STUNKARD

Descriptions of some mew and

little known Trematodes of Turtles

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1914

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DESCRIPTIONS OF SOME NEW AND LITTLE KNOWN TREMATODES OF TURTLES

BY

HORACE WESLEY STUNKARD

B.S. Goe College, 1912

THESIS

Submitted in Partial Fulfillment of the Requirements for the

Degree of MASTER OF ARTS

IN ZOOLOGY

IN

THE GRADUATE SCHOOL

OF THE

UNIVERSITY OF ILLINOIS

1914

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UNIVERSITY OF ILLINOIS

THE GRADUATE SCHOOL

| HEREBY RECOMMEND THAT THE THESIS PREPARED UNDER MY SUPERVISION BY |

HORACE WESLEY STUNKARD |

DESCRIPTIONS OF SOME NEW AND LITTLE KNOWN ENTITLED .

TREMATODES OF TURTLES

BE ACCEPTED AS FULFILLING THIS PART OF THE REQUIREMENTS -FOR- THE

MASTER OF ARTS

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INTZODUCTION While considerable attention has been paid to the a8 study of the parasites of man and domestic eh inete, less work

has been done on those organisms which infest animals of little economic importance. From a purely scientific point of view, a knowledge of these forms is essential, since it affords a more complete morphological basis for the determination of relationships and contributes to the solution of taxonomic problems.

During the year 1913-14, I collected parasites of turtles, and the present paper contains descriptions of some of the trematodes found. The material was obtained from seventy specimens collected from a wide extent of territory. The number of individuals of each species examined, the locality from which they were procured, and the parasites found are listed in the following table. For assistance in securing the turtles, crateful acknowledgments are due to Dr. N-.A.Cobb of Washington, D-C., Professor A.-W-Orcutt of Denison University, Professor J.E.Ackert of Kansas State Agricultural College, and Professor W.EK.Burge of the University of Illinois. The work

was done under the direction of Professor denry B. Ward, to

whom the author is indebted for inspiration and suggestions.

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Name of Number lULocality Organ Trematoda turtle examined ' infécted Polystoma Pe has- P.orbic- sali ulare

Chrysemys marginata Walker,Ia. lung

“4 Urbana, Ill. intestine

8 Chicago,I11. lung

Chelydra serpentina Granville,O- lung Urbana, Ille none

Chrysemys (lung elegans 2 Havana, Ill. (esophagus (intestine

ny Newton, Tex. he

Pseudemys troosti intestine

Malacoclemmys geographicus

Cistudo carolina Falls Church,Vae none

Aromochelys carinatus Newton, Tex. Uninary bladder

Chrysemys concinna intestine

Trionyx (esophagus ferox (intestine

Malacoclemnys leseurii 10 (esophagus (intestine

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»matoda (cont'd) Nematoda Acanthocephala docotylea Distoma = Monostoma dina n.spe. Tecorti nespe-1x 2x 3x

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Methods. The parasites were studied alive, in toto mounts; and in serial sections. For the live study, the worm ‘was placed on a slide in a drop of water or normal saline solu- tion and examined with the low powers of the microscope. The water supports the cover-glass sufficiently to permit the animal to move freely. The importance of the live study can not be over-emphasized as it is the best method of tracing the excretory system, and the only way to observe the movements of the animal. The material was fixed in Gilson's, Zenker's, Vom Rath's, Tellyesniczky's,and Kleinenberg's fluids, as well as in a saturated solution of corrosive sublimate to which 2% of

glacial acetic acid had been added- Sections were cut in trans-

verse, sagittal,and frontal planes from specimens stained in

toto in an aqueous solution of Ehrlich'’s acid haematoxylin, destained in 70% alcohol, and counterstained with eosin, Balsam mounts were made of toto specimens stained with Mayer's haem-alum, Conklin's picro-haematoxylin, ihrlich's acid haem- atoxylin, and Mayer's paracarmine. Other stains were tried

but the results were less satisfactory.

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In seven of the ten specimens of Malacoclemmys leseurii from Newton, Texas, were found specimens of an undescribed aspidocotylean, the numbers present in each turtle varying from four to twenty five.

Description. The worm (Figs-1,3) is about 1-5 mm in length by O-7 mm in width, and consists of two distinct parts, an anterior forebody and a posterior, .ventral adhesive disc.

The forebody has the shape of a cornucopia, the larger end being attached obliquely to the central two-thirds of the dorsal sur- face of the adhesive disc. When extended (Fig.1) it manifests an elongate form, projecting beyond the adhesive disc a distance equal to the length of that structure; when in a retracted, com- pact condition, it may not project beyond the disc. The total length of the worm varies therefore with the state of extension of the forebody, from the length of the adhesive disc to twice that distance.

The adhesive disc is a muscular organ, used for attach- ment and locomotion. It has a crenulate oval outline, the dorsal surface is arched and the ventral surface is flattened. There is a limiting membrane separating the musculature of the dise from the parenchyma of the body. The ventral surface is divided by two longitudinal and eleven cross ridges into thirty two acetabula, which are arranged in three rows, there being

ten median and twenty two peripheral alveoli. These compart-

ments changé in shape with the movements of the animal, becom-

ing circular, oval or quadrangular.- The size and Shape of the

disc are relatively constant, a series of measurements of

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6 twenty mounted toto specimens show that it varies only from 1.2 mm to 1-4mm in length, and from 0-58 mm to 0-78 mm in width.

Since in functional capacity and superficial form, this structure

recalls the molluscan foot, it has often been termed the foot,

altho the morphological comparison is not precise.

Movement consists of extension and retraction of the forebody, which may be turned in any direction, and in the less striking and more restricted movement of the disc. The disc has a tendency to turn up at the edges, especially at the anterior and posterior ends.- In adhesion the organ may act as a unit or the different alveoli may function separately- In locomotion there is a regular series of movements, the forebody is extended and attached by the sucking action of the mouth funnel, then the dise is loosed and the forebody contracted, bringing the anterior part of the disc near the mouth, when the disc is attached and the series of movements repeated. The worm moves rapidly across the stage of the microscope.

Details of Structure. Externally the worm is covered by a non-cellular cuticula, which is thichest on the dorsal side of the body, and thinnest on the ventral surface of the adhesive dise. It is without hooks or Spines, and on the dorsal side reaches five micra in thickness, while on the disc it is only One micron in thiekmess. This cuticula is deflected inward at

the external openings. Its outer part is darker colored and appears more dense than that near the dermomuscular wall, altho there is no line of separation. Immediately inside the cuticula is the three layered dermomuscular wall, the circular, longitud-

inal and oblique muscles occurring in the order mentioned, the

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7 circular lying next to the cuticula, and in all parts of the body being better developed than the others. In some places the long-~ itudinal and oblique muscles are very scanty, however those of the ventral side of the forebody are continued caudad in a thin sheet, the so called septum, (Fig-8), which lies just above the limiting membrane of the musculature of the dise and extends posteriad as far as the hinder end of the cirrus sac.

The parenchymous muscles of the forebody are long, often much branched, and most abundant in locations where they

connect different parts of the body wall with each other or with adjacent internal structures. In the anterior part are many well developed muscles, used in the movement of that region. Running longitudinally among the vitellaria also, there are many muscle fibers. Sphincters and dilators occur at the genital pore, excre- tory pore and at the opening between the pharynx and intestine.

As previously mentioned, the adhesive dise is separated from the forebody by limiting membrane, (Figs.3,4). This mem- brane runs parallel to the general course of the external, ventral surface, projecting ventrad at each ridge. Extending between the membrane and the ventral wall, there are vertical muscles, often branched, especially at the ends. The ventral projection of the limiting membrane into the ridges of the disc form two sides of long triangular prisms, which extend longitudinally and trans- versely above the musculature of the disc.e One face of this

prism is dorsal and the opposite angle extends ventrad, increasing

the size and prominence of the ridges which separate the disc into fossettes.

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8 The Alimentary Tract. The mouth funnel (Fig.10) is a cup shaped muscular structure, which functions as an organ of attachment, there being no oral sucker. It is 0.08 mm to O-l mm in diameter, sub-terminel in position. There is no prepharynx, the mouth funnel opening directly into the lumen of the pharynx. The latter is a spherical muscular orgen 0.09 mm to 0.1 mm in diameter. It is followed by a short esophagus, an anterior

) section of the digestive tube,differing only in that it is lined with flattened epithelial cells. The intestine is of the rhabdo- coele type, extending on the dorsal side of the body 0-1 to 0.2 mm posterior to the hinder end of the testis. It varies but slightly

in caliber, averaging about 0.075 mm in diameter. The wall con-

sists of two layers, a muscular layer comoosed of outer longitud-

inal and inner circular fibers, and inside this a single layer

of large epithelial cellse A portion of the wall of the intestine is shown in Fig. 9, and represents the large deeply staining nuclei of the epithelial cells lying in the proximal, basal part of the cell, while many delicate, elongate processes extend out into the lumen of the canal.

