Revision of the Genus Anguillicola Yamaguti, 1935 (Nematoda: Anguillicolidae) of the Swimbladder of Eels) Including Descriptions of Two New Species, A

FOLI.-\. PARASITOLOGICA 35: 125-146,198B.

REVISION OF THE GENUS ANGUILLICOLA YAMAGUTI, 1935 (NEMATODA: ANGUILLICOLIDAE) OF THE SWIMBLADDER OF EELS) INCLUDING DESCRIPTIONS OF TWO NEW SPECIES, A. NOVAEZELANDIAE SP. N. AND A. PAPERNAI SP. N.

F. MORAVEC and H. T.ARASCHEWSICI

Ins~itute of Parasitology, Czechoslovak Academy of Soiences, (Jeske Budejovice, and Instituto of Special Zoology and Parasi~ology, Ruhr University, Boehum

Abstract. The genU£! Anguillicola is reviewed On the basis ofthe examination ofsome type specimens and other material. Two new species are c1esoribed: A. novaezelandiae sp. n. from Anguilla australis (type host) from New Zealand and A. an(IUilla from Italy (oonspecifio nematodes were oI'iginally reported from Italy as A. australien8is) and A. papernai sp. n. from Anguilla m088ambica from South Africa. At present the genus Anguillicola comprises the following five valid species: A. globiceps Yamaguti, 1935, A. australiensis .Tohnston at Mawson, 1940, A. cras8USKuwahara, Niimi at Itagaki, 1974, A. novaezelandia6 sp. n., and A, papernai sp. n, Two Anguillicola speaies, A. ora88U8 and A. n01'aezelandiae, were apparently introduced in Europe from other continents a few years ago. The genus AnguillicoZa is divided int.o two s\1bgenera: An(Juillicola (type species A. (A.)globiceps) and Angw"Zlicoloides subgen. n. (type species A. (A.) ara881As). All Anguillicolaspeeies are briefly described and illustrated. The paper is supplemented by a key to the identifica~ion of AngtdZlicola speoies.

The genus Anguillicola Ya,ma.guti, 1935 of the lllonotypic family Anguillicolic1ae (Dl'a.~unculoidea) was established by Yama.gu1ii (1935) for A. globiceps from the swimbladder of Japanese eels (Anguilla }aponica) in Japan. Later two additional congenerio species were described: A. australiensis Johnston e"t Ma.wson, 1940 from Anguilla reinhardtii from Austra.lia and A. cra88U8 Kuwahara., Niimi et Itagaki, 1974 from Anguilla anguilla and A. japonica raised in eel fa.rms ill Japan. The species were distinguished prinoipally by lihe form of the oesophagus, number of large unioellular reotal glands, and humbet of ca.uda.l papillite in the 11180le (see Pe1iel's a.nd Hattmanu 1986). Until recently, all the three a.bove mentioned species were considered to be limited in t·heir distribnt~on to East Asia and Australia. a.nd New. Zealand. The gene:ral interest in these pathogen,c nematodes has inoreased considerably since Angwillicola appeal'ed in Europe. In 1982, P aggi etaI. reported Anguillicola australiensis from AngUilla anguilla from Lake Braociano neal' Rome in Italy. At about the sallie time Anguillicola nematodes were recorded from eels from the open wa.ters of the Weser-Ems Rivet region in Northern Getmany (Neumann 1985); since then, "they have spread throughout many countries of WesteI'n1 Northern and Central Europe with grea.t rapidity. Although the nematodes from Westem Europe (Holland, Belg~um) were identified 80S A. crassus by V an Bann~ng at a.!. (1985) and De Charlexoy (1986), Peters and Hartmann (1986) sta.te that those from NOI'thern Germany cannot be 11l1eqllivocally assigned to allJ of the known species. According to De Oha:rlel.'oy et a!.

(1987), the Belgian. nematodes seem to be conspecific with A. cra88U81 but the authors mention that the l'edescription of A. australiensis based on Italian nematodes (Paggi -et a!. 1982) a.pproximates very closely to that of tlle helminths foun.d in Belgium. .consequently, they consider it possible that A~ Cra88'lt8 and .A. aU8tralien8~8 ar'e one and

125 the same speoies. This uncertainty in speoies identification of European Anguillioola members js responsible to the fa-at that these highly pathogenic nema.todes are frequently reported from Europe as only Anguillicola sp. (see e.g. Peters and Hartmann 1986, Molna.r 1986, K"ie 1987, Koops 1987). In an attempt to resolve the ptoblem of the species identity of European members of Anguill'£cola, we examined a number of the type and non-type spe~imens of all three previously recognized species in this genus. SOIlle of the results obtained pertaining to the European members 'of AnguilUcola have been published by Taraschewski et a1. (1987), who showed clearly that there are actually two species occurring in eeLo; in Europe. Since, however, some new and important data onthe morphology of Anguillicola members of other continents wete obtained during this study, it proved possible to undertake a. revision of the whole genus. It is hoped that this will help in relia.ble species identifica'tiolJ, and will fOl'm the basis for future work on, th0!3 biology of these interesting a:nd economically important helminths. In addition to the pre-viously recognized species, the ma.terials at our disposal contained two new, hitherto tmdescribed A.nguill1:cola. species which aYe described in the present paper.

MATERIALS AND METHODS

The following Anguillicora specimens were studied: A. globiceps - 3 ~i! + 2 ~~ from unknown host {Anguilla japonica?) f.com Ja.pan {leg. Dl" H, Hirose); - 1 ~, 2 juv. !i1~ + lllarvae from unknown host (Anguilla japonica?) from China (leg. Prof. P. Q. Wang and Prof. H. S. Wu); A. awtralierulis - type speeimens (~+ ~) from Anguilla reinhardtM from Sou~h Australia (Sou~h .Australian Museum, Adelaide, Call. No. V 1392); A. Ct'a98'U8 - syntypes (~+ !j:!) from Anguilla anguiUa from Ja.pan (Shizuka Pref., 1912, leg..Dr, A. Kuwahara) (Magmo Parasi~ological Museum, Tokyo, Call. No. MPM 19093); - specimens (21 d'd' + 29 !j!!?) from (IUltmed Anguilla japonica (Shiz1.ua Prof., M:aroh 1979, leg. Dr. E, Hirose); - () c1'Cf + 7 ~~ from un.knownhos~ (Anguilla iapon~'caf) from China (leg. Prof. . P. Q. Wang and Frof. H. S. Wu); -numerous speoimens from Anguilla anguilla from the Feeleral Republic of Germany (Ruhr Lake, River Weger, R. Elba estuary, June-Septembex: 1986, leg. Dr; H. Tarasohewski); A. novaezelaJtliUae ~ specimens (4 (f~ + 3!j?!j:! - types) from Anguilla australis from New Zealand (Matahina dam, 17 May 1978, leg. Dr. B. Jones); - speoimens (3 (fr! + 3 ~~) from Anguilla anguilla from I~aly (B:racciano Lake, leg. Prof. L. Paggi); ...4. papernai - specimens (2 ~r1 + 4 !j?!i!) from Anguilla m08sambica from Sou~h Africa (21 Sep~ember 1984, leg. Dr. I, Paper~a). The nematodes wElre mos~ly :fixed in 70% ethanol.or 4% formaldehyde. All the speoimens were cleared and examined in glycerine. En face views were prep!llred according to Anderson's (1958) method. Drawingswere made with the aid of a Zeiss microscope dra.wing attaohment. Allrneasure menta are in nun.

