KEDESCRIPTION OF TWO SPECIES OF TREMATODE WORMS FROM THE MacCALLUM COLLECTION, WITH A NOTE ON THE FAMILY PRONOCEPHALIDAE
By Emmeit W.
PriceZoological Dimsion, Bureau of Animal Industry, United States Department of Agriculture
Recently Dr. G. A. MacCallimi deposited his extensive
and
valu- able collection of helminthsand
other parasites in the collections of theUnited States NationalMuseum,
thusmaking
the material avail- able for study.In
goingover this collection the writer foundmany
of the original specimens of species described
by
Doctor MacCallura.In
view of the fact thatsome
of thespecieswhich
he described were placed in generano
longer recognized as valid except as collective genera, the writer has undertaken to reclassify the trematodes of this collectionand
to redescribe those ofwhich
the original descrip- tions are inadequatefrom
the viewpoint of present-day standards.This paper contains a redescription of
two
species ofmonostomes
belonging to the family Pronocephalidae,and
in this connection an attempt hasbeenmade
to revisethe family.MacCallum
(1916) described amonostome from
the intestine of theFrench
angel fish,Pomacanthus
'paru, forwhich
heproposed thename Monostomum
pomacanthi.An
examination of the available specimens of this species shows that it belongs to the genus Pleuro- gonius Looss, 1902,and
represents the only species of this genusknown from
a fish host.The
specimens available for study are as follows:One
specimen, a totomount
stained with carmine (U.S.N.M. No. 8087), labeled"Monostome,
int., Angelichthys ciliaris (blue angel fish). Aquar- ium, N. Y., Feb. 5, '14"; 10 specimens (3 slides), stained withhema-
toxylin (U.S.N.M. No. 8088), labeled ''•Monostomum 'pomacanthi., int.,French
angel fish, Pomaca7ithus jmru. Aq.May
29, '15";1 badly
damaged
specimen, a totomount
stained with carmine (U.S.N.M. No.8089),labeled"Distome
?,int.,Angelichthysisdbelita.Aquarium. May
27, '16"; 1 specimen, a totomounted
stained with carmine (U.S.N.M. No. 8090), labeled "Distome, int.,Pomacanthus
No. 2365— Proceedings U. S. National Museum. Vol. 78. Art. 22.
23921—31 1
lOOHfrr*
2 PROCEEDINGS OF THE NATIONAL MUSEUM
vol.78arcuatus.
Black
angel fish. Aq. Jany. 13, '17";and
1 specimen, a totomount
stained with carmine (U.S.N.M. No. 8091), labeled"
Monostomum
fomacantlii^ int., PoTnacanthus paru.Aq. Ap.
18,'17." All of thesespecimens
show
evidence of having been flattened before fixationand
certain details can not bemade
out.The
meas- urements given in the following description are doubtless consider- ably in excess ofwhat
theywould
have beenhad
a different technique been followed in thehandlingand
preserving of the specimens.PLEUROGONIUS POMACANTHI (MacCallum. 1916)
Synonym. — Monostomum pomacanthi MacCallum,
1916.Specific description.
—
Pleurogonius:Body
elongated, 3.5 to 6mm.
long
by
1.25 to 2mm. wide
in the region of the ovary; anteriorend
bluntlypointedand
posteriorend
rounded.A
shortdistancecaudad
of the oral sucker, the ventral surface
shows
a deep depression overwhich
projecttwo
shoulderlike structures, one on each side,which
are not connected dorsallyby
a ridge as in Pronocephalusand
related genera.The
cuticle issmooth and
without spines.The
oral sucker is terminal, 153//, to 236/a in diameter; esophagus 310/x to 465ju, long; intestinal ceca extend to posterior
end
of body, the posterior portions bending medially, passing between the testes,and
provided with pocketlike diverticula throughout their entire length.The
excretory pore is oval in outlineand
situated dorsally near the posteriorend
of thebody, butowing
to the extreme flattening of the specimens the remainder of the excretory system can not bemade
out.
The
genital pores are close together, 1.5 to 2.5mm. from
the anterior end,and
situated near themargin
on the left side of the body.The
testes are ovoid, 524|u to 620/x longby
356/*to 387/i wide, with irregular outline, situated near the posteriorend
of thebody and
separatedfrom
each otherby
the terminal portions of the ceca.The
cirruspouch
isfrom
0.93 to 1.5mm.
longby
155/* to 310/twide and
directed obliquelybackward;
it is strongly muscularand
con- tains a well-developed prostateand
a simple,unarmed
cirrus.The
seminalvesicle isstrongly convolutedand
liesfree in theparenchyma
atthebase of thecirruspouch.
