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

REVISION OF THE GENUS ANGUILLICOLA YAMAGUTI,1935 (NEMATODA: ANGUILLICOLIDAE) OF THESWIMBLADDER 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 Institutoof Special Zoology and Parasi~ology,Ruhr University, Boehum

Abstract. The genU£! Anguillicola is reviewed On the basis of the examination of some type specimensand 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'iginallyreported from Italy as A. australien8is) and A. papernai sp. n. from Anguilla m088ambica from SouthAfrica. At present the genus Anguillicola comprises the following five valid species: A. globicepsYamaguti, 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. Thegenus AnguillicoZa is divided int.o two s\1bgenera: An(Juillicola (type species A. (A.)globiceps) andAngw"Zlicoloides subgen. n. (type species A. (A.) ara881As). All Anguillicolaspeeies are briefly describedand illustrated. The paper is supplemented by a key to the identifica~ionof 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 theswimbladder of Japanese eels (Anguilla }aponica) in Japan. Later two additionalcongenerio species were described: A. australiensis Johnston e"t Ma.wson, 1940 fromAnguilla reinhardtii from Austra.lia and A. cra88U8 Kuwahara., Niimi et Itagaki, 1974from Anguilla anguilla and A. japonica raised in eel fa.rms ill Japan. The species weredistinguished prinoipally by lihe form of the oesophagus, number of large unioellularreotal glands, and humbet of ca.uda.l papillite in the 11180le (see Pe1iel's a.nd Hattmanu1986). Until recently, all the three a.bove mentioned species were considered to be limitedin 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 sinceAngwillicola appeal'ed in Europe. In 1982, P aggi etaI. reported Anguillicola australiensisfrom AngUilla anguilla from Lake Braociano neal' Rome in Italy. At about the sallie timeAnguillicola nematodes were recorded from eels from the open wa.ters of the Weser-EmsRivet region in Northern Getmany (Neumann 1985); since then, "they have spreadthroughout many countries of WesteI'n1 Northern and Central Europe with grea.trapidity. Although the nematodes from Westem Europe (Holland, Belg~um) wereidentified 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 be11l1eqllivocally 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. cra88U8 1 but the authorsmention 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 Anguillioolamembers js responsible to the fa-at that these highly pathogenic nema.todes are frequentlyreported 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 ofAnguill'£cola, we examined a number of the type and non-type spe~imens of all threepreviously recognized species in this genus. SOIlle of the results obtained pertaining tothe European members 'of AnguilUcola have been published by Taraschewski et a1.(1987), who showed clearly that there are actually two species occurring in eeLo; inEurope. Since, however, some new and important data on the morphology of Anguillicolamembers of other continents wete obtained during this study, it proved possible toundertake a. revision of the whole genus. It is hoped that this will help in relia.ble speciesidentifica'tiolJ, and will fOl'm the basis for future work on, th0!3 biology of these interestinga: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~hAustralia (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 ofGermany (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 (21Sep~ember 1984, leg. Dr. I, Paper~a).

The nematodes wElre mos~ly :fixed in 70% ethanol.or 4% formaldehyde. All the speoimens werecleared and examined in glycerine. En face views were prep!llred according to Anders on's (1958)method. Drawingswere made with the aid of a Zeiss microscope dra.wing attaohment. Allrneasure­menta are in nun.

RESULTS

Gen.us Angnillicola YamagutiJ 1935

Diagnosis: AnguilHcol'dae. Cuticle very finely spinose or a.spinose, sometimes withmarked, irregularly scattered exol'esceno.es of fibrous structure on. a.n.tetior a.nd posteriorends 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 andwide, consisting of antel'ior muscular and posterior muscular-glandular sections. Vslvulara.ppa,fatus well developed. Intestine dal'k, distended. Rectal glands Htl'ge. Male: Spiculesabsent, Testisbeginntng near tail end; seminal vesicle well developed. DuctusejacuIa.torius opening outside on prominent caudalprocesB. Six pairs of sessile caudalpapillae present. Female: Vulva opening on tip of prominent cone in posterior Pal't ofbody. 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 AnguiUicoletspecies in some taxonomically -very ixnporta.nt features (structure of oesophagus, spinosecuticle)" 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 A nguillicolaspecies.

