THE TREMATODE PARASITES OF MARINE MAMMALS
By Emmett W. PkiceParasitologist, Zoological Division, Bureau of Animal IndustryUnited States Department of Agriculture
The internal parasites of marine mammals have not been exten-sively studied, although a fairly large number of species have beendescribed. In attempting to identify the trematodes from mammalsof the orders Cetacea, Pinnipedia, and Sirenia, as represented byspecimens in the United States National Museum helminthologicalcollection, it was necessary to review the greater part of the litera-ture dealing with this group of parasitic worms. In view of thefact that there is not in existence a single comprehensive paper onthe trematodes of these mammals, and that many of the descrip-tions of species have appeared in publications having more or lesslimited circulation, the writer has undertaken to assemble descriptionsof all trematodes reported from these hosts, with the hope that sucha paper may serve a useful purpose in aiding other workers in de-termining specimens at their disposal.In addition to compiling the descriptions of species not availableto the writer, two new species, one of which represents a new genus,have been described. Specimens representing 10 of the previouslydescribed species have been studied and emendations or additionshave been made to the existing descriptions; in a few instances thespecies have been completely reclescribed.Three species, Distoinwni pallassil Poirier, D. vaUdwim von Lin-stow, and D. am/pidlacewni Buttel-Reepen, have been omitted fromthis paper despite the fact that they have been reported from ceta-ceans. These species belong in the family Hemiuridae, and since allspecies of this family are parasites of fishes, the writer feels thattheir reported occurrence in mammals may be regarded as eithererrors of some sort or cases of accidental parasitism in which fisheshave been eaten by mammals and the fish parasites found in themammal post-mortem. Buttel-Reepen (1902) has pointed out, inconnection with the reported occurrence of D. ampullaceum in acetacean, that sailors commonly designate species of mackerel as
"Delphin" or "Dolphyn (holliindisch)." It appears likely, there-No. 2936?Proceedings U. S. National Museum, Vol. 81. Art. 13118893?32 1 1
2 PROCEEDINGS OF THE NATIONAL MUSEUM vol.81
fore, that in this case these terms for fishes have been interpreted tomean a dolphin and that these fish parasites have been erroneouslylisted as parasites of cetaceans.In the compiled descriptions all measurements when less than 1 mmhave been given in microns, when more than 0.5/x the fraction hasbeen counted as 1^, and if less the fraction has been discarded.The trematodes of marine mammals which may be considered validor recognizable forms comprise 30 species belonging to nine families
:
Fasciolidae Railliet, Echinostomatidae Looss, TroglotrematidaeOdhner, Opisthorchiidae Braun, Heteroplwidae Odhner, Paramphis-tomatidae Fischoeder, Notocotylidae Liilie, Opisthotrematidae Poche,and Rhabdiopoeidae Poche. These families may be differentiatedby the following key
:
KEY TO FAMILIES OF TEEMATODE PARASITES OF MABINE MAMMALS
1. Body provided with two suckers; eggs without polar filaments 2.Body provided with only one sucker ; eggs with polar filaments 7.2. Acetabulum or ventral sucker situated at posterior end of body.Paraniphistomatidae (p. 41).Acetabulum situated in anterior half of body 3.3. Anterior end of body provided with a kidney-shaped collar armedwith one to two rows of large spines Echinostomatidae (p. 21).Anterior end of body without collar as above 4.4. Usually large flukes, either flat and leaflike, slightly flattened,or cylindrical ; cirrus pouch and cirrus present ; seminal re-ceptacle very small or absent Fasciolidae (p. 3).Medium-sized to very small flukes, usually oval and flattened
;
cirrus pouch and cirrus absent; seminal receptacle large andconspicuous 5.5. Body spindle shaped ; occurring in cysts, usually in pairs.Troglotrematidae (p. 23).Body flat, not occurring in cysts 6.6. Body covered with small, scalelike spines ; acetabulum inclosedin the genital sinus ; parasites of the intestinal tract.Heterophyidae (p. 33).Body usually without spines ; acetabulum not inclosed in thegenital sinus; parasites of the gall bladder or bile ducts.Opisthorchiidae (p. 25).7. Genital pore situated at extreme posterior end of body.Opisthotrematidae (p. 48).Genital pore near anterior end of body 8.S. "Ventral surface of body usually provided with longitudinal rowsof glands, or rugae ; vitellaria pretesticular ; without probo-scid complex in posterior part of body Notocotylidae (p. 44).Ventral surface of body without rows of glands, or rugaevitellaria posttesticular ; proboscid complex in posterior partof body Rhabdiopoeidae (p. 47).
AUT. 13 TREMATODE PARASITES OF MARINE MAMMALS PRICE OFamily FASCIOLIDAE Railliet, 1895Synonyms.?Fasciolopsidae Odhner, 192G, p. 4; CampulidaeOdliner, 1926, p. 5 ; Brachycladiidae Faust, 1929, p. 88.Family diagnosis.?Large flat forms; suckers relatively close to-gether; cuticle with or without spines. Intestinal ceca simple orwith lateral dendritic branches. Excretory bladder simple and tube-like, or profusely branched. Genital pore preacetabular ; cirruspouch present; cirrus armed or unarmed. Testes usually pro-fusely branched, but may be without branches or lobes. Ovarybranched or entire; seminal receptacle reduced or absent; Laurer'scanal present. Vitellaria profusely developed, consisting of numer-ous follicles situated along the sides of the body and becomingconfluent posteriorly. Uterus with relatively few coils; eggs large,either circular or triangular in cross section.Type genus.?Fasciola Linnaeus, 1758.
KEY TO SUBFAMILIES OF FASCIOLIDAE CONTAINING SPECIES PARASITIC IN MARINEMAMMALS
1. Body flat, leaflike; testes and ovary profusely branched; eggswithout thickening at posterior pole, circular in cross section.Fasciolinae (p. 3).Body elongated and slightly flattened, but not leaflike; testesand ovary usually unbranched ; eggs with thickening at pos-terior pole, usually triangular in cross section Campulinae (p. 5).Subfamily Fasciolinae Stiles and Hassall, 1898Subfamily diagnosis.?Fasciolidae: Body flat and leaflike. In-testinal ceca profusely branched, the branches being dendritic andmainly lateral. Excretory vesicle with lateral branches. Testesprofusely branched; cirrus pouch not extending beyond posteriormargin o,f acetabulum; cirrus unarmed. Ovary branched. Eggswithout thickening at posterior pole, circular in cross section. Para-
sites of herbivorous or omnivorous mammals.J'ype ge7ius.?Fasciola Linnaeus, 1758.
Genus FASCIOLA Linnaeus, 1758
Gene7'io diagnosis.?Fasciolinae : Body large, broad, flat, and leaf-like; anterior end conical, forming a cephalic cone, which is set offfrom the wider, flattened, leaflike portion. Cuticle armed with scale-like spines. Oral sucker subterminal ; acetabulum at base of cephaliccone. Prepharynx short ; pharynx well developed ; esophagus short
;
intestinal ceca long, extending to posterior end of body and providedwith numerous, long, dendritic lateral branches, and fewer, shorter^
4 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 81
median branches. Excretory vesicle long and slender, with numer-ous lateral branches, which anastomose to form an extensive dorsaland ventral network. Genital pore preacetabular, situated at baseof cephalic cone; cirrus pouch not extending caudally beyond pos-terior margin of acetabulum; testes profusely branched, tandem inposition, and situated in the equatorial zone. Ovary branched, pre-testicular, situated to one side of median line ; seminal receptacle ab-sent; Laurer's canal present. Vitellaria very profusely developed,extending from base of cephalic cone to posterior end of body, com-pletely filling posttesticular area, and extending both dorsal and ven-tral to intestinal ceca. Uterus coiled in the form of a rosette, sit-uated between acetabulum and ovary. Eggs large, without thicken-ing of posterior pole, circular in cross section. Parasitic in bile ducts
o,f herbivorous and omnivorous mammals.Type species.?Fasciola hepatica Linnaeus, 1758.FASCIOLA HEPATICA Linnaeus, 1758Plate 1, Figure 1Synonyms.?Planario) latiuscula Goeze, 1782, p. 169; Distonia he-paticutn (Linnaeus, 1758) Abildgaard (?) ; Fasciola humana Omoiin.,1790, p. 3053; Fasciolaria hepntica Encycl. Metropolitana, 1845, p.141; Distomum caviae Sonsino, 1890, p. 100; Gladocoelium hepaticuTn(Linnaeus, 1758) Stossich, 1892, p. 7.Description.?Fasciola: Body flat and leaflike, 13 mm long by 3.7mm wide in region of ovary ; cephalic cone 1 mm long. No cuticularspines were present on the specimen examined by the writer. Oralsucker subterminal, 387/^ long by 542/x wide ; acetabulum transverselyoval, 542|ii long by 697/* wide, situated 1.2 mm from anterior end ofbody. Digestive tract as described for F. hepatica by other writers.Testes profusely branched, occupying the entire intercecal field fromthe level of the ovary to about one-fourth of the body length fromthe posterior end. Cirrus pouch ovoid, not extending caudad be-yond center of acetabulum ; cirrus strongly muscular, not protruded.Ovary branched, situated to right of median line immediatelycephalad of testis ; Mehlis's gland globular, 341/* in diameter, situatedin median line. Vitellaria profusely developed, extending to levelof acetabulum on the right side, but stopping abruptly about 1 mmcaudad of acetabulum on the left side ; caudall}' the vitelline folliclescompletely fill the posttesticular area. Uterus closely coiled andoccupying the intercecal field between the anterior testis and acetabu-lum; genital pore 186/* cephalad of anterior margin of acetabulum.Eggs oval, 120/t to 127/* long by 60/* to 67/* wide.Hosts.?Herbivorous mammals and man; marine mammals{Orcinus orca^^Orca gladiator; Balaenoptera acutorostrata?B
.
Tostrata)
.
AUT. 13 TREMATODE PARASITES OF MAEINE MAMMALS PRICE O
Location.?Bile ducts for marine mammals, and usually for otheranimals.Distribution.?Europe (for B. acutorostrata and Orcinus orca).Remarks.?The description of Fasciola hepatica given here is basedupon a carmine-stained toto mount, which Dr. C. W. Stiles kindlyloaned the writer for study, it being one of the specimens referredto below. Aside from a few slight differences, such as body size anda slightly smaller ^gg, this fluke so closely resembles F. hepatica fromruminants that it must be regarded as the same species.There appear to be only two records of Fasciola Jiepatica frommarine mammals. Stiles and Hassall (1894) record this speciesfrom Orca gladiator., and later Stiles (1894, p. 302) states : " Leuckartonce gave me two s]:)ecimens of a fluke, which I still have in mypossession, labeled ' Leber, Schwert-fisch.' I am unable to distin-guish this fluke from Fasciola hepatica. I assume that this ' Schwertfish ' is Orca gladiator rather than Xiphias gladius., as all the otherhosts of F. hcpatica are mammals." Odhner (1905) notes that hefound a fragment of F. hepatica in a vial labeled " Distoma goliathvan Ben.," in the Kopenhagen Museum. Odhner believed that thiswas a case of mislabeling and regarded it as unthinkable that awhale could have become infested with this fluke.The two records cited above may have been the result of mislabel-ing, but if so it is quite remarkable that two such instances shouldhave occurred in which the hosts cited were cetaceans.Subfamily Campulinae Stunkard and Alvey, 1930Synonym..?Brachycladiinae Odhner, 1910c, p. 165.Suhfamily diagnosis.?Fasciolidae : Elongated, medium-sized tovery large flukes; body relatively narrow and thick. Cuticle armedwith relatively large, pointed spines. Suckers close together insome genera and rather widely separated in others. Digestive sys-tem H shaped, except in Odhnericlla; anterior ceca short; posteriorceca long, sinuous and extending to posterior end of body; divertic-ula, when present, are both median and lateral. Excretory vesicletubelike, without lateral branches. Genital aperture preacetabular;cirrus pouch frequently extends far caudad of acetabulum; cirrusarmed or unarmed. Testes slightly or deeply lobed, or smooth,tandem in position, and situated not far caudad of ovary. Ovarysmooth or slightly lobed ; seminal receptacle much reduced in size orabsent; Laurer's canal present. Vitellaria well developed. Uteruswith relatively few loops; vagina with or without spines. Eggswith thickened posterior pole, usually triangular in cross section.Parasites of marine mammals.Type genus.?Campula Cobbold, 1858.
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KBTT TO THE GENERA OF THE SUBFAMILY CAMPUUNAE
1. Cirrus unarmed 2.Cirrus armed 4.2. Body slender, 60 mm to 80 mm long; ovary deeply lobed ; vitella-ria in more or less rectangular masses of follicles. Lecithodesm.us (p. 11).Body more robust, less than 20 mm long ; ovary not lobed ; vitel-laria not as above 3.3. Oral sucker much larger than acetabulum ; eggs circular in crosssection Zalophotrema (p. 13).Oral sucker and acetabulum about equal in size ; eggs triangularin cross section Campula (p. 6).4. "Vagina unarmed; vitelline follicles in distinct masses 5.Vagina armed ; vitelline follicles not in distinct masses 6.5. Anterior ceca absent; in liver of pinnipeds Odhneriella (p. 20).Anterior ceca present; in intestine of cetaceans Hadw^enius (p. 17).6. Testes deeply lobed ; intestinal ceca with median and lateraldiverticula Synthesium (p. 16).Testes not deeply lobed ; intestinal ceca without median andlateral diverticula Orthosplanchnus (p. 14).
Genus CAMPULA Cobbold, 1858Synonym.?Brachycladium Looss, 1899, p. 558.Gener'w diagnosis.?Campulinae: Body elongated and slightlyflattened; posterior end usually more pointed than anterior end.Cuticle completely covered with large, pointed spines. Oral suckerslightly smaller than acetabulum. Intestinal ceca with short medianand lateral diverticula. Excretory vesicle tubelike, extending an-teriorly to ovary. Genital pore immediately cephalad of acetabulum
;
cirrus pouch short; cirrus unarmed; testes with or without lobes,never dendritic. Ovary entire, pretesticular ; seminal receptacle re-duced; Laurer's canal present. Vitellaria profusely developed, ex-tending anteriorly to level of pharynx. Uterus with few coils. Eggstriangular in cross section.Type species.?Gmnpula ohlonga Cobbold, 1858.
KEY TO SPECIES OF THE GENUS OAMPULA
1. Testes lobed 2.Testes not lobed 3.2. Cirrus pouch not extending caudad of acetabulum; intestinalbranches without anal openings palliata (p. 9).Ciriiis pouch extending caudad of acetabulum ; intestinalbranches with anal openings oblonga (p. 7).3. Suckers close together; testes preequatorial rochebruni (p. 11).Suckers widely separated; testes in posterior third of body_ delphini (p. 9).
ART. 13 TREMATODE PARASITES OF MARINE MAMMALS PRICE 7CAMPULA OBLONGA Cobbold, 1858Plate 1, Figures 2-5Synonyms.?Distonmm ohlongum (Cobbold, 1858) Braun, 1892,p. 99; Distomum {Brachylahnus) oblongwiii (Cobbold, 1858) Stos-sich, 1892, pp. 16-17; Brachycladkim ohlongum (Cobbold, 1858)Looss, 1902, p. 716; Distomum tenuicolle Rudolphi of Olsson, 1893,p. 9.Description.?CatnpvJa: Body elongate, 4 to 7 mm long by 1 to3 mm wide, anterior end obtuse, posterior end rounded. Cuticlearmed with pointed spines, 44/x, long by 14/a wide, arranged inalternating transverse rows. Oral sucker subterminal, 310)u, to 340/*in diameter; acetabulum 430^ to 4C5/A in diameter, situated a littleless than one-fourth of the body length from the anterior end.Pre23harynx very short and wide; pharynx piriform, 310/a to 360/along by 170/i to 220^ wide; esophagus about IOO/a long. The short,anteriorly directed, intestinal ceca extend beyond posterior marginof the oral sucker and are not provided with lateral diverticula;the posteriorly directed branches are more or less zigzag andextend to the posterior end of the body, where they open into adepression in common with the excretory vesicle; the posteriorbranches of the intestine are provided with short median and lateraldiverticula. The excretory vesicle is tubelike and extends anteriorly,dorsal to testes, to the level of the posterior margin of the ovary;it is slender at the posterior end, but becomes progressively wideranteriorly. The genital aperture is situated immediately cephaladof the acetabulum; genital sinus small. Cirrus pouch somewhatpestle shaped, slightly curved, and extending caudad to a pointabout midway between the posterior margin of acetabulum and theanterior margin of ovary. The greater part of the cirrus pouch isfilled with a sinuous seminal vesicle. Cirrus unarmed, protrusible.The testes are deeply lobed, tandem in position, and occupy theequatorial third of the body ; the anterior testis is 620ju, to 770/i, longby 770/A to 990/x, wide, and the posterior testis in 620/a to 1 mm long by770/x, to 1.2 mm wide. Ovary transversely oval, 186/x to 372/a longby 310/A to 527yu, wide, situated immediately cephalad of anteriortestis and to the right of the median line. Seminal receptaclegreatly reduced in size or (?) absent (Nicoll described a smallseminal receptacle, but the writer has been unable to demonstratethis structure in the specimens at his disposal) ; Laurer's canalpresent. Vitellaria abundantly developed, the follicles being dis-tributed over the entire dorsal surface from the level of the pos-terior end of the esophagus to the posterior end of body; ventrally
8 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 81
the follicles do not extend medially much beyond the inner limits ofthe intestine in the testicular and pretesticular zones, but com-pletely fill the posttesticular zone. The uterus is relatively short,consisting of a few loops confined to the intercecal field between theovary and acetabulum; vagina about one-half the length of cirruspouch, muscular, and without spines. Eggs 90/* to 97)u, long by 45jliwide, oval in outline, but more or less triangular in cross section;shell yellow, thickened to form a knoblike projection at posteriorpole ; opercular pole flat.Host.?Phocaena. fhocoena {=Phocaena communis).Location.?Bile ducts.Distribution.?Europe; North America (United States).Remarks.?The above description is based upon specimens(U.S.N.M. Helm. Coll. No. 8415) collected by Dr. G. A. MacCallum,June 27, 1925, from the liver of Phocaena yhocoena at Woods Hole,Mass. Two additional lots of specimens have also been examined.The first of these (U.S.N.M. Helm. Coll. No. 3379) was collectedby Prof. Max Braun from the liver of Phocaena commu/nis atWarnemiinde, and identified as CampuJa ohlonga. The date of col-lection is not given, but it is probable that this is a part of the ma-terial upon which his (1900) description of this species is based.The second lot of specimens (U.S.N.M. Helm. Coll. No. 16682) islabeled " Campula ohlonga.^ liver, Phocaena co')n7nunis, Millport.August 15, 1908, determined by Wm. Nicoll." In the case of thismaterial, there appears to be no doubt that this represents a partof the specimens described by Nicoll in 1909. The specimens com-prising both of these lots agree in all essentials with those from theMacCallum collection.One of the outstanding characters which distinguish C. ohlongafrom all other species of the genus is the presence of anal openings.These structures are quite distinct and were found to occur in allspecimens examined.Anal openings have been reported as occurring in species belong-ing to the family Echinostomatidae by Leiper (1908) and by Odhner(1910c) ; in the Steringophoridae by Odhner (1911) ; in the Accacoe-liidae by Looss (1912) ; in the Azygiidae and AUocreadiidae, and inJSchistorchis cameus (syn. Pleorchis oligorchis; family uncertain) byOdhner (1928) ; in the Opecoelidae by Ozaki (1925; 1928) ; and inthe Diploproctodaeidae by LaRue (1926) and by Ozaki (1928).Stunkard (1930) also demonstrated the presence of these openingsin Distomum sp. of Linton, 1899, and proposed a new genus andspecies, Bianium conca^u/m^ for this trematode.The genetic significance of the occurrence of anal openings inthese worms is not clear. In two of the families, Opecoelidae andDiploproctodaeidae, these structures are present in all species in-
ART. 13 TREMATODE PARASITES OF MARINE MAMMALS PRICE 9
eluded in these groups ; in other famihes, however, this does not holdtrue. So far as the family Fasciolidae is concerned, G. oblonga isthe only species known to possess these structures ; it appears, there-fore, that anal openings in some species are characters of specificrather than generic or family significance.CAMPULA PALLIATA (Looss, 1885) Looss, 1901Plate 2, Figures G-7Syno7iy7ns.?Distonvumi palliatum Looss, 1885, pp. 390-427;Brachycladiwni palliatum (Looss, 1885) Looss, 1899, p. 558; Glado-coeliu/m palliatum (Looss, 1885) Stossich, 1892, pp. 10-11.Description.?Campula: Body elongated, 9 mm to 10 mm long by1.5 mm to 2 mm wide and 750/x to 1 mm thick; the anterior end ismore rounded than the posterior end, and there is a definite constric-tion of the body in the vicinity of the acetabulum. Cuticle armedwith closely set rows of pointed spines, 62ju, to 76/i, long by 5/i, to 8/xwide, which completely cover the body. Suckers about equal in sizeand situated 2.5 mm to 3.5 mm apart. Pharynx ovoid, 380/x long by293;u wide; esophagus 540jt* wide. The intestinal tract consists of apair of anteriorly directed ceca, one on each side, which extend to thelevel of the oral sucker, and a pair of posteriorly directed ceca,which extend to the posterior end of the body, both pairs of cecabeing provided with short median and lateral diverticula. Excre-tory vesicle tubular, extending anteriorly dorsal of testes and di-viding into two branches. Genital aperture median and situateda short distance cephalad of acetabulum. Cirrus pouch stronglymuscular and situated mostly in front of acetabulum ; it contains theseminal vesicle, ejaculatory duct, and cirrus. Testes lobed, tandemin position, situated in posterior part of middle third of body.Ovary irregular in outline, 489/x in greatest diameter, situated to leftof median line and cephalad of anterior testis; seminal receptaclesmall ; Laurer's canal present. Vitellaria abundantly developed andconsisting of grapelike follicles, which extend from region ofpharynx to posterior end of intestinal ceca. Uterus consists ofnumerous coils situated dorsal to acetabulum. Eggs 59/j, long by 43ju,wide, ellipsoidal, the opercular pole blunter than the posterior pole.Host.?Delphinus delfliis.Location.?Liver (bile ducts).Distribution.?Europe.CAMPULA DELPHINI (Poirier, 1886) Bittner and Sprehn, 1928Plate 2, Figures 8-10Synonyms.?Distonvwin delphini Poirier, 1886, pp. 34?36; Clado-coelium delphini (Poirier, 1886) Stossich, 1892, p. 10; Brachycla-diuTn, delphini (Poirier, 1886) Looss, 1899, p. 558.
