Skip to main navigation Skip to main content
  • KSPTM
  • E-Submission

PHD : Parasites, Hosts and Diseases

OPEN ACCESS
ABOUT
BROWSE ARTICLES
FOR CONTRIBUTORS

Articles

Original Article

New Definitive Hosts and Differential Body Indices of Isthmiophora hortensis (Digenea: Echinostomatidae)

The Korean Journal of Parasitology 2017;55(3):287-294.
Published online: June 30, 2017

1Department of Parasitology and Tropical Medicine, and Institute of Health Sciences, Gyeongsang National University School of Medicine, Jinju 52727, Korea

2Department of Parasitology, College of Veterinary Medicine, Chonnam National University, Gwangju 61186, KoreaJFIFddDuckydqhttp://ns.adobe.com/xap/1.0/ Adobed     ! 1AQa"q 2#w8B36v7XRr$9bCt%u&Ws'(xy4T5fH  !1AQaq"2B Rbr#u67Ѳ3sTt5v8Sc$4ĂCÔ%UӅFV ?_Aנj- H>>,m*>fzp"TrKkr^r.|_&]|*vPuܶvoQ1mwVJUhu-I"=LniAƕ8"۲ k*ҿ[yu:.vUQ+)%F DHyVBk>Hy8jݹ q~9D4KRmzQ)^ʔ.J%k_tVi5NTjg!'ky|5asOȻ)R۸ߩFMԿ3L4j6dڜ#NIwUF]JqB/(FafJRzq3\G՛ ?~\ 6)6W4m[O^L0E&rRMض*C .]Unl-1 1r#Rj/&QɈ׉˩s6Rj=5Tg.y.·Pӡ:JJS:C8-2u]d&vUz;7p9 5VnL֢"y)">iי(IDDd| Yj0; LRfS:ktYK%*N2^m|&dğth":ey)uPQZW)gcC3Pv&MMWd&Ŵ۲mvTRoժM03*F3Yd6\8,\hݻ kߔi<k NTwSԪmljj[>->ptU%'LR>&EBH$MQAUx[$Z6vi&_a.KIQ{hyƒ j"JOC9eFҝfj;˚Ω<[3_m% lQ@4g=5$(J]Yc-OMq<Ǎ wSzڗ)k$7VIP붾ͯnV+卵*t]iЎD31~SA1éC2u)ʼnQn-Uoi3:grI8ؓWm*G zܕ)ZקJ}Y YlGeJ6cB2I NS3Q>k=KTBT]W6+SOXQgGR? telˊ%-Re\hѯ2TF"C/OJΩ6r[N.0{SpljjX1“jOsӥ;ҭhe}xu`Ք&.)yO̒ Fߑ.$Qw;9Iw2o+RVJMSOj[SoҌZ%;`d$blQ{Ro{Imڌ>3egf\O֝Uzx"䢸g+mv%Gʆ:|V[N'&ס-ޝ'kfE|K,G&˳98Juin/\\Qݿ̋v~Ǩ!rtWU d|E߫R4d}.qPw*Ӭv5YEcn~f5c%MTMkb-F>5JT,})QHg%{("ӔȸWMsYyWNRrkkJr0XドnͫT}r-jj,Ŕʍ\Q2Ri>v$5!]"JB2WɅ)]VԜUc8i|.jeRO6^V.¸ Q&#|ܶ-*uOG%JAtRZRr]FFG\۩w+?'zչSѧt jz>KW&ot{7P&2D;&\\>Q2JzܗAKSfeNn[jRrԕf6,q,F1tRfԗ>vֶևj-&R'Zi2=xv~Elbsvm8=ӛ"ū񕜈BȩlWau[]ٷBߨF~J!|Ipr3R̴#Yp)={7:G{+:\W}n|Q#%)7^-h"Ƒq:M*%J&$T軨I333׎g_- ucBwwjp[6i25$̏bU’ٱRv?G\~#Iͪb7<<}Ezt" q_Inw,7-d,G÷%T* Wg1"䥱kq/A.,_KhqŒxwvo u2ۥۧ.bQ}XκA$֣ +K״ZUNmڸII{.v{5z5ѮRme[moyƾd~cRݾK'j.\i&/S6f|b=5: p!6i_ 4j6=.si˧eƾtS^c.Y^RJVS-Vi3,esi08?H$GvZgg?gi䤟2adw릿:"۪lkSN>q-4kI܋ێe̊qۅgDoѨ9; #T.Q;7#~_Ufstb_'w~Xw1Xk,vcOt._}v}8"(4Z\ۘgk?J?bm_c!g{HZV]Fkk%~gEt)b秴vΰB|꽸}mp~E6ݹv;7P٤v+ri*3Ԣ|'O14_~7nP{7ZU\Vű[ +7󖱅o#:ǥŬ\|3r%TJX]V7ez¨Y]lc|O3V! R zbJ'PnGqVJ"19WVeOF埜EaEJωqCN5Z g-9[S<$sUK5b|7sn\7x qmv##FF\ w[=-43$^ooVSiXօv7iB۴yg>]Vf"r$J3""32!Zh[K%7GvNLs+4nB/B{vlsobJaҺJR:0g%&zR\ S3T[&ִor*ⷳc3ʊO[iozW٨%$gn:ܶWwFBԹjHP&z u&F2\f;ipW73 [; '_̽b;vib!oec dC-tS__$Xs]l9&z$2/N>%'[}b{h/{`{Ji׉׏ YJB/X%}.|+{(S:qz]4_Kѵo`^tY_4S#* ^zvݾMr+TrkQ g.8Ͽ^i>ӈǙvix>$o( ^qt*&t1oJVu-ql5U6jCЉmĻ*"?JT=K'O/|=Vo}l0b}}f?X[?/\JSBe,kP8ETJ==?.p5ފgbU9}ǶdNKk—_$8̸͓ۍ8Di\BԿ-1v{FF]|.^ۅ{vl12׏z7-R7wE?\nh\jN/Kձr_oBw"N QMBZqe-m:ӨSn6j4%!hQ;sv'm4kcM=!8\m[M4{SMliۇ%eֽR&N:{2A8)THLK3Zj[jPBx#BگMf:G1\`edcʮ?|w(-̮vXt,bW2;.ιNHRR#YwTM"<;mk\.foIDjmlJ;vxy7o7i\,KQŊ9d^Mmgc L*.T6tLeIuOH3SJQ3=F/ʿ<9\JM6mN6=<{xkP!F1QR[I$6ُimXu2An2yԒMU q f[IB-'䤯jYm52&JG\zд\~vdg QtHGXw&1Lw+nDEdC1w|YJmvP)HZ>i0BPβә?R:QO["]I_Jʏۍ>QKyu^bycBq4lXF~l [\*N>-J6,Gq(Zr5h]CwYӤU~ʶߑ u*SIv%ZfJ7)! FS*s_\|IŸZ)J ]ܜi4"z[+Z,MOZ))}|Ʀ(RUNIII.S'ˍO~˨rn}M)xxӕ0 eyҵ7YMAB]ӣU:/ѭ*6bcwP͵ "+qēVjŹO|GtY4V j[mLV M -m>",B$ GD1~j6O4|LxnNmqATNR3ε|DŽa[fmn-ڭ+FiK7Pcm;r5 l8r{#-]'nrFh2ruycb;pW=njRqRJ(d mnpckNnʹ+6]tz~E=ʕ l ZZ5jSi3#47.Lcfe`9؏v囜.F\-UZ:*0_<Νu9Lӵm&)_3\^ҹ3"1n1v_|uRʞͫr'iȧN_kH׺8xXrj=\МH)V\ˬ.Xʸ oVRC}ySU9/OBY먌5 ٿwޞ)rw8Ӫi5*5ZΗcGƱ !ZۄlmpjJ -l <R̵/JAպZuq\IdUS 48wXJJtcg4cI~aqߓwŷrm-v)G7yS^7H^-\mŌAq|"m9IBnF㏉9[N+mmy/!KKۉ%n +BdddfFF6FQRN-U5;Sv'm4kcM=Mn)\qιqUd9F%",6MGdT%-+~ f%+y֛^3SrF>6lc(֪vۊN;g._0Sѧ]ETWرkQKzGe9ʨsKA"yC y2\[5 rԭ7Gk5Mzw_4sM3hxЊ'oÍ5jsub )ͪ~tR2H]R͍>̋m6=%(˿(Wrr-܅y5(ܔJ޺YunW̹븹NsqK ]/QR#"ZMDfD|43Qw|._ԡSqTZBg??O Ϥ)/E_U|i}2 9Z?¹0:x'3,whǣ?C y-A~=daJј&M?D1_PS+Oi&;a @;Dž7[ zZC"bv:jjMQk$M RԸ3uA\=wI.AwC"^.{?-\NSiˏ"b}T/}q/ o.1M}R%:-ZniʒL$SgrBW*,Mw'N\ɇ{s\j]VryG'8f`}'N<*/`U숻z CwHq18J+vԕKss4R53/&XTt1bZƟo\=%nO)h$rBi-nKĪ^ ջڜlwkYm[̑+/QrZo%TQ;TLs($2C:s.%+eoNttq۰kK7O0m_t_pZ1SsSM7"mevFZ[w -FJ*T*jФQRg BSu|]g:ɵzjqwmltL.e3sRMچkSmjkmWœިm++¦'tILk*բQ D,PB\lI[9{%Gb R6öۍmX-MaʉA931cs..G4CujQտ[9 }G-xwl)IQz j Ó"rqe&=]꾧֎c)<kӳ+0JrRR3'TnXi^xMF Bު*tIL.[h"2"nKzZe'ZV/RrNYz]8죝n]Ķܩ>^Ժ]u-7^\mZjܣ9+Rmn ߑv?oꋘ?&ƪy^N4o=3-ؔ̿*`}V݁ ƒPu8%$ ݗ]wt;\y\>='OjPIp/nJU8{϶FNMsf"ίNqƹ(+ ݮF2Km |jܴZs%zf*eȫ?]4)I۵nR&FX + [jDh(#哑9q9Eծj8noǕZf\J-l&Z˫}`ӎhyrΉn\űn]9pʌӣ"׮Wt?N4_I_~54#/my1Xr*척aS#DT >q ssΛW;3oUaJSRMDgQnt:Ql,/ ܷfRqiM Ȼ>Cob;A>ڦWقM9X~/!'MW.}Vrߔꔵ!5|iB(0-zF=}okڢE$^wW~nokY߮\6՜̌{i-AF*9)\t9IV6۸5ZUF6R$ŨQIq砳YUZ]eyv >hI櫥N )&l JulwE1GDOuFN2| }馥uC1rޫV+^gdb&W[4<^e4YW,d|htͮsUM)۸8:{3d{AѢ)~ \#J=NdƮꮓ90 |1K$v*?мS ]i$J,C,SG?/_՜pMSƯM|mG1V1$~K>CSvkuj=&) -,yLjuFHK{c駗.SOua;BrSqj-ۍZ#'Jys7[g2z/.u4+XV2VQ.ޕ)$"(%)#Z7suZ%j }BǬݕe)Jvz8zJf:hIN|svO1O#IEcۍjݽ:SdὮvu^@:o^5cs>i/VqmVm]ؔܢn6'vޑ̗J4Wn@OlKbX ;n:hgJ9ŻyǑz8f܌q&Y fN0N;[69 rbׅC2/#kE l&2~èMR.*%g=Ft.%؝e8<.e=Uv{~㻏"EˑnvDѭ͜Lu3u0:U֝$[M5<:oi+V4V9 6nXvx&_ q Qqw3W:uϔ2yb/(ɳ|5zQiJ#r|Hw#.W?4aDŲ\ugWG;Cw鐢K|xg)##=O.dF˟jMUvWĻsr.z]kPc9"]R)mkfOd*uYf١RsB Aîh=k]ʳUrrZsq`d#r$/Ը3o^&lRWȍyuW̦Y4QDUMJ65ƒ[+ygk XK_±k#y:8(TJOSQhJt2.DR}"5[) r)6V6u5k:eXZmv𭤔!