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: cylindticaLType species: A. (A.) globiceps :yamaguti, 1935

n. Subgenus AnguiUicoloides subgen. n.

Diagnosis: Anguillicola. Body fusiforni or filiform, cuticle aspinose. Anterior end ofoesophagus 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,adend rounded, buccal capsule very large, well sclerotized; some 80 smaH circumoralteeth present. Anteri01'! end of oesophagus bulbously inflated" posteri,or part of oesopha­gus cylindrical. Valvular apparatus well developed; Tlu:~ee latge and oue small unicellularrecta.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 thickness0.OO~0.Ol5. Length o£ oesopha.gus 1.25-1.50; length of its anterior bulb 0.408 to0.544, 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 excretorypore from anterior extremity 0.571-0.775 and 1.27-1.37, respeptively. Commoncloacal duct opening on prominent prooess 0.068-0.109 long. Size of larger recta.l glands0.109-0.312 X 0,082--0.109, that of SillBlUer one O.068~0.081X 0.030-0.041. Sixpa.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 width1.44--1.54. Length of buccal capsule 0.109-0.122, width 0.408, maximum thicknessof its wall 0.009. Length of oesophQlgus 1.50; length of its anterior bulb 0.049, width0.571-0.639, maximllm width of its posterior cylindrical part 0.299--0.313. Lengthra.tio of oesophagus and body 1: 24.9-25.3. Dis,tance of nerve ring and ex.oretory por'a0.74:8-0.762 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 6.99-8.43from posterior extremity, this representing approximately 1/5 of body length. Uter-L"lSnot rea.'Jhing an.te1'iorly oesophagca.l region; utel'Us containing numerous eggs (size0.078-0.081 x O.069), developing embl'ya and fully formed" sheathed larvae sou:1.e0.180 long and 0.015 wide. Rectum thin hyaline tube, ,anus slightly eleva.ted: Ta..ilshort, blunt, 0.122 long.

cO.2mm

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

B

D

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, tothe angInal descrlptlOll of A .. globiceps given by Yamaguti (1935);. even though weclid not succeed in obtaining the type specimens for study, it is clear that our nematodesbelong to this species. Some slight differenoes concern only measurements that areappa.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 whichis, in our opinion, an important specific taxonomio feature in Anguillicola. spp. Un­f01·tUhately; we were also unble to establish tbe exact number of these teeth from headend sections because of the limited number of horrowed iSIlecimeus available; in latera.lview it was possible to determine that the buccal capsule of the specimens of the presentma.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 variabilityin 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 morphologyfrom other congeneric species, principally in the struct1.1re of the oesopha.gus which isusually considered one of the most important features in nema.tode taxonomy. Othermarked differences are the spinose epicuticlea;nd the form of body. Therefore, wepr.opose to pla.ce A. globz'c,eps and the rema.ining AnguilUcola species into two sepa.ra.tesubgenera (see a.bove).