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Description.?Campula: Body flat, 14 mm long by 2 mm wide,slightly attenuated at the extremities. Cuticle covered with smallslender spines. Suckers about equal in size; acetabulum 700/x indiameter, oral sucker slightly smaller; distance between suckers 7mm. Prepharynx short and wide; pharynx piriform in shape, TOO/xlong by 400/x wide; esophagus very short; intestine with short, an-teriorly directed ceca, one on each side, which extend to the levelof the oral sucker, and long posterior ceca, which extend to the pos-terior end of the body. The anterior ceca are provided with threemore or less well-developed lateral diverticula, and the posteriorceca are provided with short diverticula throughout their course,especially along their lateral margins. Genital aperture median, situ-ated a short distance cephalad of acetabulum ; cirrus pouch short andwide, containing the seminal vesicle, short prostatic canal, and a poor-ly developed ejaculatory duct, and situated entirely in front of ace-tabulum. Testes large, ovoid, tandem in position, and situated in an-terior part of the posterior third of body. Ovary small, spherical,situated cephalad of anterior testis and to the right of median line
;
Mehlis's gland well developed, to left of ovary ; Laurer's canal pres-ent. Vitellaria well developed and occupying almost the entire sur-face of body, both dorsally and ventrally; they are composed oftubular glands entangled to form a compact network; the vitellineducts unite cephalad of the genital aperture and caudad of posteriortestis, and also between ovary and anterior testis where they formthe vitelline reservoir. Uterus sinuous, situated between Mehlis'sgland and genital aperture. Eggs elliptical, 60/i, long by 45/i, wide,slightly pointed posteriorly.Host.?Delphinus delphis.Location.?Liver (bile ducts).Distribution.?Europe.Remarks.?Odhner (1905) was of the opinion that this species wasprobably identical with Brachycladium palUatum Looss {= Cain-pula palliata) , since he stated : " Ich bin nlimlich nicht vollig iiber-zeugt, dass Br. delphini nicht mit dem demselben Wirte entstam-menden Br. palUatum am Ende zusammenfallen konnte. Die Differ-enze in der Form der Hoden diirfte fiir das Auseinanderhalten derbeiden Arten kaum geniigen." Odhner's point regarding the formof the testes is well taken, but there are other characters whichappear definitely to eliminate the likelihood of the two species beingidentical. In Campula palliata the uterus forms a rosette mass ofcoils dorsal to the acetabulum, the egg does not show a definite polarprolongation or thickening, and the anterior and posterior vitellineducts do not anastomose. In C. delphini the uterus lies caudad ofthe acetabulum, the egg shows a marked polar thickening or pro-longation, and the vitelline ducts anastomose anterior to the genital
ART. 13 TREMATODE PARASITES OF MARINE MAMMALS PRICE 11
aperture and posterior to the posterior testis. Despite the fact thatPoirier's (1886) description of G. delphini is so somewhat incomplete,the writer feels that a restudy of specimens of this species will showit to be specifically distinct from C. falliata and perhaps to belongto a different genus. Aside from the shape of the ovary and testes,it appears to be more closely related to the genus Lecithodesmus thanto the genus Campula, but the writer prefers to leave it in the lattergenus until an examination of specimens of these forms is possible.CAMPULA ROCHEBRUNI (Poirier, 1886) Bittner and Sprehn, 1928Plate 3, Figures 11-12Synonyms.?Bistomuin rodiebruni Poirier, 1886, pp. 36-37;Cladocoelium rochehruni (Poirier, 1886) Stossich, 1892, p. 11;Brachycladium roche'bruni (Poirier, 1886) Looss, 1899, p. 558.Description.?Camfula: Body narrow, 10 mm long by 1 mm wide;ventral surface flat, dorsal surface slightly convex. Cuticle besetwith very slender spines, which are especially abundant and closetogether on the anterior part of body. Suckers equal in size, 380/xin diameter (Poirier gives the diameter as 0.038 mm, but this un-doubtedly is an error for 0.38 mm) and 700/x. apart. Pharynx 490)nlong by 180/i wide ; esophagus very short ; intestine as in C. delphini.Genital aperture immediately cephalad of acetabulum; cirrus pouchshort and wide, preacetabular. Testes large, ovoid, tandem in posi-tion, and situated near equator of body. Ovary small, spherical,situated cephalad of anterior testis and to right of median line;Mehlis's gland more elongated and situated to left of ovary. Vitel-laria composed of anastomosing tubular glands and extending overthe greater part of body ; the vitelline ducts do not anastomose in theanterior and posterior parts of body as in C. delphini. Uterus withfew loops, situated between Mehlis's gland and genital sinus. Eggsoval, 82/x long by 45/x wide, strongly pointed posteriorly.Host.?Delphinus delpJiis.Location.?Liver.Distribution .?Europe.Genus LECITHODESMUS Braun, 1902Generic diagnosis.?Campulinae: Body long, slender, and flat-tened dorsoventrally. Suckers widely separated. Intestinal cecaprovided with median and lateral dendritic diverticula. Cirruspouch extending slightly beyond posterior margin of acetabulum;cirrus unarmed; testes branched. Ovary deeply lobed; vitellaria inquadrangular groups of follicles extending from region of pharynxto posterior end of body. Parasites of cetaceans.Type species.?Lecithodesmus golinth (van Beneden, 1858)Odhner, 1905.
12 PROCEEDINGS OF THE NATIONAL MUSEUM vol.81LECITHODESMUS GOLIATH (van Beneden, 1858) Odhner, 1905Plate 3, Figures 15-16Synonym.?Distomuiii goliath van Beneden, 1858, pp. 95-97.Description.?Lecithodesmus : Body ribbonlike, 60 to 80 mm longby 8 mm wide and 1.6 to 1.8 mm thick (90 mm long by 9 mmwide, according to Lonnberg, 1891), anterior end bluntly roundedand posterior end slightly attenuated. Spines present on anteriorpart of body. [Odhner (1905) believes that the absence of spineson the posterior part is clue to the effects of maceration for severaldays of habitual delay before specimens can be collected after thehost has been killed.] Oral sucker 2.3 mm in diameter by 2 mmdeep, subterminal in position ; acetabulum 1.8 mm in diameter by 1.6mm deep, according to Odhner (1.3 mm in diameter, according toBraun, 1902b), situated a little more than one-third of the bodylength from the anterior end (28 mm from oral sucker, accordingto Lonnberg). Prepharynx short; pharynx 1.5 mm long by 950/awide (TOO/A wide, according to Braun); esophagus very short; in-testinal ceca extend to posterior end of body and are provided withmedian and lateral dendritic diverticula. The anteriorly directedcecal appendages extend to the level of the middle of the pharynx,each being provided with four lateral diverticula. Excretory vesicletubular and extending to level of ovary. Genital pore preacetabu-lar; cirrus pouch club shaped, containing a large seminal vesicleand an unarmed cirrus 3 mm to 4 mm long. Testes branched, tan-dem in position, and situated in the posterior half of the body.Ovary star shaped, situated immediately cephalad of the anteriortestis, slightly to right of median line. Laurer's canal present;seminal receptacle (?). The vitellaria consist of quadrangulargroups of follicles extending both dorsally and ventrally from thelevel of the pharynx to the posterior end of body. According toBraun, the vitelline ducts consist of a long, slender, anterior duct,which bifurcates to form two lateral ducts that extend to a shortdistance caudad of ovary where they join the transverse ducts, andof a shorter, unpaired, posterior duct which bifurcates to form an-terior lateral ducts which are sometimes connected by transverseanastomoses; the transverse ducts are formed by the union of theanterior and posterior lateral ducts. Uterus convoluted and termi-nating in a well-developed vagina, which opens into the genitalsinus near the male genital aperture. Eggs 120/x long by 75/x wide,triangular in cross section.Hosts.?Balaenoptera acutorostrata {=B. rostrata)., B. horealis,and Balaena mysticetus.Location.?Liver.Distribution.?Europe.
ART. 13 TREMATODE PARASITES OF MARINE MAMMALS PRICE 13Genus ZALOPHOTR'EMA Stunkard and Alvey, 1929Generic diagnosis.?Campulinae : Body flattened, somewhat atten-uated posteriorly and rounded anteriorly. Oral sucker terminal,larger than acetabulum. Intestinal ceca as in Gampula; posteriorceca without anal openings. Excretory pore terminal ; excretoryvesicle tubular and extending to ovary. Cirrus pouch extendingcaudad of ovary; cirrus unarmed. Ovary indented but not lobed,pretesticular and slightly to right of median line; seminal receptacleabsent; Laurer's canal present. Vitellaria as in Camjyula. Uteruscoiled. Eggs with thickening at posterior pole, circular in crosssection. Parasites of pinnipeds.Type species.?Zalophotreina hepaticum Stunkard and Alvey,1929. ZALOPHOTREMA HEPATICUM Stunkard and Alvey, 1929Plate 3, Figures 13, 14Description.?Zalophotrema: Body elongate and flattened dorso-ventrally, 11 to 13 mm long by 3 to 3.6 mm wide, and from600/A to 1 mm thick, the thickness decreasing abruptly imme-diately caudad of testes. Cuticle armed with spines, which varyfrom 14^ to 35/i, in length. Oral sucker 700/^ to 800//, long by 800/ato 1 mm wide; oral opening slightly subterminal. Acetabulum570/A to 620/A in diameter, situated one-fifth to one-sixth of the bodylength from the anterior end. Pharynx piriform, 550/u, to 600/x longby 380ja to 420/A wide. The digestive system bifurcates a short dis-tance caudad of pharynx into two laterally directed branches, eachof which bifurcates into a short anterior and a long posterior cecum
;
the anterior ceca terminate near the anterior end of pharnyx; theposterior ceca are provided with many median and lateral diverticula,and terminate near the posterior end of the body. Excretory poreterminal ; excretory vesicle extending anteriorly as a simple sac tothe region of the ootype. Cirrus pouch relatively weakly developed,extending from genital pore to level of ovary, according to Stunkardand Alvey (1930), and containing a coiled seminal vesicle, prostate
cells, and an unarmed cirrus. Testes large and deeply lobed, tandemin position, and occupying the equatorial third of body. Ovarylobed, 500/A long by 1 mm wide, situated at caudal end of anteriorthird of body ; Mehlis's gland posterior and dorsal to ovary ; seminalreceptacle absent; Laurer's canal present. The vitellaria consist ofmasses of follicles, which extend in the lateral areas of the bodyfrom the level of the pharynx to the posterior end ; they tend to bedorsal in position in the anterior part of body, almost meeting inthe median line cephalad of genital pore, while caudad of testesthey are both dorsal and ventral in position and invade the median
14 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 81
field. The uterus consists of masses of coils filling the central partof body from the ovary to the acetabulum. The vagina is aboutone-half the length of the cirrus pouch. Eggs oval, 68/a to 79ja longby 43/A to 52ju wide, thickened at posterior pole and circular incross section.Host.?Zalophus californianus.Location.?Bile ducts.Distribution.?North America?United States (New York Aqua-rium and National Zoological Park, Washington, D, C).Remarks.?The foregoing description is slightly modified fromthat given by Stunkard and Alvey (1930).Four specimens (U.S.N.M. Helm. Coll. No. 29731) of what ap-pear to be this species were collected, March 17, 1930, from theliver of a specimen of Zalophus californianus^ which died in theNational Zoological Park, Washington, D. C. These specimensagree in all essential characters with the description given by Stunk-ard and Alvey except as regards the branching of the anterior cecaand in the length of the cirrus pouch. As regards the anterior ceca,Stunkard and Alvey state that " each of the anterior branches gives
oif two or three diverticula," but in the specimens at the writer's dis-posal no such diverticula were found. Two of the specimens showeda slight irregularity in the diameter of these ceca, but this appearedto be due to distention with ingesta. The cirrus pouch is describedby Stunkard and Alvey as extending caudally to the level of theovary, but in the writer's specimens this structure was found toextend to about midway between the posterior margin of the acetab-ulum and the anterior margin of the ovary. These differences,however, appear too slight to warrant considering the possibility ofthe writer's specimens representing a species distinct from Z.hepaticvmi. Genus ORTHOSPLANCHNUS Odlmer, 1905Generic diagnosis.?Campulinae : Body elongated and slightlyflattened; cuticle covered with pointed spines. Intestinal ceca with-out lateral or median diverticula. Genital pore preacetabular ; cirruspouch long and pestle shaped, extending beyond posterior marginof acetabulum; cirrus armed with strong pointed spines. Testeswith indented margins, postequatorial and tandem in position.Ovary without lobes ; seminal receptacle very small ; Laurer's canalpresent. Vitellaria profusely developed, the follicles being dis-tributed both dorsally and ventrally from region of pharjaix to pos-terior end of body. Uterus with few coils; vagina well developedand lined with strong pointed spines. Eggs triangular in crosssection. Parasites of pinnipeds.Type species.?Ortliosplaivchnus arcticus Odhner, 1905.
AUT. 13 TKEMATODE PAEASITES OF MAKINE MAMMALS PRICE 15
KEY TO SPECIES OF ORTHOSPLANCHNUS
1. Body almost cylindrical ; ventrally the vitelline follicles encroachon the testicular and uterine fields but do not extend ante-riorly as far as pharynx fraterculus (p. 16).Body somewhat flattened ; ventrally the vitelline follicles donot encroach on testicular and uterine fields but extendanteriorly beyond level of posterior end of pharynx arcticus (p. 15).ORTHOSPLANCHNUS ARCTICUS Odhncr, 1905Plate 4, Figxjkes 17, 18
Desc7^ption.?Orthosplanchnus : Body slightly flattened, 3.5 mmto 7 mm, usually 4.5 mm to 6 mm long by 850/a to 1.15 mm wide,tapering toward both ends, the posterior part of body being slightlymore attenuated than the anterior part. Cuticle spiny throughout,the spines on the anterior part of body being about 40/x long, whilethose at both ends are somewhat smaller. Oral sucker 480/a to 600/iin diameter, subterminal in position; acetabulum 450;U, to 530/a indiameter, situated about one-fourth of body length from anteriorend. Prepharynx about 200/t long, pharynx about 400|U, long by SOO^awide, esophagus 120ju to IbOjx long. Intestine with short anteriorlydirected ceca, one on each side, which extend to level of anterior endof pharynx, and long posterior ceca, which extend to end of body;both the anterior and posterior ceca are simple and without diver-ticula. Excretory pore terminal ; excretory vesicle simple and tubu-lar, and extending anteriorly to level of anterior margin of anteriortestis. Genital aperture preacetabular, median ; genital sinus spa-cious. Cirrus pouch pestle shaped and extending caudally to aboutmidway between acetabulum and anterior testis (about two-fifthsof the distance from acetabulum to ovary, according to Cooper,1921) ; seminal vesicle more or less spherical and about 250/^ indiameter when filled with sperms, more tubelike when empty ; parsprostatica cylindrical, 170/^ to 200/x long, and set off from the seminalvesicle by a sharp constriction; ejaculatory duct about the samelength as pars prostatica; cirrus protrusible and spiny, the spine?40/A to 45/x long by 7ju, wide at their bases and inseited into a basaidisk ISju, to 19/t in diameter by 12/a thick. Testes elongated oval inshape, with indented margins; the anterior testis is usually slightlysmaller than the posterior; they are tandem in position and situ-ated in the anterior half of the posterior part of the body. Ovaryentire, transversely oval in shape, situated cephalad of anteriortestis and to the right of the median line; seminal receptacle verysmall; Laurer's canal present. Vitellaria profusely developed andextending both dorsal ly and ventrally of intestinal ceca from thelevel of the pharynx to the posterior end of the body ; anterior to
16 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 81
acetabulum the follicles form a transverse dorsal band across body
;
ventrally the follicles do not encroach on the testicular and uterinefields; caudad of the testes the follicles coalesce in the median lineand completely fill the posttesticular zone. The uterus extendsanteriorly from Mehlis's gland in transverse loops to the acetabulum,where it terminates in a well-developed vagina, the cuticular liningof which is armed with spines similar to those of the cirrus. Eggs91/x to lOOfx long by 54/* to 58fi wide, triangular in cross section, andhaving a thickened posterior pole.Hosts.?Erignathus harbatus (= Phoca harhata) ; Phoca klspida.Location.?Gall bladder and liver.Distribution.?Europe (west coasts of Greenland and Spits-bergen) ; North America (Canada?Bernard Harbor, Northwest Ter-ritories?according to Cooper, 1921).ORTHOSPLANCHNUS FRATERCULUS Odhner, 1905Plate 4, Figxtre 19
Description.? rthosplanchnus : Body almost cylindrical, 3 mm to4 mm long by 500/a to 600//, wide. Cuticular spines more abundantthan in O. arcticus. Oral sucker 370/^ to 440/a in diameter; aceta-bulum 400/i, to 500/x in diameter. Pharynx 300/a to 330/a long by 200/i,wide; esophagus 220/a long. Cirrus pouch as in 0. arcticus. Testesdeeply indented. Vitellaria do not unite to form a dorsal band an-terior to acetabulum, and ventrally they encroach on the testicularand uterine fields. Other characters as in O. arcticus.Hosts.?Odohenus rosniarusj Erignathus barbatu^.Location.?Gall bladder.Distribution.?Europe (Spitsbergen).Genus SYNTHESIUM Stunkard and Alvey, 1930Generic diagnosis.?Campulinae : Body long and slender. Suckersof equal size. Digestive system as in Campula. Cirrus pouch longand slender, inclosing seminal vesicle, pars prostatica, and spinycirrus; testes deeply lobed as in Lecithodesmus. Ovary spherical,pretesticular ; vitellaria consist of small, grapelike clusters of fol-licles and do not extend anteriorly as far as acetabulum ; uterus withfew coils; vagina lined with spines. Eggs with polar thickening.Parasitic in intestine of cetaceans.Type species.?Synthesium tursionis (Marchi, 1873).SYNTHESIUM TURSIONIS (Marchi, 1873). new combinationPlate 5, Figures 20-21Synonyms.?Distomuin tursionis Marchi, 1873, p. 304; Distonvmnlongissimum- Poirier, 1886, pp. 29-30, not D. longissimum von Linstow,
ART. 13 TEEMATODE PARASITES OF MARINE MAMMALS PRICE 17
1896; D. {Dicrocoelium) tursionis (Marchi, 1873) Parona, 1896, p.162.Description.?Synthesium: Body whitish, 20 mm long by 1.5 mmwide, flattened ; cuticle spiny. Suckers equal in size, 800/i in diam-eter; acetabulum situated 3 mm to 4 mm caudad of oral sucker.Prepharynx 1 mm long ; pharynx 720/* long by 300/x wide ; intestinalceca as in BrachycJadiuvi { = Campula)., according to Odhner (1926).Genital aperture preacetabular ; cirrus pouch long and tubular, andcontaining the seminal vesicle, a long slender pars prostatica, andcirrus (cirrus as in Orthosplanchnus., according to Odhner). Testesas in Lecltliodesmus^ according to Odhner; anterior testis with fivelobes and posterior with six lobes, situated in posterior third of body,according to Poirier. Ovary small, almost spherical, pretesticular,and situated near the equator of body ; Mehlis's gland a little largerthan ovary. Vitellaria well developed and consisting of small grape-like clusters of follicles ; they extend from a short distance cephaladof ovary to the posterior end of body. Uterus sinuous ; vagina as inOrtlios'planchnus. Eggs oval, 56|U, long by 33/* wide, with thickeningat posterior pole.Host.?Delphlnus tursio {^vohahly^Turslops truncatus).Location.?Intestine.Distribution.?Europe.Remarhs.?This description is taken largely from that given byPoirier (1886) for Distomum, longisshnuvi. The description givenby Marchi (1873) for D. tursionis is very incomplete and the figureaccompanying it is very sketchy. Parona (1896) restudied speci-mens from Marchi's original material and pointed out that D. tur-sionis Marchi and D. longissirawm Poirier were identical, but addednothing to the original descriptions. Odhner (1926) reported thatthis species had the digestive tract of Brachycladiuvi {^ Campula)
.,the lobed testes of Lecithodesmus., and the characteristic copulatoryorgans (charakterischen Endapparate der Geschlechtswege) of Or-tJiosplanchnus., and suggested that it might be included in the lattergenus. Kecently Stunkard and Alvey (1930) proposed the genusSynthesium for this species. The writer is in accord with this action,since the inclusion of this species in the genus Ortliosplanchnus.,or in any of the related genera, would necessitate extensive revisionof the diagnoses of these groups; this is regarded as inadvisableuntil more species have been described.HADWENIUS, new genusGeneric diagnosis.?Campulinae : Body very long and slender,cylindrical, suckers close together; oral sucker slightly larger thanacetabulum. Cuticle of anterior part of body spiny. Intestinal cecawithout diverticula. Excretory vesicle tubular, extending anteriorly118893?32 2
18 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 81
to near posterior margin of ovary. Cirrus pouch pestle shaped,extending caudad of acetabuhim ; cirrus spiny as in Orthosplanchnus.Testes oval, tandem in position, situated in anterior fourth of body.Ovary transversely oval, pretesticular ; seminal receptacle absent;Laurer's canal present. Vitellaria composed of rosette masses ofradiating cords of follicles, which extend from vicinity of anteriortestis to posterior end of body. Uterus with few coils confined tointercecal space between ovary and acetabulum; vagina well devel-oped, unarmed. Eggs triangular in cross section, slightly thickenedat posterior pole. Parasites of cetaceans.Type species.?Hadwenius seymouri^ new species.HADWENIUS SEYMOURI, new speciesPlate 6, Figures 23-25
Description.?Hadwenius : Body slender, 27 mm to 60 mm long by1.5 mm to 2 mm wide, and almost circular in cross section. Thecuticle of the interior part of the body is spiny, the spines beingabout 27)a long by 7/x wide and arranged in alternating transverserows; they are deep-set in the cuticle so that only the tips projectabove the surface. The rows of spines are close together in theregion immediately caudad of the oral sucker, but become fartherapart as they approach the region of the anterior testis, where theydisappear completely. The oral sucker is cup shaped, 1.8 mm to2 mm long by 1.7 mm to 2 mm in diameter, the thickness of the wallbeing about 190/i, ; the oral aperture is 540jm to 900|U, in diameter andslightly subterminal. The acetabulum is transversely oval, 930/1,to 1.2 mm long by 1.3 mm to 1.5 mm wide, and the walls are about180^ thick; the distance between the suckers is from 800/x to 1.6 mm,depending upon the degree of contraction. The length of the pre-pharynx is variable; in some specimens it is about 810/x long, whilein others the pharynx is drawn into the base of the oral sucker sothat the prepharynx is very wide and short. The pharynx is 900ju,to 1 mm long by 620/^ to 900/x wide. The esophagus is very short andwide. The intestine is H shaped as in other members of the sub-family; the ceca are straight and without lateral or median diver-ticula. The excretory pore is situated at the summit of a papillalikeprominence, which projects into a deep pit or depression at theposterior end of the body. The excretory vesicle is similar to that inother members of the subfamily. The genital aperture is situatedimmediately in front of the anterior margin of the acetabulum;it communicates with a spacious genital sinus. The cirrus pouch ispestle shaped, about 1.8 mm long by 560/x wide ; it extends caudad alittle more than half the distance between the acetabulum and
ART. 13 TREMATODE PARASITES OF MARINE MAMMALS PRICE 19
ovary. The seminal vesicle is about 830/* long bj^ 290/>i wide andalmost fills the posterior part of the cirrus pouch; pars prostaticaslender, about 700/a long, and separated from the seminal vesicle bya sharp constriction; ejaculatory duct relatively short. The cirrusis protrusible and armed with strong spines. The spines are about40/t long and are inserted into a basal disk which is about 16/i, indiameter. Testes oval in shape and situated in the anterior fourthof the body; the anterior testis is 900/a to 1.5 mm long by 620/* to930/* wide, and the posterior 1 mm to 1.6 mm long by 620/a to 850,awide, the distance between them being 310/x to 1.2 mm. The ovaryis transversely oval, 232/* to 387/* long by 465/* to 590/* wide, situateda short distance cephalad of the anterior testis and to the right of themedian line. Seminal receptacle not observed. Laurer's canal isslender and sinuous, and opens in the mid-dorsal line at the levelof the ovary. Mehlis's gland is large and is situated median anddorsal to the ovary. The vitellaria consist of chainlike rows offollicles, which radiate to form rosettelike masses, and extend fromthe anterior testis to the posterior end of the body; the masses offollicles are distributed on all sides and form a continuous layer be-neath the dermomuscular layer of the body. The uterus consists ofsix or more transverse coils confined to the intercecal field betweenMehlis's gland and the acetabulum. The vagina is well developed,unarmed, and about one-half the length of the cirrus pouch ; it opensat the base of the genital sinus to the left of the male genital aper-ture. The eggs are oval, 97/* long by 52/* wide, with a short prolonga-tion at the posterior pole, triangular in cross section.Host.?White whale {Delphinapterus leucas).Location.?Intestine.Distribution.?North America (Alaska).Type sfecimens.?U.S.N.M. Helm. Coll. No. 30807; paratypes,No. 26157. Collected by Dr. Seymour Hadwen, September 9, 1921,at Golovin, Alaska.Remarks.?Iladicenius seyiihouri appears to be more closely relatedto Synthesiuin tursionis than to any of the other species of Campu-linae. Both are parasites of the intestinal tract of cetaceans and aresimilar in body fonu. They differ, however, in two principal charac-ters, which are considered generic, viz, the copulatory organs anddistribution of the vitellaria. In Iladwenius seymouri the cirrus isarmed but the vagina is not, and the vitelline follicles are arrangedin rosettelike masses similar to those in Lecithodesmus^ while inSynthesiv/tn tursionis both cirrus and vagina are armed and the vitel-line follicles are distributed in small grapelike groups. Other dif-ferences are present, but these appear to be only of specific value.