The
ovary ismore
orless triangular in outline, 310/ato 520/i longby
232/i to248/* wide, pretesticular,and
situated to the right of themedian
line.The
shell gland is about one-half the size of the ovaryand
lies in themedian
linecaudad
of the ovary.The
vitellaria consist of ovoid folliclesand
are situated immediately anterior to thetestes.(The
masseswhich
areshown
inMacCallum's
(1916) figure of this species, lying in the extreme pos- teriorend and
in the anterior half of the body,and
described as vitellinefollicles, are intestinal diverticula filled with ingesta.)The
ART.22
TWO
SPECIES OFTREMATODE WORMS—
PRICE3
uterus runs anteriorlyfrom
the shell gland inmore
or less regular coils extendingbeyond
the lateral limits of the ceca, and terminates in a well-developed metratermwhich
is about two-thirds as long as the cirruspouch.The
eggsare oval,30/.longby
18/x wide, provided with a slender filament ateach pole.Hosts.—
Angelichthys ciliaris, A. isabelita, Poviacanthus arcuatus,and
P. paru.Location.
—
Intestine.Dlstrihiition.—\JmiQ({ States
(New York Aquarium).
In 1917 a peculiar species of trematode
was
described byMac- Callum from
the oviducts of Trionyx aspidonectes ferox towhich
the
name Paramphistomum
aspidonecteswas
given.Poche
(1926) in discussing his supersuperfamilyParamphistomida
writes that"in no case does this species belong to the genus
Paramphistomum
or to
any
of the other genera ofParamphistomida
which are difi'er-Figure1.
—
Pledrogoniuspomacanthi (MacCallum,1916). Ventralview entiatecl at present."
Fukui
(1929) states: "The
generalanatomy
isapparently that of an amphistome, as the author pointed out.
But
the situation of the genital pore is entirely differentfrom
that of all the other amphistomes, the arrangement of the vitellaria is pe- culiarand
the position of the excretory poreis also remarkable. In general appearance thisworm
issomewhat
like a distome stood onits head." Because of these characters
which
are so outstandingly differentfrom
all other amphistomes, he proposed anew
genus, Opisthoporus.1and
family, Opisthoporidae, for thisspecies, assuming, as he says: "that there are no mistakes in the original description."The
spechnensof this specieswhich
are available for .study consist of five specimens representingtwo
different lots.The
first consists offour specimensstainedwith carmineand mounted
on a single slide (U.S.N.M. No. 8092) labeled ''''Paramphistomum aspidonectes., ovi- duct, Aspidonectes ferox. Soft-shelled turtle. Aq.May
10, '16,"PEOCEEDINGS OF THE NATIONAL MUSEUM
VOL.78and
the second,onespecimen, also a totomount
stained with carmine (U.S.N.M. No. 8093), labeled '"''Paramyhistomwii
aspidonectes n. sp., int.,Trionyx
sjrbiifer, soft-shelled turtle. Firstfound May
10, '16.W.
H.,Aug.
16, '23."As
in the case of Pleurogonius pomacanthi^the specimens have been flattened
and
aresomewhat
distorted,and some
characters can not bemade
out with certainty.OPISTHOPORUS ASPIDONECTES (MacCallum, 1917) Fukui, 1929i
Synonym. —Parmn/phisfoniutn aspidonectes MacCallum,
1917.
Specific description.
—
-Oplsthoporus:Body
spindlelike in outline,more
attenuated posteriorly than anteriorly, 2.4 to 3.2mm.
long by620/Ato 930/A
wide
at the equator. Cuticlesmooth and
without spines.Oral sucker subterminal, 310/x to 403/a in diameter.
There
is a low ridge orcollar immediatelyposterior to the oral suckerwhich
resem- bles thatin thegenusEucotyle. This collaris indistinct inthe speci-//n/n.
Figure 2.