RESULTS

Gen.us Angnillicola YamagutiJ 1935 Diagnosis: AnguilHcol'dae. Cuticle very finely spinose or a.spinose, sometimes with marked, irregularly scattered exol'esceno.es of fibrous structureon. a.n.tetior a.nd posterior ends o£ body. Head end bearing foUl' cephalic pa.pilla.e a.nd two la.teral a.mphids. :Suo oal capsule present, provided with row of small circumoral teet,h, Oesophagus short and wide, consisting of antel'ior muscular and posterior muscular-glandular sections. Vslvular a.ppa,fatus well developed. Intestine dal'k, distended. Rectal glands Htl'ge. Male: Spicules absent, Testisbeginntng near tail end; seminal vesicle well developed. Ductus ejacuIa.torius opening outside on prominent caudalprocesB. Six pairs of sessile caudal papillae present. Female: Vulva opening on tip of prominent cone in posterior Pal't of body. Uterus opposed. Ovovivipa.rous. Parasites of swimbladder of eels. Type species: A. globiceps Yamaguti, 1935 Remark: Since the type spedes A. globiceps differs from the rerna.illing AnguiUicolet species in some taxonomically -very ixnporta.nt features (structure of oesophagus, spinose cuticle)" we consider it useful to sepa.ra.te the genus Anguill't'cola into two subgenera. which, in out' opinion, will reflect more accurately relationships among Anguillicola species.

I. Subgenus Anguillicola Yamaguti, 1935 Diagnosi s: Anguil.licola. Body filiform., outicle vel'Y :finely spinose throughout. Oesophagus ,conspicuously infla.ted at antel'iol' end, its posterior' part: cylindticaL Type species: A. (A.) globiceps :yamaguti, 1935 n. Subgenus AnguiUicoloides subgen. n. Diagnosis: Anguillicola. Body fusiforni or filiform, cuticle aspinose. Anterior end of oesophagus lackinginfla.tion, na.rrow; posterior part of oesophagus distinotly expanded. Type species: A. (A.)eras8U8 Kuwahara, Niimi et Itagaki, 1974 - o.

;REVIEW OF ANGUILLICOLA SPECIES

1. ..4nguillicola (Anguillicola) globiceps Yamaguti, 1935 Fig. 1 Descl'~:Ption: Body filiform. Ep~cuti(lle densely oovered by minute spines. He,ad end rounded, buccal capsule very large, well sclerotized; some 80 smaH circumoral teeth present. Anteri01'! end of oesophagus bulbously inflated" posteri,or part of oesopha gus cylindrical. Valvular apparatus well developed; Tlu:~ee latge and oue smallunicellular recta.lghmds present,. Tail very short, usually blunt. Male (5 specimens): Length of body 24: .. 28--32.91, maximum width 0.952--1.115. Length of hucoal capsule 0.068-0.109, itBwidth 0.272-0.408, its maximum thickness 0.OO~0.Ol5. Length o£ oesopha.gus 1.25-1.50; length of its anterior bulb 0.408 to 0.544, width 0.4713---0.707, maximum widthofit,s postedor cylindricalpart 0.258-0.299. Lengthrntio of oesophagus and body 1: 21.5-25.4. Distanoe of nerve ring and excretory pore from anterior extremity 0.571-0.775 and 1.27-1.37, respeptively. Common cloacal duct opening onprominent prooess 0.068-0.109 long. Size of larger recta.l glands 0.109-0.312 X 0,082--0.109, that of SillBlUer one O.068~0.081X 0.030-0.041. Six pa.irs of c&udalpapillae present: 3 preanals and 3 postanals. Tail 0.095-0.244 long. Female (2 speoimens): Length of body of gravid females 37.31-37.81, maxi:mum width 1.44--1.54. Length of buccal capsule 0.109-0.122, width 0.408, maximum thickness of its wall 0.009. Length of oesophQlgus 1.50; length of its anterior bulb 0.049, width 0.571-0.639, maximllm width of its posterior cylindrical part 0.299--0.313. Length ra.tio of oesophagus and body 1: 24.9-25.3. Dis,tance of nerve ring and ex.oretory por'a 0.74:8-0.762 and 1.56-1.59, :respectively. Size of 1801'ger recta.l gla.nds 0.163-0.204 by

127 0.109-0.136, that o£smaller one O.068xO.041. Vulva elevated, situated 6.99-8.43 from posterior extremity, this representing approximately 1/5 of body length. Uter-L"lS not rea.'Jhing an.te1'iorly oesophagca.l region; utel'Us containing numerous eggs (size 0.078-0.081 x O.069), developing embl'ya and fully formed" sheathed larvae sou:1.e 0.180 long and 0.015 wide. Rectum thin hyaline tube, ,anus slightly eleva.ted: Ta..il short, blunt, 0.122 long.

B ·1:::: -. ·""4', ~ .• 4.11I1 I. "101' ,. j,,"Ioo~.11 II., li~..."OIIU, Hili•••• IJ H,~ ."'tUUI ~.. ~.II.U~

O.2mm c

Fig. 1. AnguillicoZa UZobi~ep8 Yamagnti; 1935. A- head end of male; B- buocal capsule of male. a .,..- eauclal end of gravid female; D- posterior end of male; E- tail ~ip of male; F - vulva: 128 Ho st: An(Juilla japonica Terrl/ninck at Schlegel. Localization: swimbladder., . Distl'ibution: Japan. (C'entraI1fonshu) and China (Fujian, FIubei and Henan Provinces). Comm?~ts: - Th~ ~otphology of the present spe:::imen'S corresponds, more or less, to the angInal descrlptlOll of A .. globiceps given by Yamaguti (1935);. even though we clid not succeed in obtaining the type specimens for study, it is clear that our nematodes belong to this species. Some slight differenoes concern only measurements that are appa.rently vvithin the normallill1its of intraspeci.fic varia.bility of this species. Yamaguti (1935,) provided a relatively very origin~l description of A. gloMceps, but he did not study the exa.ct number of circumoral teeth in the buccal capsule which is, in our opinion, an important specific taxonomio feature in Anguillicola. spp. Un f01·tUhately; we were also unble to establish tbe exact number of these teeth from head end sections because of the limited number of horrowed iSIlecimeus available; in latera.l view it was possible to determine that the buccal capsule of the specimens of the present ma.teria.l was a.rmed with some 80 teeth. According to Wang and Zhao (1980), A. globiceps possesses 76 teeth, but it is probable that there is some individual variability in the number of these teeth,as suggested by data on other congeneric species. It has been mentioned above that A. globz'ceps diffe '8 considerably in its morphology from other congeneric species, principally in the struct1.1re of the oesopha.gus which is usually considered one of the most important features in nema.tode taxonomy. Other marked differences are the spinose epicuticlea;nd the form of body. Therefore, we pr.opose to pla.ce A. globz'c,eps and the rema.ining AnguilUcola species into two sepa.ra.te subgenera (see a.bove). A. globicep8 was odg,naJly described by Y amaguti (1935) from Lake Hama.na. from Japan (Central Honshu); he mentions tha.t this species is vm:y COUlmon in Anguilla. }aj')onica in which hea.vy infections ca.uae a. considerable thickening of the host's swimbla.dder. wall. Latel'this species Wa.B reported from Japan by Egusa et a.!. (1969), but in fact another speo~es, A.Cfa.88'US was misident~fied as A. globicep8 in this case (soo Hirose et a.!. 1976). According to recentinfoz-mation from Dr. H. Hirose, in Japan this species is muoh leBscommon in A. japonz:cathan A. C'l'aS8U8. In 1956, Wu reported A. ylobicep8 from pond-cultured A.japonicafrom China, butthe accompanying drawings suggest that besides A .. globioepa, A. cra.S8'U8 might also ha.v1e beenincluded in the material examined. A. globiceps was also reported by W u (1984) from th6 ,same host species from the Liaoho River (China, Prov. Rel1a.n) , buv the drawing of the anterior end of uhe nematode (Fig. 17, p. 198) is more similar to A. Cra88'U8 tha.n A. globiceps. The A.nguill?~cola material from Ohina provi,ded by Prof. P. Q. Wang and Prof. H. S. Wu contai.ned both A. gloMceps and A. craS8U8. Acco~ding to Wang and Zhao (1980), A. globiceps is a common parasite of Anguilla. japonica in the Chinese Provinces of Fujia.n and Hubei. .It appears from the present data. tha.t A. globiceps is restricted in its distribution to the region of the Fa.r East only, btlt its future accidental introducliion in other geogra.phical regions cannOli be excluded.*