The Excretory System. (Fig-4) Most of the observations on this system were made on the living animal. As the water evaporated from the mount, the worm was flattened, and the larger excretory tubules could be easily followed. The pore is dorsal, at the posterior end of the forebody. There is a single excretory vesicle, which is situated between the large flask like ends of the collecting ducts and the pore. In the pulsations of this

organ, the anterior part contracted and the constriction passed

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9 posteriad, expelling the fluid thru the pore. Two collecting ducts extend cephalad from the excretory vesicle, one on either side of the forebody, median to the vitellaria- Just posterior

to the pharynx, each vessel divides, sending a brach cephalad on either side of the digestive tube, and a median branch which turns caudad,sub-dividing into a branch extending to the region of the genital pore, and a larger, longer branch which paxses posteriorly

receives to the region of the testis andAsmaller side branches. Cross sections show the collecting ducts to be dorsal in position, but the smaller tubes do not show in sections. Female Reproductive Organs. The ovary (Figs-.5,7) is a

small organ, triangularly ovoid in shape, 0-16 mm in length by

O.1 mm in width, and o-5 mm in thicknesse It is located at the

right of the median line, about midway between the cephalic and caudal ends of the forebody.- In well extended specimens, it is nearer the posterior end, and in much contracted ones nearer the anterior end. Cells in various stages of mitotic division were Observed in sections. The duct arises at the posterior, lateral margin of the ovary and passes posteriorly, receiving a common vitelline duct, and then expanding into several enlarzements- It continues posteriorly on the lateral side of the collecting duct of the excretory system,as far as the hinder part of the testis, where it turns to the median line- Here it passes ventrally and anteriorly beneath the testis; in front of the testis it turns dorsad and toward the ovary, but just before reaching the ovary it turns squarely across to the opposite side of the body and passes with little deviation directly to the pore. Eggs with

Shells were found as far from the pore 2s the region below the

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testis, and when the worms were placed in tap water, the eggs

near the pore were extruded. The eggs are few in number, not more than six were found in any one specimen. In size they ranged from 0.071 mm to 0.086 mm in width and from 0.137 mm to 0.145 mm length.- The average of twenty five eggs measured was 0.141 mm in length by 0-075 mm in width. The vitellaria (Figs.1,3) are arranged along the sides of the forebody, extending from the posterior end to the cirrus sace They are more numerous ane closer together in the hinder region, gradually becoming fewer in the anterior part of the vitelline zone. They lie just above the limiting membrane which forms the dorsal boundry of the mus- culature of the adhesive disc, and number up to forty on each Side. In size they vary from small follicles 0.01 mm in diameter toxslobes 0.03 or 0.04 mm in diameter.

Male Reproductive Organs. The testis (Figs.1,3,9) is @ large,single, median organsituated 0-25 mm to 0-35 mm from the posterior end of the forebody, and in the anterior third of the posterior half of the region of the adhesive disc. It is almost Spherical and measures from 0.25 to 0-35 mm in diameter. Cells in all stages of division and mature spermatozoa are to be seen in sections. The sperm duct arises at the anterior, ventral part of the testis and turns to the left, entering the side of a long, much coiled seminal vesicle. The seminal vesicle is a large tube, O.l1 mm to 0-175 mm in diameter, extending from the testis to the cirrus sac. It is coiled eight to sixteen times and filled with masses of spermatozoa. Wear the pore it enters a large muscular walled cirrus sac, 0.2 mm long and 0-145 mm wide, which is filled

with large unicellular, prostate glands. These gland ce!ls are

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Inside the cirrus sac there is a dilated curved portion

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nucleated epithelial cells, (Figs.7,8)- The cirrus was observed in the extruded condition.

The common genital pore (Fig-1l1) is in the median line,

on the ventral side of the forebody, above the point of attach- ment of the adhesive disc. The opening of the cirrus is on the

right side while the metraterm opens on the left.

Sensory Structures. There is a dorsal nerve commissure

crossing the anterior part of the pharynx, and lateral nerves

were traced running cephalad and caudad from it. In about one

third of the specimens, a pair of black eye spots wre present on

the dorsal commissure, situated just dorsad of the lateral walls

of the pharynx. At the ends of the cross partitions of the adhesive dise are the "Marginal Organs",(Randktrper, Looss). These structures (Figs.2,8) occur in the angles where the transverse ridges meet the musculature of the margin of the disc. Leading to the exterior, there is a narrow canal, which is surrounded by strong muscles,,and above this a large circular cavity, which

was empty in the specimens sectioned. In the study of the living

animals, these organs were everted and retracted as the animal

moved. They were moved rapidly and when extended looked like

membraneous sacs. The rapid, connected and accurate movement would suggest a good nerve supply, sltho nerve fibers were not demonstrated. Most workers regard them as sensory, while many

believe them to be glandular as well.

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Comparisons.

This is the third aspidocotylean described

from turtles, the two previously reported forms being Cotylaspis

lenoiri Poirier (1886), and Lophotaspis vallei Stossich (1899),

both African forms. Poirier described C. lenoiri from the intes-

tine of Tetrathyra vaillanti of Senegal, and Looss(1902)reports

it as occurring also in Trionyx notilica of the Nile- Lophotas-

pis vallei lives in the stomach of Thalassochelys corticata, a

turtle of Egypt. The species describes in this paper is very different from Lophotaspis, but shows considerable resemblance to C. lenoiri and exact agreement with the generic description as given by Leidy for Cotylespis, at once places it in that genus.

The characters of the genus Cotylaspis as given by

Leidy (1858) are "Body curved infundibuliform, anteriorly cylin- dro-conical, posteriorly expanding into a subcircular or oval ventral disc with numerous acetabula arranged in a triple series.

Mouth infero-terminal, with prominent upper lip,and protractile

‘into a cup or disc like acetabulum. Intestinal anparatus as in

Aspidogaster, eyes two, distinct, black, situated on either side

of the head. Generative apertures inferior,between the head and

ventral disc."

Leidy's type species, insignis, is described by him as "Translucent white or pink white, upper lip snout like,conical, ventral dise crenate at the margin;acetabula 29, oblong quadrate,

the outer rows continuous in front and behind forming a circle.

Length one half to one line, ventral dise one fourth to one half line. Adheres to the outer surface of the renal organ and upper

margin of the foot, within the cleft of the upper branchisl cavity

of Anadonta fluviatilis and A. lacustris." Further information

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Chautaucua, N.Y-, ,parasitic in various species of Anadonta, and

in two cases in Unio luteolus- However Osborn's description does

not contain complete measurements, and since there are no other

descriptions, many of the following comparison's are drawn from

his figures.

A tabulated comparison of the species of Cotylaspis

shows their specific distinctions.

Characters C. insignis C. lenoiri ¢. tfhadina (Leidy, 1.5-3.0mm

length (Osborn, 1.2-1-8mm 1.7mm 1-2--3emn

width

0.6-lmm 1mm 0.58-0.78mm

ventral disc broadly oval eireular twice as long as wide

acetabula 9 median,

7 median, 20 peripheral

10 médian, 18 peripheral

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marginal organs 20 ? 22

pharynx

no size given Oval,ilmm long,

spherical, 0.9 to 1-5mm wide

lmm in dia.

ovary O0.1-0-3mmlong O0.22mmlong 0.69mm long 0.07-0.13mmwide O-1Ommwide QO-16mm wide ergs 0.£2-0.35mm long 0.43mm long 0.15-0.14mm long

O-1--O-l6mm wide 0.14mm wide 0.07-0.08mm wide

testis 0.18-0.26mm ° Hien 5— 0.35mm

j cirrus sac about size of ovary ? 0.2 by 0-145mm

The above comparison shows decided differences in the |} Size and shape of the worms, the size and shape of the disc, number of alveoli and marginal organs,size of ovary and testis, as

well as size of the cirrus sac and eggs.

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Classification. Revisions or summaries of the Aspidocotylea have been made by Diesing (1859), Taschenberg (1879), Hoyle (1888), Monticelli(1892), Braun (1889-93), and Nickerson (1902). Present information seems to support the valid- ity of the following genera.

I. Aspidogaster von Baer 1827.

Type species, Ae conchicola von Baer 1827.

Oval adhesive disc, four rows of alveoli, margin- al organs present, mouth subterminal,no oral sucker, one testis. This genus contains A.conchicola, which infests the kidney and pericardium of various species of Unionidae in Europe and North America. It also is found in gasteropods and in the immature condition in the intestine of Unionidae. The other species in- cluded in this genus is A- limacoides, a form which Stafford (1896)

and Kofoid (1899) suspect of being identical with A. conchicola.

A. limacoides occurs in the intestine of different species of

leuciscus in Europe. Another species, macdonaldi, was placed

, in this genus by Monticelli (1892) and removed to Lophotaspis by

Looss (1902),

II. Cotylaspis Leidy 1857.

Type species, Ce insignis lLeidy 1857.

Oval adhesive disc, three rows of alveoli, mereinal organs present,mouth subterminal, no oral sucker, one testis. This genus contains C. insignis, C- lenoiri, and C. rhadina,nespe A description of C. insignis has been given.