A. globicep8 was odg,naJly described by Y amaguti (1935) from Lake Hama.na. fromJapan (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'sswimbla.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 Hiro se et a.!. 1976). According to recentinfoz-mation from Dr. H. Hirose, in Japanthis species is muoh leBscommon in A. japonz:cathan A. C'l'aS8U8. In 1956, Wu reportedA. ylobicep8 from pond-cultured A.japonica from China, but the accompanying drawingssuggest that besides A .. globioepa, A. cra.S8'U8 might also ha.v1e been included in the materialexamined. A. globiceps was also reported by W u (1984) from th6 ,same host species fromthe Liaoho River (China, Prov. Rel1a.n) , buv the drawing of the anterior end of uhenematode (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 bothA. gloMceps and A. craS8U8. Acco~ding to Wang and Zhao (1980), A. globiceps isa 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 theregion of the Fa.r East only, btlt its future accidental introducliion in other geogra.phicalregions 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. In our opml~n.0. re-examination of these. speoim.ena and the verificati.on of their species applU'te21a.nce is qUltenecessal'Y, because t,heir identifioation was based em Dr. il'rIoItir's presumption that A. {/lobicepswas 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.dend bulbously jnflated" spherical, neck constrICtIOn m front of nerye rmg marked; bodytapering at postedol' end to pointed tail. Buccal ca.psule small, thm.walled, its anterioredge a.t"lued with numerous minute circumoral teeth; numher' of teeth not established.Oesophagus expanded at its posterior half. Valvular app~&.tuB of oesophagus welldeveloped. NeI've ring situated somewhat below neck constrICtIOn, excretol'y pore nearjunction 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 conspicuouslylarge oval mllcellula.r rental gla.nds present, one dorsal and two subventral; additiona.lsmall ventral rectal gland also present. Tail conical, pointed.Male (1 specimen): Length of body 32.30, maximum width 0.911; length of infla.tedhead end 0.163, its width 0.218; width of body at neck constriction 0.136. Buccal capsule0.009 long a.nd 0.030 wide; maximum thickness of its wall 0.003. Length of oesophagusO.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 anteriorextremity 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 duotopening on prominent prooess 0.075 long. Size of large rectal glands 0.313-0.340 by0.190-0.299, 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 constriction0.095. 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 valves0.054. Length ratio of oesopha.gus and body J.: 30.3. Distance of nerve ring and excretorypore from anterior extremity 0.272 andO.789} respectively. Vulvaelevated,prominent~situated in posterior part of body, 6.96 from posterior extremity, this repI'esentingapproximately 1/4 of body length. Uterus 'still without eggs. Anterior ova.:ry reachinganteriorly almost mid-length of oesophagus. Size of large rectal glands 0.245-0.272 by0.231-0.245, of smeJl one 0.068 XO~O54:. Posterior end of body at l'ect9J. regiongradually 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 thelong-finned 001, Anguilla reinkardtii, from PI'ospect Reservoir Ilear Sydney, New South.WaJes, Australia. Aocording to the original desoription, the gravid fem.ales of thisspecies a.re 60-70 mm long &nd 1.5 mm wide, whereas the body of young fema.les is25-30 mm long and 0.5 mm wide; the m.ale is 40 rom. long ,and 1 mm wide. This cor­responds, 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 andnarrow body. Although Johnston and Mawson (1940) reported only four pairs ofoa.udal papillae in the male of this species, we found six pai:r:s; it is possible that the abovementioned· authors overlooked the adanal and first preBdlal pairs of papillae, that areless distinct than others. .

130

F

c

D

E

EE

r>lci

c, :

: :

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

131

A. australiensi8 has also been reported from Anguilla dieffenbachii and A. austral£Sfrom :Xew Zealland (Bruns don 1956, Rid 1973, Hine 1978, Boustead 1982, Blai r1984) 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 andillustrated a.nd we could not obtain the specimens relating to these records, we considerthem 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): Bodyda.rkly coloured, fusiform,ra.ther plump, ta.pering to both ends. Epicllticle almost

O.OSmm

F

G

E]~.:.'.E .."'"'; .' . -;'

0,

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

132

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

E

c

~]O.OSmm

F

EE."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 pairsof caudal papillae).