20 PROCEEDINGS OF THE NATIONAL MUSEUM vol. SIGenus ODHNERIELLA Skrjabin, 1915Generic diagnosis.?CamjDulinae : Body fiat and ribbonlike ; cuticleof preacetabular part of body armed with spines. Oral suckerslightly smaller than acetabulum. Digestive tract without anteriorlydirected ceca. Excretory vesicle as in Canipula. Cirrus pouch sac-like, extending beyond posterior margin of acetabulum ; cirrus armedwith spines as in Orthosplanchnus ; testes entire. Ovary entire, pre-testicular. Vitellaria consisting of grapelike masses of follicles, sit-uated laterall}^ and not invading median field, and extending fromabout midw^ay between ovary and acetabulum to level of terminationof ceca. Uterus relatively short; vagina unarmed. Eggs triangu-lar in cross section. Parasites of pinnipeds.Type species.?Odhneriella rossica Skrjabin, 1915.ODHNERIELLA ROSSICA Skrjabin, 1915Plate 5, Figure 22
Description.?Odhneriella: Body flat and ribbonlike, 9 mm longby 760/x wide at acetabulum ; sides of body almost parallel. Cuticleof preacetabular part of body armed with spines. Oral sucker di-rected ventrally, 500ja long by 480/1, to 530/*, wide; acetabulum 500/j.long by 680/1 wide, slightly elevated above surface of body, and sit-uated in anterior part of body. Prepharynx 119/t long; pharynx325/t long by 290/i wide; esophagus 230/t long; intestinal ceca simpleand extending to po.sterior end of body; anteriorly directed ceca,characteristic of other members of the subfamily, absent. Excretorypore terminal; excretory vesicle as in other members of the sub-family. Genital pore median, near anterior margin of acetabulum.Cirrus pouch saclike, extending beyond posterior margin of acetab-ulum; cirrus strong and armed with spines. Testes oval, 935/i longby 390/1 wide, entire, tandem in position, and situated in the posteriorfourth of the anterior half of the body. Ovary globular, 300/i, to340/1 in diameter, pretesticular. Vitellaria consisting of grapelikemasses of follicles distributed along sides of body and extendingfrom about midway between ovary and acetabulum to level of endsof ceca. Uterus short, with few coils, situated in the intercecal fieldbetween ovary and acetabulum; vagina straight and unarmed.Eggs oval, 100/t long by 60/* wide, thickened at posterior pole, tri-angular in cross section.Host.?Odohenus rosmarus.Location.?Bile ducts.Distribution.?Europe (Russia).
AUT. 13 TREMATODE PARASITES OF MARINE MAMMALS PRICE 21
Remarks.?The foregoing description, taken from Skrjabin(1915)/ was based upon specimens collected by Doctor Staroka-domsky, February 8, 1912, in north Russia, near KaluchinskayaBay. Family ECHINOSTOMATIDAE Loess, 1902Family diagnosis.?Body more or less elongate, small or verylarge, usually much flattened anteriorly, less so, or even cylindrical,posteriorly. Oral sucker small and weak, surrounded dorsally andlaterally, but not ventrally, by a collarlike fold, bearing one or tworows of spines, which are continued laterally to ventral corners,the corner spines usually large or specialized ; acetabulum large andpowerful, usually preequatorial and near oral sucker. Cuticle usu-ally spinose, especially anteriorly. Excretory vesicle Y shaped, withlateral twiglike branches. Pharynx and epithelial " pseudoesoph-agus " present ; intestinal ceca extend to posterior end of body.Genital aperture preacetabular; genital sinus present or absent;cirrus pouch usually present. Testes postequatorial, usually tandemin position. Ovary pretesticular, usually to right of median line;Laurer's canal present. Vitellaria lateral, rarely extending anteriorto acetabulum. Uterus in transverse coils, rarely extending beyondintercecal field. Parasites of intestines or bile ducts of vertebrates,especially birds.Type genus.?Echinostoma Rudolphi, 1809.
Genus ECHINOSTOMA Rudolphi, 1809 (sensu lato)Generic diagnosis.?Characters of the family.ECHINOSTOMA ACANTHOIDES (Rudolphi, 1819) Cobbold, 1860SynonyTn.?Distoma acanthoides^ Rudolphi, 1819, p. 114.Description.?Ecldnostoma: Body elongated and flattened (4 mmto 6 mm long, according to Dujardin, 1845), divided into a somewhatslender (271/x wide) anterior part and a broader (520/* wide) poste-rior part (Dujardin gives the maximum width of the body as 750/1.)the two portions being united at the level of the acetabulum. Oralsucker 15G/x long by 135/i wide; acetabulum 375/t long by 396/a wide,situated about one-third of the body length from the anterior end.Cephalic collar provided with four spines, 73ju, long, on each ventrallobe, with 16 to 18 smaller spines, 59/i long, arranged around the mar-gin in a single row, uninterrupted dorsally, and with one small spine,
^ The writer Is indebted to Dr. R. Ed. Schulz, of the School of Veterinary Medicine,Moscow, for a tjpt'writton cojiy of Professor Skrjabin's paper, the publication in whichthis paper appeared being unavailable in this country.
22 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 81
26/x long, on each side between the larger spines of the ventrallobes and the smaller marginal spines. Pharynx 145/a long by114/x wide, situated almost immediately caudad of the oral sucker.Testes globular 159/a in diameter, tandem in position and situatedin the posterior part of the body. Viteliaria lateral, not extendinganteriorly beyond acetabulum.Host.?Phoca vitulina.Location.?Intestine.Distribution.?Europe (Berlin, Germany).Remarks.?Rudolphi (1819) reported finding two specimens ofthis species in the above host in Berlin. Braun (1901a) redescribedthis form on the basis of the original material and stated that thespecimens were immature. The above description is compiledlargely from that given by Braun.It is not certain that this species belongs to the genus Echinostoina.,sensu stricto. The arrangement of the spines of the cej)halic collartends to exclude it from the genus, but the other characters are notsufficiently well described to permit a definite generic assignmentat present. Genus STEPHANOPRORA Odhner, 1902Synonyms.?Mesorchis Dietz, 1909a, p. 183; Monilifer Dietz,1909a, p. 183.Generic diagnosis.?Echinostomatidae : Body elongated and sub-cylindrical. Cuticle of anterior part of body armed with spines.Oral sucker surrounded by a well-developed reniform collar bearinga single row of spines, which is interrupted dorsally by a spaceabout as wide as oral sucker. Acetabulum situated approximatelyone-fourth of body length from anterior end. Cirrus pouch welldeveloped, containing seminal vesicle, pars prostatica, and a short,strong cirrus. Testes tandem in position, situated near equatorof body. Ovary pretesticular. Viteliaria almost filling posttesticu-lar part of body and extending anteriorly to anterior testis. Uterusmoderately long and containing relatively few eggs.Ty^e species.?Stepkanoprora omata Odhner, 1902.STEPHANOPRORA DENTICULATA (Rudolphi, 1802) Odhner, 1910Plate 7, Figures 20-30Synonyms.?Fasciola denticulata Rudolphi, 1802, p. 91 ; Distomadenticulatwn (Rudolphi, 1802) Rudolphi, 1809, p. 424-425; Echin-ostama denticulatum (Rudolphi, 1802) Cobbold, 1860, p. 36;Echinostoma {Mesorchis) denticulatmn (Rudolphi, 1802) Dietz,1909a, p. 183; Mesorchis denticulatus (Rudolphi, 1802) Dietz, 1909b,p. 31.
ART. 13 TEEMATODE PARASITES OF MARINE MAMMALS PRICE 23
Description.?Stephanoprora: Bodj^ elongated, 2.08 mm to2.64 mm long by 288/x to 352/* wide at acetabulum. Cuticle of an-terior part of body closely beset with spines arranged in alternatingtransverse rows ; these rows are close together in the region anteriorto acetabulum, but caudad of this point they are more widelyseparated and finally disappear in the region of the anterior testis.Oral sucker subterminal, 74/* to 92/x in diameter, surrounded by awell-defined reniform collar, 213/i to 237/* wide. The collar bears22 spines arranged in a single row, which is interrupted dorsallyby a space almost as wide as the oral sucker. Four of these spines,two on each ventral lobe, are slightly smaller and more aboral thanthe marginal spines; they measure 29/* to 37/t long by 11/* to 14/iwide; the marginal spines are 37/i to 44/* long by 11/* to 15/i wide.Acetabulum almost circular, 185/* to 222/t in diameter, situated 370/*to 560/1 from anterior end of body. Prepharynx 37/* long; pharynx81/* to 110/t long by 74/1 wide; esophagus 92/i to 166/t long; intestinalceca slender. Genital pore situated about midway between intestinalbifurcation and anterior margin of acetabulum. Cirrus pouchovoid, 185/1 to 222/t long by 48/i to 129/i wide, containing a largeseminal vesicle, pars prostatica, and short cirrus. Testes tandemin position, situated in equatorial zone; anterior testis globular,166/* to 229/1 in diameter; posterior testis ovoid, 203/* to 259/1 longby 148/1 to 222/t wide. Ovary transversely oval, 85/t to llO/i long by100/t to 122/t wide, situated cephalad of anterior testis and to rightof median line. Vitellaria consisting of large follicles and extend-ing from level of posterior margin of anterior testis to near pos-terior end of body. Uterus short and with few coils. Eggs 74/ito 77/t long by 52/t to 55/t wide.Host.?Zaloph us californiatviis.Location.?Small intestine.Distrlhufion.?North America?United States (Washington,D. C).jSpecimens.?U.S.'N.M. Helm. Coll. No. 28147. Collected by E. W.Price, June 1, 1928, from a California sea lion, which died in theNational Zoological Park.^to^Family TROGLOTREMATIDAE Odhner, 1914Family diagnosis.?Body compact, more or less flattened ventrallyand convex dorsally. Cuticle with pointed spines. Body muscula-ture as well as that of suckers weakly developed. Excretory vesicleY shaped or tubular. Pharynx present; esophagus short; intestinalceca extend to near posterior end of body. Genital pore nearacetabulum, either immediately in front or behind, median or slightlyto the left. Cirrus pouch absent except in Troglotrenna; pars prosta-
24 PROCEEDINGS OF THE NATIONAL MUSEUM vol.81
tica and seminal vesicle always distinct. Testes opposite eachother, equatorial or postequatorial. Ovary usually lobed, dextral,pretesticular ; seminal receptacle and Laurer's canal present. Vitel-iaria usually well developed, exclusively or mostly dorsal, leavingonly the median dorsal area of body unoccupied. Uterus eithervery long, with relatively few loops, or shorter and more convoluted.Parasites of carnivorous mammals and birds, usually occurring inpairs in cystlike cavities.Type genus.?Troglotreiiia Odhner, 1914.
Genus PHOLETER Odhner, 1914Generic diagnosis.?Troglotrematidae : Body more or less spindleshaped. Cuticle armed with small pointed spines, the spines not ingroups. Excretory vesicle Y shaped, the bifurcation occurring infront of testes, and branches extending to acetabulum. Genitalaperture at anterior border of acetabulum, slightly to left of medianline ; genital sinus moderately deep and wide ; cirrus pouch absent
;
pars prostatica short, directed dorsoventrally ; seminal vesicle tube-like, undivided, extending under the dorsal surface to near the ovary.Testes elliptical, situated opposite each other a short distance fromthe posterior end of body. Ovary deeply lobed; seminal receptaclepresent; Laurer's canal moderately long. Vitellaria strongly de-veloped, dorsal in position, and having a tendency to occur in grape-like bunches. Uterus long and convoluted, occupying entire bodywidth and extending from ovary to genital pore. Parasites ofcetaceans.Type species.?Pholeter gastrophilus (Kossack, 1910) Odhner,1914. PHOLETER GASTROPHILUS (Kossack, 1910) Odhner, 1914Plate 7, Figure 26Synonym.?Distomuni gastrophiluni Kossack, 1910, pp. 118-120.Description.?Pholeter : Body spindle shaped, 1.5 mm to 3.33 mmlong by 1.7 mm to 2.1 mm wide according to Odhner (1914), or3.15 mm to 3.66 mm long by 1.8 mm to 2.25 mm wide according toKossack (1910), the thickness being about one-third of the width.Oral sucker l70/x to 200/a in diameter; acetabulum 250/i to 300/a indiameter, situated about one-third of the body length from the ante-rior end. Pharynx 150/x to 170;u, in diameter; esophagus of samelength as pharynx ; intestinal ceca simple and terminating about themiddle of testes. Genital pore situated at the anterior margin ofacetabulum; cirrus pouch absent; pars prostatica short and directed
AKT. 13 TREMATODE PARASITES OF MARINE MAMMALS PRICE 25dorsoventrally ; seminal vesicle tubelike; undivided. Testes ellip-tical, situated side by side in the posterior third of the body. Ovarydeeply lobed, situated immediately cephalad of testes and slightly toone side of the median line; seminal receptacle moderately large,dorsal to ovary. Vitellaria dorsal and extending from about halfwaybetween intestinal bifurcation and acetabulum to the level of the endsof the ceca. Uterus long and strongly looped, occupying almost theentire width of the body from the ovary to the genital pore ; vaginashort. Eggs oval, 23/i* to 25/a long by 14/x wide.Host.?Fhocaena phocoena {?P. communis).Location..?Stomach (encysted in the mucosa of the pylorus).Distribution.?Europe.
Family OPISTHORCHIIDAE Braun, 1901Family diagnosis.?Medium-sized to small forms; body elongate,flat, thin, transparent, with weak musculature. Suckers relativelyweakly develoiDcd. Intestinal ceca simple, extending to posteriorend of body. Excretory vesicle Y shaped with long stem and shortbranches. Genital aperture median, immediately preacetabular ; nogenital sinus containing suckerlike structures or gonotyls. Cirruspouch and cirrus absent; seminal vesicle coiled. Testes in post-equatorial region, situated more or less obliquely. Ovary pretes-ticular; seminal receptacle voluminous; Laurer's canal present.Vitellaria lateral of ceca, moderately developed, not reaching poste-rior end of body. Uterus long, with numerous loops, usually confinedto intercecal space between ovary and acetabulum. Eggs small andnumerous. Parasites of bile ducts and gall bladder of mammals,birds, fishes, and reptiles.Type genus.?Opisthorchis R. Blanchard, 1895.
KEY TO SUBFAMILIES OF OPISTHORCHUDAB
1. Vitellaria and uterus do not extend cephalad of acetabulum.Opisthorchiinae (p. 25).Vitellaria and uterus extend cephalad of acetabulum Tletorchiinae (p. 30).Subfamily Opisthorchiinae Looss, 1899
, Subfamily diagnosis.?Opisthorchiidae : Excretory pore terminal
;
stem of excretory vesicle long. Vitellaria do not extend anteriorlybeyond level of acetabulum. Uterine coils usually confined to inter-cecal space and not extending anteriorly beyond acetabuhnn.Type genus.?Opisthorchis R. Blanchard, 1895.
26 PROCEEDINGS OF THE NATIONAL. MUSEUM vol.81
KEY TO GENERA OF OPISTHORCHnNAB OCCURRING IN MARINE MAMMALS
1. Uterine coils extend laterally beyond limits of intestinal ceca.Cyclorchis (p. 28).Uterine coils do not extend laterally beyond limits of intestinalceca 2.2. Vitellaria divided into two distinct regions by a break at thelevel of the ovary Amphimerus (p. 29).Vitellaria not divided into tvpo regions Opisthorcliis (p. 26).Genus OPISTHORCHIS R. Blanchard, 1895Generic diagnosis.?Opisthorchiinae : Body elongated, flattened,anterior end attenuated, posterior end broader. Cuticle generallysmooth, without spines. Excretory vesicle Y shaped, with S-shapedstem and short branches. Cirrus pouch absent; testes usually lobed,situated in posterior part of body and placed obliquely to the longaxis of the body. Ovary simple or lobate, pretesticular ; seminalreceptacle prominent, postovarial; Laurer's canal present. Uteruswith numerous transverse loops confined to intercecal field, not ex-tending anteriorly beyond acetabulum. Vitellaria moderately de-veloped, extracecal, not extending anteriorly beyond level of aceta-bulum or posteriorly beyond level of ovary. Parasites of bile ductsof mammals, birds, and fishes.2'ype species.?Opisthorchis felineus (Rivolta, 1884) R. Blanchard,1895=6*. tenuicollis (Rudolphi, 1819).OPISTHORCHIS TENUICOLLIS (Rndolphi, 1819) Stiles and Hassall, 1896Plate 7, Figures 27, 28Synonyms.?Distoma tenuicollis Rudolphi, 1819, p. 93; D. conusGurlt, 1831, p. 193, not Creplin, 1870; D. lanceolatmn von Siebold,1836, p. 113, not Rudolphi, 1803 ; D. felineum Rivolta, 1884, pp. 20-28; D. viverrini Poirier, 1886, pp. 116-130; Opisthorchis felineus(Rivolta, 1884) R. Blanchard, 1895, p. 217; 0. viverrini (Poirier,1886) Stiles and Hassall, 1896, p. 155; O. tenuicollis-felineus Looss,1899, p. 678.Description.?Opisthorchis: Body flat, 6.5 mm to 8.5 mm long by2.1 mm to 2.2 mm wide, anterior end somewhat more attenuatedthan posterior end. Cuticle smooth and without spines. Oral suckersubterminal, 320/* to 340/x. in diameter; acetabulum 260;a to 320/1, longby 300/A to 360/A wide, situated 1.4 mm to 1.6 mm from the anteriorend of body. Pharynx ovoid to piriform in shape, 200/x long by140/A to 160/z, wide; esophagus 80/a to 140/x, long; intestinal cecaslender and extending to near posterior end of body, the left cecumbeing slightly shorter than the right. Excretory vesicle Y shaped,with a long sigmoid stem and short branches. Genital aperture im-mediately cephalad of acetabulum. Seminal vesicle more or less
ART. 13 TEEMATODE PARASITES OF MARINE MAMMALS PRICE 27
spirally coiled and free in parenchyma, situated posterior to ace-tabulum and to the right of the median line. Testes lobed andsituated in the posterior third of body; the anterior testis has fourlobes and measures 500//, to 600/x long by 660/a to 700/x, wide; theposterior testis has five lobes and is 540/* to 700/a long by 680/a toTOOjU, wide. Ovary more or less trilobed, 160/x to 200/* long by 400/ito 440/1, wide, situated slightly to right of median line and about400/A to 440/A cephalad of anterior testis. Mehlis's gland diffuse,dorsad and cephalad of ovary; seminal receptacle large, somewhatovoid or retort shaped, situated to the right and caudad of ovary;Laurer's canal long and slender. Vitellaria extracecal, each one com-posed of eight poorly defined groups of follicles which extend froma short distance caudad of acetabulum to level of ovary. Uteruswith closely packed loops confined to intercecal field between ovaryand acetabulum. Eggs oval, 27/x, to 31/* long by 13/* to 15/* wide.Hosts.?Erignathus harhatus {^Phoca barbata), Halichoerusgrypus^ Phocaena phocoena {^Delphinus phocoena?PhocaenacomTnunis)^ Gulo borealis, Fells vivei^ina, domestic cat, dog, andman.Location.?Liver (bile ducts).Distribution.?Europe; Asia (Siberia).Remarks.?The above description is based upon specimens(U.S.N.M. Helm. Coll. No. 3357) labeled '' Opisthorchis felineus(Riv.), Haliclioei^s grypus., Konigsberg Thiergarten, collected anddetermined by Mtihling," which were donated to the helminthologi-cal collection by Prof. Max Liihe, June, 1902. These specimens areconsiderably smaller than the measurements given by various authorsfor C. tenuicollis^ but so far as can be determined from the litera-ture, this species exhibits considerable variation as regards size.Whether Opisthorchis tenuicolUs and O. felineus are identicalspecies appears to be a moot question. Braun (1893) stated: "Soreiht sich Dist. tenuicolle Rud. aus Phoca barbata dem Dlst. feVmemnRiv. und verwandten Arten an." Miihling (1896, 1898a, and 1898b)was convinced of their morphological identity, as was Looss (1899).Barker (1911) noted that there was a lack of specific characters thatwould definitely separate the two species. Morgan (1927) also statesthat several species of the genus, including O. tenwicoUis and 0.felineus^ are very similar and of questionable validity, and points outthat widely different hosts, when feeding on the same intermediatehost, may become infested with the same species of fluke. Afterstudying the descriptions and figures of (9. tenuicolUs and O. felineus
.^the writer is convinced that they are the same morphological speciesand should no longer be regarded as distinct. There also appearsto be no good reason for considering O. viverrini as valid, especiallysince the figure given by Fuhrmann (1928) shows that the uterine
28 PROCEEDINGS OF THE NATIONAL. MUSEUM vol. 81
coils are more closely packed than those figured by Poirier (1886) forthis species, the distribution of the uterine coils being essentially theonly character by which this species could be differentiated fromO. tenv/icoUis { = 0. felineus). It is also possible that O. entzi^ de-scribed by von Ratz (1900) from the gall bladder of Ardea purpu-rea^ and 0. geminus, described hj Looss (1896) from the liver ofMilvus parasiticivs, are species identical with O. tenuicoUis, sincethe characters given fall within the range of variation exhibited inthe latter species.It appears that in the case of O. teiiuicoUis there is a lack of hostspecificity, as is also the case for Gryptocotyle lingua and certainother trematodes.