—
Opisthopords aspidonectes (MacCallum, 1917). Ventral view
mens from
Aspidonectes ferox but is quite evident in the specimenfrom Trionyx
spinifer.There
isno
acetabulum.The
esophagus isfrom
230/A to 310/* long; intestinal ceca simpleand
without diver- ticula,and
terminating near the posterior edge of the anterior testis.The
excretory pore is simpleand
terminal, but the details of the remainder of the excretorysystem can not bemade
out.The
genital pore is situated inthemedian
line atthe bifurcation of the intestine.It is impossible to determine Avhether the
male and
female genital openings are separate or whether theyopen
into acommon
sinus.The
cirruspouch
is pestle-shaped, 685/* to 774/* long, stronglymus-
cular, containing a w^ell-developed prostate
and
a long cirrusarmed
with needlelike spines.The
seminal vesicle is a convoluted dilation of the vas deferensand
lies free in the parenchyma.The
testes are^While this paper was in course of publication a paper has appeared by Stunkard (Trans. Amer. Microsc. Soc, vol. 48, no. 3, 1930), in which a description ofthis species is given. Stunkard had access to an abundance of well-preserved material and his de- scription of thisform is, therefore, more complete thanthatgiven in this paper.
AET.22
TWO
SPECIESOF TREMATODE WORMS
PRICE5
situated in the posterior end of thebody and
are unequal in size.The
anterior testis is rounded, 248/^by
217/x; the posterior testis isabout200/xin diameter
and
liessomewhat
postero-dorsal of the ante- rior.Ovary round
to oval, 125^ to 155jii in diameter, pretesticular,and
situated to the right of themedian
line.The
shell gland isaboutone-half the size of the ovary
and
is situatedsomewhat
dorsal to it.The
uterus runs anteriorly in irregular transverse coilsand
terminates in ametraterm which
is about one-half the length of the cirrus pouch.The
vitellaria are extracecaland composed
of large, discrete,rounded
follicles extendingfrom
the level of the posterior edge of the ovary to the level of the origin of the metra- term.Eggs
oval, 23julongby
VSjxwide,provided with a longfilament ateach pole.Hosts.
— Amyda ferox and
A. spiniferar
Location.
—
Oviductsand
intestine.Distribution.
—
United States(New
York, N. Y.,and Woods
Hole, Mass.).From
the above description the genusmay
be tentatively defined as follows:Genus
OPISTHOPORUS
Fukui, 1929Genericdiagnosis.
—
Pronocephalidae:Body
spindlelikein outline,more
attenuated posteriorlythananteriorly;cuticlesmooth and
with- out spines; cephaliccollarpresent. Oral suckersubterminal;esopha- gus simple; intestinal ceca without lateral diverticula. Genital poremedian;
cirruspouch
pestle-shaped, well developed; cirrus spiny;testes situated in posterior
end
ofbody
with their fields coincidingand
zones slightly overlapping, the posterior testis being slightly dorsal of anterior; ovary pretesticular, toright ofmedian
line; shell gland dorsaland
slightlymedian
of ovary; vitellaria extracecal, ex- tendingfrom
ovary to level of origin of metraterm; egg with long filament at each pole. Parasiticin turtles.Type
species.—
Opisthoporus aspidonectes(MacCallum,
1917) Fukui, 1929.The
genus Opisthoporus shows affinitieswith boththeNotocotylidaeand
Pronocephalidae, but its resemblances to the lattergroup
are greater than to the former.The
position of the genital pore is sug- gestive ofthe Notocotylidae,butthe presence of a cephalic collarand
the pretesticular, dextral position of the ovary are characteristic of mostmembers
of the Pronocephalidae; Opisthoporus is therefore regarded asbelonging to the latter family.Up
to the presenttime the following genera have been included in the family Pronocephalidae: Pronocephalus Looss, 1899; Charaxi-2Thewriter is indebted to Miss Doris M. Cochran, of the U. S. National Museum, for the correctnamesof these hosts.
6 PROCEEDINGS
OFTHE NATIONAL MUSEUM
vol.7Scephalus Looss, 1901; Pleurogonlus Looss, 1901; C'ricocephalus Looss, 1899; Glyphicephalus Looss, 1901; Epibathra Looss, 1902; Pyeloso- mur)iLooss, 1899;
Desmogonius
Stephens, 1911;DiaschistorchisJohn- ston, 1913; Wilderia Pratt, 1914; Syn.echorchis Barker, 1922;Hip-
pocrepis Travassos, 1923;and
Astrorchis Poche, 1926.The
genera Diaschistorchis Johnston, Synechorchis Barker,and
'Wilderia Pratt are so similar in shapeand
arrangement of the in- ternal organsthatSynechorchisand
Wilderia should be considered assynonyms
of Diaschistorchis,and
D.pandus and
S.megas
apparently should be regarded as being specifically identical as well as con- generic. Diaschistorchis ellipticus (Pratt) differsfrom
D.pandus
in the absence of a cephalic collarand
polaregg
filaments,and
for the present theymay
be considered as distinct species.The
genus Hippocrepiswhich was
proposedby
Travassos (1922) forMonostoma
hippocrepis Diesing, a parasite ofHydrochoerus
capyhara, has been placed in the family Pronocephalidaeby Poche
(1926).From
a study of the descriptionsand
figures of this species as givenby Braun
(1901)and
Diesing (1855), it is the writer's opinion that the genus Hippocrepis should be retained in the family Notocotylidaewhere
itwas
placed originallyby
Travassos. Flip- pocrepis hippocrepis is characterizedby
thepresence oftwo
muscular flaplike lobes at the anteriorend
of the body, a pretesticular ovary, a preovarial shell gland,and
a genital pore situated in themedian
line in the anterior part of the })ody.