C. lenoiri was described by Poirier(1886) as a species of Aspid-

ogastere Monticelli (1892) created a new genus, Platyaspis, to

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contain Poirier's species, evidently overlooking the the simil- arity between it and the form reported by Leidy- He declined to

accept the genus Cotylaspis, suggesting that insignis was a spec- ies of Aspidogaster. Braun (1889-93) ascribes the species insig-

nis to Aspidogaster. Kofoid (1899) established the validity of

Leidy's genus, but contended that genus Platyaspis should be

retained for Poirier's species; first because of the difference in the number of alveoli, insignis having 29 and lenoiri 25; second, Cotylaspis has eyes while Poirier did report such struct-

ures for lenoiri; and lastly, because of the difference in habit,

insignis being an ectoparasite, while lenoiri occurs in the intes- tine. Nickerson argues that these cifferences are not of generic importance. To substantiate his position he states that the

number of alveoli is variable in every genus in the family, and

often in the same species. Both insignis and lenoiri have three longitudinal rows of alveoli and the number of fossettes varies

with the number of transverse ridges, which latter incresse in

some forms with the age of the animal. Developmental differences in the number of alveoli are reported for Cotylespis by Osborn,

Macraspis by Jdgerski6bld, Stichocotyle by Odhner, and Aspidogaster

by Stafford. In regard to the presence or absence of eyes, Nick- erson recalls that not all specimens of insignis have eyes, and

that Poirier's silence on the subject does not necessarily imply

that such structures are absent in lenoiri. Nickerson continues the argument by saying that the presence or absence of eyes is of

no more importance than the presence or absence of marginal organs

which structures are not mentioned by Poirier. Concerning the

ectoparasitic habit of one and the endoparasitic habit of the

thi a a

*

ee se ges ae alll dehy Lhe

“a

it SOLF: As Me cd ong

a ty WEG g! ‘345 Gt

Thaler. (howe a eee

‘4 om, Poet & le ~—apit pene ih tyessebacrSacrsce / ¥ Sy mir? ia by & Wee fh. ay | ror |, it ae Seal , " , & ae

‘vet

coanen sorbic tt nebo i es TR wat’ ban es ou r

ie ‘us Bayne t99, sag

Ciiiw weve AOR eee BEE CL ; F . ' a 7

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= a5

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ara | BP ee.

16

other, he says that habit is really of less importance than

habitat, and more significance might be attached to the fact that one is an African and the other a North American form. Morpholog- ically Poirier's species is similar to Cotylaspis and the evidence appears sufficient to justify the suppression of Monticelli's genus Platyaspis and Aspidogaster lenoiri,Poirier 1886, Platyaspis lenoiri (Poir.'86) Monticelli 1892, become synonomous with Cotyl-. aspis lenoiri Poirier 1886.

Cotylaspis then contains the species,

insignis Leidy 1857, a ectoparasite occurring in the mantle cavity of Unionidae in North America.

lenoiri Poirier 1886, from the intestine of the turtle Tetrathyra vaillanti of Senegal.

rhacina n.- spe, from the intestine of Malacoclemmys

leseurii of North America.

III. Macraspis Olsson 1868.

Type species, M. elegans Olsson 1868.

This genus has @ single row of confluent acetabula in the adhesive organ, sense organs present, mouth terminal, one testis. The single species is parasitic in the call bladder of Chimaera monstrosa, a fish from the coast of Europe.

IV. Stichocotyle Cunningham 1884

Type species, S- nephropis Cunningham 1884.

There is a single row of more or less distinct acetabula sense organs lacking, mouth audieininal. oral sucker absent, two testes.

Cunningham's original description was of the lerva and

iE ae

ae

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ane | ml enh fF me lok CU emthedt

he aie * 9 Bit eR OE 1 ohegon | ¢ a = i oy >} Ms tio et ign iA 7 ; Ly i

rey", aL ae mth ye Ae ARI ¥ ? ue? oe Fes ag ey ~ é 4 NY Fn 4 "" nd a " ihe he ot 47 Oe \ ? 7 hie @ | nA f $4 . a + ¢ ¢ i lee

4 vs } { 1 “4 { : ii ah: +] soy * aed i) bis matt 5 eee ~ 9 Pi im" a Se ne aS } : i aN : am r f ee

i

+ ah Se Sea te ' nn? bee ; P 4 a F b a we Siped at er taee dent 2 Pain ar sks 3 = si are 4 py; ma 1) f quer ae: Ms Rais Westy meri | eek Abia We! Z) :

1? Monticelli (1893) declined to recognize its generic importance, thinking it might be an immature form of Macraspis. Odhner (1898) however, by discovering the adult and tracing the life history, established the genus. Adults live in the bile ducts of the liver of rays; larvae occur encysted in the wall of the intestine of the larger marine Crustacea. Cunningham described it from the Norweg- Higes lobster, Nephrons, and Nickerson (1895) reports it from the American lobster, Homarus americanus. 3 V. Sibedé pater Monticelli 1892. Type species, C. michaelis Monticelli 18692.

Adhesive disc with three rows of alveoli; sense organs

present, mouth terminal; oral sucker present; two testes.

There are two species.

C. michaelis occurs in the intestine of Cantharus vul-

paris, a European fish.

@. occidentalis ~Nickerson.1899, occurs in the intestine

of Aplodinotus grunniens of liorth America.

VI. Lophotaspis Looss 1902.

Type species, L. vallei (Stossich) 1899.

Adhesive organ with three rows of alveoli: eversible sense organs present at all the intersections of the ridges of the the adhesive dicc: cirrus absent.

Looss (1901) reported Lophotaspis adhaerens as a worm belonging to new genus of the \spidobothridze, but was not aware that Stossich (1899) had described the same form ag Aspidogaster vallei- lLooss (1902) described and figured the form ag Lophotas-

pis vallei.

In the same paper(1902) Looss compared Aspidopaster

i a “a 7 Mads = ¥ A, a% is 1 | Vm > ' ¥ % 5 ; ¥ . Ae i v th ary hy ‘i j ay we ie eas A ¥ Mu ft A ne 2 + ¥ i a lf f. : ; (. we Nd \ So} mo he is 8 L = La) a i ; Pas | ain? co's ; : AS ie: eer ¥ ; st cRO ON yf f fy al wits bay ii

{

q > te

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oh be ewes see oe *4 i i ie sea)

Vpou) cbaia) by aera

. "4 ce i” ‘fe wed ue eavdden he

* »

' a ut, ae) Y, im oe “a4 , . ~~ am Ot “on Os epee gre es

ee CC aT ee

4" . pa = eons agua Sawer! baie sve, S se ¢ oe & Cine of a |

; RE Pate oka! (ee Ne ie en

é 4 nie i «xh i Ai! hi PMT mid re!

18 macdonaldi with Lophotaspis and placed it in that genus. This tremotoce was reported by Macdonald(1878) but not named by hin, and Monticelli (1892) named the forn as a species of Aspidogaster. Nickerson (1902) declared it to be an aspidobothrid, but different from all other known aspidobothrids, and predicted that a new genus would have to be created for it when its structure is better -known. Macdonald reported 180 extensile structures, like tentacles of a snail, occurring at the margins and intersections of the rid- ges of the adhesive disc- Nothing is known of the internal struct- uree Looss in placing the worm in the genus Lophotaspis states "Mit ihrer tentakeltragenden Bauchschiebe bildet die Art aber ganz zweifellos einen freméen EKindringling in dcr Gattung Aspidogaster, da dessen typischen Art jedenfalls solche Tentakel nicht besitzt. Gerade diesen auffallenden Character aber tielt sie mit Lophotaspisy ich bin geneigt, A. macdonaldi Monticelli, trotsdem wir von seiner inner Organization noch nichts wissen und trotsdem bei ihm die Genitalédffnung wejter Rickwarts liegt als bei Lophotaspis vallei, aus dem Genus Aspidogaster herauszunehmen und zu Lophotaspis zu Stellen". This form certainly does not belong in the genus Aspid- Oogaster, and until its internal structure is known, it may well be placed with Lophotaspis.

| I agree with Jagerski8ld that it is much too early to undertake a final systematis revision of the Aspidobothridae. The discovery of the sexual form of Stichocotyle by Odhner (1898) establishes the fact that at least one species of the Aspidocotylea has an intermediate host, and the family is thus both monogenetic

and digenetic. Nickerson observes “owing to the well known ten-

dency of fresh water conditions to obliterate larval life, 1% may

eM Cy

Ww, Le es td

a | ‘*

bash th sav | px : ee ry uriges ‘wit he

? : : 4 + 2098s Graen 4

Ue

tT Peye

4 ys s = 4 : - T f , (vee ae ie 4 * ua > - Co ¢ ‘ie t - w yA wf , me li ib’ at : 7

ont we OSL. Se - 2 0F

r Pee s+

j p wor) ae it ik cae

4 es . ! > } i Te ~ ee ap -oemenge apeirety mee fant ST ee es asian +

adi

Ld

well be that Aspidogester has secondarily lost a more or less

complicated series of changes, which have been retained by its

relatives inhabiting salt water". The presence of both monogen-

etic and digenetic development within the family, together with

other characters common to both the Heterocotylea and Malacocotylea

designate it as an intermediate group. The morphological structure

is like that of the digenetic trematodes, but whether or not the

Aspidocotylea are primitive or secondarily degenerate forms is as

yet undecided.

‘Gen ibd it si ai fiigepa it cate ane

Beene | . rr rhe ; : i 7 eas re EMP i on

} i. , i t ay,

Polystoma orbiculare, Sp. nov.

Two Beeravenes were obtained from the esophagus of a single specimen of Trionyx ferox, from Newton, Texas, and another similar worm was found in the esophagus of Malacoclemmys leseurii, from the same region. These trematodes were very smell, the same color as the lining of the esophasus, and so firmly attached that they were removed only with great difficulty.