Genus CYCLORCHIS Luhe. 1908
Generic diagnosis.?Opisthorchiinae : Body more or less spindleshaped, the maximum width being near the equator. Cuticle with-out spines. Suckers about equal in size. Digestive tract and excre-tory vesicle as in Opisthorchis. Testes globular, situated in poster-ior fourth of body. Ovary and adjacent structures as in Opisthor-chis. Vitellaria lateral, situated in posterior half of body, but notextending caudally beyond level of seminal receptacle. Uterine coilsloosely arranged and extending laterally beyond limits of ceca.Type species.?Cyclorchis amphileucus (Looss, 1896) Liihe, 1908.
CYCLORCHIS CAMPULA (Cobbold, 1876) Luhe, 1908
Plate 8, Figure 31Synonyms.?Distoma campula Cobbold, 1876, p. 40; Metorchiscampula (Cobbold, 1876) Looss, 1899, p. 565; Opisthorchis campula(Cobbold, 1876) Looss, 1899, p. 559.Description.?Cyclorchis: Body elliptical, about 3 mm long by 1mm wide, slightly more attenuated anteriorly than posteriorly. Oralsucker subterminal; acetabulum about the same size as oral sucker,situated about one-fourth of the body length from the anterior end.Esophagus short; intestinal ceca relatively wide and sinuous, ex-tending to posterior end of body. Genital pore preacetabular ; testesovoid and situated diagonally to the long axis in the posterior fourthof body. Ovary small; seminal receptacle large. The body whichCobbold says is " apparently the ovary " is probably the distendedseminal receptacle, and the smaller body immediately in front of itis probably the ovary. Vitellaria (?). The uterus passes poster-iorly and forms a transverse loop between the ovary and testes, andthen passes anteriorly in transverse loops which extend laterally be-
AiiT. 13 TEEMATODE PARASITES OF MARINE MAMMALS PRICE 29yond the inner limits of the ceca. Eggs " 1/1000 of an inch frompole to pole by 1/2100 inch in breadth."Host.?PJatanista gangetica.Location.?Bile ducts.Distribution.?Asia (India).Remarks.?The foregoing description is taken largely from Cob-bold's (1876) figure of D'lstotna campula. The description whichhe gives for this form is very incomplete, and almost no measure-ments are given. Cobbold confused this species with Gaiiifula oh-lo7iga^ a species which he had described earlier from Phocaena pho-coeiia i=P. coTnnmmis). his identification being based largely onthe zigzag course of the intestinal ceca. The species from Platanistagangetica is unquestionably an opisthorchid and has been placed inthe genus Cyclorchi^ by Liihe (1908) as species inquirenda. Thedisposition of the reproductive organs, so far as they have been fig-ured by Cobbold, is strikingly similar to that in C. amphileucus sothat there appears to be good reason for including it in the samegenus. Genus AMPHIMERUS Barker, 1911Generic diagnosis.?Opisthorchiinae : Body elongated and flat-tened, anterior end attenuated. Cuticle frequently covered whollyor in part with small spines. Excretory vesicle as in Opisthorchis.Cirrus pouch and cirrus absent ; testes in posterior part of body, sim-ple or lobate, situated diagonally to long axis of body. Ovary an-terior to testes, simple or lobate; seminal receptacle well developed;Laurer's canal present. Vitellaria well developed, lateral of in-testinal ceca, divided into two distinct regions by a break oppositethe ovary, not extending anteriorly beyond the acetabulum, but fre-quently extending posteriorly to or beyond the posterior testis.Uterus anterior to ovary as in Opisthorchis; the coils may extendlaterally beyond the inner limits of the ceca. Parasites of the bileducts of mammals, birds, and reptiles.Type species.?Amphimerus ovalis Barker, 1911.AMPHIMERUS LANCEA (Dicsing. 1850) Barker, 1911Plate S, Figueks 32-33Synonyms.?Distommii lancea Diesing, 1850, p. 334; Opisthorchislancea (Diesing, 1850) Braun, 1901c, p. 897.Description.?Aniphimeriis: Bod}^' lanceolate, 5.5 mm to 12.5 mmlong by 1 mm to 2.8 mm wide ; anterior end conical and shorter thanthe flattened posterior portion ; margins of posterior portion serrated.Oral sucker subterminal, 330/x to 360/x by 510/* to C60/a; acetabulum510/x to 1.2 mm in diameter, situated one-third of the body length
30 PROCEEDINGS OP THE NATIONAL MUSEUM vol.8]from the anterior end. Genital pore preacetabular and median inposition. Testes spherical or lobed, slightly oblique in position andsituated in the posterior part of the body. Ovary biscuit shaped;seminal receptacle generally spindle shaped, situated caudad and tothe right of the ovary ; Mehlis's gland indistinct. Vitellaria lateralto intestinal ceca, consisting of eight groups of follicles on each sideof body divided into two regions by a break between the fourth andfifth groups, and extending from a short distance caudad of theacetabulum to the ends of the intestinal ceca. Uterus consisting oftransverse coils which extend intercecally from the ovary to theacetabulum. Eggs oval, 29/x, to 33/x long by 12|u, to 14/i, wide.Hosts.?Delphinus tacuschi {])voh&h\y^ Sotalia tucuxi)^ (?)Orcaella hrevirostris.Location.?Not given ; probably bile ducts.Distribution.?Sou:th America (Brazil?Barra do Rio Negro),(?) Asia (India).Cobbold (1876) reported what he thought was this species fromOrcaella hrevirostris, the specimens upon which the report was basedhaving been collected in " the North-eastern Province of India " byDr. John Anderson, superintendent of the Indian Museum, Calcutta.The character which apparently caused Cobbold to regard the formfrom India as the same as that from Brazil was the "irregularlyserrated " margin of the body, since he states : " I know of no othertrematode possessing these sinuosities." The description and figurewhich he gave are quite different than those given by Diesing (1855)and by Weski (1900). The writer doubts whether the form whichCobbold calls Distoma lancea is the same as Diesing's species, but onaccount of the incompleteness of his description and figure, no opin-ion is expressed as to its probable affinities.Subfamily Metorchiinae Liihe, 1909Suhfamily diagnosis.?Opisthorchiidae : Excretory pore ventral;stem of excretory vesicle usually short, ventral to testes. Vitellariaextend cephalad of acetabulum. Uterine coils frequently overlapceca and extend cephalad of acetabulum.Type geniis.?Metorchis Looss, 1899.
KETS" TO GENERA OF METORCHIINAE OCCUREING IN MARINE MAMMALS
1. Posterior end truncate and suckerlike Pseudamphistomum (p. 31).Posterior end rounded Metorchis (p. 30).Genus METORCHIS Looss, 1899Generic diagnosis.?Metorchiinae: Body rounded posteriorly andattenuated anteriorly. Cuticle usually covered with spines. Intes-tinal ceca extend to posterior end of body. Testes large, usually
AET. 13 TEEMATODE PARASITES OF MARINE MAMMALS^ PRICE 31
lobed, and more or less obliquely placed, and almost filling posteriorpart of body. Ovary, Mehlis's gland, seminal receptacle, andLaurer's canal as in Opisthorchis. Vitellaria compact and extend-ing anteriorly beyond acetabulum. Uterine coils often extend extra-cecally and preacetabular. Parasitic in gall bladder and bile ductsof mammals and birds.Type species.?Metorchis aJhidus (Braun, 1893) Looss, 1889.
METORCHIS ALBIDUS (Braun, 1893) Looss, 1899Plate S, Figube 34Synonyms.?Distomwn albidum Braun, 1893, pp. 347-355; D. {Di-crocoeliunv) alhidum Braun, 1893, p. 353; Opistliorcliis albidus(Braun, 1893) Railliet, 1896, p. IGO.Description.?Metorchis : Body spatulate, 1.6 mm to 2.2 mm longby 800ju, to 1 mm wide ; anterior part of body narrower than the flatposterior part. Cuticle covered with small spines. Oral sucker sub-terminal, 200/A to 240)a in diameter ; acetabulum 200/^ in diameter andsituated 600/i, to 900/i, from the anterior end of the body. Pharynxovoid, 75/x to 90/x long by 47/a to 85^ wide ; esophagus 28fx long ; in-testinal ceca extend to posterior end of body. Genital aperture pre-acetabular; seminal vesicle relatively short. Testes lobed, obliquein position, the left testis anterior to right, and situated in the pos-terior jDart of the body; the anterior testis is 300/a to 34:0ju, long by300/x to 380/A wide, and the posterior 320/a long by 380/* wide. Ovarysomewhat triangular in shape, lOOju, to ISOjx long by 140ja to 180/xwide, situated a short distance in front of anterior testis. Seminalvesicle large and situated posterolateral of ovary; Mehlis's glanddiffuse. Vitellaria lateral and extending from near level of intestinalbifurcation to level of ovary. Uterus greatly convoluted and occupy-ing the greater part of the intervitellarian field from the ovary toa short distance cephalad of acetabulum. Eggs 27/i to 32/x, long by13/x to 16/1 wide.Hosts.?Ualichoerus g7'ypus, Felis domestica^ Vulpes vulpes, andCanis fatniliaris.Location.?Gall bladder and bile ducts.Distribution.?Europe.
Genus PSEUDAMPHISTOMUM Luhe, 1908
Generic diagnosis.?Metorchiinae : Body conical in shape, anteriorend pointed ; posterior end truncate and surrounded by a ridge giv-ing it a suckerlike appearance. Cuticle beset with fine spines. Ex-cretory pore in center of posterior suckerlike structure. Intestinalceca slightly sinuous, extending to posterior end of body. Genital
32 PROCEEDINGS OF THE NATIONAL MUSEUM vol.81
pore preacetabular ; cirrus pouch and cirrus absent; seminal vesicleconvoluted, free in parenchyma. Testes in posterior third of body,placed slightly obliquely to long axis of body. Ovary median in po-sition, situated about midway between anterior border of testes andacetabulum; seminal receptacle voluminous, postovarial. Vitellariaextracecal and consisting of relatively large groups of follicles ex-tending from level of seminal receptacle to level of genital pore orslightly beyond. Uterus greatly convoluted, extending laterally overceca and anteriorly beyond acetabulum. Parasites of bile ducts ofmammals.Type species.?PseudamphistoTnuin truncatwm (Rudolphi, 1819)Luhe, 1908.
PSEUDAMPHISTOMUM TKUNCATUM (Rudolphi. 1819) Luhe. 1908
Plate 8, Figure 35
Synonyvis.?Amphistoma truncatum Rudolphi, 1819, p. 91; Bis-ioma coivm Creplin, 1825, pp. 50-53; Distomum lanceolafuni Mehlisof Diesing, 1858, p. 332 ; Distoma cannpanulatmn Ercolani, 1875, pp.432-439; Metorchis truncatus (Rudolphi, 1819) Looss, 1899, p. 565.Description.?Pseudainpliistomwm: Body conical, 2 mm long, an-terior end pointed, posterior end truncate and suckerlike. Cuticlethickly and regularly beset wdth fine spines. Oral sucker 132)ii to137ja in diameter ; acetabulum about same size as oral sucker and sit-uated slightly preequatorial. Excretory pore situated in the de-pression of the posterior suckerlike structure. Pharynx 91/x long;esophagus very short ; intestinal ceca slightly sinuous and extendingto posterior end of body. Genital aperture preacetabular; seminalvesicle convoluted and free in parenchyma. Testes globular orslightly ellii^tical, 172/* to 376/x long, situated in the posterior thirdof the body, one slightly in front of the other. Ovary globular,situated about midway between the anterior border of the anteriortestis and the acetabulum, usually obscured by the uterine coils;seminal receptacle voluminous, postovarial. Vitellaria situated lat-erally, chiefly in the middle third of body, each composed of 10 to 12groups of follicles. Uterus greatly convoluted, occupying the greaterpart of the body width between testes and acetabulum, and extend-ing somewhat in front of acetabulum. Eggs 29ju, long by 11/x wide.Hosts.?Phoca vitulina, P. groenJandica, P. hispida= Halichoerusfoetidus, Halichoerus gryjyus, Gido horeaUs.) Felis domestica, CanisfaTniliaris, C. vulpes.Location.?Bile ducts.Distribution.?Europe (Germany, Holland, Italy, France).
AET. 13 TEEMATODE PARASITES OF MAEINE MAMMALS?PRICE 33Family HETEROPHYIDAE Odhner, 1914Family diagnosis.?Small or very small forms, usually not exceed-ing 2 mm in length. Anterior portion of body thinner, usuallymore slender and more movable than the posterior portion. Surfaceof body covered with small scalelike spines that become reducedposteriorly and may disappear tovt^ard the posterior end of the body.Intestinal ceca simple, usually extending to the posterior end of thebody. Genital pore in the immediate neighborhood of the aceta-bulum; genital ducts usually open into a genital sinus, which maybe variously modified and contain a cirruslike body or gonotyl(genital sucker). Acetabulum usually median in position, but maybe displaced to the right of the median line ; in some instances theacetabulum may be joartially or completely atrophied and inclosedin the genital sinus. Cirrus pouch absent. Seminal vesicle well de-veloped, U or S shaped, the vas deferens surrounded proximally bya mass of prostatic cells. Testes oval, globular, or slightly lobed,near the posterior end of the body, side by side, or obliquely one infront of the other. Ovary oval, globular or slightly lobed, pretes-ticular, usually to the right of the median line. Seminal receptacleand Laurer's canal present near the ovary, usually in relation withits posterior border. Vitellaria, located mainly in the lateral fields,may extend anteriorly to or beyond the genital aperture. Uterususually restricted to the intercecal field between the ovary andgenital pore, but may extend to posterior end of body {Galacto-somum). Adults parasitic in the intestine of birds and mammals.Type genus.?Heterophyes Cobbold, 1866.KEY TO THE GENEEA OF HETEKOPHYIDAE OCCUERING IN MARINE MAMMALS
1. Acetabulum absent; uterus extending to posterior end of body.Galactosomum (p. 39).Acetabulum present ; uterus not extending caudally beyond an-tei'ior border of testes 2.2. Acetabulum not contained in the genital sinus; seminal recep-tacle median and slightly preovarial ; vitellaria not extendingto acetabulum Phocitrema (p. 38).Acetabulum contained in the genital sinus ; seminal receptaclelateral and postovarial ; vitellaria extending cephalad ofacetabulum 3.3. Genital sinus large; genital ducts open into sinus caudad ofacetabulum Cryptocoyle (p. 33).Genital sinus small ; genital ducts open into sinus cephalad ofacetabulum Apophallus (p. 35).
Genus CRYPTOCOTYLE Luhe, 1899Synonyms.?Tocotrem^ Looss, 1899, p. 585; Uallum Wigdor, 1918,p. 254; Ciureana Skrjabin, 1923, p. 67.118893?32 3
34 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 81Generic diagnosis.?Heterophyidae : Body ovoid to linguiform inshape. Prepharynx very short; esophagus short; intestinal bifurca-tion nearer to oral sucker than to acetabulum ; intestinal ceca extendinto posterior end of body and terminate caudad of testes. Aceta-bulum rudimentary, in anterior wall of the spacious, more or lessmuscular, genital sinus; genital ducts open into sinus at base ofa single papillalike gonotyl ; genital aperture postacetabular, incenter of genital sinus. Seminal vesicle well developed, curved ina more or less S-like manner, dorsal to uterine coils. Testes nearposterior end of body, irregularly oval or slightly lobed, either sideby side or the right testis obliquely behind left. Ovary irregularlyoval or lobed, situated to right of median line and cephalad of semi-nal receptacle. Vitellaria fill posttesticular space and extend an-teriorly to acetabulum or beyond. Uterus with few loops, confinedto intercecal space between ovary and genital sinus.Type species.?Cryptocotyle concava (Creplin, 1825) Fischoeder,1903. CRYPTOCOTYLE LINGUA (Creplin, 1825) Fischoeder, 1903Plate 9, Figure 36Synonyms.?Distoma lingua Creplin, 1825, i^p. 27-38; Tocotremalingua (Creplin, 1825) Looss, 1899, p. 586; Distomum maci^orhinisMacCallum, 1916, p. 34; Hallum caninum Wigdor, 1918, pp. 254-257.Description.?Cryptocotyle: Body linguiform, 550/x to 2 mm longby 200/x, to 900ju. wide. Cuticle covered with scalelike spines, 2/a to4/A long by about 1/x wide. Oral sucker 66/x to 110/x in diameter;prepharynx shorter than pharynx ; pharynx 40/a to SO/x long by 30ju,to 48/A wide. Esophagus short, about 40^ to 60/a long. Genital sinus120/tA to 250ja in diameter, situated near equator of body ; acetabulumrudimentary, in anterior wall of sinus. Seminal vesicle long andcoiled in an S-like manner, extending caudally to about the levelof the anterior border of ovary. Testes irregularly globular orovoid in shape, 120/x to 250/i, by 70ja to 130ju,, margins uneven orslightly lobed. Ovary lobed, 70/a to 120/j, long by 140ft to 180fi wide,situated to right of median line; seminal receptacle ovoid in shapeand situated caudad of ovar3^ Vitellaria extend anteriorly beyondacetabulum and caudally to posterior end of body. Uterus confinedto intercecal space between ovary and genital sinus. Eggs oval, 49ftto 50fi long by 18ft to 2ofi wide.Hosts.?Birds {Golynibus aurltus., Gavia iviniev., Larus marinus,L. argentatus, L. fuscus, L. atriciUa, Nycticorax nycticorax.^ RissatHdactyla, Alca torda^ Steriui dougalli, S. hirundo) and mammals
ART. 13 TEEMATODE PARASITES OF MARINE MAMMALS PRICE 35{Canis familiaris^ Vulpes fulva, Phoca vituUna, and Miroungaangustirostris)
.
Location.?Small intestine.Distrihution.?Europe and North America (United States,Canada)
.
Remarks.?Cryptocotyle lingua appears to have been reported buttwice from pinnipeds. Ransom (1920) reported this species fromPhoca vitulina, the report being based upon specimens (U.S.N.M.Helm. Coll. No. 4280) collected by Dr. Albert Hassall, December 21,1905, at Washington, D. C. MacCallum (1916) described a trem-atode, Distomum macrorhinis., from specimens collected from amelephant seal, Macrorhinus angustirostris {=Mirouriga angustiros-tris) , which died at the New York Aquarium. The writer has exam-ined the specimens upon which MacCallum based his descriptionof D. macrorhlnis and finds no differences which would warrant re-garding this form as a species distinct from Gryptocohjle lingua.
Genus APOPHALLUS Luhe, 1909Synonyms.?Rossicotrema Skrjabin and Lindtrop, 1919, p. 40;Cotylophallu-s Ransom, 1920, p. 529.Generic diagnosis.?Heteroph3adae : Body ovoid to very elongatedin shape. Prepharynx short; esophagus long; intestinal bifurcationusually nearer to acetabulum than to oral sucker; intestinal cecaslender, terminating as in Cryptocotyle. Acetabulum relativelywell developed, opening into a small, nonmuscular genital sinus;genital ducts open into sinus at the base of two papilliform gonotyls
;
genital aperture cephalad of acetabulum. Seminal vesicle well de-veloped, C or S shaped, dorsal to uterine coils. Testes ovoid orglobular, situated near posterior end of body, the right usually be-hind left. Ovary ovoid or globular, situated to right of medianline cephalad of seminal receptacle. Vitellaria fill posttesticularspace and extend usually to acetabulum or beyond. Uterus as inCryptocotyle.Type species.?Apophallus miihlingi (Jagerskiold, 1899) Liihe,1909.
KEY TO SPECIES OF THE GENUS APOPIIAT.LUS OCCURRING IN MARINE MAMMALS
1. Esophagus longer tliaii prepharynx; seminal vesicle slender, Sshaped ; vitelline follicles relatively small, not extendinganteriorly as far as pharynx donicus (p. 3G).Esophagus shorter than prepharynx ; seminal vesicle wide, Cshaped ; vitelline follicles relatively large, extending to levelof pharynx _ zalophi (p. 36).