133

present being frequently overh.pped by large ones and, accordingly, indistinct. T8Jloonica.l, short. . .Male (21 specimens): Length of body 6.54-~I.76, m~xllnum ~dth 0.55.8-1.22. Bucca.lcapsul9 0.021-0.027 long and 0.048-0.63 wIdej maXImum thIckness of Its wan 0.003 to0.006. Length of oesophagus 0.721-0.843, its maximum width 0.190-..0.218; lengthof 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.nd0.694-0.924, respectively. Seminal v68i.c1e well dev.eloped, variable in size. Commoncloacal duct opening on prominent process 0.048-0.090 long. Size of large rectal glands0.122:-0.394x0.082-0.204, of small one 0.042-0.109xO.027-Q.068. Mostly abcpa,irs of oa.udal pa,pilla.e present: 2-3 preansJs, 1 adanal, a.nd 2-3 postana-Is. In onemale 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 thicknessof 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 excretorypore 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 posteriorextremity, this repl'esenNng a.pproximately 1/4-1/10 of body length. Ovarian tubesd'eaching anteriorly to some dist,a:nce below oesophagus end level and posteriady toregion in front of rflctal gla.nds. Uterus occupying most space of body, containingIl,umerous eggs, developing embry80, Bald fully formed, sheathed larva.e 0.244--0.258long 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 Ku waha.ra et aI. (1974) from pond­oultured 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 in­adequate and inaccurate in some respects and this was the ma,n reason for doubts about.the species iden.ti:6.cat~on of conapeoifio Anguillioola nematodes reco:rded later from eelsin Europe. Only recently, Ta,l'&siohew,ski c,t, at (1987) carried out a detailed comparisonof 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; theyoonoluded. 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,veobtained a.dd~tio.nalnumerous 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 trheoonsiderable biometrical va.ria.bility of A. crassu8 the size of the buccal capsule remainsmuoh 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 I1pecifiofeatures of .Anguilli·cola; the present study shows th80tin addition the position of th6 buocalloapsule in relation to the nematode's anterior extremity and the number of circumoralteeth may be ta~onQmicallyimport,ant. The number of circumoral teeth in A. eras.SU8wa.s not established by Kuwaha.ra at al. (1974:) in the oI'iginaJdescription, but seems

134

........W~

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

it !? ~~ ~!j? ~~ I' !?~II

_.~--

Length of body' 34.94 ' 42.94- 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

Length ratio of oesophagus andbody 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 of vulva. frompos~el'iorend - ? ........ 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

._"-.-.--

to bo rather va,riable in this species. Wang and Zhao (1980) report-e,a 25-26 big teethin 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 Taraschewskiet 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 byKuwaha.ra. at aI. (1974), but this species was found in Japan a.s ea.rly as in 1969 byEgusa et al., who ha.d mistaken it for A. globicep8 (see Hirose et al. 1976). Lat,er, thisspecies was also recocded from Ja.pan by Hirose eli .Ill. (1976) and Egusa. (1979); thelatter 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 culturedeels being several tens of per cent ilond ooca.sionally neatly 100% inA. anguilla and10-40 % in A, jarponica). This species wa.s also present in the Afi(Ju~1lu'ola materialfrom 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¥ (Ne1] 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 (riverbasins of the Elbe, Weser and Rhine) and in the ,vater bodies of Berlin. It occurs alsoin Holland (Van Banning at at 19'85), Belgium (Belpaire et al. 1987, De Charle­royet 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 (personalcommunioation 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-westernand 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 constrictionjust in front of nerve ring. BuoGal CRpsule smaIl,feebly sclerotized, with minutecircumoral teeth; exa.ct number of teeth not established (32 in Ita.lian specimens accord­ingto Paggi et aI.1982). Oesophagus expa.nded at its posterior h&1f. Valvular appa.ratusof oesophagus well developed. Ne1'Ve ring situa.ted immed~ately below neck constriction,excretory pore near junotion of oesophagus and intestine. Intestine da.rk, almost straight,.broad. Three conspic'llously large oval unicellular reotal glands present; additiona.l sma.llrectal 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 end0.111-0.132(0.120), its width 0.087--0.111 (0.093); width of body a.t nook constriction0.078-0.099 (0.078). Buccal capsule 0.006 (0.006) long and 0.021 (0.021) wide, maximumthickness of its wall 0.002 (0.002). Length of oesophagus 0.476-0.530 (0..530), itsmaximum width 0.095-0.120 (0.109). Length rat~o of oesopha.gus and body' 1: 10.4 to16,2 (1: 16.2). Distance of nerve ring and exoretory pore :£tom anterior extremit,y0.15,9-0.165 (0.15H) and 0.462--0.585 (0.585), respecti,vely. Common oloacal ductopening on prominent proces~ 0.087-0.0916 (0.087) long. Size oi large rectal glands0.272--0.354 X 0,136-0.231 (0.272-0.340 X 0.136), that of small one 0.033-0.039 by0.030-0.033 (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 to13.63 (13.63), illtmmum Width 0.653-1.18 (1.18); length of sligthly enlarged hea.d end0.123-0.135 (0.132), its Width 0.090-0.102 (0.102); width of body a.t n.eck oOll..'ltI'ictionO.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