The
cephalic lobesand
pre- testicular position of the ovary give this species a su]Derficial re-semblance to certain
members
of the Pronocephalidae.The
struc-ture of these lobes, however, according to the description of
them
as givenby Braun
(1901), does not appear to be comparable to that of the cephaliccollar ofmembers
of the Pronocephalidae.The median
position of the ovaryand
preovarial position of the shell gland are characteristic of all genera of Notocotylidae, while in the genera ot Pronocephalidae the ovary is situated to the right of themedian
lineand
theshellglandiseither dorsal,median,or posteriortothe ovary.^The
host relationship of Hippocrepis, although not necessarily a decidingfactor,isinsupport oftheopinionthat thisgenus should be placed in thefamilyNotocotylidae, as allmembers
oftliis family are parasitic inwarm-blooded
vertebrates while those of the Pronoce- phalidaeare parasiticin cold-bloodedvertebrates.Fuhrmann
(1928) doubtfully places Hippocrepis in the subfamily Nudacotylinae, butit seems to the writer that it should be placed in the subfamily Notocotylinae on account of the position of the genital pore
and
the^Baylis (1928) described a species, Notocotylus chionis, in wliicli he says that the shell gland liesbehindtheovary. Thebody whichhefiguresas being the shellglandmay
be this structure, but if so the postovarial position of the shell gland in a species of Notocotylusisunusual anda notable exception to the rule.
Ai'.T.2-1
TWO
SPECIES OFTEEMATODE WORMS
PRICE 7distribution of the uterine coils.
In
the genera of Nudacotylinae the genital pore is postuterineand
to the side of themedian
line,and
the uterinecoilsextendaconsiderable distance beyondthe lateral limits of the ceca, while in the Notocotylinae the genital pore ispreuterine
and median
in position,and
the uterine coils are confined for themostpart within thelateral limitsof the ceca.Moiiostomhumsphargidis
MacCallum,
1921, isconsideredasynonym
of Astrorchis renicapite (Leidy).
A
comparison of specimens (U.S.N.M. No. 8094) of this speciesfrom
theMacCallum
collection with specimensfrom
theLeidycollectionshowsno
essential differences.Linton (1910) described
two
trematodes, Barlsovium er-ubescens,new
genusand
species,and Hlmmomum.
cmiclidvlwin^new
genus and species,which
unquestionably belong tothe Pronocephalidae.So
far as it is possible to determinefrom
the descriptions of these trema- todes, there appears to beno
justification for regarding these species as belonging to separate generaand Himasomuon
is considered asynonym
ofBaHsomum, H. candidulum
Linton, therefore,becoming Bansoinunn cmididulum
(Linton).The
genusBarlsomwm
is appar- ently closely related to GlyphicepJiaJus Loossand
Epibathra Looss, the principal difference between thesegenera being intheir excretory systems. InBarisomwni
the limbs of the excretory vesicle unite at the anterior end of the body, in Glyphlcephalus the limbs do not unite anteriorly,and
inEpibathra
the limbsdo
not unite but are connectedby means
of anumber
of transverse anastomoses.Whether
the excretory systemis a character of sufficientimportance to justify the separation ofthese otherwise identical genera is a matter of per- sonal opinion. Further investigation is necessary before this point can be definitely settledand
for the present these generamay
be regarded as valid.In
considering the classification of this family certain difficulties arise in regard to the validity of thename
Pronoceplialidae. Looss (1899) proposed the subfamily Pronocephalinae, but he designated no type genus, although he apparently intended the genus Prono- cepTialus as such, sincehebases thename
on itand
gives the diagnosis of this genus immediately following tlie subfamily diagnosis. In 1902,he elevatedthissubfamily to therankof familyand designatedGlypMcephalus
as type genus. This is in violation of article 4 of the International Kules of Zoological Nomenclaturewhich
says:"The name
of a family isformed by
adding the ending idae. thename
of a subfamilyby
adding inae^ to the stem of thename
of itstype genus."