푊Q[qQ}ҹLE- 8qIZG|UM4j}Mܕ[Vwm{} Naqµ"ԈM zOpKѰ?IAD3Ir0'/q1itoB5{%wkOBn-ۜduqIzYK60{+DʕܞqIt";r1mG/\/ym[6JƫR \L=S=OT@Ix[TMm{>ݾտ֒ݸӉLYIx>+"JVNzx||5rI?C{oz8۹e\R-^\A2F R+N9 vlT]"ۭ d)t֞i #E2jB@׵=#/N+!ĕhx}I!cM`ąZ*ŻɄҒ߮Y.Z}='/oۙ3IpW̮hT7cTSuz9>B}΄&h!>lӵn~j˅IvU.'v'CSZw8QK3G> ,J59ٷ+HSg䧎hJdzvwv-cvxS5[̊n~ؿ%ַX?O0\6ne 6kn9.ϯ} *h 8_QhLݣ7q +=XBҲ5?[[)+F`=4 }B,sNg==u*Nj9k_GJ)+R~GSPBȒZ:(K]heL=vKPӢwq(NrG^ثϣ?#tC?.ͼ[ۅo؞y#%ǛjVyLSw%T*s92JTM%"YkQО.q)gCͲn8cgi6j1MѾ[{9h^vƘǚםidfi.^RHmg&rׇz:}݃}xT$ضk'5s-狶,\vpbPD،=Okf.c#cdz2FK5T!&)|ntD<+OŹU i-G[EE*FDfeaf2QƤM\UG_{ǹm%\yrGy:.\4wjPGUJޕUV7Do\7Vy_13w;[?c]H\$IJ,*L]3b%L{y.JRKG2sq,B6T}(#nW|km+q5] r㪍bJ@y{byz,b踊3ϻJ,'^xd،)JVw#.Vټc''ÝպWtbRؒJz۠8!o9IۄS95E9ؔ-e9JR{dmnッ<[~n${~Њ$W?&ՐY_? #a.ߑv?oꋘ?&ơ|y^N4o=3t=~7!/M3>n8W홎2M`Qx+ z qy8%]7_~540ۦ彷]Wq CѡwkďyF5Dum_}~P(5.(X,K9vᯐ?leB9;Jhm#3{CxGE-S{;@Fz˙]=O'!ɿ]' r`:7'2bЖ>Iy,/eTy/V<.H?UYY{\^#ѣr9^7?xoRȆ7EoS_&??zϾM?(~Q-K&>"~aߨ t7Emsϛ+?;fCr)fY+>z$tIkjn_>vnrֳki-˹l= t;'EyC¥|/BLwBJdgjۛ$s S1|ɍV%JI6KvəhzIlBYɒ|0"Sy0F>eo5W)O+X˻u';v)2vVq۳kۮws?UʑBǴYO漪e2MIjPAک\b1)DDؚKm6ZWΨgȕ۶yjڳ 2ضN[C[|r@9Jfo<_eI7q.|cÊV߷:i.:$ȋ)1%%)ADZCEBxJ0MJۥy(bNsKM9k43IwNt.\%N簤I'.j|ƃ2$grBEٌ\}9:v*!n7M(ɽ]7c@XxƱԨ37īf62cTTfFK]9wntQHͮvٱI/f|j=7}\_V5U^+:uljSȃY(XI.ȱmo1甅jڎIZ2>#\*:gY|4k\8ZwSqtyA!+];бޞKծË¥e)#5ap.QK^8VdU{*ѽL\=qmjnB5>{ Ӟ`v±5 ^k&O~Oshɷ,;6nOW>u6{RqS`)S%jp\ipdEBLfTWy$GIYw~䲭J.1vSY5z.V>^+Ǎvc.I[R{QsNR3ӎfhd>y?UJ*}~[e\i5U^͛E]G_FS(Iɿ]i8:4zj~շsW,ˆsy:%O}iur]iF5~3M:Ӟ#N06)4ߧgdawIotiz:1r5YDZLHBSi;NQc44la=Y kQIT*ըl:tq2(է9VO4뒳܂~2rq'nrVZŦ[t7\oլfb/mlpc.I8콚q^1iE~䰳mi[dۧw֤ICfdFeCsg:i| 6擣׋* 96lust^{%99UNRvaMܽo ammi$em4D6DD\nA%$$#}۷/ݕr99JMն[oT޲E"KTaP+HGkŴj5TM5xƱOS-k`ۛkٝWz;{kS}F;~q|~^_|euwnE'pSupUP)V]vE+t =ZRaVdG6= *.ϼnj9:UɷbېmF_tޫgHjVS'śǕًdkkѻ_]Kv?nT>)^e=Ar1'3ԔILyD?:-^in):{7.؂\.:V }#뺾.3r̸*xbFM aȵz 6SQ:ײj[ 8nn iFMw rR"5M5I旘35f^j='j:nNW.ʭocZvZKV^ɚJ.cM1ZI7E'6rg탸5oZ=[m Z`\hbMUR١Ȗĉ):Jin!_7Dй+f̷eKҷvͨBPR(V`y6tw*MRΝcB.ڭTnc;P$8nFvm4(D(R#R-L -2:FP lxZKQc6I("Km%$E, 78uXIFA$RQI$JbInG]c[ֹ:ZM+n^')JmJMJRu{e)7jQDw~%yQl}BZujSSf۩QZ+Dzhd5o%BIc'GZ?}΍:>Ɵivז-%݌J5MqGWTVʦh݇ܟ~Օ_6 n'{3~mϬj'J11OȻn߃r Qr\3y٘+WӍ'WxEs^O3 o~[|7>]]H9݇ZomT@]?5B:Z߂'`V_+/MSKX߆ޠk3?o7y:4R/7þ] iG߬aBRU&?r&/} cQߥGj2?C5Yśe7hU=?+ x龳f-܈czW^7p%-(\D4h{UK&ӡn^m]Fݢ:`δvj俜F+) y[{{ 7 tu>gvrěOj'5 iRg[ͶFjGe n~qT$ci ۚ0oԹc*jL[sVWqj\ݻ&6"WoK:cnWmrv)o>66(F>=W^bf#c zzʞtپy%mՉPël e}J.\Zk4ttt>oEM=q)hJjI=ͥ(%]脼_88ф;͛gWG;Cw~˘$4=uWdĜTثNDkiQL9U*O"4XP`02,Ge-k5$h>ܼ]3vr6!9RQPIVSnM(ۓ{>;/Qͱv{3&-[rc)ܚI$n{Sv3[j00)-D3z}MRzVQпj,T[uVs0\}Sid;r(ݝJ>æʺL&c[jPK0~d(FKÝW\m]GTcF|Iׁ)I3~#oX%vҦEݑؼ5Żv2qAZTE^..M{ʐfȏ2##.R}*KʛZz^ӞN*lPťLf\G6[WVQquV]XAi)5J!,$iJ6o$tPZc;Kjx_n3`qIelV~vLy{fn匋Ѿn%;zV.n'-ұdd2߽1bZksPe3TI9)$ԩIN9Vơ\=2885N\ p)/a柛w9g_lױo8ݷ iixJV& ғRi{N^_oAŮE6Y7I$Nk$|Q)-*4Z)^¸%4Qm [I%.c-OV+C֧R#%ѨCe3i;w$G+_dy| Fzj$DI(=OA gj%v/]8qԯNIS*֩',Q%\44ZZ%D|Ǧʴ6&vֵI$%8(ԬƾS&#Z. }6z?b/|Jl{ץv&mpx4Z$”ڝ4-H%dGKfM:sKSRWeJAn]>s6应-W9'H]'uȫYvgK^\czp|My\鏩w/ËQ.)]\QiS`8uL뚛̸=J"ܻi\å'-)54Ue]:K\퓡vK xwBqrH\*֕TnzC.mT=t-H]SČ~Nu╏NÅ3f|͡G~B+Xm[Q7U{9"~jgK Zoʰ7"qJ,ekSeNGgϳ] ^.6:s}_,%eRg<5⿨z{ZPun#jRІ.6g T.!]xa c#jN$Zpl̋H WZu8WmMRýsĮ?Mco~sx TU҆Q :KDG4n42.<3/'^?6/ܠڒ^yrrÿr2\D}}B]^E~^T cɛ7϶Y[<֞[7d}2%QPqOLEQR\CIsj1?\}%tJ0e~ *sk"*)&ۓEi#{1J8Hrt|'ܝRr8)=ƔN'RVz:cf]F7bZyZUȘ4x8,#JG̒?.W9XnO]KO]%]ƻ O5Γ/3qÓj؍/r̺rƵ 5\&m6h.xoeX[=<3%< lZ"2h\Z[&jW3ejm?k&[]ųj+{N{66leu_+lj]q* 7g*knأYv= q ەdxЬZ|%GUrQ3jLŒqET]1% qkXYūYc[7Ś]QY\jko\</Lc7+'hMSUc6qXyؙ~6#ѯv.0$BQi5YyIhɍiy=KD!n3Vm[V%W-B%swa97ajۗ m+9~]fKq|Ddaˑ0A]_v޺mM5* F-BYHJ5}q>ʉ.6hyDmpD׬'-_v5;5[8K[viJ.3dR:oYHHh9I7:۽fi+wm^ [)odPѱ52CZUJicSw\&_s0uBȍh32džzQflcd^m|7GѹE!fO5]]H9݇ZomT@]?5B:Z߂'`V_+/MSKX߆ޠk3?o7y:4R/7þ] iG߬aBRU&?r&/} cQߥGj2?C5Yśe7hU=?+ x龳f-܈czW^7p%5|Y:SJE\U-(a_cƣUǽXXKiȞNlmۊڭڄR!**ܤMeȽ$|X5(Ź\rJ~ܮ]>'HB0cp XFr_c?f?7<ukSgov¥iG>>䙗i.+t+bOjIܶ . i^:nm}s}(3>NZ$2Qg([".>i.ƾ)B̋M8+"- >eE6DݥJnJˣt׻ 5.˅nJGwZD~!i۶a,Db3ZQ3O#KO5/֍ozuK'GbRi᝘NV_ҝcvם ZoX}F6z 7e5_e:ۓj=AB+iܔERadMBq*ԯ DwI/Gy*mĥiRKg6skY/#SN4e$-yXM YL?^ĸNNӪ{$r1JJRSLO]Aqm>V/s[~i/j+m>z}eI"Qvp]{ZԼ:{vPAG2=T͡@ڐ#u"E*>C;o$~C#_d/HBq^YRٽzIKbOm\~żjFFGdiQ(*/i*#.FF]©m=BmpQQQSP&Ҫ!T&^>:y)$ˑÐFčI Bӡ-t!bM WҦŶ'UZ=}zvn~oT/\ǒ'nr8 AJIӆz<^uߖ4eFC1i+v!3qNyߕni?4JZlmYFXFۼO0B\m[ tʄU3s"Sr(NJ;SKW72L4̏BVdf^Ҹj\]ȱ۪(ӷm?J-KEmWڽ^4<8qu%9pŹW~877ܾeVгS(յe^C]yX͹! םm4FGȋ\y'Z FX7e)|Gjt߹#gb\ŧq_([R8[qU$Z (ʻezV2V!iQ,i$JE˂٩ a(GK'O{vnBvryRd-RK4=qxZJMl_CuuIz @Rt㮽޳!|68\-l[џ84-2Pu" RJ_^OL>G1~XnBŬw6J0*Uvlږ1N G1q9IUm*'oWu][&UyYZbBZRZNfEJf"+2nF~Eû7n1xv.RUM$6 lAxSQJ&n5ܞwlEói"#>4׿Q.nEq7Oko[1wg8ZQwZYiqtm&~">Bo?w͡ni2峋NCEy Ҕ+%ZJ ʩq*fpˤl,~^Mχk1+:ݕ z&Y`KLӪУDr3[*Z :(SL&ݻ۬Vqsyԭs x|iI߽zZrg.:mp%6ԜvgmpIUt;QbS.Է) ǨKSV,*lڌ|5Jt3#NP.=+OZ~/G سIgbꥹJnl_DUM\iM!֔wVZuԺ,yV.Q>f v:݇WiaŸN5Ҕ[M7SsrvǣrMW= \8ZW-jsnڕ.ZnF2qt ً[ٻޘY۷Zm"Jxr&NAfA-݌to9s359݆mZ+N1-qS$D=17 x׵+%_ ve4ir6Z$FDڗnFtOr'7'{9C˨ꤡaYoace{Refnft RR"4%ʌm:Sj3)OdInTO>X'vxV#jܮw9Fog;5.~Y5\~18YQܹvj4+~t7S ﬕs %^۵ڴDZV69R^Y+rj$ԇoJKR5wB9C>Y:l+EǎS{ʲ{T6Wi* ^^9k/y/Cs\g*qڵgn4T8mERr|Ti+iPe;;.i\EBEJ 丬i9ɧM-ԼsGDrZ>r#R>~X9y4b棇9JwV۔%m(b[Tjvl}۩~nDԺ{Zo-YuK1vx.nWuO+jN [ٮ0%"΢CdTJK-RަH"$I(*ve &҉FzB,_Vpqp9m8werv')E;o&QE׵^d9˦j\_,ڵugZȻ̧8k+jK{wmr@3ӭ2 wFkzFVqs1؛.v'I%$[iT]D5Dl2 nk7qUxԫLS+sا3/ΖeZYK<["%-g/kRs:f3;*E ت wJ%)5&+&rw*霣i|sMҴ|;R+fm䡩.!**dӶ-6s6,]zAXMWjmnz%SJߴm2UXw7MQ%<!tKys#P,W>s;3IYwx<+i_\\\U6 u7P|xbn_k&ӓVOe䦒 VUr,-㘘"-LZeOSҠթrEvq8Kf%5%&K"#%vD/.ZYYŏ+p$nZkvއuW9㓱Z G wYIFyf)?