cB

F

~I

E

A

Fig 5 A.nguillicola. 'I1ovaezelamdiae sp.n. from A. autitraUs from New Zeabmd. A, B - heia.d e~d.ofmaie ~nd gI'a.vid female; O. D -,- buccal ca,psule ofmale and young fema.le; E --:- vulva; F - pas ,enorend 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. Vulvaelevat.ed, prominent, situated in pos.terior part of body, 1.84-2.08 (2.08) from postedoirextI'emity~ this )'epreoonting approxima.tely 1/4--1/6 (1/6) of body length. Ovarialtubes 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 onlynear vulva. of two larger females (body length 9'.6 and 13.6). Size of large rectal glands0.204-0.275 )<0.136-0.245 (0.272 x 0.217-0.245), small rectal gland indistjnct. Ta.ilconica.l, 0.186-0.225 (0.186) long, pointed.

A

!!d

c

,~

H

F G

Fig.6. AnlJ'Uillicola novaezela71dia,e Bp. n, - specimens from A. anguilla from Ita.Iy. A _ bea.d endof 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

I Specimens from New Zealand I European specimens(own data) (own data)

_ . M I w I M I w I

~

1:.<3co

Length of body.

Max. width of bony

Length of buccal capsule

Width of buccal capsule

Length of oesophagus

Max. width of oesophagus

Length ratio of oesophagu.Cland body

No. of caudal papillae in male

Distance ofvu1vtl. from polilterior end

Length of tail

Hosts

Oocurrence

! 5.54 -8.5'1 I 1.11 -Ia.63,

0.476-0.680 0.653- 1.183

0.006 0.006- a.OOg

0.021 0.018- 0.021

0.476-0.530 0.544- 0.625

·0.095-0.120 0.130- 0.177

1 : 10.4-16.2 1 : 13.2-21.&

6 ,,~

- 1.84: - 2.08

0.156-0.195 I 0.186- 0.225

A. C&mtrali8

New Zealand

11.63-14.92

1.41- l.oG

0.007

0.024

O.ti39-0.6lHi

0.136-0.163

I : 18.2-22.S

{)

O.HlO

A. anguilla

Italy

24.82 -28.70

2.00- 3.00

0.007- 0.009

0.024--- 0.027

0.6153- 0.843

O.204--0.24G

1 : 37.0--38.0

4.42 -- 4.7G

0.204- 0.272

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'

