STUDIES ON THE TREMATODE PARASITES OF DUCKS IN
MICHIGAN WITH SPECIAL REFERENCE TO THE MALLARD

BY
W. CARL GOWER

A THESIS
P r e s e n t e d t o the G ra duate School o f M ichigan S t a t e
C o l le g e o f A g r i c u l t u r e and A p p lie d S c ie n c e i n P a r t i a l
F u l f i l l m e n t o f R e q u irem e n ts f o r th e
Degree o f D o c to r o f P h i l o s o p h y

Departm ent o f B a c t e r i o l o g y , H ygiene , and P a r a s i t o l o g y
and D epartm ent o f Zoology
E a s t L a n s i n g , M ic higan
1937

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•Foreword and Acknowledgments

T his s t u d y was a t f i r s t i n t e n d e d to be an i n t e n s i v e
s t u d y o f t h e s p e c i e s o f P rosthogonimu3 o c c u r r i n g i n s o u t h ­
e r n M ic h ig a n .

Soon a f t e r b e g i n n i n g , however, i t became

a p p a r e n t t h a t , p e r h a p s , a more i m p o r t a n t c o n t r i b u t i o n m ight
b e made a t t h i s tim e t h r o u g h a s u r v e y o f th e tr e m a t o d e s
found i n ducks i n M ic h ig a n .

The obvious need f o r , and th e

e x c e l l e n t o p p o r t u n i t y a f f o r d e d a t th e W. K. K e llo g g B i r d
S a n c t u a r y f o r such a s t u d y , were i n s t r u m e n t a l i n t h e c h a n g e .
Too, a l t h o u g h P ro sth ogonim us i s h i g h l y p a t h o g e n i c i n c h i c k e n s ,
c e r t a i n o t h e r t r e m a to d e s seemed to be o f more im p o r ta n c e i n
ducks.
The t h e s i s i s d i v i d e d i n t o t h r e e main p a r t s :

Intro d u ctio n

P a r t I , d e a l i n g w i t h t h e forms s t u d i e d i n d e t a i l a t t h e
S a n c t u a r y ; and P a r t I I , which c o m p ris e s a key to t h e g e n e r a ,
a d e s c r i p t i o n o f t h e f a m i l i e s and g e n e r a , and a c h e c k - l i s t
o f th e s p e c i e s o f t r e m a t o d e s t h a t have b ee n r e p o r t e d from
N o rth American d u c k s .
The W* K. K e llo g g B i r d S a n c t u a r y o f M ichigan S t a t e
C o l l e g e o f f e r s an e x c e l l e n t o p p o r t u n i t y to p u r s u e a s t u d y
o f th is kind.

The S a n c t u a r y i s l o c a t e d n e a r B a t t l e C reek,

and c o m p ris e s a b o u t 80 a c r e s which i n c l u d e s W i n te r g r e e n Lake,
t o t a l i n g a b o u t 20 a c r e s , and a b o u t f i v e a c r e s o f s w a l e .

ii
A r e l a t i v e l y h i g h duck p o p u l a t i o n i s p r e s e n t d u r i n g mo3t
o f t h e y e a r , th e s m a l l e s t number b e i n g i n e a r l y summer and
m id-w inter.

A c a p t i v e f l o c k o f swans, g e e s e , and ducks

i s m a i n t a i n e d , b u t th e m a j o r i t y o f t h e ducks p r e s e n t a r e f u l l ­
winged m a l l a r d s and b l a c k d u c k s .

An a v e r a g e o f from SO to 35

m a l l a r d s n e s t e v e r y y e a r a t th e S a n c tu a r y *

Experim ental pens,

b o t h a t t h e Lake and away from i t , have b e e n a v a i l a b l e .
Through th e h a n d l i n g o f ducks f o r b a n d i n g , i t h a s been p o s s i b l e
to examine some t h i n and s i c k b i r d s .

With t h e c o o p e r a t i o n

o f Dr. H. R. H unt, head o f t h e Department o f Zoology, a
c o m p le te t e c h n i q u e l a b o r a t o r y was s e t up a t th e S a n c t u a r y ,
so t h a t i t has b e e n p o s s i b l e t o do e v e r y t h i n g e x c e p t l i b r a r y
work t h e r e .
In March 1937, th ro u g h Mr. E. D. R u h l , c h i e f o f t h e
Game D i v i s i o n , M ichigan Departm ent o f C o n s e r v a t i o n , one
hundred m a l l a r d ducks were o b t a i n e d f o r e x p e r i m e n t a l p u r ­
poses.

They were o f th e " w i l d ” t y p e , and were used i n

p la ce o f dom esticated m a lla rd s .
A p p r e c i a t i o n i s e x p r e s s e d h e r e f o r th e many p r i v i l e g e s
ex ten d ed by Dr. M. D. P i r n i e , D i r e c t o r o f th e W. K. K e llo g g
B i r d S a n c t u a r y , who, t h r o u g h a f e l l o w s h i p h as made t h i s
s t u d y p o s s i b l e , and who has o f f e r e d many h e l p f u l s u g g e s t i o n s
t h r o u g h o u t t h e work on th e h a n d l i n g , r e a r i n g o f d u c k s , e t c .
To Dr. W. L. C h a n d l e r , A s s o c i a t e i n P a r a s i t o l o g y , M ichigan

ii i

S t a t e C o l l e g e , u n d er whose g u id a n c e t h i s work h a s b een
d o n e, t h e w r i t e r i s e x t r e m e l y in d e b t e d f o r many h e l p f u l
s u g g e s t i o n s and c o n s t r u c t i v e c r i t i c i s m t h r o u g h o u t .
th e f o l l o w i n g p e o p le th a n k s a r e a l s o e x p r e s s e d :

2o

D r. H. R.