ƎUm5ԉ/'k84{KO:rQI}XRuԪ|*lu)3qZ[mSm5R3".Xcَ5c®ࢫI*۳~wRϿQWޝ(EJrri&ۥ^ʶ齲Im|[yb;mnm֩uiܘq>E+Ikx߄3r33-5𹻖09ϖ9[Tz~mr5NsWl$oPusޛ^{Z;);sڹf\3oٹZmԉ/'k84{NO:rQIBø8Bݱ3n֤DiK4u& ofSȒܩx<˘|N0Fչ]qsp"}! QWw@t4ӭ+cO5%]'*{eM߲DRO1y*q8w++e!c߶ܪlZWّM欼 CQ̼빶lX{vib/V/ ai;x6~]+z]MWB>re-:lgk}պ!#9?%܋V-c[z!W?c7YNm/jRr[HOzԻefճ0q15Zp#rkQQ0tU-AmڵP/cȕ?0cZYj;:0ZM=D6g ?'UN+ձ[K ܖB2'xq9{|۫N0ku 7xaj;n\ 2[VznMlWiKbSk))f..)Km)&bGZ=>OR܍W:j'rM'wYz&/鶧{Sʵb"vջq[I-ՌZH._x*BagC'T(Q:$ͳQcMCKy?3g'ߝqnT);qs #ؤZ}OOI:cfnc8W~qy.;^pVl]Hԓ>^H^@7-AA܃nmL(uWܻS߿ Td95Bdh4t6*dDh!EhI[iŨ\L.&Nc ܮf^;$R)\rip9I|ٺ?#R.ZDZ;/]nݻqs\QE9M&Bd ]N mN*D>tgbK>+ˏ.!23]BȔR1ɝ^j'k2ƮqBQq[$di]icV/e`޵B.FIIJqbi>Ӥ|p; 6${)RU>_e}^dzdfzi %ekRVUS?6'hׂ)5.\+qUgzE2C˷ecŏ^֔ibk shesFWJ#~> Wk~ݨ}ڶ>ơǚ)׽ZƉo~B-ڼrvoE:Ʃ3ۣK7+Y`WirS):{>ڛ}:wԨ(J_";6R%[u&ƫdZ_\'np| RJwNeTW,=rrbnkڄ[M3ܴz)3- R.?:okۼ0TU'w{6&w7j1z3ON'fGoO?)S_bQ_¿R(^ԴԴG.EtMڇ&RUiW uQjU> Kiu1d<ѥIQ'RQ1:O/lŗᏩiʂv&Jc{D5 Tt)1.n[n۶X}RjqnOʽ(~[Ns{ސ⛌uO,kgo֢dRNQȄ .'6W!׌P朼tdZjFGE"]K@'i۪N;sI[{SOzk>`rRR+!σj8&TjlvA̷Q?HyjyLHNտJMjܶT۽lG?SnKN%<‘ nq[N0Sq[Ta(&t(|HGO~gvkݻTR4&Z$#ViOY1r$6YF?e4U/Mvxų:zbU^gQQ+NW_'4jfz^c'#`rvrڡ(IJ/J ݦ6 ]-CW |_{v*_q3^DZ}Ic6Uڌ8p7{crZq5ki`)mU6|-Z5^iEz3P=:Cu7DF'k%}<C-޹ֲ̱#\,(f88%X-N(ck0VLR~} G"-8ӏ/ϰKq?(#nrVTmZ;zióM4 m |UT'C^_1X.gXM{%ʤd 4\ovN":"y-,T)fLQgۢr=/CƹǨJVr[a+!rT|%Y\ٱzsS>jͱ.oOc6f$q% ǒGo;n[];ߎjrk{~\VۓNIGn:iqxo |~t5)Rxעri{Vi&NUOl_ѮMfsޕkЄay.0P{7N((BaIP$ K"U6Gl ݙqJRu+qN$ m#*p<|{:>-Ev=86N*MM긭U*uѾ?/^o7;'u,h4݌xښRM:5.(/ \իU.{F^rmF-Jɷ.>Q"[4xT^OZ~mK}T0ݛ^SAo9u?lX(' qj%=X}"^e4wˠ|rܫ 6I\Ķ;Ӻw!'ڍWg{ i U_9Avhۣƾ+:vs/MK[ɭīe{`Zgb}r[i'GE2J7Nez579wRq+Un ]J.cJ4M:h箽Wxxm^ pc\wcN%'My $$| :$Fqɏ¾^қP9J6Wxvu}ݵP>Z'FFdg"-; [¢cmWkÎT8nG%ݣ7*\խCLRYZͤiD&J#'ehbSyXK|y*ӞpS̍R`[pTr/Eg)K+92{_ n3zwz'oŸۤ+sOj J:`T>Cf*lwd\fYOP"R E֢̔L4ɥ :;.b(B02rJ蠟9>V'9M%)IqnhP<%,r'P/vNSwr#w"ݨaqc(|{kd=^0jTMR2ULNz|.<|^PfY22##!,K~E BEJۜ&jRNsHަޛg\r,v؜.jK3)[EJ2ii{KEiHP^&]Gn8x=K}Wx/KI9-ϵwQ%spܾ[^R}S3$qvq8M[ ozKxcqmJ/ӿ{_}7&ݨ\f6ZSyQz& 7ۉ[8~UNn|nkiTB+4RI8'Nc%tn{!]Ȋo.nEmʱn𵵥J A+wy#+ikǒڂ;՛s85'KmE:Ђu""Iģ5p=БbTY-ͽڔ詻ngL2Q}$de# fs^o{DUUsfwӶ;s1T,ǤtޒQ\෼J=.tKU,7čJ5 N$y3kdSMQU~mO[03 $zAڟsF5^뜞"Կ QHmrR"ӳηer+ҔZ]hE-6Jmt'ޒ=O[sQj)6K}?e4v_KfZheޓ=BV[bY}lݒTTЬ{ȫvO_qpRApVŗ 6ju=*BR)g "O1yhb=tqJ gtm\b3RY+JQ^Ō֍\յ\>+uSi{=x ^w;uӘ#ĸzLn*$anok߷CBӷ}5Yqvdž<( "_OWit5:EZj2 B ρ1̊fi[n!HQF82q1牙nqnEpT(2RMoM4ϳOu ':֧_Xjsg jP^(ڙ{2%E͖j^}ZU[Q$'U) <܂%!s"m R'G5M0<+zM6qYm$ڕ$3ǧH]?o2N<8F1̻r_my[Rf59NjpzBnl7*{.QP 3N&^BLJPjAHCK2Q}$#~YMq8 k(MFMU)8MEqTy+Tʞ-ar5yܕOXw!e;q-Jqܶ䓊Y:LC UE{/t>r"lI9)3KJjϤA 6SEE$d߇3KG*En|P\ԭTn6I-ƍKTj<1H_zwGr19wF N8ݝ+a9ɫM6mhePi%mmD! """"""*1bRKrD"vnrM۫mmĽm]ӡiG~e"˩ lhRTMk^MX["Jݱk7_ޕ*DqĒ&flՒ}`W}~SմZ{ĕ~wm*/{{ѹ_-0ط#P]xlڱ~Tn5wi*lڪ (JxioϏbqKYR|!|KN53 OS222$jzww%i}>N)E+rۥ7c$Ofl/LNث\6H9: FY󡈾I)fB֔JI_ ֣^: 9mY{66㒢7Uj]:.-os[R&gMF3˸#໹kmjq^8W"PΦURjʄWa˧T!͋ lW48JB2ko+ /Nw QwQzQ ے%$ޓ7^YL|r7!v%Trܥ &|M8~ybrn[RV gSn{{*#2#ԽᢏӴHak" ӌcwҜw&RJ07ױ>Ļ =^ BɆ)v32.M1=#6%̠tҤnzqMwԣ~s*%-j|_m*.Yx9Sz=)qE4 3pk+,`=kNRڥ=B=nŔNAx)Q$ԩȧ4z3t#Z2lҮYn$S%y- JzGpu|LBV7ZW#;Wwipܷ%(6jFG5#{$D"uۭ~]֫SrD܃fҎӾ+Tu>-ZTQ& N|$沸ii>eRWݳu'[O̻j8JۻEѩ[]vni= ڒ,[_%kC7I3Nv$4ɎЈeٸoUu:[}Do5|zNq=Tre%ɧ6&~DȍF]ƞG5q m]/w/ \ʲr8=oʔe9U(W"|S]uZd#?Se[W"ֿh][-7Nu:T=)R}.;ml*5Dlf $fF(̏T hiIUU4Szɕ t(%_|2 ~6eM;TƗK[f&]LK^CE2[ȏBOd;Mi|cx,^6;sیGpQ\NuJIFTJ~đArh* B"$H쉩eXPRj?sl"ԥ)su]xpԴY%VESH"ЋJǰ K&5^Ukzׄ8kEgS2h&Se\ Yl]WҶp-ZUvi7QS:4byqOo+[̺腋[6-_Fo.6[7$p&^ _GZԸߍkc.qqoI[9m߸YxOZЦ1uoiSH)P9Uʄjcq= S>֙NeR><;+ڌk%_qT].srNO?s[=vH[]RZHRMtᩗVؾ:/~u)ԍdg%=edVrISb{6vSu=(ܥ)mTv/J}̇8 S3ad:^hBSf؉OɔLhI_1d8,L><_A0y3rXq"'(۱;mFNII.v5_(^q~X>y{3צ I*Vܛv/jW' T'NR'j%ꔩ:mJ3SB}΋!-H-RJBТQoedi9tjENenPpke.%4]#{:>mkEɱdYWl\\\'nRM4&U>?Ќˉk÷!𴪛]]5}UqG~ݏI"O~s6(Ļ)qO~h}uԕd}Q~G,oE!&G&/]_H-O=o{k\̭bkv.Ô܈+;arZx)m?M\3lU$mk-CFXjTv6u' g:Vn_*qk:VC A%'4JV%EY)#BғO4<e׿jQQ]yUr4=wm[K1r׵%Iũ-O}|kC;/VcݩWZ)EHdžTru]8hgĵ-;=>U_ InvTm_jBM+QiF"9*{DI/iuo(=TzϖmPQl_v4z>T*ȴ>YF;ε\t]EH4ꌇ[VrLzef 2T^V>g2~kg5~Nק;{~Z~W}&ŒBӿS2$J?~(Yœ"˲ߩ\O]: J׉ښT{mmIѩn3˧)4LdFZ/zUG>U> n 5& ϴ-KJi2o]uKljvK3$bԔҚV旧iY5.ίfi96v7!v))FJM4{jG~Jt/lUE%pTAFe4qQk\ve۽/u/Im+W')v{\-E|Pms7߮DZRr۞/mu*1ՙaB܆ -xg3#6ۥtRogʌU)׎]ZҞNnŞr}F1Nnޞ;cZ{N}ۿMiuxʉ*3qi'9KHQ$WJxXyرŔe~[v5~/jN9Q4o6rJv FrdxM*iRjMzUinHdн7ᾞS=S'7 } ̽zt7K|_g J=Lq+/Bw_\ۧx\HJUPzQ<hqF[V0x==CsU7q|^ {)Iq38$_A(VgcKu06Ƅ"%i~_ˉk QCܣB8Ku/񋇵u([w}$F|8TՠI.E !;RJ^}MɒD_q2];Ɖ{5}*n7nEInO{Mwv}&q+v [V}Ĝ@%>#dXQ$f;iep.GquixVt x6bj͵mlKقQ[T]zs/&yەnM'W}!Fp_d^Tu N{ɻ'l{խ2.sTu{W^H&;1s)Pӛ6>$mě;Łnj= fLT)>׸+qReɴ[UR\L*P/!$Ӊ3Q 'K=m~6XqW3^W+ųO_[F$rR*u"T%@O +%# ]˽!aܽz{ͷvQh쩎]hGތ5ɇ*DzJDRNLi 4:{~2FmXY-zzĽ^f=]uū{/+&c:Ma{ĝDp2m܍kHș/(--m_vݮK(V{R}.k&yƴ7i^4@3f sK3^Ř˸B=]?gt5KbZB<e;kQLpxuWC}n 5ҴepB##~q= `x]KWF {GfŲ}?G.I9pjWkU]>={7q{kO/^I3==f1ɏ%nnʫ/Zu_yXN<57ۍ'vy/"8넭M2eԷ&Y,в33%IkjMr7xf nmQkX4踼>a-GcIeތw&U=-:qnW)z¥j :WqSZvԒ#j"KrIU)%qrmRoDGQ~SYRsu*V)  ,/x)MFD6O#]z 96[Ui(JRfw'y$GeUީkdMF-ݻ98F2d[o{Rn0n-xsV6Dh|Eb2E:KCOӪv4SJCr"J!!m,hRLD| ZYFm/X~ΧfrN&4Ƒ=Z9Mh.Mܵw/BdrܥniŪ8ɧ|y%œ[M=_tj?F!z5\evM:\ ~F-sg钬OWq“iiȍ<Gi%%n2rqͻllƑ)okw7}\Uk-:&fj솘XerV9yZuʼşdFC=rmo%~ZN78X(N)_7.Εn1MpJ}62jjJdI";R5&iLԸc:jmqiQj$ujp\{;v5B񥍪Xn Ą4qOERjzN(Ga٠䌡)p*v(J7#ZۻZ8O W uONb+^Qipv9GvֽƼϯrYƖKGJQDNPhRJjᡧC"21"9ѓS1;R_O7/WGz)8fE%F2ukmvSov/iZ&/]~KmI[:^~ͤ\kMi稜\ywJt3W7 8Ʒ~ݥeFgѼw"8VVSج\뻆}ݭ/J6Q)d|)zU3>k\L=;ow֯gN3pKѫ|wmkZ$z^2R:E)f>ς нd|#׆?