'" Only l'ecen~ly. when this paper w,as M:ept.erl. for publication. t'he recon~ pUbli/J.ation of Ghi ~t~no (1981S), xepOl'ting the ~ecol'cl of A. rJlotncBp8 III Italy. has come t;o the authors a,t-ten,tlOn. Inour opml~n. 0. re-examination of these. speoim.ena and the verificati.on of their species applU'te21a.nce is qUlte necessal'Y, because t,heir identifioation was based em Dr. il'rIoItir's presumption that A. {/lobiceps was the only valid species of Ang1.tillico~a,.

129 2. Anguillicola (Anguillicoloides) australiensis Johnston at IVlawson, 1940 Fig. 2

Description (based on type specimens):.~arge siz?d, .brown coloured,nematodes~ Body filiform, relatively Ila.ttow for AngmZ~~c~la; .eplcutwle sl11ootl~, asplnose. IIea.d end bulbously jnflated" spherical, neck constrICtIOn m front of nerye rmg marked; body tapering at postedol' end to pointed tail. Buccal ca.psule small, thm.walled, its anterior edge a.t"lued with numerous minute circumoral teeth; numher' of teeth not established. Oesophagus expanded at its posterior half. Valvular app~&.tuB of oesophagus well developed. NeI've ring situated somewhat below neck constrICtIOn, excretol'y pore near junction of oesopha.gus and intes,tine. Intestine thick~waned, dark, being :first narrow" t,hen becoming very broad, occupying a.lmost whole width of body. Three conspicuously large oval mllcellula.r rental gla.nds present, one dorsal and two subventral; additiona.l small ventral rectal gland also present. Tail conical, pointed. Male (1 specimen): Length of body 32.30, maximum width 0.911; length of infla.ted head end 0.163, its width 0.218; width of body at neck constriction 0.136. Buccal capsule 0.009 long a.nd 0.030 wide; maximum thickness of its wall 0.003. Length of oesophagus O.93B} its maximum width 0.204; length of oesophageal valves 0.054:. Length :ratio o~ oesopha.gus and body 1: 34.4-. Distance of nerve ring and excretory pore it'om anterior extremity 0.258 .and 1.05, respectively. Testis reaching anteriorly slightly below end o£ oesopbagu8. Seminal vesicle well developed, size O.816xO.476. Common cloaca.l duot opening on prominent prooess 0.075 long. Size of large rectal glands 0.313-0.340 by 0.190-0.299, of small one 0.122 X 0.082. SixpaJrs of caudal papUlae present: 3 preanals, 1 adana!, and 2 posta-nals; adana.l and first pI'eanaJ pail'S less distinct than. others. Tail conical, 0.204 long. Female (1 specimen): Length of body of nongravid fema.le 27.25, maximum width 0.721 ; length of inflated head end O.150 j its width 0.204; width of body at neck constriction 0.095. Buccal ca.psule 0.009 long ,and 0.033 wide; maxi,mum thiokness of its waJI 0.003_ Length of oesophagus 0.898, its maximum width 0.177; length of qeSOp.hageal valves 0.054. Lengthratio of oesopha.gus and body J.: 30.3. Distanceofnerve ring and excretory pore from anterior extremity 0.272 andO.789} respectively. Vulvaelevated,prominent~ situated in posterior part of body, 6.96 from posterior extremity, this repI'esenting approximately 1/4 of body length. Uterus 'still without eggs. Anterior ova.:ry reaching anteriorly almost mid-length of oesophagus. Size of large rectal glands 0.245-0.272 by 0.231-0.245, of smeJl one 0.068 X O~O54:. Posterior end of body at l'ect9J. region gradually tapering to end of tail. Ta.U conical, relatiyely long (0.42.2), pointed.

Eost: Anguilla feinharrltii Strd. Localization: swimbladder. DiB~ribution: South Australia..

Comments: -This speoies wa.s described by Johnst,o.n and Mawson (1940) froul the long-finned 001, Anguilla reinkardtii, from PI'ospect Reservoir Ilear Sydney, New South. WaJes, Australia. Aocording to the original desoription, the gravid fem.ales of this species a.re 60-70 mm long &nd 1.5 mm wide, whereas the body of young fema.les is 25-30 mm long and 0.5 mm wide; the m.ale is 40 rom. long ,and 1 mm wide. This cor responds, more or less, to our iindings. In contrast to most other speoies of Anguillicola (except A. globic.eps), A. aUBtralien.81,·s B68ms to be oharacterized by a. relatively long and narrow body. Although Johnston and Mawson (1940) reported only four pairs of oa.udal papillae inthe male of this species, we found six pai:r:s; itis possible that the above mentioned· authors overlooked the adanal and first preBdlal pairs of papillae, that are less distinct than others. .

130 c

D E E r>l ci : :

c, :

Fig. 2. Anguillioola australiensi.s Johna~on e~ Mawson, 1940 - type speoimens. A- head end of gravid female; B- buccal capsule ofgravid female; 0 - head end ofmale; D ~ oaudal end offe male; E - posterior end of male; F ~ vulva.

131 A. australiensi8 has also been reported from Anguilla dieffenbachii and A. austral£S from :Xew Zealland (Brunsdon 1956, Rid 1973, Hine 1978, Boustead 1982, Blai r 1984) and from A •. anguilla from Ita.ly (Paggi et 801. 1982, Sarti et a1. 1985, Sarogli a. at aI. 1985). Although the nematodes from New Zealand have not been described and illustrated a.nd we could not obtain the specimens relating to these records, we consider them conspecific with the newly collected specimens from A ... australis from New Zea. land and those from A. anguilla from Italy that are now desoribed as a new species ~ A. novaezelandiae sp. n.T'he nematodes reported as A. a~t8trab;ensi8 by Hartmann (1987) from the FRG belonged probably to A. Gras8~l8.

8.AnguilUcola (Anguillicoloides) crassus Kuwahara., Niimi et Itagaki, 1974 Figs. 3,.4 Description (ba.sed on non-type specimens from A. japom'ca from Japan): Body da.rkly coloured, fusiform,ra.ther plump, ta.pering to both ends. Epicllticle almost

O.OSmm

EE]~.:...'. "'"'; .' . -;' 0,

Fig. 3. An{l'uillicola ora88UB Ku.wahara, Niimi eli Itagaki, 1974 - tYJ?e specimens. A, B_ head end of male and fe.ma.le; C - buooal capsule offamaJe; D- oaudal end of£emale; ]l ~ posterior end of male; F- taIl of ma.ls; G - larva from u~el'us.