H u n t, head o f D epartm ent o f Z oology, Dr. H. J . S t a f s e t h ,
Departm ent o f B a c t e r i o l o g y , and Mr.

W. S t a c k , D e p a r t ­

ment o f Z oology, a l l o f M ichig an S t a t e C o l l e g e , who w i t h
Dr. R i r n i e and Dr. C ha ndler have a c t e d a s th e g u id a n c e
co m m itte e ; Mr. H. D. R u h l , C h i e f , Game D i v i s i o n , M ichigan
D epartm ent o f C o n s e r v a t i o n , f o r f u r n i s h i n g one hundred
ducks a s w e l l a s o t h e r f a v o r s , and t o Mr. Homer B r a d l e y , o f
t h e W. K. K e llo g g B ird S a n c t u a r y s t a f f , who tu r n e d o v er to
me t h e v i s c e r a o f f o r t y ducks ta k e n a t Houghton Lake and
Muskegon Lake.

Table o f C o n te n ts

Introduction
I n t r o d u c t o r y r e m a r k s ........................................................................1
The D ucks..................................................

*.3

C o l l e c t i o n and P r e p a r a t i o n o f S pecim ens.............................5

Part I .

Trematodes from ducks i n Michigan

Z y g o c o ty le lunatum ( B i e s i n g , 1 8 3 6 ) ......................

9

P r o s thogonimus m a c r o r c h i s Macy, 19.34................................. 14
P ro stho gonlm us r u d o l p h i i S k r j a b i n , 1919.......................... 18
P r o s thogonimus a n a t i n u s Markov/. 1 9 0 2 ................................. 19
Amphimerus e l o n g a t u s n . s p ..................................................... . 2 4
M a r it r e m a n e t t a e n . s p .................................................................32
S p e lo tr e m a pygmaeum( L e v in a e n . 1 8 8 1 ) ................................... 38
Typhlocoelum cucumerinum ( R u d o lp h i, 1 8 0 9 ) ............... . . . 3 9
Typhlocoelum c.ymbium ( B i e s i n g , 1 8 5 0 ) ...............

39

N o to c o t y l u s u r b a n e n s i s ( O o r t , 1 9 1 4 ) ................................... 47
Echinostoma r e v o lu tu m ( F r o e l i c h , 1 8 0 2 ) .............................49
S te p h a n o p ro r a s p l n o s a (Odhner, 1 9 1 0 ) ................................. 52
S p a e r i d i o t r e m a g l o b u l u s ( E u d o l p h i , 1 8 1 4 ) ........................ 52
C o t y l u r u s f l a b e l l i f o r m i s ( F a u s t , 1 9 1 7 ) . . . . . ................. 53
L e u c o c h lo r ld io m o rp h a m ao ro o o ty le n . g . , n . s p .............59

Part I I .

Key to t h e g e n e r a , and d i a g n o s i s o f the

f a m i l i e s and g e n e r a , and l i s t s o f the s p e c i e s o f
t r e m a t o d e s r e p o r t e d from N orth American Bucks.

Key to t h e g en e ra o f tr e m a to d e s r e p o r t e d from
N orth American ducks ...........

.62

D i a g n o s i s o f f a m i l i e s and g e n e ra and l i s t s o f
s p e c i e s o c c u r r i n g i n North American d u c k s ......................66

Trematodes o f ducks a r r a n g e d a c c o r d i n g to h o s t .......................117
Summary and C o n c l u s i o n s ..................................................................... 128
L ite ra tu re .

...................................................................................

I n d e x .............................................................................................................. 144

129

Introduction
Among t h e e a r l i e s t d e s c r i p t i o n s o f tr e m a to d e worms
a r e four t h a t a re p a r a s i t i c in ducks.

Bloch d e s c r i b e d

Hypoderaeum conoideum i n 1782, and a l i t t l e l a t e r (1789)
F ro e lic h described C a ta tro p ls v e rru o o sa .

E a r l y i n th e

n i n e t e e n t h c e n t u r y th e work o f Z e d e r , F r o e l i c h , and
e s p e c i a l l y R u d o l p h i , added a number o f new s p e c i e s to the
list.

More r e c e n t l y t h e works o f L o o s s , Lube, B r a u n ,

D i e t z , S k r j a b i n , Odhner, and N i c o l l have added a g r e a t d e a l
to our knowledge o f th e t r e m a to d e s o f t h i s group o f b i r d s .
L i t t l e work has b ee n done i n North America on th e treraato d e s o f d u c k s , a s s u c h , and t h a t i s h i g h l y s c a t t e r e d .
T h is s t u d y h a s been done w i t h t h r e e o b j e c t i v e s i n mind:
( 1 ) to make a s com plete a s u r v e y a s p o s s i b l e o f th e tr e m a ­
t o d e s found i n M ic higan d u c k s , (2) to d e t e r m i n e , a s f a r
a s p o s s i b l e , h o s t - p a r a s i t e r e l a t i o n s h i p s and im p o r ta n c e
o f t h e s e v a r i o u s s p e c i e s , and (5) to a s sem b le from th e
l i t e r a t u r e , a l l th e forms t h a t have b een r e c o r d e d from
N o rth American d u c k s .
The r e l a t i o n s h i p s o f many o f t h e s e s p e c i e s and g e n e r a
a r e n o t c o m p l e t e l y u n d e r s t o o d , and c o n s e q u e n t l y we f in d
them c l a s s i f i e d i n a v a r y i n g number o f f a m i l i e s .