\ǔpV{;\$ƵE%-ͪm0S6[n< kE[}mvE4DDZ^$OZ0*$~XUv҅B@^?]so#%ojw;Y#SxxueBگy v^i-)s)zV jC{7Gt.w3v,ygg8s]aE_,*E tY5k٨h=o"m泏:\6w噓aiL׎n^c\75AGkЯ0Lf46َ`egZ˓p/k;̛]kq!ݸzpԭG"}R9Ve>ˏHUjJ-&7nrnwG*Xv\˱/vN}O)ʼn&CV͍f̵]r\PMB-6Du-#RͰtRN^)mT _}nSȕC*_xBuTkJW[`ɩ`ejvsngP ڻ.-WUtܑqԹQj)t;vN&RNũT+8%IXӃ5fK՛-d9 ]CƑm|nZ-6=Hz,*aEm W3VzRšdY~Xf׀Xx"]s;)5u*ُHB BRGS6bݶؿ 9j[1*jױga7oX CUI%0v#~\-O-Ꙛuɷ쏪&5mY٦M`LJ2qK~HZbr =N'YobI. (^ ׾{_ ?OJ`S`3BN[}5w6:ǵ/iSlt=4F*d&T4y/#. ɵim5Uֲf 眕6Y7 fơ=3dϕq뚩$qTM-%r!$@A? ޾V0c~{[{;򥧅a~ڵ»&ڄv1ek=wb MLkNAԬw-x>~/r=e73VeVN)K%Sښe"+3uXuچrn ֺVzscJ峻m}vb㶓n\YbIUBT%*,0nov=;z꣓S/nSXSpl##k9mXGrZv^Gde!ŷRԠzQyjC]`gToPov{j~KRBMY}i[߶9KL2ԉO0K#m>wB[ٍ+n[[b٦DX ݲpo] [\m5qdT()mo4Oy9Ie b][wղmM~vmi۱~t \}$яimRk(L c Cvk7r9_r1 ;zv|F@KyZ[&jEji/"6$69ml#e]9s\{ScL}Ȣؿ0q/nZ*t,CLoD߉Njǚy=Pgmu6^]l-["çUʖMlʍp-"qmU>۷uFOJ%Ǔkx 'g=睋k[3u,{³WɘݪF]ՍeFX"Oy\,cچ=w/gn Ļ]#2? vqy-gXnR.^}ݺFs{ŝG]}e|#0mjx"ƬWكm?rgU^xVB":Dt>@LRbun~ݭ,w+v⪕;\U(RYa61>#Jm˞Μ9g9XKaG='u8gf}'qy#ɉw J]We.ʲ-<+&q%s?2dњztҼn`cΤmmqMdz O[-ߩӲ&;[tmܝVnr">{x<8U+p:Ig]zjGkt,uzf}dؠoJaکqEq -(:d<պ=eKy[˗^%ZXkX[C2߱\ITTLGzANM￵i]K>UsOGDDD.ZF6* ҃V Zhz{'xp^`wo8r0h ZmJ5"jb[l=yUu7-;7IT%:jFjߖm0tzU'K)څNۧYJ)4IQ}^KWm7kSP>q;ނ#)'n7&׊r?óM{IwR\j2Qn[v pe#/tAF\ϵ225q֒om6z})6҅*oqDsMf CNIN=T S2t,_ѧ}kveMF0J\Rnnݙܹy[rUc-j{yGtkQ%s]5qB.Nw.JN1LvR Ui5J ZESQԙr):MJ+g}χ!2;q([jAud][ljVK3$ײSJI=/|&tl'*n۽f.frܥ jQO8>&Z];.|7T/C}$ڋUmP2Reҭ8hFF\L 3~e v\۫]ݝNmrnB%*]Z«hKc=BTLG :V74$=Ǘy+EX'4tn(I:Ѝ;Df8c,k1%dJ6.j6ź{N~l6&*fœI7 WAlGOu-ҢH,,(ǔe뿋쩨kM܍ZſgRvQ' 9)?n|er˭|I|-fGK.rΛp8XV1%K6mvG+tc+qE&ǸC_Nm:l=_/m5^[dߌڇ.c<%:)tQ$Ow~-aY;UJ>=F)2[nk؆?훐M=l6[4(O.]2#-H^n#->&mp5~Fӛ+|| S,xag%qkEUzUgæBhߕP(7]kFnq?֖CpruZ6*rEڊtS|*tI*E}7R<,nUU֫^I7Q*mSly%rdȓd8hE<9oHhMfNSRj[i7D[Rj݊+kდq{"$$H?p\̅S?㭻;t~R߁)^/>Qj`yt[w ԛ;²~+ߔ_ YW~|o]?x^ᯛ `ʼn;g)T@vWn]>&4lp+$D̢1l|ȨF%-}.9[}w~ ԠLM9hСablfe&QoW!s?wjLK?s7yO>(=C~_nyǜu?v3vyo oI@qV-jeES^[9WoSܝh"l2C1a͔CiJ@3:Pճw=/7ovuk+\V;lDgն<[A+rX~d;m!_s8ݖ׷;;.0llUC+?i#_crʙ1~C.\–q ul8Hܶ2m`ܻM3Tov|Bs rɵ"oLS- DКw=Tv@f'6|YlD͓Y%׵-#Ѯo%:&!3o%\J<02;K87>^vgƓ# ;ݝmz^Y6=PS39U%~ &f# }o!muH;ʲŇ˷yvP+&.7e[3'vR4Yj̗IZ`e˽3o[WU{ m[sUbۋZǾۆl6~9'V*.\S2<Sd*zY[aŶ`]C$n.v^Ʌ dng>ەZ,Mmϑ :n6nϦezWqUJ4! ۇ4R! =>>Fn|Q[{pRO17ƕ~._I''00k=b՛o}Osðc2'o\3}ݭQ^2 . R1yKȣtAݿ-uܾw!`?1Whn|gzUo[ECWwjUIן)^h#1ɭ!/Z np;o;ΗŻkXs."6E`Z1 עӐ9Kl8qd q} 2Stt;#j>;խabONŗ=fwP1j)l6J̶|gV2`y/0E˛6+ԫ1? 6}KW c\KoKͨ2ۅFw–s*TԞLיuDx .kCzWXhy۶gLu|%TnupǺl-S* PRaLnT+c+*xl.v!.U=|; !_L̎뱚U=4hm:ٯ"y)$:>%(n}X'p[ȴ ^˒4kƓmzDx \ 'NqamP7nyN݅=j7%McSڵj%STy qXymvCg{w/w=wSW5r̹u erծˊsOm=DhEҚRb#n)QOxtվQwe]I}wCa'"[ۂ-z}2UuKP$㜉ԧ:mc<Ý>RoL?wu|%ҷ&K y_!y9 ??:tq3(UU-lkS'ɸ@jdzQˬR] EVPW1DJq2n:,c|ǻ̑;y{X,ۂ.u.b˕u.tKBjQ"[S園S`ٮdNبeJ&9Ơ ~0a(Vm٘L+Jr*vڑE( x0+tp˕ n';wm-ޜMOxX>{#2%jgb2M[`K*\5@8l'e=0u+w ֘鳾{y܀:R*Ya]"Ӧ%ktynlۣ65,3gU}{GYrb;ge'TKwǘ.,rpܚV]Tr,!dp /ԺU,xՉ>s׽~W5oTh yx?xrrx?)?ilbT׬,z$Ԏ.UH٠\U1pU:]JwSrGZq8àd驐,N67QYBӢD㏙W!Q25ϸo9ms-7-%3CihO.J鯽-;MZM8ku-7k9S$8]q2E(}bۏI[DKOK}3KUB^u %Y,u.-&f#]'܆o$x`Yu,dzwM;#oKxn;\[d7}Rb+*Y䛂ZuBӱl{j0O̓}LhK;[aֶaGL{Cb#S.T[>߃F]NK"u^LUʐ_ykW?!GRj29͖qa'0[npcDvV)qz9R)PۨM^aJx W] r>];eN3vxdmĘ(5W2K1䪖weF{mE/QP6\u54x5[hۮ-Nk”i[lUgL]J}5 S:EhiUrgHl!ŒJ$pe=q^b͵Q' ?6|R\,JA ڵ"TDꈭ:ymg`B5t%M] <N_zv2_Ortٵ/i/ReӮ*7[qүqEG* m"[I:6e^p"I$jԴęh!m)]GZkcjS!{e^z}+Cѥ9;R|/ֱeiUԏCNu2Zhcٗg$ݭwvr P8*7/Lk~I'Km1+MW%Bk|oOm>-#qj*|Dbѱkn|n{v#jĮqNpMIUm(7Liz;{ҜݞڝVƚVϬ+sO!OstGvxӉ']uӎ4g_ 1^-8ۦ k!)Ύ5O;YSB#2Zzχ;<.ֵOtge~.(RC#wFZeGZٸ6FFJ4e2ˇpJT$[wgV)q6muDGJ56q\I!̗ y/I~RtJ9kJ]Iy*'FN0s.[l!fw'y(7$œ WƫgyΙdMEU JQJv̋vmrۖ.jWR_M֨djYgSj0^\y'EoECjm$ IƩK>Z28J2TiJ2N#}.s cArl嫶nB.FIJ.)۔\ZiM>/hLĸ=C1s[?YMqp|94- 鮝𦔽/k^#NT(Y LS$6˩}{;5 )B۷W$qpN)qqoot}ZDVә;7TiK|6f3h$dԄ}fqݡ>Nb򗉉+ͶO]>ߡ_VtYf79ڰիF sq~prս|QM)g%l0ocJȨHz V;Bb/kLAcfPJ,ԭ{ƍgpjNR6VSI*$!yV足jᇑ.](EܣqM\qJ2eZT).<9UB/(B0j)mtKEj#׿fDI-=rZړj|'Nڤ]k*i$5qt"ݙPM6E4ke^Z8ۏhz$Q(R Ay2zfRñnpnkbkI:=j &ΝșW?׵d{+ύM'??XqeeĽ.[o=UxFS=ӷdZwenՄ]_X=ĭVa* pKs0ބۍfJ3 gz̚i|wnxtjc¼5${(1fXQ65ȼb̶Zkn>%FQMJXӡ{TZEVNᖣimT/37cNJUPnP҂ZOE~"-Rc4^b- FEͧtf5[)S!OZIښݲ͑;tvܡ+N)AR=hCNn;wL16-:特7M$=Tҕ-.R[HٷnXk sn[ҞD-0WS9p9:-Ϸ-jѬNu{ҹfv)[Ľvwfg(ٷfe+0mYj8Q1\ݧg]Eǎvڿc!4#j5̋C2"}BRriFp7=ô\TZ:\BLfj#I22װ<;صZl j 6:l"6]۸ K'6RTѯ^ئOԓV\?$x7s#r:Oh{ց=MmuHԷd{pN /܅:UE#Yy+(SgQ(Щ)RHzw>^Ѿݻ>mK&^ '$Jۻ&w%F|xfz%˳ L~3N?Cy9 v w/{ƿ kz3x> sXv}vP"@WyC z`'톽Dw%-tt yVY\wmuPYQA0iG-2JP,6/gˢ]u.-n!Zw.N7Q]Df}Q0({a\@=i_X7gFǘ8^⻲}G MZ1)WEfO12G+=-B@z\`||w6ċj߬m}UwRox֢I &c~XGP6Qndpvܻul'V7^FJt^{b^B(L~sѣ6@߿^xqU!ڙ5|Vpvef-uӥ^3  FSDɯKD%0r}FF穛r7 +o"V8tv̖NQU!5uFd"bCr^bJ=֤fM#ʳԷP0O-9xRBm\=`r-:;~3Tl(nXtXi%2Vٛ#vwqƴ`L@"H‹qW.j,JM5B[)WܺUeZFqc'V˷1W7V̾-MHФwn8N;HPSdݷC7&2j.W\τGŎ'Vb]c.x+Rx1%C2T{myg[qU|+m:M:շ8҉yWd)ՋWS%%:iqlʹmGwݹ WnNŤѩ5(9hTٵDdGUi-)vSs2 2{OnT$Xck n:¶(lASLeȔBjμPpTb2~N2~%^k[ܗ[Jzs0ӓHBKq[}JَA-$dFQgjxxFv4r/x*Rm% `4J(&iv7SkԲmSH1YWmx 8n.k']:Z˭_W >ڃXЩ. jTq%Aā[E}amc]D:rmHRiu:uӚӢ\p(5-q%e)(۬ҖȽIf<߽pr&ݫVfY91q2ĭEQgYbTGQ&,yL+N$[q*RVۉQ=FuTܻ>f>f㋳8N6$܌n)9&»iˤsX,݅܍ȩv+sRTpO}d?Wn/Inpȸ%O]StQO|v5\}7Zwb.AIVK^:wb{[uݯcytO߶S<{8KSRׁH̏N7ۚ[xkwYy_'ZӵF+>쌛ZUĦreE9F[24De{}@:ExWs-\ǻ7K-\JNvEk%:s˙#κ].oͳ;լ7wB6nwu:$L; DkI#Wz.:Xp(˅v$Sq,wn\qIN-e<5Oe+vuYTpcojUI_ާP8 O 7&VL8z$_B-H-[uh]T{|8=qVRN-:Ij:7PUtXϷmy鉿:RIM~33ӸS2#׳GdŲ5+/Bx{(WzȨ5Y㞎#|˖+ ط.