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Species of Discocelis (Platyhelminthes: Polycladida) from Queensland, with description of a new species PDF

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Preview Species of Discocelis (Platyhelminthes: Polycladida) from Queensland, with description of a new species

SPECIES OFDISCOCELIS(PLATYHELMINTIIES; POLYCt^ADlDA)FROM QUEENSLAND, WITHQfiSCRIPTlON QFANEWSPECIES Beveridgt1.20(H)0630 speciesol7^/;vcoL-L-7/>(Plat}helriiinthes: \cludiJajfrorTiQueens- laad»withdescriptionofanewspecies.MemoirsojtheQueenslandMuseum45(2);205-213. ^sbane.ISSN007^8835. Jwo :^pccies ofDiscocciisare describedfromint&iS$diwatersfromnorthernQtieenslanii. Thefirstspeciesischaraderisedbythemarginaleyesextendingtotheposteriorregionofthe bod^fthepresenceofaseminalvesicle,cerebraleyesdistributedmoreorlessinanteriorand mMQtWfi^miamate-amnia^withcOmRlexlobesjtrtlorst^'venttdviews^Ttiui^f^!^ difBsfStwnyfr^Hidll«^g6rt6r6ahd named O.pGtrvilmxcutata ht!AcTh^seccmdhas marginal e>'e5 extending tothe [eve! ofthe cerebral organ, cerebral eyesarranged moreor lessintwogroups. lacksaseminal vesicleandhasprosuiioids inihe^wallofthemaleantrum aswellaji iniliepenispapillaitself.The speciesisclosely reIaitd /^M'V/i/Kato, 193Silt isprobabi) distinct l->iii isnotnamedowingtothepoordescription ui / ' p>''\,-''la.Therecords prcscnled suggestthat SL'Vcral species oJ DiscoceUsarc present in Austi'alian coastal waters and that the distribuiiun of prosiaioids seen in dorso-\entral views ofthe male antrum O provides usetui characters for distinguishing ^ieC)6^Withlt^ gl^Sw <Pc)/)'C/«)k/^ DLsnKelis, ;;cvt spt'cics. taxonomy. I. Beverhige, DepartmenfofVetenmryScience, UmversityoJMelbourne. Parb/Ule3052, MeihauPm*Australia;24AprU Thepolyclad^amlyDcseocelidaeLaiiilatv;I9D3 esttended, ^(^]^pe!rM<a&Tt^dlyplacedpiEta Uacosmopolitan family ofcssentiaily iniertidal blockoffrozenfixative,either4%foBivaldebyd^ polyciads characterised by the presence of in sea-water or lormaldehyde-calcium acetafc- iiKiruinal eyesand.smallsecretoryorgans,lemied piopylenc ghcol- propylenephenoxetoi. Following proslatoid.s,associatedwith tlicmalereproductive fixation, worms were dehydrated in a graded ^StefO (fiiubel, 19S3). The family is curretitly series of ethanols, cleared in methyl salicylate represented by a single Australian speeieSj aad mounted m Canada balsam The median Discocelis aiis&ah's Uytnsin. 1959, found'Uttdei* i^osteriorsecfionfi^ofrndividtial polyciads ^Vere roeka in die intertidal regit-n close to Sydney removed using a scalpel blade, embedded in (Hyman, 1959)andfrom Wesi I.. SouthAustralia paraffinand serial longuudinal sections, cul al a CPrudhoe, 1982). Faubel (l')S:v) iransferred /). thickness of 7|,im, were stained with Gill's j^mralis^ to thf related Thalamoplatia haeinatoxylin and cosin. Drawings were made Vaiidlaw;. l$Q4,/(t&lingidslied from Distocetfs usingadi^'ftgtubeattachedtotoOlyttipusBM Elues^bei^183$-'ftypossessingseparatemaleand microscope. Measurements are presented in ferhate gon6p&}«fi» By contrast, Prudhoe (i985j millimetres asthe range followed by the meanin considered th^ thiilarfic^lart&ViTdnso^ OfAy parentheses; sub-genericrank. Ailspec^i^ii^caUQCtedhavebeepdepositedin Tber piFesi^noe- 6fity a single Atistraliari fheQuefenslaridMuseumXQM). rcpresentalive«ftfi]BfinuIjrisprobably theresult Type specunens ot D. a/<Mruli\ from Nc'^n oflackofcollectingratherthan thefamily being South Wales tAu&iraJianMuseum W3685) were pv»or]\ representedinAustralianinlertitj^waters. cotnpdtsdVfiSxthenewmaterial. Thivpaperr^)Qrt$the{jfcssenceoftw^addit^Qnal ipQws^J>lsso0p To^ville,Queens^ FOLYCLADIDALana 1884 land,oneofwBichiscieaily specbs. ACOTYLEALang, 1884 DISCOCELIDAE Laidlaw. 190> METHODS ^ Oiscocetisparvimaculftta sp. luov. Polyciads were coilegt^f) low-tide ironi. under rocks on exposed imld-'flats. Kixation followed the technique ofNewman &. Cannon MATERIAL.HOLOTNPH: RowcsBa).To\mis\iile.