The worms (Fig.12) measured 4, 3.75, and 3.25 mm in length by 1, 0-85, and 0-8 mm in width at the widest. part. The body is oval, flattened dorso-ventrally, altho the shape varies greatly, depending on the state of contraction of the animal. In an extended condition it narrows at either or both ends, and the

contracted form may be broadly oval to quadrate. On the ventral ( gide, at*the-postérior end of the body is an adhesive disc, carry- ing six acetabula, which are arranged compactly in a circle. The dise is slightly wider than the greatest width of the body, being 1.09 mm to 1-21 mm in width, while each bothrium measures about O-4 mm in diameter.

The cuticular covering of the body is 0.014 mm in’ helekuess, and is usually because of the contraction of the body, thrown into minute folds and furrows. Externally the cuticula is dotted with very small chitinous granules.

The anterior sucker is an oval structure measuring 0-2 to 0-22 mm in length by 0-2775 mm in width.e It is merely a mouth funnel, which functions as an adhesive organ, there being no oral sucker. At the base of this anterior sucker is the pharynx :

(Fig-14), a spherical muscular structure 0.3 mm in diameter.

There is a broad dorsal nerve commissure crossing the anterior

i

ie! i. uF 7 » me at ew De I A oo eae ; 7 - ; fa 4 J iv ye ) Py aw A q ae 1h eae A ee WOE Se hic haiti! ie y ' 7 \e a's

ha Se ory a hae

fp hee | be ea ‘Pah ge ae honitthe : a

? Py. £4 yFOT arencetiny foe acy re aoe a, a’

Tet eres 6 seo eee

P “iar a? Pee wae iE fae ak (tx y mi FY i wary on

« ' 7 = ° =< 4 i 5s OF - i = Po oe 4. i . + “ht : ¢ * j \* * \fl" ~ 2 . © { hs tim ADD , + ie £45) 4 +i » om tu if

ue . asf ac} Nish T BhiM 3 FT hisd nN fa JB a a Bole a | ee it: ene

+

ve ae *y ‘a> 4 WO - mriD ei se ag wetted rf sa yt yee ou wv a

4 1) ae ae a OMe dee Ne ae ATA,

21

part of the pharynx, which contains large ganglion cells (Fig. 12).

The dorsal commissure is continuous with a nerve ring that passes ventrad on either side of the pharynx.

The digestive tract is of the triclad type, the pharynx is followed by a short esophagus, 0-17 mm in length and the diver- ticula of the intestine extend as simple tubes to the posterior end of the body, being approximately 0.15 mr in diameter, and ending blindly just dorsal to the middle pair of bothriae The caeca are lateral in position, but are quite close together, being only about 0-2 to 0-25 mm apart. They have the usual muscular coat and epithelial lining, and were empty in the specimen that was ce uticoned.

Reproductive Organs. (Fig-20) The testis (Big-16) is a large spherical organ, 0-4 to 0-5 mm in diameter, situated in the median line of the body. It is in the posterior part of the cephalic half of the worm, and is composed of a large number of small lobes, compact and contained in a membraneous capsule.

Cells in all stages of mitotic division and mature spermatozoa were observed in sections, the spermatozoa measuring approximately 0.014 mm in length. Two tubes, one from either side, arise from the anterior ventral region of the testis and unite to form the sperm duct, which curves dorsad, then after passing cephalad, it turns ventrad into the cirrus sac which opens to the surface at the common genital pore. The ovary is a comma shaped organ, located on the left side of the body. In dorsal view it is from 0.16 to 0.2 mm in length, and 0.08 to 0.12 mm in width, while in

the specimen that was sectioned it is 0-08mm in width and 9.3 mm

(aoad bah pe line nan | peng a ik 3 ‘bara Bala ath aa Seu si whRe, Jaap

ni pene 6 x) Ti Ww oT, oad 74 s-Tx aati

ik Aye ? oy) ge ot ee |

2 ERO et : he te acu: ah oS ae hh be ' ¥ Fond il

‘Fierce ome Gl, ORae Rieke

id « A a » ? AO PP f S7 es iz 2 oh eat Ee ' eik , i “0: 4 4 aftersg iA}, ay £4 ' t ' 2 = * t A . id Ait

Ff - t he 1 44 Pa

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yn MOD" Ase 1 a me 7 Cn sto th Sih aa tone

sy f

Les

{- it oa fe a ( cf ts th sae Groat Stas pon pucrioap lp ieee gn teed eee “a ae rd nd : Re yey J wee) WW aot) ¢ vt ae of ' Pa alee y

22 in thickness. The oviduct arises at the posterior point of the Ovary, and after receiving a short duct from the vitello-vaginal canal, it coils dorsally and anteriorly, passing thru a small shell

gland and crossing to the opposite side of the body, where it

expands into a large uterus. There are two vaginae, one opening

to the surface on either side, at the ventral, lateral margin of the body, just ovposite to the posterior part of the ovary. Near the external opening each vagina receives a duct from the common vitelline' canal, and the vaginae meet in the median line of the body, where a short tube connects with the oviduct. The vaginae thus form a tube, leading thru the body from one side to the other. In each of the Specimens, there was 2 single large egg in the uterus, and in the one sectioned the uterus extended cephalad of the pore and to a point only 0-03 mm from the bifurcation of the intestine. The eggs are. broadly oval, 0-25 mm long by 0-2 mm wide. They are very refractive to light, do not stain, and in sections the ecg looks like a mass of homogeneous yellow yolk.

The uterus and cirrus sac open into the cloaca, the open- ing of the cirrus is anterior and at the left of that of the uterus The common genital pore is situated in the median lineof the body about 0.12 mm caudad of the bifurcation of the intestine, and is surrounded by a coronet of spines, there being 32 in one mounted specimen and 33 in the other. These cirrus spines or hooks (Fig.18) are sickle shaped, and have branched processes which are embedded in the musculature of the walls and in entire length they measure 0.05 mm, the hook proper comprising a little over half the total

length.

hy Gai

es { r

4 P

Levenuedigeaeninicd Oneik mew Alain aa a -, coer e » hal

3 wots (eas wiles ar Bs epitconsety ai i eye St vg fo ui: eeu i fe

2 st RR

a A ae SS ee > fos «+ eed 2 semua —>

) j ys a : , ee tole Bye ee eT

20

The vitellaria are large compact follicles underlying the entire dorsal surface of the body, except the region over the uterus and pharynx. Common collecting ducts run longitudinally along the body, lateral to the intestinal diverticula, and these open on either side to the vaginae.

Armature of the Adhesive Disc. Each bothrium has a chitinous skeletal structure, (Figs.12,17), cup or bell shaped, on the inside walls of which there are 32 parallel cuticular ridges, extending to the exterior. Between the anterior pair of bothria, the chitinous granules which dot the cuticula of the body have been developed into many minute spines. In one of the mounted speci- mens two of the spines were larger than the others and possessed hooked points; they were 0-007 mm in length. There is no regul- arity in the arrangement or characteristic size for the spines. Between the posterior pair of bothria, there are two large chitin-

ous rods, i.e. without sharp hooked tips, 0-075 mm in length, and

between these a smaller pair, 0.046 mm in length. Near the post-

erior margin of the disc in one specimen, there was one well developed hook, and in the other three small chitinous spines. In the base of each bothrium there is a small hook 0.009 mm in

length (Fig.19).

He hie pny

| Hat 2s booting’ "ee

M4 ' , t= ven f bad on ; Aa: ray » hie Py t a tr a Af ays Bee tes z

t

i ~ > a - - =

=e 1

> 5 i

~ 3 > ~ do oe a

. * : . 7 * 4 3

a = Pt

: - NY 1 h s es <r ie = ut See aT ar a - ' “es i @ 2 oi Pa i 2 ane ad) ah Whee he alee ae y ‘(a é ‘a f iy \ a, => é r wr) 7 im Wee da; 4 ¥ ‘ae. 2

24 Comparisons. Of the described polystomes, two species, P. integerrimum and P. wncinatum are parasitic in the urinary bladder of Huropean frogs. The other Buropean form P-.ocellatum occurs in the pharyngeal cavity of Emys europa and Halichelys atra.

The three previously known American forms are all from turtles,

P. oblongum occurring in the urinary bladder of Aromochelys, odora-

tus, P.ecoronatum in the fauces of Chrysemys rugosa, and P-hassali in the urinary bladder of Kinosternum pennsylvanicum. In size and external characters, P. orbiculare agrees closely with Leidy's description of P.coronatum.

A comparison of P. orbiculare with one of the type specimens of P.coronatum, however, shows undoubted specific differ- ences in the number and arrangement of the vitellaria, the size and shape of the adhesive disc, and the number and arrangement of the hooks which occur on the disc. The vitelline lobes of P. Orbiculare are larger, more numerous and more compact, and the differences in the adhesive disc are shown in a tabulated compar- tson.

Characters Pecoronatum P.-orbiculare Shape of disc cordiform circular

Width at anterior bothria Lect mm 0.68 mm

Width at the middle bothria 1-18 mm 1-04 mm

Width at posterior bothria 0.78 mm 0.72 mm

Gential hooks 32, 0-037 mm long 52, 0-05 mm long distal ends bifid. distal ends hooked.