|e<o/rߔX>7s}VE.OVti׽ .5nNJO"95{#q}Ay9do]R"M6z\tnNS-D!@3N_jicWsy*5uٮRcWv/.,j}=S)j5C^> Ie =gu9ӛqjtz]۪TMoߧI!Ǧ¶m:,"[L!{qAv-o 3{"KʼnrIkfٶj2ƙ؄S`7` k6jzޞ?e5G&6uʷ2%ԒRKE*G\Npom F/V |C0.q_eenƣ<5Oh'67ɪn[SĽ{ڔjǘzs;~׌(ۂ`ܢ1ƣ` _l9Va6%UQWh~P~\F^ZHR@:ۧCJ{ôGeBh;~ۧnU J\O+n2 RҠ)ng}Kh{5+S×ܛ.1ZjG)iRȤIN 4%{oΜ/eO[Nffd ĹK?nnԼMqX'܌nZvq<ķbFnͪaQ`5 s,M_լ?-@_{w{ӺձJ}GF[%v\5[ŒGkOw/ΜM9rjË%2+rd~+󲕛C9U۳r[aJǭm|˒LAʨSCq[XMۺoubfp:t+ΤĻo ][ zt-*67kvS7D·MMCQXm;)܎n_h%]4ܙnRk!]ڵsDUF"`R, &#R_*[z*ZqFXɻ]7|۵w+'pFDەs=r./ᐚm3Hשy yD"jHCr':sA65نѮ^o1V/ f;nFr3VM)e*- s D'H݅fӧ\*޷[k<7u<-]֍Q8R h|p=WlW3s%Q %3l}@U-K6f-NϿu|ڴmWN׮[׸F*mW\%r! C78:޳vBG7ŵ.JթԚ2x)ST!řn~9 W:Wpܢ件{xf8ٳwKE ҰWxVB\qBZ 2wMb[lGSnyԚ~z9ZmያvoN2Afnݽjf>)j3 !;gOYʹK" Wftڎ+׭b*2ϻK>ۢӱeyԪXISUm[z+ugX%0lϏnvg!;t{BqPj>PyvR7Cj]O%+ݲ :qiMj6W}3vC/R=4Som]ŗ=ю, TF6U_-\6MyskwMr&Q\wjKܩyMϣUj0*}RZܷSdY3>Zjqj6TgzpA/M`/Cmл,޻feE[/+uk^Vs1W$G(JsW2ٰu*߻q*Y޵.Wi:ur5T),=0uRmho.twܖiYwrWHntvEj8qhf`Ͻpf(R&>Ki%I7$QӖm-2 ~yߗQ-앑/ x[k8nw.c㩵k}]FkbJl:{.(˩n0Hqvαp7 귎.Gupx[N`Yq'+ruU7[ү+>!xrȫoSo]OC# d^Q]\>!ƛGw^Mx"-+%vdX-:M2UR%d>%l ioSu6lsj7D P>XxHz Ukà(n^Q V>5cVtWj SEiJdznyej[lE' 3kuٌNn4JW)gB {4 j6&]' m-(ZMEz8cz>WZ6#7+[,MR-Z!4ܓtCyE|umj1ƽvƷV\;%>Q :#Le(iVz5 4ũۤUWxX ^(ҔsլB2w-V ^R+; ˂M\z+Uwr+RWY⺧~ Q*JcYSNSλUd8in=v K낫k\IRרSUaCFmϿ5̗P|u ZTԕ}>oYѲ1sfP+sQkX8Gb~6r,s>^\,mGL+7[n-E\.Fqḕcl*Jmjb5 ,m]c}NXfeVlǸJ5eˡ$4%g~N p4Y*WwW٧<8v#;qԩTut,m"#Y D\5V`\\Lȋ];LȇiS6ϝZ l>LruR\v=ǘϔDg=ԈdFZ+M{=|,[;0>RiSi4,S5}yxw&(E7&fݙ4UՕ! ~'Id)]ǽu2K-fޭ \08Vڅ쓬=Vy^^ IhyKR-B#Ըr=]mܻӾ'*Umkoy rTqT_i,/8Q^<ݤ|4ԻO(܄"'5N~#m.(Ҿ2i6Uev&I*<}҄$eNtÛzyWJubW^iBW.܅Wڮg]irO6Ve90sgv.+sV޿aޔ[p?3q*FutUo*eL\KM'EG*ZcAFfG5J 5jj=MJ3OK:k˝'NMB7m3uFҕ\-Ywg%PRqMIyZGY9|μvn߻5cWݷa^+X֥vnݘ\v7m>Fgzv"-;Ew֝}1|RjN𿊀7g#֟*GQQ|#/bo]p$>_Un9гUbn9׃ErQBU-^vDmVh'<R[fdHT]*~}3j;nvjc7s-rӳ Y8[n[1pJx kX[Jk9Mn!_Nю6x:iZ˦U |߉^Ԛ݃hYxk &U^bwKk.[jE+P(˞=9j@snCv7%c_7=xǁ<l {t'酚+1F‹l׭:ݻILruǶkL-L(K0L1&>wXB(pm;1fpnlp֓%Skidkt(U +xulo'/ڕeN r=^pZZ:Pnj8Hf"48ijY[ N[yZٻ+=  ø:3 ?^ܷ^Sr#YK[UF?CuhC b]GM')mڏsNrܗI]ljq6VB. W,UK"YX5{c >Iqā> T:n!,5l2VzCl|+I[*SrjnS6٨y+x,@>П.g+!rn9>N|W>OZT_ut Y""v7|sfި;Pclm EùN,{'fNT%U&LfH8~1v>Il}统u6P˗c(WV~H^bMU.o*oOF0N:_:6Smr_.b+|ݶYY غF,mwjv>f*>QM뭱Sd:`N{l/⎱;n-z~"Gze퇎J5S KG9!Gn;N1 ݎ h6m|S?ɂ5'WOÞ 7|7^ao @mxGmi^jϽ>01Mf0լD3-2T. VXR"ɥV Kl J O7|u?bvа;6.eߓ|[1bmRr,eRz`z 6܎-ͨku͹Fː dPhYgZUj}nvX;z=gVեTv_J }\1n7w2J?ޘγc\E 1Aޑzq;\r]]\Y&[nsNei\uURje*Qk2CSl*xJz-xٶlm+|UjUؓ`Ladqiĩ!Gd\W~fz;Tn*PdRM&T4`չSWq5k훶(N"Ӎ% V]֦wb.nUO!u*J&Oӕ2e|Z=eV쫚΅g#+/RW:طnbi*Wyo)p{:ETKؚR(RY+r웓r(IF) VmȵNB:h Q1ғ|u8E]{,'$-TR[j49l*3"I鯴zhd>Q+\BkNF=.$ZR4Nwհ(IpNi.(Gi33#33e$FXK*NdWrud[r{xnk$v2ıh+J1TQ[#JQl[tRO]LHKٮ NӍnF񨔤֞Em'MILB"ԋ%dBŋ+p̿_17jzT~4pc Vo\ƹb9Rq-'1j;8ܗ)hE%DZKS<璸Bu*%*Yw5ڻ9ۣ^z4U; Ñk\U(o~G?VUĎ:?P?_F_Kߤ~ᓾI |pr.Ok\SklRhҪz{­P .}SktZ7UQ4ڌIM8̈eaӊJZ%FFZu,KZvln廐SNFIVtuNi?CM5]+Ph,{jN JSR$IS^tSUVrORYu.9WyP6 [Kiu m!X|]Y79ӄ)\ģ)pbڳr%*&ꑶ_-H*dzk)1 V3')UAϹٶWRxe'պn۫h7AR9 EAJeGLms!%D| A 5]/Q3eb̄vnVn%za\m kZnv([emqrIҕij|""><hjJשvvǕ|Pޟs}V~2&Z?+2N&Z4w@)4iSڪ_>/JN9Hiۏuf8'It[ȲR.hZ$ȋ_Y ~U<UUO*6b)Ovzڜj\R̋.$FsQuҊj^נ䈈y<zZIuP[}Qm=C?zN(Exqu/kn S-FzKZzOסӽjJ\)F3b!r5ٝ|;6 o=-3*λ]αb\abqRi-w޵⦪~b8Kpo)Z=>)ғ"5/GTZLE-輵f7ݘ۹~+&+w/7GFI:l33fg.N~۲\2|*cnermnnM+Fq"ѪIz%j =YW8@~gc/~?N'?)«qȸs➟n=k" X“m֮VreMh2[uݖ] *FܖN)MȐ`f0 g,C9̑o;ddudJ=In13:ݒvvdMUEJLp^,6t-@͐9'{7m{-3,>hnF;ѰM)->>+Ěz!R* :`e--m7nB\u{b U>[8֪]6^ߤLʦ\DFNo$$dͶlgno8OrsQ\l̯hRo8tuNo+ CTxu!2[>ctFpeޓƻֶR"3QrQuOѳgwQr;S~)6HhZw/GgVTmUf_yt7%$];zLWF̰xy2Ʉu!MCmš_0[W6jf#a-KLi+3Q7c^qg%s<1aYIQeZf+}>;S6L0]Yu_h9߻<ƅpmiM$AVvŚ,*#t2.8Y)-Zhshü97/#Oro"u^/uFgWɺ,p:6a,^x%$Yve^3PƗMnTP&yS}OJ '덫MH^:rXԴJۋ/rI;S*,+yz1hv)Qw^ڍJ2oL׊q(\fDj:^T%vOadɂnS}ZO)N*λdaȜkG_PIEO}нa(^iQX᯦-7^)%g'SJx(.S9zVɴZ{E ))ۅi/s7 VIV-|sj0*UBTHIqRf>FP$KqN0 R̻8j\GcC}IUz\i 6F)Q{Gҧ3qSzKj-Az VЛS-zy:8*mNk|D鿓ND2u+0Yŝ7kqm·?8Ib]u>˗^_>(]vӋzv+ݩ){vZrJ2RQ몋C$z [,pp,8mڊbR]Il .f~d/ݓs㓓mͶ{mgjQwn=Oic9ܚm4Q/6ݨ[TƧ?nԶoytf{@AzT{e{[O'ZRZt~AGD?s3􌿂ՉIw'|~U\ w~di:Kޱ)U/sU%njѩ&GSP^ǝd)..!^U` 1wX[aԇSxoFV6_扐)T 2Mfd=ۖͭiZ7KK Bi9%7@<3<ճԻU,},a}FRqɛr i@ONJvK KLN M, ʖv0n-]DwlI-X6ܶ$Jʴh5O+mOI+Ra瞠\ MG7BفjYo1#͖0V`Ѱ2M?c8>-Crt*JkIGS:e#hPKx[鱼>{5m;wcն&>j-M֥^َ) 6yȜl_w{-ō̱r> U=]iw3)r*]:K]6BdCTZ|>gf}LW}[$'Y5 &c -j.z6R 67MԷFMnÌwI7w5E}o޽+K ֵy4܌ȥW"COyR[q5Ӱ͙f[v"_#q{MV6܍3"u9BK(41ӯqˇc${ߝCi6I(OmθzҜ5k^:>Jzw.>qV8{vU[ڶEm|DžBz].KHjI]x;Mɗ{m,qZXr忇2u^RO2Z}ZێS[2Jen!*NDcrBUً4<ǼMҲs1Zw57c3&ĖڻzmP*FuJG1-dN:|OU}ҵgi2t~F^^Z.VxjvŧnNNh<:]^~NN+ge^g.SԔGFe߯'[vn'(ScJ]kܗ7eJOlRrfziݮq̋S"\*U<*W]k$FջV}? 7g#֟*GQQ|#/bo]p$>_Un9;l S VvQU%OLU{οmU6bZ1MTx%!֙Q7, J=!3 ;Q,ڌ;6ͱ݅q^&ߔ·n #WbwӖX.HtG)N&d̵zpI,n cu ޖUj+VXUp[w]N o.J6Z8Ts&utxln;~HPHS/xw`G\ʡ¿rj Z^vt"[L:SD\h0sUwR,}[x^X,R2Vn< ]2YDr[SRKs8tXb̷G?Ps Tv 3be,zVz D[/I.KOEQrm'$7|[J>r S`5յwT#\w1FTz\Ԛ &"ׅhSHrD\'r]~/>p;:Piuu:"9ő=tTaS7V2rӷk7mb[^WmPp*[y.Þ6f]cizJCgRR@UVl큝.WJP1N{/\whZ ػϧӱE7|E֫Sί.x-Y&pi%v''-x6r'Ws*6=DwwUu]=C?MK [yrtܒG$!WGqJ*%SAz ED[^)/tė/g=#Omd.|^n/sl׉g DZqemqowݮRzUܜ=ڽ-o/Iۖ;qVʘgPp|mm;6zGl9.8pwWgsJ2qPbe}}UpNjٯ}7TMQKrؽtEx%v w߾8%|j;~|}pK]ơ/ w߾8%|j;~|}pK]ơ/ w&~e_H 8PL7:%ʭ5Kw&U2vwR_+rm'}C7#rWoO&HoG?M$UR7{FU]u ;# !Wk`|W>׹潇9Vn)6)*ҹ{%qV4q>W1vi#T"Qk&GwxcJBJ- Ϸ^ˁxkU}ԣ/3.;]J=<*)cS)ROK9H=,r zX @)cS)Da^ԽQ gxJI=w֣gf*TRj