(.)ld (I995J inwhich polyciads were placed on filter (19''16'S, 146'^49^E),17.vi.l997,coll,I.Bexpridge.whole paper in a dish of searw?^^ and wljen fiilly mount, unstattied (QM G217321); 2 colour slides. 206 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 2. Discocelis parvimaculata sp. nov., cerebral organ,tentacularandcerebraleyesshowingvariation between individual specimens(see Fig. 3). Scalebar = 0.1mm. ocellidiminishesposteriorly; inmostspecimens, includingholotype,eyesreachlevelofgonopore; in some specimens, eyes encircle body; cerebral eyes arranged in elongate groups, on either side of mid-line, 41-65 ocelli anterior to cerebral organ, 5-20 posterior to cerebral organ, anterior FIG. 1. Discocelis parvimaculata sp. nov., entire and posterior groups usually but not invariably polyclad, dorsal view, showingpatternofpigmented maculae and extent ofmarginal eyes (ME arrows). separated (Figs 2,3); tentacular eyes with 25-40 Scaie bar= 1mm. ocelli per cluster; rutTled pharynx in mid-body, with 10-12 lateral folds, 11 inholotype;mouthat PARATYPES: 5 entire specimens and fragments of 1 posterior end ofpharynx, 4.9 from posterior end specimen, whole mounts; 1 set ofsections stained with in holotype, 7.0-8.1(7.6) in paratypes; single haematoxylin and eosin, Rowe's Bay, Townsvilie, Qld, gonopore 3.13 from posterior end in holotype, coll.I.Beveridge, l.vii.l994,29.vi.1995(QMG217322-7, 5.5-6.8(6.0) in paratypes; antrum masculinum serialsectionsG217328). volimiinous, folded in both dorso-ventral views DESCRIPTION. Large,ovalpolyclads;holotype and sagittal sections; in ventralviews (Figs 4,5), non-gravid specimen 13 long, 10 wide; gravid antrum with prominent anterior lobe containing specimens, 18-21(19) long, 7-12{10)wide; dorsal penis papilla and two lateral lobes each partially surface fawn, darker in centre, covered with subdivided; in sagittal section (Fig. 6), several numeroussmall browncircularareasofpigment, muscular lobes descend from dorsal surface of larger brown patches in central regions, becom- antrum; antrum with numerous pyriform ing smaller towards periphery (Fig.l); ventral prostatoids opening into lumen; in ventral view, surface pale grey; nuchal tentacles absent; prostatoids arranged in subcircular cluster on cerebral organ0.41 x 0.55 inholotype, 0.39-0.48 penis papilla and on posterolateral margins; in (0.43)X0.46-0.56(0.51) in paratypes, 2.58 from sagittal sections, prostatoids present on all anteriorextremityinholotype,2.94-4.60(3.77 )in pendantprocesses;noprostatoidspresentin wall paratypes; marginal eyes 3-4 deep, extend to ofantrum; prostatoids oftwo histological types; posterior quarter of body, number of rows of mostwith faintlyeosinophic content; prostatoids . DISCOCELIS SPECIES FROM QUEENSLAND 207 9°% FIG. 3. Discoce/is parvimaculata sp. nov., cerebral organ,tentacularandcerebraleyesshowingvariation between individual specimens. CE = cerebral eyes; CO=cerebralorgan;TE=tentaculareyes. Scalebar =0.1mm. FIG. 4. Discoce/isparvimaculata sp. nov., gonopore FIG. 5. Discocelis parvimaculata sp. nov., ventral andgenitalcomplex,ventralview.G=gonopore;L= aspect showing cerebral organ, eyes, pharynx and Lang'svesicle;P^prostatoids; U^uterineduct;VD genitalcomplex.C=cementglands;G^gonopore;L =vas deferens. Scale bar=0.1mm. = Lang's vesicle; M ^ mouth; PH = pharynx; U = uterineduct;VD=vasdeferens.Scalebar=0.1mm 208 MEMOIRS OF THE QUEENSLAND MUSEUM FIG. 6. Discoce/isparvimaculafa sp. nov., median sagittal section showing mouth, gonopore and histological details ofgenital ducts. C =cementglands; IN = intestine; L = Lang's vesicle; M ^ mouth; PI =eosinophilic prostatoids, P2 =basophilic prostatoids; PH=pharynx; SV=seminal vesicle. Scalebar=0.1mm. at anterior extemity ofantrum and on ventral or 0.42-0.44, 2.5-4.6 from anterior extremity; anterior surfaces