36 PROCEEDIlSrGS OF THE NATIONAL MUSEUM vol.81APOPHALLUS DONICUS (Skrjabin and Lindtrop, 1919) Price, 1931Plate 9, Figubb 37Synonyms.?Rossicotrema donicuin Skrjabin and Lindtrop, 1919,pp. 41-42; R. simile (Kansom, 1920) Ciurea, 1924, p. 14; Cotylo-phallus venustus Eansom, 1920, p. 555; G. sirnilis Eansom, 1920, p.555.Description.?Apophallus: Body ovoid to linguiform in shape,500/A to 1.14 mm long by 200fji to 'SdOfi wide. Cuticular scalelikespines 4/x to 7.5/x long by 1.5ju to 3^^ wide. Oral sucker 65/x to 85/u, indiameter; prepharynx very short; pharynx 30/x to 44/x, in diameter;esophagus slender, bifurcating 135ja to 265ja from the anterior end ofbody ; intestinal ceca simple, extending into posterior fourth of body.Acetabulum 45/i, to 58/x long by 48/x to 60/i wide, situated 185/x to560/1 from the anterior end of body. Testes oval or globular inshape, 80fi to 200/x by 60)U to 200/i,, situated obliquely in extendedspecimens, more or less opposed in more contracted specimens, andoccupying the posterior third of the body. Ovary 65ja to 140/i by40/* to 120/A, situated 200/* to 750/t from the anterior end of body.Seminal receptacle 60/i to 130/* wide by 35/* to 90/x long, situatedbetween the posteromedian border of the ovary and the antero-median border of the left testis. Vitellaria well developed, extend-ing from posterior end of body to slightly beyond the bifurcationof the ceca. Uterus with few coils and occupying the intercecalsjDace between anterior border of the left testis and the anteriormargin of acetabulum. Eggs 30fi to 35/* long by 16/c to 20/* wide.Hosts.?Canis familiaris, Felis domestica.^ Viilpes lagop^is^ andPhoca vitulina.Location.?Small intestine.Distribution.?Europe and North America (United States).Reniarhs.?This description is taken from that of Ransom (1920)for Gotylophallus sifnilis, the specimens (U.S.N.M. Helm. Coll. No.4279) upon which the description was based having been collectedfrom the harbor seal by Dr. Albert Hassall, in Washington, D. C,December 21, 1905. The writer has compared these specimens withspecimens from dogs and cats and agrees with Witenberg (1929) thatthere is no reason for regarding this form as a species distinct fromA. donicus {= R. donicum).APOPHALLUS ZALOPHI, new speciesPlate 9, FiGxmE 38
Desc7'iption.?Apophallus : Body elongated piriform in shape, 435/ilong by 215/* to 263/i wide at the level of the ovary. The cuticle isbeset with small scalelike spines, 4/* long by 2/i wide, arranged in al-
ART. 13 TEEMATODE PAIJASITES OF MARINE MAMMALS PRICE 37
ternating tranverse rows. Oral sucker slightly subterminal in posi-tion, 60/x to 75/x, in diameter. Prepharynx SOjw, to 33/x long ; pharynxovoid to spherical in shape, 29/x, to 33/* wide; esophagus 18/a long;intestinal eeca relatively wide and extending to near the posteriorend of the body, their blind ends being hidden by the testes. Theacetabulum is circular, 52/a to 60/1 in diameter, situated from 235/* to259/A from the anterior end of the body and inclosed in the shallowgenital sinus. The genital ducts open into the anterior part of thesinus and two elliptical gonotyls are present, one on each side of thegenital aperture. The seminal vesicle is voluminous, more or less Cshaped, and lying to the right of the acetabulum; there is a sharpconstriction of the vesicle near the level of the posterior margin ofthe acetabulum which divides it into an anterior piriform part anda posterior globular part. The testes are somewhat triangular inoutline, 81/i to 96/t by Sl/i to llO/i, and are situated side by side in theposterior fourth of the body. The ovary is more or less triangularin outline, 55/i to To/i by 67/a to 92/x,, situated a short distance cephaladof the right testis. The seminal receptacle is spherical, 44ju, in di-ameter, and situated dorsal to the ovary and right testis. The vitel-laria consist of large, closely packed follicles, which extend from thelevel of the acetabulum to the level of the anterior margin of thetestes ; the follicles are distributed over the entire dorsal surface butventrally they are chiefly lateral except near the intestinal bifurca-tion where they form a distinct band across the body. The uterusconsists of a few loops confined to the intercecal field between theanterior margin of the testes and the genital aperture. The eggs are33/i long by 18/* wide, golden yellow, and slightly piriform in shape.Host.?Zalophus califomianus.Location.?Small intestine.Distribution.?North America (United States?National Zoo-logical Park, Washington, D. C).Type specimens.?U.S.N.M. Helm. Coll. No. 30808; paratypes,No. 26652.Remarks.?Apophallus zalophi is easily distinguished from A.donious, the only species of the genus with which it might possiblybe confused, by its size, relative length of the prepharynx andesophagus, and by the distribution of the vitellaria. A. zalophi ison the whole a much smaller species than A. donicus and the body issomewhat thicker. The prepharynx is longer than the esophagus inA. zalophi, while in A. donicus the reverse is true. The vitelline fol-licles are relatively larger in A. zalophi and extend from the levelof the pharynx to the anterior margin of the testes, while in A.donicus the follicles extend from about midway between thepharynx and intestinal bifurcation to the posterior end of the body.The arrangement of the genital glands is essentially the same in
38 PROCEEDINGS OF THE NATIONAL MUSEUM vol.81both species; there is, however, a greater tendency for the testes tobe opposed in A. zalophi than in A. donicus.
Genus PHOCITREMA Goto and Ozaki, 1930Generic diagnosis.?Heterophyidae : Body fusiform in shape, cu-ticle spiny. Oral sucker terminal; acetabulum not inclosed in agenital sinus. Prepharynx shorter than esophagus; intestinal cecasimple and terminating at the anterior border of testes. Genitalpore median, immediately preacetabular ; testes almost directlyopposite each other in j^osterior part of body; seminal vesicleslender, C shaped, lying free in parenchyma. Ovary oval, pre-testicular, situated to right of median line ; seminal receptacle largeand situated anteromedian of ovary; vitellaria lateral, in posteriorhalf of body ; uterus v^ith few transverse coils extending intercecallyand extracecally between testes and genital pore.Type species.?Phocitrema fusiforme Goto and Ozaki, 1930.Remarks.?Goto and Ozaki (1930) place this genus in the familyOpisthorchiiclae. It appears to the writer, however, that its affini-ties are with the Heterophyidae rather than with the Opisthorchiidae,and it is included in this family in spite of the apparent absence ofsome characters which would definitely determine its systematicposition. PHOCITREMA FUSIFORME Goto and Ozaki. 1930Plate 9, Figxjee 40Description.?Pliocitreina: Body fusiform in shape, 1.16 mm to1.4 mm long by 500/>i to 620jii wide. Cuticle covered with minutespines. Oral sucker terminal, SO/x long by 60/i wide; acetabulum120/x in diameter, slightly preequatorial in position. Prepharynx80^ long by 60/>i wide ; esophagus 120ja to ISO/n long, bifurcating aboutmidway between pharynx and acetabulum; intestinal ceca terminateat the level of the anterior border of the testes. Genital pore medianand situated immediately cephalad of the acetabulum; seminal vesicleelongate, cylindrical, forming a transverse loop between ovary andacetabulum ; vas deferens slender, surrounded by glandular cells, ex-panding at anterior margin of acetabulum to form a bulbous parsprostatica which is surrounded by conspicuous gland cells. Testesoval or reniform in shape, 160/li to 240)ii by 120/i, to 140/x, situatedopposite each other in the anterior part of the posterior fourth ofbody. Ovary ovoid, about the size of one of the testes, situated tothe right of the median line in front of the right testis. Seminalreceptacle large, situated obliquely anterior to ovary. Vitellariaconsist of five to seven groups of small follicles on each side of the
ART. 13 TEEMATODE PAEASITES OF MAEINE MAMMALS PRICE 39
posterior half of body. The uterus begins with irregular transversecoils on the left side of ovary, then passes to the right around theovary and forms two transverse loops caudad of the acetabulum, theloops extending into the extracecal fields; vagina nearly straight,14:0fx long, opening close to the male aperture. Eggs 23 to 28ju, longby 13 to 15/i, wide.Host.?Phoca hisplda.Location.?Intestine.Distribution.?Asia (Japan?Hanayashiki Zoological Garden,Tokyo). Genus GALACTOSOMUM Looss, 1899Synonyms.?Mic7'olistruni Braun, 1901b, p. 5G3 ; Gercarioides Wit-enberg, 1929, p. 138.Generic diagnosis.?Heterophyidae : Body elongated and spindle-like in outline, or expanded anteriorly and more or less cylindricalposteriorly. Prepharynx usually distinct, longer than esophagus;intestinal ceca simple, extending to posterior end of body. Genitalaperture situated medially, about one-third of the body length fromthe anterior end. Genital sinus complicated, containing a spiny,spheroidal, more or less muscular body (gonotyl ?) imbedded in thedorsal wall of the sinus. Seminal vesicle well developed, free inparenchyma, the terminal portion of the vesicle being provided witha muscular wall (expulsor). Testes ovoid, spherical or slightlylobed, tandem or slightly oblique in position, and situated in theposterior half of the body. Ovary spherical or ovoid in shape, situ-ated slightly to the right of the median line and cephalad of theseminal receptacle. Vitellaria extracecal or expanding medially inthe posttesticular space, and extending anteriorly as far as the an-terior testis or beyond. Uterus with a descending and an ascendinglimb, both passing between testes.Type species.?Galactosonium lacteum (Jiigerskiold, 1896) Looss,1902.Remarks.?The genus Galactosomum was proposed by Looss(1899) to contain Monostomum lacteum Jiigerskiold. Braun (1901b)proposed the genus MieroUstrutn to contain three species, Distomutncoclileariforme Kudolphi, 1819, D. cochlear Diesing, 1850, and M.spinetum Braun, 1901b. Odhner (1910a) pointed out that the struc-ture which Braun interpreted as an acetabulum in the species in-cluded in the genus Microlistrum was the same structure as thatdescribed by Jiigerskiold (1896) as the " stacheligen Korper " inGalactosom/wm lacteum.. Other similarities were also noted betweenthe two genera. Pratt (1911) gave a redescription of M. cochleari-forme and showed that the character of the genitalia was essentially
40 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 81
the same as that described by Jagerskiold for Monosto^num lacteuvi;on the basis of this similarity he transferred M. cochlearifornie to thegenus Galactosomum^ making the two genera synonymous. Travas-sos (1929) apparently recognizes this arrangement, but Witenberg(1929) regards the two genera as distinct "because of differences inthe arrangement of the genital glands." The writer (Price, 1931)has shown that the arrangement of the genital glands is subject toconsiderable variation within a genus and is a character of no ge-neric importance in the Heterophyidae.In the same paper, Witenberg proposed a new genus, Gercaur-ioides^ as the basis for a new subfamily, Cercarioidinae, the typespecies, G. aharonii, being characterized by having a dilated anteriorpart of the body set off by a slight constriction from the morecylindrical posterior part. To this new genus, Nazmi (1930) addedan additional species, G. l)aylisi. A comparison of the descriptionsshows no essential differences between these species and those belong-ing to the genus Oalactosomum. The anterior widening of thebody in G . aharonii and in G. haylisi is onlj^ slightly more pronouncedthan that in G. cochlearifoi-ine; the arrangement of the genitalglands, the course of the uterus, and the terminal portions of thegenital ducts also appear to be similar. Therefore, the writerregards Gercarioides Witenberg as a synonym of Galactosoiii'wnLooss, the two species G. aharonii Witenberg and G . haylisi Nazmibecoming G. aharonii (Witenberg) and G. haylisi (Nazmi), re-spectively.GALACTOSOMUM ERINACEUM (Poirier, 1886) Bittner and Sprehn, 1928Plate 9, Figure 39/Synonyms.?Distonvwm erinacevmi Poirier, 1886, pp. 37-38; Astio-trema erinacea (Poirier, 1886) Stossich, 1904, p. 2.Description.?Galactosonvmn : Body elongated, 3 mm long by 800/awide, the anterior half of the body being wider than the posteriorhalf. Cuticle beset with spines, which become less numerous towardthe posterior end of body. Oral sucker 300/a in diameter. Excretoryvesicle tubelike, curving between the testes, and extending anteriorlyto the vicinity of the ovary. Pharynx small, ITjH long, the widthbeing almost equal to the length, and situated about 10/x from theoral sucker. Esophagus short and narrow; intestinal ceca simpleand extending to the posterior end of the body. Genital pore 100/* to800ju, caudad of the oral sucker ; seminal vesicle long, slender, and withmuscular walls. Testes globular, 300/t in diameter, situated diago-nally in the posterior third of the body. Ovary globular, 150/a indiameter, situated near the equator of the body ; receptaculum seminis
ART. 13 TEEMATODE PARASITES OF MARINE MAMMALS PRICE 41
large and pedunculated; Laurer's canal present. Vitellaria andMehlis's gland not yet developed. The uterus passes posteriorlybetween the testes and then extends anteriorly to the genital pore.Host.?DelfMnu^ delphis.Location.?Intestine.Distribution.?Europe.Remarks.?The specimens upon which Poirier (1886) based thedescription of the foregoing species were still encysted, the cystsbeing free in the intestine; they were described as spherical andmeasured 1 mm in diameter. Looss (1899) referred this species tothe genus Asfla, which he later (1900) renamed Astiotrema.Jiigerskiold (1908) pointed out that this form was similar to thespecies G. laeteum^ which he had previously described from speci-mens encj^sted in the brain of Cottus scorpius Bloch. Odhner(1911) was of the opinion that this species was closely related toG. lacteum^ and Bittner and Sprehn (1928) actually placed it in thegenus Galactosommii by making the combination of the generic withthe specific name.It seems extremely doubtful whether G. ennacewin is a parasite ofDel'phinus in view of the fact that all other members of the genus areparasites of birds. This was pointed out by Jagerskiold (1908, p.317), who states: " Wahrscheinlich ist der Delphin nicht der wirk-liche Wirt, sondern die von Poirier gefunden Kapseln sind mit ir-gendwelchen Fische in den Delphin hineingekommen. Der wirk-liche Wirt, falls hier wirklich ein Galatosomum vorleight, ist da-gegen wahrscheinlich ein fischfressender Meeresvogel."
Family PARAMPHISTOMATIDAE Fischoeder, 1901
Family diagnosis.?Medium-sized to large trematodes, with orwithout ventral jDOuch. Cuticle without spines. Oral sucker termi-nal or in some cases retracted into the body; with or without dorsalpocketlike evaginations ; acetabulum at posterior end of body.Pharynx absent; intestinal ceca spacious. Excretory vesicle saclike,opening dorsally a short distance from posterior end of body.Lymph system present. Genital opening ventral, in anterior partof bod}^, with or without genital sucker; cirrus pouch present orabsent; testes relatively large, frequently lobed, usually cephaladof ovary. Vitellaria usually well developed. Uterus dorsad oftestes. Eggs without filaments. Parasites of mammals, birds, fishes,amphibians, and reptiles.Type geniis.?Paramphistomum Fischoeder, 1901.
42 PROCEEDINGS OF THE NATIONAL MUSEUM vol.81Genus CHIORCHIS Fischoeder, 1901Generic diagnosis.?Paramphistomatidae : Body ovoid, convexdorsally and flat to slightly concave ventrally; anterior end some-what attenuated, posterior end rounded ; ventral pouch absent. Oralsucker with paired evaginations, usually retracted into an oral canalby the action of strong muscles attached to the oral sucker; aceta-bulum circular, near posterior end of body. Esophagus relativelyshort, thick walled ; intestinal ceca terminate at middle of acetabularzone. Excretory vesicle large, cephalad of acetabulum. Genitalpore at intestinal bifurcation, surrounded by a poorly developedgenital sucker; cirrus pouch absent; seminal vesicle much coiled.Testes X shaped, tandem in position. Ovary posttesticular ; Mehlis'sgland dorsad of ovary; Laurer's canal opens dorsally in front ofexcretory pore. Vitellaria extracecal, except near tips of ceca, wherea few follicles may be intercecal, and extending from a short distancecephalad of intestinal bifurcation to level of anterior margin ofacetabulum. Uterus slightly sinuous, in median line dorsad of testes.Eggs large, thin shelled. Parasites of Sirenia.Type species.?CMorchis fdbaceus (Diesing, 1838) Fischoeder,1901. CHIORCHIS FABACEUS (Diesing, 1838) Fischoeder. 1901Plate 10, Figures 41-45Synonym..?A7iiphistomum fahacev/tn Diesing, 1838, p. 189 ; Schiz-amphistonia nianati Sokoloff and Caballero, 1932, pp. 163-167.Description:?CMorchis: The body is ovoid in outline, 9 mm to11 mm long by 5 mm wide, ventral surface flat or slightly concave,dorsal surface strongly convex. The oral opening is slightly sub-terminal and is followed by an oral canal, the length of whichdepends upon the degree of retraction of the oral sucker. The oralsucker is about 465/>(, to 800/i in diameter, strongly muscular, and pro-vided with two muscular dorsal pouches. The oral sucker is usuallyretracted so that its position is about midway between the intestinalbifurcation and the anterior end of the body, the retraction beingdue to contraction of about 30 muscular bands inserted into thewall of the oral sucker, which radiate posteriorly and attach to thebody wall. The acetabulum is ventral, 1.3 mm to 1.8 mm in diam-eter, situated near the posterior end of the body. The esophagusis relatively long and thick walled, its posterior portion being thick-ened to form a bulblike structure. The intestinal ceca are spaciousand thick walled and extend to about the center of the acetabularzone. The excretory pore is situated in the mid-dorsal line slightlycephalad of the level of the ovary; the excretory vesicle is pouchshaped and relatively thick walled. The lymph system consists oftwo longitudinal vessels which lie dorsally and medially of the
ART. 13 TREMATODE PARASITES OF MARINE MAMMALS?PRICE 43
intestinal ceca and two longitudinal vessels which lie ventrally ofthe ceca. The dorsal vessels extend the full length of the body andterminate in a number of short, dilated branches. Each vessel givesoff about 15 primary branches, which extend lateroventrally andbifurcate to form secondary branches. The secondary branches usu-ally bifurcate to form tertiary branches, each of which terminatesin a bulbous swelling near the ventral surface of the body. A num-ber of ventral branches are also given off from the main dorsalcanals, but their number and course can not be determined with cer-tainty. The ventral longitudinal canals extend the full length ofthe intestinal ceca and give off a number of branches, both mediallyand laterally, which branch again and again and finally terminatein large bulbous swellings; these branches and swellings cover theentire ventral surface of the body beneath the dermomuscular layer.The genital pore is situated immediately caudad of the intestinalbifurcation and is surrounded by a weakly muscular genital sucker.In immature specimens the suckerlike arrangement of the muscularfibers is not so distinct as in more mature specimens. The genitalopening communicates with a genital sinus into which projects theprominent genital papilla. The greatly convoluted seminal vesiclelies free in the parenchyma dorsad of the genital pore. The proxi-mal portion of the ejaculatory duct is inclosed in a muscular saclikestructure, but distally this sac is very feebly developed or absent.The ejaculatory duct unites with the vagina or metraterm to forma hermaphroditic canal, which opens into the genital sinus at thesummit of the genital papilla. The testes are X shaped in maturespecimens and almost spherical in immature specimens; they aresituated in the median field and are tandem in position. The ovaryis almost spherical, about 30/^, in diameter, median in position, andsituated about midway between the posterior testis and acetabulum.Mehlis's gland well developed, dorsad of ovary. Laurer's canal isvery slender and opens in the mid-dorsal line a short distancecephalad of the excretory j^ore. The vitellaria are extracecal, exceptfor a few follicles, whicli are distributed intercecally immediatelyin front of the acetabulum, and extend anteriorly beyond the intes-tinal bifurcation. The uterus extends anteriorly in the median line,dorsal to testes, to the level of the cephalic margin of the anteriortestis, and then turns ventrally and terminates in a weakly muscularvagina or metraterm. The proximal portion of the uterus may befilled with spermatozoa, and constitutes a receptaculum seminisuterinum. The eggs are oval, about 150;a long by 90/x wide, and thinshelled.Hosts.?Manatus exunguis Natterer ( = ? Trlchechus inunguisPelzeln), Trlchechus latirostrls (syn. Manatus latirostris) , and T,senegalensis.
44 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 81
Location.?Large intestine.Distribution.?North America (Philadelphia Zoological Garden;National Zoological Park, Washington, D. C. ; Mexico), SouthAmerica (Brazil), and Africa (Belgian Congo).Remarks.?The above description is based largely upon specimens(U.S.N.M. Helm. Coll. No. 24716), collected at Washington, D. C,July 16, 1921, by Dr. E. A. Chapin, these flukes being well pre-served and fully mature. Other specimens available for comparisonconsisted of three lots as follows: U.S.N.M. Helm. Coll. No. 5775,from the Leidy collection, probably a part of the specimens describedby Leidy (1891) ; U.S.N.M. Helm. Coll. No. 18425, collected atWashington, D. C, November 14, 1916, by Dr. L. T. Giltner; andU.S.N.M. Helm. Coll. No. 8416, labeled " F. 1063, ChiorcMs fabaceusDies., stomach and intestine, Manatus senegalensis, Banana, Aug.1915." The last-named material is from the MacCallum collectionand represents a part of the lot of specimens described by Stunkard(1930).All the specimens examined from these different collections agreein general with the description given for ChiorcMs fabaceus byFischoeder (1903). The majority of the specimens, however, wereimmature and smaller than the ones upon which the above descrip-tion is based, and considerable variation was found to exist, espe-cially as regards the shape of the testes. The Leidy specimenswere the most immature, and in these the testes varied in shape fromspherical to very slightly lobed, none showing the typical X-shapedtestes characteristic of the species. They agree, however, in all otherrespects and must be regarded as the same species as those of theother lots which correspond more closely to the description givenby Fischoeder.
Family NOTOCOTYLIDAE Liihe, 1909Family diagnosis.?Small to medium-sized monostomes; bodyusually elongate, tapering anteriorly and rounded posteriorly.Ventral surface usually concave, with or without longitudinal rowsof glands or ridges. Oral sucker terminal; pharynx absent; esoph-agus short; intestinal ceca slender, usually provided with shortdiverticula. Excretory pore dorsal, near posterior end of body; ex-cretory vesicle with short stem and long branches, which unite nearanterior end of body. Genital pore median and situated in anteriorpart of body, except in Nudacotyle where it is lateral and in pos-terior part of body; cirrus pouch elongate; testes postequatorial,in same transverse plane, usually extracecal. Ovary between testes
;
Mehlis's gland complex preovarial; vitellaria lateral, pretesticular
ART. 13 TREMATODE PARASITES OP MARINE MAMMALS PRICE 45
uterine coils transverse, regular, pretesticular. Eggs sn;iall andprovided with a long filament at each pole. Parasites of birds andmammals.Type genus.?Notocotylus Diesing, 1839.
Subfamily Ogmogasterinae Kossack, 1911Subfamily diagnosis.?Notocotylidae : Ventral surface of body pro-vided with longitudinal ridges or rugae. Uterine coils extendanteriorly beyond base of cirrus pouch. Parasites of cetaceans andpinnipeds.Type genus.?Ogmogaster Jagerskiold, 1891.
Genus OGMOGASTER Jagerskiold, 1891Generic diagnosis.?Ogmogasterinae: Body oval, flattened, andleaflike, margins fluted ; ventral surface provided with longitudinalribs or rugae. Oral sucker terminal; esophagus short; intestinalceca slender and extending to posterior end of body. Genital aper-ture median, a short distance caudad of oral sucker; cirrus andvagina open into a short genital sinus. Cirrus pouch long, situatedin median line and extending posteriorly to near equator of body;testes deeply lobed, situated in the same transverse plane in theposterior fourth of body. Ovary lobed, median in position and sit-uated at the level of the posterior margin of testes; Mehlis's glandmedian, preovarial; Laurer's canal present; vitellaria lateral, con-sisting of isolated follicles and extending from anterior margin oftestes to level of the base of the cirrus pouch; uterus greatly con-voluted, the coils extending laterally beyond intestinal ceca. Para-sites of cetaceans and pinnipeds.Type species.?Ogmogaster plicatus (Creplin, 1829) Jagerskiold,1891. OGMOGASTER PLICATUS (Creplin, 1829) Jagerskiold, 1891
Plate 12, Figure 52Synonym.?Monostomum plicatum Creplin, 1829, pp. 878-880.Desa^iption.?Ogmogaster: Body oval, 6 mm to 14 mm long by anaverage width of 4 mm, flat and leaflike; the margins of the bodyhave a fluted or pleated appearance, and the ventral surface is pro-vided with 15 to 17 longitudinal rugae. Oral sucker terminal, 500/xin diameter, according to Leiper and Atkinson (1915) ; esophagusshort; intestinal ceca sinuous and terminating near the posteriorend of the body. Excretory pore dorsal, about 700/* from theposterior end of body, according to Jagerskiold; excretory vesicle
46 PROCEEDINGS OF THE NATIONAL MUSEUM vol.81Y shaped, situated ventral to ovary and testes. The limbs of thevesicle extend anteriorly and unite at the level of the intestinal bi-furcation at which point two branches are given off, one on each side,which extend posteriorly in the lateral fields, then turn and i^assanteriorly to the level of the oral sucker, where they again turn andpass backward and terminate near the posterior end of the body.Long branches are given off here and there along the course of theexcretory ducts. Genital aperture median, situated a short distancecaudad of the oral sucker; the male and female genital pores aresituated side by side at the base of a short genital sinus. Cirruspouch cylindrical, 3 mm long by 300ju, wide, containing a seminal
vesicle, 1.4 mm long by 200/a wide, and a long convoluted ejaculatoryduct. The ejaculatory duct is lined with a membrane, which isclosely beset with small papillalike projections ; it may be protrudedas a slender cirrus the length of which may equal the length of thebody, according to Creplin (1829). Testes deeply lobed, 1 mmlong by 1.1 mm wide, situated in the same transverse plane in theposterior fourth of the body ; vasa efferentia short, uniting to form arelatively wide vas deferens which extends anteriorly in the medianline to the base of the cirrus pouch ; it then passes to the right of thecirrus pouch where it makes several loops, four to five accordingto Jiigerskiold, and then passes backward and enters the base of thecirrus pouch. Ovary deeply lobed, 500/a long by 1 mm wide, situatedbetween the testes ; Mehlis's gland preovarial ; Laurer's canal present,Vitellaria lateral, composed of 12 to 16 isolated follicles on each side,lying ventral to ceca, and extending from the anterior margin of thetestes to about the level of the base of the cirrus pouch. Uterusslender and greatly convoluted, extending laterally beyond the cecaand from the testes anteriorly to about one-fourth of the bodylength from the anterior end. Vagina muscular and lined withspines which are about 35/x long. Eggs elongate oval in shape, 25/*long, and provided with a long filament at each pole.Hosts.?Cetacea {Balacnoptera horealls^ B. musculus = B. physa-libs, B. acutorostrata= Balaeiia rostrata) ; Pinnipedia - {Leptonycho-tes weddellii., Lohodon carcinophaga)
.
