <77617320> Tj 1 0 0 1 132.96 200.88 Tm <64657465726D696E656420> Tj 1 0 0 1 182.40 201.12 Tm <284669672E20> Tj 1 0 0 1 204.24 200.64 Tm <322920> Tj 1 0 0 1 215.28 200.64 Tm <666F7220> Tj 1 0 0 1 229.44 200.40 Tm <6561636820> Tj 1 0 0 1 250.80 200.16 Tm <636C6F6E65> Tj 0.48 Tc 1 0 0 1 56.88 188.88 Tm <7769746820> Tj 1 0 0 1 78.48 188.88 Tm <6E6F20> Tj 1 0 0 1 92.16 188.64 Tm <646966666572656E63657320> Tj 1 0 0 1 142.80 188.40 Tm <6265747765656E20> Tj 1 0 0 1 181.44 188.16 Tm <636C6F6E65732E20> Tj 1 0 0 1 214.32 188.16 Tm <6265747765656E20> Tj 1 0 0 1 253.20 187.92 Tm <696E64692D> Tj 0.72 Tc 1 0 0 1 300.72 774.00 Tm <76696475616C20> Tj 1 0 0 1 330.24 774.00 Tm <6C69636520> Tj 1 0 0 1 348.00 773.52 Tm <6F7220> Tj 1 0 0 1 359.04 773.52 Tm <6265747765656E20> Tj 1 0 0 1 397.68 773.28 Tm <73697465732E20> Tj 1 0 0 1 421.68 773.28 Tm <54686520> Tj 1 0 0 1 440.64 773.28 Tm <6D6F737420> Tj 1 0 0 1 464.16 772.80 Tm <73696D696C617220> Tj 1 0 0 1 496.32 772.80 Tm <637275732D> Tj 0.72 Tc 1 0 0 1 300.72 761.52 Tm <74616365616E20> Tj 1 0 0 1 335.76 761.52 Tm <31385320> Tj 1 0 0 1 357.12 761.52 Tm <70726576696F75736C7920> Tj 1 0 0 1 409.68 761.04 Tm <7265636F7264656420> Tj 1 0 0 1 454.32 760.56 Tm <697320> Tj 1 0 0 1 467.04 760.56 Tm <7468617420> Tj 1 0 0 1 489.60 760.32 Tm <6F6620> Tj 1 0 0 1 504.24 760.32 Tm <746865> Tj 0.48 Tc 1 0 0 1 300.72 748.80 Tm <6272616E6368697572616E20> Tj 1 0 0 1 355.68 749.04 Tm <706172617369746520> Tj 1 0 0 1 392.64 748.56 Tm <417267756C757320> Tj 1 0 0 1 431.04 748.08 Tm <6E6F62696C697320> Tj 1 0 0 1 464.16 747.84 Tm <7769746820> Tj 1 0 0 1 487.20 747.60 Tm <6120> Tj 1 0 0 1 495.60 747.60 Tm <73696D692D> Tj 0.72 Tc 1 0 0 1 300.96 736.56 Tm <6C617269747920> Tj 1 0 0 1 327.12 736.32 Tm <6F6620> Tj 1 0 0 1 340.08 736.08 Tm <38332E39322520> Tj 1 0 0 1 376.32 736.32 Tm <2839313420> Tj 1 0 0 1 399.12 735.84 Tm <6E742920> Tj 1 0 0 1 414.96 736.08 Tm <284162656C652C20> Tj 1 0 0 1 450.48 735.60 Tm <4B696D20> Tj 1 0 0 1 473.04 735.36 Tm <2620> Tj 1 0 0 1 484.56 735.36 Tm <46656C67656E2D> Tj 0.72 Tc 1 0 0 1 300.48 724.08 Tm <68617565722C20> Tj 1 0 0 1 333.84 723.84 Tm <31393839292E20> Tj 1 0 0 1 366.00 723.60 Tm <54686520> Tj 1 0 0 1 387.84 723.36 Tm <6E75636C656F7469646520> Tj 1 0 0 1 438.96 723.12 Tm <73657175656E636520> Tj 1 0 0 1 484.08 722.88 Tm <666F7220> Tj 1 0 0 1 501.36 722.64 Tm <74686973> Tj 0.72 Tc 1 0 0 1 300.48 711.84 Tm <7061727469616C20> Tj 1 0 0 1 332.16 711.12 Tm <73657175656E636520> Tj 1 0 0 1 375.12 710.88 Tm <6F6620> Tj 1 0 0 1 387.36 710.88 Tm <74686520> Tj 1 0 0 1 405.60 710.88 Tm <6C385320> Tj 1 0 0 1 424.32 710.64 Tm <7269626F736F6D616C20> Tj 1 0 0 1 471.12 710.40 Tm <524E4120> Tj 1 0 0 1 497.28 710.64 Tm <67656E65> Tj 0.48 Tc 1 0 0 1 300.48 698.88 Tm <68617320> Tj 1 0 0 1 318.24 698.64 Tm <6265656E20> Tj 1 0 0 1 342.48 698.64 Tm <7375626D697474656420> Tj 1 0 0 1 388.32 698.40 Tm <746F20> Tj 1 0 0 1 400.32 698.40 Tm <74686520> Tj 1 0 0 1 417.12 698.16 Tm <47656E42616E6B20> Tj 1 0 0 1 461.28 697.92 Tm <4461746120> Tj 1 0 0 1 485.28 697.92 Tm <4C696272617279> Tj 0.72 Tc 1 0 0 1 300.48 686.40 Tm <756E64657220> Tj 1 0 0 1 328.80 686.40 Tm <74686520> Tj 1 0 0 1 345.84 686.16 Tm <6163717569736974696F6E20> Tj 1 0 0 1 396.96 685.92 Tm <6E756D62657220> Tj 1 0 0 1 435.36 685.68 Tm <41463034333937392E> Tj 0.48 Tc 1 0 0 1 300.48 648.96 Tm <622920> Tj 1 0 0 1 312.00 648.96 Tm <496E7465726E616C20> Tj 1 0 0 1 348.96 648.72 Tm <5472616E7363726962656420> Tj 1 0 0 1 401.04 648.24 Tm <53706163657220> Tj 1 0 0 1 434.16 648.72 Tm <286C66732D3129> Tj 0.72 Tc 1 0 0 1 300.24 623.76 Tm <4120> Tj 1 0 0 1 312.00 623.76 Tm <746F74616C20> Tj 1 0 0 1 335.52 623.52 Tm <6F6620> Tj 1 0 0 1 348.48 623.52 Tm <7477656E747920> Tj 1 0 0 1 382.32 623.28 Tm <73656120> Tj 1 0 0 1 400.08 623.28 Tm <6C69636520> Tj 1 0 0 1 420.00 623.04 Tm <66726F6D20> Tj 1 0 0 1 445.20 622.80 Tm <73697820> Tj 1 0 0 1 461.76 622.80 Tm <736974657320> Tj 1 0 0 1 485.04 623.52 Tm <283320> Tj 1 0 0 1 498.00 622.56 Tm <77696C64> Tj 0.48 Tc 1 0 0 1 300.48 611.28 Tm <616E6420> Tj 1 0 0 1 320.64 611.04 Tm <3320> Tj 1 0 0 1 330.48 611.28 Tm <6661726D2920> Tj 1 0 0 1 358.80 610.80 Tm <7765726520> Tj 1 0 0 1 384.24 611.04 Tm <707265706172656420> Tj 1 0 0 1 427.20 610.56 Tm <666F7220> Tj 1 0 0 1 444.24 610.32 Tm <636C6F6E696E672E20> Tj 1 0 0 1 484.32 610.32 Tm <54776F20> Tj 1 0 0 1 508.32 609.84 Tm <6F72> Tj 0.48 Tc 1 0 0 1 300.24 598.80 Tm <746872656520> Tj 1 0 0 1 326.64 598.56 Tm <636C6F6E657320> Tj 1 0 0 1 359.04 598.56 Tm <66726F6D20> Tj 1 0 0 1 384.48 598.32 Tm <6561636820> Tj 1 0 0 1 408.96 598.32 Tm <6C6F75736520> Tj 1 0 0 1 436.32 597.84 Tm <7765726520> Tj 1 0 0 1 461.76 598.08 Tm <707265706172656420> Tj 1 0 0 1 504.72 597.60 Tm <666F72> Tj 0.48 Tc 1 0 0 1 300.48 586.32 Tm <73657175656E63696E672E20> Tj 1 0 0 1 352.08 586.08 Tm <466F727761726420> Tj 1 0 0 1 389.76 585.84 Tm <616E6420> Tj 1 0 0 1 406.56 585.84 Tm <7265766572736520> Tj 1 0 0 1 439.44 585.36 Tm <73657175656E63657320> Tj 1 0 0 1 483.84 585.36 Tm <666F7220> Tj 1 0 0 1 498.00 585.12 Tm <65616368> Tj 0.72 Tc 1 0 0 1 300.24 573.84 Tm <636C6F6E6520> Tj 1 0 0 1 328.32 573.84 Tm <7765726520> Tj 1 0 0 1 353.76 573.60 Tm <6F627461696E65642C20> Tj 1 0 0 1 398.64 573.36 Tm <616C69676E656420> Tj 1 0 0 1 435.36 573.12 Tm <616E6420> Tj 1 0 0 1 455.28 573.12 Tm <74686520> Tj 1 0 0 1 473.28 572.88 Tm <73657175656E636573> Tj 0.48 Tc 1 0 0 1 300.24 561.36 Tm <636F6D70617265642E20> Tj 1 0 0 1 348.72 561.12 Tm <576865726520> Tj 1 0 0 1 380.64 560.88 Tm <646966666572656E63657320> Tj 1 0 0 1 432.00 560.64 Tm <6F6363757272656420> Tj 1 0 0 1 473.04 560.64 Tm <696E20> Tj 1 0 0 1 484.32 560.40 Tm <74686520> Tj 1 0 0 1 500.88 560.40 Tm <666F722D> Tj 0.48 Tc 1 0 0 1 300.00 548.88 Tm <7761726420> Tj 1 0 0 1 324.00 548.64 Tm <616E6420> Tj 1 0 0 1 341.76 548.64 Tm <7265766572736520> Tj 1 0 0 1 375.60 548.40 Tm <73657175656E636520> Tj 1 0 0 1 417.12 548.16 Tm <616E6420> Tj 1 0 0 1 434.88 548.16 Tm <2F20> Tj 1 0 0 1 440.40 547.92 Tm <6F7220> Tj 1 0 0 1 451.20 547.92 Tm <6265747765656E20> Tj 1 0 0 1 489.84 547.68 Tm <636C6F6E6573> Tj 0.72 Tc 1 0 0 1 300.24 536.40 Tm <66726F6D20> Tj 1 0 0 1 324.96 536.16 Tm <6561636820> Tj 1 0 0 1 348.72 536.16 Tm <6C6F7573652C20> Tj 1 0 0 1 378.24 535.92 Tm <74686520> Tj 1 0 0 1 395.76 535.68 Tm <73657175656E63657320> Tj 1 0 0 1 443.52 535.68 Tm <7765726520> Tj 1 0 0 1 468.00 535.44 Tm <72656A65637465642E20> Tj 1 0 0 1 509.04 535.20 Tm <41> Tj 0.48 Tc 1 0 0 1 300.00 523.92 Tm <746F74616C20> Tj 1 0 0 1 322.80 523.68 Tm <6F6620> Tj 1 0 0 1 334.80 523.68 Tm <746869727465656E20> Tj 1 0 0 1 371.52 523.44 Tm <6C69636520> Tj 1 0 0 1 390.48 523.44 Tm <66726F6D20> Tj 1 0 0 1 414.48 523.20 Tm <666F757220> Tj 1 0 0 1 435.84 522.96 Tm <736974657320> Tj 1 0 0 1 458.40 523.44 Tm <283220> Tj 1 0 0 1 471.12 522.72 Tm <776C6C6420> Tj 1 0 0 1 494.16 522.48 Tm <616E6420> Tj 1 0 0 1 511.68 522.48 Tm <32> Tj 0.48 Tc 1 0 0 1 300.24 511.44 Tm <6661726D2920> Tj 1 0 0 1 325.92 511.20 Tm <7765726520> Tj 1 0 0 1 349.20 510.96 Tm <73656C656374656420> Tj 1 0 0 1 385.68 510.72 Tm <616E6420> Tj 1 0 0 1 403.20 510.72 Tm <636F6D706172656420> Tj 1 0 0 1 447.60 510.48 Tm <616E6420> Tj 1 0 0 1 465.12 510.24 Tm <6120> Tj 1 0 0 1 473.04 510.24 Tm <34353420> Tj 1 0 0 1 491.04 510.24 Tm <6E75636C652D> Tj 0.72 Tc 1 0 0 1 300.00 498.72 Tm <6F7469646520> Tj 1 0 0 1 325.44 498.72 Tm <73657175656E636520> Tj 1 0 0 1 368.40 498.48 Tm <77617320> Tj 1 0 0 1 388.80 498.24 Tm <6465726976656420> Tj 1 0 0 1 425.04 498.48 Tm <284669672E20> Tj 1 0 0 1 449.52 498.00 Tm <33292E20> Tj 1 0 0 1 464.64 498.00 Tm <4265747765656E20> Tj 1 0 0 1 506.16 497.52 Tm <616C6C> Tj 0.72 Tc 1 0 0 1 300.24 486.24 Tm <73616D706C657320> Tj 1 0 0 1 336.72 486.72 Tm <286E20> Tj 1 0 0 1 347.76 488.64 Tm <3A20> Tj 1 0 0 1 357.12 486.24 Tm <31332920> Tj 1 0 0 1 372.48 486.00 Tm <746865726520> Tj 1 0 0 1 396.24 485.76 Tm <77617320> Tj 1 0 0 1 415.20 485.76 Tm <616E20> Tj 1 0 0 1 427.68 485.52 Tm <6F766572616C6C20> Tj 1 0 0 1 460.32 485.52 Tm <39322E31346F2F6F20> Tj 1 0 0 1 496.08 485.28 Tm <73696D692D> Tj 0.72 Tc 1 0 0 1 300.00 474.00 Tm <6C617269747920> Tj 1 0 0 1 326.16 473.76 Tm <696E20> Tj 1 0 0 1 337.20 473.76 Tm <6E75636C656F7469646520> Tj 1 0 0 1 385.92 473.28 Tm <73657175656E63657320> Tj 1 0 0 1 432.72 473.04 Tm <7769746820> Tj 1 0 0 1 455.04 473.04 Tm <323820> Tj 1 0 0 1 468.72 473.04 Tm <7472616E736974696F6E732C> Tj 0.72 Tc 1 0 0 1 299.76 461.52 Tm <323820> Tj 1 0 0 1 314.40 461.28 Tm <7472616E7376657273696F6E732C20> Tj 1 0 0 1 379.44 461.04 Tm <6C3720> Tj 1 0 0 1 392.88 461.04 Tm <696E64656C7320> Tj 1 0 0 1 422.64 460.56 Tm <616E6420> Tj 1 0 0 1 441.84 460.56 Tm <3320> Tj 1 0 0 1 450.96 460.56 Tm <696E76657273696F6E7320> Tj 1 0 0 1 499.44 460.80 Tm <2862652D> Tj 0.72 Tc 1 0 0 1 299.76 449.04 Tm <747765656E20> Tj 1 0 0 1 330.24 448.80 Tm <3620> Tj 1 0 0 1 339.84 448.80 Tm <6E75636C656F7469646520> Tj 1 0 0 1 389.28 448.80 Tm <706F736974696F6E73292E20> Tj 1 0 0 1 439.68 448.08 Tm <57697468696E20> Tj 1 0 0 1 473.76 447.84 Tm <74686520> Tj 1 0 0 1 491.52 447.84 Tm <4954532D31> Tj 0.48 Tc 1 0 0 1 300.00 436.32 Tm <73657175656E63652C20> Tj 1 0 0 1 344.40 436.08 Tm <746865726520> Tj 1 0 0 1 368.88 435.84 Tm <61726520> Tj 1 0 0 1 384.24 435.84 Tm <746872656520> Tj 1 0 0 1 408.24 435.84 Tm <686967686C7920> Tj 1 0 0 1 438.72 435.60 Tm <636F6E73657276656420> Tj 1 0 0 1 484.80 435.36 Tm <726567696F6E73> Tj 0.96 Tc 1 0 0 1 299.52 423.84 Tm <6265747765656E20> Tj 1 0 0 1 341.76 423.84 Tm <6E75636C656F746964657320> Tj 1 0 0 1 398.88 423.60 Tm <312D33322C20> Tj 1 0 0 1 428.64 423.12 Tm <39313139342D3138332F31383920> Tj 1 0 0 1 500.16 423.12 Tm <28782F79> Tj 0.72 Tc 1 0 0 1 299.76 411.60 Tm <64656E6F74657320> Tj 1 0 0 1 335.76 411.36 Tm <74686520> Tj 1 0 0 1 352.32 411.36 Tm <72616E676520> Tj 1 0 0 1 379.68 411.12 Tm <6163726F737320> Tj 1 0 0 1 409.92 410.88 Tm <616C6C20> Tj 1 0 0 1 424.32 410.88 Tm <73706563696D656E73292C20> Tj 1 0 0 1 478.80 410.64 Tm <616E6420> Tj 1 0 0 1 497.28 410.64 Tm <3431302F> Tj 0.48 Tc 1 0 0 1 300.48 399.36 Tm <3431313434363134353620> Tj 1 0 0 1 357.60 398.88 Tm <616E6420> Tj 1 0 0 1 374.88 398.64 Tm <6F6E6520> Tj 1 0 0 1 392.16 398.88 Tm <68797065727661726961626C6520> Tj 1 0 0 1 452.40 398.40 Tm <726567696F6E20> Tj 1 0 0 1 481.44 398.16 Tm <6265747765656E> Tj 0.48 Tc 1 0 0 1 299.52 386.64 Tm <6261736520> Tj 1 0 0 1 320.16 386.88 Tm <706169727320> Tj 1 0 0 1 343.20 386.64 Tm <323839313239393430316C34316C20> Tj 1 0 0 1 417.12 386.16 Tm <2E20> Tj 1 0 0 1 421.44 386.16 Tm <54686520> Tj 1 0 0 1 439.92 385.92 Tm <73696D696C617269747920> Tj 1 0 0 1 481.44 385.68 Tm <6265747765656E> Tj 0.72 Tc 1 0 0 1 299.76 374.16 Tm <616C6C20> Tj 1 0 0 1 314.40 374.16 Tm <73657175656E63657320> Tj 1 0 0 1 362.16 374.16 Tm <697320> Tj 1 0 0 1 372.96 374.16 Tm <676976656E20> Tj 1 0 0 1 400.80 373.92 Tm <696E20> Tj 1 0 0 1 412.80 373.68 Tm <5461626C6520> Tj 1 0 0 1 442.32 373.68 Tm <3120> Tj 1 0 0 1 450.48 373.44 Tm <616E6420> Tj 1 0 0 1 469.68 373.44 Tm <6120> Tj 1 0 0 1 477.84 373.68 Tm <7068796C6F67652D> Tj 0.72 Tc 1 0 0 1 299.52 361.92 Tm <6E6574696320> Tj 1 0 0 1 326.16 361.68 Tm <7472656520> Tj 1 0 0 1 347.76 361.92 Tm <70726F647563656420> Tj 1 0 0 1 394.32 361.20 Tm <66726F6D20> Tj 1 0 0 1 420.72 360.96 Tm <64657465726D696E656420> Tj 1 0 0 1 475.20 360.72 Tm <626F6F747374726170> Tj 0.72 Tc 1 0 0 1 299.52 349.44 Tm <76616C75657320> Tj 1 0 0 1 329.76 349.68 Tm <283130303020> Tj 1 0 0 1 357.36 349.20 Tm <697465726174696F6E732920> Tj 1 0 0 1 403.92 348.96 Tm <697320> Tj 1 0 0 1 413.28 348.96 Tm <676976656E20> Tj 1 0 0 1 439.92 348.72 Tm <696E20> Tj 1 0 0 1 450.96 348.48 Tm <4669672E342E20> Tj 1 0 0 1 480.72 348.48 Tm <546F20> Tj 1 0 0 1 495.12 348.24 Tm <73756D2D> Tj 0.24 Tc 1 0 0 1 300.00 336.96 Tm <6D6172697A652C20> Tj 1 0 0 1 332.16 336.72 Tm <74686520> Tj 1 0 0 1 346.80 336.96 Tm <70657263656E7461676520> Tj 1 0 0 1 394.32 336.24 Tm <73696D696C617269747920> Tj 1 0 0 1 435.84 336.00 Tm <6265747765656E20> Tj 1 0 0 1 472.80 336.00 Tm <6C69636520> Tj 1 0 0 1 489.60 335.76 Tm <77697468696E> Tj 0.48 Tc 1 0 0 1 299.76 324.48 Tm <6661726D20> Tj 1 0 0 1 322.80 324.24 Tm <736974657320> Tj 1 0 0 1 344.88 324.00 Tm <77617320> Tj 1 0 0 1 364.32 324.00 Tm <39392E3120> Tj 1 0 0 1 384.00 324.00 Tm <6C256F20> Tj 1 0 0 1 400.56 323.76 Tm <616E6420> Tj 1 0 0 1 418.32 323.76 Tm <39352E3732256F20> Tj 1 0 0 1 453.84 323.76 Tm <666F7220> Tj 1 0 0 1 469.20 323.52 Tm <736974657320> Tj 1 0 0 1 490.80 323.52 Tm <4120> Tj 1 0 0 1 501.12 323.28 Tm <616E64> Tj 0.48 Tc 1 0 0 1 299.52 312.00 Tm <4420> Tj 1 0 0 1 310.80 312.00 Tm <726573706563746976656C7920> Tj 1 0 0 1 367.44 311.52 Tm <616E6420> Tj 1 0 0 1 387.36 311.52 Tm <38362E39302520> Tj 1 0 0 1 424.80 311.28 Tm <616E6420> Tj 1 0 0 1 444.00 311.28 Tm <38362E30336F2F6F20> Tj 1 0 0 1 480.24 311.04 Tm <6265747765656E> Tj 0.72 Tc 1 0 0 1 299.52 299.76 Tm <6C69636520> Tj 1 0 0 1 317.76 299.52 Tm <66726F6D20> Tj 1 0 0 1 341.04 299.28 Tm <77697468696E20> Tj 1 0 0 1 371.28 299.04 Tm <77696C6420> Tj 1 0 0 1 393.36 299.04 Tm <736974657320> Tj 1 0 0 1 414.96 299.04 Tm <4220> Tj 1 0 0 1 424.80 298.80 Tm <616E6420> Tj 1 0 0 1 442.80 298.80 Tm <432E20> Tj 1 0 0 1 455.28 298.80 Tm <5468697320> Tj 1 0 0 1 477.12 298.56 Tm <697320> Tj 1 0 0 1 486.72 298.56 Tm <636C6561726C79> Tj 0.72 Tc 1 0 0 1 299.52 287.04 Tm <73686F776E20> Tj 1 0 0 1 331.68 287.04 Tm <696E20> Tj 1 0 0 1 343.92 286.80 Tm <4669672E20> Tj 1 0 0 1 365.04 286.80 Tm <3420> Tj 1 0 0 1 374.64 286.56 Tm <776865726520> Tj 1 0 0 1 405.12 286.56 Tm <6C69636520> Tj 1 0 0 1 425.04 286.56 Tm <66726F6D20> Tj 1 0 0 1 450.24 286.32 Tm <73616C6D6F6E20> Tj 1 0 0 1 485.04 286.08 Tm <68656C6420> Tj 1 0 0 1 507.84 286.08 Tm <696E> Tj 0.72 Tc 1 0 0 1 299.52 274.56 Tm <6661726D20> Tj 1 0 0 1 324.72 274.32 Tm <636167657320> Tj 1 0 0 1 353.76 274.08 Tm <636C757374657220> Tj 1 0 0 1 387.84 274.08 Tm <746F67657468657220> Tj 1 0 0 1 429.12 273.84 Tm <696E20> Tj 1 0 0 1 442.08 273.84 Tm <74686520> Tj 1 0 0 1 460.32 273.60 Tm <64656E64726F6772616D2C> Tj 0.72 Tc 1 0 0 1 299.28 262.08 