ofpendant processes of penis marginal eyes in rows 3-4 deep, extend around papilla with basophilic content; penis papilla anterior quarter ofbody, reach level ofcerebral fleshy, prominent, in anterior part of antnun; organ; cerebral eyesarrangedinelongategroups, ejacLilatory ductsimple,straight;prostateabsent; either side ofmid-line, 31-42 ocelli anterior to ejaculatory duct leads to pyriform seminal cerebral organ, more or less separate from 4-7 vesicle with thin but highly eosinophilic wall, posterior to cerebral organ (Figs 8,9); tentacular passes venlrally, divides; walls of spermiducal eyes with 18-30 ocelli per cluster; ruffled bulbs highly muscular; vasa deferentia pharynx in mid-body, with 10 lateral folds; thin-walled, pass anterolaterally from male mouth at posterior end of pharynx, 5.2 from complex,to levelofmouth,thendivide;posterior posteriorend; singlegonopore3.4 fromposterior branches coil posteromedially, uniting posterior end; antrimimasculinumvoluminous;prominent to Lang's vesicle. No separate female gonopore; anterior penis papilla, circular in ventral view vagina opens into male antrum immediately (Figs 10,11), with numerous prostatoids; wall of posteriortocommongonopore;vaginawiththick antrum encircling penis papilla bearing single muscular walls, ciliated lining, cur\^es anteriorly rowofprostatoids; antrumwith2 laterallydirected to short, horizontal region; uterine canals empty branches on each side, immediately anterior to into vagina immediately anterior to termination gonopore; anterior pair oflateral branches with of vagina into prominently Y-shaped Lang's row of prostatoids along posterior margin; in vesicle; uterinecanalsextendanteriorlyoneither sagittal section (Fig. 12), large muscular penis side of pharynx; cement glands prominent in papilla descends from dorsal surface ofantrum, horizontal region of vagina, extend posteriorly with numerouspyriform prostatoids; prostatoids andlaterallyintoparenchyma,brancheddistally. present in wall ofantrum, restricted to anterior ventral region; prostatoids with faintly Discocelis sp. eosinophic content; ejaculatory duct simple, (Figs 7-12) straight; prostate absent; seminal vesicle absent; ejaculatory duct divides into vasa del'erentia MATERIAL. Two specimens, Rowe's Bay Townsville, whichpassanterolaterally frommalecomplex,to Qsledc,ti1o.nvsii.s1t9a9i4n,edcolwli.tI.hBehvaeermiadtgoe,xywlhionleamnodunetosainnds(erQiaMl level of pharynx, then re-divide; posterior G217329-30, serial sectionsG217331). branches coil posteromedially, uniting posterior to Lang's vesicle. No separate female gonopore; DESCRIPTION. Oval polyclads; gravid speci- vagina opens into male antrum immediately mens 12-16 long, 5-8 wide; dorsal surface fawn, posterior to common gonopore; antrum anterior darker in centre, covered with numerous brown to vaginal opening, prominent, muscular with circular areas of pigment, larger patches in thicker epithelium; vagina with thick muscular central regions, becoming smaller towards per- walls, ciliated lining, ciu^es anteriorly; uterine iphery (Fig. 7); ventral surface pale grey; nuchal canals empty into vagina anterior to prominent tentacles absent; cerebral organ 0.33-0.45 X dorsal loop; vagina passes ventrally to enter DISCOCELISSPECIES FROM QUEENSLAND 209 FIG. 8. Discocelis sp., cerebral organ, tentacular and cerebral eyes showing variation between individual specimens (see Fig. 9). CE ^ cerebral eyes; CO = cerebralorgan;TE=tentaculareyes.Scalebar=0.1mm. Adenoplana as an ejaculatory duct lined with a glandular epithelium. Whatever the precise definition of the structures involved may be, Adenoplana differs fi"om the species described hereinpossessingdistinctlyseparategonopores. FIG. 7. Discocelis sp., entire polyclad, dorsal view, The remaining genera, Discocelis and Tha- showingpatternofpigmentedmaculaeand extentof marginal eyes (ME arrows). Scale