Location.?Intestine.Distribution.?Europe (Norway) ; Antarctic waters (vicinity ofCape Evans).Remarks.-?The above description is taken largely from an exhaus-tive description of this species given by Jiigerskiold (1891).
2 Johnston (1031) regards the form from pinnipeds as a species distinct from Ojmogas-ter plicatus. The new species, for which he proposes the name 0. antarcticus, differsfrom O. plicatus in " its smaller size, diflferent body proportions, presence of only 13rugae, more restricted vitelline zone, and relatively smaller cirrus sac."
ART. 13 TEEMATODE PARASITES OF MARINE MAMMALS PRICE 47Family RHABDIOPOEIDAE Poche, 1926Family diagnosis.?Body elongated, rounded at each end, convexdorsally and concave ventrally. Cuticle of ventral surface armedwith large, curved, hooklike spines. Posterior end of body providedwith large cavity opening dorsally, containing a number of probos-cislike, protrusible structures. Excretory pore situated in the floorof the proboscid cavity; excretory vesicle with four anteriorly di-rected limbs, two lateral and two median ; the lateral pair of branchesunite near the intestinal bifurcation and possess short lateral diverti-cula; the median branches are intercecal and terminate blindly nearintestinal bifurcation. Oral sucker subterminal; esophagus slender;intestinal ceca pass between testes and unite at posterior end of body.Genital aperture situated at side of oral sucker; cirrus pouch longand slender, containing a portion of seminal vesicle. Testes extra-cecal, situated in the same transverse plane near the posterior end ofbody. Ovary between testes; seminal receptacle and Laurer's canalabsent. Vitellaria extracecal and posttesticular. Uterus long andslender, with numerous transverse loops extending beyond laterallimits of the intestinal ceca; vagina as long as cirrus pouch. Eggssmall, provided with a long filament at each pole. Parasitic inSirenia.Tyfe genus.?Rhahdlopoeus S. J. Johnston, 1913.
Genus RHABDIOPOEUS S. J. Johnston, 1913Generic diagnosis.?Rhabdiopoeidae : Characters of the family.Type species.?Rhdbdlopoeus taylovi S. J. Johnston, 1913.RHABDIOPOEUS TAYLORI S. J. Johnston. 1913Plate 12, Figure 51
Dexcriptioh.?Rliabdiopoeus : Body elongated and lancelike, 22 mmlong by 5 mm wdde; thinner anteriorly than posteriorly; dorsalsurface convex; ventral surface flat or slightly concave anteriorlyand more deeply concave i)osteriorly. The ventral surface is coveredwith large, recurved, hooklike spines, 107/x long by 64|i, wdde at theirbases; they are set in a thick cuticula. Oral sucker almost circular,733^ in diameter, subterminal and directed ventrally; esophagusmoderately long; intestinal ceca slender, united at posterior end ofbody; small lateral ceca are present along the posttesticular por-tions, but absent elsewhere along their course. A large cavity,opening dorsally, is present near the posterior end of the body;this cavity communicates with nine tunnel-like tubular spaces,three of which are anterior and the remaining six are arranged in
48 PROCEEDINGS OP THE NATIONAL MUSEUM vol. 81two lateral groups of three, each containing a protrusible fingerlikeproboscis. The excretory pore is situated in the floor of theproboscid chamber; excretory vesicle large and branched, each ofthe branches becoming narrowed into 1 of the 4 chief trunk vessels,2 of the vessels being median and 2 lateral. The lateral vessels ex-tend anteriorly, lateral of testes, and unite immediately anterior andventral of the intestinal bifurcation; they are provided with shortlateral branches along their course from the level of the anteriormargin of the testes to the level of the intestinal bifurcation. Thetwo median vessels pass between the testes and extend anteriorly inthe intercecal field to near the intestinal bifurcation where theyterminate blindly. Genital aperture near right margin of oralsucker. Cirrus pouch long and slender, its posterior end lying inthe median line about one-fourth of the body length from the an-terior end; it contains a small portion of the seminal vesicle, anejaculatory duct surrounded by prostate cells, and a muscular cirrus.Testes deeply lobed, about 2 mm long by 940ju, wide, situated in thesame transverse plane near the junction of the third and last bodyfourths; they are so situated that their long axes are oblique to thelong axis of the body, their bases being separated by the intestinalceca. The vas deferens is expanded to form a relatively wide semi-nal vesicle which extends anteriorly in a more or less tortuous coursefrom the level of the anterior margin of the testes to the cirruspouch. Ovary oval, 890/* long by 5T0/A wide; Mehlis's gland 820/along by 490/A wide, situated to the right of ovary; both ovary andMehlis's gland lie in the angle formed by the testes. There is noseminal receptacle or Laurer's canal. Vitellaria extracecal and post-testicular in position, and consisting of grapelike groups of ovalfollicles, each lateral mass being composed of about 50 groups.Uterus long and slender, composed of numerous, closely packedtransverse loops which extend laterally beyond the limits of the cecaas far as the lateral branches of the excretory system. Eggs 26/xlong by 15/jt wide, thin shelled, and with a long filament at each pole.Host.?EaUcore dugong.Location.?Intestine.Distribution.?Australia (coast of Queensland).Family OPISTHOTREMATIDAE Poche, 1926Family diagnosis.?Body spoon shaped; ventral surface spiny.Esophagus moderately long, slender; intestinal ceca without diver-ticula. Excretory pore dorsal; excretory vesicle short and with lat-eral branches. Genital openings median and situated almost at ex-treme posterior end of body. Cirrus pouch long and slender, median,containing a strongly convoluted seminal vesicle and a protrusible
ART. 13 TREMATODE PARASITES OF MARINE MAMMALS PRICE 49
cirrus; testes intercecal or extracecal, situated in the same transverseplane in the posterior part of body. Ovary either dextral or sinistralin position, pretesticular ; Mehlis's gland postovarial ; seminal recep-tacle present; Laurer's canal present or absent; vitellaria weaklydeveloped, intercecal, pretesticular; uterus long and slender, occupy-ing the greater portion of the central part of body, usually confinedto the intercecal field. Eggs with long polar filaments. Parasitesof respiratory passages of Sirenia.Type genus.?O-pisthotrema Fischer, 1883.
KEY TO THE GENERA OF OPISTHOTREMATIDAE
1. Testes extracecal Opisthotrema (p. 49).Testes intercecal Pulmonicola (p. 55).
Genus OPISTHOTREMA Fischer, 1883Synonym.?Cochleotrema Travassos and Vogelsang, 1931.Generic diagnosis.?Opisthotrematidae : Body oval to piriform inshape, flattened dorsoventrally, dorsum strongly arched and venterstrongly concave ; margin of body may or may not be provided witha muscular rim. Cuticle of ventral surface spiny. Oral sucker ven-tral, situated a short distance from the anterior margin of body;pharynx absent; esophagus slender and of medium length; intes-tinal ceca more or less sinuous and extending into the posterior fourthof body. Excretory pore dorsal and somewhat removed from theposterior margin; excretory vesicle tubular and provided with lat-eral branches. Genital pores at posterior end of body ; cirrus pouchlong and slender, containing a slender, convoluted seminal vesicleand a long, slender, protrusible cirrus; testes lobed, situated extra-cecally in the same transverse plane in the posterior part of body.Ovary lobed, to the right or left of the median line anterior to andseparated from the testis on the corresponding side by the intestinalcecum, which passes between them; Mehlis's gland well developed;seminal receptacle present; Laurer's canal present or absent; vitel-laria intercecal and consist of a grapelike mass of follicles on eachside of median line, or of a mass of irregular follicles in the medianfield. Uterus long, slender, and convoluted, occupying the greaterpart of the central portion of the body ; vagina long, slender, slightlysinuous, and provided with relatively strong muscular walls. Eggsoval and provided with a slender filament at each pole. Parasitesof the respiratory tract and eustachian tubes of Sirenia.Type species.?Opisthotrema cochleare Fischer, 1883 {=Mono-stomum datjonis Leuckart, 1874)
.
Remarks.?The foregoing diagnosis is based on the characterscommon for the two species which are described later. The writer118893?32 4
50 PROCEEDINGS OF THE NATIONAL MUSEUM vol.81
recognizes the genus Pulmonicola^ which was proposed by Poche(1926), as distinct from the genus Opisthotrema on the ground thatthe intercecal position of the testes in Pulmonicola puhnoTialis is toogreat a difference to be regarded as being a character of only specificvalue.
KEY TO THE SPECIES OF THE GENUS OPISTHOTREMA
1. Body piriform in outline ; distinct muscular rim absent ; in-testinal ceca relatively wide, their blind ends divei-ging__ dujonis (p. 50).Body oviil in outline : di.stinet muscular rim present ; intestinalceca slendei", their blind ends converging cochleotrema (p. 52).OPISTHOTREMA DUJONIS (Lcuckart, 1874), new combination
Plate 11, Figure 46
Synonyms.?Monostommii dujonis Leuckart, 1874, p. 419; Opis-thotrema cochleare Fischer, 1883, pp. 1-42.Desci^iption.?Opisthotrema: Body piriform in outline, 9 to 11 mmlong by 5 mm wide; the dorsal surface is strongly convex andthe ventral surface concave, which gives the body the appear-ance of the bowl of a spoon. Cuticle spiny on ventral surface,according to Fischer, and also on anterior part of dorsal sur-face, according to Johnston (1913). Oral sucker ventral, 600ju, longby 850)u, wide, situated about 460/x from the anterior end of body;esophagus slender, about 1 mm long by 80)it wide; intestinal ceca rela-tively wide, slightly sinuous and extending to near the posterior endof the body, their blind ends somewhat distended and diverging.The excretory system consists of two canals, one on each side, unitedby a commissure a short distance caudad of the intestinal bifurcationand again by a similar commissure about midway between the in-testinal bifurcation and the posterior margin of the oral sucker.Each of the canals is provided with lateral branches, which extendto the margin of the body. Excretory vesicle ( ? ) ; excretory pore( ? ) . According to Fischer, the canals terminate near the ends of thececa and probably open separately. The cirrus pouch is cylindrical,2 mm long by 245/x, wide, situated in the median line in the posteriorpart of the body, containing a greatly convoluted seminal vesicle anda slender protrusible cirrus; genital pore ventral, near posteriormargin. Testes lobed, 250/a to 540/>i in diameter, situated in the sametransverse plane and extracecal in position. Ovary small, lobate,situated to the left of the median line and a short distance cephaladof the level of the testes; Mehlis's gland small, caudad of ovary;Laurer's canal jDresent, the proximal part of the canal being expanded
ART. 13 TREMATODE PARASITES OF MARINE MAMMALS PRICE 51
to form a seminal receptacle measurinc: 170/i, long by 57)u, wide.Vitellaria weakly developed, consisting of a few follicles situatedalonsr the vitelline duct at each side of the intercecal field. Uterusslender and occupying the intercecal space in the equatorial third ofthe body ; during its course the uterus describes a number of loopswhich form a treelike pattern. The distal portion of the uterus isexpanded to form an egg reservoir and continues as a slender vagina,which opens beside the male genital aperture. Eggs oval in outline,29^ long by d/x wide, provided with a long, slender filament at eachpole.Host.?Halicore dugong.Location.?Eustachian tubes and esophagus.Distrihutlon.?Philippine Islands ; Australia.Remarks.?This species was named Monostomum dujonis byLeuckart (1874) in a review of Zeiler's (1874) paper " Uber Leu-cochJoridmm paradoxum Cams and die weitere Entwickelung seinerDistomenbrut." In this review, Leuckart, in commenting on thegenital system of L. paradoocum., stated : " Und seine Generations-organe in alien ihren Theilen vollstandig und deutlich erkennenliisst. Die Ausmiindung derselben liegt, wie sonst nur bei wenigenDistomeen?Ref. kennt audi ein Monostomum, das sich ganz ahnlichverhalt, M. dujonis., das Semper in den Eustachischen Rohren desDujung der Philippinen sammelte und dem Ref. freundlichst zurUntersuchung uberlassen hat?an Hinterende des Korpers, dichtneben der des excretorischen Apparates." Fischer's (1883) descrip-tion of Opisthotrema cochleare is based unquestionably upon a studyof the specimens which Leuckart referred to, since he stated in theintroduction of his paper : " Besagte Form wurcle von Herrn Prof.Semper in Wiirtzburg auf seiner Expedition nach den Phili})pinengesammelt. Sie entstammt der Paukenhohle von Halicore Dugong."The name given to this worm by Leuckart can not be regarded asa nomen nudum because an important morphological character, thejDosition of the genital pore, is pointed out and in addition are giventhe host and habitat of the species. It is, therefore, the opinion of thewriter that the correct name for this species is OpisthotTema dujonis(Leuckart), since the rules of zoological nomenclature state: "Art.25.?The valid name of a genus or species can be only that nameunder which it was first designated on the condition: a) That thisname was published and accompanied by an indication, or a defi-nition, or a description;" etc. The description given by Leuckartis more in the form of a comparison than an actual description, butit appears sufficient to fix the specific name, since the rules do notstate how much description should be given.
52 PROCEEDINGS OF THE NATIONAL, MUSEUM vol.81OPISTHOTREMA COCHLEOTREMA (Travassos and Vogelsang-, 1931 ?) new combinationPlate 11, Figtjbb 47-49Synonyms.?AmpMstomum fdbaceum Diesing of Leicly, 1891, pp.413-414, in part ; Opisthotrema cochleare Fischer of Stiles and Has-
sall, 1894, p. 253 ; Cochleotrema cochZeotreina Travassos and Vogel-sang, 1931, pp. 143-146.Descriptio7i.?Opisthotrema: The body is oval in outline, 8.5 mmlong by 6.5 mm wide, strongly concave ventrally and convex dorsally
;
the margin of the body is surrounded by a more or less delicatemuscular rim, which is about 250/1 wide. The strongly convexo-con-cave condition of the body suggests that the entire worm attachesitself to the mucous membrane in the manner of a vacuum cup, themuscular rim serving as a sort of seal. The cuticle of the ventralsurface is covered with scalelike spines, Yyu, long by 4/* wide, arrangedin irregular alternating rows. The excretory pore is situated dor-sally, about 930/1, from the posterior end of the body, the aperturebeing surrounded by a muscular sphincter. The excretory vesicleis tubular, about 1 mm long, and is provided with two principalbranches on each side ; the first branch occurs at the level of the blindends of the ceca and the other at the anterior end of the vesicle ; twosmaller branches are given off on each side a short distance caudadof the anterior branches. The remainder of the excretory systemcould not be worked out in detail, but judged from sections the courseof the principal branches is probably similar to that described byFischer for O. cochleare. The oral sucker is transversely oval, 1 mmlong by 1.3 mm wide, strongly muscular, and situated ventrally about465/A from the anterior margin of the body ; it is deeply imbedded inthe parenchyma and projects only slightly beyond the ventral sur-face. The esophagus is slender and about 465/x long ; the intestinalceca are slender and serpentine, 155/x wide, the blind ends convergingtoward the median line. Cirrus pouch slender, about 3 mm longby 155/x wide at the level of the testes ; the walls are moderately thick
? This species was described by the writer as new in the present paper, but while themanuscript was awaiting publication a description appeared by Travassos and Vogelsang(1931) of a species from Trlchcchns maiuttus which appears to be the same form. Thedescription given herein is from the writer's specimens. Some slight differences exist be-tween this and the description given by Travassos and Vogelsang, but these differencesappear to be those of interpretation. Travassos and Vogelsang note that in their speci-mens the ventral surface is covered with small papillae similar to those occurring on theventral surface of Gastrodiscus, and that Laurer's canal is present. In the specimens atthe writer's disposal the ventral surface is covered with small triangular, scalelike spines(pi. 11, fig. 49) and no Laurer's canal is present. The absence of Laurer's canal is shownin pi. 11, fig. 48, which is a reconstruction from serial sections.The writer does not agree with Travassos and Vogelsang as to the necessity of creatinga new genus for this species, since the differences between this species and the type of thegenus are too slight to warrant such action.
ART. 13 TREMATODE PARASITES OF MARINE MAMMALS PRICE 53
and muscular, both circular and longitudinal fibers being present.The seminal vesicle is slender and greatly convoluted, and occupiesthe basal third of the cirrus pouch. A definite pars prostaticaappears to be absent although some prostate cells are present alongthe distal, less convoluted portion of the seminal vesicle. The cirrusis slender, about 2 mm long, unarmed and protrusible. The genitalaperture is ventral and situated at the posterior end of the bodyat the junction of the muscular rim with the body proper. Thetestes are deeply lobed, about 850^ by 620(0,, extracecal and oppositeeach other in the posterior third of the bod3^ The ovary is irregularin outline but not deeply lobed, about 435/x in diameter, situatedto the right of the median line, intercecal and pretesticular. Mehlis'sgland is composed of large piriform cells and is situated medianand slightly dorsal to the ovary. The seminal receptacle is mus-cular, more or less oblong, about 697fi long by 155/x wide, and situateddorsal to Mehlis's gland. There is no Laurer's canal. The vitel-laria consist of relatively few, large, irregularly shaped folliclessituated in the intercecal space between the equator of the bodyand the testes ; the follicles are not divided into two separate groupsas in 0. dujonis^ but form a single, more or less grapelike mass. Theuterus is greatly convoluted, forming a wreathlike mass of loopswhich occupies the greater part of the central portion of the body;the loops extend slightly beyond the ceca laterally, but do not extendanteriorly beyond the intestinal bifurcation. The terminal portionof the uterus is continued as a muscular walled, slightly sinuousvagina or metraterm, which runs dorsal to the cirrus pouch and opensbeside the male genital aperture. The eggs are oval in shape, 18/ilong by 11/A wide, light straw colored, and provided with a long,slender filament at each pole.Hosts.?Trichechus manatus and T. latirostrls {^Mavafus Jati-rostris.)Location.?Nasal passages and stomach.Distrihution.?North America (United States?Philadelphia, Pa,),and Tropical America.SpeeimeTis.?V.S.'^M. Helm. Coll. No. 1732.Remarks.?In a note presented at a meeting of the PhiladelphiaAcademy of Natural Sciences, Leidy (1891) reported the occurrenceof Amphistonium fabacev/m Diesing in a sea cow, Manatus latirostris,which died in the zoological garden. These specimens had been col-lected from the large intestine and submitted by Dr. H. C. Chapman.In the same note he said : " Numerous specimens, many of larger
size, up to 11 mm long by 9 mm broad, were obtained from thenasal passages of another sea cow, and were presented to theAcademy by Jacob Geismar."
54 PROCEEDINGS OP THE NATIONAL MUSEUM vol.81
Stiles and Hassall (1894) examined the trematodes of the Leidycollection and redetermined the specimens reported from the nasalpassages as Opisthotrenna cochleare Fischer. A part of the ma-terial, two specimens, was retained in the United States NationalMuseum collection, and it is upon these that the foregoing descrip-tion of Opisthotrenia cochleotrema is based.These specimens are sexually mature, but somewhat smaller thanthose reported by Leidy. One of them was stained and mountedwhole; the other was stained and sectioned. The description givenabove is a composite one, the details being obtained from the sec-tioned specimen.Opisthotre7na cochleotrema differs in a number of respects from0. dujonis (Leuckart) { = 0. cochleare Fischer), so that there ap-pears to be no doubt that they are distinct species. O. cochleotremais provided with a muscular rim similar to that described for Pid-monicola puhnonalis (von Linstow) ; the intestinal ceca are slenderand uniform in diameter, serpentine, and their blind ends converge
;
Laurer's canal is absent; and the terminal part of the uterus is notexpanded to form an egg reservoir. In O. dujonis the body is notprovided with a muscular rim ; the intestinal ceca are not of uniformwidth, only slightly sinuous, and their blind ends diverge; Laurer'scanal is present; and the distal part of the uterus is expanded toform an egg reservoir. Other differences may be easily seen by com-paring the descriptions and figures of the two species.Fischer's (1883) figure of Opisthotrema cochleare { = 0. dujonis)leaves the impression that the principal branches of the excretorysystem open separately and in this connection he states " indemwahrscheinlich nicht eine, sondern zwei Miindungsstellen vorhandendiirften, die der ventralen Seite angehoren und jederseits unterhalbder Darmschenkel liegen." Poche (1926) doubts the presence of twoexcretory pores in this species, but Fuhrmann (1929) believes thattwo openings should be present and has indicated in the figure of O.cochleare (" nach Fischer '') just where these openings should belocated. The study of serial sections of O. cochleotrema has shownthat the position of the excretory pore in this species is essentiallythe same as that in members of the famil}'^ Notocotylidae * andPronocephalidae. In view of this fact it appears reasonable to as-sume that the position of the excretory pore in O. dujonis will be
* In Pochc's (1926) classification of the trematodes, the monostomes belonging to thefamilies Notocotylidae, Pronocephalidae, Opisthotrematidae, and Rhabdiopoeidae are placedIn the super-superfamily Notocotylida. Since groups of the rank of super-superfamilyhave not been recognized by helminthologists and are unnecessary at present, the writerproposes at this time the new superfamUy Notocotyloidea, to replace the super-superfamiljNotocotylida Poche.