The

a r r a n g e m e n t used h e r e was a r r i v e d a t a f t e r a s t u d y o f th e
e x t e n s i v e works o f P o ch e, S p re h n , and Fuhrmann, a s

(2)

w e l l a s o t h e r more d e t a i l e d s t u d i e s on f a m i l i e s .

The

M i c r o p h a l l i d a e and C y a t h o o o t y l i d a e a r e c o n s i d e r e d v a l i d
fam ilies.
P a r t I d e a l s w i t h the forms r e c o v e r e d from ducks i n
M ic h ig a n .

One new genus and t h r e e new s p e c i e s a r e d e s c r i b e d .

Under each s p e c i e s d i s c u s s e d a r e g iv e n t h e h o s t s , l i f e - h i s t o r y n o t e s , and som ething o f t h e economic s t a t u s a s w e l l a s a
d e s c r i p t i o n i n some o f th e o a s e s .

P a r t I I i s a key to t h e

g e n e r a and a c h e c k - l i s t o f t h e s p e c i e s o f t r e m a to d e s r e p o r t e d
from N orth American d u c k s .

I n making up t h e c h e c k - l i s t , t h e

g e n e r a l o u t l i n e o f Bans on1s (1909) "Taenoid C e sto d e s o f N o rth
American B i r d s " h a s b ee n f o l l o w e d .

The key and c h e c k - l i s t

have b e e n compiled by means o f an e x t e n s i v e s u r v e y o f th e
l i t e r a t u r e p l u s t h e work d e s c r i b e d i n P a r t I .

The l i s t con­

t a i n s 76 s p e c i e s o f d i g e n e t i c tr e m a to d e s b e l o n g i n g to 42
g e n e r a and 17 f a m i l i e s .

The f a m i l y and g e n e r i c d i a g n o s i s a r e

g i v e n i n each c a s e , a f t e r which t h e s p e c i e s a r e l i s t e d w ith
th e r e o o r d e d l o c a l i t i e s and the h o s t s .
two i m p o r t a n t r e f e r e n c e s a r e g i v e n .

In each e a s e one o r

No a t t e m p t i s made t o

l i s t t h e synonyms o f t h e s p e c i e s , and s u b f a m i l y d e s c r i p t i o n s
have b e e n o m i t t e d .

I t w i l l be noted t h a t many o f t h e s e s p e c i e s

have n o t a c t u a l l y b ee n r e p o r t e d from ducks t a k e n i n North
A m e r ic a , b u t t h e y have b een r e c o r d e d from s p e c i e s o f ducks
which o c c u r h e r e , ( i n some c a s e s European s u b s p e c i e s ) .

While

(3)

suc h a l i s t a s t h i s could p r o b a b l y n e v e r be c o m p l e t e , i t i s
hoped t h a t i n b r i n g i n g t h i s i n f o r m a t i o n t o g e t h e r i t may be o f
v a l u e i n f u t u r e s t u d i e s on the tr e m a to d e s o f b i r d s .

The Bucks

D uring t h e c o u r s e o f t h i s s t u d y a b o u t 200 ducks have
b ee n a u t o p s i e d .

While m a l l a r d s and b l a c k ducks com prise

t h e b u l k o f t h i s number, n i n e a d d i t i o n a l s p e c i e s have been
examined.

Most o f t h e specimens were c o l l e c t e d a t W i n te r g r e e n

Lake a t th e W. K. K e llo g g B i r d S a n c t u a r y (JSalamazoo C o u n t y ),
b u t b i r d s have b ee n examined from Houghton Lake (Roscommon
C o u n t y ), Muskegon Lake (Muskegon C o u n ty ), Crooked Lake
( B a r r y C o u n t y ) , th e Seney Marshes ( S c h o o l c r a f t C o u n ty ),
Crooked Lake ( A l le g a n C o u n ty ), and Mud Lake (Washtenaw County)*
I n March, 1936, 100 one y e a r - o l d m a l l a r d s were o b t a i n e d
from t h e s t a t e game farm a t Mason, Michigan*

These ducks

were used i n v a r i o u s ways, some o f them b e i n g allow ed a c c e s s
to th e l a k e , w h i l e the o t h e r s were h e ld i n a pen e n t i r e l y
away from th e l a k e , and used more or l e s s a s c o n t r o l s f o r
the in fec ted b i r d s .

In a d d i t i o n to t h e s e b i r d s , a b o u t 25

young m a l l a r d s were h a t c h e d , and used a t d i f f e r e n t ag e s f o r
i n f e c t i o n experim ents.

The o t h e r b i r d s examined were w ild

ones o b t a i n e d by t r a p p i n g o r s h o o t i n g .

(4)

The f o l l o w i n g s p e c i e s o f ducks have b ee n examined i n this;
study:

m a l l a r d (Anas platy rh y n c h Q 3 p l a t y r h y n o h o s ) ^ . b l a c k

duck (Anas r u b r i p e s ) 2 . widgeon (Mareoa a m e r i c a n a ) , p i n t a i l
( D a f i l a a c u t a t z i t 2 i h o a ) . green-w inged t e a l ( N e t t i o n
c a r o l i n e n s e ) , b lu e -w in g e d t e a l ( Querquedula d i s c o r s ) , wood
duck (Alx s p o n a a ) , redhea d ( Nyroca a m e r i c a n a ) , oanvasbaok
( Nyroca v a l i s i n e r i a ) , g r e a t e r s c a u p ( Nyroca m a r i l a ) , l e s s e r
s c au p ( Nyroca a f f i n i s ) , r i n g - n e c k e d duck ( Nyroca c o l l a r i s ) ,
and American g o l d e n - e y e ( G l a u c i o n e t t a o l a n g u l a a m e r i c a n a ).
There i s a p p a r e n t l y c o n s i d e r a b l e c o n f u s i o n i n th e
r e c o r d i n g o f t h e names o f th e v a r i o u s duck h o s t s mentioned
in t h i s r e p o r t .