Tm <7768696C737420> Tj 1 0 0 1 330.72 261.84 Tm <73656120> Tj 1 0 0 1 348.96 261.84 Tm <6C69636520> Tj 1 0 0 1 369.12 261.84 Tm <66726F6D20> Tj 1 0 0 1 394.32 261.60 Tm <77696C6420> Tj 1 0 0 1 418.32 261.36 Tm <73616C6D6F6E69647320> Tj 1 0 0 1 466.80 261.12 Tm <61726520> Tj 1 0 0 1 484.56 261.12 Tm <646973747269622D> Tj 0.48 Tc 1 0 0 1 299.04 249.60 Tm <7574656420> Tj 1 0 0 1 321.60 249.60 Tm <7468726F7567686F757420> Tj 1 0 0 1 372.72 249.36 Tm <74686520> Tj 1 0 0 1 390.00 249.12 Tm <64656E64726F6772616D2E> Tj 0.72 Tc 1 0 0 1 309.84 237.36 Tm <4120> Tj 1 0 0 1 322.80 237.12 Tm <73657175656E636520> Tj 1 0 0 1 367.68 236.88 Tm <68617320> Tj 1 0 0 1 387.12 236.88 Tm <6265656E20> Tj 1 0 0 1 413.04 236.64 Tm <7375626D697474656420> Tj 1 0 0 1 461.28 236.40 Tm <746F20> Tj 1 0 0 1 474.96 236.16 Tm <47656E42616E6B> Tj 0.72 Tc 1 0 0 1 299.28 225.36 Tm <286163717569736974696F6E20> Tj 1 0 0 1 354.72 224.64 Tm <6E6F2E20> Tj 1 0 0 1 372.72 224.40 Tm <414630343339383020> Tj 1 0 0 1 423.60 224.16 Tm <666F7220> Tj 1 0 0 1 440.40 223.92 Tm <6120> Tj 1 0 0 1 449.76 224.16 Tm <6C6F75736520> Tj 1 0 0 1 476.88 223.92 Tm <636F6C6C6563746564> Tj 0.48 Tc 1 0 0 1 299.28 212.40 Tm <66726F6D20> Tj 1 0 0 1 323.28 212.16 Tm <6120> Tj 1 0 0 1 331.20 212.40 Tm <6661726D656420> Tj 1 0 0 1 364.80 211.92 Tm <73616C6D6F6E696420> Tj 1 0 0 1 407.52 212.88 Tm <287369746520> Tj 1 0 0 1 429.36 211.92 Tm <6E2779292E20> Tj 1 0 0 1 450.00 211.68 Tm <54686520> Tj 1 0 0 1 469.68 211.68 Tm <6D6F737420> Tj 1 0 0 1 493.92 211.44 Tm <73696D692D> Tj 0.72 Tc 1 0 0 1 299.28 200.16 Tm <6C617220> Tj 1 0 0 1 314.16 199.92 Tm <6372757374616365616E20> Tj 1 0 0 1 363.60 199.68 Tm <73657175656E636520> Tj 1 0 0 1 406.80 199.44 Tm <696E20> Tj 1 0 0 1 418.56 199.20 Tm <47656E42616E6B20> Tj 1 0 0 1 462.72 199.20 Tm <77617320> Tj 1 0 0 1 482.88 199.20 Tm <74686520> Tj 1 0 0 1 500.16 198.96 Tm <495453> Tj 0.48 Tc 1 0 0 1 299.28 187.44 Tm <73657175656E636520> Tj 1 0 0 1 342.24 187.44 Tm <666F7220> Tj 1 0 0 1 358.08 187.44 Tm <74686520> Tj 1 0 0 1 375.12 187.20 Tm <6F73747261636F6420> Tj 1 0 0 1 415.20 187.20 Tm <44617277696E756C6120> Tj 1 0 0 1 465.84 186.72 Tm 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J|JpՊcK+kaޖ֒}:OOF!0ڄM6VKMi2ڶ8.K-<_,ud0„B!@Ah iIi/KKIR}v<^%UVUS zB#""AR LBdW!_\:pب=%l%uiz_q 0 UTATUP⢘BvDDDDDE &dꩅMC UDDDDDDDDaZqMIG, (k߮?岪>[)_ V?Wz 3_-k endstream endobj 28 0 obj <>stream A.P. Shinn et al. - l BS rDNA and ITS sequences as population markers for Lepeophtheirus salmonis Novagen blunt cloning kits. Transformations werecarried out using NovaBlue competent cells andscreened on LB agar plates containing ampicillin(100 pglml final conc.), tetracycline (25 pglmlfinalconc.), X-gal (0.5p1 / ml final conc.) and IPTG(0.1 pl/ml final conc.).Dideoxynucleotide sequencing and analysisDirect colony PCR was used as a rapid screeningmethod to ensure correct insertion of the selectedproduct. Plasmid DNA samples were sequencedusing the dideoxy method (Sanger, Nicklen &Coulsen, 1977) using fluorescently labelled nucle-otides. Either a LI-COR Model 4000L sequenceror a Perkin Elmer ABI PRISM 377 was used fol-lowing amplification with Ml3 forward and re-verse primers. Sequence data was analysed usingMT Navigator ver. 1.02b3 (1994) (Applied Bio-systems Inc., Perkin Elmer) and ClustalX (1.5b)for the multiple alignment of genetic sequences.A phylogenetic tree was constructed using Saitou& Nei's Neighbour Joining Method (NJ) (Saitou& Nei, 1987) which calculates distances (% di-vergence) between all pairs of sequence from amultiple alignment. The NJ is then applied to thedistance matrix and the bootstrap tree which pro-vides confidence estimates for the groupings wasdrawn using Treeview 1.5 (Page, 1996).Resultsa) 185 ribosomql DNAThe precise sequence for a 0.9kb fragment of the18S ribosomal RNA gene was determined fromeight sea ltce (2 from each site 2 wild and 2farmed sites), using two clones per specimen (i.e.a total of l6 clones) and from both forward andreverse sequences for each clone. Further, the DNAfor this region was extracted, purified, cloned andsequenced for lice taken from the same sampleson two separate occasions placing increased con-fidence in the sequence identity. A 904 nucleotide(nt) sequence was determined (Fig. 2) for each clonewith no differences between clones. between indi- vidual lice or between sites. The most similar crus-tacean 18S previously recorded is that of thebranchiuran parasite Argulus nobilis with a simi-larity of 83.92% (914 nt) (Abele, Kim & Felgen-hauer, 1989). The nucleotide sequence for thispartial sequence of the l8S ribosomal RNA genehas been submitted to the GenBank Data Libraryunder the acquisition number AF043979.b) Internal Transcribed Spacer (lfs-1)A total of twenty sea lice from six sites (3 wildand 3 farm) were prepared for cloning. Two orthree clones from each louse were prepared forsequencing. Forward and reverse sequences for eachclone were obtained, aligned and the sequencescompared. Where differences occurred in the for-ward and reverse sequence and / or between clonesfrom each louse, the sequences were rejected. Atotal of thirteen lice from four sites (2 wlld and 2farm) were selected and compared and a 454 nucle-otide sequence was derived (Fig. 3). Between allsamples (n : 13) there was an overall 92.14o/o simi-larity in nucleotide sequences with 28 transitions,28 transversions, l7 indels and 3 inversions (be-tween 6 nucleotide positions). Within the ITS-1sequence, there are three highly conserved regionsbetween nucleotides 1-32, 91194-183/189 (x/ydenotes the range across all specimens), and 410/4114461456 and one hypervariable region betweenbase pairs 2891299401l41l . The similarity betweenall sequences is given in Table 1 and a phyloge-netic tree produced from determined bootstrapvalues (1000 iterations) is given in Fig.4. To sum-marize, the percentage similarity between lice withinfarm sites was 99.1 l%o and 95.72%o for sites A andD respectively and 86.90% and 86.03o/o betweenlice from within wild sites B and C. This is clearlyshown in Fig. 4 where lice from salmon held infarm cages cluster together in the dendrogram,whilst sea lice from wild salmonids are distrib-uted throughout the dendrogram.A sequence has been submitted to GenBank(acquisition no. AF043980 for a louse collectedfrom a farmed salmonid (site n'y). 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Tm <412E20> Tj 1 0 0 1 52.32 609.84 Tm <6E6F62696C2D6973> Tj 1.44 Tc 1 0 0 1 33.36 592.80 Tm <4C2E20> Tj 1 0 0 1 52.80 592.80 Tm <7361666D6F6E6973> Tj 1.44 Tc 1 0 0 1 33.12 584.40 Tm <412E20> Tj 1 0 0 1 52.56 584.40 Tm <6E6F626C666973> Tj 1.44 Tc 1 0 0 1 33.60 567.60 Tm <4C2E20> Tj 1 0 0 1 52.80 567.60 Tm <7361666D6F6E6973> Tj 1.20 Tc 1 0 0 1 33.12 559.20 Tm <412E20> Tj 1 0 0 1 52.56 559.20 Tm <6E6F62696C2D6973> Tj 1.44 Tc 1 0 0 1 33.60 542.16 Tm <4C2E20> Tj 1 0 0 1 53.04 542.16 Tm <73614C6D6F6E6973> Tj 1.44 Tc 1 0 0 1 33.36 533.76 Tm <412E20> Tj 1 0 0 1 52.56 533.76 Tm <6E6F62694C6973> Tj 1.20 Tc 1 0 0 1 33.84 516.48 Tm <4C2E20> Tj 1 0 0 1 53.04 516.72 Tm <73616C2D6D6F6E6973> Tj 1.20 Tc 1 0 0 1 33.36 508.32 Tm <412E20> Tj 1 0 0 1 52.80 508.32 Tm <6E6F62696C2D6973> Tj 1.44 Tc 1 0 0 1 33.84 491.52 Tm <4C2E20> Tj 1 0 0 1 53.28 491.28 Tm <7361666D6F6E6973> Tj 1.44 Tc 1 0 0 1 33.36 482.88 Tm <412E20> Tj 1 0 0 1 52.80 483.12 Tm <6E6F6269666973> Tj 1.44 Tc 1 0 0 1 33.84 466.08 Tm <4C2E20> Tj 1 0 0 1 53.28 466.08 Tm <73614C6D6F6E6973> Tj 1.44 Tc 1 0 0 1 33.60 457.68 Tm <412E20> Tj 1 0 0 1 53.04 457.68 Tm <6E6F6269666973> Tj 1.44 Tc 1 0 0 1 33.84 440.88 Tm <4C2E20> Tj 1 0 0 1 53.28 440.64 Tm <7361666D6F6E6973> Tj 1.20 Tc 1 0 0 1 33.60 432.24 Tm <412E20> Tj 1 0 0 1 53.04 432.48 Tm <6E6F62696C2D6973> Tj 1.20 Tc 1 0 0 1 34.08 415.68 Tm <4C2E20> Tj 1 0 0 1 53.28 415.68 Tm <73616C2D6D6F6E6973> Tj 1.44 Tc 1 0 0 1 33.60 407.28 Tm <412E20> Tj 1 0 0 1 53.04 407.28 Tm <6E6F6269666973> Tj 0.96 Tc 1 0 0 1 34.08 390.72 Tm <4C20> Tj 1 0 0 1 42.00 390.72 Tm <2E20> Tj 1 0 0 1 53.52 390.72 Tm <736120> Tj 1 0 0 1 66.24 390.96 Tm <6C2D6D6F6E6973> Tj 1.20 Tc 1 0 0 1 33.84 382.56 Tm <412E20> Tj 1 0 0 1 53.28 382.56 Tm <6E6F62696C2D6973> Tj 1.44 Tc 1 0 0 1 34.32 365.76 Tm <4C2E20> Tj 1 0 0 1 53.52 365.76 Tm <73614C6D6F6E6973> Tj 1.44 Tc 1 0 0 1 33.84 357.36 Tm <412E20> Tj 1 0 0 1 53.52 357.60 Tm <6E6F6269666973> Tj 0.96 Tc 1 0 0 1 34.32 340.56 Tm <4C20> Tj 1 0 0 1 42.24 340.56 Tm <2E20> Tj 1 0 0 1 53.76 340.80 Tm <736120> Tj 1 0 0 1 66.48 340.80 Tm <6C2D6D6F6E6973> Tj 1.20 Tc 1 0 0 1 34.08 332.16 Tm <412E20> Tj 1 0 0 1 53.52 332.40 Tm <6E6F62696C2D6973> Tj 1.50 Tc 1 0 0 1 34.56 315.12 Tm <4C2E20> Tj 1 0 0 1 54.00 315.12 Tm <73614C6D6F6E6973> Tj 1.44 Tc 1 0 0 1 34.32 306.72 Tm <412E20> Tj 1 0 0 1 53.28 306.72 Tm <6E6F6269666973> Tj 0.48 Tc /F3 23.0 Tf 1 0 0 1 138.72 761.76 Tm <66664839543F5474725466663F693A3A693A3A3A3A3A3A61545454543A3A693A3A543A3A5454693A3A3A54695420> Tj 1 0 0 1 470.88 761.28 Tm <3333> Tj 0.72 Tc /F3 9.0 Tf 1 0 0 1 138.48 744.96 Tm <41545441414147545447545447434747544341275555555C47435443475441475454474741545454434747544741475441414720> Tj 1 0 0 1 471.36 744.48 Tm <3939> Tj 1.50 Tc 1 0 0 1 202.08 736.32 Tm <632E2E2E2E2E2E542E2E20> Tj 1 0 0 1 360.96 736.08 Tm <2E2E63632E412E2E54474341472E4320> Tj 1 0 0 1 471.12 736.08 Tm <313030> Tj 0.00 Tc 1 0 0 1 138.96 719.28 Tm <47414720> Tj 1 0 0 1 158.16 719.28 Tm <4747475420> Tj 1 0 0 1 183.36 719.28 Tm <5420> Tj 1 0 0 1 189.60 719.28 Tm <43415420> Tj 1 0 0 1 208.56 719.28 Tm <4320> Tj 1 0 0 1 214.80 719.28 Tm <5420> Tj 1 0 0 1 221.04 719.28 Tm <5420> Tj 1 0 0 1 227.28 719.28 Tm <475420> Tj 1 0 0 1 239.76 719.28 Tm <5420> Tj 1 0 0 1 246.24 719.28 Tm <5420> Tj 1 0 0 1 252.48 719.28 Tm <47415420> Tj 1 0 0 1 271.20 719.04 Tm <475441414320> Tj 1 0 0 1 302.64 719.28 Tm <5420> Tj 1 0 0 1 308.88 719.04 Tm <474320> Tj 1 0 0 1 321.60 719.28 Tm <54475420> Tj 1 0 0 1 340.32 719.04 Tm <435420> Tj 1 0 0 1 352.80 719.04 Tm <5420> Tj 1 0 0 1 359.04 719.04 Tm <475420> Tj 1 0 0 1 371.76 719.04 Tm <435420> Tj 1 0 0 1 384.24 719.04 Tm <4320> Tj 1 0 0 1 390.72 719.04 Tm <4743475420> Tj 1 0 0 1 415.68 719.04 Tm <5420> Tj 1 0 0 1 422.16 719.04 Tm <5420> Tj 1 0 0 1 428.40 719.04 Tm <545420> Tj 1 0 0 1 441.12 719.04 Tm <4347> Tj 1.50 Tc 1 0 0 1 140.88 710.88 Tm <2E472E432E20> Tj 1 0 0 1 172.32 710.88 Tm <2E20> Tj 1 0 0 1 178.56 710.88 Tm <2E432E47432E20> Tj 1 0 0 1 216.48 710.88 Tm <2E43412E4747434747542E432E20> Tj 1 0 0 1 298.08 710.64 Tm <2E20> Tj 1 0 0 1 304.32 710.64 Tm <2E20> Tj 1 0 0 1 310.80 710.64 Tm <2E2E435443472E20> Tj 1 0 0 1 354.72 710.64 Tm <2E432E20> Tj 1 0 0 1 373.44 710.64 Tm <2E47414341544143432E432E20> Tj 1 0 0 1 449.04 710.64 Tm <2E> Tj 0.72 Tc 1 0 0 1 138.48 694.32 Tm <414341474147475420> Tj 1 0 0 1 189.60 694.08 Tm <5420> Tj 1 0 0 1 196.08 694.08 Tm <5420> Tj 1 0 0 1 202.56 696.72 Tm <2D20> Tj 1 0 0 1 208.80 696.72 Tm <2D20> Tj 1 0 0 1 215.28 696.96 Tm <2D20> Tj 1 0 0 1 221.28 694.08 Tm <4347415420> Tj 1 0 0 1 246.24 694.08 Tm <47475447434354545443414141474747544754435420> Tj 1 0 0 1 378.24 693.84 Tm <434747474154414354475443> Tj 1.50 Tc 1 0 0 1 139.20 685.68 Tm <4754542E20> Tj 1 0 0 1 166.32 685.92 Tm <2E54432E20> Tj 1 0 0 1 191.52 685.68 Tm <2E20> Tj 1 0 0 1 197.76 685.68 Tm <2E4754432E4347432E41542E20> Tj 1 0 0 1 273.12 685.68 Tm <2E54432E20> Tj 1 0 0 1 298.08 685.68 Tm <2E20> Tj 1 0 0 1 304.32 685.68 Tm <2E20> Tj 1 0 0 1 310.80 685.68 Tm <2E54432E2E20> Tj 1 0 0 1 372.00 685.44 Tm <472E20> Tj 1 0 0 1 386.16 685.44 Tm <2E2E2E20> Tj 1 0 0 1 405.12 685.44 Tm <2E47472E432E4747> Tj 0.48 Tc 1 0 0 1 139.20 669.12 Tm <4747475420> Tj 1 0 0 1 164.64 669.36 Tm <5420> Tj 1 0 0 1 170.88 669.36 Tm <54414354545420> Tj 1 0 0 1 208.80 669.12 Tm <4720> Tj 1 0 0 1 215.52 669.12 Tm <4E5555555C2C5C545441474147544743544341414147434147474354545420> Tj 1 0 0 1 397.20 669.12 Tm <5420> Tj 1 0 0 1 403.68 669.12 Tm <4141414147435454> Tj 1.50 Tc 1 0 0 1 139.44 661.92 Tm <416320> Tj 1 0 0 1 298.32 660.72 Tm <2E2E2E542E20> Tj 1 0 0 1 360.96 660.48 Tm <2E2E2E47412E2E2D2D2D2D2E2E4341> Tj 0.48 Tc 1 0 0 1 139.20 643.68 Tm <4741415441545441475420> Tj 1 0 0 1 202.56 643.68 Tm <47434154474741415441415420> Tj 1 0 0 1 277.92 643.68 Tm <47414141544147474141475420> Tj 1 0 0 1 353.28 643.44 Tm <434754545420> Tj 1 0 0 1 384.72 643.44 Tm <4354415454545447545447> Tj 1.50 Tc 1 0 0 1 170.40 635.28 Tm <412E472E2E20> Tj 1 0 0 1 329.76 635.04 Tm <2E43432E2E2E472E2E2E2E2E2E432E> Tj 0.72 Tc 1 0 0 1 139.44 618.24 Tm <47545420> Tj 1 0 0 1 158.88 621.12 Tm <2D20> Tj 1 0 0 1 164.64 618.48 Tm <5420> Tj 1 0 0 1 171.12 618.48 Tm <5420> Tj 1 0 0 1 177.36 618.24 Tm <4347474E555C5420> Tj 1 0 0 1 221.52 618.24 Tm <434741435420> Tj 1 0 0 1 252.96 618.24 Tm <5441415447415454414154414747474143414754434747474747434154544147> Tj 1.50 Tc 1 0 0 1 141.12 609.84 Tm <2E2E2E632E20> Tj 1 0 0 1 204.24 609.84 Tm <2E43412E2E2E47472E2E20> Tj 1 0 0 1 359.52 609.60 Tm <472E412E2E20> Tj 1 0 0 1 424.08 609.60 Tm <2E2E2E432E> Tj 0.48 Tc 1 0 0 1 139.44 593.04 Tm <5441545420> Tj 1 0 0 1 164.88 592.80 Tm <434147414347414341474147475447414141545420> Tj 1 0 0 1 290.64 592.80 Tm <4354544747414343475443544741414741435441414354414354> Tj 1.50 Tc 1 0 0 1 141.36 584.64 Tm <2E2E2E2E47432E2E2E2E545420> Tj 1 0 0 1 311.28 584.40 Tm <2E2E2E542E2E2E2E474320> Tj 1 0 0 1 403.68 584.40 Tm <472E2E2E20> Tj 1 0 0 1 430.80 584.40 Tm <5F2E2E2E> Tj 0.48 Tc /F3 22.0 Tf 1 0 0 1 139.68 559.44 Tm <3A3A3A543A3A6954545420> Tj 1 0 0 1 215.52 559.20 Tm <3A3A3A543A54543A5420> Tj 1 0 0 1 284.40 559.20 Tm <5454543A415454545420> Tj 1 0 0 1 347.28 558.96 Tm <3A543A543A3A593A5420> Tj 1 0 0 1 429.12 559.20 Tm <54693A69> Tj 1.20 Tc /F3 4.0 Tf 1 0 0 1 139.68 546.00 Tm <2D5E74362D2D3120> Tj 1 0 0 1 184.32 546.00 Tm <6E20> Tj 1 0 0 1 190.32 546.00 Tm <5E5E5E5E6E20> Tj 1 0 0 1 221.76 546.00 Tm <6D5E6E20> Tj 1 0 0 1 240.48 546.00 Tm <5E5E66724120> Tj 1 0 0 1 265.68 546.00 Tm <69645E643720> Tj 1 0 0 1 296.88 546.00 Tm <69366C20> Tj 1 0 0 1 315.84 546.00 Tm <5E6D6D5E6D6E20> Tj 1 0 0 1 354.00 546.00 Tm <6E20> Tj 1 0 0 1 359.76 