132 smooth. Head end rounded. Mouth aperture oircular, surrounded by four big dorsolatera.l and ventrolateral cephalic papillae and two small lateral amphids. Buccal capsule well sclerotized, its anterior rim bearing one row of 22 rather big circumoral teeth. Oesopha.gus strongly muscular, expanded at its posterior half. Valvular apparatus of oesophagus well developed. Nerve ring situated approximately at border of first and second thirds of oesophagus length, excretory poI'e neal' junction of oesophagus andintestine. Intestine dark, straight, broad, only its anterior end being usually narrowed. Three conspicuously large, oval unicellular rectal glands present; a.dditiona.l small ventral rectal gla.nd a.lso

c ~]

O.OSmm E

E E ." Q

Fig. 4. Anguillicola crassus K:uwahara. Niimi at It.agaki~ 1974 - non·type specimens fro~ Japan. A- head end of male· B -buocal capsule of female; C- vulva; D- head end of gravld female; :El - posterior end of~ale; F- caudal end of female; G;.'- tail of male (speoimen wi~h only 5 pairs of caudal papillae).

133 present being frequently overh.pped by large ones and, accordingly, indistinct. T8Jl oonica.l, short. .. Male (21 specimens): Length of body 6.54-~I.76, m~xllnum ~dth 0.55.8-1.22. Bucca.l capsul9 0.021-0.027 long and 0.048-0.63 wIdej maXImum thIckness of Its wan 0.003 to 0.006. Length of oesophagus 0.721-0.843, its maximum width 0.190-..0.218; length of oesophagea.l valves 0.054-0.095. Length ratio of oesophagus a.nd body 1: 9.1-25.8. Distance of DIlrV'6 ring an.d excretory pore from anterior extremity 0.210-0.286 a.nd 0.694-0.924, respectively. Seminal v68i.c1e well dev.eloped, variable in size. Common cloacal duct opening on prominent process 0.048-0.090 long. Size of large rectal glands 0.122:-0.394x0.082-0.204, of small one 0.042-0.109xO.027-Q.068. Mostly abc pa,irs of oa.udal pa,pilla.e present: 2-3 preansJs, 1 adanal, a.nd 2-3 postana-Is. In one male only 5 papilla.e found (Fig. 4G). Tail conical, 0.109-0.240 long. Female (29 specimens): Length of body of gravid :females 13.08-44.74, maximum width. 1.22-3.50. Buccal capsule 0.02'7 long and 0.057-0.063 wide; ma.ximulll thickness of its wall 0.006. Length of oesophagus 0.85'7-1.088, its maximum width 0.204-0.272. Length ratio of oesophagus and oody 1: 15.3-40.3. Distance of nerve ring and excretory pore from anterior extremity 0.258--0.299 and 0.857-1.142, 1'espectively. Vulva. prominent, cone-shfloped, situa,ted in posterior part of body, 3.40-4.90 from posterior extremity, this repl'esenNng a.pproximately 1/4-1/10 of body length. Ovarian tubes d'eaching anteriorly to some dist,a:nce below oesophagus end level and posteriady to region in front of rflctal gla.nds. Uterus occupying most space of body, containing Il,umerous eggs, developing embry80, Bald fully formed, sheathed larva.e 0.244--0.258 long and 0.015 wide. Rectum hyaline tube, opening usually on well developed papHla. like projection; latter indistinct in. some specimens. Tail conical, 0.272-0.299 long.

Hosts: Anguilla anguilla (L.) (type h03b)e.nd A. iaponioa Temminck at Schlegel. Localiza.t.ion: switnbladder. Distribution: Ja.pa.TI, China. anc1 Europe.

Comments: - This sp,ecies was first described by Kuwaha.ra et aI. (1974) from pond oultured Anguz'lla angui'lla and A. japonica from Japan (Hamamatsu, Shizuoka. Pref. ~ Honshu). The original description of A. craBSU8 (erroneously named a,s crassa) was in adequate and inaccuratein some respects and thiswas the ma,n reason for doubts about. the species iden.ti:6.cat~on of conapeoifio Anguillioola nematodes reco:rded later from eels in Europe. Only recently, Ta,l'&siohew,ski c,t, at (1987) carried out a detailed comparison of the mOI'JIhology of the type (syntypes) andnon-type specimens of A. craasUs frolll. Japan with Ang'u/l,'Zlicola specimens ftom three localities in West Getll18ollY; they oonoluded. that all these nematodes belonged to one and the B&me species, A. craS8U8. ~oe the pa,pe! by Tar80sohewski et al. (1987) was submitted for publication, we ha,ve obtained a.dd~tio.nal numerous Japa.nese non~type specimens of A. CraS8'U8 for· study. An exarniD80tion of these specimens showed. muoh greater biometrical varia.bility o£ A. cra88'U8 from Ja:pan~ which is similar to that found by Tarasohewski at a.I. (1987) :fol' Enropean specimens (see Table 1). It ha.s already been mentioned by Tau,schewaki €It a.1. (1987) that in spite of trhe oonsiderable biometrical va.ria.bility of A. crassu8 the size of the buccal capsule remains muoh the sa.me (Table 1). According to these authors the oharacter of the buccal capsule (size, shape, degree of sclerotization) seems to be one of the most impol'tant I1pecifio features of.Anguilli·cola; the presentstudyshows th80tin additionthe positionofth6buocal loapsule in relation to the nematode's anterior extremity and the number of circumoral teeth may be ta~onQmicallyimport,ant. The number of circumoral teethin A. eras.SU8 wa.s not established by Kuwaha.ra at al. (1974:) in the oI'iginaJdescription, but seems

134 Table l. Compa.rison of Allfl'U,micola cra88& from Japan a.nd EurQ,Pe

Pa.ro.types Non'typo specimens from Japan European specimens (own data) (new da~a) (after Tarasehewski et, a1. 1987)

I I ~~ ~!j? ~~ !?~ it !? I' I _.~-- Length of body' 34.94 ' 42.94- I 6.54: -21.76 13.08 -44..74 5.77 - 23.12 16.23 -36.72

Width of body 3.'1 4.7 0.558- 1.22 1.2.2 - 3.50 0.340- 1.'17 L3G - 5.00

Length of buccal capsula 0.024 0.027 0.021- 0.027 0.02'1 0.021- 0.02'1 0.024- 0.021

Width of buccal capsule 0.048 0.054- 0.04S- 0.063 0.057- 0.068 0.048- 0.057 0.054- 0.063

Lcnght of oesophagus 0.760 0.980 0.721- 0.843 0.867- 1.09 0.571- 0.8]6 0.775- 1.06

Width of oesophagus 0.260 0.330 0.190-- 0.218 0.204.- 0.272 o. I35- 0.258 0.258- 0.381 Lengthratio of oesophagus and body 1:43.3 1:43.7 1:9.1-26.8 1:15.3-40.3 1:10.1-29.2 1:20.D-34.v No. of caudal papillae in male () - 6 - 6 - Distance ofvulva. from pos~el'ior end - ? ...... 3.4.0 - 4.90 - 4.01 - 7.01 Length of ~ail 0.207 0.231 0.109- 0.240 0.272- 0.299 0.]20- 0.286 O.13n- 0.448