This i s d u e , c h i e f l y , to th e ch anging o f

many o f th e g e n e r i c names o f t h e s e forms i n r e c e n t y e a r s .
Many o f t h e o l d e r r e p o r t s u se Anas a s a g e n e r i c name f o r many
o f the d u c k s .

The A. 0 . U. c h e c k - l i s t sh o u ld be c o n s u l t e d

f o r th e p r o p e r name.

The names o f ducks ta k e n from the

l i t e r a t u r e have been r e t a i n e d e x a c t l y as r e p o r t e d .

1 . A c c o r d in g to D r. Glover M. A l l e n , Museum o f Comparative
Z oolog y, H arvard U n i v e r s i t y , i n a p e r s o n a l com m unication,
u s e o f d o m e s tic a a s a s u b s p e c i f i c d e s i g n a t i o n i s i n c o r r e c t ,
s i n c e d o m e s t i c a t e d s t r a i n s a r e n o t v a l i d s u b s p e c i e s . The
d o m e s tic m a l l a r d should be d e s i g n a t e d Anas p l a t y r h y n c h o s
( d o m e s t i c ) o r some s i m i l a r e x p r e s s i o n .
2. A c c o rd in g to Dr. M. D. P i r n i e (Michigan Waterfowl Manage­
m e n t, p . 12) t h e r e i s no e v id e n c e t h a t t h e r e a r e two s u b s p e c i e s
o f b l a c k duck i n M ic h ig a n .

(5)

C o l l e c t i o n and P r e p a r a t i o n o f Specimens

The r o u t i n e e x a m in a tio n o f a duck u s u a l l y f o llo w ed i n
t h i s s t u d y was f i r s t to weigh and measure th e b i r d *

The

n eck was th e n c u t open and t h e n a s a l c a v i t i e s , e s o p h a g u s ,
t r a c h e a , e t c . , were examined.

The body c a v i t y was the n exposed

and t h e v i s c e r a removed to a dish*

The body c a v i t y was c a r e ­

f u l l y examined, and th e n th e v i s c e r a s c r u t i n i z e d i n d e t a i l
under a d i s s e c t i n g m icroscope.

A l l t h e p a r a s i t e s found were

removed to a d i s h o f w a t e r , counted and g r o u p e d , a f t e r which
t h e y were f i x e d f o r f u r t h e r s t u d y .
A lth o u g h a number o f f i x a t i v e s have b een t r i e d and found
s a t i s f a c t o r y , none can compare in c o n s t a n c y and d e l i c a t e n e s s
o f f i x a t i o n w i t h B o u i n ’s p i c r o - f o r m a l s o l u t i o n .
used h o t , b u t works a l m o s t a s w e l l c o l d .

I t is be3t

I t i s n e c e s s a r y to

f l a t t e n l a r g e r worms b e f o r e f i x a t i o n by means o f two g l a s s
slid es.

A f t e r t h e worms a r e p r o p e r l y f l a t t e n e d , t h e y a r e

f lo o d e d w i t h h o t o r cold B o u i n 's f l u i d , and l e f t f o r one h a l f
t o one h o u r , a f t e r which t h e y w i l l be h ardened s u f f i c i e n t l y to
remove th e p r e s s u r e .

I t i s b e s t to l e a v e them i n th e f i x i n g

f l u i d an hour o r more a f t e r removing the p r e s s u r e , b u t even a
l o n g e r time w i l l n o t i n j u r e th e sp e cim e n .

Prom th e f i x a t i v e

t h e specim ens a r e removed to 50% a l c o h o l to which a few d r o p s
o f a s a t u r a t e d aquous s o l u t i o n o f 1100^ have been a d d e d .

(6)

A f t e r th e y e l l o w c o l o r ha3 b ee n removed (2 4-48 h o u r s ) t h e y
may be s t o r e d I n 70fa a l c o h o l , h y d r a t e d and s t a i n e d i n b u l k ,
or d e h y d r a t e d , c l e a r e d , and imbedded i n p a r a f f i n f o r s e c t i o n ­
ing.
For whole mount s t a i n i n g , M a y e r 's carmalum h a s proved
even s u p e r i o r to E h r l i c h ' s o r D e l a f i e l d * s h e m a t o x y l i n .

If

destain ed c a r e f u l l y i t gives a very d e l i c a t e d i f f e r e n t i a t i o n ,
and t h e s t a i n i s v e r y b r i g h t and p e rm e n an t.

Carmalum i s b e s t
1

d e s t a i n e d by u s i n g M ayer’s c h l o r i n e method o f b l e a c h i n g (Lee ) .
The s t a i n a p p a r e n t l y does n o t d e t e r i o r a t e w i t h a g e , and i s
e a s y to u s e .

I t may be used f u l l s t r e n g t h o r d i l u t e d w i t h a s

much a s 20 p a r t s o f w a t e r .

E h r l i c h 's acid hem atoxylin a l s o

makes e x c e l l e n t p r e p a r a t i o n s b u t was n o t found q u i t e a s con­
s t a n t i n r e s u l t s a s carmalum.

In e i t h e r c a s e th e worms a r e

p a s s e d down th e graded s e r i e s o f a l c o h o l s to 15$, a f t e r which
th e y a r e p u t i n t o t h e s t a i n *

With e i t h e r o f t h e s e s t a i n s a t

l e a s t 24 h o u r s sh o u ld be a l l o w e d , and o r d i n a r i l y any p e r io d
up t o a week w i l l n o t i n j u r e t h e s p e c im e n s .