*Corresponding author (wmsohn@gnu.ac.kr)
• Received: March 7, 2017   • Revised: April 12, 2017   • Accepted: April 30, 2017

Copyright © 2017 by The Korean Society for Parasitology and Tropical Medicine

This is an Open Access article distributed under the terms of the Creative Commons Attribution Non-Commercial License (http://creativecommons.org/licenses/by-nc/4.0) which permits unrestricted non-commercial use, distribution, and reproduction in any medium, provided the original work is properly cited.

  • 8,742 Views
  • 136 Download
  • 5 Web of Science
  • 6 Crossref
  • 6 Scopus
prev next
  • The present study was performed to record new definitive hosts of Isthmiophora hortensis, and to describe morphological characteristics derived from a variety of worm samples for clarification of its taxonomic validity. Morphological characteristics with dimensions were observed in worm samples (n=21) from naturally infected wild animals, including a raccoon dog Nyctereutes procyonoides from Gimhae-si (City), Gyeongsangnam-do, stray cats and a striped field mouse from several localities, and a wild boar Sus scrofa, from Gurye-gun (County), Jeollanam-do. In addition, adult flukes (n=45) recovered in albino rats experimentally infected with the metacercariae from a freshwater fish species were also subjected to morphological studies. The mean ratios of the body length (BL) to body width (BW) were 5.86 and 5.76 in worms from wild animals and experimental rats, respectively. Those of the ventral sucker to oral sucker were 2.92 and 3.01 in worms from 2 groups. The mean percentages of the hindbody length (HBL) to BL were 42.1 and 41.2 in 2 groups. Those of uterine fields to BL were 9.8 and 12.2 in the 2 worm groups. By the present study, the 2 species of wild animals, the raccoon dog and wild boar, have been added as new definitive hosts for I. hortensis. The morphological characteristics of adult flukes derived from a variety of host source were redescribed to support the taxonomic validity of this echinostome species.
Isthmiophora hortensis (Digenea: Echinostomatidae) was originally described as Echinostoma hortense from rats in Japan [1]. After then, this fluke was also found in rats, dogs, cats, weasels, raccoons, and striped field mice naturally infected in Korea, Japan, China, and Vietnam [210]. Human infections with this fluke were reported in Japan, the Republic of Korea (=Korea), and China [1117]. Especially in Korea, several endemic foci have been reported [1821], and some clinical cases have been sporadically diagnosed by the recovery of worms in gastroduodenal endoscopy [2226]. Therefore, this echinostome is a dominant species with clinical importance in Korea.
The generic conceptions and relationships of Isthmiophora and Euparyphium have not been clarified for a long time in the family Echinostomatidae. To clarify the validity of Isthmiophora in relation to Euparyphium, Kostadinova and Gibson [27] analyzed the morphological characteristics of related specimens in 2 genera and redefined each generic character. They also made a differential key to the species of Isthmiophora with the morphological characteristics of related worm samples. In their study, I. hortensis was redescribed with nomination of a new combination from previously described as Echinostoma hortense [1]. However, Kostadinova and Gibson [27] used only 4 worm specimens donated by a Korean parasitologist. Therefore, we seriously needed to describe the differential morphological indices derived from a variety of worm samples to support the taxonomic validity. In addition, new definitive hosts of this fluke, the raccoon dog and wild boar, were reported in this study.
We obtained a dead body of the raccoon dog, Nyctereutes procyonoides, which was captured from Jangyu-myeon (Township), Gimhae-si (City), Gyeongsangnam-do (Province), Korea. Several sets of the viscera of wild boars, Sus scrofa, which were captured from Masan-myeon in Gurye-gun (County), Jeollanam-do, Korea were donated from a local hunter. Five stray cats (Felis catus) and a striped field mouse (Apodemus agrarius) were captured in several different localities. The muscles of wild animals were examined to survey the infections of Trichinella larvae, and additionally their visceral organs were also examined to detect other helminths.
The metacercariae of I. hortensis were isolated and collected from the Korean dark sleeper, Odontobutis platycephala, from Yugucheon (a branch of Geum River) in Gongju-si, Chungcheongnam-do by the artificial digestion method. Then, 50–70 metacercariae were orally infected with a gavage needle to each of 4 albino rats. To obtain the age-controlled adult worms, rats experimentally infected with the metacercariae were killed by cervical dislocation at days 15, 18, 24, and 36 after infection. The small intestines of wild animals and rats were longitudinally opened with a scissors in 0.85% saline, and washed with same solution until the supernatant became cleared. The sediment of the intestinal content was carefully examined with the naked eyes and under a stereomicroscope. The recovered worms were fixed in 10% neutral buffered formalin under the slight pressure of a cover glass, stained with Semichon’s acetocarmine, and observed under a light microscope equipped with a micrometer (OSM-4, Olympus Co., Tokyo, Japan).
For the differential indices of I. hortensis, total 76 worm samples were subjected to this analysis. Among them, 21 samples were recovered from wild animals, including a raccoon dog, stray cats, a striped field mouse, and a wild boar. The remaining 55 specimens were recovered from experimental rats (Tables 1, 2, 4). We basically measured the body length (BL) and width (BW), the size of oral (OS) and ventral suckers (VS), pharynx, esophagus, head collar, cirrus sac, ovary, and 2 testes, and additionally the forebody length (FBL: from the anterior end to the anterior margin of the ventral sucker), hindbody length (HBL: from the posterior margin of the posterior testis to the posterior end), and uterus length (UL: from the posterior margin of the ventral sucker to the anterior margin of the ovary). Then, we also calculated the ratios of BL/BW, VS/OS, the length/width in ovary and 2 testes, and the percentages (%) of FBL/BL, HBL/BL, and UL/BL as the differential body indices of this fluke. Measurements and scales are in micrometers (μm) unless stated otherwise.
Recovery of worms from wild animals
Total 3 adult specimens of I. hortensis with some other species of echinostomes were recovered in the small intestines of a raccoon dog, N. procyonoides, and 2 adult specimens were collected from a wild boar, S. scrofa (Fig. 1). Total 16 specimens of I. hortensis were recovered from the small intestines of 5 stray cats and a field mouse.
Dimensions of worms from wild animals and experimental rats
The characteristic features of I. hortensis included the presence of a head collar with total 27 collar spines and the dextral location of the ovary. Morphological characteristics of worm samples from wild animals (raccoon dog, stray cats, striped field mouse, and wild boar) were consistent with similar basic features (Fig. 1). However, their dimensions, including the size of worms, were more or less different by the species of host animals (Table 1). Dimensions of worms recovered from albino rats at days 15, 18, 24, and 36 after infection were revealed in Table 2.
Differential indices of worms from wild animals and experimental rats
The ratios of BL to BW were 5.21–6.70 (5.86 in average) and 5.53–5.95 (5.76) in worms from wild animals and experimental rats, respectively. Those of VS to OS were 2.45–3.71 (2.92) and 2.93–3.26 (3.01) in worms from 2 groups. The ratios of length/width of the ovary were 0.92–1.05 (0.96) and 0.81–0.97 (0.88), those of the anterior testis were 0.83–1.11 (0.91) and 0.93–0.96 (0.95), and those of the posterior testis were 1.28–1.67 (1.41) and 1.28–1.49 (1.41) in worms from 2 groups, respectively. The percentages of FBL to BL were 10.7–16.6 (14.0) and 10.2–12.9 (11.6), and those of HBL to BL were 40.3–43.2 (42.1) and 38.7–42.5 (41.2) in 2 groups. The percentages of uterine field (U) to BL were 6.6–16.9 (9.8) and 9.9–15.6 (12.2) in 2 worm groups (Table 3).
Comparison of normal and abnormal worms from experimental rats
Among the worms recovered from an experimental rat at day 15 after infection, 2 morphologically different worm groups, normal slender and abnormal plump, were categorized (Fig. 2). Normal slender worms were 6,075–7,000 (6,475) long and 1,100–1,275 (1,171) wide. Abnormal plump worms were 3,875–6,050 (4,846) long and 1,125–1,575 (1,343) wide. Dimensions of 2 worm groups and those from Kostadinova and Gibson [26] were designated in detail in Table 4.
The mean ratios of BL/BW were 5.53 and 3.61 in normal and abnormal worms, respectively. Those of VS/OS were 2.95 and 2.89 in 2 worm groups. The mean ratios of L/W in the ovary were 0.94 and 0.94, those in the anterior testis were 0.95 and 0.83, and in the posterior testis were 1.49 and 1.28 in 2 worm groups, respectively. The mean percentages of FBL/BL were 12.9 and 15.8, those of HBL/BL were 38.7 and 27.6, and of U/BL were 9.9 and 14.2 in 2 worm groups (Table 5).
Redescription of I. hortensis based on specimens from experimental rats (Table 2)
Body elongate, with attenuated anterior end, maximum width at mid-level of uterus. The average ratio BL/BW was 5.76. Head collar reniform, bearing 27 collar spines with 4 end group ones on each side. Tegumental spines distributed on the body surface from the posterior level of the collar to the posterior 1/3 of body. Oral sucker small, subterminal. Ventral sucker well developed, slightly elliptical. The average ratio VS/OS 3.01. The average percentage of FBL/BL 11.6. Prepharynx very short. Pharynx muscular, well developed. Esophagus short. Ceca bifurcating between the suckers or near the ventral sucker, extending to near the posterior end. Cirrus sac elongate. Seminal vesicle in the posterior portion of the cirrus sac, pars prostatica and ejaculatory duct present in anterior portion of cirrus sac. Ovary spheroidal or rarely oval, dextro-anterior to the anterior testis. The average ratio of L/W in the ovary 0.88. Mehlis’ gland large. Testes usually lobed or rarely entire, tandem; anterior testis usually wider than long; the posterior testis usually longer than wide. The average ratios of L/W in the anterior and posterior testis 0.95 and 1.41. The average percentage of HBL/BL 41.2. Vitelline glands well developed, follicular, laterally extending from anterior level of ovary to near the posterior end. Uterine coils transverse intercecal, extending from ovary-level to anterior level of acetabulum. The average percentage of U/BL 12.0. Eggs large, yellowish-brown, 93–128 (102) by 45–68 (57).
Several species of mammals, the black-striped field mouse, dog (Canis lupus), cat, Manchurian reed vole (Microtus fortis pellceus), mouse (Mus musculus), house rat (Rattus norvegicus), and black rat (Rattus rattus alexandrinus), have been reported as the definitive hosts for I. hortensis in Korea [24,9,10]. This fluke was also found in wild animals, i.e., rats, dogs, cats, weasels (Mustela itatsi), and raccoons (Procyon lotor), in Japan, China, and Vietnam [58]. By the present study, 2 species of mammals, raccoon dog (N. procyonoides) and wild boar (S. scrofa), are added in the list of the definitive hosts of this fluke.
Total 21 echinostome species in 14 genera have been reported from humans and/or animals in Korea, i.e., Acanthoparyphium marilae, A. tyosenense, Chaunocephalus ferox, Echinochasmus japonicus, E. perfoliatus, Echinoparyphium recurvatum, Echinostoma cinetorchis, E. gotoi, E. miyagawai, E. revolutum, Euparyphium murinum, Himasthla alincia, H. kusasigi, H. megacotyle, I. hortensis, Nephrostomum ramosum, Patagifer bilobus, Pegosomum bubulcum, Petasiger neocomense, Saakotrema metatestis, and Stephanoprora sp., [2832]. Among them, 4 species, namely, A. tyosenense, E. japonicus, E. cinetorchis, and I. hortensis, are known as the causative agents of human echinostomiasis in Korea [28]. Especially, human cases with I. hortensis have been sporadically and endemically occurred in this country since the first case was reported [11,13,14,16,1821]. Some clinical cases by this fluke infection have been recently diagnosed by the recovery of worms in gastroduodenal endoscopy [2226]. However, human cases with other 3 echinostome species were rarely reported in Korea [28]. Accordingly, I. hortensis is the predominant echinostome species in Korea.
Kostadinova and Gibson [27] used only 4 worm specimens (3 from a rat and 1 from mouse) donated by a Korean parasitologist in the new nomination of this fluke. However, in the present study, total 76 worm samples were subjected to make the differential indices for the identification of I. hortensis. Among them, 21 specimens were recovered from naturally infected wild animals (raccoon dog, stray cats, striped field mouse, and wild boar), and 55 specimens from rats (15-day, 18-day, 24-day, and 36-day-old) experimentally infected with the metacercariae. Each differential index was nearly similar in 2 worm groups (from wild animals and experimental rats). However, they were more or less different from those of Kostadinova and Gibson [27]. The average ratios of BL/BW were 5.86 and 5.76 in 2 worm groups in the present study; however they were 3.75 (from rat) and 4.65 (from mouse) in a previous study. The average ratios of L/W in posterior testis were 1.41 in both worm groups, but they were 0.90 and 1.58 in Kostadinova and Gibson [27].
We recovered 2 morphologically different types of worms, normal slender and abnormal plump, from rats experimentally infected with metacercariae from a freshwater fish species, O. platycephala. Especially in the 15-day-old worms, the average ratios of BL/BW were 5.53 and 3.61 in the normal slender and abnormal plump worm groups. However, those from Kostadinova and Gibson [30] were 3.75 (worms from rat) and 4.65 (worm from mouse). The ratio of BL/BW in worms (from rat) from Kostadinova and Gibson [27] was similar to that of abnormal plump worms rather than normal slender worms obtained in the present study. The average percentage of HBL/BL was 27.6 in the abnormal worms, which did not correspond with that (>30%) of the worms of the genus Isthmiophora [27,33].
Kostadinova and Gibson [27] divided the species of Isthmiophora into 2 groups by the number of collar spine (27 and 29), and they allocated 4 species, i.e., I. beaveri, I. inermis, I. melis, and I. hortensis, in the worm group with 27 collar spines. Among the 27-collar spined group, I. hortensis is very similar to I. melis except for the location of the cirrus sac and size of eggs. The differential indices for I. melis are also closely similar to those of I. hortensis from a mouse of Kostadinova and Gibson [27] and normal worms of the present study (Table 5). The species validity between above 2 Isthmiophora is not obvious only in the morphological aspect. Therefore, the more reasonable taxonomic keys, such as molecular phylogenetic data, will be needed to support the validity of each species.
On the basis of data from 45 worms, which were recovered from experimental rats in the present study, the following diagnostic features were considered as the differential indices for I. hortensis. The worm body is long slender (the ratio of BL/BW: 5.53–5.95); the ventral sucker is larger about 3 times than the oral sucker (the ratio of VS/OS: 2.93–3.26); the ovary is transversely elliptical (the ratio of L/W in the ovary: 0.81–0.97); the anterior testis has a slightly larger transverse axis (the ratio of L/W in the anterior testis: 0.93–0.96); the posterior testis has a long vertical axis (the ratio of L/W in the posterior testis: 1.28–1.49); short forebody (% of FBL/BL: 10.2–12.9); long hindbody (% of HBL/BL: 38.7–42.5); short uterine field (% of UL/BL: 9.9–15.6); the head collar with 27 collar spines and the dextral position of the ovary.
By the present study, it has been confirmed for the first time that the raccoon dog (N. procyonoides) and wild boar (S. scrofa) are the definitive hosts of I. hortensis. Additionally, the morphological characteristics with differential indices derived from a variety of worm samples from wild animals naturally infected and rats experimentally infected could support the taxonomic validity of I. hortensis.

CONFLICT OF INTEREST

The authors declare that they have no conflict of interest related to this article.

We thank Jung-A Kim and Hee-Ju Kim, Department of Parasitology and Tropical Medicine, Gyeongsang National University School of Medicine, Jinju, Korea, for their help in the collection of metacercariae from fish and recovery of worms from animals.
Fig. 1
Isthmiophora hortensis adults (Semichon’s acetocarmine stained) recovered from the small intestines of wild animals. (A) A specimen from the raccoon dog (Nyctereutes procyonoides). (B) Another from a wild boar (Sus scrofa). (C) A worm from a stray cat (Felis catus). (D) Another from a black-striped field mouse (Apodemus agrarius). Scale bar=1,500 μm.
kjp-55-3-287f1.gif
Fig. 2
Isthmiophora hortensis adults (Semichon’s acetocarmine stained). (A) Normal slender. (B) Abnormal plump, recovered from a rat experimentally infected with metacercariae, which were collected form the Korean dark sleeper, Odontobutis platycephala, from Yugucheon (a branch of Geumgang) in Gongju-si, Chungcheongnam-do. Scale bar=1,500 μm.
kjp-55-3-287f2.gif
Table 1
Dimensionsa of Isthmiophora hortensis adults recovered from wild animals in Korea
Table 1
Organs Raccoon dog (n=3)
(Nyctereutes procyonoides)
Stray cat-1 (n=6)
(Felis catus)
Stray cat-2 (n=6)
(Felis catus)
Striped field mouse (n=4)
(Apodemus agrarius)
Wild boar (n=2)
(Sus scrofa)
Body
 (L) 4,625–6,125 (5,433) 3,925–4,775 (4,408) 5,750–8,800 (6,850) 4,675–9,950 (7,700) 4,500–4,875 (4,688)
 (W) 925–1,025 (983) 675–875 (758) 900–1,750 (1,196) 1,000–1,300 (1,150) 875–925 (900)

Oral sucker
 (L) 190–195 (193) 125–165 (152) 165–210 (190) 160–175 (168) 128–150 (139)
 (W) 190–195 (192) 175–185 (179) 185–250 (215) 150–185 (170) 133–155 (144)

Head crown (W) 350–355 (352) 315–350 (329) 375–450 (416) 300–375 (335) 225–250 (238)

Pharynx
 (L) 165–180 (173) 165–175 (171) 160–200 (183) 150–190 (165) 145–158 (152)
 (W) 140–150 (147) 135–165 (150) 145–215 (170) 155–180 (169) 110–120 (115)

Esophagus (L) 225–250 (233) 75–150 (105) 160–340 (223) 125–235 (200) 125–138 (132)

Ventral sucker
 (L) 400–490 (455) 420–500 (455) 500–710 (577) 470–550 (520) 470–530 (500)
 (W) 450–510 (487) 410–470 (450) 500–710 (613) 470–590 (540) 550–560 (555)

Cirrus sac
 (L) 325–490 (403) 270–400 (334) 450–650 (512) 420–550 (510) -
 (W) 150–200 (183) 120–200 (154) 210–350 (262) 170–300 (230) -

Ovary
 (L) 145–170 (155) 145–175 (160) 205–350 (254) 140–300 (258) 150–165 (158)
 (W) 130–165 (148) 155–185 (173) 220–350 (273) 200–310 (250) 150–190 (170)

Anterior testis
 (L) 335–500 (443) 300–400 (333) 450–720 (582) 400–790 (680) 310–350 (330)
 (W) 350–450 (400) 330–480 (398) 460–1,100 (673) 470–880 (710) - (400)

Posterior testis
 (L) 400–630 (515) 400–530 (470) 580–970 (718) 490–1,100 (898) 425–510 (468)
 (W) 350–410 (373) 300–480 (355) 400–800 (560) 380–600 (538) 270–325 (298)

a(L): length; (W): width; unit is μm.