Hooks between anterior bothria 6, 0.023 mm long many small spines

Het aan

Hooks between posterior bothria 4 pairs 2 pairs of rods

Largest pair 0.143 mm long O- O75 mm long median pair 0.074 " bi O- 046 " , posterior pair S022.” if

The shape and armature of the posterior disc of Pe cor- onatum are similar to those features in all previously figured polystomes, P.integerrimum, P. oblongum, and P-hassali, but the Shape of the adhesive disc and character of the armature of the Texas specimens is very different. Fig.17 shows the close prox- imity of the anterior bothria, and the difference in shape, togeth-

er with the departure from the usual arrangement of hooks at once

mark the Texas worms as belonging to a new species. Because of

the shave of the adhesive disc, the name orbiculare is proposed

for this new form.

The simple triclad digestive tract and the reproductive Organs as described for P. orbiculare are similar to the corres- ponding structures in all other polystomes, with the exception of the endoparasitic form of P.integerrimun. The single egg in the uterus is common for all American forms of this genus whose anatomy is known. There are no prominent "Seitenwilste" at the external openings of the vaginae, such as are described for P. integerrimum, but beneath the surface of the body at these points are spongy masses which may correspond to those structures. The vitellaria ave more numerous than in any described American form,

|} and in this respect correspond only to P. ocellatum.

or oer J a, ad Pin, ie ih! ae | AY ee ; +) ES ¢i\) = ge ae - =, 5 He ‘i rs ‘e ‘, . ; " nu v _ si UTE OE tea Sie

“68

F bul

F " ny , re . ° fr sc “a!

; ; ¥ apes | t | 5 a 8 Oia ' iP k u = cule Me. {4

2

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Ms od

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er

dae) :

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ee aa em re A "% f

ry f ty, ra id e «& / : 4 LY _ SD a" ii , re ; vi . iG ' U > ae VA. 1 Te ve el) in ne 6 a en le

rrr *

26

Polystoma hassali Goto, 1899

A single polystome (Fig.21) was found in the washings of |

the intestine and urinary bladder of Aromochelys c2rinatus from Newton, Texas. Unfortunately these organs were examined in the same dish, and the present specimen, which was found in the wash- ings, may be from either, but since Goto described the form from the urinary bladder, it is probable that the present specimen is from that organ. Upon staining and mounting the specimen it proved to be P-hassali, a species that Goto (1899) described from the urinary bladder of Kinosternum pennsylvanicum from Maryland. The worm is 1-35 mm in length, with an adhesive dise the Same Size and shape as that described for P-ehassali. It is armed in the same manner as that figured by Goto. However he reports the smaller hooks that occur in the bases of the bothria and at the anterior and posterior margins of the disc as being 0.33 mm in length, and the larger hooks with bifid bases, and that are between the posterior bothria as being 0-125 mm in length. This is evi- dently an error, since he figures the posterior hooks as about four times the size of the smaller ones. In the present specimen, the larger hooks are 1.25 mm in length and the smaller ones 0.033, which figure agrees with that of Goto by a change of one place in the decimal point. Goto makes no mention of the number or size of eggs. The Texas specimen has a single large ege in the uterus, the usual condition for polystomes. The egg is badly collapsed

but measures 0.255mm by O-11 mm.

wr atinlw ON em ee ee a _ = ; Koc “A ; sa! ert we NAN atte Phyl ss otal ee Lf , i i Ob i, te ie” iN ‘= af | vo Ge rea - ) a } : < e et iiantae o.) (s ; oy i r ri # Ny

et . saute TAN ha

i ; “On ris ei) sella AD Cae ae: te achat 1 | | \ ee oa Stroe oes ay. i ) . a3 a 1 aad 0: gate fod.', tae se wonrn i< é

H | Yon ofdadaxe at SH a

ei Ate Ani¢ b sant ‘. tats

' .]

? - a A.

~ our —_ ~~ ld en eI we ~ - on. of . dor vshemcmrhers asennad eh tater :

* rf im ry rigs Wer

27

The genus Polystoma Zeder 1800, is described by Braun

(1890), "Body long egg shaped, anteriorly tapering to a point,

posteriorly extending as a broad adhesive disc, upon which are two longitudinal rows of outward springing bothria. Between those at the posterior end are large chitinous hooks. The vagima is double] Opening at the right and left sides of the body; eggs oval,without | filaments. Occurs on the gills and in the urinary bladder of Amphibia and in the esophagus of turtles".

The genus contains seven species, among which P-integer-

rimum is the most common and best known species. The adults oceur in the gill chamber and urinary bladder of frogs and toads

in Europe; larvae occur on the gills of tadpoles and migrate to

the urinary bladder when»métamorphosis takes place. Zeller (1876) showed that if sexual maturity .is attained on the gills of the

tadpole, the structure of the worm resulting is very different

from that of the adult which matures in the urinary bladder. The form found in the urinary bladder has a lobulated testis, two vaginae, many eggs in a long coiled uterus, and a much branched

intestine, while the ectoparasitic form has a single testis, no

vaginae, single egg in the uterus, and a simple triclad intestine.

P. integerrimum is about 12 mm long,with the surface cross wrinkled There are eight spines in the coronet surrounding the common genit-| al pore. P-eocellatum Rudolphi 1819 is parasitic in the esophagus

of Emys eurodpa, Cistudo europa, and Halichelys atra, all European

species. It is flesh red in color and from 3 to 5 mm in length.

Eyes are present in young animals but later disappear. The phar-

| ynx is pyriform and there is a simple forked intestine. The testi

OP as ot oan

Fay hae

~ Oe eka; Bh aa eee : ag ae i dente: an

$) RRM a

Lee ©. 2S, Ed aA

7 Yr nfs he as, vat, Vee sa Siig:

28

is a lobed structure; the ovary is a large quadrate organ, in xbout

the middle of the body. The large lobed yolk glands are dorsal

and fill the region from the mouth sucker to the posterior disc.

Their ducts unite to form a common duct which opens into the ovi-

duct. A shell gland and ootype are present. The uterus opens

at the posterior part of the genital pore. There is no internal

vas deferens. The genital pore is in the median ventral line of

the body, posterior to the acetabulum, and is surrounded by a circle of about 40 hooks. the vaginae are anterior, about one third] of the length of the body from the cephalic end. Peoblongum Wright 1879, was described from four specimens obtained in the urinary bladder of the musk turtle, Aromochelys odoratus of Canada. Stafford (1902) also reports it from Chelydra

same

serpentina in the\locality. "The body is oblong, the mouth being

on’ the ventral surface of the rounded anterior end. The pharynx is bowl shaped, The intestinal caeca without branchings, cirrus

coronet composed of 16 alternately large and small hooks, which are

sabre shaped. Length up to 2-5 mm; breadth 1-5 mm- Viviparous.

Egg greenish, 0-255 mm by 0-195 mn. The larva is ocellate, 0.5mm

in length. The testis is a solid structure in the posterior

third of the body. Vitellaria scattered, but not numerous; hooks as in P.integerrimum." The single specimen reported by Leidy from the urinary bladder of Pseudemys rugosa of Maryland, and assigned by him to

the species oblongum Wright,has been examined by Goto (1899) and

his observations show that it is not Peoblongum. It had 16 cirrus

Spines, all the same size and shape, measuring 0.56 mm in length.

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af ct peepee! u

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PPh Lai 4 tai ra > a vi i's wah ont

* Fn at SOE + “it ee a Vi ee eer Oy

29 Each spine consists pf eorti ons, a distal awl shaped part with an expanded base, and handle like part, consisting of two slender filaments; In oblongum Wright, the spines are alternately large and small, hence Leidy's specimen is not oblongum ivright, but must be a new species. The material was in such poor condition, how- ever Goto did not attempt a description and for the present it remains unknown.

Peuncinatum Macé 1880, is about 5 mm long and was report- ed from the gall bladder of Rana temporaria from France.

P.coronatum Leidy 1888, was described from four specimens of Pseudemys rugosa, three from the throat and one from the nasal cavity. Leidy's original description is,"Body when elongated lanceolate; caudal disc wider than the body, cordiform, three pairs of bothria, the body being attached between the anterior two pairs; | changeable in form to oblong, circular or quadrate; with three pairs of minute hooks between the anterior pair of bothria, and One larger and two smaller pairs of hooks between the last pair of bothria, genitsl aperture with circular or transverse ovel coronet of 32 hooks of equal lengith. No eyes visible. Length elongated 4-6 mm, contracting to one half that length and widening propor- tionately ;

Pehassali Goto 1899, was described from the urinary bladder of Kinosternum pennsylvanicum from Maryland. Total length | of body 1-5 mm; body ovate; adhesive disc hexagonal; suckers at the angles, each with a minute hook in the center; Three pairs of | hooks between the anterior pair of suckers; three pairs between

the posterior bothria, a large pair, 0-125 mm long, bifid‘at .the

Mi : 4 f “a a ? b Vein’ a Ly ' ¥ i t p42 i] 1 mi a 4 " ie A Ps ah} ere)

| a oe » <a fat 7 (ati tPadeeie. | raaeramne otha thai a | | | oes rv is tes cui edb 2

Pee | (oak

y he Lee vu Qh bir

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bai) 4 doers ee eae |

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i Re bs ua co a abe ees

| yO deal rand Agee

f BLO a ao% , NO Rae ; ma

| in wa area io 7

~ > § : p ; ric a Leg 1 ! a a , y 1 + ia * sO MA

fom) iL! nego, &

5 fons hit hoe edie we, ‘sith Te Mbee’ ols ag sine i ad

fe! tos BSL th Le RR: jenna sane

. a ay f WAY ae tog et enegiaag ate ates = ans ee: pe! : yy” f (eas “i i : MI ea 7 hi ie) ta tls Mes iS hake ie ae / iy

30.

base, and two smaller pairs, the same size and shape as those that Occur in the suckers, possessing lateral processes and measuring 0.033 mm in length. There is a bifurcated, unbranched intestine. The genital coronet contains 15-16 spines, 0.028 mm in length, straight with wing like vrocesses in the middle. The testis is a large ovoid organ, with a lobulated margin, and situated in the posterior half of the body. The ovary is comme shaped, anterior to the testis , on either the right or left side; the oviduct

| proceeds obliquely forward, and expanding into the uterus, which is just posterior to the genital pore. The veginal onenings are lateral, without papillae, midway between the cephalic and caudal ends of the body proper, the two vaginal canals directed almost straight across the body and meeting in the median line. Genito-

| intestinal canal slightly behind the vagina on the same side of the body as the ovary. Vitelline lobes not very numerous, separated

from one another, mostly confined to the lateral portion of the

body, but present in the median portion behind the testis.