I have found

w ith E h r l i c h 's acid hem atoxylin, t h a t a very ra p id d e s ta in i n g
w i t h a 5-1 0 $ HC1 s o l u t i o n o f 70$ a l c o h o l g i v e s b e t t e r r e s u l t s

Tl

$he t e c h n i q u e s b r i e f l y o u t l i n e d h e r e w i l l be found i n
d e t a i l i n G uyer’s "Animal M io rology" and L e e ’s "The M icromati s t ' s Vade Mecum” .

(7)

t h a n a slow d e s t a i n i n g w i t h a v e r y weak a c i d s o l u t i o n *

This

however must h e c a r e f u l l y watched and i f n o t stoppe d a t j u s t
the r i g h t moment the d e s t a i n i n g p r o c e s s w i l l he c a r r i e d too
far.

The d e s t a i n i n g i s sto p p e d hy removing th e specimen to

70$ a l c o h o l c o n t a i n i n g a t r a c e o f c o n c e n t r a t e d ammonium
hydroxide.

In t h e c a s e o f c h l o r i n e d e s t a i n i n g , i t i s s u f f i ­

c i e n t to r e p l a c e th e c h l o r i n e a l c o h o l w i t h f r e s h 70$.
A f t e r the d e s t a i n i n g i s f i n i s h e d , th e worm i s c o m p l e t e l y
d e h y d r a t e d and c l e a r e d .

I t i s f r e q u e n t l y n e c e s s a r y to f l a t t e n

o u t th e a n i m a l a g a i n , f o r i t i s o f t e n c u r l e d somewhat.

This

i s done by p u t t i n g i t u nder s l i g h t p r e s s u r e i n th e 95$ a l c o h o l ,
which h a r d e n s t h e specimen i n a f l a t t e n e d p o s i t i o n .

O rdinarily

a h o u t one hour i n the a b s o l u t e a l c o h o l i s s u f f i c i e n t to com­
p le te l y dehydrate.

As a c l e a r i n g a g e n t , x y l o l has been found

e x c e l l e n t f o r c e r t a i n s p e c i e s , and w o r t h l e s s f o r o t h e r s .

For

g e n e r a l u s e cedarwood o i l and t e r p i n e o l g i v e t h e most c o n s t a n t
resu lts.
S e c t i o n s were u s u a l l y c u t a t from 10-20 m i c r o n s .

I t is

b e s t to s t a i n v e r y s m a ll worms s l i g h t l y b e f o r e imbedding.
I r o n alum h e m a t o x y l i n fo llo w e d by a c o u n t e r s t a i n i s a v e r y
good s t a i n f o r th e d e t a i l e d s t u d y o f s e c t i o n s .

For o r d i n a r y

u s e , I have found M a l l o r y ’s t r i p p l e a v e r y u s e f u l t e c h n i q u e .
I t d i f f e r e n t i a t e s th e o r g a n s b e a u t i f u l l y , and t h e e n t i r e

(8)

t e c h n i q u e r e q u i r e s o n l y a few m i n u t e s .

E h r l i c h ' s or

D e l a f i e l d 'a h e m a to x y l in f o llo w e d by e o s i n i s a l s o u s e f u l
for se ctio n s.

PART I

tr e m a t o d e s from Bucks i n M ichigan

(9)

P am ily P aram phistom ida e F i s c h o e d e r , 1901
Genus Z y g o c o ty le S t u n k a r d , 1916
Z y g o c o ty l e lun atum ( B i e s i n g , 1856)
T his amphistome was t h e o n l y member o f th e f a m i l y
P a ra m p h is to m id a e ta k e n d u r i n g t h e c o u r s e o f t h i s work,
and from a s t u d y o f t h e l i t e r a t u r e i s a p p a r e n t l y th e
o n l y r e p r e s e n t a t i v e o f th e f a m i l y t h a t h a s b e e n r e p o r t e d
from d u c k s ,

The s p e c i e s was f i r s t d e s c r i b e d by D i e s i n g

from m a t e r i a l c o l l e c t e d i n South America from Anas
m e l a n o t u s . Anas i p e c u t i r i * Hemantopus w i l s o n i l , and
Cervus d i c h a t o m u s .

These a r e r e s p e c t i v e l y , d u c k s , a

s h o r e b i r d , and t h e South American deer*

The f l u k e s

a r e found i n t h e caecum in each s p e c i e s *

Due to th e

f a c t t h a t one o f t h e h o s t s was a mammal and t h e o t h e r s
b i r d s , i t was th o u g h t by some w o rke rs t h a t D ie 3 i n g had
made a m i s t a k e , p r o b a b l y i n l a b e l i n g th e specimens*
In 1916 S tunkard c r e a t e d th e genus Z y g o co ty le * and
p la c e d i n i t h i s new s p e c i e s Z y g o o o ty le c e r a t 0 3 a . a s
w e l l a s D i e s i n g ' s Amphistoma lu n a t u m .

Ee s e p a r a t e d the

two s p e c i e s s o l e l y on t h e b a s i s o f minor d i f f e r e n c e s in
s h a p e and s i z e o f some o f th e i n t e r n a l o r g a n s , s i n c e t h e y
w ere i d e n t i c a l i n most c h a r a c t e r i s t i c s .

S tunkard had

o b t a i n e d h i s m a t e r i a l from th e i n t e s t i n e s and c a e c a e o f
Anas p l a t y r h y n o h o s platyrh.ynohos i n N e b r a sk a .