Table 2
Dimensionsa of Isthmiophora hortensis adults recovered from albino rats experimentally infected with metacercariae
Table 2
Organs 15-day-old (n=10) 18-day-old (n=10) 24-day-old (n=15) 36-day-old (n=10)
Body
 (L) 6,075–7,000 (6,475) 5,925–7,800 (6,780) 5,675–9,825 (7,447) 9,450–10,850 (9,965)
 (W) 1,100–1,275 (1,171) 1,000–1,325 (1,158) 1,050–1,625 (1,303) 1,450–1,775 (1,675)

Oral sucker
 (L) 160–190 (174) 160–185 (172) 155–215 (188) 170–220 (193)
 (W) 185–205 (196) 185–210 (204) 190–240 (216) 220–250 (238)

Head crown (W) 350–375 (368) 315–400 (371) 350–490 (398) 410–440 (426)

Pharynx
 (L) 185–200 (189) 175–210 (190) 190–240 (212) 210–240 (229)
 (W) 130–175 (153) 140–195 (168) 130–210 (174) 160–210 (187)

Esophagus (L) 90–250 (143) 110–210 (152) 75–180 (134) 110–325 (173)

Ventral sucker
 (L) 480–600 (526) 510–600 (555) 480–630 (572) 640–770 (689)
 (W) 540–670 (564) 500–610 (550) 510–680 (613) 680–770 (719)

Cirrus sac
 (L) 350–520 (450) 370–530 (441) 380–600 (459) 550–720 (606)
 (W) 160–270 (201) 120–190 (160) 190–275 (226) 230–320 (278)

Ovary
 (L) 260–310 (284) 225–340 (281) 210–385 (274) 250–310 (291)
 (W) 250–340 (303) 235–355 (290) 230–435 (329) 330–380 (360)

Anterior testis
 (L) 580–720 (657) 560–930 (668) 510–1,000 (703) 750–1,050 (936)
 (W) 530–810 (693) 510–840 (693) 520–1,030 (760) 870–1,130 (980)

Posterior testis
 (L) 770–950 (871) 600–1,160 (845) 660–1,280 (899) 1,000–1,230 (1,139)
 (W) 480–650 (586) 510–650 (577) 470–800 (632) 880–1,190 (1,016)

a(L): length; (W): width; unit is μm.

Table 3
Differential indices in the adults of Isthmiophora hortensis recovered from wild animals and experimental rats
Table 3
Items Differential indices (average) in worms from
Wild animalsa Experimental ratsb
Body length (BL)/Body width (BW) 5.21–6.70 (5.86) 5.53–5.95 (5.76)
Ratio of ventral sucker (VS)/oral sucker (OS) 2.45–3.71 (2.92) 2.93–3.26 (3.01)
Ratio of length (L)/width (W) in ovary 0.92–1.05 (0.96) 0.81–0.97 (0.88)
Ratio of L/W in anterior testis 0.83–1.11 (0.91) 0.93–0.96 (0.95)
Ratio of L/W in posterior testis 1.28–1.67 (1.41) 1.28–1.49 (1.41)
Percentage (%) of forebody length (FBL)/BL 10.7–16.6 (14.0) 10.2–12.9 (11.6)
Percentage (%) of hindbody length (HBL)/BL 40.3–43.2 (42.1) 38.7–42.5 (41.2)
Percentage (%) of uterus length (UL)/BL 6.6–16.9 (9.8) 9.9–15.6 (12.0)

Total a21 and b45 adult worms were subjected respectively.

Table 4
Comparative dimensions of Isthmiophora hortensis adults from the present study and a previous study with those of I. melis
Table 4
Organs Present studya Kostadinova and Gibson [27] from I. melisb


Normal (n=10) Abnormal (n=10) Rats (n=3) Mouse
Body
 (L) 6,075–7,000 (6,475) 3,875–6,050 (4,846) 5,376–6,463 4,043 4,564–7,346 (5,577)
 (W) 1,100–1,275 (1,171) 1,125–1,575 (1,343) 1,362–1,826 869 1,014–1,348 (1,203)

Oral sucker
 (L) 160–190 (174) 155–180 (167) 159–246 145 188–261 (221)
 (W) 185–205 (196) 170–190 (183) 203–246 159 203–261 (228)

Head crown (W) 350–375 (368) 325–375 (364) 319–391 261 377–464 (406)

Pharynx
 (L) 185–200 (189) 165–195 (180) 145–239 130 188–246 (221)
 (W) 130–175 (153) 130–180 (156) 116–217 217 130–188 (169)

Esophagus (L) 90–250 (143) 125–180 (149) 0–145 130 116–290 (200)

Ventral sucker
 (L) 480–600 (526) 460–575 (499) 551–826 406 551–797 (676)
 (W) 540–670 (564) 455–600 (513) 580–768 464 536–753 (668)

Cirrus sac
 (L) 350–520 (450) 365–450 (404) 377–420 406 246–652 (462)
 (W) 160–270 (201) 140–200 (176) 217–232 174 203–362 (303)

Ovary
 (L) 260–310 (284) 210–300 (246) 232–275 188 203–319 (256)
 (W) 250–340 (303) 205–325 (261) 275–319 188 in diameter

Anterior testis
 (L) 580–720 (657) 450–660 (576) 478–623 464 362–739 (568)
 (W) 530–810 (693) 580–860 (691) 739–942 464 420–811 (590)

Posterior testis
 (L) 770–950 (871) 570–980 (744) 609–739 594 536–1,174 (734)
 (W) 480–650 (586) 480–700 (581) 609–884 377 319–609 (476)

a15-day-old adults recovered from rats experimentally infected with metacercariae which were collected form Korean dark sleeper, Odontobutis platycephala.

bFrom Kostadinova and Gibson [27].

Table 5
Comparative differential indices of Isthmiophora hortensis adults from the present study and a previous study with those of I. melis
Table 5
Index Present studya Kostadinova and Gibson [27] from I. melisb


Normal (n=10) Abnormal (n=10) Rats (n =3) Mouse
Body length (BL)/Body width (BW) 5.53 3.61 3.75 4.65 4.64

Ratio of ventral sucker (VS)/oral sucker (OS) 2.95 2.89 3.18 2.86 2.99

Ratio of L/W in the ovary 0.94 0.94 0.86 1.0 1.0

Ratio of L/W in the anterior testis 0.95 0.83 0.66 1.0 0.96

Ratio of L/W in the posterior testis 1.49 1.28 0.90 1.58 1.54

Percentage (%) of forebody length (FBL)/BL 12.9 15.8 12.5 14.7 14.8

Percentage (%) of hindbody length (HBL)/BL 38.7 27.6 40.6 37.6 41.0

Percentage (%) of uterus length (UL)/BL 9.9 14.2 7.4 5.7 3.0

a15-day-old adults recovered from rats experimentally infected with metacercariae which were collected form Korean dark sleeper, Odontobutis platycephala.

bFrom Kostadinova and Gibson [27].

  • 1. Asada J. On a new echinostomatid trematode and its life history. Trans Jpn Soc Pathol 1926;16:293-294. (in Japanese).
  • 2. Park JT. A rat trematode, Echinostoma hortense Asada, from Korea. Keijo J Med 1938;9:283-286.
  • 3. Seo BS, Cho SY, Hong ST, Hong SJ, Lee SH. Studies on parasitic helminths of Korea 5. Survey on intestinal trematodes of house rats. Korean J Parasitol 1981;19:131-136.
  • 4. Cho SY, Kang SY, Ryang YS. Helminthes infections in the small intestine of stray dogs in Ejungbu City, Kyunggi Do, Korea. Korean J Parasitol 1981;19:55-59.
  • 5. Chen HT. Fauna Sinica. Platyhelminthes, Trematoda, Digenea (I). Beijing, China. Science Press; 1985, pp 333-497 (in Chinese).
  • 6. Fan SQ, Sun MF. Echinostoma hortense. Bull Harbin Med Univ 1989;23:161.
  • 7. Sato H, Suzuki K. Gastrointestinal helminths of feral raccoons (Procyon lotor) in Wakayama Prefecture, Japan. J Vet Med Sci 2006;68:311-318.
  • 8. Nguyen LA, Madsen H, Dao TH, Hoberg E, Dalsgaard A, Murrell KD. Evaluation of the role of rats as reservoir hosts for fishborne zoonotic trematodes in two endemic northern Vietnam fish farms. Parasitol Res 2012;111:1045-1048.
  • 9. Sohn WM, Na BK, Song HJ, Kim CM, Nam GJ. Intestinal helminthic infections in striped field mice, Apodemus agrarius, from two southern regions of Korea. Korean J Parasitol 2014;52:419-423.
  • 10. Shin SS, Oh DS, Ahn KS, Cho SH, Lee WJ, Na BK, Sohn WM. Zoonotic intestinal trematodes in stray cats (Felis catus) from riverside areas of the Republic of Korea. Korean J Parasitol 2015;53:209-213.
  • 11. Seo BS, Hong ST, Chai JY, Lee SH. Studies on intestinal trematodes in Korea. VIII. A human case of Echinostoma hortense infection. Korean J Parasitol 1983;21:219-223.
  • 12. Miyamoto K, Nakao M, Inaoka T. Studies on the zoonoses in Hokkaido, Japan. 5. On the epidemiological survey of Echinostoma hortense Asada, 1926. Jpn J Parasitol 1983;32:261-269. (in Japanese).
  • 13. Ryang YS, Ahn YK, Lee KW, Kim TS, Hhan MH. Two cases of natural human infection by Echinostoma hortense and its second intermediate host in Wonju area. Korean J Parasitol 1985;23:33-40.
  • 14. Lee SK, Chung NS, Ko IH, Ko HI, Chai JY. Two cases of natural human infection by Echinostoma hortense. Korean J Parasitol 1986;24:77-81. (in Korean).
  • 15. Chen XQ, Feng GX, Qian ZF. Survey on infection due to Echinostoma hortense in Liaoning Province. Chinese J Parasitol Parasit Dis 1993;11:226. (in Chinese).
  • 16. Huh S, Lee SU, Huh SC. A follow-up examination of intestinal parasitic infections of the army soldiers in Whachon-gun, Korea. Korean J Parasitol 1994;32:61-63.
  • 17. Hamamoto T, Kawasaki H, Maejima J, Hirai K. A case of Echinostoma hortense infection diagnosed by the upper gastrointestinal endoscopy. Nihon Shokakibyo Gakkai Zasshi 1997;94:487-491. (in Japanese).
  • 18. Ryang YS. Studies on Echinostoma spp. in the Chungju Reservoir and upper stream of the Namhan River. Korean J Parasitol 1990;28:221-233. (in Korean).
  • 19. Son WY, Huh S, Lee SU, Woo HC, Hong SJ. Intestinal trematode infections in the villagers in Koje-myon, Kochang-gun, Kyongsangnam-do, Korea. Korean J Parasitol 1994;32:149-155.
  • 20. Ahn YK, Ryang YS. Experimental and epidemiological studies on the life cycle of Echinostoma hortense Asada, 1926 (Trematoda: Echinostomatidae). Korean J Parasitol 1986;24:121-136. (in Korean).
  • 21. Lee SK, Chung NS, Ko IH, Sohn WM, Hong ST, Chai JY, Lee SH. An epidemiological survey of Echinostoma hortense infection in Chongsong-gun, Kyongbuk Province. Korean J Parasitol 1988;26:199-206. (in Korean).
  • 22. Chai JY, Hong ST, Lee SH, Lee GC, Min YI. A case of echinostomiasis with ulcerative lesions in the duodenum. Korean J Parasitol 1994;32:201-204.
  • 23. Lee OJ, Hong SJ. Gastric echinostomiasis diagnosed by endoscopy. Gastrointest Endosc 2002;55:440-442.
  • 24. Cho CM, Tak WY, Kweon YO, Kim SK, Choi YH, Kong HH, Chung DI. A human case of Echinostoma hortense (Trematoda: Echinostomatidae) infection diagnosed by gastroduodenal endoscopy in Korea. Korean J Parasitol 2003;41:117-120.
  • 25. Chang YD, Sohn WM, Ryu JH, Kang SY, Hong SJ. A human infection of Echinostoma hortense in duodenal bulb diagnosed by endoscopy. Korean J Parasitol 2005;43:57-60.
  • 26. Park CJ, Kim J. A human case of Echinostoma hortense infection diagnosed by endoscopy in area of southwestern Korea. Korean J Med 2006;71:229-234.
  • 27. Kostadinova A, Gibson DI. Isthmiophora Lühe, 1909 and Euparyphium Diet, 1909 (Digenea: Echinostomatidae) redefined, with comments on their nominal species. Syst Parasitol 2002;52:205-217.
  • 28. Chai JY. In Fried B, Toledo R eds, Echinostomes in Humans. The Biology of Echinostomes. 2009, New York, USA. Springer; pp 147-183.
  • 29. Choe S, Lee D, Park H, Oh M, Jeon HK, Lee Y, Na KJ, Kim Y, Lee H, Eom KS. Three echinostome species from wild birds in the Republic of Korea. Korean J Parasitol 2014;52:513-520.
  • 30. Choe S, Lee D, Park H, Jeon HK, Lee Y, Kim E, Na KJ, Eom KS. Two echinostome species, Pegosomum bubulcum and Nephrostomum ramosum (Digenea: Echinostomatidae), from an Eastern cattle egret, Bubulcus ibis coromandus, in Republic of Korea. Korean J Parasitol 2016;54:485-496.
  • 31. Choe S, Lee D, Park H, Jeon HK, Lee Y, Na KJ, Park SR, Eom KS. A case of chaunocephalosis by Chaunocephalus ferox (Digenea: Echinostomatidae) in an oriental white stork, Ciconia boyciana, in Korea. Korean J Parasitol 2016;54:659-665.
  • 32. Lee YI, Chung OS, Seo M. Recovery of Oswaldotrema nacinovici from whimbrels (Aves) in Korea. Korean J Parasitol 2016;54:809-812.
  • 33. Kostadinova A. Family Echinostomatidae Looss. In Jones A, Bray RA, Gibson DI eds, Keys to the Trematoda. 2:Wallingford, UK. CABI; 2005, pp 9-64.