Hosts Anguilla anguilla Anguilla jtvpOnica A ngtdlla an(fu£lla

Ocourrence 'Japan Japan· F.R. Germany ._"-.-.--

...... W ~ to bo rather va,riable in this species. Wang and Zhao (1980) report-e,a 25-26 big teeth in Far.easter.u A. c.raS8US, while only 21-25 Meth were found l1l the conspecific: nematodes from Belgium (De Chatl,eroyeli aI. 1987); according to Taraschewski et a!. (1987) the specimens of A.. c.raS8U8 from West Germany possess 22-28 teeth; in ona non-type specimen from Japan we found 22. t~h. . It ha,f) been lnOOl,tioned that .A. cmS8'll8 was orlgmaJly descrIbed irom Japan by Kuwaha.ra. at aI. (1974), but this species was found in Japan a.s ea.rly as in 1969 by Egusa et al., who ha.d mistaken it for A. globicep8 (see Hirose et al. 1976). Lat,er, this species was also recocded from Ja.pan by Hirose eli .Ill. (1976) and Egusa. (1979); the latter author memions that tihe introduced European eel, AngM'lla anguilla,.is, ~n Japan,. much more susceptible to A. craS8U8 infection than A. japonica (prevalenoe in cultured eels being several tens of per cent ilond ooca.sionally neatly 100% inA. anguilla and 10-40 % in A, jarponica). This species wa.s also present in the Afi(Ju~1lu'ola material from China provided by Prof. P. Q. Wang and Prof. H. S. Wu. Recently A. cmS8US has been reported from Japan by 8alati (1987). In Europe, A. Cfa88U8 was first recorded as Anguillicola SPA from eels (Anguilla anguilla) of the Wes,er-Ems River region in North Germany in 198¥ (Ne 1] mann 19'85, Mann 1986). According to Peters and Hartmann (1986) and Taraschewski et al. (1987), it now occurs frequently in other localities in the F. R. of Germany (river basins of the Elbe, Weser and Rhine) and in the ,vater bodies of Berlin. It occurs also in Holland (Van Banning at at 19'85), Belgium (Belpaire et al. 1987, De Charle royet a.l. 19'87) and the record of Aguillicola sp. in eels in Denmark (Keie 1987) apparently relates to this species; recently it has been recorded also fro.m eels in Italy (fish farm in the Po delta) (Canestri-Trotti 1987) and Great Britain (personal communioation of Prof. C. R. Kennedy). It seems that A. c.ra88U8 wa.s introduced here .only a few years ago and is now quickly spreading mainly throughout North-western and Central Europe.

4. Anguillicola (Anguillicoloides) novaezelandiae sp. n. Figs. 5, 6

Description (based on speoimensfrom A. australis from New Zealand): Medium. sized" darkly coloured nematodes. Epicuticle aspmose. Heald end with slight neck constriction just in front of nerve ring. BuoGal CRpsule smaIl,feebly sclerotized, with minute circumoral teeth; exa.ct number of teeth not established (32 in Ita.lian specimens accord ingto Paggi et aI.1982). Oesophagus expa.nded at its posterior h&1f. Valvular appa.ratus of oesophagus well developed. Ne1'Ve ring situa.ted immed~ately below neck constriction, excretory pore near junotionofoesophagus andintestine. Intestine da.rk, almost straight,. broad. Three conspic'llously large oval unicellular reotal glands present; additiona.l sma.ll rectal gland, often overlapped hy large ones, also present. Tail conioal, pointed. 1\faJe (4 speoimens; measurements of holotype in brackets): Length of body 5.5~8.57 (8.57), maximum width 0.476-0.680 (0.558), length of sllghtly enJarged head end 0.111-0.132(0.120), its width 0.087--0.111 (0.093); width of body a.t nook constriction 0.078-0.099 (0.078). Buccal capsule 0.006 (0.006) long and 0.021 (0.021) wide, maximum thickness of its wall 0.002 (0.002). Length of oesophagus 0.476-0.530 (0 .. 530), its maximum width 0.095-0.120 (0.109). Length rat~o of oesopha.gus and body' 1: 10.4 to 16,2 (1: 16.2). Distance of nerve ring and exoretory pore :£tom anterior extremit,y 0.15,9-0.165 (0.15H) and 0.462--0.585 (0.585), respecti,vely. Common oloacal duct opening on prominent proces~ 0.087-0.0916 (0.087) long. Size oi large rectal glands 0.272--0.354 X 0,136-0.231 (0.272-0.340 X 0.136), that of small one 0.033-0.039 by 0.030-0.033 (0.039 X O.OBO). Six pairs of caudal papillae present: 3 pl.'eanals, 1 &da.na.l, and 2 pOistanals. Ta.il conical, 0.156--0.195 (0.156) long.

136 Female (3 speoimens; measurements of allotype in bracketis): Lengtl1 of body 7.17 to 13.63 (13.63), illtmmum Width 0.653-1.18 (1.18); length of sligthly enlarged hea.d end 0.123-0.135 (0.132), its Width 0.090-0.102 (0.102); width of body a.t n.eck oOll..'ltI'iction O.06~O.078 (0.078). Buocal ca.psule 0.006-0.009 (0.009) long and 0.018-0.021 (0.021) wide; maximum thickness of its wa.ll 0.002-0.003 (0.003). Length of oeBophagus

A B c

E F ~I

Fig 5 A.nguillicola. 'I1ovaezelamdiae sp.n. from A. autitraUs from New Zeabmd. A, B- heia.d e~d.of maie ~nd gI'a.vid female; O. D -,- buccal ca,psule ofmale and young fema.le; E --:- vulva; F - pas ,enor end of male; G- cauda.l end of female; H -la.rvadrom. ut'6ruS.

137 0.544-0.625 (0.625), its maximum width 0.136--0.177 (0.150). Length ratio of oesopha. gus and body 1: 13.2-21.8 (1: 21.8) .. Distance of nerve ring and excretory pote frOlll anterior extremity 0.165-0.195 (0.195) and 0.558-0.680 (0.680), respectively. Vulva elevat.ed, prominent, situated in pos.terior part of body, 1.84-2.08 (2.08) from postedoir extI'emity~ this )'epreoonting approxima.tely 1/4--1/6 (1/6) of body length. Ovarial tubes not reaching anteriorly level of oesophagus. Uterus conta.ining mostly ,eggs, se\'era,} sheathed larvae, 0.249-0.255 (0.255) long and Q.015 (0.015) wide, present only near vulva. of two larger females (body length 9'.6 and 13.6). Size of large rectal glands 0.204-0.275 )<0.136-0.245 (0.272 x 0.217-0.245), small rectal gland indistjnct. Ta.il conica.l, 0.186-0.225 (0.186) long, pointed.

A c ,~

!!d H

F G

Fig.6. AnlJ'Uillicola novaezela71dia,e Bp. n, - specimens from A. anguilla from Ita.Iy. A _ bea.d end of mde; B- head end, apical view {re·drawn from Paggi \'It a1. 1982); 0 ~ head end of female; D. E -. buceal capfmle of male and female; F- oaudal end of fem.ale; G --'- v1.11v8;H ~ postel'iol' end of male. '

138 Table. 2. Comparison of AnguillicDJa flDroezelandiae sp.n. from Now Zealand and Italy Specimens from New Zealand I European specimens I (own data) (own data)

_ . M I w I M I w I

Length of body. 5.54 -8.5'1 I 1.11 -Ia.63 11.63-14.92 24.82 -28.70 !, Max. width of bony 0.476-0.680 0.653- 1.183 1.41- l.oG 2.00- 3.00

Length of buccal capsule 0.006 0.006- a.OOg 0.007 0.007- 0.009

Width of buccal capsule 0.021 0.018- 0.021 0.024 0.024--- 0.027

Length of oesophagus 0.476-0.530 0.544- 0.625 O.ti39-0.6lHi 0.6153- 0.843

Max. width of oesophagus ·0.095-0.120 0.130- 0.177 0.136-0.163 O.204--0.24G

Length ratio of oesophagu.Cland body 1 : 10.4-16.2 1 : 13.2-21.& I: 18.2-22.S 1 : 37.0--38.0

No. of caudal papillae in male 6 ,,~ {) Distance ofvu1vtl. from polilterior end - 1.84: - 2.08 4.42 -- 4.7G Length oftail 0.156-0.195 I 0.186- 0.225 O.HlO 0.204- 0.272