Localiza tion: 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 reportedby 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 of Parasitology.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 theorigin of type specimens.Comments: -A. novaezelandiae sp. n. is most similar to A. a'UStmlz"enais Johnstonet Mawson, 1940. However, it differs from the latter speoies mainly in the :shape of thehead end which is bulbously infla.ted, almost spherioal, and followed by a marked neckconstriction in A. australlenaisJ whereas it ~s only slightly expanded in .A.. novaezelandiaesp. n.; the anterior ova.ry in A. australiensis females extends anteriorly to about themid-length of the oesophagus, while it does hot reach the end of oesophagus in A. novae­zelaniUa.e sp. n. Both the sp'eoiea differ also in the size and form of the body. While thebody 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 longand 0..72 mm wide) was still non-gravid, without eggs in hs uterus. The shape of theposterior 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. anIJuillafrom 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 considerthem conspecific with A. novaezelandiae sp. n. It is probable that this .new species wasintroduced in Ualyalong with the transfers o.f live eels Anguilla australis from Ne wZealand; 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, anotherAng1.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 parasiteswere provided, it may well be that they belonged in the fact to the species A. novae­zelandiae sp. n. and that A. aust'iah'ensis is restricted in distribution to the territoryof 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'sintest,ine), i~orm, l'atherplump, tapering to both ends; a.n.1ierior end of body bottle­shaped, posteriOr one na..rrowed, conica.l. Epicuticle finely Wl'inkled, with networkstruo­tore" Mpinose, forming irregular fine transparent coating. Cuticle of anter'io),' andposte~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&onsi1!dis~inct on female po~teriOI~ end. Head end rounded, bain~ sepaI'ated by slight cotl,Btrio­tlOn 1~ front of nerve rmg level; bo~y of son:e specimens somewha.t infl.a:ted in IJegion ofpo~terlOr half o£oesophagus. EntIre antel'lOr nal'1'owed part of body appl'oximatelytWIoe &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 ofgravid female and male; C - anterior end of female; D - bucoal capsule of male; :m - head end offemale, apical view; F - buccal oapsule of female, apical view; G - female, general view; H, I ­larva and egg from uterus; J - vulva.

141

rI

l/II

Ii

A

EE..,.d

E

Fig. 8. Anguillicolapapemai Bp. n. fro.OlA. m08sambica from Sonth Africa. A - posterior end of male ~

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 buccalca.psule being 0.027-0.030 from anterior extremity; mouth opeID:ng circular. Buccalcapsule well sclerotized, trapezhull-shapedin lateral view, its a.nterior rim beating onetow ~f 26 circu~o~al teeth: Oesophagus strongly muscular, expanded at its posteriorhalf; Its lumen dIstlllctly trIangular. Valvular apparatus of oesophagus well developed.Nerve ring situated immediately below neck constriction, excretory pore near junctionof oesophagus and intestine. Intestine da.rk, almoslJ stra.ight, broad, being narrowed ata.nterior and posterior ends only. Three conspicuously large oval unicellular l.'ectalglands 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 entirenarrowed anterior part of body (1.22); width of body at head constriction (0.060), atend of naITowed anterior body part (0.272). Bucca.! capsule 0.011-0.012(0.011) longand 0.030 (0.030) wide; maximum thickness of its wall 0.003-0.004 (0.004). Lengthof oesophagus 0.530-0.598 (0.530), its maximum width 0.150--0.163 (0.163). Lengthratio of oesophagus and body (1: 24.5). Distance of nerve ring and excretory pore froUlanterior extremity (0.180) aud (0.667), respe~tively. Testis reaching anteriorly to somedistance 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 oflarge rectal glands (0.105-0.120 X 0.054-0.057), that of small one (0.039 X 0.030). Sixpairs 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 gravidfemales 15.78-28.29 (28.29), maxiullun width 1.64--2.18 (1.64); length of head end0.081-0.099 (O.099), of entire narrowed anterior pari; of body 1.43-2.07 (1.64); widthof body at head constriction 0.060-0.068 (0.063), at end of nal'rowed anterior body part0.408-0.571 (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), its maximum 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 extremity0.147-0.171 (0.147) and 0.692-0.695 (0.695), respectively. Vulva elevated, prominent,situated in postetior part of body, 3.20-3.74 from posterior extremity, this repr"esentingapproxbnauely 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 reachingto narrowed part of body. Uterus occupying most space of body, containing numerouseggs (size 0.081-·0~105X 0.075-0.084), developing embrya, andfully formed, sheathedlarvae O.189~.255 long and 0.018-0.021 wide. Uterus extending anteriorly to endOt narrowed anterior. body part and posteriorly reaohing almost level of rectal glands.Rectum thin hyaline tube, anus not elevated. Size of large rectal glands 0.120-·0.180 ~y0.063-0.090 (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 oftail 0.180-0.204 (0.180). .