Ee named

(10)

Z. c e r a t o s a a s t h e ty p e o f t h e genus*
P r i c e (103)^ h a s d i s c u s s e d t h e taxonomy o f t h e genus
and c o n c lu d e d t h a t Z. c e r a t o s a and Z. lunaturn a r e synonymous*
Thus Z. lu n a tu m {D ies*, 1836) becomes t h e s o l e and type
s p e c ie s in the genus.

P r i c e a l s o r e p o r t s t h e s p e c i e s from

t h e cow (Bos t a u r u s ) from t h e Panama Canal Zone, and i n
a d d i t i o n from t h e f o l l o w i n g b i r d h 0 3 t s :

Anser a n s e r d o m e s t i o u s .

M a r i l a a m e r i c a n a . H e t t i o n c a r o l i n e n s i s , and G a l l i n g o d e l i o a t a .
Ee g i v e s a v e r y good c o m p a r a tiv e d i s c u s s i o n o f t h e m a t e r i a l
from th e d i f f e r e n t h o s t s .
In t h e p r e s e n t s t u d y Z y g o c o ty le lunatum h as been r e c o r d e d
from t h e m a l l a r d (Anas jd. p l a t y r h y n c h o s ) and t h e d o m e s tic
m a l l a r d , th e b a l d p a t e (H areoa a m e r i c a n a ) , t h e b lu e - w in g e d
t e a l ( Q uerquedula d i s c o r s ) , redhead (flyroca a m e r i c a n a ) ,
l e s s e r s c a u p duck (Ayroca a f f i n i s ) , and the b l a c k duck ('Anas
ru b rip e s ).

Each o f t h e s e s p e c i e s , w i t h th e e x c e p t i o n o f th e

m a l l a r d and r e d h e a d , r e p r e s e n t s a new h o s t r e c o r d .

M ic h ig an

i s a new l o c a l i t y r e c o r d f o r t h e o c c u r r e n c e o f t h e p a r a s i t e .
S in c e S tu n k a r d (130) and P r i c e (103) have d i s c u s s e d
t h e anatomy o f t h i s form i n d e t a i l , and a s h o r t d e s c r i p t i o n
o f the flu k e i s contained in a n o th er s e c tio n o f the p re s e n t
p a p e r , no d i s c u s s i o n o f t h e anatomy o f t h i s form i s deemed
necessary here.

1*

lu m b e rs i n p a r e n t h e s i s r e f e r to l i t e r a t u r e number.

(11)

The l i f e h i s t o r y o f t h i s s p e c i e s was d i s c o v e r e d q u i t e
by a c c i d e n t .

While s e i n i n g f o r minnows i n W i n te r g r e e n Lake

to o b t a i n i n f e c t i v e s t a g e s o f a d i f f e r e n t f l u k e , a s i n g l e
l a r g e E e li s o m a t r i v o l v i s was ta k e n i n t h e n e t .

This s n a i l

was b r o u g h t i n t o th e l a b o r a t o r y and d i s s e c t e d .

The h e p a t i c

t i s s u e s were c o m p l e t e l y f i l l e d w i t h r e d i a and c e r c a r i a o f
an a m p h isto m e , a s w e l l a s a few f e r o a c e r c o u s c e r c a r i a .
The amphistome c e r c a r i a e were l a r g e , a c t i v e , many o f them
b e i n g f u l l y d ev e lo p ed and r e a d y to l e a v e th e s n a i l .

A fter

a b o u t two h o u r s t h e s e c e r c a r i a began e n c y s t i n g on the s i d e s
and bottom o f t h e d i s s e c t i n g d i 3 h .

When r e a d y to e n c y s t

t h e c e r c a r i a a t t a c h e d th e m se lv e s to t h e g l a s s , v i b r a t e d t h e
body i n a c i r c u l a r m o t i o n , th e t a i l b e i n g l o s t in th e p r o c e s s .
They g r a d u a l l y d eve loped a h e a v y , brown c y s t w a l l , which i s
roughly h em isp h erical in shape.
I n o r d e r to o b t a i n th e a d u l t f l u k e , f i f t y c y s t s were
p l a c e d i n s i d e a s m a l l p i e c e o f b r e a d and fed to a h a l f - g r o w n
m a l l a r d duck.

This duck had b ee n r e a r e d u n d e r s c r e e n , w i t h o u t

a c c e s s to w a t e r o t h e r th a n w e l l w a t e r , and was r e g a r d e d a s
p a r a s i t e f r e e a t t h e time o f i n f e c t i o n .

The duck was examined

two weeks a f t e r i n f e c t i o n a l o n g w i t h a c o n t r o l .

The i n f e c t e d

duok c o n t a i n e d 46 immature Z y g o co ty le i n th e low er i n t e s t i n e
and caecum, t h e c o n t r o l b e i n g n e g a t i v e .

(12)

The a b o v e , o f c o u r s e , g i v e s o n l y an i n d i c a t i o n o f th e
d i r e c t i o n o f th e l i f e h i s t o r y o f t h i s form*

More work i s

b e i n g done a t th e p r e s e n t t i m e , and i t i s hoped a d e t a i l
r e p o r t on t h e l i f e h i s t o r y o f t h i s s p e c i e s w i l l be r e a d y
w i t h i n a s h o r t time*

From t h e f a c t a found r e l a t i v e to the

l i f e h i s t o r y , and t h e known l i f e h i s t o r i e s o f t h e P a ra m p h is­
tomidae i n g e n e r a l , i t i s p r o b a b l y c o r r e c t to assume t h a t
th e l i f e - c y c l e o f Z y g o c o ty l e f o l l o w s p r a c t i c a l l y th e same
c o u r s e a s C o t y lo p h o ro n c o ty lo p h o ru m .