546.00 Tm <695E6E20> Tj 1 0 0 1 378.72 546.00 Tm <6D6E20> Tj 1 0 0 1 391.68 546.00 Tm <6E20> Tj 1 0 0 1 398.16 546.00 Tm <3120> Tj 1 0 0 1 403.92 546.00 Tm <5E2D6E20> Tj 1 0 0 1 422.88 546.00 Tm <6D5E5E5E6E> Tj 0.48 Tc 1 0 0 1 139.68 542.40 Tm <555520> Tj 1 0 0 1 153.36 542.40 Tm <6620> Tj 1 0 0 1 159.84 542.40 Tm <4920> Tj 1 0 0 1 165.12 542.40 Tm <555566666955555555414920> Tj 1 0 0 1 228.00 542.40 Tm <5541554120> Tj 1 0 0 1 254.16 542.40 Tm <5420> Tj 1 0 0 1 258.72 542.40 Tm <4155555555555520> Tj 1 0 0 1 304.32 542.40 Tm <6620> Tj 1 0 0 1 309.12 542.40 Tm <415520> Tj 1 0 0 1 323.04 542.40 Tm <4920> Tj 1 0 0 1 329.28 542.40 Tm <4920> Tj 1 0 0 1 334.56 542.40 Tm <5520> Tj 1 0 0 1 342.00 542.40 Tm <4920> Tj 1 0 0 1 346.80 542.40 Tm <6C2D66695555414920> Tj 1 0 0 1 384.48 542.40 Tm <6666695555415420> Tj 1 0 0 1 429.12 542.40 Tm <55555541> Tj 0.72 Tc /F3 9.0 Tf 1 0 0 1 139.92 516.72 Tm <474354414743474154434347434147545420> Tj 1 0 0 1 246.96 516.48 Tm <4754545441545420> Tj 1 0 0 1 290.88 516.72 Tm <5420> Tj 1 0 0 1 297.12 516.72 Tm <54544143474143544354474347474741414743545443434747> Tj 1.50 Tc /F3 7.0 Tf 1 0 0 1 140.40 510.24 Tm <41542D54722D43542D412741723A4154542D723A4141722D> Tj 1.20 Tc /F3 4.0 Tf 1 0 0 1 139.92 495.36 Tm <2D6E20> Tj 1 0 0 1 153.12 495.36 Tm <6E20> Tj 1 0 0 1 159.36 495.36 Tm <6E5E5E6E20> Tj 1 0 0 1 184.56 495.36 Tm <6E20> Tj 1 0 0 1 191.04 495.36 Tm <745E5E6D6D6D6D5E2D2D66726D69692D2D2D5E2D6E20> Tj 1 0 0 1 316.56 495.36 Tm <6E5E6D746D5E5E6D6D5E6E6C20> Tj 1 0 0 1 391.92 495.36 Tm <6E20> Tj 1 0 0 1 398.16 495.36 Tm <6E5E5E6D5E6E20> Tj 1 0 0 1 436.08 495.36 Tm <6E20> Tj 1 0 0 1 442.56 495.36 Tm <6E5E> Tj 0.24 Tc 1 0 0 1 139.92 491.76 Tm <5C4A66666955556666695C4A5520> Tj 1 0 0 1 210.48 491.76 Tm <4920> Tj 1 0 0 1 216.72 491.76 Tm <4920> Tj 1 0 0 1 222.96 491.52 Tm <4920> Tj 1 0 0 1 229.20 491.52 Tm <4A2D20> Tj 1 0 0 1 234.48 491.52 Tm <555C665C6620> Tj 1 0 0 1 254.40 491.52 Tm <4920> Tj 1 0 0 1 260.64 491.52 Tm <4A2E20> Tj 1 0 0 1 265.92 491.52 Tm <55555C375C545C4A5C4A5C66496966695C4A20> Tj 1 0 0 1 329.76 491.52 Tm <4C20> Tj 1 0 0 1 334.56 491.52 Tm <66742E4C20> Tj 1 0 0 1 347.52 491.52 Tm <5C4A5C4A20> Tj 1 0 0 1 361.20 491.52 Tm <4920> Tj 1 0 0 1 367.44 491.52 Tm <7420> Tj 1 0 0 1 372.72 491.52 Tm <5C72557D31746C7D495C4A5520> Tj 1 0 0 1 417.84 491.52 Tm <4920> Tj 1 0 0 1 422.88 491.76 Tm <5C4A2731277466315C2D> Tj 1.50 Tc 1 0 0 1 203.04 486.96 Tm <6D6964> Tj 0.48 Tc /F3 9.0 Tf 1 0 0 1 139.92 466.32 Tm <54544141414747414154544741434747414147474743414343414343414747414754475F20> Tj 1 0 0 1 366.72 468.96 Tm <2D20> Tj 1 0 0 1 372.72 466.08 Tm <4743435420> Tj 1 0 0 1 397.92 466.08 Tm <474347474354544141> Tj 1.50 Tc 1 0 0 1 360.00 457.68 Tm <47412E20> Tj 1 0 0 1 380.64 457.68 Tm <2E2E> Tj 1.20 Tc /F3 4.0 Tf 1 0 0 1 139.92 444.96 Tm <66726D6D5E6E20> Tj 1 0 0 1 171.60 444.96 Tm <5E6D6E20> Tj 1 0 0 1 191.04 444.96 Tm <6E20> Tj 1 0 0 1 197.28 444.96 Tm <3120> Tj 1 0 0 1 203.28 444.96 Tm <694120> Tj 1 0 0 1 215.76 444.96 Tm <5E5E5E2D3120> Tj 1 0 0 1 247.68 444.96 Tm <6E20> Tj 1 0 0 1 253.92 444.96 Tm <6E20> Tj 1 0 0 1 259.68 444.96 Tm <6D5E6D6E6E20> Tj 1 0 0 1 291.12 444.96 Tm <5E643120> Tj 1 0 0 1 310.08 444.96 Tm <6969414164465E5E20> Tj 1 0 0 1 360.24 444.96 Tm <413120> Tj 1 0 0 1 373.20 444.96 Tm <69667246695E20> Tj 1 0 0 1 404.64 444.96 Tm <6E20> Tj 1 0 0 1 410.64 444.96 Tm <5E5E6C20> Tj 1 0 0 1 429.36 444.96 Tm <6D6D5E6E> Tj 0.24 Tc 1 0 0 1 141.12 441.12 Tm <4A2D20> Tj 1 0 0 1 147.36 441.12 Tm <4920> Tj 1 0 0 1 153.60 441.12 Tm <492E20> Tj 1 0 0 1 159.12 441.12 Tm <552E497B5520> Tj 1 0 0 1 179.04 441.12 Tm <4920> Tj 1 0 0 1 184.08 441.12 Tm <5C2D666966745C2D66745C2D5C4A4C545C4A666C7D742E7474272E4C20> Tj 1 0 0 1 265.92 441.12 Tm <552E4C20> Tj 1 0 0 1 278.64 441.12 Tm <55415C2D55414C5D284A5C2D5C2D554C66554155415520> Tj 1 0 0 1 379.92 441.12 Tm <2D7420> Tj 1 0 0 1 385.44 441.12 Tm <5C4A5C4A667B66495C4A5C4A7D6C2E20> Tj 1 0 0 1 430.56 441.12 Tm <4920> Tj 1 0 0 1 436.80 441.12 Tm <4920> Tj 1 0 0 1 442.08 441.12 Tm <4C6A5469> Tj 1.50 Tc 1 0 0 1 260.16 436.56 Tm <646D2D> Tj 1.50 Tc 1 0 0 1 361.92 432.48 Tm <2E2E2E2E312E392E2E2E> Tj 1.20 Tc 1 0 0 1 140.16 419.04 Tm <5E6C20> Tj 1 0 0 1 152.64 419.04 Tm <6E6C20> Tj 1 0 0 1 165.36 419.04 Tm <546D616C20> Tj 1 0 0 1 190.80 419.04 Tm <616E20> Tj 1 0 0 1 203.28 419.04 Tm <615E6D5E6D6D6D6E6E6C6120> Tj 1 0 0 1 272.88 419.04 Tm <4120> Tj 1 0 0 1 278.64 419.04 Tm <6E344620> Tj 1 0 0 1 297.36 419.04 Tm <6120> Tj 1 0 0 1 303.84 419.04 Tm <2D6D6120> Tj 1 0 0 1 322.56 419.04 Tm <6120> Tj 1 0 0 1 328.80 419.04 Tm <613420> Tj 1 0 0 1 341.52 419.04 Tm <6120> Tj 1 0 0 1 347.76 419.04 Tm <616D6C20> Tj 1 0 0 1 366.48 419.04 Tm <746D6120> Tj 1 0 0 1 385.44 419.04 Tm <6920> Tj 1 0 0 1 392.16 419.04 Tm <5471616620> Tj 1 0 0 1 416.88 419.04 Tm <7420> Tj 1 0 0 1 423.12 419.04 Tm <7420> Tj 1 0 0 1 429.84 419.04 Tm <616E4672> Tj 1.20 Tc 1 0 0 1 140.16 415.92 Tm <753667367220> Tj 1 0 0 1 172.80 415.92 Tm <66676E756E757572756620> Tj 1 0 0 1 235.68 415.92 Tm <6620> Tj 1 0 0 1 241.92 415.92 Tm <61397275676E7220> Tj 1 0 0 1 285.84 415.92 Tm <72757575727575757275757275757467756E7275757575757220> Tj 1 0 0 1 443.28 415.92 Tm <3175> Tj 1.50 Tc /F3 9.0 Tf 1 0 0 1 236.40 407.28 Tm <2E2E2E2E542E2E2E2E2E41> Tj 0.24 Tc 1 0 0 1 140.16 390.96 Tm <5420> Tj 1 0 0 1 146.40 390.96 Tm <5441475420> Tj 1 0 0 1 171.84 390.96 Tm <5420> Tj 1 0 0 1 178.08 390.96 Tm <474720> Tj 1 0 0 1 190.80 390.96 Tm <5420> Tj 1 0 0 1 197.04 390.96 Tm <474741475420> Tj 1 0 0 1 228.48 390.96 Tm <47415420> Tj 1 0 0 1 247.44 390.96 Tm <5420> Tj 1 0 0 1 253.68 390.96 Tm <5420> Tj 1 0 0 1 259.92 390.96 Tm <475420> Tj 1 0 0 1 272.40 390.96 Tm <4320> Tj 1 0 0 1 278.88 390.96 Tm <5420> Tj 1 0 0 1 284.88 390.96 Tm <4720> Tj 1 0 0 1 291.36 390.96 Tm <475420> Tj 1 0 0 1 303.84 390.96 Tm <5441415420> Tj 1 0 0 1 328.80 390.96 Tm <5420> Tj 1 0 0 1 335.28 390.96 Tm <4320> Tj 1 0 0 1 341.52 390.96 Tm <4320> Tj 1 0 0 1 347.76 390.96 Tm <47415441414320> Tj 1 0 0 1 385.44 390.96 Tm <4741414320> Tj 1 0 0 1 410.64 390.96 Tm <474147414320> Tj 1 0 0 1 442.08 390.96 Tm <5420> Tj 1 0 0 1 448.56 390.96 Tm <43> Tj 1.50 Tc 1 0 0 1 222.48 386.40 Tm <2D61> Tj 1.50 Tc /F3 5.0 Tf 1 0 0 1 205.20 382.80 Tm <2E2E2E752E> Tj 0.72 Tc /F3 9.0 Tf 1 0 0 1 140.40 366.00 Tm <54475420> Tj 1 0 0 1 159.36 366.00 Tm <434354474320> Tj 1 0 0 1 191.04 366.00 Tm <5441414154414741475454414143475443415420> Tj 1 0 0 1 310.32 366.00 Tm 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G. ... .GG.C.GGGGGT T TACTTT G NUUU\,\TTAGAGTGCTCAAAGCAGGCTTT T AAAAGCTTAc ...T. ...GA..----..CAGAATATTAGT GCATGGAATAAT GAAATAGGAAGT CGTTT CTATTTTGTTGA.G.. .CC...G......C.GTT - T T CGGNU\T CGACT TAATGATTAATAGGGACAGTCGGGGGCATTAG...c. .CA...GG.. G.A.. ...C.TATT CAGACGACAGAGGTGAAATT CTTGGACCGTCTGAAGACTAACTACT....GC....TT ...T....GC G... _...:::T::iTTT :::T:TT:T TTT:ATTTT :T:T::Y:T Ti:i-^t6--1 n ^^^^n m^n ^^frA id^d7 i6l ^mm^mn n i^n mn n 1 ^-n m^^^nUU f I UUffiUUUUAI UAUA T AUUUUUU f AU I I U I l-fiUUAI ffiUUAT UUUA GCTAGCGATCCGCAGTT GTTTATT T TTACGACTCTGCGGGAAGCTTCCGGAT-Tr-CT-A'Ar:ATT-r:AAr--n n n^^n n t^^mmmm^--frmii---^-n n^mtm^^mm^nl n n^^m^n n n^\JffiUUffi\JU I I I J- U\f\f I J. UU\7\T\J\J\fIifi\J L ft.L \J\J I t \rU}1tl}I\JU I \J'1'tf1\-midTTAAAGGAATTGACGGAAGGGCACCACCAGGAGTG_ - GCCT GCGGCTTAAGA. ..frmm^n ^mn n 1 iA ^^^-1 n n m^mnn ^d1 iiAAdF^^ A1 ifrFi^ n ^^l mm^nJ- I I. U.I{U I \-fift\-ft\-\JLT\Jfl}t.tt'.L U.L UA\-UAL](J\-\-ULfUAUAU -t \J\Jf{fI\J\J}l. I I LjTidm-....1.9...^l nl Tmal an a^m^mmmnnla A n4F a -ma a a4 a aml tma i Tqaf t t anFru6g6r fgnunuuruf f a9rugnr ruuuruuuruuruutgunruuuuur 1u....T.....AT TAGT T GG T GGAGT GAT T T GT C T G GT TAAT T C C GATAAC GAAC GAGAC T C-a...u.TGT CCTGC TAAATAGAGTTAACGTCAT TTTTT T TGCGTTAACT T TT CTTCT CCCC r]A r- aA r-^" ^JUf{\fJ.\fi,U....\J..T TAGAGGGACT GGT GGC GT CTAGC CACACGAGAT T GAGCAATAACA@TC.....A ...T. G.... .-TGTGATGCCCTT r49150195200245246295296344346394396444446494496 544546594596642'o{ooYz696142146 792196842845892895904907 Fig. 2. The determined partial 18S ribosomal RNA gene sequence from Lepeophtheirus salmonis is given and compared to Argulusnobilis (Branchiura: Crustacea). Letters in bold and underlined represent the primers used for PCR amplification. Pinto & Kotzian (Schon, Butlin, Griffiths & Mar-tens, 1998) with a similarity of 47 .94oh (3 l5 nt).DiscussionThe ability to identify different sea lice popula-tions will make a maior contribution to understand- ing characteristics including the dispersion andepidemiology of sea lice infections and contributeinformation to the ongoing dispute between salmonfarming and wild salmonid fishery interests. Tobegin investigating the population characteristicsof L. salmonis, two regions of the sea louse ge-nome, namely l8S rDNA and ITS-1 were targeted.No variation in a partial fragment (904 base pairs) endstream endobj 36 0 obj <> endobj 30 0 obj <>/ProcSet[/PDF/Text/ImageB/ImageC/ImageI]/Subtype 35 0 R/Font<>>>/MediaBox[0 0 595.20 841.68]>> endobj 38 0 obj <>stream % CANON_PFINF_TYPE0_TEXTON endstream endobj 39 0 obj <>stream q 595.20 0 0 841.68 0.00 0.00 cm /Obj2 Do Q endstream endobj 40 0 obj <>stream BT 3 Tr 0.48 Tc /F3 9.0 Tf 1 0 0 1 59.52 797.28 Tm <393420> Tj 1 0 0 1 85.68 797.28 Tm <412E502E20> Tj 1 0 0 1 107.04 797.28 Tm <5368696E6E20> Tj 1 0 0 1 134.64 797.04 Tm <657420> Tj 1 0 0 1 144.96 797.04 Tm <616C2E20> Tj 1 0 0 1 158.64 799.44 Tm <2D20> Tj 1 0 0 1 169.44 797.04 Tm <6C425320> Tj 1 0 0 1 186.00 797.04 Tm <72444E4120> Tj 1 0 0 1 214.08 796.80 Tm <616E6420> Tj 1 0 0 1 232.56 796.80 Tm <49545320> Tj 1 0 0 1 250.32 796.56 Tm <73657175656E63657320> Tj 1 0 0 1 296.64 796.32 Tm 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<726D20> Tj 1 0 0 1 96.48 333.36 Tm <6E2D> Tj 0.48 Tc /F3 7.0 Tf 1 0 0 1 157.20 773.52 Tm <545454434347544147475447414143435447434747414147474154434154544141544743434141545441544143544143544147434754434354432D20> Tj 1 0 0 1 467.28 774.96 Tm <2D20> Tj 1 0 0 1 492.00 772.56 Tm <3520> Tj 1 0 0 1 497.28 772.56 Tm <38> Tj 0.48 Tc 1 0 0 1 157.20 763.44 Tm <545454434347544147475447414143435447434747414147474154434154544141544743434141545441544143544143544147434754434354432D20> Tj 1 0 0 1 467.28 764.88 Tm <2D20> Tj 1 0 0 1 492.00 762.48 Tm <3520> Tj 1 0 0 1 497.28 762.48 Tm <42> Tj 0.48 Tc 1 0 0 1 157.20 753.60 Tm <545454434347494147475447414143435447434747414147474154434154544141544743434141545441544143544143544147434754434354435F20> Tj 1 0 0 1 467.28 754.80 Tm <2D20> Tj 1 0 0 1 492.00 752.40 Tm <3520> Tj 1 0 0 1 497.28 752.40 Tm <42> Tj 0.48 Tc 1 0 0 1 157.20 743.28 Tm <545454434347544147475447414143435447434747414147474154434154544141544743434141545441544143544143544147434754434354432D20> Tj 1 0 0 1 467.28 744.48 Tm <5F20> Tj 1 0 0 1 492.00 742.08 Tm <3520> Tj 1 0 0 1 497.28 742.08 Tm <42> Tj 0.48 Tc 1 0 0 1 157.20 732.96 Tm <545454434347544147475447414143435447434747414147474154434154544141544743434141545441544143544143544147434754434354432D20> Tj 1 0 0 1 467.28 734.40 Tm <2D20> Tj 1 0 0 1 492.00 731.76 Tm <3520> Tj 1 0 0 1 497.28 731.76 Tm <42> Tj 0.48 Tc 1 0 0 1 157.20 722.88 Tm <54545443434754414747544741414343544743474741414747415443415454414754474343414154544154544354414354414743475443475454544320> Tj 1 0 0 1 492.24 721.68 Tm <3620> Tj 1 0 0 1 497.28 721.68 Tm <4F> Tj 0.48 Tc 1 0 0 1 157.20 712.80 Tm <545454434347544147475447414143435447434747414147474154434154544141544743434141545441544143544143544147434754434354432D20> Tj 1 0 0 1 467.28 714.00 Tm <2D20> Tj 1 0 0 1 492.00 711.36 Tm <3520> Tj 1 0 0 1 497.28 711.36 Tm <42> Tj 0.48 Tc 1 0 0 1 157.20 702.72 Tm <54545443434754414747544741414343544743474741414747415443415454414754474343414154544154544354414354414743475443475454544320> Tj 1 0 0 1 492.24 701.52 Tm <3620> Tj 1 0 0 1 497.28 701.52 Tm <4F> Tj 0.48 Tc 1 0 0 1 157.20 692.64 Tm <545454434347544147475447414143435447434747414147474154434154544141544743434141545441544143544143544147434754434354432D20> Tj 1 0 0 1 467.28 693.60 Tm <5F20> Tj 1 0 0 1 492.00 691.20 Tm <3520> Tj 1 0 0 1 497.28 691.20 Tm <42> Tj 0.48 Tc 1 0 0 1 157.20 682.56 Tm <54545443434754414747544741414343544743474741414747415443415454414754474343414154544154544354414354414743475443475454544320> Tj 1 0 0 1 492.24 681.36 Tm <3620> Tj 1 0 0 1 497.28 681.36 Tm <4F> Tj 0.48 Tc 1 0 0 1 157.20 672.48 Tm <54545443434754414747544741414343544743474741414747415443415454414754474343414154544154544354414354414743475443475454544320> Tj 1 0 0 1 492.48 671.28 Tm <3620> Tj 1 0 0 1 497.28 671.28 Tm <4F> Tj 0.48 Tc 1 0 0 1 157.20 662.40 Tm <54545443434754414747544741414343544743474741414747415443415454414754474343414154544154544354414354414743475443475454544320> Tj 1 0 0 1 492.24 661.20 Tm <3620> Tj 1 0 0 1 497.28 661.20 Tm <4F> Tj 0.48 Tc 1 0 0 1 157.20 652.56 Tm <54545443434754414747544741414343544743474741414747415443415454414754474343414154544154544354414354414743475443475454544320> Tj 1 0 0 1 492.24 651.36 Tm <3620> Tj 1 0 0 1 497.28 651.36 Tm <4F> Tj 1.50 Tc /F3 4.0 Tf 1 0 0 1 159.60 644.64 Tm <2A20> Tj 1 0 0 1 164.88 644.64 Tm <2A20> Tj 1 0 0 1 170.16 644.64 Tm <2A2A20> Tj 1 0 0 1 180.48 644.64 Tm <2A2A2A2A20> Tj 1 0 0 1 201.60 644.40 Tm <2A20> Tj 1 0 0 1 206.88 644.40 Tm <2A20> Tj 1 0 0 1 212.16 644.40 Tm <2A20> Tj 1 0 0 1 217.44 644.40 Tm <2A20> Tj 1 0 0 1 222.72 644.40 Tm <2A20> Tj 1 0 0 1 227.76 644.40 Tm <2A20> Tj 1 0 0 1 233.04 644.40 Tm <2A20> Tj 1 0 0 1 238.32 644.40 Tm <2A20> Tj 1 0 0 1 243.60 644.16 Tm <2A20> Tj 1 0 0 1 248.88 644.40 Tm <2A2A20> Tj 1 0 0 1 259.20 644.40 Tm <2A20> Tj 1 0 0 1 264.48 644.16 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0.48 Tc /F3 8.0 Tf 1 0 0 1 157.44 614.64 Tm <2D20> Tj 1 0 0 1 162.96 614.64 Tm <2D2D2D4143474747434743544754414141435454544E55555555555555555555555C695C4741275C275C275C5454545443414143474754474741544154435420> Tj 1 0 0 1 492.24 611.28 Tm <312E4934> Tj 0.48 Tc 1 0 0 1 157.68 604.56 Tm <2D20> Tj 1 0 0 1 162.96 604.56 Tm <2D20> Tj 1 0 0 1 168.24 604.56 Tm <2D20> Tj 1 0 0 1 173.52 604.56 Tm <2D4143474747434743544754414141435454544169555C695C69555C69555C4E5C695555555C4741695C275C275C5454545443414143474754474741544154435420> Tj 1 0 0 1 492.24 601.20 Tm <5C4934> Tj 0.48 Tc 1 0 0 1 157.68 594.48 Tm <2D20> Tj 1 0 0 1 162.96 594.48 Tm <2D20> Tj 1 0 0 1 168.24 594.48 Tm <2D20> Tj 1 0 0 1 173.52 594.48 Tm <2D41434747474347435447544141414354545420> Tj 1 0 0 1 273.12 591.84 Tm <412C695555555C20> Tj 1 0 0 1 299.28 591.60 Tm <414120> Tj 1 0 0 1 309.84 591.60 Tm <344A5C2C555C6955555C47414141415454545443414143474754474741544154435420> Tj 1 0 0 1 492.24 591.12 Tm <313720> Tj 1 0 0 1 502.80 591.12 Tm <34> 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<2D2D2D4143474747434743544754414141435454544E55555555555C414A555C2D2D2D2D47414141415454545443414143474754474741544154435420> Tj 1 0 0 1 492.24 551.28 Tm <3120> Tj 1 0 0 1 497.52 551.28 Tm <3120> Tj 1 0 0 1 502.80 551.28 Tm <4F> Tj 0.48 Tc 1 0 0 1 156.96 542.16 Tm <4143472D414347472D434743544754414141435454544E555C695C434E555C695555554120> Tj 1 0 0 1 336.00 541.44 Tm <4E5C47414141415454545443414143474754474741544154435420> Tj 1 0 0 1 492.48 541.20 Tm <3120> Tj 1 0 0 1 497.52 541.20 Tm <3120> Tj 1 0 0 1 502.80 540.96 Tm <42> Tj 0.48 Tc 1 0 0 1 157.68 534.48 Tm <2D2D2D2D4143474747434743544754414141435454544E5C69555555555C275C2C555C2D2D2D2D474E5C69555C5454545443414143474754474741544154435420> Tj 1 0 0 1 492.48 531.36 Tm <3120> Tj 1 0 0 1 497.52 531.36 Tm <3120> Tj 1 0 0 1 502.80 531.36 Tm <4F> Tj 0.48 Tc 1 0 0 1 156.96 522.24 Tm <41434747414347472D20> Tj 1 0 0 1 204.96 521.76 Tm <43474354475420> Tj 1 0 0 1 236.64 521.76 Tm <4E555C4354545420> Tj 1 0 0 1 273.36 521.76 Tm <4E55555C43412C55555555554120> Tj 1 0 0 1 336.00 521.52 Tm <4D47667555555C5454545443414143474754474741544154435420> Tj 1 0 0 1 492.48 521.28 Tm <3120> Tj 1 0 0 1 497.52 521.28 Tm <3120> Tj 1 0 0 1 503.04 521.04 Tm <39> Tj 0.48 Tc 1 0 0 1 157.20 512.16 Tm <41434747414347474743474354475441414143545454414141414341414141414141414147414141415454545443414143474754474741544154435420> Tj 1 0 0 1 492.24 511.20 Tm <313220> Tj 1 0 0 1 502.80 510.96 Tm <4F> Tj 0.48 Tc 1 0 0 1 156.96 502.08 Tm <4143472D414347472D4347435447544141414354545420> Tj 1 0 0 1 273.36 501.84 Tm <4E5C3A5C4A5C43414A55555C6A555555555C47414141415454545443414143474754474741544154435420> Tj 1 0 0 1 492.48 501.12 Tm <3120> Tj 1 0 0 1 497.52 501.12 Tm <3120> Tj 1 0 0 1 502.80 501.12 Tm <38> Tj 0.48 Tc 1 0 0 1 156.96 492.00 Tm <4143472D414347472D4347435447544141414354545420> Tj 1 0 0 1 273.36 491.52 Tm <4E55555C434E555555555555555C47414141415454545443414143474754474741544154435420> Tj 1 0 0 1 492.24 491.04 Tm <3120> Tj 1 0 0 1 497.76 491.04 Tm <3120> Tj 1 0 0 1 502.80 490.80 Tm <42> Tj 0.48 Tc /F3 7.0 Tf 1 0 0 1 156.96 451.68 Tm <41474754544354434754414343474147474141474141434743414743474141435447434741544141474341415447434741415454474341474141434720> Tj 1 0 0 1 492.48 450.72 Tm <313120> Tj 1 0 0 1 502.80 450.72 Tm <34> Tj 0.48 Tc 1 0 0 1 156.96 441.60 Tm <41474754544354434754414343474147474141474141434743414743474141435447434741544141474341415447434741415454474341474141434720> Tj 1 0 0 1 492.48 440.64 Tm <313120> Tj 1 0 0 1 502.80 440.40 Tm <34> Tj 0.48 Tc 1 0 0 1 157.20 431.52 Tm <41474754544354434754414343474147474141474141434743414743474141435447434741544141474341415447434741415454474341474141434720> Tj 1 0 0 1 492.48 430.56 Tm <493120> Tj 1 0 0 1 502.80 430.56 Tm <34> Tj 0.48 Tc 1 0 0 1 156.96 421.68 Tm <41474754544354434754414343474147474141474141434743414743474141435447434741544141474341415447434741415454474341474141434720> Tj 1 0 0 1 492.48 420.72 Tm <37273120> Tj 1 0 0 1 502.80 420.72 Tm <34> Tj 0.48 Tc 1 0 0 1 157.20 411.84 Tm <41474754544354434754414343474147474141474141434743414743474141435447434741544141474341415447434741415454474341474141434720> Tj 1 0 0 1 492.48 410.88 Tm <31223120> Tj 1 0 0 1 502.80 410.88 Tm <34> Tj 0.48 Tc 1 0 0 1 156.96 402.00 Tm <41474754544354434754414343474147474141474141434743414743474141435447434741544141474341415447434741415454474341474141434720> Tj 1 0 0 1 492.48 401.28 Tm <3120> Tj 1 0 0 1 497.52 401.28 Tm <3720> Tj 1 0 0 1 502.80 401.04 Tm <38> Tj 0.48 Tc 1 0 0 1 156.96 391.92 Tm <41474754544354434754414343474147474141474141434743414743474141435447434741544141474341415447434741415454474341474141434720> Tj 1 0 0 1 492.48 391.20 Tm <3120> Tj 1 0 0 1 497.52 391.20 Tm <3720> Tj 1 0 0 1 502.80 391.20 Tm <4F> Tj 0.48 Tc 1 0 0 1 156.96 382.08 Tm <41474754544354434754414343474147474141474141434743414743474141435447434741544141474341415447434741415454474341474141434720> Tj 1 0 0 1 492.48 381.12 Tm <3120> Tj 1 0 0 1 497.52 381.36 Tm <3720> Tj 1 0 0 1 502.80 381.12 Tm <42> Tj 0.48 Tc 1 0 0 1 156.96 372.00 Tm <41474754544354434754414343474147474141474141434743414743474141435447434741544141474341415447434741415454474341474141434720> Tj 1 0 0 1 492.48 371.28 Tm <3120> Tj 1 0 0 1 497.52 371.28 Tm <3720> Tj 1 0 0 1 502.80 371.04 Tm <4F> Tj 0.48 Tc 1 0 0 1 156.96 361.92 Tm <414747545420> Tj 1 0 0 1 184.08 361.68 Tm <4354434754414343474147474141474141434743414743474141435447434741544141474341415447434741415454474341474141434720> Tj 1 0 0 1 492.72 361.20 Tm <312D3120> Tj 1 0 0 1 503.04 360.96 Tm <39> Tj 0.48 Tc 1 0 0 1 157.20 352.08 Tm <41474754544354434754414343474147474141474141434743414743474141435447434741544141474341415447434741415454474341474141434720> Tj 1 0 0 1 492.48 351.12 Tm <3120> Tj 1 0 0 1 497.76 351.12 Tm <4220> Tj 1 0 0 1 502.80 351.12 Tm <4F> Tj 0.48 Tc 1 0 0 1 157.20 342.00 Tm <41474754544354434754414343474147474141474141434743414743474141435447434741544141474341415447434741415454474341474141434720> Tj 1 0 0 1 492.48 341.04 Tm <3120> Tj 1 0 0 1 497.76 341.28 Tm <3F20> Tj 1 0 0 1 503.04 341.04 Tm <42> Tj 0.48 Tc 1 0 0 1 157.20 331.92 Tm <41474754544354434754414343474147474141474141434743414743474141435447434741544141474341415447434741415454474341474141434720> Tj 1 0 0 1 492.48 331.20 Tm <3120> Tj 1 0 0 1 497.76 331.20 Tm <3720> Tj 1 0 0 1 503.04 330.96 Tm <42> Tj 0.48 Tc /F3 9.0 Tf 1 0 0 1 301.20 266.16 Tm <657665722C20> Tj 1 0 0 1 324.24 266.16 Tm <7765726520> Tj 1 0 0 1 347.04 265.92 Tm <7661726961626C6520> Tj 1 0 0 1 383.52 265.92 Tm <6265747765656E20> Tj 1 0 0 1 421.68 265.68 Tm <736974657320> Tj 1 0 0 1 442.56 265.68 Tm <7769746820> Tj 1 0 0 1 463.92 265.92 Tm <6772656174657220> Tj 1 0 0 1 495.84 265.68 Tm <736C6D6C2D> Tj 0.48 Tc 1 0 0 1 301.20 253.92 Tm <6C617269747920> Tj 1 0 0 1 327.36 253.68 Tm <64656D6F6E7374726174656420> Tj 1 0 0 1 388.80 253.44 Tm <6265747765656E20> Tj 1 0 0 1 428.40 253.44 Tm <696E646976696475616C20> Tj 1 0 0 1 475.68 253.20 Tm <6C69636520> Tj 1 0 0 1 494.40 253.20 Tm <74616B656E> Tj 0.48 Tc 1 0 0 1 301.20 241.44 Tm <66726F6D20> Tj 1 0 0 1 324.48 241.44 Tm <6661726D656420> Tj 1 0 0 1 357.36 241.20 Tm <73616C6D6F6E69647320> Tj 1 0 0 1 403.20 241.20 Tm <7468616E20> Tj 1 0 0 1 424.32 240.96 Tm <74686F736520> Tj 1 0 0 1 449.52 240.96 Tm <74616B656E20> Tj 1 0 0 1 475.44 240.96 Tm <66726F6D20> Tj 1 0 0 1 498.48 240.72 Tm <77696C64> Tj 0.72 Tc 1 0 0 1 301.20 228.72 Tm <73616C6D6F6E6964732E20> Tj 1 0 0 1 349.44 228.72 Tm <566172696174696F6E20> Tj 1 0 0 1 392.40 228.48 Tm <77697468696E20> Tj 1 0 0 1 422.16 228.48 Tm <74686520> Tj 1 0 0 1 437.76 228.48 Tm <4954532D6C20> Tj 1 0 0 1 464.88 228.48 Tm <726567696F6E20> Tj 1 0 0 1 494.88 228.24 Tm <697320> Tj 1 0 0 1 504.00 228.24 Tm <6E6F74> Tj 0.48 Tc 1 0 0 1 300.72 216.48 Tm <756E636F6D6D6F6E20> Tj 1 0 0 1 353.76 216.24 Tm <616E6420> Tj 1 0 0 1 373.68 216.24 Tm <68617320> Tj 1 0 0 1 392.40 216.24 Tm <6265656E20> Tj 1 0 0 1 416.88 216.24 Tm <7265706F7274656420> Tj 1 0 0 1 457.68 216.00 Tm <66726F6D20> Tj 1 0 0 1 482.64 216.00 Tm <5363686973746F2D> Tj 0.48 Tc 1 0 0 1 300.72 204.00 Tm <736F6D61776865726520> Tj 1 0 0 1 352.56 204.00 Tm <626F746820> Tj 1 0 0 1 373.92 203.76 Tm <73697A6520> Tj 1 0 0 1 391.92 203.76 Tm <616E6420> Tj 1 0 0 1 409.44 203.76 Tm <73657175656E636520> Tj 1 0 0 1 449.76 203.76 Tm <766172696174696F6E20> Tj 1 0 0 1 489.12 203.52 Tm <616D6F6E67> Tj 0.72 Tc 1 0 0 1 300.96 191.76 Tm <636F7069657320> Tj 1 0 0 1 332.40 191.52 Tm <6F6620> Tj 1 0 0 1 344.64 191.52 Tm <74686520> Tj 1 0 0 1 361.68 191.52 Tm <4954532D6C20> Tj 1 0 0 1 390.48 191.52 Tm <72657065617420> Tj 1 0 0 1 420.72 191.28 Tm <77697468696E20> Tj 1 0 0 1 452.16 191.28 Tm <696E646976696475616C7320> Tj 1 0 0 1 504.00 191.04 Tm <6F632D> Tj 0.72 Tc 1 0 0 1 300.96 179.28 Tm <6375727320> Tj 1 0 0 1 324.96 179.52 Tm <2856616E20> Tj 1 0 0 1 352.32 179.04 Tm <48657277657264656E20> Tj 1 0 0 1 406.80 178.80 Tm <657420> Tj 1 0 0 1 420.24 178.80 Tm <616C2E2C20> Tj 1 0 0 1 440.64 179.04 Tm <31393938292E20> Tj 1 0 0 1 474.00 178.56 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Shinn et al. - lBS rDNA and ITS sequences as population markers for Lepeophtheirus salmonis F: rm A,F: rm A^E'^rm A-I'^ rm A,wild 81wild 82Iili I d f -v{ild c2F:rm Fl,F^rm n-F^rm n-F-: rm Il.E-rrm Fl-E-^ rm A^F: rm A,vl]ild B1hli I rl R^wild c1wild c2I-: rm n.E:rm Fl^Errm D-F:rm O-E:rm Fl-E': rm A.E-:rm A-F-erm A,I^li I /.l P-wild 82!\1il-d c1wifd c2F ^ rm n.F: rm n^F:rfr f)-!-: rm f),E': rm n- TTTCCGTAGGTGAACCTGCGGAAGGATCATTAATGCCAATTATACTACTAGCGTCCTC- - 5 8TTTCCGTAGGTGAACCTGCGGAAGGATCATTAATGCCAATTATACTACTAGCGTCCTC- - 5 BTTTCCGIAGGTGAACCTGCGGAAGGATCATTAATGCCAATTATACTACTAGCGTCCTC_ - 5 BTTTCCGTAGGTGAACCTGCGGAAGGATCATTAATGCCAATTATACTACTAGCGTCCTC- _ 5 BTTTCCGTAGGTGAACCTGCGGAAGGATCATTAATGCCAATTATACTACTAGCGTCCTC- - 5 BTTTCCGTAGGTGAACCTGCGGAAGGATCATTAGTGCCAATTATTCTACTAGCGTCGTTTC 6 OTTTCCGTAGGTGAACCTGCGGAAGGATCATTAATGCCAATTATACTACTAGCGTCCTC- - 5 BTTTCCGTAGGTGAACCTGCGGAAGGATCATTAGTGCCAATTATTCTACTAGCGTCGTTTC 6 OTTTCCGTAGGTGAACCTGCGGAAGGATCATTAATGCCAATTATACTACTAGCGTCCTC- _ 5 BTTTCCGTAGGTGAACCTGCGGAAGGATCATTAGTGCCAATTATTCTACTAGCGTCGTTTC 6 OTTTCCGTAGGTGAACCTGCGGAAGGATCATTAGTGCCAATTATTCTACTAGCGTCGTTTC 6 OTTTCCGTAGGTGAACCTGCGGAAGGATCATTAGTGCCAATTATTCTACTAGCGTCGTTTC 6 OTTTCCGTAGGTGAACCTGCGGAAGGATCATTAGTGCCAATTATTCTACTAGCGTCGTTTC 6 O* * ** **** * * * * * * * * * ** * * * ** ** * ** ** * ** * * * * * * * * * * * * * * * * * * x -- ---ACGGGCGCTGTAAACTTTNUUUUUUUUUUU\i\GA'\'\'\TTTTCAACGGTGGATATCT 1.I4- - - -ACGGGCGCTGTAAACTTTAiU\i\iU\iU\N\iUUU\GAi\'\'\TTTTCAACGGTGGATATCT \I4- - - -ACGGGCGCTGTAAACTTT A,iUUU\ AA 4J\,U\iUU\GAAAATTTTCAACGGTGGATATCT 17 4----ACGGGCGCTGTAAACTTTAJUU\'\'\'\'UUUUUU\GAAAATTTTCAACGGTGGATATCT 114- - - _ACGGGCGCTGTAAACTTTNUUUUUA NUU\iU\'\GAAAATTTTCAACGGTGGATATCT 1 1 4ACG-ACGG-CGCTGTAAACTTTAiU\ACA'U\iUUUUU\GAiUU\TTTTCAACGGTGGATATCT 1 1 8- ---ACGGGCGCTGTAAACTTTNUUUUU\AJU\----GAAAATTTTCAACGGTGGATATCT 1 1 OACG-ACGG-CGCTGTAAACTTTNU\i\CNU\iUUUA N\GAAAATTTTCAACGGTGGATATCT 1 1 B----ACGGGCGCTGTAAACTTTN\iUUUU\'\,U\----GN\iU\TTTTCAACGGTGGATATCT 1 1 OACGGACGG- CGCTGT NU\CTTT NUU\CA,UUUUUA MGfuUU\TTTTCAACGGTGGATATCT 1 1 9ACGGACGGGCGCTGTAAACTTTAAAACAAAAAAAAAGAAAATTTTCAACGGTGGATATCT 12 OACG-ACGG-CGCTGTAAACTTT N\:\J\CAJUU\jUUUU\GAAAATTTTCAACGGTGGATATCT 1 1 8ACG-ACGG-CGCTGTAAACTTT NUU\CNUUUUUUU\GAAAATTTTCAACGGTGGATATCT 1 1 B AGGTTCTCGTACCGAGGAAGAACGCAGCGAACTGCGATAAGCAATGCGAATTGCAGAACG 11 4AGGTTCTCGTACCGAGGAAGAACGCAGCGAACTGCGATAAGCAATGCGAATTGCAGAACG 11 4AGGTTCTCGTACCGAGGAAGAACGCAGCGAACTGCGATAAGCAATGCGAATTGCAGAACG I1 4AGGTTCTCGTACCGAGGAAGAACGCAGCGAACTGCGATAAGCAATGCGAATTGCAGAACG 7'1 4AGGTTCTCGTACCGAGGAAGAACGCAGCGAACTGCGATAAGCAATGCGAATTGCAGAACG 1"1 4AGGTTCTCGTACCGAGGAAGAACGCAGCGAACTGCGATAAGCAATGCGAATTGCAGAACG 1 7 8AGGTTCTCGTACCGAGGAAGAACGCAGCGAACTGCGATAAGCAATGCGAATTGCAGAACG 1 7 OAGGTTCTCGTACCGAGGAAGAACGCAGCGAACTGCGATAAGCAATGCGAATTGCAGAACG 1 7 BAGGTTCTCGTACCGAGGAAGAACGCAGCGAACTGCGATAAGCAATGCGAATTGCAGAACG 1 7 OAGGTT CTCGTACCGAGGAAGAACGCAGCGAACTGCGATAAGCAATGCGAATTGCAGAACG 1-1 9AGGTTCTCGTACCGAGGAAGAACGCAGCGAACTGCGATAAGCAATGCGAATTGCAGAACG 1 B OAGGTTCTCGTACCGAGGAAGAACGCAGCGAACTGCGATAAGCAATGCGAATTGCAGAACG 1 ? BAGGTTCTCGTACCGAGGAAGAACGCAGCGAACTGCGATAAGCAATGCGAATTGCAGAACG 1 7 B ever, were variable between sites with greater slml-larity demonstrated between individual lice takenfrom farmed salmonids than those taken from wildsalmonids. Variation within the ITS-l region is notuncommon and has been reported from Schisto-somawhere both size and sequence variation amongcopies of the ITS-l repeat within individuals oc-curs (Van Herwerden et al., 1998). Similarly, Fig. 3. The determined 454 nucleotide sequence for the internal transcribed spacer (lTS I ) from different populations of Lepeophtheirussalmonis. Letters in bold and underlined represent the primers used for PCR amplification. Stars represent areas of identity between of the l8S rRNA gene was found between samplesoflice collected from four different sites. 