Hosts A. C&mtrali8 A. anguilla

Oocurrence New Zealand Italy

~ 1:.<3 co Bosh: Short.finned eel, A'IU'ldlia o'U8tralia Richard.son (type host.) and European eel, A. anguilla (L.);. probably also in long-filmed eel, A. dleDenbachii Gray' Localization: swimbladder. Type locality: Matahina dam, New Zealand (171tla~ 1975., call. Dr: ~.~ones); other localHies: Lake Bracaiano near Rome, Italy; probably also ill other localltlBs In New Zealand reported by Brunsdon (1956) .Riel (1973), Hine (1978) and Bous t:ead (1982),. Deposition of ~ypes: holQtYP? (0'). a.IlotYP.8 (~) an,cl ~aratyp8ls \r3 + ~) in Institute ofParasitology. Czechoslovak Academy of SClences. Ceske BudoJovlce (Cat,. :N o. N-263). E,tymology: The speoifio name of this nematode speoies relates to the countt'y of the origin of type specimens. Comments: -A. novaezelandiae sp. n. is most similar to A. a'UStmlz"enais Johnston et Mawson, 1940. However, it differs from the latter speoies mainly in the :shape of the head end which is bulbously infla.ted, almost spherioal, and followed by a marked neck ~s constriction in A. australlenaisJ whereas it only slightly expanded in .A.. novaezelandiae sp. n.; the anterior ova.ry in A. australiensis females extends anteriorly to about the mid-length of the oesophagus, while it does hot reach the end of oesophagus in A. novae zelaniUa.e sp. n. Both the sp'eoiea differ also in the size and form of the body. While the body of A. austrah"en8Z8 is long (30-40 111ill in malesaud 60-70 mm in gra.vid females) and reJla.tively slender (at most 1.5 mm in gtavid fema.les), uha.t, of A. novae~elandiae iSP' n. iiS much shorter (6-15 mm in males and 10-31 mm in gravid females) and wider (up to 3 rom in graVid females); the type A. a'l.l.3traliensis female examined (27 mm long and 0..72 mm wide) was still non-gravid, without eggs in hs uterus. The shape of the posterior end of the female body seems also to be different in these two species. The specimens reported asiA. australiensis by Paggi et a!. (1'982) from A. anIJuilla from Ita.ly are somewhat lal'ger than those from A. au.stralis fJ.'om New Zealand (see 'T,a.bl'e 2), but they are m'orphologicaUy indistinguishable and, therefore, we consider them conspecific with A. novaezelandiae sp. n. It is probable that this .new species was introduced in Ualyalong with the transfers o.f live eels Anguilla australis from Ne w Zealand; a stock of A. australis was introduced into Lake Bracciano in 19'75 (Wel~ comme 1981, Paggi at aI. 1982). Aocording to the ptesent dat.a., A. novaezeland~'ae sp. n. is proba.bly indigenol.1B to New Z;ealand; from where it has recently been introduoed to Italy (Paggi et &1 1982, 8aroglia et al. 1985,Di Cave 1986, Sarti 1986). From New Zealand, another Ang1.till·£eola species, A. australiensis, hais so far been recorded from Anguilla dieUen- .bachii and A. australis (Brunsdon 1956, Rid 1973, Hewitt and Hine 1972, Hine, 1978, Boustead 1982); however, since no description or drawings of these parasites were provided, it may well be that they belonged in the fact to the species A. novae zelandiae sp. n. and that A. aust'iah'ensis is restricted in distribution to the territory of Australia only.

5. Anguillicola (Anguillicoloides) papernai sp. n. Figs. 7, 8

Description: Body spira.lly coiled, da.rkly coloured (due to eel blood inside nema.tode's intest,ine), i~orm, l'atherplump, tapering to both ends; a.n.1ierior end of body bottle shaped, posteriOr one na..rrowed, conica.l. Epicuticle finely Wl'inkled, with networkstruo tore" Mpinose, forming irregular fine transparent coating. Cuticle of anter'io),' and poste~or nDil'l'owe~ pa.:-ts of body bea.ring severallnarked, up to 0.030 high, irregularly ,scattered dull pap~lla-like excrescenoes of fibrous st:rnctu'OO' sometimes these format&ons i1!dis~inct on female po~teriOI~ end. Head end rounded, bain~ sepaI'ated by slight cotl,Btrio tlOn 1~ front of nerve rmg level; bo~y of son:e specimens somewha.t infl.a:ted in IJegion of po~terlOr half o£oesophagus. EntIre antel'lOr nal'1'owed part of body appl'oximately tWIoe &S long as oesophagtls. FoUl' dorso- and ventro-Iateral papiHae and two sma.ll

140 A B

O,1mm .J J

Fig. 7. Anguillicola papernai sp. n. from A. mossambioa from South Africa. A, B- head end of gravid female and male; C- anterior end of female; D - bucoal capsule ofmale; :m - head end of female, apical view; F- buccal oapsule of female, apical view; G - female, general view; H, I larva and egg from uterus; J- vulva.

141 r A I l / I I

E ..,.E d

Fig. 8. Anguillicolapapemai Bp. n. fro.OlA. m08sambica from SonthAfrica. A- posterior end ofmale ~ B, C - caudal end of female; D- posterior end of male; E -poa~eriorend of fema.le.

142 lateral anl.phids present. Mouth conSpiC11011sly depressed, anterior margin of buccal ca.psule being 0.027-0.030 from anterior extremity; mouth opeID:ng circular. Buccal capsule well sclerotized, trapezhull-shapedin lateral view, its a.nterior rim beating one tow ~f 26 circu~o~al teeth: Oesophagus strongly muscular, expanded at its posterior half; Its lumen dIstlllctly trIangular. Valvular apparatus of oesophagus well developed. Nerve ring situated immediately below neck constriction, excretory pore near junction of oesophagus and intestine. Intestine da.rk, almoslJ stra.ight, broad, being narrowed at a.nterior and posterior ends only. Three conspicuously large oval unicellular l.'ectal glands present, one dorsal and two subventra.l; additional small ventral rectal gland, often overlapped by large ones, also present. Tail conical, short. Male (1 specimen and 1fragment of another one; measurements of holotype in brackets): Length of body (12.99), maximum width. (1.50); length of head end (0.109), of entire narrowed anterior part of body (1.22); width of body at head constriction (0.060), at end of naITowed anterior body part (0.272). Bucca.! capsule 0.011-0.012(0.011) long and 0.030 (0.030) wide; maximum thickness of its wall 0.003-0.004 (0.004). Length of oesophagus 0.530-0.598 (0.530), its maximum width 0.150--0.163 (0.163). Length ratio of oesophagus and body (1: 24.5). Distance of nerve ring and excretory pore froUl anterior extremity (0.180) aud (0.667), respe~tively. Testis reaching anteriorly to some distance below end of narrowed a.nterior part of body. Seminal vesicle well developed, rather long. Common cloacal duct opening On prominent process (0.048) long. Size of large rectal glands (0.105-0.120 X 0.054-0.057), that of small one (0.039 X 0.030). Six pairs of caudal pa;pillae present: 3 prea.nals, 1 adanal, and 2 postanals. Tail conical, (0.141) long. Female (4 specimens; measurements of allotype in brackets): Length of body of gravid females 15.78-28.29 (28.29), maxiullun width 1.64--2.18 (1.64); length of head end 0.081-0.099 (O.099), of entire narrowed anterior pari; of body 1.43-2.07 (1.64); width of body at head constriction 0.060-0.068 (0.063), at end of nal'rowed anterior body part 0.408-0.571 (0.408). Buccal capsnle 0.009-0.012 (0.012) long and 0.030 (0.030) wide; maximulIl thickness of its wall 0.003-0.005 (0.005). Leng~h of oesophagus 0.598-0.625 (0.625), itsmaximum width 0.135-0.177 (0.135). Length rat~oofoesophagusand body 1: 24.2-45.3 (1: 45.3). Distance of nerve ring and excretory pore from anterior extremity 0.147-0.171 (0.147) and 0.692-0.695 (0.695), respectively. Vulva elevated, prominent, situated in postetior part of body, 3.20-3.74from posterior extremity, this repr"esenting approxbnauely 1/5-1/8 of body length. Postvulva,r part of body markedly tapered. Ovaria.l tubes forming numerous· coils in anterior hal£ of body, anteriorly not reaching to narrowed part of body. Uterus occupying most space of body, containing numerous eggs (size 0.081-·0~105 X 0.075-0.084), developing embrya, andfully formed, sheathed larvae O.189~.255 long and 0.018-0.021 wide. Uterus extending anteriorly to end Ot narrowed anterior. body part and posteriorly reaohing almost level of rectal glands. Rectum thinhyaline tube, anus not elevated. Size of large rectal glands 0.120-·0.180 ~y 0.063-0.090 (0.150-0.180 X0.090), that of small one 0.039-0.045 X0.030-0.033 (O.045XO.033). T~il oonical, its tip sharply pointed or somewhat rounded; length of tail 0.180-0.204 (0.180). .