Two
solutions of this problem, therefore, appear pos- sible: (1) Pronocephalus isintended astype genus of the subfamily Pronocephalinaeand
the elevation of this subfamily to the rank of family retainsPronocephalus as typein spite of the later designation8 PROCEEDINGS
OFTHE NATIONAL MUSEUM
vol.78 ofGlyphicephalusastype,or (2),thename
Pronocephalidaeisinvali- datedby
the designation of Glyphicephalus astypeand
anew
familyname must
be proposed. Pronocephalus^ unfortunately,is one of the least typical of the genera ofthisgroup, but in view of the factthat the adoption of the second alternativewould
introduce anew and
unfamiliarname
for onewhich
has been in use for almost 30 yearsand would
serve no usefulpurpose, it is the writer's opiniontliat the family Pronocephalidae should be retained in spite of its atypical type genus.The
genera assigned to the family Pronocephalidae appear to fall into three distinct groupswhich
are regarded as constituting sub- familiesand
the following classification is, therefore, proposed with keys as formulated here for the separation of the genera:
Family PRONOCEPHALIDAE
Looss,1902 Synonym. —Opisthoporidae Fukui, 1929.
Family
diag7iosis.—
Elongated monostomes,more
or less deeply concave ventrallyand
convex dorsally; anteriorend
usually pro- vided with a collarlike enlargement; posterior end eitherrounded
or truncate, or provided withtwo
papiilalike processes. Oral sucker simple, terminal, or subterminal; intestinal ceca simple or provided with diverticula,and
extending to posteriorend
of body. Excretory pore dorsal, near posteriorend
of body; excretor}^ vesicle Y-shaped, the stem extending to the shell-glandcomplex and
the limbs extend- ing to the anterior end of body. Genital pore in the anterior half of the body, usually to the left of themedian
line. Cirruspouch
well developed, containing cirrus, ductus ejaculatorius,and
pars prostatica; seminalvesicleconvolutedand
lying free inparenchyma
;
testes usuallypostovarial, entire or divided into follicles, zones coin- ciding or overlapi^ing,
and
fields usuallyseparate.Ovary
pretesticu- laror posttesticular, to theright ofmedian
line; shell-glandcomplex
median, postovarial; Laurer's canal present; receptaculum seminis absent; vitellaria moderately developed, extracecal, in posterior half ofbody
; uterus consisting of regular transversecoils terminating in well-developed metraterm; eggs relatively small, numerous, usually providedwith polar filaments. Parasites of cold-bloodedvertebrates.Type
genus.—
PronocephalusLooss, 1899.KEY TO THE St'EFAMtLIES OF PROKOCEPHALIDAE
1. Testespostovarial and tandem in position; genital pore in median line.
Opisthoporinae.
Testespreovarial or postovarial,latei'alandoppositeinposition; genitalpore
to left of median line 2.
2. Testes preovarial Charaxicephalinae.
Testes postovarial Pronocephalinae.
Ar.T.22
TWO
SPECIESOF TREMATODE WORMS
PRICE 9Subfamily Pronocephalinae
Looss, 1899Subfamily
diagnosis.—
Pronocephalidae: Cephalic collar present.Genitalporetoleftof
median
line. Testespostovarial, zonescoincid- ing or overlapping, fields separate or touching. Parasites of marine turtlesand
fishes.Type
genus.—
Pronocephalus Looss, 1899.KEY TO THE GENERA OF THE SUBFAMILY PRONOCEPHALINAE
1. Testes intercecal,situated obliquelytolong axis of body.
Pronocephalus Looss, 1899.
Testes largely extracecal, situated opposite each other 2.
2. Testes divided into several distinct follicles Diaschistorchis Johnston, 1913.
Testes not divided into follicles.
3. Posterior end of body more or less truncate and bearing two small, conical, lobe.ikeprojections Cricocephalus Loosf>, 1899.
Posteriorend ofbody roundedand withoutlobelike projections 4.
4. Subventral lobes of cephalic collarnot connected by a dorsal ridge.
Pleurogonius Looss, 1901.
Subventral lobes of cephalic collar connected by a dorsal ridge 5.
5. Ventral surface ofbody provided with longitudinalrowsof glands.
AdenogasterLooss, 1901.
Ventral surfacenotprovidedwith glands as above 6.