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70.32 574.32 Tm <522E> Tj 0.96 Tc /F3 7.0 Tf 1 0 0 1 70.56 542.88 Tm <4431> Tj 0.96 Tc 1 0 0 1 70.56 532.80 Tm <4432> Tj 0.96 Tc 1 0 0 1 70.56 524.64 Tm <6E5E> Tj 0.96 Tc 1 0 0 1 70.56 512.88 Tm <4461> Tj 1.20 Tc 1 0 0 1 70.56 504.24 Tm <6E2D> Tj 0.48 Tc /F3 8.0 Tf 1 0 0 1 131.76 774.24 Tm <43474147544341545447474154545454434741414347434141435447474347475443544741434743414143414154434147414343415447544754475420> Tj 1 0 0 1 466.08 773.28 Tm <323334> Tj 0.48 Tc 1 0 0 1 131.76 764.16 Tm <43474147544341545447474154545454434741414347434141435447474347475443544741434743414143414154434147414343415447544754475420> Tj 1 0 0 1 466.32 763.20 Tm <323334> Tj 0.48 Tc 1 0 0 1 131.76 753.84 Tm <43474147544341545447474154545454434741414347434141435447474347475443544741434743414143414154434147414343415447544754475420> Tj 1 0 0 1 465.84 752.88 Tm <323334> Tj 0.48 Tc 1 0 0 1 131.76 743.76 Tm <43474147544341545447474154545454434741414347434141435447474347475443544741434743414143414154434147414343415447544754475420> Tj 1 0 0 1 465.84 742.80 Tm <323334> Tj 0.48 Tc 1 0 0 1 131.76 733.68 Tm <43474147544341545447474154545454434741414347434141435447474347475443544741434743414143414154434147414343415447544754475420> Tj 1 0 0 1 466.32 732.72 Tm <323334> Tj 0.72 Tc 1 0 0 1 131.76 723.60 Tm <43474147544341545454474154545454434741414347434154544347474347475443434741434754414743414154434747414343415447544754475420> Tj 1 0 0 1 466.56 722.64 Tm <323338> Tj 0.48 Tc 1 0 0 1 131.76 713.52 Tm <43474147544341545447474154545454434741414347434141435447474347475443544741434743414143414154434147414343415447544754475420> Tj 1 0 0 1 466.56 712.32 Tm <323330> Tj 0.72 Tc 1 0 0 1 131.76 703.20 Tm <43474147544341545454474154545454434741414347434154544347474347475443434741434754414743414154434747414343415447544754475420> Tj 1 0 0 1 466.56 702.00 Tm <323338> Tj 0.48 Tc 1 0 0 1 131.76 693.36 Tm <43474147544341545454474154545454434741414347434141434347474347475443434741434754414743414154434747414343415447544754475420> Tj 1 0 0 1 466.56 692.16 Tm <323330> Tj 0.72 Tc 1 0 0 1 131.76 683.28 Tm <43474147544341545454474154545454434741414347434154544347474347475443434741434754414743414154434747414343415447544754475420> Tj 1 0 0 1 466.08 682.32 Tm <323339> Tj 0.72 Tc 1 0 0 1 131.76 673.44 Tm <43474147544341545454474154545454434741414347434154544347474347475443434741434754414743414154434747414343415447544754475420> Tj 1 0 0 1 466.08 672.24 Tm <323430> Tj 0.72 Tc 1 0 0 1 131.76 663.36 Tm <43474147544341545454474154545454434741414347434154544347474347475443434741434754414743414154434747414343415447544754475420> Tj 1 0 0 1 466.56 662.40 Tm <323338> Tj 0.48 Tc 1 0 0 1 131.76 653.28 Tm <43474147544341545454474154545454434741414347434154544347474347475443434741434754414743414154434747414343415447544754475420> Tj 1 0 0 1 466.56 652.32 Tm <323338> Tj 1.50 Tc /F3 10.0 Tf 1 0 0 1 134.16 645.84 Tm <2A2A2A2A2A2A2A2A2A20> Tj 1 0 0 1 186.72 645.60 Tm <2A2A2A2A2A2A2A2A2A2A2A2A2A2A20> Tj 1 0 0 1 275.76 645.36 Tm <2A2A2A2A2A2A2A20> Tj 1 0 0 1 317.52 645.12 Tm <2A2A2A2A20> Tj 1 0 0 1 343.68 645.12 Tm <2A20> Tj 1 0 0 1 354.24 645.12 Tm <2A2A2A2A2A20> Tj 1 0 0 1 385.68 645.12 Tm <2A2A2A2A2A2A2A2A2A2A2A2A> Tj 0.48 Tc /F3 8.0 Tf 1 0 0 1 131.76 623.52 Tm <5420> Tj 1 0 0 1 137.04 623.52 Tm <5443414754475420> Tj 1 0 0 1 174.00 623.28 Tm <4341414320> Tj 1 0 0 1 195.36 623.28 Tm <4E555C69555C5441434341415443545441415420> Tj 1 0 0 1 289.20 622.80 Tm <475447415454474141435420> Tj 1 0 0 1 346.80 622.80 Tm <5420> Tj 1 0 0 1 352.08 622.56 Tm <434147545420> Tj 1 0 0 1 378.24 622.56 Tm <435420> Tj 1 0 0 1 388.80 622.56 Tm <435443415447414141544120> Tj 1 0 0 1 466.32 622.32 Tm <3220> Tj 1 0 0 1 471.84 622.32 Tm <3920> Tj 1 0 0 1 476.88 622.32 Tm <34> Tj 0.48 Tc 1 0 0 1 131.76 613.44 Tm <5420> Tj 1 0 0 1 137.04 613.44 Tm <5443414754475420> Tj 1 0 0 1 174.00 613.20 Tm <43414143414141414154414343414154435454414154475420> Tj 1 0 0 1 299.76 612.72 Tm <47415454474141435420> Tj 1 0 0 1 346.80 612.72 Tm <5420> Tj 1 0 0 1 352.08 612.72 Tm <4341475454435420> Tj 1 0 0 1 388.80 612.72 Tm <435443415447414141544120> Tj 1 0 0 1 466.08 612.48 Tm <3220> Tj 1 0 0 1 471.84 612.24 Tm <3920> Tj 1 0 0 1 476.88 612.24 Tm <34> Tj 0.48 Tc 1 0 0 1 131.76 603.12 Tm <5420> Tj 1 0 0 1 137.28 603.12 Tm <5443414754475443414143414141414154414343414154435454414154475420> Tj 1 0 0 1 299.76 602.64 Tm <474154544741414354544341475454435420> Tj 1 0 0 1 388.80 602.40 Tm <435443415447414141544120> Tj 1 0 0 1 466.32 601.92 Tm <3220> Tj 1 0 0 1 471.84 601.92 Tm <3920> Tj 1 0 0 1 476.88 601.92 Tm <34> Tj 0.48 Tc 1 0 0 1 132.00 593.28 Tm <54544341475447544341414341275555555C5441434341415443545441415447544741545447414143545443414754544354435443415447414141544120> Tj 1 0 0 1 466.32 592.08 Tm <323920> Tj 1 0 0 1 476.88 592.08 Tm <34> Tj 0.48 Tc 1 0 0 1 131.76 583.20 Tm <5420> Tj 1 0 0 1 137.28 583.20 Tm <544341475420> Tj 1 0 0 1 163.68 583.20 Tm <47544341414320> Tj 1 0 0 1 195.36 582.96 Tm <414A5555555C54414320> Tj 1 0 0 1 237.12 582.72 Tm <434141544320> Tj 1 0 0 1 263.28 582.72 Tm <5454414154475420> Tj 1 0 0 1 299.76 582.48 Tm <47415420> Tj 1 0 0 1 315.60 582.48 Tm <5447414143545443414754544354435420> Tj 1 0 0 1 399.36 582.24 Tm <43415420> Tj 1 0 0 1 415.20 582.24 Tm <47414141544120> Tj 1 0 0 1 466.32 582.00 Tm <3220> Tj 1 0 0 1 471.84 582.00 Tm <3920> Tj 1 0 0 1 476.88 582.00 Tm <34> Tj 0.48 Tc 1 0 0 1 131.76 573.12 Tm <545443414754475443414143414141414354434343414154435454414154475447415454474141545454434741545443544354434154474E555C434120> Tj 1 0 0 1 466.32 572.16 Tm <3220> Tj 1 0 0 1 471.84 571.92 Tm <3938> Tj 0.48 Tc 1 0 0 1 132.00 563.28 Tm <54544341475447544341414341414141415441434341415443545441415447544741545447414143545443414754544354435443415447414141544120> Tj 1 0 0 1 466.32 561.84 Tm <323930> Tj 0.48 Tc 1 0 0 1 132.00 552.96 Tm <545443414754475443414143412755555C435443434341415443545441415447544741545447414154545443474154544354435443415447414141434120> Tj 1 0 0 1 466.80 551.76 Tm <323938> Tj 0.48 Tc 1 0 0 1 132.00 542.88 Tm <54544341475447544341414341414141435443434341415443545441415447544741545447414154545443474154544354435443415447414141434120> Tj 1 0 0 1 466.32 541.68 Tm <323930> Tj 0.48 Tc 1 0 0 1 132.00 532.80 Tm <5454434147544754434141434127555C275C435443434341415443545441415447544741545447414154545443474154544354435443415447414141434120> Tj 1 0 0 1 466.32 531.84 Tm <323939> Tj 0.48 Tc 1 0 0 1 132.00 522.72 Tm <54544341475447544341414341414141435443434341415443545441415447544741545447414154545443474154544354435443415447414141434120> Tj 1 0 0 1 466.32 521.52 Tm <3320> Tj 1 0 0 1 471.60 521.52 Tm <4F20> Tj 1 0 0 1 476.88 521.52 Tm <4F> Tj 0.48 Tc 1 0 0 1 132.00 512.88 Tm <54544341475447544341414341275C27555C435443434341415443545441415447544741545447414154545443474154544354435443415447414141434120> Tj 1 0 0 1 466.80 511.68 Tm <323938> Tj 0.48 Tc 1 0 0 1 132.00 502.56 Tm <54544341475447544341414341414141435443434341415443545441415447544741545447414154545443474154544354435443415447414141434120> Tj 1 0 0 1 466.80 501.36 Tm <323938> Tj 0.72 Tc /F3 7.0 Tf 1 0 0 1 131.76 472.56 Tm <414354475447432D5454474347474354434747414354474143474141475441475443414343434754435443544741434747434147414741545454434720> Tj 1 0 0 1 466.32 471.36 Tm <333533> Tj 0.72 Tc 1 0 0 1 131.76 462.48 Tm <414354475447432D5454474347474354434747414354474143474141475441475447414343434754435443544741434747434147544741545454434720> Tj 1 0 0 1 466.56 461.28 Tm <333520> Tj 1 0 0 1 476.88 461.28 Tm <33> Tj 0.72 Tc 1 0 0 1 131.76 452.16 Tm <414354475447432D5454474347474354434747414354474143474141475441475443414343434754435443544741434747434147544741545454434720> Tj 1 0 0 1 466.32 450.96 Tm <333520> Tj 1 0 0 1 476.88 450.96 Tm <33> Tj 0.72 Tc 1 0 0 1 131.76 442.32 Tm <414354475447432D5454474347474354434747414354474143474141475441475443414343434754435443544741434747434147544741545454434720> Tj 1 0 0 1 466.56 441.12 Tm <333520> Tj 1 0 0 1 476.88 441.12 Tm <33> Tj 0.72 Tc 1 0 0 1 131.76 432.48 Tm <41435447544743435454474347474354434747414354474143474141475441475443414343434754435443544741434747434147544741545454434720> Tj 1 0 0 1 466.56 431.28 Tm <333520> Tj 1 0 0 1 477.12 431.28 Tm <34> Tj 0.72 Tc 1 0 0 1 131.76 422.64 Tm <41474147474747435454474347474343545443544354474143474141475441475443414343434743545447434747412D2D20> Tj 1 0 0 1 389.28 423.84 Tm <2D20> Tj 1 0 0 1 394.56 423.84 Tm <2D2D544741545454434720> Tj 1 0 0 1 466.56 421.44 Tm <3320> Tj 1 0 0 1 471.84 421.44 Tm <3533> Tj 0.72 Tc 1 0 0 1 131.76 412.56 Tm <41435447544743435454474347474354434747414354474143474141545441475443414343434754435443544741434747434147544741545454434720> Tj 1 0 0 1 466.56 411.60 Tm <333530> Tj 0.72 Tc 1 0 0 1 131.76 402.72 Tm <41474141474747435454474347474343545443544354474143474141545441475443414343434743545447434747412D2D20> Tj 1 0 0 1 389.28 404.16 Tm <2D2D2D544741545454434720> Tj 1 0 0 1 466.56 401.52 Tm <3320> Tj 1 0 0 1 471.84 401.52 Tm <3533> Tj 0.72 Tc 1 0 0 1 131.76 392.88 Tm <41474147474747435454474347474343545443544354474143474141475441475443414343434754435443544741412D2D2D2D2D544741545454434720> Tj 1 0 0 1 466.56 391.68 Tm <333420> Tj 1 0 0 1 477.12 391.68 Tm <35> Tj 0.72 Tc 1 0 0 1 131.76 383.04 Tm <41474147474747435454474347474343545443544354474143474141475441475443414343434743545447434747412D2D2D2D2D544741545454434720> Tj 1 0 0 1 466.56 381.84 Tm <333534> Tj 0.72 Tc 1 0 0 1 131.76 372.96 Tm <41474147474747435454474347474343545443544354474143474141475441475443414343434743545447434747412D2D2D2D2D544741545454434720> Tj 1 0 0 1 466.56 371.76 Tm <3320> Tj 1 0 0 1 471.84 371.76 Tm <3520> Tj 1 0 0 1 477.12 371.76 Tm <35> Tj 0.72 Tc 1 0 0 1 131.76 363.12 Tm <41474147474747435454474347474343545443544354474143474141475441475443414343434743545447434747412D2D2D2D2D544741545454434720> Tj 1 0 0 1 466.56 361.68 Tm <3320> Tj 1 0 0 1 471.84 361.68 Tm <3520> Tj 1 0 0 1 477.12 361.68 Tm <33> Tj 0.72 Tc 1 0 0 1 131.76 352.80 Tm <41474141474747435454474347474343545443544354474143474141545441475443414343434743545447434747412D2D2D2D20> Tj 1 0 0 1 399.84 354.24 Tm <2D544741545454434720> Tj 1 0 0 1 466.56 351.60 Tm <3320> Tj 1 0 0 1 471.84 351.60 Tm <3520> Tj 1 0 0 1 477.12 351.60 Tm <33> Tj 0.72 Tc 1 0 0 1 44.40 472.56 Tm <4661726D20> Tj 1 0 0 1 69.84 472.56 Tm <4131> Tj 0.72 Tc 1 0 0 1 44.40 462.48 Tm <4661726D20> Tj 1 0 0 1 69.84 462.72 Tm <4132> Tj 0.72 Tc 1 0 0 1 44.40 452.16 Tm <4661726D20> Tj 1 0 0 1 69.84 452.40 Tm <4133> Tj 0.72 Tc 1 0 0 1 44.40 442.32 Tm <4661726D20> Tj 1 0 0 1 69.84 442.32 Tm <4134> Tj 1.20 Tc 1 0 0 1 43.68 432.48 Tm <77696C6420> Tj 1 0 0 1 70.32 432.48 Tm <4269> Tj 1.20 Tc 1 0 0 1 43.68 422.64 Tm <77696C6420> Tj 1 0 0 1 70.32 422.64 Tm <3832> Tj 1.44 Tc /F3 9.0 Tf 1 0 0 1 43.68 412.56 Tm <7769726420> Tj 1 0 0 1 70.32 412.56 Tm <6331> Tj 1.20 Tc 1 0 0 1 43.68 402.72 Tm <77696C6420> Tj 1 0 0 1 70.32 402.48 Tm <6332> Tj 0.96 Tc /F3 7.0 Tf 1 0 0 1 44.64 392.88 Tm <4661726D20> Tj 1 0 0 1 70.80 392.88 Tm <4431> Tj 0.72 Tc 1 0 0 1 44.64 383.04 Tm <4661726D20> Tj 1 0 0 1 70.80 383.04 Tm <4432> Tj 0.72 Tc 1 0 0 1 44.64 372.96 Tm <4661726D20> Tj 1 0 0 1 70.80 372.96 Tm <4433> Tj 0.72 Tc 1 0 0 1 44.64 363.12 Tm <4661726D20> Tj 1 0 0 1 70.80 363.12 Tm <4461> Tj 0.96 Tc 1 0 0 1 44.64 352.80 Tm <4661726D20> Tj 1 0 0 1 70.80 352.80 Tm <4435> Tj 0.72 Tc 1 0 0 1 44.64 322.80 Tm <4661726D20> Tj 1 0 0 1 70.08 322.80 Tm <4131> Tj 0.72 Tc 1 0 0 1 44.64 312.72 Tm <4661726D20> Tj 1 0 0 1 70.08 312.72 Tm <4132> Tj 0.72 Tc 1 0 0 1 44.64 302.40 Tm <4661726D20> Tj 1 0 0 1 70.08 302.64 Tm <4133> Tj 0.72 Tc 1 0 0 1 44.64 292.56 Tm <4661726D20> Tj 1 0 0 1 70.08 292.56 Tm <4161> Tj 1.20 Tc 1 0 0 1 43.92 282.48 Tm <77696C6420> Tj 1 0 0 1 70.56 282.48 Tm <3831> Tj 1.20 Tc 1 0 0 1 43.92 272.64 Tm <7769666420> Tj 1 0 0 1 70.56 272.64 Tm <3832> Tj 0.72 Tc 1 0 0 1 43.92 263.76 Tm <492F2E5D696C6420> Tj 1 0 0 1 70.56 262.56 Tm <6331> Tj 1.20 Tc 1 0 0 1 43.92 252.48 Tm <77696C6420> Tj 1 0 0 1 70.56 252.24 Tm <6332> Tj 0.96 Tc 1 0 0 1 44.64 242.40 Tm <4661726D20> Tj 1 0 0 1 70.80 242.40 Tm <4431> Tj 0.72 Tc 1 0 0 1 44.64 232.56 Tm <4661726D20> Tj 1 0 0 1 70.80 232.56 Tm <4432> Tj 0.72 Tc 1 0 0 1 44.64 222.48 Tm <4661726D20> Tj 1 0 0 1 70.80 222.48 Tm <4433> Tj 0.72 Tc 1 0 0 1 44.64 212.64 Tm <4661726D20> Tj 1 0 0 1 70.80 212.64 Tm <4461> Tj 0.96 Tc 1 0 0 1 44.64 202.56 Tm <4661726D20> Tj 1 0 0 1 70.80 202.56 Tm <4435> Tj 0.48 Tc 1 0 0 1 33.12 175.68 Tm <4669672E20> Tj 1 0 0 1 49.92 175.68 Tm <332E20> Tj 1 0 0 1 58.80 175.68 Tm <436F6E74696E7565642E> Tj 1.50 Tc /F3 10.0 Tf 1 0 0 1 134.88 345.36 Tm <2A20> Tj 1 0 0 1 160.80 345.12 Tm <2A20> Tj 1 0 0 1 176.64 345.12 Tm <2A2A2A2A2A2A2A> Tj 1.50 Tc 1 0 0 1 192.48 194.88 Tm <2A2A2A2A2A20> Tj 1 0 0 1 223.68 194.64 Tm <2A20> Tj 1 0 0 1 234.24 194.64 Tm <2A20> Tj 1 0 0 1 244.80 194.64 Tm <2A20> Tj 1 0 0 1 255.12 194.64 Tm <2A2A20> Tj 1 0 0 1 276.24 194.64 Tm <2A2A2A2A20> Tj 1 0 0 1 302.16 194.40 Tm <2A2A2A2A2A20> Tj 1 0 0 1 349.44 194.40 Tm <2A2A20> Tj 1 0 0 1 365.28 194.40 Tm <2A20> Tj 1 0 0 1 375.60 194.40 Tm <2A2A2A2A2A2A2A20> Tj 1 0 0 1 417.60 194.40 Tm <2A20> Tj 1 0 0 1 428.16 194.40 Tm <2A2A2A2A> Tj 0.48 Tc /F3 7.0 Tf 1 0 0 1 132.00 322.80 Tm <434347474141414741475454474147414147475420> Tj 1 0 0 1 237.36 322.32 Tm <5443414741474141414143435443545454544741474154434141474143435447414141544147474320> Tj 1 0 0 1 466.80 321.60 Tm <3420> Tj 1 0 0 1 471.84 321.60 Tm <3120> Tj 1 0 0 1 477.12 321.60 Tm <33> Tj 0.48 Tc 1 0 0 1 132.00 312.72 Tm <43434747414141474147545447414741414747545443414741474141414143435443545454544741474154434141474143435447414141544147474320> Tj 1 0 0 1 466.56 311.52 Tm <3420> Tj 1 0 0 1 471.84 311.52 Tm <3120> Tj 1 0 0 1 477.12 311.52 Tm <33> Tj 0.48 Tc 1 0 0 1 132.24 302.40 Tm <43434747414141474147545447414741414747545443414741474141414143435443545454544741474154434141474143435447414141544147474320> Tj 1 0 0 1 466.80 301.44 Tm <3420> Tj 1 0 0 1 472.08 301.44 Tm <3120> Tj 1 0 0 1 477.12 301.44 Tm <33> Tj 0.48 Tc 1 0 0 1 132.24 292.32 Tm <43434747414141474147545447414741414747545443414741474141414143435443545454544741474154434141474143435447414141544147474320> Tj 1 0 0 1 466.56 291.60 Tm <3420> Tj 1 0 0 1 471.84 291.60 Tm <3720> Tj 1 0 0 1 477.12 291.36 Tm <33> Tj 0.48 Tc 1 0 0 1 132.00 282.48 Tm <43434747414141474147545447414741414747545443414741474141414143435443545454544741474154434141474143435447414141544147474320> Tj 1 0 0 1 466.80 281.52 Tm <3420> Tj 1 0 0 1 472.08 281.52 Tm <4C20> Tj 1 0 0 1 477.12 281.52 Tm <34> Tj 0.48 Tc 1 0 0 1 132.00 272.40 Tm <47434147414743544747435447414741474743544343474741474341414143545443545454414143414154414143474143435447414741434147474320> Tj 1 0 0 1 466.80 271.44 Tm <3420> Tj 1 0 0 1 472.08 271.44 Tm <4C33> Tj 0.48 Tc 1 0 0 1 132.24 262.56 Tm <4343474741414147414154544741474141472D20> Tj 1 0 0 1 232.08 262.32 Tm <545443414741474E55555C43435443545454544741474154434141474143435447414141544147474320> Tj 1 0 0 1 466.80 261.84 Tm <3420> Tj 1 0 0 1 471.84 261.60 Tm <4F20> Tj 1 0 0 1 477.36 261.60 Tm <39> Tj 0.48 Tc 1 0 0 1 132.24 252.24 Tm <47434147414143544747435447414741414743544343474741474341414143545443545454414143414154414143474143435447414741434147474320> Tj 1 0 0 1 466.80 251.52 Tm <3420> Tj 1 0 0 1 472.08 251.52 Tm <3120> Tj 1 0 0 1 477.12 251.52 Tm <33> Tj 0.48 Tc 1 0 0 1 132.24 242.40 Tm <43434747414141474147545447414741414747545443414741474141414143435443545454544741474154434141474143435447414141544147474320> Tj 1 0 0 1 466.80 241.68 Tm <3420> Tj 1 0 0 1 472.08 241.68 Tm <4F20> Tj 1 0 0 1 477.12 241.68 Tm <35> Tj 0.48 Tc 1 0 0 1 132.24 232.56 Tm 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L!h0> :sGPB%Hޞ}IIIIi>N">y͍y '#输\D9"k.