Type hoatl Anguillamo8sambica Peters. LocalizaMon: swimbladder. Type looality: Amalynda fish farm near East London, Cape Province, Rep. of SO\lth Afrioa (21 September 1984; 0011. Dr. I. Paperna). .,. Deposi:tion of types: holotype (&,), allotype (~) and 2 paratypes(~~) In InstItute of ParaSItology. Czeohoslovak Academy of Sciences, Ceske Budejovice (Oat. No. N-279); 3 paratypes (1 male trag. + 2 ~~) in Na~ional Animal Pa.rasi~e Laborat.ory, Maryland, USA (Ca~. No. 80007).

143 Etymology: This Bpecies is named in honour of Dr. Ilan Paperna, the Hebrew University of Je:rusalem, who contributed greatly to the knowledge of fish parasites. Comments: - The unique morphological features of .A.. papernai sp. n. by which this species differs dis.tinctly from its congeners are Tihe presence of ll1atked cu.ticular ex~ crcscences on the anterior and posterior ends of the body and the location of the buocal capsule deeply inside the head end. This species is the first Anguillicola member known from Africa.

KEY TO THE SPECIES OF ANGUILLICOLA: 1 Epicuticle finely spinose. Oesophagus conspicuously infla1:€d at anterior end, its posterior part cylindrical. Buccal capsule large, 0.068-0.109 X 0.272-0.408 lllll.l in male and 0.109-0.122 X0.408 mm in female, with about 80 very small circumoral teeth. Ta.il rela.tively short. In eels of East Asia. (Japan, Ohina) ... (subgen. A'I1{}uillicola)...... A. globiceps - Epicuticle aspinose. Anterior end of oesophagus la.cking infla.tion, nal'I'OW; posterior part of oesophagus distinctly expanded. Buccal capsule small or medium sized, less than 0.030 mm long and O.070mm wide; maximum number of circumoral teeth 32. Tail rela.tively long. In eels of East Asia, Australia, New Zealand, Africa. or Europe .••

(subgen. Anguillicolooz'de8). . +•+•••••••••••••••••••• 2 2 Buccal capsule medium sized, 0.021-0.027 mm long and 0.048-0.063 mm wide, with 21-28 ra.thet' big oircumoral teeth. Head end noti expanded, wiTihOllt distinct neck constriction. Body fusiform. East Asia and Europe A. crassuS - Bucca.l ca.psule sma.ll, at most; 0.012 mm long and 0.033 mm wide; circumoral teetih very small. Hea.d end bulbously inflated or at least slightly expanded with distinc't neck constriction. Body fusiform or filiform. Austl'alia, New Zeala.n.d, Africa. or Europe ; ...... 3 3 Body fusiform, anterior and sometimes also posterior ends of body with marked pa.pila-like cuticular excrescenoes. Head end slightly expanded. Buccal capsule located deeply inside head end, with 26 minute teeth. Mrica .... A. papernai sp. n. -. Body filifol'm of fusiform, c'Uti

ACknowledgements. Our thanks are due to the following persons who kindly provided us with the Anguillicola specimens and thus made tho present work possible: Dr, H. Hirose, Nihon University, Toky:o, Japan; Dr. B. Jones, Fisheries Research Centre, Miniatry of Agricult'llI'e and Fisheries, Welhngton, New Zealand; Dr. D. C. Lee, South Australian Musetun, Adelaide, Australia; Prof. L. Paggi, Institute of Parasitology, University of Rome, Rome, Italy; Prof. I. Paperna, Department ofAnimal Sciences, Hebrew University ofJerusalem, Israel; Prof. F. Q. Wang, Parasitology :Research L.a~oratory, Fujian.Normal Universi~y, Chh~a; Prof. H. S. Wu, Institu.te ofHydrobiology, Academia Sllllca, Wu~an, Chma; and Dr.. S. Kamegtu, Maguro Parasitological MUEl6Um, Tokyo, Japan. Saxne nec~ssary hterature waS provided by Prof. P. Ghittino, Instituto Zooprofllactioo Sperimentale. Tonno, Italy. We are also very grateful to Prof. a. R. Kennedy, Department of Zoology, University of Exeter, UK, for his kindness in correcting the English translation of the manuscript. • 144 PEBII3I1fI POAA ALVGUILLICOLA YAl\fAGUTI. 1935 (NEMATODA: ANGUILLICOLIDAE) - I1APA3H:TOB ILHABATEJIbHOrO HY3blPH ~T r PE tt C 0 IT IICAH 1'1 E M: ,II; B "Y X HOB bI X 131'1 P; 0 B _ A. NOVAEZELANDIAE SP. N. 1'1 A. PAPERNAI SP. N.

(J). j\IopaBeI~ Hr. TapallIeBeIUI

Pe3IOllte. I-I a OCHOEe TIIHOEbIX SH3eMIIJIHpOB II APyrnx MIlTeplIu.n:oB CJ~eJHI;IlI peBu3uro pOAIl Anguillicola. OmrcaHr,r ,[l;Ba HOBbIX BI:l:,r~a: A. novaezelancliae sp. n. OT )'l'peii Anguilla australis (TlmoBoii: x03lum) ll3 HOBoR 3eJU1HJ.:uur II A. anguilla 113 IhamlII (nOMa'l'OiJ,bI ::lTOI'O BlIAH PUHbUIe rrpIIBoglIJIIIC-h 1I3 I1TaJIlHr nop; Ha3BaHfreM A. australiensis) II A. papernai sp. n. OT Anguilla mossamb iea li3 IOamoii: AepPliHH. B rrUCTomu;eo BpeMfI H POAY Anguillicola rrupHaJVW)I

CJI8P,YIOIIl.UX BRAOH: A. globiceps Yamaguti j '1935, A. australiensis Johnston et Mawson, '1040, A. craSBUS Kuwahara, Niimi et ltugaki, '1974, A. novaezelandiae sp. n. l{ A. paper/wi sp. n. 3a lIOCJIe):{Hne lIJiTb neT 6bIJIH aUBoaOH:&I B EBpony 113 APyrnx I{QHTHHOHTOB ADU Bu.n;a pOAR An guillicola ~ A. crassus II A. novaezelancliae sp. n. POA Anguillicola pa3HCJIeU lin nBll no/wo )'ill - Anguillicola (nm013oii: DlIl.( A. (A.) globiceps) II Anguillicoloides subgen. n. (TlInoBoii BHil; A. (A.) crasslLs). TIpnBc;r.cHJJI RopoTRue omfcalIlUI II pneylIJ{H D('CX BlI/lon p..A nglLillico!a, a THIORC onpCACJIHTeJIbH1UI Tu6mlIl;a AmI HA8HTucIlIlH:aIJ;InI BIll\OB :3Toro POj\H.