ART. 13 TEEMATODE PARASITES OF MARINE MAMMALS PRICE 55found to be the same as that in O. cochleotreina^ as it would beunusual to find such marked differences in two species so closelyrelated. Genus PULMONICOLA Poche, 1926
Generio diagnosis.?Opisthotrematidae : Entire margin of body,except space occupied by oral sucker, provided with a muscular rim.Cuticle without spines. Oral sucker ventral, near anterior marginof body; intestinal ceca straight, uniform in diameter. Excretorysystem ( ?), probably similar to that in Opisthotrema. Testes entire,intercecal, situated near ends of ceca. Ovary median. Other charac-ters as in OpisthotT^nia.Type species.?Pulmonicola puhno7ialis (von Linstow, 1904)Poche, 1926.
PULMONICOLA PULMONALIS (von Linstow, 1904) Poche, 1926
PLATE! 11, FiGUKB 50Synonyms.?Opisthotrenia pulmonale von Linstow, 1904, pp.67^680.Desc7'iption.?Pulmonicola': Body oval in outline, 5.13 mm longby 3.95 mm wide; margin of body, except for the portion occupiedby the oral sucker, modified to form a muscular rim. Oral suckersituated on ventral surface, 330/i, from the anterior margin ; esopha-gus very slender, 180/a long by 28/[a wide; intestinal ceca uniform indiameter and extending into the posterior fourth of body. Cirruspouch slender, spirally coiled distally, and containing a greatly con-voluted seminal vesicle. Testes oval, entire, 390/i, long by 280yii wide,situated intercecally in the same transverse plane. Ovary small,median in position, situated a short distance caudad of the equatorof body; Mehlis's gland smaller than ovary and situated immedi-ately caudad of it ; receptaculum seminis present ; Laurer's canal pres-ent. Vitellaria in intercecal field and consisting of a few isolatedfollicles situated between the ovary and testes. Uterus slender, inter-cecal, occupying the equatorial third of body in immature specimens,but in mature specimens more convoluted and occupying the secondand third fourths of the b(jdy ; vagina dorsal to cirrus pouch; genitalpore at posterior end of body. Eggs 16|u. long by 9jii wide, yellow incolor, and provided with a long filament at each pole.HostH.?Halicore australe^ U. diigong.Location.?Lung.Distrihution.?Australia (Torres Straits).
56 PROCEEDINGS OF THE NATIONAL MUSEUM vol.81SPECIES OF UNCERTAIN POSITIONDISTOMA ANDERSONI Cobbold, 1876Plate 12, Figuke 53Description.?" Body oblong, smooth externally, uniform in thick-ness, six times as long as broad; head with lateral projections; ven-tral sucker large and prominent ; neck much constricted ; tail evenlyrounded off, blunt. Length 1/8", breadth about 1/50". The testesare globular and placed high up in the middle line of the body.The small lobed gland immediately above them is probably the ovary.The clear narrow line extending from the border of the lower testisto the end of the tail seems to mark the limit of the vitellogene organson either side below. These glands in all likelihood extend upwardsto the neck, being apparently very largely developed in this species."Host.?Platanista gangetica.Location.?Small intestine.Distribution.?Asia (India)
.
Remarks.?The foregoing description, copied from Cobbold(1876), was based upon a drawing sent to him by Dr. John Anderson,superintendent of the Indian Museum, Calcutta. Odhner (1905)believed that this worm was probably related to Brachycladium{==Campula) , as he stated: " Endlich finde ich sehr wahrscheinlich,dass das sehr ungeniigend bekannte Dist. andersoni Cobb, aus Pla-tanista gangetica sich bei genauerer Untersuchung als eine Brachy-cladium-K.vt entpuppen wird." It is probably unwise to venture anopinion as to the systematic position of this species, but certain fea-tures as shown in the drawing, assuming that the sketch was notmade from a mutilated specimen, suggest that this form might be anechinostome. The anterior end is not unlike the cephalic collar ofthe echinostomes ; the small oral sucker and the large acetabulum, aswell as the relative size and position of the ovary and testes, suggestsuch affinities. On the other hand, what is figured as the oral suckermay be only the oral aperture, and the structure which resembles acephalic collar may in reality be the oral sucker which has been tornloose from the body.DISTOMUM PHILOCHOLUM Creplin, 1845
Description.?None.Host.?DelpJiinus delphis.Location.?Liver.Distribution
.
?Europe.Remarks.?Creplin (1845) gives only the name and habitat ofthis species, and for this reason it must be regarded as a nornennudum.
AUT. 13 TREMATODE PARASITES OF MARINE MAMMALS?PRICE 57AGAMODISTOMUM DELPHINI (Diesing, 1S50). new combinationSyiionyms.?Monostoinum delphini Diesing, 1850, p. 390; Monos-tomum hlainvillei Cobbold, 1860, p. 39; Monostomulutn delphini(Diesing, 1850) Brandes, 1892, p. 510.Description.?The following is the only existing description ofthis worm and is quoted verbatim from de Blainville (1825) :En eulevant la gniisse de cet animal [a whale], on a trouve, euferme dansson epaisseiir, et contenu dans ime sorte de kyste k parois lisses en dedans,mals nou distinctes en dehors, un ver assez singulier, que M. de Blainvillecrut d'abord pouvoir rapporter au genre Monostome, mais qui en differesensiblement, comme on va le voir: il etait repli6 dans sou kyste, et vivant,quoique le dauphin f(it mort depuis cinq ou six jours. Mis dans de I'eau froide,il se contractait dans tous les sens, de maniere a presenter une form extr&me-ment variable, quelquefois globuleuse, d'autres fois ovale-alongee, ctrangl^eau milieu ou nou^e, avec une sorte de queue en arriere ou de tube en avant;sou extremite anterieure souvent attenuee et cylindrique, prfeentait un orificeevident de form circulaire. II en existait aussi un autre a I'extremite pos-terieure, mais beaucoup plus petit, et au milieu d'une sorte d'aureole plusgrise ; eufiu, sur un individu, M. de Blainville a vu, h peu pres a la moitede la longueur et en dessous, une petite masse blanche, ovale, saillante endehors, un peu comme dans les fascioles, ou distomes. Ce ver, d'une couleurblanche mate, etait forme d'une sorte d'enveloppe epaisse de cette couleur,et d'une autre intestinale comme gelatineuse.Host.?Delphinus dalei {^rohiihly= Mesoplodon hiden^).Location.?Encysted in fat.Distribution.?Europe (Havre, France).Remarks.?Diesing (1850) regarded this species as an immaturemonostome, as did Cobbold (1860) and Brandes (1892). The de-scription and habitat, however, do not sustain this assumption. Sofar as known, the true monostomes do not encyst as larvae in thebody of vertebrates, but encyst in water and become attached toherbage or other objects, or remain floating. Certain features ofthis form as given in de Blainville's description, especially the small,oval projection on the ventral surface situated about the middleof the body (a peu pres a la moite de la longeur et en dessous, unepetite masse blanche, ovale, saillante en dehors), suggest that thisworm is the metacercaria of a species of Alaria or that of a relatedgenus, the oval body corresponding to the holdfast organ of theseforms. This species, therefore, has been transferred to the collec-tive distome genus Agaryiodistomum.
58 PROCEEDIlSrGS OF THE NATIONAL MUSEUM VOL. 81LIST OF TREMATODES ARRANGED ACCORDING TO HOSTS'Order CETACEASuborder Mysticeti
Family BALAENIDAEBalaena mysticetus:LecitJiodesmiis goliath (van Ben-eden).FamUy BALAENOPTERIDAEBalaenoptera acutorostrata (syn. B.rostrata)
:
Fasciola hepatica Linnaeus.Lecithodesmus goliath (van Ben-eden).Ogmogaster plieatus (Creplin).Balaenoptera borealis:Lecithodesmus goliath (van Ben-eden).Ogmogaster plicatus (Creplin).Balaenoptera physalus (syn. B. mus-cuius Companyo) :Ogmogaster plica tus (Creplin).
Suborder Odontoceti
Family DELPHINIDAEDelphinus delphis:Campula delphini (Poirier).Campula palliata (Looss).Campula rochehruni (Poirier).Galactosomuni erinaceum (Poir-ier).Distomuni philocholum Creplin.Tursiops truncatus (syn. Delphinustursio)
:
Sifiitli cs'nnn tursionis ( Mf.rchi )
.
Orcinus orca (syn. Orca gladiator):Fasciola hepatica Linnaeus.Orcaella brevirostris:fAmphimcnis laueea (Diesing).Phocaena phocoena (syn. P. com-munis) :Campula oblonga Cobbold.Opisthorchis tenuicollis (Rudol-ph!).Pholeter gastrophilus (Kossack).
Sotalia tucuxi (syn. ? Delphinustacuschi).Amphimerus lancea (Diesing).Delphinapterus leucas:Hadivcnius seymonrl, new species.
Family PLATANISTIDAE
Platanista gangetica:Cyclorchis ca?ipula (Cobbold).DistomM andersoni Cobbold.
Family ZIPHIIDAEMesoplodon bidens (syn. ? Delphinusdalei)
:
Agatnodistomum delphini (Dies-ing).
Order SIRENIA
Family TRICHECHIDAE
Trichechus inunguis (syn. ? Manatusexunguis Natterer) :Chiorchis fabaceus (Diesing).Trichechus latirostris
:
Chiorchis fabaceus (Diesing).Opisthotrema cochleoirema (Tra-vassos and Vogelsang).Trichechus manatus:Opisthotrema cochlcotrema (Tra-vassos and Vogelsang).Trichechus senegalensis:Chiorchis fabaceus (Diesing).
Family HALICORIDAE
Halicore dugong (syn. H. cetacea)
:
Opisthotrema dujonis (Leuckart).Rhabdiopocus taylori S. J. John-ston.Pulmonicola pulmonale (von Lin-stow).Halicore australe:Pulmonicola pulmonale (von Lin-stow).
s The writer is indebted to Dr. Remington Kellogg, assistant curator of the division ofmammals of the United States National Museum, for suggesting the probable identifica-tions of the hosts.
ART. 13 TREMATODE PABASITES OF MARINE MAMMALS PRICE 59
Order PINNIPEDIA
Fair.ily OTARIIDAEZalophus californianus:ApophaUiis zalophi, new species.Zaiophotrcma hepatictim Stuu-kard and Alvey.Stephauoprora denticulata (Ru-dolphi).Family PHOCIDAEPhoca groenlandica:PseudampJUstomum t r u n c a t um(Kiulolphi).Phoca hispida (syii. HaUclioerusfoetidus) :Orthosplanchnus arcticus Odliner.Phocitrema fusiform c Goto andOzaki.Psendamphistomum trunoatuni(Rudolph!).Phoca vitulina :Apophallus donicus (Skrjabin andLindtrop )
.
Cryptocotyle lingua (Creplin).Eohinostoma acanthoides (Ru-dolphi).Pseudamphu'^tomuni t run cat u m(Rudolphi).
Erignathus barbatus (syn. Pliocabarhata) :OpisthorcM^ t e n u icoll i s ( Ru-dolphi )
.
Orthosplanclinii^s arcticus Odhner.Orthosplatwhn us fraterculusOdhner.Halichoerus grypus:Mctorchis albidus (Braun).Opisthorehis t c n u i colli s (Ru-dolphi).Pseudamphistommn t r u neat it, m(Rudolphi).Leptonychotes weddellii:Ogmo'jastcr pUeatus (Creplin).Lobodon carcinophaga:Ogmogaster plicatus (Creplin).
Family ODOBENIDAE
Mirounga angustirostris (syn. Macro-rhiniis angustirostris) :Cryptocotyle lingua (Creplin).Odobenus rosmarus:Odhneriella rossica Skrjabin.Orthosplanchnus frateroulusOdhner.
BIBLIOGRAPHYAfeica, Candido M.1929. On two German Heterophyidae with notes on the variability ofcertain structures. Centralb. fiir Bakt., Parasit. und Infekt., Abt.1, Orig., vol. 114, nos. 1-2, pp. 81-86, figs. 1-4; German summary,p. 86.Babkee, Franklin D.1911. The trematode genus Opisthorchis E. Blanchard, 1895. Studies Zool.Lab. Univ. Nebraska, no. 103, pp. (513) -561, 3 tables, pis. 17-20.(Advance separate from Arch, de Parasitologie, vol. 14, no. 4, pp.513-561, pis. 17-20.) Paris.BeNEDEN, p. J. VAN.1858. Note sur une nouvelle espece de distome, le g(5ant de sa famille.habitant le foie d'une baleine, nommee Distoma ffoliath v. Ben,Bull. Acad. Roy. Sci., Lettres, et Beaux-arts Belgique, ser. 2, vol.5, no. 7, pp. 95-97, 1 pi., figs. 1-5. Brussels.BiTTNER, H., and SpBEHN, C.1928. Trematodes. Saugwlirmer. Biologic der Tiere Deutschlands(Schulze), vol. 27, pt. 5, pp. 1-133, figs. 1-56. Berlin.Blainville, Maeie Henri Ducbotay de.1825. Note sur un c6tac6 4choue au Havre, et sur un ver trouv6 dans sagraisse. Nouveau Bull. Sci. Soc. Philom. Paris, Sept., pp. 139^141.Blanchard, Raphaex.1895. Animaux parasites (preliminary notice). Bull. Soc. Zool. France,vol. 20, nos. 8-9, p. 217.Brandes, G.1892. Revision der Monostomiden. Centralb. fiir Bakt., Parasit, und In-fekt., vol. 12, no. 15, pp. 504^511.Braun, Max.1892. Verzeichniss von Eingeweidewiirmern aus Mecklenburg. Arch. Ver.Freunde Naturg. Mecklenburg, vol. 45, no. 2, pp. 97-117.1893. Die Leberdistomen der Hauskatze {Felis catn^ domesticus und ver-wandte Arten). Centralb. fur Bakt., Parasit. und Infekt., vol. 14,no. 12, pp. 381-392 ; no. 13, pp. 422-428, figs. 1-4.1900. Tiber Campula oblonga Cobb. Centralb. fiir Bakt., Parasit. und In-fekt., Abt. 1, vol. 28, nos. 8-9, pp. 249-254, figs. 1-3.1901a. Zur Kenntuiss der Trematoden der Siiugethiere. Zool. Jahrb. (Abt.Syst), vol. 14, no. 4. pp. 311-348, pis. 19-20, figs. 1-17.1901b. Zur Revision der Trematoden der Vogel. I. Contralb. fiir Bakt,Parasit. und Infekt., Abt. 1, vol. 29, no. 13. pp. 560-568.1901c. Zur Revision der Trematoden der Vogel. II. Centralb. fiir Bakt.,Parasit. und Infekt, Abt. 1, vol. 29, no. 23, pp. 895-897.1902a. Fascioliden der Vogel. Zool. Jahrb. (Abt. Syst.), vol, 16, no. 1, pp.1-162, pis. 1-8, figs. 1-99.1902b. tJber Distoma goliath P. J. Van Ben,, 1858. Centralb. fiir Bakt,Parasit, und Infekt., Abt. 1, Orig,, vol. 32, no, 11, pp, 800-803, 1pl? figs. 1-2.Buttel-Reepen, H.1902. Zur Kentniss der Gruppe des Distomum clavatum, insbesondere desDist. ampullaceum und des Dist. siemersi. Zool. Jahrb. (Abt.Syst,), vol, 17, no. 2, pp. 165-236, pis. 6-10, figs. 1-54,60
AnT. 13 TREMATODE PARASITES OF MARINE MAMMALS PRICE 61
ClURBlA, J.1924. Heterophyides de la faune parasitaire de Rouinauie. Parasitology,
vol. 16, no. 1, pp. 1-21, pis. 1-5, figs. 1-19.CoBBOLD, Thomas Spencer.1858. Observations on Eutozoa, with notices of several new species, includ-ing an account of two experiments in regard to the breeding ofTaenia serrata and T. cucumerUia. Ti-ans. Linn. Soc. Loudon, vol.22, pt. 3, pp. 155-172, pis. 31-33, tigs. 1-85.1860. Synopsis of the Distomidae. Journ. Proc. Linn. Soc. Loudon, Zool.,no. 17, vol. 5, pp. 1-58.1876. Trematode parasites from the dolphins of the Ganges, Platmiistcugangetica and Orcella 'b7-evirost7'is. Jouni. Linn. Soc. London, Zool.,no. 65, vol. 13, pp. 35-46, 1 fig., pi. 10, figs. 1-3.Cooper, A. R.1921. Trematodes and cestodes of the Canadian Arctic Expedition 1913-18.Rep. Canadian Arctic Exped., vol. 9, part GH, pp. (3) -27, pis. 1-2,figs. 1-15. Ottawa.Cbeplin, Fridrich Christian Hbnrich.1825. Observations de entozois, 86 pp., 1 pi., 17 figs. Gryphiswaldiae.1829. Filariae et Monostouii speciem novam in Balaena rostrata repertam.Nova Acta Phys.-Med. Acad. Nat. Curios., vol. 14, no. 2, pp. 871-882,pi. 52, figs. 1-11. Bonn.1845. Nachtrage zu Gurlt's Verzeichuiss der Thiere, bei welchen Entozoeugefunden worden sind. Arch, fiir Naturg., Jahr. 11, vol. 1, pp.325-336.DiESiNG, Karl Moritz.1838. [Abbildungeu neuer Gattungen brasilianischer Binnenwiirmer (En-tozoeu)] (Secretary's abstract). Ber. Versamml. deutsch. Naturf.und Aerzte in Prag, Sept., 1837, p. 189.1889. Neue Gattungen von Binnenwiirmeru nebst einem Nachtrage zurMouographie der Amphistomen. Ann. Wien Mus. Naturg., vol. 2,no. 2, pp. 219-242, pis. 14-20.1850. Systema helruinthum, vol. 1, 679, pp. Vindobouae.1855. Neunzehn Arten von Trematoden. Denkschr. Kais. Akad. Wiss.math.-naturw., vol. 10, pt. 1, pp. 59-70, pis. 1-3. Wien.1858. Revision der Myzhelminthen. Abtheilung: Trematoden. Sitz. Akad.Wiss. Wien, Math.-naturw. CI., vol. 32, no. 23, pp. 207-390, pis.1-2.DiETZ, Eugene.1909a. Die Echinostomiden der Vogel. Zool. Auz., vol. 34, no. 6, pp.183-192.1909b. Idem. Diss. 37 pp., 8 figs. Konigsburg in Preussen.DUJABDIN, F^LIX.1845. Histoire naturelle des helminthes ou vers intestinaux, 654 pp., 12pis. Paris.Ercolani, G. B.1875. Osservazioni elmintologische sulla dimorfobiosi nei nematodi, sullaFilaria immitis e sopra una nuova specie di Distoma dei cani.Mem. Acad. Sci. 1st. Bologna, ser. 3, vol. 5, no. 3, pp. 391-441,1 pi., figs. 1-9.Faust, E. C.1929. Human helminthology. A manual for clinicians, sanitarians andmedical zoologists, 618 pp., figs. 2-297, 1 pi. Philadelphia.
62 PEOCEEDINGS OP THE NATIONAL MUSEUM VOL.81
Fischer, P. M.1883. tJber den Bau von Opisthotrema cochleare nov. genus ; uov. spec. EinBeitrag zur Kenntnis der Trematoden. Diss. 42 pp., 1 pi.1884. Idem. Zeitsclir. fiir wiss. Zool., vol. 40, no. 1, pp. 1-41, pi. 1.FiscHOEDEE, Franz.1901. Die Parampliistomiden der Saiigethiere. Zool. Anz. vol. 24, pp.367-375.1902. Die Paramphistomiden der Saiigetiere. Diss. 59 pp., 4 figs. Konigs-berg.1903. Idem. Zool. Jahrb. (Abt. Syst.), vol. 17, nos. 4^6, pp. 485-660, figsj.A-Q, pis. 20-31, figs. 1-104.Ftthrmann, Otto.1929. Zweite Klasse des Cladus Platyhelminthes: Trematoda. Haudbuch derZoologie (Kiikenthal und Krumbach), vol. 2, Lief. 3, Teil 2, Bogen1-8, pp. 1-128, figs. 1-171. Berlin and Leipzig.Gmexin, Johann Friedrich.1790. Caroli a Linne . . . Systema naturae per regna tria naturae, secun-dum classes ordines, genera, species cum characteribus, difler-entiis, synouymis, locis, vol. 1. Editio decima tertia, aucta, re-formata, cura Jo. Fred. Gmelin, pt. 6 (Vermes), pp. 3021-3910.GoBZE, Johann August Ephraim.1782. Versuch einer Naturgeschichte der Eingeweidewiirmer thierisclierKorper, 471 pp., 44 (35) pis. Blankenburg.Goto, S., and Ozaki, Y.1930. Brief notes on new trematodes III. Jap. Journ. Zool., vol. 3, no. 1,pp. 72-82, figs. 1-7.GURLT, E. F.1831. Lehrbuch der patbologischen Anatomie der Haus-Saiigethiere. Nebsteineu Auhange, welcher die Beschreibung der bei den Haus-Saiige-thieren vorkommenden Eingeweidewiirmer enthalt, vol. 1, 399 pp.Berlin.Jagerskiold. Leonard Axel Krister Eduard.1891. t'ber den Bau des Ogmogaster plieatus (Creplin) {Monostomumplicatum Creplin). K. Svenska Vetensk.-Akad. Handl., new ser.,
vol. 24, pt. 1, no. 7, 32 pp., 2 pis., figs. 1-16. Stockholm.1S96. tJber Monoslomnm lactenm n. sp. Zoologiska Studier. Festskrift Wil-helm Lilljeborg, pp. 165-177, 1 fig., pi. 9, figs. 1-9. Upsala.1908. Kleine Beitriige zur Kenntnis der Vogeltrematoden. Centralb. fiirBakt.. Parasit. und Infekt., Abt. 1, Orig., vol. 48, no. 3, pp. 302^17,figs. 1-7.Johnston, S. J.1913. On some Queensland trematodes, with anatomical observations anddescriptions of new species and genera. Quart. Journ. Micr. Sci.,new ser., vol. 59, no. 3, pp. 361-400, pis. 22-27, figs. 1-42.Johnston, T. Harvey.1931. New trematodes from the subantarctic and antarctic. Austral. Journ.Exper. Biol, and Med. Sci., vol. 8, no. 2, pp. 91-98.KOSSACK, W.1910. Neue Distomen. Centralb. fiir Bakt., Parasit. und Infekt., Abt, 1,Orig.. vol. 56, no. 2, pp. 114-120, figs. 1-i, 1 pi., figs. 1-8.LaRue, George R.1926. A trematode with two ani. Journ. Parasitol., vol. 12, no. 4, pp. 207-209, 1 fig.