较N!'aZ.0{i M T _4ONZA ]h #pD~ @I}A8` q`5$@͠A;p@.^!մouN'Iק^Ӡl!Tפ5 :tuZZO5V+kTCTmXZ4Cڊեii ;֓T4+UJZwq_ZVowJ_^7^Z]S8_5]oU}P@f&Oz]~_y;k_ƅ&b~f&|'biYSuu׫]z_K!CwCB1}׫׆ _cFsyEoh_ƅOL/ޝg'f'~1u7׽ҿ~F?cяw?޾и㿋Cм'ׇ$G#~}cюsяF#~C5!u)ڥw붖}}5~ti6 яs=F>яڜ8:[ }Cl5aV뷯O{kOQ%{dCUi^][ӐV'B 8hBp1AA`'a %l5խT-mknH>;;3[Aa~lSb  a 6) (r0IaX4a;l/ݤMڷimڭ}DD`0LR4! lP@`b8B@F B & 00zi _{ $ + /m /^"""""""C@` S$=_RMI  `&1@B  C `&HrR-P+fYBGN w &4[^-6+V) QVHDDDDDDCDB""kC"B!&c2!h!TFdJVb%٪%.u^׵U%hȌȌ",DtKvک+^DdFDrўo k78L.̈YgߑU_)Ў##)yf27//U 0@ M1LFTEDqGq"2;3x);3yvfԻ;"";1Fvc4Vc4Y/Ӱ ` 0A)6#؎"8#8#B4#B4쁻#;.ݑٛT@TaA * aBi v#b8#6;@G"?iD};ih0@0 @ 0A a Mi:MIghތwh@G"<Tjݦ STtM6Mi:3whv"=pD}#ʁ">}zznI[I^FzVFwhw;=>ZNM6ji6tڶDDDDDDD^\@ ³x[AzMc_=9,_T^}3s?3lN'PPdiT{ & (pTpٶī~i>?m_%WׯK=Z_KozJU=Ϊ4_[ISdVLn.a[ҶI^튴:۠Ȅ L L mSA/qHF}Ec!d)\ (:@iWU>#DZZū)Z@m   endstream endobj 49 0 obj <>stream Contributions to Zoology, 69 (1/2) - 2000 Farm A1F i rm A^tr:rn A^tr:rm A-wild 81witd 82vfild c1wild c2tr:rm n-E': rm n^I'.rm n-F'arm fl,trarm n- FarmFarmFarmFarmWiIdv1]ildlqildwi 1dFarmFarmFarmFarmFarm 95 A.A.R.R.D1D2n^Dan- CGAGTCATTGGATTTTCGAACGCAACTGGCGGTCTGACGCAACAATCAGACCATGTGTGT 234CGAGTCATTGGATTTTCGAACGCAACTGGCGGTCTGACGCAACAATCAGACCATGTGTGT 234CGAGTCATTGGATTTTCGAACGCAACTGGCGGTCTGACGCAACAATCAGACCATGTGTGT 234CGAGTCATTGGATTTTCGAACGCAACTGGCGGTCTGACGCAACAATCAGACCATGTGTGT 234CGAGTCATTGGATTTTCGAACGCAACTGGCGGTCTGACGCAACAATCAGACCATGTGTGT 234CGAGTCATTTGATTTTCGAACGCATTCGGCGGTCCGACGTAGCAATCGGACCATGTGTGT 238CGAGTCATTGGATTTTCGAACGCAACTGGCGGTCTGACGCAACAATCAGACCATGTGTGT 230CGAGTCATTTGATTTTCGAACGCATTCGGCGGTCCGACGTAGCAATCGGACCATGTGTGT 238CGAGTCATTTGATTTTCGAACGCAACCGGCGGTCCGACGTAGCAATCGGACCATGTGTGT 230CGAGTCATTTGATTTTCGAACGCATTCGGCGGTCCGACGTAGCAATCGGACCATGTGTGT 239CGAGTCATTTGATTTTCGAACGCATTCGGCGGTCCGACGTAGCAATCGGACCATGTGTGT 240CGAGTCATTTGATTTTCGAACGCATTCGGCGGTCCGACGTAGCAATCGGACCATGTGTGT 238CGAGTCATTTGATTTTCGAACGCATTCGGCGGTCCGACGTAGCAATCGGACCATGTGTGT 238**** **** ** **** * **T TCAGTGT CAAC NU\iU\TACCAATCTTAAT GTGATTGAACT T CAGTT CT CTCATGAAATA 2 9 4T TCAGTGT CAACAAAAATACCAATCTTAATGT GATTGAACT T CAGTTCT CTCATGAAATA 2 9 4T TCAGTGTCAACAAAAATACCAATCTTAATGT GATTGAACTTCAGTTCT CTCATGAAATA 2 9 4TTCAGTGTCAACA'UUU\TACCAATCTTAATGTGATTGAACTTCAGTTCTCTCATGAAATA 29 4T TCAGT GTCAAC AJUUU\TAC CAATC TTAATGT GAT TGAACTTCAGTTCTCT CAT GAAATA 2 9 4TTCAGTGTCAACAAAACTCCCAATCTTAATGTGATTGAATTTCGATTCTCTCATGNU\CA 2 98TTCAGTGTCAACAAAAATACCAATCTTAATGTGATTGAACTTCAGTTCTCTCATGAAATA 290TTCAGTGTCAACA'UU\CTCCCAATCTTAATGTGATTGAATTTCGATTCTCTCATGAAACA 298TTCAGTGTCAACAAAACTCCCAATCTTAATGTGATTGAATTTCGATTCTCTCATGAAACA 290TTCAGTGTCAACA'U\'\CTCCCAATCTTAATGTGATTGAATTTCGATTCTCTCATGAAACA 299TTCAGTGTCAACAAAACTCCCAATCTTAATGTGATTGAATTTCGATTCTCTCATGAAACA 3 O OTTCAGTGTCAACA'\'U\CTCCCAATCTTAATGTGATTGAATTTCGATTCTCTCATGAAACA 298TTCAGTGTCAACAAAACTCCCAATCTTAATGTGATTGAATTTCGATTCTCTCATGAAACA 298 ACTGTGC-TTGCGGCTCGGACTGACGAAGTAGTCACCCGTCTCTGACGGCAGAGATTTCG 353ACTGTGC-TTGCGGCTCGGACTGACGAAGTAGTGACCCGTCTCTGACGGCAGTGATTTCG 35 3ACTGTGC-TTGCGGCTCGGACTGACGAAGTAGTCACCCGTCTCTGACGGCAGTGATTTCG 35 3ACTGTGC-TTGCGGCTCGGACTGACGAAGTAGTCACCCGTCTCTGACGGCAGTGATTTCG 35 3ACTGTGCCTTGCGGCTCGGACTGACGAAGTAGTCACCCGTCTCTGACGGCAGTGATTTCG 35 4AGAGGGGCTTGCGGCCTTCTCTGACGAAGTAGTCACCCGCTTGCGGA-- - --TGATTTCG 3 53ACTGTGCCTTGCGGCTCGGACTGACGAATTAGTCACCCGTCTCTGACGGCAGTGATTTCG 350AGAAGGGCTTGCGGCCTTCTCTGACGAATTAGTCACCCGCTTGCGGA-- ---TGATTTCG 3 53AGAGGGGCTTGCGGCCTTCTCTGACGAAGTAGTCACCCGTCTCTGAA-----TGATTTCG 34 5AGAGGGGCTTGCGGCCTTCTCTGACGAAGTAGTCACCCGCTTGCGGA-----TGATTTCG 354AGAGGGGCTTGCGGCCTTCTCTGACGAAGTAGTCACCCGCTTGCGGA-----TGATTTCG 3 5 5AGAGGGGCTTGCGGCCTTCTCTGACGAAGTAGTCACCCGCTTGCGGA-----TGATTTCG 3 5 3AGAAGGGCTTGCGGCCTTCTCTGACGAATTAGTCACCCGCTTGCGGA---- -TGATTTCG 3 5 3 Farm A1Farm A2Farm A3Farm A4wild Biwild 82wird c1wild c2Farm D1Farm D2Farm D3Farm DaFarm D5Farm A1Farm A2Farm A3Farm Aawild 81wifd 82I/.]ild c1wild c2Farm D1Farm D2Farm D3Farm DaFarm D5Fig. 3. Continued. * * ** * * * ** **** ** * ** * **** CCGGAAAGAGTTGAGAAGGT TCAGAGAAAACCTCTTTTGAGATCAAGACCTGAAATAGGC 4 1 3CCGGAAAGAGTTGAGAAGGTTCAGAGAAAACCTCTTTTGAGATCAAGACCTGAAATAGGC 4 1 3CCGGAAAGAGTTGAGAAGGTTCAGAGAAAACCTCTTTTGAGATCAAGACCTGAAATAGGC 4 1 3CCGGAAAGAGTTGAGAAGGTTCAGAGAAAACCTCTTTTGAGATCAAGACCTGAAATAGGC 4 7 3CCGGAAAGAGTTGAGAAGGTTCAGAGAAAACCTCTTTTGAGATCAAGACCTGAAATAGGC 4 L 4GCAGAGCTGGCTGAGAGGCTCCGGAGCAAACTTCTTTAACAATAACGACCTGAGACAGGC 4 L3CCGGAAAGAATTGAGAAG- TTCAGAGNUU\CCTCTTTTGAGATCAAGACCTGAAATAGGC 4 O 9GCAGAACTGGCTGAGAAGCTCCGGAGCAAACTTCTTTAACAATAACGACCTGAGACAGGC 4 1 3CCGGAAAGAGTTGAGAAGGTTCAGAGAAAACCTCTTTTGAGATCAAGACCTGAAATAGGC 4 O 5GCAGAGCTGGCTGAGAGGCTCCGGAGCAAACTTCTTTAACAATAACGACCTGAGACAGGC 47 4GCAGAGCTGGCTGAGAGGCTCCGGAGCAAACTTCTTTAACAATAACGACCTGAGACAGGC 4 1 5GCAGAGCTGGCTGAGAAGCTCCGGAGCAAACTTCTTTAACAATAACGACCTGAGACAGGC 4 1 3GCAGAACTGGCTGAGAAGCTCCGGAACAAACTTCTTTAACAATAACGACCTGAGACAGGC 4 I3 endstream endobj 50 0 obj <> endobj 44 0 obj <>/ProcSet[/PDF/Text/ImageB/ImageC/ImageI]/Subtype 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Shinn et al. - 185 rDNA and ITS sequences as population markers for Lepeophtheirus salmonis Farm A1Farm A2Farm A3Farm Aawild B1viild 82wifd c1wild c2Farm D1Farm D2Farrn D3Farm DaFarm D5.Fig. 3. Continued AAGATAACCC GC CCAAC T TAAGCATATSCTAAGCGEGqAAGATAACCC GC CCAAC T TAAGCAg4ECACTAAGCGGAGGAAAGATAAC C C GC CCAAC T TAAGCATATACTAAGCGqGEAAGATAAC CC GC C CAAC T TAAGCATATCACTAAGCGGAGGAAAGATAACCC GCCCAACT TAAGCATTAAGCGGAGGAAAGATAACCC GCCCAACT TAAGCATATCACTAAGCGGAGGAAAGATAAC CCGC CCAAC T TAAC CATATCACTAAGCGGAAGAAAGATAAC CC GC C CAAC T TAAGCATATACTAAGCGqAEAAGATAACCC GCCCAACT TAAGCAIATCACTAAGCGGAGGAAAGAT AAC C C GC C C AAC T T AA GCAETEdEiEffiGAAGAT AA C C C G C C CAAC T T AAG CAEiEd6EEM6AAGAT AAC C C G C C C AAC T T AAC CAiiEAffiAEMdAAGATAAC CC GC CCAAC T TAAGCATATCACTAAGCGGAAGA* * ** * * * * * * * * * * * *** * * * * * * * ** * * * ** * * * * * * * * AF^454AqAAqA454AtrI446/ qq456AAAAtrt A4 Al 76.3 A2 A,19.3B1 68.386.4 D4 58.6 100.0 D3 91.1D262.0B2 Fig. 4. Unrooted phylogenetic tree of Lepeophtheirus salmonis showing the relationship between the samples based on the ITSlsequence constructed using Saitou & Nei's Neighbour Joining Method (NJ). Samples A,-A. and D,-D. represent lice taken from twoAtlantic salmon farms and samples B,-Brand C,-C, represent lice taken from two wild salmon collection sites. Nodes show bootstrapdetermined confidence estimates for each grouping expressed as percentages. -Ds93.6c2 Diggles & Adlard (1997) found I I variable nucle-otides in the 169-170 sequence representing ITS-1 between 16 isolates of Cryptocaryon. Further,they reported genetic drift amountingto 2.9-3.5ohdivergence when the original samples of 3 isolateswere compared to those maintained in a labora- tory over a 36 month period. The 0.29-4.28% di-vergence between samples of lice collected fromfarmed salmonids when compared to backgroundlevels (i.e. wild lice) of 13.10-13.97%o divergencebetween samples, may serve to illustrate that cagesites represent a bottleneck to genetic flow. 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<616E6420> Tj 1 0 0 1 51.12 470.88 Tm <74686520> Tj 1 0 0 1 68.40 470.88 Tm <6672657175656E637920> Tj 1 0 0 1 115.20 470.64 Tm <6F6620> Tj 1 0 0 1 127.92 470.64 Tm <6368656D6F746865726170657574616E7420> Tj 1 0 0 1 214.08 470.16 Tm <6170706C6963612D> Tj 0.72 Tc 1 0 0 1 31.92 458.64 Tm <74696F6E2E20> Tj 1 0 0 1 55.92 458.40 Tm <5768696C6520> Tj 1 0 0 1 86.88 458.40 Tm <66616C6C6F77696E6720> Tj 1 0 0 1 132.48 458.40 Tm <697320> Tj 1 0 0 1 144.00 458.16 Tm <616E74696369706174656420> Tj 1 0 0 1 195.84 458.16 Tm <746F20> Tj 1 0 0 1 208.80 457.92 Tm <656C696D696E617465> Tj 0.48 Tc 1 0 0 1 31.92 446.16 Tm <74686520> Tj 1 0 0 1 47.28 446.40 Tm <7265736964656E7420> Tj 1 0 0 1 83.76 446.16 Tm <6C69636520> Tj 1 0 0 1 101.28 446.40 Tm <706F70756C6174696F6E2C20> Tj 1 0 0 1 152.40 445.92 Tm <697420> Tj 1 0 0 1 160.80 445.92 Tm <697320> Tj 1 0 0 1 169.92 445.92 Tm <6E6F7420> Tj 1 0 0 1 185.76 445.92 Tm <6B6E6F776E20> Tj 1 0 0 1 217.20 445.68 Tm <746F20> Tj 1 0 0 1 227.76 445.68 Tm <77686174> Tj 0.48 Tc 1 0 0 1 32.16 433.92 Tm <657874656E7420> Tj 1 0 0 1 61.68 433.92 Tm <74686520> Tj 1 0 0 1 78.48 434.16 Tm <67656E6520> Tj 1 0 0 1 101.76 434.16 Tm <706F6F6C20> Tj 1 0 0 1 124.80 433.92 Tm <697320> Tj 1 0 0 1 134.88 434.16 Tm <70726F7061676174656420> Tj 1 0 0 1 186.72 433.44 Tm <6163726F737320> Tj 1 0 0 1 216.96 433.44 Tm <74686520> Tj 1 0 0 1 233.76 433.44 Tm <66616C2D> Tj 0.48 Tc 1 0 0 1 32.16 421.92 Tm <6C6F77696E6720> Tj 1 0 0 1 65.76 422.16 Tm <706572696F6420> Tj 1 0 0 1 97.20 421.68 Tm <746F20> Tj 1 0 0 1 109.68 421.68 Tm <74686520> Tj 1 0 0 1 126.72 421.68 Tm <6E65787420> Tj 1 0 0 1 149.76 421.44 Tm <73746F636B20> Tj 1 0 0 1 176.16 421.44 Tm <6F6620> Tj 1 0 0 1 189.36 421.44 Tm <6669736820> Tj 1 0 0 1 209.28 421.20 Tm <627920> Tj 1 0 0 1 223.92 421.20 Tm <656974686572> Tj 0.72 Tc 1 0 0 1 31.92 409.68 Tm <74686520> Tj 1 0 0 1 52.32 409.68 Tm <696E746572616374696F6E20> Tj 1 0 0 1 106.32 409.44 Tm <6F6620> Tj 1 0 0 1 121.92 409.44 Tm <74686520> Tj 1 0 0 1 142.08 409.68 Tm <706C616E6B746F6E696320> Tj 1 0 0 1 195.84 409.20 Tm <6C617276616520> Tj 1 0 0 1 229.20 408.96 Tm <77697468> Tj 0.48 Tc 1 0 0 1 31.92 397.44 Tm <6E65696768626F7572696E6720> Tj 1 0 0 1 95.28 397.44 Tm <6661726D20> Tj 1 0 0 1 120.48 397.20 Tm <736974657320> Tj 1 0 0 1 144.48 397.20 Tm <6F7220> Tj 1 0 0 1 157.92 397.20 Tm <74686520> Tj 1 0 0 1 176.16 396.96 Tm <616374697669747920> Tj 1 0 0 1 213.84 396.72 Tm <6F6620> Tj 1 0 0 1 227.52 396.96 Tm <6C6F63616C> Tj 0.48 Tc 1 0 0 1 32.16 385.20 Tm <73616C6D6F6E6964732C20> Tj 1 0 0 1 81.12 384.96 Tm <626F746820> Tj 1 0 0 1 103.44 384.96 Tm <6F6620> Tj 1 0 0 1 114.72 384.96 Tm <776869636820> Tj 1 0 0 1 144.00 384.96 Tm <6D617920> Tj 1 0 0 1 165.60 384.72 Tm <736572766520> Tj 1 0 0 1 191.04 384.72 Tm <617320> Tj 1 0 0 1 202.80 384.72 Tm <6120> Tj 1 0 0 1 210.00 384.72 Tm <7265736572766F6972> Tj 0.72 Tc 1 0 0 1 32.16 372.72 Tm <6F6620> Tj 1 0 0 1 47.04 373.20 Tm <67656E6574696320> Tj 1 0 0 1 84.96 372.96 Tm <6964656E746974792E20> Tj 1 0 0 1 127.68 372.72 Tm <5573696E6720> Tj 1 0 0 1 159.60 372.72 Tm <74686520> Tj 1 0 0 1 179.28 372.48 Tm <666561747572657320> Tj 1 0 0 1 220.32 372.24 Tm <6F6620> Tj 1 0 0 1 235.20 372.24 Tm <746865> Tj 0.72 Tc 1 0 0 1 31.92 360.48 Tm <68797065727661726961626C6520> Tj 1 0 0 1 95.76 360.48 Tm <726567696F6E20> Tj 1 0 0 1 127.92 360.48 Tm <6964656E74696669656420> Tj 1 0 0 1 173.76 360.24 Tm <77697468696E20> Tj 1 0 0 1 205.92 360.00 Tm <74686520> Tj 1 0 0 1 223.68 360.00 Tm <4954532D31> Tj 0.72 Tc 1 0 0 1 31.92 348.48 Tm <726567696F6E20> Tj 1 0 0 1 63.36 348.24 Tm <696E20> Tj 1 0 0 1 75.12 348.24 Tm <7468697320> Tj 1 0 0 1 94.56 348.24 Tm <73747564792C20> Tj 1 0 0 1 124.08 348.24 Tm <697420> Tj 1 0 0 1 133.44 348.24 Tm <6D617920> Tj 1 0 0 1 155.52 348.00 Tm <626520> Tj 1 0 0 1 168.96 348.48 Tm <706F737369626C6520> Tj 1 0 0 1 208.32 347.76 Tm <746F20> Tj 1 0 0 1 220.56 347.76 Tm <64657369676E> Tj 0.48 Tc 1 0 0 1 31.92 336.48 Tm <706F70756C6174696F6E20> Tj 1 0 0 1 81.12 336.00 Tm <737065636966696320> Tj 1 0 0 1 117.12 336.00 Tm <7461677320> Tj 1 0 0 1 137.28 335.76 Tm <6F7220> Tj 1 0 0 1 148.56 336.00 Tm <7072696D65727320> Tj 1 0 0 1 184.56 335.76 Tm <7468617420> Tj 1 0 0 1 203.52 335.52 Tm <636F756C6420> Tj 1 0 0 1 230.40 335.52 Tm <7375622D> Tj 0.48 Tc 1 0 0 1 32.16 323.76 Tm <73657175656E746C7920> Tj 1 0 0 1 77.28 323.76 Tm <626520> Tj 1 0 0 1 91.68 323.52 Tm <7573656420> Tj 1 0 0 1 114.72 323.76 Tm <746F20> Tj 1 0 0 1 127.20 323.52 Tm <61736365727461696E20> Tj 1 0 0 1 169.44 323.52 Tm <74686520> Tj 1 0 0 1 186.96 323.28 Tm <6F726967696E20> Tj 1 0 0 1 216.72 323.28 Tm <6F6620> Tj 1 0 0 1 229.44 323.28 Tm <6E61752D> Tj 0.48 Tc 1 0 0 1 31.92 311.76 Tm <706C696920> Tj 1 0 0 1 50.88 311.52 Tm <616E6420> Tj 1 0 0 1 70.56 311.52 Tm <636F7065706F646974657320> Tj 1 0 0 1 126.48 311.28 Tm <6F6E20> Tj 1 0 0 1 141.60 311.28 Tm <6661726D656420> Tj 1 0 0 1 176.16 311.04 Tm <616E6420> Tj 1 0 0 1 195.84 311.04 Tm <77696C6420> Tj 1 0 0 1 219.60 311.04 Tm <73616C6D6F2D> Tj 0.72 Tc 1 0 0 1 31.92 299.28 Tm <6E6964732E20> Tj 1 0 0 1 56.40 299.28 Tm <54686520> Tj 1 0 0 1 76.80 299.28 Tm <6163717569736974696F6E20> Tj 1 0 0 1 127.92 299.04 Tm <6F6620> Tj 1 0 0 1 140.64 299.04 Tm <7375636820> Tj 1 0 0 1 163.68 299.28 Tm <706F70756C6174696F6E20> Tj 1 0 0 1 213.84 298.80 Tm <6D61726B657273> Tj 0.48 Tc 1 0 0 1 31.92 287.04 Tm <636F756C6420> Tj 1 0 0 1 59.28 287.04 Tm <6D616B6520> Tj 1 0 0 1 86.40 286.80 Tm <7369676E69666963616E7420> Tj 1 0 0 1 135.60 286.80 Tm <696E726F61647320> Tj 1 0 0 1 170.88 286.80 Tm <696E746F20> Tj 1 0 0 1 191.28 286.56 Tm <6F757220> Tj 1 0 0 1 209.28 286.56 Tm <6162696C69747920> Tj 1 0 0 1 240.24 286.56 Tm <746F> Tj 0.48 Tc 1 0 0 1 31.92 274.80 Tm <6D6F6E69746F7220> Tj 1 0 0 1 70.32 274.80 Tm <74686520> Tj 1 0 0 1 88.08 274.56 Tm <64697370657273616C2C20> Tj 1 0 0 1 133.68 274.56 Tm <6661746520> Tj 1 0 0 1 154.32 274.32 Tm <616E6420> Tj 1 0 0 1 174.24 274.56 Tm <696D7061637420> Tj 1 0 0 1 207.84 274.08 Tm <6F6620> Tj 1 0 0 1 220.80 274.56 Tm <706C616E6B2D> Tj 0.72 Tc 1 0 0 1 31.92 262.56 Tm <746F6E696320> Tj 1 0 0 1 57.60 262.56 Tm <6C617276616520> Tj 1 0 0 1 87.84 262.32 Tm <6E6F7420> Tj 1 0 0 1 105.60 262.32 Tm <6F6E6C7920> Tj 1 0 0 1 128.88 262.08 Tm <6F6E20> Tj 1 0 0 1 143.28 262.08 Tm <77696C6420> Tj 1 0 0 1 166.80 262.08 Tm <73616C6D6F6E6964732C20> Tj 1 0 0 1 216.96 261.84 Tm <62757420> Tj 1 0 0 1 234.72 261.84 Tm <62652D> Tj 0.48 Tc 1 0 0 1 31.92 250.32 Tm <747765656E20> Tj 1 0 0 1 61.44 250.32 Tm <6661726D20> Tj 1 0 0 1 85.44 250.08 Tm <736974657320> Tj 1 0 0 1 108.00 250.08 Tm <616E6420> Tj 1 0 0 1 127.20 250.08 Tm <6163726F737320> Tj 1 0 0 1 157.92 250.08 Tm <66616C6C6F7720> Tj 1 0 0 1 188.64 250.08 Tm <706572696F64732E> Tj 0.48 Tc 1 0 0 1 31.92 219.60 