Pian be < ' 5 oe Satna Need iaiensinry: sir ot otecoeae ¥ 2 Chant, ; f oy ae s ae *

add pu soy, “dee wd iw

me

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a a ie aoe eRe:

bs

sts |), Oe ee ite

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airerne? "ee : ‘kok

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.

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j

Telorchis corti, Sp. nov.

In the intestine of sexennof the ten specimens of Malacoclemmys leseurii from Newton, Texas, there were found some fifty distomes, most of them immature worms; and another adult of the same kind was obtained from the intestine of a single individ- ual of Chrysemys elegans from the same region.

These worms proved to be of a single species and later were found to identical with others Dr. H.J.VanCleave had collected in June 1910 at Havana, Ill. from the intestine of Malacoclemmys georraphicus. Dre W.eWeCort, to whom this latter material had been given, turned it over to me, together with specimens of Telorchis aculeatus and T.-robustus, and in appreciation of his kindness, the worms which proved to be a new species, I shall name Telorchis corti.

The largest specimens (Fig24) ‘measured 7.15 mm in length, by 0-5 mm in width at the widest part, and the smallect eee mature forms i-e- those with eggs in the uterus, were about 4 mm long and 0-35 mm wide at the widest part. In the adult the great- est width is at the region of the acétabulum, the worm tapering Slightly toward either end, ,being narrower at the caudal end of the body. In immature specimens the section of greatest width is further anterior, being in the pharyngeal region. The worm is flattened dorsoventrally, being thickest at the median line and

becoming thinner toward the lateral edges.

The body is covered by a heavy cuticula, 0.004 to 0.008

mm in thickness, and in the living worms it was armed with a

covering of minute spines arranged in rows. ‘These Spines were

ee re a as = ED Ob em ne se a aE PA TN a 3 peat

= by oe is iss a cae a idee. re ee shane ma ne

git : oe aay Of Peete wae hy 4 | aoe Dr ae | - reball noe ‘Ny vse ae

tc Mosk" e Q ; : y AAS ‘4 it evr? Ae Leedq) qa

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af ce Lovo coy aera. €n Gis

. mo

i

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4 ; : C7 ie 5 Ae BO34 VOR ay eee

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sigad “lomo ah bestia ye +i8ah Bai rea

oy Seen ee SS ee seeeeneme= y ; afi Mare 4 (x 4

a) > eS rive

32

Wlost in the process of technique, and do not show in the mounted

specimens. A few were found in sections and measured 0.014 mm in length. The acetabulum is in the median line, one sixth to one

seventh of the length of the body from the anterior end- It is

circular, 0-14 mm in diameter. The oral sucker is sub-terminal

in position and measures 0-14 mm in diameter, being the same size and shape as the oral sucker. Here aS in all distomes, there is @ true oral sucker and not a mere sucking mouth funnel such as occurs in the Aspidocotylea.

The oral sucker is followed by the pharynx, a spherical muscular structureO.07 to 0.08 mm in diameter. The esophagus is &@ short tube, 0.05 mm long and 0-025 mm in diameter,leading from the pharynx to the diverticula of the intestine. The esophagus is longer proportionately in immature worms than in the adult con- dition. The intestine is bifurcate, the diverticula meeting anteriorly at an acute angle, and extending to the posterior end of the body. There are only slight variations in caliber, and these are probably due to pressure from the genital organs, or the contraction of the musculature of the body. On either side of the body the branches of the intestine lie between the uterus and the vitellaria, forming lines of separation between these organs.

The wall of the intestine is composed of scant longitudinal and circular fibers, while the lumen is lined with the usual digestive epithelium. The caeca were empty with the exception of a few

partly decomposed fragments.

| |

Ligon mie se

oe me et as Ee

Aga x ba | , a ii r wf ce % Fath ice. =p) E wy be. is yes eer y me arc 1a fh. Deer sie we (2: ig ars po 7 a } aah i; era? a TA i P e* e Liga yh = | - heme | “a t ‘vg { EY 7a, P r ako , f mh ; f ° vG Latte oo 7 x | Us 04 ame iia . - +) 3 , a SLES 8 Prt e ¥ ca Ys i) pele Fi t f . - ; 1 ra! i 7 red?’ 2 peat Ena ; © Sy). RM OBIE AtT oF Ge ‘aadat trav ‘te nodonee Py t £ ant af | fi ; ay va ae! oe * |

fe tly! ok icin eres "

Pe a RG A eh a ee ee a gah, bes on

edly haga

33

Reproductive Organs. (Figs.22;24) The ovary is a broadly

Oval structure, slightly left of the median line, about three eighths of the length of the body from the anterior end. It is 0.147 mm by 0-117 mm in the smaller specimena and 0-176 by 0-147 mm | in the largest ones. The long axis is parallel to that of the body. The oviduct arises from the median, dorsal, posterior part of the ovary and :soon expands into the fertilization space. Then on turning ventrad, it receives the common vitelline duct, and

this entire region is surrounded by the large secretive cells of the shell gland. After receiving the efferent ducts of the shell gland, Laurer's canal is given off. It turns posteriad and dorsad, opening to the surface 0-15 mm caudad of the ovary. This canal is 0.004 mm in diameter and was empty in the sectioned worms.

The uterus is a much coiled tube extending caudad as far

as the anterior testis, where it turns cephalad and continues as a

convoluted tube to the genital pore. In most of the specimens, it extends caudad on the left side of the body, returning on the ‘right side. In several specimens however, after extending about one third of the distance to the testis, it crosses to the opposite Side of the body, passing posteriorly on the right side, and after teturning on the left sice to the point of crossing winds anterior- ly on the right side of the body to the common genital pore. The uterus lies almost entirely between the intestinal caeca, but in specimens that are very much congested with eggs, the coils of

the uterus may overly the intestine half the distance from the Ovary to. the anterior testis on one or both sides.

The vitellaria are arranged in nine lobes on the right,

and twelve lobes on the left side of the body, laterad of the

oH i matics witht dat \taagiteat ree) = 7 i e ; 'DGn Sut ; +h (ein

ae

Di ue ote Lt ee To pbs

deem Vs 2 eee | an ts

5 Le sgn: Vaan tae hag eg!

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Pe ei.

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a)

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34

intestinal caeca. Often the separate lobes are not distinct, but

are massed together and overlap. Anteriorly they extend about

one third of the distance from the ovary to the acetabulum, termin- |

just cephalad of the posterior end of the cirrus sac, and poster-

iorly they extend about five sixths of the distance from the ovary

to the anterior testis. There are 20 to 40 follicles in each

lobe, and in stained material, the nuclei of the cells are con-

Spicuous among the yellow vitelline globules.

The genital pore (Fig-28) is on the ventral surface, just anterior to, and slightly left of the acetabulum. The opening of

| the uterus is anterior and slightly at the left of that of the

| cirrus sac.

The eggs occur in enormous numbers, filling the uterus.

They average 0.051 by 0-015 mm in size, but those near the ovary

are more spherical, being 0-031 by 0-019 mm. In most of the eggs

the reagents had caused the shell to collapse, and in many cases

| the cap had come off. The contents were granular in appearance.