ART. 13 TREMATODE PAEASITES OF MARINE MAMMALS PRICE 63
Leidy, Joseph.1891. Notices of Entozoa. Proc. Acad. Nat. Sci. Philadelphia, vol. 42 (ser.3, vol. 20), no. 3, pp. 410-418, Oct.-Dec., 1890.LEIPE31, ROBEKT T.1908. An account of some helminthes contained in Dr. Wenyon's collectionfrom the Sudan. 3d Rep. Wellcome Research Lab., pp. 187-199,figs. 44-50, pis. 21-22. London.Leipee, Robert T.. and Atkinson, E. L.1915. Parasitic worms, with a note on a free living nematode. BritishAntarctic Expedition, 1910. Nat. Hist. Rep., Zool., vol. 2, no. 3,pp. 19-GO, figs. 1-11, pis. 1-5, figs. 1?42. London.Leuckart, Rudolph.1874. Bericht fiber die wissenschaftlichen Leistungen in der Naturgeschichteder niederen Thiere wiihrend der Jahre 1872-1875. Arch, furNaturg., Jahr. 40, vol. 2, pp. 401-505.Linnaeus, Carolus.1758. Systema naturae per regna tria naturae, secundum classes, ordines,genera, species, cum chai'acteribus, differentiis, synonymis, locis.Editio decima, reformata. Vol. 1, 823 pp. Holmiae.Lin STOW, Otto FRiEOEiticn Bern hard, von.1904. Neue Helmiuthen. Centralb. fiir Bakt, Parasit. und Infekt., Abt.1, Orig., vol. 37, no. 5, pp. 678-683, figs. 1-11.LONNBERG, E.1891. Mitteilungen fiber einige Helmiuthen aus dem zool. Museum derUniversitiit zu Kristiania. Biol. Foren. Forhandl., Verb. Biol.Ver. in Stockholm, vol. 3, nos. 4-6, pp. 64-78, pi. 2, figs. 1-9.Looss, Arthur.18S5. Beitrjige zur Kenntnis der Trematoden. Dlstoinum palUatum nov.spec, und DUtomum rcticulafiim nov. spec. Zeitschr. fur wiss. Zool.,vol. 41, no. 3, pp. 390-446, pi. 23, figs. 1-30.1896. Recherches sur lu faune parasitaire de I'Egjpte. Premiere partie.Mem. Inst, i^gypte, vol. 3, pp. 1-252, pis. 1-16, figs. 1-193.1899. Weitere Beitriige zur Kenntnis der Trematoden-Fauna Aegyptens,zugleich Versuch einer naturlicheu Gliederung des Genus DistomumRetzius. Zool. Jahrb. (Abt. Syst.), vol. 12, nos. 5-6, pp. 521-784,figs. 2r-b, pis. 24-32, figs. 1-90.1900. Nachtragliche Bemerkungen zu den Namen der von mir vorgeschla-genen Distomidengattungen. Zool. Anz., vol. 23, pp. 601-608.1901. Natura doceri, ein Erklarung und Begrundung einiger Grundsatze,welehe mich hei meinen " Versuche einer naturlicheu Gliederungdes Genus Distomum Retzius geleitet haben." Centralb. furBakt., Parasit. und Infekt., Abt. 1, vol. 29, no. 5, pp. 191-210.1902. Uber neue und bekiinnte Trematoden aus Seeschildkroten. NebstErorterungen zur Sytematik und Nomenclatur. Zool. Jahrb. (Abt.Syst.), vol. 16, no. 3-6, pp. 411-894, figs. A-B, pis. 21-32, figs. 1-181.1912. rber den Bau einger anscheinend seltner Trematoden-Arten. Zool.Jahrb.. Suppl. 15, vol. 1, Festschr. 60. Geburtst. J. W. Spengel,pp. 323-366. pis. 17-19, figs. 1-22.LuHEi, Max.1899. Zur Kenntnis einiger Distomen. Zool. Anz., vol. 22, pp. 524-539.1908. Zur Systematik und Faunistik der Distomen. 1. Die GattungMctorchis Looss, nebst Bemerkungen fiber die Familie Opisthor-chiidae. Centralb. f. Bakt., Parasit. und Infekt, Abt. 1, Orig., vol.48, no. 4, pp. 428-436, figs. 1-6.
64 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 81
LtJHE, Max?Continued.1909. Parasitische Plattwiirmer. 1: Trematodes. SiisswasserfaunaDeutschlands (Brauer), vol. 17, 217 pp., 188 figs. Jena.MacCallum, G. a.1916. Some new species of parasitic trematodes of marine fishes. Zoopatho-logica, vol. 1, no. 1, pp. 3-38, figs. 1-16.Marchi, Pietko.1873. Sopra una specie nuova di Distomum trouvata nelle intestina delDelphinns tursio. Atti Soc. Ital. Sci. Nat. (1872-73), vol. 15, no.4, p. 304, pi. 5, fig. B. Milan.MOEGAN, D. O.1927. Studies on the family Opisthorchiidae Braun, 1901, with a descrip-tion of a new species of Opisthorchis from a Sarus crane {AntigoneanUgone). Journ. Helminth., vol. 5, no. 2, pp. 89-104, figs. 1-11.MtJHLiNG, Paul.1896. Beitrage zur Kenntnis einiger Trematoden. Centralb., fiir Bakt.,Parasit. und Infekt., Abt 1, vol. 20, no. 16-17, pp. 588-590.1898a. Studien aus Ostpreussens Helminthfauna. Vorlaufige Mittheilung.Zool. Anz. vol. 21, pp. 16-24.1898b. Die Helminthen-Fauna der Wirbeltiere Ostpreussens. Arch, fiirNaturg., Jahr. 64, vol. 1, no. 1, pp. 1-14, pis. 1-14, figs. 1-28.Nazmi, M.1930. A new trematode parasite from the domestic goose. Ann. Mag. Nat.Hist., ser. 10, vol. 6, pp. 377-380, figs. 1-3.NicoLL, William.1909. Studies on the structure and classification of the digenetic trema-todes. Quart. Journ. Micr. Sci., new ser., vol. 53, no. 3, pp. 391-487, pis. 9-10, figs. 1-28.Odhner, Tbodor.1905. Die Trematoden des arktischen Gebietes {In Romer and Schaudinn,Fauna Arctica, vol. 4, pt. 2, pp. 291-372. 4 figs., pis. 2-A. Jena.)1910a. Microlistrum Braun, eine angebliche Distomengattung. Zool. Anz.,
vol. 35, nos. 12-13, pp. 350-356, 1 fig.1910b. tjber Distomen, welche den Excretionsporus als Anus verwendenkonnen. Zool. Anz., vol. 35, nos. 14-15, pp. 432-433.1910c. Nordostafrikanische Trematoden, grosstenteils vom Weissen Nil (vonder Schwedischen Zoologischen Expedition gesammelt), 170 pp..14 figs., 6 pis. (Results of the Swedish Zoological Expedition toEgypt and the White Nile 1901, under the direction of L. A.Jiigerskiold). Uppsala.1911. Zum natiirlichen System der digenen Trematoden. 3 (Ein weitererFall von sekundarem Anus). Zool. Anz., vol. 38, no. 4, pp. 97-117.figs. 1-8.1914. Die Verwandtschaftsbeziehungeu der Trematodengattung Paragon-inius Brn. Zool. Bidrag Uppsala, vol. 3, pp. 231-(246), figs. 1-5.1926. Protofasciola n. g., ein Prototypus der grossen Leberegels. Ark. forZool., vol. 18, no. 3. pt. A, art. 20, pp. 1-7, figs. 1-2. Stockholm.1928. Weitere Trematoden mit Anus. ArW. for Zool., vol. 20, no. 2, pt. B,art. 2, pp. 1-6, figs. 1-5. Stockholm.Olsson, Peter.1893. Bidrag till skandinaviens helminthfauna. K. Svenska Vetensk.-Akad. Handl., new ser., vol. 25 (1892), no. 2, art. 12, 41 pp.,pis. 1-5. figs. 1-81. Stockholm.
ART. 13 TREMATODE PARASITES OF MARINE MAMMALS-?PRICE 65
OZAKI, Y.1925. Preliminary notes on a trematode with anus. Journ. Parasite!., vol.12, no. 1, pp. 51-53, pi. 7, figs. 1-2.1925. On some trematodes with anus. Jap. Journ. Zool., vol. 2, no. 1, pp.5-33, figs. 1-17.Paeona, Coerado.1896. Interno ad alcuni distomi nuovi o poco noti. Atti Soc. Ligust. Sci.Nat. e Geogr., vol. 7, no. 3, pp. 162-180, figs. 1-7. Geneva.PocHE, Franz.1926. Das System der Platodaria. Arch, fur Naturg., vol. 91 (1925), Abt.A, no. 2, pp. 1-240, figs. 1-5, pis. 1-3, figs. 1-95 ; no. 3, pp. 241-458,figs.POIKIER, J.1886. Trematodes nouveaux ou peu connus. Bull. Soc. Philom. Paris, ser. 7,vol. 10 (1885-86), pp. 20-40, pis. 1-4, 30 figs.Pratt, H. S.1911. On Galactosomum cochleari^orme Kudolphi. Zool. Anz., vol. 38, nos,5-6, pp. 143-148, figs. 1-5.Price, Emmett W.1931. A new species of trematode of the family Heterophyidae, with a noteon the genus Apophallus and related genera. Proc. U. S. Nat. Mus.,
vol. 79, art. 17, pp. 1-6, 1 fig.Railliet, Alcide.1896. Quelques rectifications 3 la nomenclature des parasites. Rec. M4d.V6t., V. 73, ser. 8, vol. 3, no. 5, pp. 157-161.Ransom, Buayton Howard.1920. Synopsis of the trematode family Heterophyidae with descriptionsof a new genus and five new species. Proc. U. S. Nat. Mus., vol. 57,lip. 527-573.Ratz, S. von.1900. Parasitologiae jegyzetek. Veterinarius, vol. 23, no. 19, pp. 525-534,figs. 1-4.RiVOLTA, Sebastiano.1884. Sopra una specie di Distoma ncU gatto e nel cane. Giorn. Anat.,Fisiol. e Patol. Anim., vol. 16. no. 1. pp. 20-28, pi. 1, figs. 1-5. Pisa.RxJDoLPMi, Cakl Asmund.1802. Fortselzung CiQv Beobachtun;;en fiber die EingeweidewUrnier. Arch.fiir Zool. uiid Zoot., vol. 3, no. 1, pp. 61-125, pi. 2.1809. Entozoorum sive vermium intestinalium historia naturalis, vol. 2, 457pp., i:ls. 7-12. Amsterdam.1819. Entozoorum synopsis cui accedunt mantissa duplex et indices locu-pletissimi, 811 pp., 3 pi.";;. Berlin.SiEBOLD, Carl Thbodob Ernst von.1836. Helminlhologische Beitrape. Zweiter Beitrag. Sinii/ainns trachealis.Ein doppelleibiger Eingev.'eidewurm. Arch, ffir Naturg., Jahr. 2,vol. 1, pp. 105-116. pi. 3, figs. 1-2.Skrjabin, K. I.1915. Odhneriella rossica nov. gen. n. sp.?Bo.'JOVAHTejiB neHenoqiio-rjiHCTHOft6oJie3mi MopjKcfi {Odhneriella rossica n. g., n. sp., as the cause of ahelminthiasis of the liver in tlie walrus, Odobaenus rosmarus,~i[Russian text.] Arkh. Vet. Nak, vol. 45, no. 11 pp. 1057-1064, 1 fig.113893?32-^5
66 PROCEEDINGS OF THE NATIONAL MUSEUM vol.81
Skbjabin, K. I.?Continued.1923. Gtioabi no nsyieHHio napasHTHnecKHX lepBeft njioxaHaHtix. II. Ciureanaquinqueangularis n. g. n. sp., Hosaa KHraeinafl TpeMaTOAa kohikh[Contributions to the study of parasitic worms of carnivores. II.Ciureana quinqueangularis n. g., n. sp.. a new intestinal trematodeof the cat.] [Russian text.] Trudy Gosudarstv. Inst. Eksper. Vet.,
vol. ], no. 1, pp. 67-71, 1 fig. Moscow.Skejabin, H. I., and Lindtrop, G. T.1919. TpeMaTOAH KHraennHKa co6aK J^ohckoS oSjiacTn (Trematodes intesti-nales des chiens du Don.) [Faissian text] Izvest. Donsk. Vet.Inst., vol. 1, no. 1, pp. 30-43, 3 figs., 1 pi., 3 figs. ; Frencli summary,pp. 43-44.SoKOLOFF, Demetkio, and Cabajlleeo y C, Eduardo.1932. Una nueva especie de trematodo parasito del intestino del manati,8chlzamphistoma manati sp. n. Ann. Inst. Biol., Univ. Nac. Mexico,
vol. 3, no. 2, pp. 163-167, figs. 1-5.SONSINO, Peospeko.1890. Studi e notizie elimintologiche. Atti Soc. Toscana di Sci. Nat., Pisa,proc. verb., vol. 7, pp. 99-114.Stiles, Chaexes Wardell.1894. The anatomy of the large American fluke (Fasciola magna), and acomparison with other species of the genus {Fasciola, s. st.). Con-taining also a list of the chief epizootics of faseioliasis (distomato-sis) and a bibliography of Fasciola hcpatica, by Albert Ilassall.Journ. Comp. Med. and Vet. Arch., vol. 15, no. 3, pp. 161-178;no. 4, pp. 225-243, pis. 1-2 ; no. 5, pp. 299-313, pis. 3-4 ; bibliogra-phy of F. licpatica, by Albert Hassall, no. 6, pp. 407-417; no. 7,pp. 457-162.Stiles, C. W., and Hassall, Albert.1894. A preliminary catalogue of the parasites contained in the collectionsof the United States Bureau of Animal Indu.?!try, United StatesArmy Medical Mu.seum, Biological Department of the University ofPennsylvania (Coll. Leidy) and in Coll. Stiles and Coll. Hassall.Vet. Mag., vol. 1, no. 4, pp. 245-253; no. 5, pp. 331-354.1896. Notes on parasites 42-46. Vet. Mag., vol. 3. no. 3, pp. 151-161.1900. A muscle fluke (Agamodistomnm sp.) in American swine: the lungfluke {Paragonimus westcrmanvi) in swine and its relation toparasitic haemoptysis in man (pp. 560-611, pis. 2.3-24, figs. 24-28) ;the conical fluke {Amphistoiva ccrvi) of cattle slaughtered in theUnited States (p. 611, 1 fig.). (Notes on parasites, 50-52.) 16thAnn. Rep. Bureau of Animal Industry, U. S. Dept. Agr., 1899,pages and figs, cited.1908. Index-catalogue of medical and veterinary zoology. Subjects: Tre-matoda and trematode diseases. Hyg. Lab. Bull. 37, U. S. Pub.Health and Marine Hosp. Serv., 401 pp.1926. Key-catalogue of the worms reported for man. Hyg. Lab. Bull.142, U. S. Pub. Health Serv., 196 pp.Stossich, Michele.1892. I distomi del mammiferi. Estratto dal programma delta CivicaScuola Reale Superiore, 42 pp. Trieste.1904. Alcuni distomi delta collezione elmintologica del museo zoologico diNapoli. Ann. Mus. Zool. Univ. Napoli, new ser., vol. 1, no. 23, pp.1-14, pi. 2, figs. 1-3.
AET. 13 TREMATODE PARASITES OF MARINE MAMMALS PRICE 67
SruNKAED, H. W.1929. The parasitic worms collected by the American Museum of NaturalHistory Expedition to the Belgian Congo, 1909-1914. Bull. Amer.Mus. Nat. Hist., vol. 58, art. 6, pp. 233-289, figs. 1-37.1930. Another trematode with two anal openings (abstract of paper to beread before Amer. Soc. Zool., Cleveland, Ohio, Dec. 30). Anat.Rec, vol. 47, no. 3, p. 363.Stunkakd, H. W., and Alvet, C. H.1929. A new liver fluke, Zalophotrema hepaticum, from the California sealion, Zalophus californianus (abstract of paper to be read beforeAmer. Soc. Parasitol., Des Moines, Dec. 27-31). Journ. Parasitol.,
vol. 16, no. 2, pp. 106-107.1930. The morphology of Zalophotrema hepaticum, with a review of thetrematode family Fasciolidae. Parasitology, vol. 22, no. 3, pp.326-333, pi. 33, figs. 1^.Travassos, Lauro.1929. Contribuicao ao conhecimento dos Heterophyidae. These vet. 31 pp.,1 fig.Travassos, Laxjko, and Vogelsang, Enrique.1931. Novo tipo de trematodeo Opisthotrematidae. Bol. Biol., Rio deJaneiro, no. 19, pp. 143-147.Vinogradov, K. N.1892. O H0B0M1. BH^-fe AByycTKii (Distomum sibiricum) b-b ne-^ienH He.^OBfeKa[On a new species of distome {Distomum sibiricum) in the humanliver.] [Russian text.] Izvest. Imp. Tomsk. Univ., vol. 4 (TrudyTomsk. Obsh. Estestvois. (1890-91), vol. 3), pp. 116-130, figs. 1-5;Weski, Oskae.1900. Mitteilungen iiber Distomum lancea Dies. Centralb. fiir Bakt., Para-
sit, und Infekt., Abt. 1. vol. 27, nos. 16-17, pp. 579-583, 1 fig.Wigdor, Meter.1918. A new fluke from the dog. Journ. Amer. Vet. Med. Assoc, vol. 54,(new ser., vol. 7), no. 3, pp. 254-257, fig. 1^.WiTENBERG, G.1929. studies on the trematode-family Heterophyidae. Ann. Trop. Med.and Parasit., vol. 23, no. 2, pp. 131-239, figs. 1-33.1930. Corrections to my paper " Studies on the trematode family Hetero-phyidae." Ann. Mag. Nat. Hist., ser. 10, vol. 5, pp. 412-414.Zelixr, Ernst.1874. iiber Lcucochloridium paradoxum Cams und die weitere Entwicke-lung seiner Distomenbrut. Zeitsch. fiir wiss. Zool., vol. 24, no. 4,pp. 564-578.
ABBREVIATIONS USED ON PLATES
ac.
U. S. NATIONAL MUSEUM PROCEEDINGS. VOL. 81 . ART. 13 PL 1
1. FiiHcioUi hepnticd: Froin Orciims (irrn. Original.2-b. Campula oblonya: 2, Aher ('(jhoohl, 18.')8; 3, original; 4, section tlirougli iio.sltMior t'lnl of bodyshowing opening.s of intestinal ceca, original; 5, egg: (a) Cross section, (h) lateral view, original.
U. S. NATIONAL MUSEUM PROCEEDINGS. VOL. 81. ART. 13 PL. 2
ni\
3v
'?^
'^ W
\ -'-V^
Mh
6-7. Campula piiUiata: 6, After Looss, 1885: 7, egg, after Looss, 1885.8-10. C. delphini: 8, Ventral view, after Poirier, 1886; 9, dorsal view showing digestive system andarrangement of vitelline ducts, after Poirier, 1886; 10, egg, after Poirier, 1886.
U. S. NATIONAL MUSEUM PROCEEDINGS, VOL. 81, ART. 13 PL 3
SO^
'^??
/3.
//r^ff!
11, 12. Cuini)iila rochi hi inii: 11. .\fter Fdirier, issti; 12, egg, after I'ciirier, Lssfl.13, 14. ZaUiphotrernii htimt'icii in: i:i. Original; 14, egg: (a) Cross section, (b) lateral view,original.15, ITi. LecithodfxmuK yolinlh: ITi, After Odhner, lliO.'); Ifi, egg: (a) Cross section, (b) lateralview, after Odhner, 1905.
U. S. NATIONAL MUSEUM PROCEEDINGS. VOL 81 . ART. 13 PL. 4
i&r.''.
8^,v;vi
) II t \yy.\\>
??'
mt *l2?
?*y V^*
[?**>5r?>
?^*'
n
^^*'??*
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-Wr
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??
?"6j^?9 fiiJi-Sj
9.fl-?
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;.*<>f /s.
17, IS. Ortliosplaiichniis arcticiis: 17. After Odhner. 190.5; 1,S, egg: (a) Cros.s section," (b)lateral view, after Odhner, 190.'i.li). (). fiiit(rriilu.s: After Odiiner, 1905.
U. S. NATIONAL MUSEUM PROCEEDI NGS. VOL. 81 . ART . 1 3 PL. 5
^?vr,
'Si
',!
li
20.
20,21. S!/ii!h(Kiitiu tursiotJix: 20, After I'oirier, IssCi; 21, ckk, after I'oirier, 18S6.22. Odhneridla ronska: After Skrjabin, ISJl.'J.
U. S. NATIONAL MUSEUM PROCEEDINGS, VOL. 81 , ART. 13 PL. 6
',
. !?: ;i'i::1
m
.',/i
m
Hadwenius seymouri. New Genus. New Species23,'Orifcinal; 24, .sagittal .sec-tion through male copulatory organ.s, original; 25, egg: (a) Cross section,(b) lateral view, original.
U. S. NATIONAL MUSEUM PROCEEDINGS, VOL 81. ART. 13 PL, 7
'2C>. Phnliirr ijiistriijihilii>>: .Mtcr Ko.'^sack, i'.llO.27, 2H. Opixthorchis hniikoUis: After Braun, IS!):); 2S, from Ilntkhocriis grypiis, original.
2',), 30. Sli phnnoprora dentkulata: 29, From yjilophus californianus, original; 30, anterior end.
oiiKiiial.
U. S. NATIONAL MUSEUM PROCEEDINGS. VOL. 81. ART. 13 PL. 8
05mm
3L Cyclorchis aiinjiitln: After (\ibbold, 1879.32. Amphimerus lancea: After Weski, 1900.38. ? AmphimeTus lancea: After Cobbold, 1879.34. Metorchis albidus: From dog, original.35. Pseadamphistomii m truncatiim: From Phoca rittilirin.
U. S. NATIONAL MUSEUM PROCEEDINGS. VOL. 81. ART. 13 PL. 9
3C-. C'ri/ptncnlijte lingua: Fr.mi Phncn riliilina, oriK'iiinl.37. Apophrilliis ilonictin: From Phficu liliilina, iingiind.
.'W. A. zdlnphi: (.)i-iginal.31). Oitlacto.mmiim erinaceum: After Poirier, 188G.40. Phocitrema fusifonne: After CJoto and Ozaki, I'M).
U. S. NATIONAL MUSEUM PROCEEDINGS, VOL. 81, ART. 13 PL. 10
? //
/ -::
<^-^/77/77
Chiorchis fabaceus
41, Mature specimen, ventral view, original: 42, l.vmi)h system, dorsal view, original: 43, sagittalsection, original: 44, frnntal section through genital sucker, original; 45, immature specimen show-ing protruded oral sucker, original.
U- S. NATIONAL MUSEUM PROCEEDINGS, VOL 81. ART. 13 PL. 11
\J W W \j '^\y \y w w w y
^ ^ Vy \j\J\J ^9.
46. Opisihotremn diijnnix: After Fischer, 1883.47-49. O. cochleotremn: 47, Original; 48, section tlirougli female genitalia, oiiginal; 4!), .spines fromventral surface, original.50. Piilmonic'ild piihiwimlc: After von Lin.stow, 1904.
U. S. NATIONAL MUSEUM PROCEEDiNGS, VOL. 81, ART. 13 PL. 12
51. Rhabdiopnciif; taylori: After S, .T. .lohnston, 1913.52. OgmofjnKter plicaius: From Dalncnoplera acuturnslrain. Original.53. Bistoma andersoni: After C'obbohi, 1S79.