bai'= 1mm. lamoplana, are distinguishable on the basis of gonopores, with the former possessing a single gonopore and two gonopores in the latter. Y-shaped Lang's vesicle; uterine canals extend However, D. australis, which Faubel (1983) anteriorly on either side of pharynx; cement assigned to Thalamoplana, possesses a single glands prominent, extend posteriorly and gonopore, a feature which was confirmed by laterally into parenchyma. examination of the type specimens, while D. DISCUSSION insularis Hyman, 1955 has the male and female systems opening at essentially the same point, Both species described above belong to the which as Prudhoe (1985) has observed, is familyDiscocelidae since they possess marginal intermediatebetweentheconditionpresentinthe eyes and prostatoids opening into the male type species ofthe two genera. For the present, antrum (Faubel, 1983; Prudhoe, 1985). Generic Faubel's (1983) separation of Discocelis from distinctions within the family are not well Thalamoplana is accepted but australis is defined, and although both Faubel (1983) and considered, following Prudhoe (1985), to be a Prudhoe(1985)acceptthevalidity ofDiscocelis^ Adenoplana Stummer-Traunfels, 1933 and Coronadeua Hyman, 1940, their definitions of these genera differ. In addition, Thalamoplana Laidlaw, 1904, accepted by Marcus & Marcus (1966),deBeauchamp(1961)andFaubel(1983), was not accepted as a valid genus by Prudhoe (1985). Both species described here differ from Coronadena in lacking the 7-11 large prostatic organs arrangedradially aroundthemaleantrum in addition to the more numerous small prostatoids. Adenoplana was characterised by Stummer-Traumfels (1933) as having an FIG. 9. Discocelis sp., cerebral organ, tentacular and interpolated prostatic organ. Faubel (1983) by cerebral eyes showing variation between individual contrast interpreted the prostatic organ of specimens. Scalebar=0.1mm. 210 MEMOIRS OF THE QUEENSLAND MUSEUM FIvGe.ntr1a0l.Dviisecwo.ceGli=sgsop.nogpoonroep;orLe~anLdangge'nistavlesciocmlep;lePx=, prostatoids;VD=vasdeferens. Scalebar=0.1mm. member of Discocelis. Both species described abovearetherefore assignedtoDiscoceliswhich consists ofD. australis, D. tigrina (Blanchard, & 1847), D.fiilva Kato, 1944, D. japonica Yeri Kaburaki, 1918 and D. pusilla Kato, 1938. The type species, D. lichenoides (Mertens, 1832), is considered unrecognisable (Hyman, 1959; Faubel, 1983; Prudhoe, 1985) andwas treatedas a species inquirendaby Faubel (1983). Within Discocelis, the first species described FIoGr.ga1n1,.Deiysecso,ceplihsarspy.n,xvenatnrdalgaesnpietcatlshcoowmipnlgexc.ereCbra=l above is immediately distinguishable from all cementglands; G=gonopore;L^Lang'svesicle; M congeners on the basis ofthe extent ofthe eyes, = mouth; PH =pharynx; U = uterineduct; VD =vas which in other species extend only as far as the deferens. Scale bar= 0.1mm. region of the cerebral organ but in this species extend to, or almost to, the posterior end ofthe possessing a seminal vesicle, though this was body. The marginal eyes also extend to the described as amuscular organ in D. australis by posterior part of the body in Adenoplana and Hyman(1959)buthasathin, highlyeosinophilic Coronadena. The species described here differs wall in the specimens described above. The from all congeners except D. australis in specimens described here differ from D. tigrina DISCOCELISSPECIES FROM QUEENSLAND 211 FK-I. 12. Disci'cclis sp., median sagittal section allowing mouth, uonopoie and histological details ot genital duels.C =cementglands; IN = intestine; L=Lang's vesicle; P=prosiatoids; VD^ vasdeferens. Scalebar^ andD. mishiitisinb^VingamaleAntrum wtiicti speciesiTtwhkirtfiepattpms^onifiedor^l^tirfece foniis fivedistinctloheslndoi*soventralviews. In ha\e been adequatofydtescribed. boih ofthe oTlier species the antrum is rounded, The second species described above is based on plale l."