REFERENCES

ANDERSON R. C., 1958: Methode pour l'exa Working Party on Eel, Bristol. UK, 13-16 men des nematodes en vue apioale. Ann. April 1981, 7 pp. Parasit. Hum. Comp. 33: 171-172. Dr CAVE A., 1986: Anguillicola australiensis BELPAIRE 0., DE CHARLEROY D., THO Johnston et Mawson, 1940 parasite. eli MAS K., 1987: Etfeo~s of res~ooking eels Anguillcb anguilla: consic1erazioni sullo. com· on the distribution of the nematode Anguil pal'sa in Italia e suI cicIo biologico. Boll. licola crassa in Flanders, Belgium. European Inf. Aoc. It. Anguilla. 6; 18-21. Inland Fisher.· Adv. Oommission, EIFAC EGUSA S., 1979:.Notes all the culture of the Working Party on Eal, Bris~ol, UK, 13-16 European eel (Anguilla anguilla L.) in Japa April 198'1, 6 pp. nese eel.farming ponds. Rapp. P.-v. Reun. BLAIR D., 1984: A cheoklist and bibliography Cons. into Explor. Mer 1'14: 51-58. of parasites of New Zealand freshwa~er -'-, IURAK., WAKABAYASm H., 1969: fish. Mauri Ora 11: 5-50. On the occurrence of Anguillicola globiceps )30USTEAD N. C., 1982: Fish diseases recorded Yamaguti, a swimbla.dder roundworm, in in New .Zealand. with a discussion on po~n pondoultured eels. Fish Pathol. 4: 52-58. tial 801.11'0eS and oertification procedures. (In Japanese.) New Zealand Ministry of Agricult. and GHITTINO P., 1985: Tecnologia e patalogia Fisher.• Fisheries Res. Div., Wellington, in acquaooltura. Vol. 1, 2. I'.rivate press, Occasional PubL No. 34, 19 pp. Torino. (Ex Canestl'i-Trotti 1987.) BRUNSDON R. V., 1956: Studies on nerna.toc1e HARTMANN S., 1987: Schwim,mblassenwftrmer parasites of New Zealand fishes. Ph.D. beim Aal. Fischer & Teichwirt 38: 2-3. thesis. Victoria University of Wellington. HEWITT G. C., HINE P. M., 1972: Checklist (Ex Blair 1984). of parasites of New Zealand fishes and dANESTRI·TROTTI G., 1987: Occurrence of of their hosts. N. Zeall1nd J. Marine and the nematode Anguillicola cmssa Kuwahara, Freshwater Res. 6: 69-114. 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Australia 64: on Eel, Perpignan, September 11-18, 340-352. 1985,5 pp. KOOPS H .• 1981: Untersuohungen zum Befall SARTI l\f., 1986: Signi6cato patologico e 2'000 von Farmaalen mit Anguillicola. Inf. f. d. tecmico di Anguillicola australiensi8 e stato Fisohwirt 4: 114-171. delle ricerehe sulla terapia. Boll. lng. Ace. KalE M., 1987: Eel parasi~es in freahwa.~er It. Anguilla 6: 22-25. anci marine habitats in Denmark. 2nd -, GIORGETTI G., BRISINELLO VF. , Inter. Symp. Ichthyoparasitol., Sept. VANELLI M., 1985: Anewprableminint.an 27-0c~ob. 3, 1987, Tihany, Hungary, Abs· sive Italian eeloultur,e. The parasite An(Juil~ tracts of papers and posters. p. 4:B. licola australiensis.Post:el' paper presented KUWAHARA A .• :mIMI A., ITAGAKI R,. at the 2nd EAFP Inter. Oomerenoe, Mont 1974: Studies ona nemat:ode parasitio pellier, 2-5 Sept. 1985. (Ex Saroglia in the air bladd.er of the Ellal I. Description et a1. 1985.) of Anguillicola ,era8sa n. sp. (Philome~:ddea; TARASCHEWSKI H., MORAVEC F., LA~ Anguillicolidae). Jap. J. Parasitol. 23: MAR T., ANDERS K., 1987: Distribution 275-279. and morphology or two helminths recently ~IANN H., 1986: Aalpa:raBit~ entdeekt.. Fisch introduced into European eel populations: und Fang 8: il. . Anguillicola cra88U8 (Nematoda., Dracunau~ MOLNAR K., 1986: Solving para:site-related loidea) and Paratenui:sentis (J/mbiguu8 (Acan problems in cultured freshwater £sh. Proc. thocephala., Tenuisen~idae). Dis. aquat. Vlth ICOPA, Australian Aoad. Slli .. , Canber Org. ::I: 167-176. ra, 'pp. 819-326. VAN BANNING P., HE'ERMANS W., VAN NEUMANN W., lQBa: &hwimmblasenparasi~ WILLIGEN J. A., 1985: Anguillicola crassa, Anguillicola bei Aslen. Fischer wId 'Teioh· een ni.euwe aalparasiet in de Nederlandse wirt 11: 322. wateren. Visserij 38: 237-240. PAGOI L., OREOOHIA P., MINERVINI R., WANG P .• ZHAO Y., H180: Observations on the MA.TTlUCC! S., 1982: SuIls, comporsa life history of Anguillicola globiceps (Nema di Anguillicola australien.sia Johnston €It , toda: Ariguillicolidae). Aota Zool. Siuioa 26: Mawson, 1940 (Dracunculoidea: Angun. 243-249. (In Chinese, Eng!. summary.) licolidae) in Anguilla anguilla del lago di WELCO:MME R. L., 19B1: Register 'of inter Bra-oamno. Parassitologia 24: 1391--144. national ~ransfers of inland fish species. PETERS Go, HARTMANN F., 1986: AnguU FAO Fish. Tech. Pap. No. 213, p. 3. licola, a parasHic nematode of ~he swim "\OVU H. S., ·1956: Studies on the parasitio bladder spreading among eel populations nematodes of freshwater fishes in Ohma 1. in Europe. Dis. aqua:t. arg. 1: 2:29-230. Acta Hydrobiol. Sinion 1: 99-10t\ + 3 PIta. RID L. Eo, 1973: Hehninth parasites of ~he (In Chinese, Engl. summary.) long-fumed. eel, Anguilla dtieUenbarihii, and -. 1984: Theparasitio nematodes of :fishes ~he short·finned eel, A. australis. Mauri from Liao He in China. In: Parasiticorga~ Ora 1: 99-106. niems of freshwater :fish of China (Ed. by SALATI F., 1987: La paraasitosi da Anguillicola :met. of Hydrobiology, Aca.dem~a, Sinica,.), sp,in Giappone.. Riv. It. Piscic. lttiop. 22: Agri.oult. PubI. House, Beijing, China, pp. 115-117. 177":-200. (In Chinese, Eng!. abstraot SAROGLIA M. G., FANTIN P., ARLATI G., separately on p. 29.) 1980: Eel production in Italy-Problems YAMAGU'l'l S., 1935: Studies on the helminth and perspectives. European Inland Fish. fauna of Japan, Pt. 9,. Nematodes of fishes, Adv. Commission (FAO), Working Party 1. Jap. J. Zool. B; 337-'-386. .

Received 80otober1987 F. M.• Parazito1ogicky ustav CSAV. Bra,niAovska.:n, . 370 05 Ceske Bl.1dfSjovioe, CaSR

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