Type hoatl Anguillamo8sambica Peters.Lo caliza Mon: 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 maletrag. + 2 ~~) in Na~ionalAnimal Pa.rasi~e Laborat.ory, Maryland, USA (Ca~. No. 80007).

143

Etymology: This Bpecies is named in honour of Dr. Ilan Paperna, the HebrewUniversity of Je:rusalem, who contributed greatly to the knowledge of fish parasites.

Comments: - The unique morphological features of .A.. papernai sp. n. by which thisspecies 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 buocalcapsule deeply inside the head end. This species is the first Anguillicola member knownfrom Africa.

KEY TO THE SPECIES OF ANGUILLICOLA:

1 Epicuticle finely spinose. Oesophagus conspicuously infla1:€d at anterior end, itsposterior 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 circumoralteeth. 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; posteriorpart of oesophagus distinctly expanded. Buccal capsule small or medium sized, lessthan 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 distinctneck 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 teetihvery small. Hea.d end bulbously inflated or at least slightly expanded with distinc'tneck constriction. Body fusiform or filiform. Austl'alia, New Zeala.n.d, Africa. orEurope ; . . . . . . . . . . . . . . . . 3

3 Body fusiform, anterior and sometimes also posterior ends of body with markedpa.pila-like cuticular excrescenoes. Head end slightly expanded. Buccal capsulelocated deeply inside head end, with 26 minute teeth. Mrica .... A. papernai sp. n.

-. Body filifol'm of fusiform, c'Uti<mlar excrescences on anterior and posterior ends ofbody lacking. Head end bulbously infla.ted or slightly expanded. Buccal capsulelocated near nema.tode a..utel'iOL- extremity. In eels of other geographical regions. . 4

4 Head end bulbously inflated. Anterior ovary in females extending to about mid-length of oesophagus. Body filiform, 30-40 mm long in males and 60-70 mm ingravid females. South Austl'alia . . . . . . . . . . . . . . A. australiensis

- Head end slightly expanded. Antel'ior OVal'y in females not extending to end ofoesophagus. Body fusiform, only 6-15 mm in males and 10-31 mm in gravid.females. New Zealand and Europe (Italy) . . . . . . . A. novaezelandiae sp. n

ACknowledgements. Our thanks are due to the following persons who kindly provided us with theAnguillicola 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, Departmentof Animal Sciences, Hebrew University of Jerusalem, Israel; Prof. F. Q. Wang, Parasitology :ResearchL.a~oratory,Fujian. Normal Universi~y,Chh~a; Prof. H. S. Wu, Institu.te of Hydrobiology, AcademiaSllllca, Wu~an, Chma; and Dr.. S. Kamegtu, Maguro Parasitological MUEl6Um, Tokyo, Japan. Saxnenec~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, Universityof 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~Tr P E tt C 0 IT II CA H 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 pOAIlAnguillicola. 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 PUHbUIerrpIIBoglIJIIIC-h 1I3 I1TaJIlHr nop; Ha3BaHfreM A. australiensis) II A. papernai sp. n. OT Anguillamossamb iea li3 IOamoii: AepPliHH. B rrUCTomu;eo BpeMfI H POAY Anguillicola rrupHaJVW)I<aT IUlTbCJI8P,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 An­guillicola ~ 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 THIORConpCACJIHTeJIbH1UI Tu6mlIl;a AmI HA8HTucIlIlH:aIJ;InI BIll\OB :3Toro POj\H.

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