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