Tm <41636B6E6F776C656467656D656E7473> Tj 0.72 Tc 1 0 0 1 31.92 198.72 Tm <5468697320> Tj 1 0 0 1 52.56 198.72 Tm <776F726B20> Tj 1 0 0 1 76.08 198.48 Tm <77617320> Tj 1 0 0 1 94.80 198.48 Tm <737570706F7274656420> Tj 1 0 0 1 134.88 198.48 Tm <627920> Tj 1 0 0 1 148.80 198.48 Tm <6120> Tj 1 0 0 1 157.68 198.48 Tm <424253524320> Tj 1 0 0 1 191.04 198.72 Tm <6772616E7420> Tj 1 0 0 1 214.56 198.24 Tm <746F20> Tj 1 0 0 1 226.56 198.24 Tm <447220> Tj 1 0 0 1 240.72 198.00 Tm <432E> Tj 0.48 Tc /F3 7.0 Tf 1 0 0 1 32.16 188.40 Tm <536F6D6D657276696C6C6520> Tj 1 0 0 1 79.68 188.40 Tm <616E6420> Tj 1 0 0 1 94.56 188.40 Tm <4D756E627573686F20> Tj 1 0 0 1 134.40 188.16 Tm <74756E64696E6520> Tj 1 0 0 1 164.16 188.16 Tm <6177617264656420> Tj 1 0 0 1 196.08 187.92 Tm <746F20> Tj 1 0 0 1 205.20 187.92 Tm <50726F66276573736F7220> Tj 1 0 0 1 240.96 187.92 Tm <542E> Tj 0.96 Tc 1 0 0 1 31.92 178.32 Tm <416F6B692E> Tj 0.48 Tc /F3 9.0 Tf 1 0 0 1 274.32 531.36 Tm <5265666572656E636573> Tj 0.48 Tc /F3 7.0 Tf 1 0 0 1 274.32 507.60 Tm <4162656C6520> Tj 1 0 0 1 297.12 507.60 Tm <4C472C20> Tj 1 0 0 1 312.96 507.36 Tm <4B696D20> Tj 1 0 0 1 330.24 507.36 Tm <572C20> Tj 1 0 0 1 341.28 507.36 Tm <46656C67656E686175657220> Tj 1 0 0 1 388.32 507.12 Tm <42452E20> Tj 1 0 0 1 403.68 507.12 Tm <313938392E20> Tj 1 0 0 1 423.84 507.12 Tm <4D6F6C6563756C617220> Tj 1 0 0 1 460.56 506.88 Tm <65766964656E6365> Tj 0.48 Tc 1 0 0 1 285.12 496.80 Tm <666F7220> Tj 1 0 0 1 297.36 496.80 Tm <74686520> Tj 1 0 0 1 310.32 496.80 Tm <696E636C7573696F6E20> Tj 1 0 0 1 344.16 496.56 Tm <6F6620> Tj 1 0 0 1 353.52 496.56 Tm <74686520> Tj 1 0 0 1 366.48 496.80 Tm <7068796C756D20> Tj 1 0 0 1 395.04 496.56 Tm <50656E746173746F6D69646120> Tj 1 0 0 1 443.52 496.32 Tm <696E20> Tj 1 0 0 1 452.16 496.32 Tm <74686520> Tj 1 0 0 1 465.36 496.08 Tm <4372757374612D> Tj 0.72 Tc 1 0 0 1 285.60 486.24 Tm <6365612E20> Tj 1 0 0 1 300.72 486.24 Tm <4D6F6C2E20> Tj 1 0 0 1 319.44 486.24 Tm <42696F6C2E20> Tj 1 0 0 1 337.92 486.00 Tm <45766F6C2E36363A3638352D3639312E> Tj 0.48 Tc 1 0 0 1 274.56 475.68 Tm <426172746120> Tj 1 0 0 1 297.84 475.44 Tm <4A522C20> Tj 1 0 0 1 312.96 475.44 Tm <436F6C657320> Tj 1 0 0 1 335.76 475.44 Tm <42416F20> Tj 1 0 0 1 352.08 475.20 Tm <53636869746F20> Tj 1 0 0 1 377.28 475.20 Tm <4D4C2C20> Tj 1 0 0 1 395.52 475.20 Tm <4665726E616E646F20> Tj 1 0 0 1 433.44 474.96 Tm <4D2E412C20> Tj 1 0 0 1 454.08 474.96 Tm <4D617274696E20> Tj 1 0 0 1 482.40 474.96 Tm <412C> Tj 0.48 Tc 1 0 0 1 285.12 464.88 Tm <44616E666F72746820> Tj 1 0 0 1 323.28 464.64 Tm <48442E20> Tj 1 0 0 1 342.24 464.64 Tm <313939382E20> Tj 1 0 0 1 365.28 464.64 Tm <416E616C7973697320> Tj 1 0 0 1 399.36 464.40 Tm <6F6620> Tj 1 0 0 1 410.40 464.40 Tm <696E667261737065636966696320> Tj 1 0 0 1 459.12 464.16 Tm <766172696174696F6E> Tj 0.48 Tc 1 0 0 1 284.88 454.08 Tm <616D6F6E6720> Tj 1 0 0 1 310.32 454.08 Tm <6669766520> Tj 1 0 0 1 326.16 453.84 Tm <73747261696E7320> Tj 1 0 0 1 350.40 453.84 Tm <6F6620> Tj 1 0 0 1 359.28 453.84 Tm <456972726572696120> Tj 1 0 0 1 389.28 453.60 Tm <6D6178696D6120> Tj 1 0 0 1 418.56 453.60 Tm <66726F6D20> Tj 1 0 0 1 437.28 453.60 Tm <4E6F72746820> Tj 1 0 0 1 458.88 453.60 Tm <416D65726963612E> Tj 0.24 Tc 1 0 0 1 284.64 443.52 Tm <496E742E20> Tj 1 0 0 1 298.56 443.28 Tm <4A20> Tj 1 0 0 1 306.72 443.52 Tm <506172617369746F6C2E20> Tj 1 0 0 1 342.48 443.28 Tm <32383A20> Tj 1 0 0 1 356.64 443.28 Tm <3438352D3439322E> Tj 0.48 Tc 1 0 0 1 274.32 432.96 Tm <426C61697220> Tj 1 0 0 1 295.20 432.72 Tm <442C20> Tj 1 0 0 1 304.80 432.72 Tm <4167617473756D6120> Tj 1 0 0 1 343.92 432.48 Tm <542C20> Tj 1 0 0 1 352.32 432.48 Tm <5761746F6E6F626520> Tj 1 0 0 1 391.20 432.48 Tm <542C20> Tj 1 0 0 1 400.32 432.24 Tm <4F6B6F6D6F746F20> Tj 1 0 0 1 436.80 432.24 Tm <4D2C20> Tj 1 0 0 1 449.04 432.24 Tm <49746F20> Tj 1 0 0 1 461.04 432.24 Tm <412E20> Tj 1 0 0 1 471.60 432.24 Tm <313939372E> Tj 0.48 Tc 1 0 0 1 284.88 422.16 Tm <47656F67726170686963616C20> Tj 1 0 0 1 333.36 422.40 Tm <67656E6574696320> Tj 1 0 0 1 360.24 421.68 Tm <73747275637475726520> Tj 1 0 0 1 392.40 421.68 Tm <77697468696E20> Tj 1 0 0 1 416.16 421.68 Tm <74686520> Tj 1 0 0 1 428.64 421.68 Tm <68756D616E20> Tj 1 0 0 1 454.08 421.68 Tm <6C756E6720> Tj 1 0 0 1 471.12 421.44 Tm <666C756B652C> Tj 0.48 Tc 1 0 0 1 284.64 411.60 Tm <50617261676F6E696D757320> Tj 1 0 0 1 333.60 411.36 Tm <7765737465726D616E692C20> Tj 1 0 0 1 379.20 411.12 Tm <646574656374656420> Tj 1 0 0 1 411.60 411.12 Tm <66726F6D20> Tj 1 0 0 1 431.76 411.12 Tm <444E4120> Tj 1 0 0 1 453.60 410.88 Tm <73657175656E6365732E> Tj 0.48 Tc 1 0 0 1 284.88 401.04 Tm <506172617369746F6C6F676C2C20> Tj 1 0 0 1 331.44 400.80 Tm <6C6C353A20> Tj 1 0 0 1 348.72 400.80 Tm <343120> Tj 1 0 0 1 358.32 400.56 Tm <3120> Tj 1 0 0 1 361.92 402.24 Tm <2D3431312E> Tj 0.48 Tc 1 0 0 1 274.32 390.24 Tm <4368696C746F6E20> Tj 1 0 0 1 304.56 390.24 Tm <4E422C20> Tj 1 0 0 1 321.36 390.00 Tm <47617373657220> Tj 1 0 0 1 349.20 390.00 Tm <52422C20> Tj 1 0 0 1 365.76 390.00 Tm <42657665726964676520> Tj 1 0 0 1 404.64 390.00 Tm <492E20> Tj 1 0 0 1 413.04 389.76 Tm <313939352E20> Tj 1 0 0 1 434.88 389.76 Tm <446966666572656E63657320> Tj 1 0 0 1 477.60 389.76 Tm <696E20> Tj 1 0 0 1 486.96 389.52 Tm <61> Tj 0.48 Tc 1 0 0 1 284.88 379.68 Tm <7269626F736F6D616C20> Tj 1 0 0 1 321.36 379.68 Tm <444E4120> Tj 1 0 0 1 341.76 379.44 Tm <73657175656E636520> Tj 1 0 0 1 375.12 379.20 Tm <6F6620> Tj 1 0 0 1 384.24 379.44 Tm <6D6F7270686F6C6F676963616C6C7920> Tj 1 0 0 1 443.28 379.20 Tm <696E64697374696E67756973682D> Tj 0.24 Tc 1 0 0 1 284.88 368.88 Tm <61626C6520> Tj 1 0 0 1 300.72 368.88 Tm <7370656369657320> Tj 1 0 0 1 326.40 368.88 Tm <77697468696E74686520> Tj 1 0 0 1 361.68 368.64 Tm <4879706F646F6E74757320> Tj 1 0 0 1 404.64 368.40 Tm <6D6163726F706920> Tj 1 0 0 1 434.88 368.40 Tm <636F6D706C657820> Tj 1 0 0 1 464.88 368.88 Tm <284E656D612D> Tj 0.48 Tc 1 0 0 1 284.88 358.32 Tm <746F64613A20> Tj 1 0 0 1 304.80 358.08 Tm <5374726F6E67796C6F69642C6561292E20> Tj 1 0 0 1 360.48 358.08 Tm <496E742E20> Tj 1 0 0 1 374.16 357.84 Tm <4A2E20> Tj 1 0 0 1 382.32 358.08 Tm <506172617369746F6C2E20> Tj 1 0 0 1 418.08 357.84 Tm <32353A20> Tj 1 0 0 1 432.48 357.60 Tm <3634372D3635312E> Tj 0.48 Tc 1 0 0 1 274.32 347.52 Tm <43756E6E696E6768616D20> Tj 1 0 0 1 325.20 347.52 Tm <434F2E20> Tj 1 0 0 1 343.20 347.52 Tm <313939372E20> Tj 1 0 0 1 364.80 347.28 Tm <5370656369657320> Tj 1 0 0 1 393.36 347.28 Tm <766172696174696F6E20> Tj 1 0 0 1 426.48 347.04 Tm <77697468696E20> Tj 1 0 0 1 450.96 347.04 Tm <74686520> Tj 1 0 0 1 464.16 347.04 Tm <696E7465726E616C> Tj 0.24 Tc 1 0 0 1 284.88 336.96 Tm <7472616E7363726962656420> Tj 1 0 0 1 324.24 336.72 Tm <73706163657220> Tj 1 0 0 1 347.52 337.44 Tm <286C54532920> Tj 1 0 0 1 367.44 336.72 Tm <726567696F6E20> Tj 1 0 0 1 390.48 336.48 Tm <6F6620> Tj 1 0 0 1 399.36 336.48 Tm <4779726F64616374796C757320> Tj 1 0 0 1 445.92 336.72 Tm <284D6F6E6F67656E65613A> Tj 0.48 Tc 1 0 0 1 284.88 326.16 Tm <4779726F64616374796C696461652920> Tj 1 0 0 1 342.00 326.16 Tm <7269626F736F6D616C20> Tj 1 0 0 1 378.00 325.92 Tm <524E4120> Tj 1 0 0 1 397.20 326.16 Tm <67656E65732E20> Tj 1 0 0 1 420.00 325.68 Tm <2D7B20> Tj 1 0 0 1 427.20 325.68 Tm <506172757369746F6C2E20> Tj 1 0 0 1 462.00 325.68 Tm <38333A20> Tj 1 0 0 1 474.96 325.68 Tm <3231352D> Tj 1.44 Tc /F3 10.0 Tf 1 0 0 1 284.88 315.60 Tm <3272392E> Tj 0.72 Tc /F3 7.0 Tf 1 0 0 1 274.32 305.04 Tm <43756E6E696E6768616D20> Tj 1 0 0 1 326.64 304.80 Tm <434F2C20> Tj 1 0 0 1 344.88 304.80 Tm <4D6347696C6C697672617920> Tj 1 0 0 1 393.84 304.56 Tm <444D2E20> Tj 1 0 0 1 413.52 304.56 Tm <4D61634B656E7A696520> Tj 1 0 0 1 458.88 304.32 Tm <4B2E20> Tj 1 0 0 1 471.60 304.32 Tm <313939352E> Tj 0.48 Tc 1 0 0 1 284.88 294.48 Tm <5068796C6F67656E6574696320> Tj 1 0 0 1 333.36 294.00 Tm <616E616C7973697320> Tj 1 0 0 1 364.56 294.00 Tm <6F6620> Tj 1 0 0 1 375.60 294.00 Tm <4779726F64616374796C757320> Tj 1 0 0 1 425.04 293.76 Tm <73616C6172697320> Tj 1 0 0 1 452.40 293.76 Tm <4D616C6D626572672C> Tj 0.48 Tc 1 0 0 1 285.60 283.68 Tm <3139353720> Tj 1 0 0 1 303.60 283.68 Tm <626173656420> Tj 1 0 0 1 325.20 283.44 Tm <6F6E20> Tj 1 0 0 1 335.28 283.44 Tm <74686520> Tj 1 0 0 1 348.00 283.44 Tm <736D616C6C20> Tj 1 0 0 1 368.40 283.44 Tm <737562756E697420> Tj 1 0 0 1 395.52 283.68 Tm <2820> Tj 1 0 0 1 399.60 283.44 Tm 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B]/]p/Ÿ]FHjҦ\h]~ڠ^ dtUMaJ*IRtjm?WooUR ^h%][ _{zJ$p]}q?wQlRpRL[A?o_vF-=viu]}z__o~z60ל]eZm5ٚ3VfcU/][^ji6KZT8~A+C}Ժ׽봺.3IkJq>3Iz(uZTߥ*AA 0nh0i0aABM6)iZVv;`TAvGO][^C(yuC Za/m][KjﯥS5^a!`P@ iMAmp%:A121l4m%jҴ!kcc8lwڑeWj݅9~Y.K( ( a004b 29{ DU K 4=6`˻V!0FPq(pՊtҰ#GKJG ?[D#b"""S‚ ! M7 h ڄi 6d^vl0J*E~f\0*G !`B,&a4В6MU mB.xAH zA !C a/FaZ\C!6HkZW}\@kt%?GtJo)AMGp^Մoa8WI*խ/zz]%Iz,Ut/L: nJ굾Ӷz_]n5I;OJxb見JiУVm/0bMo ^?5q 4ʩr[gka*)@f-w@ endstream endobj 63 0 obj <>stream Contributions to Zoology, 69 (l/2) 2000 Table L Percentage similarity between nucleotide sequences for the internal transcribed spacer (ITS- I ) obtained from four populationsof Lepeophtheirus salmonis. Lice collected at sites A and D were from farmed salmonids whilst those lice collected from sites B andC were from wild salmonids. 97 Sample Ar A2Az 99.34Ar 99.78 99.784.4 99.56 99.78Br 99.56 99.56Bz 86.49 86.68cr 97.58 97.80cz 86.49 86.68D1 92.75 92.95D2 86.74 86.49Dr 85.90 86 07D+ 86.49 86.68Ds 86.06 86 24 100.0099.78 99.7886.90 86.90 86.9098.02 98.02 98.2486.90 86 90 86.6893.17 93.17 93.1786.71 86 71 86.7186.30 86.s2 86.3086.90 86.90 86.9086.46 86.46 86.24 Ds Da D3 D2 D1 L2 LI R,, Bq A4 A: it.y99.12 86.0391.39 93.30 el.l799.78 85.19 98.90 91.1999 t2 84.78 98.02 90.77 99.8299.56 85.81 99.56 9t.61 99.34 98.6898.45 85.59 99.56 90.73 98.53 97 .58 98.90 Summary statistics for lice within sites: within A (n - 6) 99.71 + 0.23% similarity; within B (n: 1) 86.90% similarity; within C (n -1) 86.03% similarity; and within D (n = l0) 95.72 + 4.04% similarity. Across all lice within farm sites (n = 36) there was 91.91 +5.70%similarity; withinwildsites(n-6)therewas90.39+6.450 similarity: andacrossallliceandallsites(n:78)therewas92.14+ 5.88% similarity. tion in the genetic variability or population isola-tion within farm sites, may stem from a combina-tion of environmental and anthropogenic inducedfactors including farming intensity, loch flush rates,fallowing history, the retention of lice larval stagesand the frequency of chemotherapeutant applica-tion. While fallowing is anticipated to eliminatethe resident lice population, it is not known to whatextent the gene pool is propagated across the fal-lowing period to the next stock of fish by eitherthe interaction of the planktonic larvae withneighbouring farm sites or the activity of localsalmonids, both of which may serve as a reservoirof genetic identity. Using the features of thehypervariable region identified within the ITS-1region in this study, it may be possible to designpopulation specific tags or primers that could sub-sequently be used to ascertain the origin of nau-plii and copepodites on farmed and wild salmo-nids. The acquisition of such population markerscould make significant inroads into our ability tomonitor the dispersal, fate and impact of plank-tonic larvae not only on wild salmonids, but be-tween farm sites and across fallow periods.AcknowledgementsThis work was supported by a BBSRC grant to Dr C.Sommerville and Munbusho tundine awarded to Prof'essor T.Aoki. ReferencesAbele LG, Kim W, Felgenhauer BE. 1989. Molecular evidencefor the inclusion of the phylum Pentastomida in the Crusta-cea. Mol. Biol. Evol.66:685-691.Barta JR, Coles BAo Schito ML, Fernando M.A, Martin A,Danforth HD. 1998. Analysis of infraspecific variationamong five strains of Eirreria maxima from North America.Int. J Parasitol. 28: 485-492.Blair D, Agatsuma T, Watonobe T, Okomoto M, Ito A. 1997.Geographical genetic structure within the human lung fluke,Paragonimus westermani, detected from DNA sequences.Parasitologl, ll5: 41 1 -411.Chilton NB, Gasser RB, Beveridge I. 1995. Differences in aribosomal DNA sequence of morphologically indistinguish-able species withinthe Hypodontus macropi complex (Nema-toda: Strongyloid,ea). Int. J. Parasitol. 25: 647-651.Cunningham CO. 1997. Species variation within the internaltranscribed spacer (lTS) region of Gyrodactylus (Monogenea:Gyrodactylidae) ribosomal RNA genes. -{ Parusitol. 83: 215-2r9.Cunningham CO, McGillivray DM. MacKenzie K. 1995.Phylogenetic analysis of Gyrodactylus salaris Malmberg,1957 based on the small subunit ( I 8S) ribosomal RNA gene.Mol. Biochem. Parasitol. 7l: 139-142.De Meetis T, Marin R, Renaud F,1992. Genetic heterogeneitywithin populations of Lepeophtheirus europaensis(Copepoda: Caligidae) parasitic on two host species. lnt. -{Parasito l. 22t 1 119- 1 181Diggles BK, Adlard RD. 1997. Intraspecific variation inCtl,ptocaryon irritans. J. Eukaryotic Microbiol. 44: 25-32.Ellis RE, Sulston JE, Coulson AR. 1986. The rDNA of C.elegans'. sequence and structure. Nucl. 