^. fiu. 1 of Long ( 18H4) for /.). distinguishable from D, australis and D. iign'fuiandobscr\ationsofthetypespeciincnbin purvim&culata in lackingaseminal vesicle and the case ofD.maus^raSfff. Themorphology ofthe ftom Ifietetter-speEiBS in li^ng the marguial molelantoa c(otKbveii!tral view has not ey^s restricted to the anterior region ofthe bodyx describra for the remaititng species'. The It differs from /-), (igrioa in ha\ ing the mouth al separation ofthe cerchra! eyes into Iavo elusters the posteriorend ofthepharynx ratherthan inthe sepai'ates the species described here from D. middle and in having Ll>e cerebral eyes divided tigrino^ D. unsfralis and D fulva and the colour intoanWrf*?ratidp09t?riOrgroups,inaddition,the pauern of the dprsal surface, with numerous arrangement of the prostatoids in ventral view bf&wn'cirtular aieas separates^-fepecfes from (FigJO) dilTcrS from thai found in D. ii^rinti in D.Jiilva which lacks a disiinctive pattem (Kato, wliich thev are arranged in a U-.shaped cluster 1944). Tltc TWO typc'^ of prostatoids. one with around the ainenorhalfofthepeniS-papilla, vvjlli eosinophilic content and tlie ulher with bn:^o~ two lateral rows extending pOSterfoWy (iMg^ philie content may also distinguish this species ISX4, pi, I V n,y- i )- frOJUallcongSKtetSyaftbW&llKat (194R4, fig. 2) 1lie _specics is therefore most closely related to iUuste&t^ed types ^prbst^ids^u) fyWch D fulva, D. iitponicif and D- />nsiIl(L all from butdidnotdescribethediffferertccsshowninthe .lapan. The specimens are distinguishable trom illustration.Themorphologicaldifferencesnoted D Jn/va stA^thisSpecies-hds rtO dorsal coldUf therefore indicate that tl)e described specimenAs pattern,hasrruitt^rouscerebral^esacrangedina represent a new species for which the natne parvimacuicfia is proposed based on the small si2eot^tdbr^Liiid^olar-i^p^dWithDtber FIG, T?.SehBI1iaticreprescntalianofgenilalatriUffiof Dtscocelist^ina^redrawn from Lang(1S84). 212 MEMOIRS OF THE QUEENSLAND MUSEUM However, Kato's(1938)specimensofD.pusilla, were evidently immature as he describes the prostatoids as rudimentary and Lang's vesicle as beingrepresented merely by a mass ofnuclei. As a conseqence, the number and distribution of prostatoids may not have been reliably determined in D. pusilla. The current specimens may therefore be D. pusilla or may represent a FIG. 15. Schematicrepresentationofgenital atriumof newspecies. However, sinceonlytwospecimens Discocelisfulva, redrawn from Kato (1944). are available and since D. pusilla has been single elongate group and, according to the inadequatelydescribed,nonewnameisproposed forthem. illustrationsofthespecies,hasprostatoidsoftwo distinct sizes (Kato, 1944, fig. 2) (Fig. 15). The descriptions presented here indicate that Discocelis is represented in Australiaby several D, japouica differs in having 15-16 eyes in species rather than the single species, D. eachposteriorcerebral clusterratherthan the4-7 australis, currently known (Hyman, 1959). in the present specimens, and differs in the Whileoneofthetwoadditional speciesfoundcan anatomy ofthe antrum masculinum and distrib- unequivocallybeidentifiedasnew, limitationsin ution of prostatoids (Fig. 16). In D. japonica ^ the descriptions of existing species prevent a thereareanimrberofprojections intothe antrum definitive name being applied to the second apart from the penis papilla, while in the current species. specimensonlythe penispapillaprojects into the antrum. Inaddition, inD. /a/70A?