B e n n e t t (1936) d e s ­

c r i b e d t h e l i f e - o y o l e o f t h a t form i n g r e a t d e t a i l ,

That

E eliso m a t r i v o l v i s may a c t a s t h e s n a i l h o s t f o r Z ygo co ty le
h a s u n d o u b t e d l y b e e n shown, and no doubt th e m e t a e e r e a r i a
e n c y s t on w a t e r v e g e t a t i o n where t h e y f i n d t h e i r way i n t o
the f i n a l h o s t .

Water b i r d s p r o b a b l y a c t a s t h e u s u a l h o s t

f o r t h i s s p e c i e s , th e r u m in a n ts b e i n g a c c i d e n t a l h o s t s .
No c a s e o f p a t h o g e n i c i t y h a s been a t t r i b u t e d

to the

genus Z y g o c o t y l e . e i t h e r i n t h e l i t e r a t u r e or i n t h i s s t u d y .
They a r e p r e s e n t , u s u a l l y o n ly i n v e r y s m a ll numbers, in
t h e c a e c a , and a r e p r o b a b l y more c o m m e n s a l i s t i c th a n p a r a ­
s i t i c in h a b i t .

In no c a s e have more tha n f o u r o f t h e s e

worms b ee n p r e s e n t i n any h o s t o t h e r th a n t h e a r t i f i c i a l l y
i n f e c t e d o n e.

A lthough t h e y have been r e p o r t e d from th e

i n t e s t i n e s o f b i r d s , I am i n c l i n e d to c o n s i d e r t h i s an
abnormal h a b i t a t , t h e p a r a s i t e s p r o b a b l y w orking t h e i r Wfcy
o u t o f t h e caecum a f t e r th e d e a t h o f th e b i r d .

These p a r a -

(13)

s i t e s a r e v e r y r e s i s t a n t , and f r e q u e n t l y a r e a l i v e 24-36
hours a f t e r the death of the h o s t.

Fam ily P l a g i o r c h i d a e Luhe, 1901
Genu3 Prosthogonim us Luhe, 1899
A lthough s i x t e e n v a l i d s p e c i e s from v a r i o u s r e g i o n s
o f t h e world a r e r e c o g n i z e d by v a r i o u s w orkers a s b e l o n g ­
i n g to th e genus P r o s t h o g o n i m u s » o n l y two o f t h e s e s p e c i e s
have to d a t e b ee n r e p o r t e d from N orth A m erica.

These worms,

b e l o n g i n g t o th e f a m i l y P l a g i o r c h i d a e , ( E a p l o r o h i d a e McMullen,
1 9 3 7 ) , a r e i n h a b i t a n t s o f t h e b u r s a o f F a b r e c i u s and th e
oviduct of v a rio u s b i r d s .

A l l th e r e s e a r c h e s on th e l i f e

h i s t o r y o f t h i s genus i n d i c a t e t h a t d r a g o n - f l y n a i a d s s e r v e
a s h o s t s f o r t h e e n c y s te d m e t a c e r c a r i a l s t a g e .
Luhe e r e c t e d th e genus P ro sthogonim us i n 1899, h i s
name t a k i n g p r e c e d e n c e over L o o s s ' s Prymnoprion (1899) by
one d ay, and made P. o v a t u s ( R u d ., 1809) ty p e o f the g e n u s .
L a t e r Luhe (64) e r e c t e d the s u b f a m i l y P ro s th o g o n im in a e t o
c o n t a i n th e g e n e r a P rosthogonim us and S c h i s t o g o n i m u s .

The

o n l y d i f f e r e n c e in the two i s t h a t in th e form er the male
and f e m a le d u c t 3 open t o g e t h e r , w h i l e i n th e l a t t e r th e y a r e
s e p a r a t e d one from the o t h e r .

Macy (68) h a s g i v e n a v e r y

good summary o f t h e g e n u s , and a key f o r the s e p a r a t i o n o f
t h e a d u l t s o f th e d i f f e r e n t s p e o i e s .

(14)

P roa thogonimus m a c r o r c h i a Maey, 1934
3fhe f i r s t r e c o r d o f a member o f t h i s genus from North
America was by l i n t o n (1887)*

Ee i d e n t i f i e d a f l u k e a s

b e l o n g i n g to t h i s genus from th e egg o f a hen from B e l o i t ,
Wisconsin*

f h e r e were o t h e r i n s t a n c e s from time to time

o f t h e o c c u r r e n c e o f f l u k e s o f t h i s genus i n t h e eggs o f
hens i n t h e n o r t h e r n U nited S t a t e s and Canada*

About 1920

B r . W. L* C h a n d le r i d e n t i f i e d f l u k e s o f t h i s genus i n the
o v i d u c t o f a w in g - p i n i o n e d m a l l a r d duck from Buck l a k e ,
n e a r Muskegon, M i c h ig a n , i n which a v e r y s e r i o u s p a t h o ­
l o g i c a l c o n d i t i o n was noted i n th e body c a v i t y .

l a t e r he

( E o t l a n and C h a n d l e r , 1925) found t h a t t h e r e were s e r i o u s
l o s s e s a t th e r a n c h on t h i s l a k e among the l a y i n g h e n s , due
t o t h e same f l u k e s noted i n the m a l l a r d duck*

In t h e hens

a v e r y much more s e r i o u s c o n d i t i o n was noted and d e s c r i b e d .
I n 1927 th e same a u t h o r s made a p r e l i m i n a r y r e p o r t on t h e l i f e
h i s t o r y o f t h i s f l u k e , showing t h a t th e i n f e c t i v e s t a g e
o c c u r r e d i n t h e n a i a d s and a d u l t s o f d r a g o n - f l y a o f t h e genus
T etragoneurla.