Download Citation

Download a citation file in RIS format that can be imported by all major citation management software, including EndNote, ProCite, RefWorks, and Reference Manager.

Format:

Include:

New Definitive Hosts and Differential Body Indices of Isthmiophora hortensis (Digenea: Echinostomatidae)
Korean J Parasitol. 2017;55(3):287-294.   Published online June 30, 2017
Download Citation

Download a citation file in RIS format that can be imported by all major citation management software, including EndNote, ProCite, RefWorks, and Reference Manager.

Format:
Include:
New Definitive Hosts and Differential Body Indices of Isthmiophora hortensis (Digenea: Echinostomatidae)
Korean J Parasitol. 2017;55(3):287-294.   Published online June 30, 2017
Close

Figure

  • 0
  • 1
New Definitive Hosts and Differential Body Indices of Isthmiophora hortensis (Digenea: Echinostomatidae)
Image Image
Fig. 1 Isthmiophora hortensis adults (Semichon’s acetocarmine stained) recovered from the small intestines of wild animals. (A) A specimen from the raccoon dog (Nyctereutes procyonoides). (B) Another from a wild boar (Sus scrofa). (C) A worm from a stray cat (Felis catus). (D) Another from a black-striped field mouse (Apodemus agrarius). Scale bar=1,500 μm.
Fig. 2 Isthmiophora hortensis adults (Semichon’s acetocarmine stained). (A) Normal slender. (B) Abnormal plump, recovered from a rat experimentally infected with metacercariae, which were collected form the Korean dark sleeper, Odontobutis platycephala, from Yugucheon (a branch of Geumgang) in Gongju-si, Chungcheongnam-do. Scale bar=1,500 μm.
New Definitive Hosts and Differential Body Indices of Isthmiophora hortensis (Digenea: Echinostomatidae)

Dimensionsa of Isthmiophora hortensis adults recovered from wild animals in Korea

Organs Raccoon dog (n=3)
(Nyctereutes procyonoides)
Stray cat-1 (n=6)
(Felis catus)
Stray cat-2 (n=6)
(Felis catus)
Striped field mouse (n=4)
(Apodemus agrarius)
Wild boar (n=2)
(Sus scrofa)
Body
 (L) 4,625–6,125 (5,433) 3,925–4,775 (4,408) 5,750–8,800 (6,850) 4,675–9,950 (7,700) 4,500–4,875 (4,688)
 (W) 925–1,025 (983) 675–875 (758) 900–1,750 (1,196) 1,000–1,300 (1,150) 875–925 (900)

Oral sucker
 (L) 190–195 (193) 125–165 (152) 165–210 (190) 160–175 (168) 128–150 (139)
 (W) 190–195 (192) 175–185 (179) 185–250 (215) 150–185 (170) 133–155 (144)

Head crown (W) 350–355 (352) 315–350 (329) 375–450 (416) 300–375 (335) 225–250 (238)

Pharynx
 (L) 165–180 (173) 165–175 (171) 160–200 (183) 150–190 (165) 145–158 (152)
 (W) 140–150 (147) 135–165 (150) 145–215 (170) 155–180 (169) 110–120 (115)

Esophagus (L) 225–250 (233) 75–150 (105) 160–340 (223) 125–235 (200) 125–138 (132)

Ventral sucker
 (L) 400–490 (455) 420–500 (455) 500–710 (577) 470–550 (520) 470–530 (500)
 (W) 450–510 (487) 410–470 (450) 500–710 (613) 470–590 (540) 550–560 (555)

Cirrus sac
 (L) 325–490 (403) 270–400 (334) 450–650 (512) 420–550 (510) -
 (W) 150–200 (183) 120–200 (154) 210–350 (262) 170–300 (230) -

Ovary
 (L) 145–170 (155) 145–175 (160) 205–350 (254) 140–300 (258) 150–165 (158)
 (W) 130–165 (148) 155–185 (173) 220–350 (273) 200–310 (250) 150–190 (170)

Anterior testis
 (L) 335–500 (443) 300–400 (333) 450–720 (582) 400–790 (680) 310–350 (330)
 (W) 350–450 (400) 330–480 (398) 460–1,100 (673) 470–880 (710) - (400)

Posterior testis
 (L) 400–630 (515) 400–530 (470) 580–970 (718) 490–1,100 (898) 425–510 (468)
 (W) 350–410 (373) 300–480 (355) 400–800 (560) 380–600 (538) 270–325 (298)

a(L): length; (W): width; unit is μm.

Dimensionsa of Isthmiophora hortensis adults recovered from albino rats experimentally infected with metacercariae

Organs 15-day-old (n=10) 18-day-old (n=10) 24-day-old (n=15) 36-day-old (n=10)
Body
 (L) 6,075–7,000 (6,475) 5,925–7,800 (6,780) 5,675–9,825 (7,447) 9,450–10,850 (9,965)
 (W) 1,100–1,275 (1,171) 1,000–1,325 (1,158) 1,050–1,625 (1,303) 1,450–1,775 (1,675)

Oral sucker
 (L) 160–190 (174) 160–185 (172) 155–215 (188) 170–220 (193)
 (W) 185–205 (196) 185–210 (204) 190–240 (216) 220–250 (238)

Head crown (W) 350–375 (368) 315–400 (371) 350–490 (398) 410–440 (426)

Pharynx
 (L) 185–200 (189) 175–210 (190) 190–240 (212) 210–240 (229)
 (W) 130–175 (153) 140–195 (168) 130–210 (174) 160–210 (187)

Esophagus (L) 90–250 (143) 110–210 (152) 75–180 (134) 110–325 (173)

Ventral sucker
 (L) 480–600 (526) 510–600 (555) 480–630 (572) 640–770 (689)
 (W) 540–670 (564) 500–610 (550) 510–680 (613) 680–770 (719)

Cirrus sac
 (L) 350–520 (450) 370–530 (441) 380–600 (459) 550–720 (606)
 (W) 160–270 (201) 120–190 (160) 190–275 (226) 230–320 (278)

Ovary
 (L) 260–310 (284) 225–340 (281) 210–385 (274) 250–310 (291)
 (W) 250–340 (303) 235–355 (290) 230–435 (329) 330–380 (360)

Anterior testis
 (L) 580–720 (657) 560–930 (668) 510–1,000 (703) 750–1,050 (936)
 (W) 530–810 (693) 510–840 (693) 520–1,030 (760) 870–1,130 (980)

Posterior testis
 (L) 770–950 (871) 600–1,160 (845) 660–1,280 (899) 1,000–1,230 (1,139)
 (W) 480–650 (586) 510–650 (577) 470–800 (632) 880–1,190 (1,016)

a(L): length; (W): width; unit is μm.

Differential indices in the adults of Isthmiophora hortensis recovered from wild animals and experimental rats

Items Differential indices (average) in worms from
Wild animalsa Experimental ratsb
Body length (BL)/Body width (BW) 5.21–6.70 (5.86) 5.53–5.95 (5.76)
Ratio of ventral sucker (VS)/oral sucker (OS) 2.45–3.71 (2.92) 2.93–3.26 (3.01)
Ratio of length (L)/width (W) in ovary 0.92–1.05 (0.96) 0.81–0.97 (0.88)
Ratio of L/W in anterior testis 0.83–1.11 (0.91) 0.93–0.96 (0.95)
Ratio of L/W in posterior testis 1.28–1.67 (1.41) 1.28–1.49 (1.41)
Percentage (%) of forebody length (FBL)/BL 10.7–16.6 (14.0) 10.2–12.9 (11.6)
Percentage (%) of hindbody length (HBL)/BL 40.3–43.2 (42.1) 38.7–42.5 (41.2)
Percentage (%) of uterus length (UL)/BL 6.6–16.9 (9.8) 9.9–15.6 (12.0)

Total a21 and b45 adult worms were subjected respectively.

Comparative dimensions of Isthmiophora hortensis adults from the present study and a previous study with those of I. melis

Organs Present studya Kostadinova and Gibson [27] from I. melisb


Normal (n=10) Abnormal (n=10) Rats (n=3) Mouse
Body
 (L) 6,075–7,000 (6,475) 3,875–6,050 (4,846) 5,376–6,463 4,043 4,564–7,346 (5,577)
 (W) 1,100–1,275 (1,171) 1,125–1,575 (1,343) 1,362–1,826 869 1,014–1,348 (1,203)

Oral sucker
 (L) 160–190 (174) 155–180 (167) 159–246 145 188–261 (221)
 (W) 185–205 (196) 170–190 (183) 203–246 159 203–261 (228)

Head crown (W) 350–375 (368) 325–375 (364) 319–391 261 377–464 (406)

Pharynx
 (L) 185–200 (189) 165–195 (180) 145–239 130 188–246 (221)
 (W) 130–175 (153) 130–180 (156) 116–217 217 130–188 (169)

Esophagus (L) 90–250 (143) 125–180 (149) 0–145 130 116–290 (200)

Ventral sucker
 (L) 480–600 (526) 460–575 (499) 551–826 406 551–797 (676)
 (W) 540–670 (564) 455–600 (513) 580–768 464 536–753 (668)

Cirrus sac
 (L) 350–520 (450) 365–450 (404) 377–420 406 246–652 (462)
 (W) 160–270 (201) 140–200 (176) 217–232 174 203–362 (303)

Ovary
 (L) 260–310 (284) 210–300 (246) 232–275 188 203–319 (256)
 (W) 250–340 (303) 205–325 (261) 275–319 188 in diameter

Anterior testis
 (L) 580–720 (657) 450–660 (576) 478–623 464 362–739 (568)
 (W) 530–810 (693) 580–860 (691) 739–942 464 420–811 (590)

Posterior testis
 (L) 770–950 (871) 570–980 (744) 609–739 594 536–1,174 (734)
 (W) 480–650 (586) 480–700 (581) 609–884 377 319–609 (476)

a15-day-old adults recovered from rats experimentally infected with metacercariae which were collected form Korean dark sleeper, Odontobutis platycephala.

bFrom Kostadinova and Gibson [27].

Comparative differential indices of Isthmiophora hortensis adults from the present study and a previous study with those of I. melis

Index Present studya Kostadinova and Gibson [27] from I. melisb


Normal (n=10) Abnormal (n=10) Rats (n =3) Mouse
Body length (BL)/Body width (BW) 5.53 3.61 3.75 4.65 4.64

Ratio of ventral sucker (VS)/oral sucker (OS) 2.95 2.89 3.18 2.86 2.99

Ratio of L/W in the ovary 0.94 0.94 0.86 1.0 1.0

Ratio of L/W in the anterior testis 0.95 0.83 0.66 1.0 0.96

Ratio of L/W in the posterior testis 1.49 1.28 0.90 1.58 1.54

Percentage (%) of forebody length (FBL)/BL 12.9 15.8 12.5 14.7 14.8

Percentage (%) of hindbody length (HBL)/BL 38.7 27.6 40.6 37.6 41.0

Percentage (%) of uterus length (UL)/BL 9.9 14.2 7.4 5.7 3.0

a15-day-old adults recovered from rats experimentally infected with metacercariae which were collected form Korean dark sleeper, Odontobutis platycephala.

bFrom Kostadinova and Gibson [27].

Table 1 Dimensionsa of Isthmiophora hortensis adults recovered from wild animals in Korea

(L): length; (W): width; unit is μm.

Table 2 Dimensionsa of Isthmiophora hortensis adults recovered from albino rats experimentally infected with metacercariae

(L): length; (W): width; unit is μm.

Table 3 Differential indices in the adults of Isthmiophora hortensis recovered from wild animals and experimental rats

Total a21 and b45 adult worms were subjected respectively.

Table 4 Comparative dimensions of Isthmiophora hortensis adults from the present study and a previous study with those of I. melis

15-day-old adults recovered from rats experimentally infected with metacercariae which were collected form Korean dark sleeper, Odontobutis platycephala.

From Kostadinova and Gibson [27].

Table 5 Comparative differential indices of Isthmiophora hortensis adults from the present study and a previous study with those of I. melis

15-day-old adults recovered from rats experimentally infected with metacercariae which were collected form Korean dark sleeper, Odontobutis platycephala.

From Kostadinova and Gibson [27].