/r(:/,aparticularly The descriptions presented above suggest that elongate projection, lying dorsal to the vagina in addition to the distribution ofmarginal eyes, bearsnumerousprostatoidsonbothsurfaces(Kato, theoccurrenceofcerebraleyesinasinglebandor 1937, fig. 2), while in the present specimens, the twogroups,andthepresenceofaseminalvesicle, region of the antrum anterior to the vaginal the distribution ofprostatoids within the antrum opening is devoid ofprojections andprostatoids. masculinum as seen in ventral views ofcleared Unfortunately, no ventral views ofthe antrum of specimens provide useful taxonomic characters. D.japonica have been published. Finally, there In D. tigrina, the prostatoids are arranged in an areprostatoidsintheventralwalloftheantrum in arc anterior to the gonopore (Lang, 1884), in D. the current species and these are lacking in D. parvimaculata, the prostatoids are arranged in a japonica. cluster in the anterior lobe of the antrum and ThespeciesdescribedhereismostsimilartoD. aulno-nngatmheedposstpeerco-ileastertahlemaprrgoisntsatwohiidlse oincctuhre pusillaincolourpattern,havingeyesrestrictedto throughout the penis papilla and are present the anterior part of the body, mouth at the along the posterior margin ofone pair oflateral posterior end ofthe pharynx and cerebral eyes diverticula within the male antrum. The type divided into anterior and posterior clusters with specimens of D. australis were examined but only one or two ocelli in the posterior clusters they are now very dark and the distribution of (Kato, 1938). Thegenitalatrium isalsosimilarin prostatoids cannot be determined. In the that there is, according to the illustration ofthe remaining species, this character has not been species (Kato, 1938, fig. 3) a large penis papilla investigated, but current observations suggest parlotjheocutgihngKianttoot(h1e9a3n8t)ruvSmtamteadscinultihneumde(sFcirgi.pt1i4o)n, that it might provide additional features for the that there were many muscular villus-like separationofspecieswithinthegenusDiscocelis. projections, as in D. japonica. Furthermore, there are no prostatoids in the posterior region of the antrum. The most obvious differences between the present specimens and D. pusilla are that there appear to be very few pros- tatoids in the antrum ofD. pusilla andthatprostatoidsdo notoccurin FIG. 16. Schematic representation of genital atrium of Discocelis the ventral wall of its antrum. japonica, redrawnfrom Kato(1944). DISCOCEUSSPECIES FROM QUEENSLAND 213 FIG, 17*SchematicrepresentationofgenitalatriumofDiscocelisaustrctlis,redrawnfixmiHyman(1959). AdKNOWLEDOEMENTS LANCi, A. 1884. Die Polycladen (Seeplanarien) des Golfes von Neapel und der angren/enden DrL.R.G Cannon ofthe Queensland Museum MfciVNahschniUe. Hine Monographic. Fauna und is thanked for comments on a draft of this Flora des Golfes von Neapel, Leipzig 11: i-xi, manuscript 1-688, LITERATURECITED MARCUS, E. & MARCUS, E. 1966. Systematische UebersichtderPolycladen.ZoologischeBeitrage de BEAUCHAMP, P. 196L Classe des Turbellaries. 12: 319-343. (Tewdb.e)llTraariit9sdQSeitZrb^ohluoir^^e41(813)1:),35I-n21G2ra(ssNffia,ssPo>nP:. NEWMAN, L.J & CANNON, L.R.G 1995. The Paris). importanceofthe fixationofcolour, pattern and FAiFBEL, A. 1983. The Polycladida, Turbcllaria, form in tropical Pseudocerotidae Proposal andestablishmentofanewsystem.Part (Platyhelm'inthes, Polycladida). Hydrobrologia I, The Acotylea. Mitteilungen aus dem 305: 141-143. haraburgischen zoologischcn Museum und PRUDHOB, S. 1982. Pohclad lurbellarians from the InstitutSO: 17-121. southerncoastsofAustralia. RecordsoftheSouth HYMAN, L. 1959. Some Australian polyclads AustralianMuseum IS: 361-384. KATO2(,T5u:Krb.Mei7l1.a9r3i7a.).PRoelc>ocrladdssofcotlhleeActuesdtrailniaInduM,uJsaepaunm, 1$&(ST,rusAtemesonoofgrtahpehBrointispholMyuclsae^utmurb(eNlaltaurriaalt History): London). JapaneseJournal ofZoolog\ 7; 211-232. 1938. Polyclads from Amakusa, Southern Japan^ STUMMER-TRAUNFELS, Rvon.1933.Polycladida JapaneseJournal ofZoology 7: 559-576, (continued).Pp. 3485-3596.InBronn,H,G (ed.) 1944. Polycladida of Japan. Journal tif the Klassen und Ordnungen des Tier-Reichs (IV). SigenkagakuKenkyusyo 1; 257-318. (Vermes)(Leipzig).

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