6. Testes deeply lobed; cirruspouch shortandhardlyextending mediallybeyond innerlimitof left cecum; ceca inwideserpentineundulations 7.
Testes smooth or slightly indented but not deeply lobed; cirrus pouch long andextendingtomedian line; ceca not asabove 8.
7. Body strongly concaveventrally,wider posteriorlythananteriorly;ceca pass
between testes Pyelosomum Looss, 1899.
Body not strongly concaveventrally, sidesofbody almost parallel; ceca pass
dorsad of testes Astrorchis Poche, 1926.
8. Limbs ofexcretoryvesicleunite nearanteriorendof body.
Barisomum Linton, 1910.
Limbs of excretory vesicle do not unite anteriorly 9.
9. Limbsof excretoryvesicleconnectedbytransverseanastomoses.
EpibathraLooss, 1902.
Limbs ofexcretory vesiclenot connectedby anastomoses.
GlyphicephalusLooss, 1901.
Charaxicephalinae, new
subfamilySutfamily
diagnosis.—
Pronocephalidae: Cephalic collar present or absent. Posteriorend
ofbody
provided withtwo
conical projec- tions. Genital pore to left ofmedian
line. Testes preovarial, con- sisting oftwo
rows of follicles, either intercecal or extracecal. Para-sites of marine turtles.
Type
genus.— Charaxicephalus Looss, 1901.
KEY TO THE GENERA OF TUB SUBFAMILY CHARAXICEPHALINAB
Cephaliccollar present; testes intercecal Charaxicephalus Looss, 1901.
Cephalic collar absent; testes extracecal Desmogonius Stephens, 1911.
10 PROCEEDINGS
OFTHE NATIONAL MUSEUM
vul.78Opisthoporinae, new
subfamilySubfamily
diagnosis.—
Pronocephalidae:Body
spindle-shaped in outline. Oral sucker subterminal. Cephalic collar present. Genital pore median. Testes with zones slightly overlappingand
fields co- inciding. Parasites of soft-shelled turtles.Type
genus.—
O'pisthopoi'us Fukui, 1929.REFERENCES
Baylis, H. a.
192S.
— A
new species of Notocotylus (Tfomatoda), with some remarks on the genus. Ann. Mag. Nat. Hist.,Lend.,ser. 10,vol. 2,December, pp. 582^585, figs. 1-2.Beaun, Max.
1901.
—
ZurKenntuiss der Trematoden der Saugethiere. Zool. Jahrb., Jena, Abt. f. Syst., vol. 14, March IS, pp. 311-348, pis. 19^20,figs. 1-17.DiEsiNG, Karl Mobitz.
1855.
—
Neunzehn Arten von Trematoden,Denkschr.d.k.Akad.d.Wissensch., Wien., Math-naturw. Gl., vol. 10, Al.t. 1, pp.59-70, pis. 1-3.Fuhbmann, Otto.
1928.
—
Zweite Klasse des Cladus Platyhelmmthes: Trematoda. Handb. d.Zool. (Krumbach), Borl. &. Leipz., vol. 2, December 6, pp. 1-128, figs. 1-171.
FuKtri, Tamao.
1929.-
—
Studies on Japanese amphistomatous parasites, with revision of the group. Japan. Journ. Zool., Tokyo, vol. 2, August 20, pp.219-351, figs. 1^5.
Linton, Edwin.
1910.
—
Helminth fauna of the Dry Tortugas. 2. Trematodes. (In Papers from the Tortugas Laboratory, Dept. Marine Biol., vol. 4) Car- negie Inst. Wash. Publication (133), pp. 11-98, pis. 1-28, figs.1-241.
Looss, A.
1899.
—
Weitere Beitrage zur Kenntniss dor Trematoden-Fauna Aegyptens, zugleich Yersuch einer natiirlichen Gliederung des Genus Dis-tomum Retzius. Zool. Jahrb., Jena, Abt. f. Syst., vol. 12,
December28, pp. 521-784, figs. A-B, pis. 24-32, figs. 1-90.
1902.
—
Ueber neue und bekannte Trematoden aus Seeschildkroten. Nebst Erorterungen zur Systematik und Nomenclatur. Zool. Jahrb., Jena, Abt. f. Syst., vol. 16, November 24, pp. 411-894, figs. A-B,pis. 21-32, figs. 1-181.
MacCallum, G. a.
1916.
—
Some new species of parasitic trematodes of marine fishes. Zoo-pathologica, N. Y., vol. 1, August, pp. 3-38, figs. 1-16.
1917.
—
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