There are two testes; They lie one behind the other at

the posterior part of the body, altho in some cases they are

Slightly on opposite sides of the median line. They are close together being only only 0-O05to 0.1 mm apart, but did not overlap

jin any case. The testes are about the same size and vary from

0.2 to 0.29 mm in length by 0-16 to 0.24 mm in width. In one immeture worm there were three testes (Fig.25), the posterior one being of normal size and in the usual position, while there were

two anterior testes, slightly smaller than normal, one on either

side of the median line.

ee ; ' Ap opnep-pt ee ye £4:

} I - ; eye y ) Pee eee etary anit Ty i

| | : >! o wheekeveri ‘ota be rr | ay ans. stant b iteat. oy

yam don tote; ;

a he as A be rie lle

ae ote s ie

' = Wea : va 4a PLEO! ! VI a aged,

a

7 hd

: ¢ Be Jed. Bai, ; wT oe Fees ila. ve et ay

7 | : { . a 14 ; CJ hi EO >, MORON LAM 9 Sia + « > wy . ! , e pQaryae rt ui Pas SPOT td i" F Pt SOT ST SS SS ete eee <r gatis y ty = id NSS) w A i } of it ae ls eS Sala

35 The duct from the anterior testes arises at the anterior ventral margin and runs cephalad ventrad of the caecum of the left side. The duct from the posterior testes arises at the anterior lateral margin and passes cephalad ventrad of the caecum of the right side. The sperm ducts pass cephalad on the median sides of

the diverticula, becoming more median and dorsal as they pass for-

ward; at the region of the ovary they cross dorsal to the excretory |

tubes and then pass on the median side of these tubes to the post- erior end of the cirrus sac, where they unite to form the seminal vesicle. The cirrus sac (Fig.26) is 1-18 to 1-295 mm in length and 0.088 mm wide at the widest part. It extends caudad from the genital pore about three fourths of the distance to the ovary. The two sperm ducts unite at the posterior end and form a large, dilated seminal vesicle which lies entirely within the cirrus sac. This seminal vesicle stains heavily, due to the masses of sperm- atozoa it contains, Anteriorly the tube is reduced in diameter and passes thru the region of the prostate gland, being surrounded | by the large cells of that organ. Then after a much coiled portio it passes es a straicht tube to the pore The posterior tip of the body is often slightly invagin- ated, the excretory pore being situated at the base of this shallow invaginatione There is a single large median collecting duct, | which extends cephalad to the region of the ovary where it divides into two branches, extending one on either side to the region of the genital pore. From these, smaller branches were traced anter-

iorly but the smaller ducts of the excretory system were not follow- ed.

;) ae (i ae Ree eat). ah) . nae e é a an cy a> ns # ay me sot hoo TOE Ot ie Rie wae cee

med vt hoereny (od erie

I My ; tary 4 = ® my iy ety MD : ; | yu,

¥ >

os

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: ; oh

j i .

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' Pe si Ate. e By 7 fi id f f 4 at Lita eh alee 5 hae

356

In the material of this species there were many immature

| forms. The smallest mounted (Fig.23) is 0-65 mm in length and

6.16 mm in width, the younger worms are wider proportionately than the adults. In this specimen the oral sucker is 0-025 mm in dia-| meter and is followed by an esophagus of equal length, the esoph- agus being more prominent and distinct than in the adult. The caeca of the intestine extend to the posterior end of the body. The excretory system is well developed and similar to that of the adult. The acetabulum is very small, 0-03 mm in diameter, and the anlage of the cirrus sac and those of the ovary and testes are merely masses of heavily staining cells, those which are to form the testes not yet having become separated into two parts.

| In a larger specimen (Fig.27) 1.7 mm long, the body has

become more elongatee It is O-2 mm wide at the widest part. The suckers have increased in size, the oral to 0-09 mm and the acetabulum to 0.63 i te diameter. Simultaneous with these changes, the anterior part of the intestine has taken on the char- acteristic shape of the adult. The ovary has assumed definite Shape, the testes have now become separates as two circular struct- ures and the cirrus sac is well defined at the anterior end altho the. posterior end is extended as line of deeply staining cells, reaching to and apparentlyconnected with the ovary. No traces

of uterus or vitellaria could be distinguished.

in

A brief specific diagnosis of Tecorti.

Sizes; 4 to 7 mm long, greatest width 0-5 mn.

Shape; elongate, flattened dorsoventrally.

Cuticula;spined.

Oral sucker; sub-terminal, 0-14 mm in diameter.

Acetabulum; one sixth to one seventh of length from anterior end.

Esophagus; presente

Ovary; 0-15 mm in diameter, three eighths of length from anterior ende

Laurer's canal; present.

Uterus; coils usually between caeca, crossing in some cases.

| Cirrus sac; extends 2 of distance from acetabumum to ovary.

Vitellaria; 9 lobes on the right, 12 on the left side of body.

Genital Pere: anterior and left of the acetabulum. Eggs; numerous, 0.031 mm by 0-015 mm.

Stossich (1904), Goldberger (1911), and Barker and Covey (1911) have compiled careful and detailed keys for the identifica-

tion of the species of the genus Telorchis. Their work is so complete that further comparisons in this paper are unnecessary. T.corti agrees closely only with T.eaculeatus von Linstow ma f.solivagus Odhner, the nearest resemblance being with T.

aculeatus. However it can not be assigned to either of these

Species because of certain specific differences.

A comparison of the Species can best be made in a

tabulated form.

ie |

an \ ai “hl ae ade. oe as) . =) iy i) ee RX Feegeh Vols eee ehh Oy Loe 7 eT ae 4 De as a hed 2 ae dipnistehn oo Nat .y er ale 2 | fo (PTS errs insite: ating ; Aduvt . ae tay Seg gael’ Oh es ae r wh oh SORE eo over 4 Ka onety . me tery , { ; ite ae rel . rodet : “oT gy 7 rsa 3 . _ oo i , l i nsf ' - . ot buvee a “4 - * i J i] a] ; ‘Ate = 3 oh NOt bs AiG dy eae? ¥ ae dagen 6.8 af vag!) sPASAD ; 6 } 4 La hy } : Ses i . es Yr Oe Vy : a Pe 7 ys AUR

PRCT MRS en + ee eer

Characters Length Width

Oral sucker Acetabulum Pharynx Esophagus

Ovary

Laurer's canal

Uterus

Testés

Cirrus sac

Genital pore

Vitellaria

Eggs Host

T.eaculeatus

5 to 7 mm

L mm 0.16 to 0.24mm 0-14 to 0.18mm O-11 mm 0-13-17 mm

O.17 mm in dia.

at posterior end of cirrus

in median line .

2

right limb descen ding, left Limb

ascending.

0-27-0-36mm separated

T.solivagus

4 to % mm

0-8 Go 0.9 mm

0-19 to 0.24mm 0.28 mm 0-146 mm 0.35 mm 0-27 mm in dia. under posterior

end of cirrus left of median

?

crosses

0.5-0-42mm contiguous

fT corti

4-tTo 7 mms

0.4 to 0-5 mm. O- 14 mm in dia. Oel4 mm tt WW . Orers, to 0.8mm MS 0.5 mm long. 0-15mn Te9 a diae O.3mm behind post end of cirrus. left of median.

present.

crosses

0.2-0.25mm separated

reaches to ovary/extends posterior/extends 2 of

anterior to acetabulum

8-9 on right 12-13 on left

0.046-0 019mm

Testudo graeca Europe

to ovary

anterior to acetabulum

indefinately arranged

0.03-0.015mm

Clemmys caspia Europe

length to ovary.

anterior and left of acetabulum.

9 on right side 12 on left side.

0.031-0-015.

M. leseurii Ne Ae

The forms are about the same length, Tecorfi being

narrower and thicker than the others;

are smaller in T.corti and the esophagus is shorter.

or absence

character, ure of the

The cirrus

but both conditions are found in T.corti.

the oral sucker.and pharynx

The crossing

of crossing of the uterus has been used as a specific

The struct-

ootype is unknown in Teaculeatus and T.solivagus.

sac is shorter in T.corti, extending only three fourths

A A A A TCT eA A Seeeeemnenemnetinedtieimediineeemmt LS A REY

ian #4) dae ¢:

132 0e hae outeeie lal

SD I nh hehe ae ney mei ey

39 of the distance from the genital pore to the ovary, while in the other forms it reaches to the ovary. The arrangement of the vitellaria is similar to that in T.aculeatus but different from T, solivagus. The ecgs of T.corti are about the same size as those of solivagus, but are smaller than those of T.-aculeatus.

Classification. In 1899, Luehe and Looss, working independently, each created a new genus, Telorchis , to contain a certain group of distomes from reptiles, which seemed to have common characters, so constant and unique as to warrant their sep- aration into a new genus. | Luehe took Distoma clava as a type, and included the species poirieri (gelatinosum), linstowi (aculeatus), ercolanii ( nematoides), bifurcus and pleriticum. As generic characters he gave," Testes median, one behind the other, at the posterior end of the body, cirrus sac very long, opens anteriorly and at the left of the acetabulum. Ovary immediately behind the cirrus sac, uterus coils in the body anterior to the testes, vitellaria of many foll- icles occupy sides of the body, caeca extend to the posterior end of body; with the exception of T.poirieri , body is spined; excre- tory bladder long, branching anteriorly; oral sucker larger than acetabulum in ercolanii, usually about equal size".

Looss took D. linstowi ( aculeatus) as a type, and in- cluded the species poirier, nematoides and arrectum. As generic characters he designates the following,™" Body long, smaller anter- iorly; spines present or absent; short esophagus, diverticula extending to the posterior end of the body; genital opening anter-

ior to the acetabulum; cirrus present, long, extending far poster-

iorly; testes behind one another, at the posterior end of the body;

f . eee 4s TEE Sat OF va se

iit! Te aRae :

' v0 4 Bi: * , i/ hog @

“F inh

tHe ote ‘Sur ae

40 ovary far distant, somewhat lateral, lying in the region at the end of the cirrus sac; Laurer's canal present."