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<506174686F6C2E20> Tj 1 0 0 1 96.96 656.64 Tm <31373A20> Tj 1 0 0 1 111.12 656.64 Tm <31372D32322E> Tj 0.48 Tc 1 0 0 1 58.80 646.08 Tm <4C6974746C65776F6F6420> Tj 1 0 0 1 99.84 646.08 Tm <44544A2C20> Tj 1 0 0 1 119.52 646.08 Tm <4A6F686E73746F6E20> Tj 1 0 0 1 154.80 646.08 Tm <44412E20> Tj 1 0 0 1 170.64 646.08 Tm <313939352E20> Tj 1 0 0 1 191.28 646.08 Tm <4D6F6C6563756C617220> Tj 1 0 0 1 228.00 646.08 Tm <7068796C6F67656E6574696373> Tj 0.48 Tc 1 0 0 1 69.60 635.52 Tm <6F6620> Tj 1 0 0 1 78.96 635.52 Tm <74686520> Tj 1 0 0 1 91.68 635.52 Tm <666F757220> Tj 1 0 0 1 108.00 635.52 Tm <53632F756973746F736F6D6120> Tj 1 0 0 1 153.36 635.76 Tm <67726F75707320> Tj 1 0 0 1 179.04 635.28 Tm <64657465726D696E656420> Tj 1 0 0 1 220.08 635.28 Tm <7769746820> Tj 1 0 0 1 237.36 635.76 Tm <7061727469616C20> Tj 1 0 0 1 262.08 635.28 Tm <323853> Tj 0.24 Tc 1 0 0 1 69.60 624.96 Tm <7269626F736F6D616C20> Tj 1 0 0 1 105.84 624.96 Tm <524E4120> Tj 1 0 0 1 125.28 625.44 Tm <67656E6520> Tj 1 0 0 1 143.28 624.96 Tm <73657175656E6365732E20> Tj 1 0 0 1 181.92 624.96 Tm <506172617369746F6C6F677920> Tj 1 0 0 1 227.52 624.72 Tm <6C6C6C3A20> Tj 1 0 0 1 245.28 624.72 Tm <31363720> Tj 1 0 0 1 258.24 626.40 Tm <2D313720> Tj 1 0 0 1 269.04 624.48 Tm <352E> Tj 0.48 Tc 1 0 0 1 59.04 614.40 Tm <4D6F7267616E20> Tj 1 0 0 1 90.24 614.40 Tm <4A41542C20> Tj 1 0 0 1 108.48 614.40 Tm <426C61697220> Tj 1 0 0 1 129.12 614.40 Tm <442E20> Tj 1 0 0 1 139.44 614.40 Tm <313939382E20> Tj 1 0 0 1 160.32 614.40 Tm <52656C617469766520> Tj 1 0 0 1 190.80 614.40 Tm <6D657269747320> Tj 1 0 0 1 214.56 614.16 Tm <6F6620> Tj 1 0 0 1 223.44 614.16 Tm <6E75636C65617220> Tj 1 0 0 1 250.80 614.16 Tm <7269626F736F2D> Tj 0.48 Tc 1 0 0 1 69.84 604.08 Tm <6D616C20> Tj 1 0 0 1 84.72 603.84 Tm <696E7465726E616C20> Tj 1 0 0 1 112.80 603.84 Tm <7472616E7363726962656420> Tj 1 0 0 1 153.36 603.84 Tm <7370616365727320> Tj 1 0 0 1 180.48 603.84 Tm <616E6420> Tj 1 0 0 1 194.64 603.84 Tm <6D69746F63686F6E647269616C20> Tj 1 0 0 1 245.04 603.60 Tm <434F6C20> Tj 1 0 0 1 263.04 603.60 Tm <616E64> Tj 0.48 Tc 1 0 0 1 69.60 593.52 Tm <4E443120> Tj 1 0 0 1 89.28 593.76 Tm <67656E657320> Tj 1 0 0 1 111.60 593.52 Tm <666F7220> Tj 1 0 0 1 124.56 593.28 Tm <64697374696E6775697368696E6720> Tj 1 0 0 1 176.40 593.28 Tm <616D6F6E6720> Tj 1 0 0 1 202.32 593.28 Tm <456368696E6F73746F6D6120> Tj 1 0 0 1 251.28 593.04 Tm <73706563696573> Tj 0.24 Tc 1 0 0 1 69.84 583.20 Tm <285472656D61746F6461292E20> Tj 1 0 0 1 117.12 582.96 Tm <506172617320> Tj 1 0 0 1 136.56 582.96 Tm <69746F20> Tj 1 0 0 1 145.68 582.96 Tm <6C6F677920> Tj 1 0 0 1 163.20 582.96 Tm <313136272E20> Tj 1 0 0 1 181.20 582.96 Tm <3238392D32393720> Tj 1 0 0 1 210.24 582.72 Tm <2E> Tj 0.48 Tc 1 0 0 1 59.04 572.64 Tm <4E6F7264686167656E20> Tj 1 0 0 1 101.76 572.40 Tm <4A522C20> Tj 1 0 0 1 116.16 572.64 Tm <486575636820> Tj 1 0 0 1 142.32 572.64 Tm <50412C20> Tj 1 0 0 1 156.72 572.40 Tm <53636872616D20> Tj 1 0 0 1 186.72 572.40 Tm <54412E20> Tj 1 0 0 1 202.08 572.40 Tm <313939382E20> Tj 1 0 0 1 223.44 572.40 Tm <53697A6520> Tj 1 0 0 1 240.00 572.40 Tm <617320> Tj 1 0 0 1 249.12 572.16 Tm <616E20> Tj 1 0 0 1 259.20 572.40 Tm <696E64692D> Tj 0.48 Tc 1 0 0 1 69.84 562.08 Tm <6361746F7220> Tj 1 0 0 1 89.76 562.08 Tm <6F6620> Tj 1 0 0 1 99.36 562.08 Tm <74686520> Tj 1 0 0 1 112.56 562.08 Tm <6F726967696E20> Tj 1 0 0 1 135.84 562.08 Tm <6F6620> Tj 1 0 0 1 145.92 562.08 Tm <73616C6D6F6E20> Tj 1 0 0 1 173.28 562.08 Tm <6C69636520> Tj 1 0 0 1 188.40 562.08 Tm <4C6570656F70687468656972757320> Tj 1 0 0 1 244.56 561.84 Tm <73616C6D6F6E6973> Tj 0.48 Tc 1 0 0 1 69.84 551.76 Tm <28436F7065706F64613A20> Tj 1 0 0 1 113.76 551.28 Tm <43616C696769646165292E20> Tj 1 0 0 1 155.76 551.52 Tm <50726F63656564696E677320> Tj 1 0 0 1 202.56 551.28 Tm <616E6420> Tj 1 0 0 1 217.92 551.28 Tm <41627374726163747320> Tj 1 0 0 1 254.64 551.28 Tm <6F4A27746865> Tj 0.24 Tc 1 0 0 1 69.36 540.96 Tm <466F7572746820> Tj 1 0 0 1 94.80 540.96 Tm <496E7465726E6174696F6E646C20> Tj 1 0 0 1 143.28 540.72 Tm <4372757374616365616E20> Tj 1 0 0 1 185.52 540.72 Tm <436F6E73726573732E20> Tj 1 0 0 1 221.52 540.72 Tm <416D7374657264616D20> Tj 1 0 0 1 264.00 540.72 Tm <32302D> Tj 0.48 Tc 1 0 0 1 69.84 530.40 Tm <323420> Tj 1 0 0 1 81.36 530.40 Tm <4A72726C792C20> Tj 1 0 0 1 98.40 530.88 Tm <702E3130322E> Tj 0.48 Tc 1 0 0 1 59.04 519.84 Tm <4F6B616D6F746F20> Tj 1 0 0 1 95.52 519.84 Tm <4D2C20> Tj 1 0 0 1 106.56 519.84 Tm <4167617473756D6120> Tj 1 0 0 1 145.44 519.84 Tm <542C20> Tj 1 0 0 1 154.08 519.84 Tm <4B75726F7361776120> Tj 1 0 0 1 192.48 519.84 Tm <542C20> Tj 1 0 0 1 201.84 519.84 Tm <6C746F20> Tj 1 0 0 1 213.36 519.84 Tm <412E20> Tj 1 0 0 1 223.92 519.84 Tm <3139393720> Tj 1 0 0 1 240.72 519.60 Tm <2E20> Tj 1 0 0 1 244.80 519.60 Tm <5068796C6F67652D> Tj 0.48 Tc 1 0 0 1 69.60 509.28 Tm <6E6574696320> Tj 1 0 0 1 89.28 509.28 Tm <72656C6174696F6E736869707320> Tj 1 0 0 1 136.08 509.28 Tm <6F6620> Tj 1 0 0 1 145.92 509.28 Tm <746872656520> Tj 1 0 0 1 166.08 509.28 Tm <68796D656E6F6C65706964696420> Tj 1 0 0 1 220.56 509.28 Tm <7370656369657320> Tj 1 0 0 1 248.16 509.28 Tm <696E666572726564> Tj 0.48 Tc 1 0 0 1 69.84 498.96 Tm <66726F6D20> Tj 1 0 0 1 88.56 498.96 Tm <6E75636C65617220> Tj 1 0 0 1 115.92 498.96 Tm <7269626F736F6D616C20> Tj 1 0 0 1 152.88 498.72 Tm <616E6420> Tj 1 0 0 1 167.28 498.96 Tm <6D69746F63686F6E647269616C20> Tj 1 0 0 1 217.92 498.72 Tm <444E4120> Tj 1 0 0 1 238.32 498.72 Tm <73657175656E6365732E> Tj 0.24 Tc 1 0 0 1 69.36 488.40 Tm <506172617320> Tj 1 0 0 1 89.28 488.40 Tm <6920> Tj 1 0 0 1 91.68 488.40 Tm <746F20> Tj 1 0 0 1 98.40 488.40 Tm <6C6F20> Tj 1 0 0 1 105.84 488.64 Tm <67317420> Tj 1 0 0 1 115.92 488.40 Tm <6C6C3520> Tj 1 0 0 1 128.88 488.40 Tm <3A20> Tj 1 0 0 1 134.16 488.40 Tm <36363120> Tj 1 0 0 1 146.64 490.08 Tm <2D3636362E> Tj 0.48 Tc 1 0 0 1 59.04 478.08 Tm <5061676520> Tj 1 0 0 1 78.48 478.08 Tm <52444D2E20> Tj 1 0 0 1 103.20 478.08 Tm <313939362E20> Tj 1 0 0 1 123.84 478.08 Tm <54524545564945573A20> Tj 1 0 0 1 171.60 477.84 Tm <416E20> Tj 1 0 0 1 184.08 477.84 Tm <6170706C69636174696F6E20> Tj 1 0 0 1 224.40 477.84 Tm <746F20> Tj 1 0 0 1 233.28 477.84 Tm <646973706C617920> Tj 1 0 0 1 259.68 478.08 Tm <7068792D> Tj 0.48 Tc 1 0 0 1 69.84 467.52 Tm <6C6F67656E6574696320> Tj 1 0 0 1 103.92 467.52 Tm <747265657320> Tj 1 0 0 1 123.36 467.52 Tm <6F6E20> Tj 1 0 0 1 134.64 467.76 Tm <706572736F6E616C20> Tj 1 0 0 1 167.04 467.28 Tm <636F6D7075746572732E20> Tj 1 0 0 1 208.32 467.28 Tm <436F6D702E20> Tj 1 0 0 1 232.56 467.28 Tm <4170706C69632E20> Tj 1 0 0 1 260.16 467.28 Tm <42696F2D> Tj 0.72 Tc 1 0 0 1 69.60 456.96 Tm <736369656E63657320> Tj 1 0 0 1 101.28 456.96 Tm <31323A20> Tj 1 0 0 1 114.96 456.96 Tm <3335372D3335382E> Tj 0.48 Tc 1 0 0 1 59.04 446.40 Tm <52696E64657220> Tj 1 0 0 1 87.12 446.40 Tm <482C20> Tj 1 0 0 1 97.92 446.40 Tm <4B61747A77696E6B656C2D576C6164617273636820> Tj 1 0 0 1 185.52 446.16 Tm <532C20> Tj 1 0 0 1 194.40 446.40 Tm <4C6F736368657220> Tj 1 0 0 1 225.60 446.40 Tm <542E20> Tj 1 0 0 1 235.20 446.16 Tm <31393937612E20> Tj 1 0 0 1 260.64 446.16 Tm <4576692D> Tj 0.48 Tc 1 0 0 1 69.84 435.84 Tm <64656E636520> Tj 1 0 0 1 93.84 435.84 Tm <666F7220> Tj 1 0 0 1 107.52 435.84 Tm <74686520> Tj 1 0 0 1 121.92 435.84 Tm <6578697374656E636520> Tj 1 0 0 1 158.40 435.84 Tm <6F6620> Tj 1 0 0 1 169.20 436.32 Tm <67656E65746963616C6C7920> Tj 1 0 0 1 211.68 435.84 Tm <64697374696E637420> Tj 1 0 0 1 241.68 435.60 Tm <73747261696E7320> Tj 1 0 0 1 268.56 435.60 Tm <6F66> Tj 0.48 Tc 1 0 0 1 69.60 425.52 Tm <456E7465726F6379746F7A6F6F6E20> Tj 1 0 0 1 126.00 425.52 Tm <6269656E657573692E20> Tj 1 0 0 1 158.88 425.52 Tm <506172617369746F6C2E20> Tj 1 0 0 1 194.64 425.52 Tm <5265732E20> Tj 1 0 0 1 211.92 425.28 Tm <38333A20> Tj 1 0 0 1 225.84 425.28 Tm <3637302D3637322E> Tj 0.48 Tc 1 0 0 1 59.04 415.20 Tm <52696E64657220> Tj 1 0 0 1 86.64 415.20 Tm <482C20> Tj 1 0 0 1 97.20 415.20 Tm <52617573636820> Tj 1 0 0 1 126.24 415.20 Tm <524C2C54616B61686173686920> Tj 1 0 0 1 180.96 414.96 Tm <4B2C20> Tj 1 0 0 1 191.52 414.96 Tm <4B6F707020> Tj 1 0 0 1 213.60 414.96 Tm <482C54686F6D7363686B65412C> Tj 0.48 Tc 1 0 0 1 69.84 404.64 Tm <4C6F736368657220> Tj 1 0 0 1 102.72 404.88 Tm <542E20> Tj 1 0 0 1 113.76 404.88 Tm <31393937622E20> Tj 1 0 0 1 140.88 404.64 Tm <4C696D6974656420> Tj 1 0 0 1 171.84 404.64 Tm <72616E676520> Tj 1 0 0 1 194.88 404.64 Tm <6F6620> Tj 1 0 0 1 205.44 404.88 Tm <67656E6574696320> Tj 1 0 0 1 234.24 404.64 Tm <766172696174696F6E20> Tj 1 0 0 1 268.80 404.64 Tm <696E> Tj 0.48 Tc 1 0 0 1 69.36 394.32 Tm <456368696E6F636F6363757320> Tj 1 0 0 1 119.76 394.32 Tm <6D756C74696C6F63756C617269732E20> Tj 1 0 0 1 172.32 394.32 Tm <4A2E20> Tj 1 0 0 1 180.48 394.32 Tm <506172617369746F6C2E20> Tj 1 0 0 1 216.48 394.08 Tm <38333A20> Tj 1 0 0 1 231.36 394.32 Tm <313034352D313035302E> Tj 0.48 Tc 1 0 0 1 59.04 383.76 Tm <526F6D7374616420> Tj 1 0 0 1 94.56 383.76 Tm <412C20> Tj 1 0 0 1 105.60 383.76 Tm <47617373657220> Tj 1 0 0 1 134.16 383.76 Tm <52422C20> Tj 1 0 0 1 150.96 383.76 Tm <4E616E73656E20> Tj 1 0 0 1 181.20 383.76 Tm <502C20> Tj 1 0 0 1 190.32 383.76 Tm <506F6C65726D616E20> Tj 1 0 0 1 228.24 383.76 Tm <414D2C20> Tj 1 0 0 1 247.68 383.52 Tm <4368696C746F6E> Tj 0.48 Tc 1 0 0 1 69.84 373.44 Tm <4E422E20> Tj 1 0 0 1 87.12 373.44 Tm <313939382E20> Tj 1 0 0 1 108.96 373.44 Tm <4E656361746F7220> Tj 1 0 0 1 139.92 373.20 Tm <616D65726963756E757320> Tj 1 0 0 1 183.60 373.44 Tm <284E656D61746F64613A20> Tj 1 0 0 1 226.56 373.20 Tm <416E63796C6F73746F6D612D> Tj 0.48 Tc 1 0 0 1 69.84 362.88 Tm <74696461652920> Tj 1 0 0 1 91.68 362.88 Tm <66726F6D20> Tj 1 0 0 1 109.92 362.88 Tm <41667269636120> Tj 1 0 0 1 134.40 362.64 Tm <616E6420> Tj 1 0 0 1 149.04 362.88 Tm <4D616C617973696120> Tj 1 0 0 1 182.16 362.88 Tm <6861766520> Tj 1 0 0 1 200.40 362.64 Tm <646966666572656E7420> Tj 1 0 0 1 232.08 362.64 Tm <4954532D3220> Tj 1 0 0 1 254.16 362.64 Tm <72444E41> Tj 0.72 Tc 1 0 0 1 311.52 771.84 Tm <73657175656E6365732E20> Tj 1 0 0 1 350.88 771.84 Tm <496E742E20> Tj 1 0 0 1 364.56 771.60 Tm <4A20> Tj 1 0 0 1 372.96 771.60 Tm <506172617369746F4C20> Tj 1 0 0 1 408.48 771.60 Tm <32533A20> Tj 1 0 0 1 422.64 771.36 Tm <366C20> Tj 1 0 0 1 432.24 771.60 Tm <6C2D3631352E> Tj 0.48 Tc 1 0 0 1 300.96 761.28 Tm <5361696B6920> Tj 1 0 0 1 321.12 761.28 Tm <524B2C20> Tj 1 0 0 1 337.44 761.28 Tm <47656C66616E6420> Tj 1 0 0 1 368.88 761.28 Tm <44482C20> Tj 1 0 0 1 385.44 761.04 Tm <53746F6666656C20> Tj 1 0 0 1 410.88 761.04 Tm <532C20> Tj 1 0 0 1 419.28 761.04 Tm <53636861726620> Tj 1 0 0 1 445.20 761.04 Tm <534A2C20> Tj 1 0 0 1 457.68 761.04 Tm <4869677563686920> Tj 1 0 0 1 488.64 761.04 Tm <526F20> Tj 1 0 0 1 498.24 761.04 Tm <486F726E> Tj 0.48 Tc 1 0 0 1 311.52 750.72 Tm <47542C20> Tj 1 0 0 1 326.64 750.96 Tm <4D756C6C697320> Tj 1 0 0 1 351.12 750.72 Tm <4B422C20> Tj 1 0 0 1 366.96 750.72 Tm <45726C69636820> Tj 1 0 0 1 391.20 750.72 Tm <48412E20> Tj 1 0 0 1 408.00 750.72 Tm <313938382E20> Tj 1 0 0 1 427.92 750.72 Tm <5072696D65722D646972656374656420> Tj 1 0 0 1 482.64 750.48 Tm <656E7A796D61746963> Tj 0.48 Tc 1 0 0 1 311.52 740.16 Tm <616D706C696669636174696F6E20> Tj 1 0 0 1 359.52 740.16 Tm <6F6620> Tj 1 0 0 1 368.16 740.16 Tm <444E4120> Tj 1 0 0 1 388.08 740.16 Tm <7769746820> Tj 1 0 0 1 405.12 739.92 Tm <6120> Tj 1 0 0 1 410.40 740.16 Tm <746865726D6F737461626C6520> Tj 1 0 0 1 456.48 739.92 Tm <444E4120> Tj 1 0 0 1 476.16 740.40 Tm <706F6C796D65726173652E> Tj 0.48 Tc 1 0 0 1 311.28 729.60 Tm <536369656E636520> Tj 1 0 0 1 340.56 729.60 Tm <3233397420> Tj 1 0 0 1 359.04 729.60 Tm <3438312E> Tj 0.48 Tc 1 0 0 1 300.96 719.28 Tm <53616D62726F6F6B20> Tj 1 0 0 1 341.52 719.04 Tm <4A2220> Tj 1 0 0 1 350.64 719.28 Tm <4672697473636820> Tj 1 0 0 1 379.44 719.04 Tm <45462E20> Tj 1 0 0 1 394.32 719.04 Tm <4D616E696174697320> Tj 1 0 0 1 429.36 719.04 Tm <542E20> Tj 1 0 0 1 439.68 719.04 Tm <6620> Tj 1 0 0 1 443.52 718.80 Tm <3938392E20> Tj 1 0 0 1 461.04 718.80 Tm <4D6F6C6563756C617220> Tj 1 0 0 1 499.92 718.80 Tm <636B746E2D> Tj 0.48 Tc 1 0 0 1 311.52 708.72 Tm <696E673A20> Tj 1 0 0 1 326.64 708.72 Tm <4120> Tj 1 0 0 1 334.80 708.72 Tm <6C61626F7261746F727920> Tj 1 0 0 1 373.44 708.48 Tm <6D616E75616C2E32226420> Tj 1 0 0 1 415.68 708.48 Tm <65646974696F6E2E20> Tj 1 0 0 1 444.24 708.48 Tm <4E657720> Tj 1 0 0 1 461.76 708.48 Tm <596F726B3A20> Tj 1 0 0 1 482.88 708.24 Tm <436F6C6420> Tj 1 0 0 1 501.84 708.24 Tm <4861722D> Tj 0.48 Tc 1 0 0 1 311.28 698.16 Tm <626F7220> Tj 1 0 0 1 325.44 698.16 Tm <4C61626F7261746F727920> Tj 1 0 0 1 366.24 698.16 Tm <50726573732E> Tj 0.48 Tc 1 0 0 1 300.96 687.60 Tm <53616E67657220> Tj 1 0 0 1 328.08 687.84 Tm <462C20> Tj 1 0 0 1 336.00 687.60 Tm <4E69636B6C656E20> Tj 1 0 0 1 366.24 687.60 Tm <532C20> Tj 1 0 0 1 374.40 687.60 Tm <436F756C736F6E20> Tj 1 0 0 1 405.36 687.60 Tm <41522E20> Tj 1 0 0 1 421.68 687.60 Tm <313937372E20> Tj 1 0 0 1 442.32 687.60 Tm <444E4120> Tj 1 0 0 1 462.24 687.36 Tm <73657175656E63696E6720> Tj 1 0 0 1 502.08 687.36 Tm <77697468> Tj 0.48 Tc 1 0 0 1 311.52 677.04 Tm 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Ecol.2l0:251-274.Van Herwerden L, Blair D, Agatsuma T. 1998. lntra- and in-ter-specific variation in nuclear ribosomal internal transcribedspacer 1 of the Sciistosoma japonicum species complex.Parusitology ll6: 3 I l-3 17.Received: 4 January 1999 endstream endobj 71 0 obj <> endobj 65 0 obj <>/ProcSet[/PDF/Text/ImageB/ImageC/ImageI]/Subtype 70 0 R/Font<>>>/MediaBox[0 0 595.20 841.68]>> endobj 8 0 obj <> endobj 72 0 obj <> endobj 73 0 obj <> endobj xref 0 74 0000000000 65535 f 0000098567 00000 n 0000000015 00000 n 0000000089 00000 n 0000000180 00000 n 0000025511 00000 n 0000094732 00000 n 0000098479 00000 n 0001003904 00000 n 0000223040 00000 n 0000098775 00000 n 0000098850 00000 n 0000098942 00000 n 0000129768 00000 n 0000218494 00000 n 0000222951 00000 n 0000300330 00000 n 0000223254 00000 n 0000223329 00000 n 0000223421 00000 n 0000240738 00000 n 0000297806 00000 n 0000300241 00000 n 0000414762 00000 n 0000300545 00000 n 0000300620 00000 n 0000300712 00000 n 0000330148 00000 n 0000410566 00000 n 0000414673 00000 n 0000506881 00000 n 0000414977 00000 n 0000415052 00000 n 0000415144 00000 n 0000439345 00000 n 0000503573 00000 n 0000506792 00000 n 0000624243 00000 n 0000507096 00000 n 0000507171 00000 n 0000507263 00000 n 0000530990 00000 n 0000619962 00000 n 0000624154 00000 n 0000738319 00000 n 0000624458 00000 n 0000624533 00000 n 0000624625 00000 n 0000643578 00000 n 0000734216 00000 n 0000738230 00000 n 0000798818 00000 n 0000738534 00000 n 0000738609 00000 n 0000738701 00000 n 0000754706 00000 n 0000796668 00000 n 0000798729 00000 n 0000904808 00000 n 0000799033 00000 n 0000799108 00000 n 0000799200 00000 n 0000829176 00000 n 0000900528 00000 n 0000904719 00000 n 0001003689 00000 n 0000905023 00000 n 0000905098 00000 n 0000905190 00000 n 0000932726 00000 n 0000999492 00000 n 0001003600 00000 n 0001004030 00000 n 0001004076 00000 n trailer <]/Info 73 0 R/Size 74>> startxref 1004254 %%EOF