Because of the difference of T.clava from the other mem- bers of his genus Telorchis, Luehe (1900) divided the genus into two sub-genera, Telorchis, with T.clava as a type, and Cercorchis, with T.linstowi as a type. To the sub-genus Telorchis he ascribes those forms with no esophagusand with uterine coils not confined between the caeca, while to Cercorchis the forms with an esophagus and with the coils of the uterus confined between the intestinal diverticula. Regarding these identifying characters, he says the Seeaeeaae or absence of an esophagus is of major importance.

Looss (1902) characterized Luehe's genus Telorchis as an unnatural group, but fails to find adequate ground to justify the division into two sub-genera. Luehe certainly was unfortunate in the selection of identifying characters for his sub-genera, for an intracaecal coiling of the uterus is not necessarily assoc- iated with the presence of an eSopnhagus, since in T-pleroticus and f.bifurcus the coils of the uterus are confined between thenand the esophagus is absent. In his separation into sub-genera Luehe's sub-genus Cercorchis corresponds identically with Looss genus Telorchis, except that Looss did not make the error of associating the presence or absence of an esophagus, which itself is a variable condition, with an indefinite character like the extent of the coiling of the uterus. The extent of coiling depends to a large extent on the fecundity of the worm; in Tecorti, in which the uterine coils are usually contained within the diverticula, in

specimens that are very much congested with eggs, the coils of the

uterus overly the intestine.

q ]

i. ;

ae 4a * 1) ad?

1h eae

fortes vice " Lanka} Le. ert

ve ey!

2

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é Ar et : eltges.

ay Te 5 ; nd * ad bt ~t ie a t 6 i i > Oa ite, . rere: ait ? j st anenew ek bd

41 Luehe's description of Telorchis was published in the Zool. Anz. of Dec.28, 1899, and Looss': account was printed in the

Zool. Jahrb. of the same date. Most workers have credited the

genus to Luehe, altho Odhner(1902) in his description of T.solivag-|

us ascribes the genus Telorchis to Looss.

As regards the two sub-genera of Luehe, since the extent of the coiling of the uterus is not associated with the presence or absence of an esophagus, the coiling of the uterus must be abandoned as a diagnosing character, and the only difference between the sub-genera becomes that of the presence or absence of an esophagus. Such a variable and uncertain character as the presence or absence of an esophagus certainly does not warrant the

separation of a genus into two sub-genera.

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ant

Bibliography. Barker, F.De and Covey, GW. 1911. A New Species of Trematode from the Painted Terrapin, Chrysemys marginata.

Nebraska Univ. Stud. ,11:472-961;3pl.

Braun, M. 1890. Plathelminthes Minot 1877- In Bronn, H.-G. Xlassen|

und Ordnungen der Thier-Reichs. vol-4,pt.1,A,:303-925;28pl. 1901. Trematoden der Chelonier. Mitt. Zool. Muse Berli 2:13-20;lpl.

Goldberger, J. 1911- On Some New Parasitic Trematode Worms of the Genus Telorchis. Bull. No.71, Hyg-e Labe, UeS- Pube Health & Mar.-Hosp. Serv., Washe :36-48.

Jagerskidld, L.A. 1899. Ueber den Bau von Macraspis elegans. Ofvers. Vet. Akad. Fordh., 3:197=214;1pl.

Kofoid, C-A- 1899. The Identity of Cotylaspis pauaiew he Leidy and Platyaspis anadontae Osborn, JZool- Bull.,5:179-185.

Leidy, J- 1857. Observations on the Entozoa found in Naids. Proce Acad. Nat. Sci. ,Phila.,.1857:18.

e 1858. Contributions to Helminthology. Proc. Acad. Nat. Sci. Phila. ,1858:110.

e 1888. Entozoa of the Terrapin. Proce Acad. Nate. Sci. Patia., 1888:127.

Looss, A. 1899. Weitere Beitréce zur Kenntniss der Trematoden

~

Me Aecypienee Zook. Jeux. u2:521-794; 6pl. 1902. Ueber Neue und Bekannte Trematoden aus Schild- kroten. Zool. Jarbe,Syst., 16:411-794. Luehe, M. 1899. zur Kenntnis einiger Distomen. Zool. AnZe,

22: 624-539.

1900. Ueber einige Distomen aus Schlangen und Eidech-

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ig %; SOC ame ‘herd rey ea TM" menda te: out aR st

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2 nee ies suit iy

; cree ore ‘Lait ‘ul ea

sene. Centr. Bakt.,1 Abt. ,28;524-539. & | Monticelli, Ss 2692. Cotylogaster michaelis. Fest. Leukart: 166-214; 2pl.

Nickerson, W-S. 1902. Cotylogaster occidentalis nesp.- and a Revis-]

= a ion of the family Aspidobothridae. Zool. Jahrb#,5:597-624;2pl.

| Odhner, Th. 1898. Ueber die geschlechtsreife Form von Stichocot- Jyle- Zool. Anz.-,21;509-513. 7 Osborn, H.-L. 1904. On the Habits and Structure of Cotylaspis insignis Leidy. JZoole Jahrb. ,21:200-240;3pl. Poirier, 1886. Trematodes Nouveaux ou peu Connus. Bulle Sees Phil. Paris, 5:10;lpl. |} Stafford, J.- 1896. Structure of Aspidogaster conchicola. Zool. Jahrb. ,9:477-537;4pl. Stossich, Me 1895. I distomi dei rettili. Boll. Soc. Adriat. di Sc. nat. in Trieste,16:2135-293;2pl. 1904. Alcuni distomi della collezione elmintologica del museo zoologico di Napolise Mus.Zo00l- Univ. Napoli,1:1-10. Taschenberg, E.0- 1879. Zur Systematik der Monogenetischen Trem- atoden. zeit. gesammt. Naturwissensch. ,52:232-265- Willemoes Suhm, R.- 1872. Zur Naturgeschichte des Polystoma integ- errimum and des P.eocellatum Rud. Zeit. wiss. Zool. ,22:29-39. Wright, R.R. 1879. Contributions to American Helminthology.

Proc.Canad. Inst., yf Beha Be

4eller, Ee. 1872. Untersuchen ueber die Hntwicklung und den Bau des Polystomum integerrimum Rud. Zelty wiss. Zool. ,22:1-28. 1876. Weiterer Beitrag zur Kenntniss der Polystomen.

Zeit. wiss. Zool. ,27:2358-275-4

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Abbreviations used in Plates.

a acetabulum e esophagus f fertilization space g genital pore i intestine 1 Laurer's canal Im limiting membrane m mouth mt metraterm mo marginal organ os oral sucker fe) ovary oc eye od oviduct p prostate gland Sv seminal vesicle Sp septum } t testis } ut uterus Vv vitellaria vas deferens

Explanation of Plates. Plate I

Figel.- Cotylaspis rhadina somewhat extended, dorsal view.x40.

Fig.2.

Cotylaspis rhadina, ventral view, showing the position

of the marginal organs and the divisions of the adhesive disc. x40.

Fig.3. Contracted specimen of C- rhadina, dorsal view. x 40.

Fig.4. Diagrammatic representation of the excretory system in C. rhadina, dorsal view. x 40.

Fig-5. Cross section thru the region of the ovary of C. rhad-

ina, showing a section of an ege in the uterus. x 85.

Fig. Be

Frontal section thru the adhesive disc of C. rhadina,

showing the musculature. x 95,

Fig.7. Reconstruction of the reproductive organs of C. rhad-

ina,from frontal sections. x 80.

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Fige8. Sagittal section thru the anterior end of Ce. rhadina,{

showing musculature, digestive, reproductive, and marginal organs. The section is at the right of the median line. x 200.

Fiz.9. Sagittal section thru the dorsal body wall,showing the cuticula, the muscle layers, a section of body parenchma, and the wall of the digestive tube. x 550.

Fig-10.- Frontal section thru the mouth funnel of C. rhadina, showing the absence of an oral suckere x 300.

Fig-ll- Frontal section thru the genital pore, showing the

relations of the cirrus sac and metraterme x 180.

Plate III

Fige12.- Polystoma orbiculare, ventral view. x 20.

Fig-13. Frontal section thru the dorsal commissure, showing the large ganglion cells, x 60.

Fig.14. Frontal section thru the pharynx and mouth funnel. The esophageal nerve ring and anterior vitellaria are shown. x 60-

Fig-15. Cross section P-eorbiculare, at the region of the genital pore. x 60.

Fig-16.- Cross section thru the testes. x 60.

Fige-17- Cross section thru the anterior bothria. x. 60.

Figel8. Hook from the genital coronete x 550.

Fig-19. Hook from the base of a bothrium. x 550.

Fig.20. Reconstruction of the reproductive organs of P. orbiculare, ventral view from frontal sections. x 40.

Fig-21.- Polystoma hassali, ventral view. x 40-

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Plate IV Fig.22-. Telorchis corti, reconctruction of the female organs at the region of the ovary,from sagittal sectionse x 120. Fige23- Immature worm, 0-65 mm longe x 100. Fig.24.- Telorchis corti, ventral view. x 15.

Fige25- Abnormal specimen, three testes. x 65.

Fig.26. Cirrus sac. x45.

Fige27. Immature worm, 1-7 mm long. x 45.

Fig.28. Cross section thru the genital pore. x 100.

Fig.29. Cross section thru the posterior part of the ovary, showing the shell gland, the anterior part of the uterus on the right and the coils returning on the left, the excretory tubes are

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