L a c k l e a (1931) i n f e c t e d v e r y young c h i c k e n s

and ducks w ith t h e s e f l u k e s .

Macy (68, 69) d e s c r i b e d t h e

common N orth American form a s a new s p e c i e s ( Proa thogonimus
m a c r o r c h i s ) and worked o u t t h e l i f e h i s t o r y i n d e t a i l .

Ee

found t h a t th e sm a ll s n a i l s Amnicola lim n o sa p o r a t a (Say)

(15)

s e r v e d a s th e f i r s t i n t e r m e d i a t e h o s t , w h ile n a i a d s b e l o n g i n g
t o f o u r d i f f e r e n t g e n e r a o f d r a g o n f l i e s were found i n f e c t e d
w ith the c y s t s .

The d r a g o n f l i e s a r e l e u o o r r h i n i a . T e t r a -

g o n e u r i a . E p i c o r d u l i a . and M e so th em is.

Macy (69) h a s a l s o

given an e x te n s iv e d is c u s s io n o f the h o s t - p a r a s i t e r e l a t i o n ­
ships of th is sp e cie s.
S in c e K o tla n and C h a n d l e r ' s a s w e l l a s Macy's work
show th e extreme p a t h o g e n i c i t y o f P ros thogonimus m a o r o r o h i s .
a s a p a r a s i t e o f c h i c k e n s , and s i n c e t h e r e a r e numerous
r e p o r t s o f th e p a t h o g e n i c i t y o f c e r t a i n European s p e c i e s ,
e s p e c i a l l y P. p e l l u o i d u s . t h e r e i s l i t t l e doubt t h a t t h i s
i s a n e x t r e m e l y i m p o r t a n t genus from the s t a n d p o i n t o f
econom ics.

E x p erim e n ts have shown t h a t o n ly a few f l u k e s

a r e n e c e s s a r y to s t o p egg p r o d u c t i o n c o m p l e t e l y , and th e
d e a t h o f t h e b i r d o f t e n f o l l o w s a heavy i n f e s t a t i o n .

Chickens

a r e e s p e c i a l l y s u s c e p t i b l e because the oviduct i s f u n c tio n a l
during the e n t i r e y e a r.

Z o t l a n and Chandler (1925) have

d e s c r i b e d i n d e t a i l t h e c o n d i t i o n b r o u g h t ab o u t by a heavy
in festatio n .
Macy s t a t e s t h a t th e duck i s p r o b a b l y th e normal h o s t ,
whereas t h e c h i c k e n i s a c c i d e n t a l or a b n o r m a l.

This o p i n i o n

i s b ased on th e extrem e p a t h o g e n i c i t y o f t h e s e p a r a s i t e s
f o r c h i c k e n s , and th e f a c t t h a t t h e y rem ain o n l y a r e l a t i v e l y

(16)

s h o r t time i n t h e s e h o s t s .

C onsidering o th e r f a c t s , i t i s

h ig h ly probable t h a t th ese p a r a s i t e s are norm ally p a r a s i t i c
i n t h e b u r s a o f F a b r e e i u s o f f i r s t y e a r b i r d s , and t h a t th e
o v id u c t is only a f a c a l t a t i v e l o c a t i o n .

This i s i n d i c a t e d

by t h e f a c t t h a t t h e p a r a s i t e s w i l l n o t d e v e lo p i n a q u i e s ­
c e n t o v i d u c t , and i n w ild ducks the o v i d u c t i s a c t i v e f o r
a p e r i o d o f p r o b a b l y n o t o v e r one month o u t o f th e y e a r ,
w hereas i n t h e young b i r d s the b u r s a i s p r e s e n t u n t i l 7-8
months o f a g e , th u s g i v i n g a much l o n g e r p e r i o d o f p o s s i b l e
parasitism *

tty a u t o p s i e s on f i r s t y e a r ducks show a v e r y

h ig h p e r c e n t a g e o f them i n f e s t e d w i t h worms o f t h i s g e n u s ,
w h i l e the o n ly c a s e s o f n a t u r a l i n f e s t a t i o n o f a d u l t f e ­
m a les came w i t h i n a r e l a t i v e l y s h o r t p e r io d o f l e s s th a n
t h r e e weeks.
O b s e r v a t i o n s on a number o f fem ale ducks i n f e s t e d
w i t h P rosthogonim us m a c r o r c h i s have shown th e p a r a s i t e
to be r e l a t i v e l y n o n - p a t h o g e n i c i n old ducks i n th e w ild
state.

In one c a s e , a fem ale m a l l a r d was ta k e n on May

7, 1936, i n a t r a p a t W i n te r g r e e n Lake.

The a u t o p s y o f

t h i s b i r d r e v e a l e d 47 l a r g e Prostho gonim us m a c r o r c h i s in
the o v id u c t.

B e f o r e open in g the o v i d u c t t h e o n ly i n d i c a t i o n

o f t h i s h ea v y i n f e s t a t i o n was th e s l i g h t l y anemic a p p e a r a n c e
o f th e w a l l o f the o v i d u c t .

I n th e lumen o f t h e o v i d u c t ,

a s i d e from th e f l u k e s t h e m s e l v e s , o n ly a v e r y few, s m a ll

(17)

albumen m asses c o n t a i n i n g eggs were p r e s e n t .
A n o th e r female m a l l a r d t r a p p e d on May 1 1 , 1936, and
k e p t over n i g h t i n a p en , l a i d an egg d u r i n g th e ti m e ,
and when a u t o p s i e d showed f i v e v e r y l a r g e P rosthogonim us
i n t h e u p p e r p o r t i o n o f t h e o v i d u c t , and t h r e e Eohinostomum
r e v o lu tu m i n t h e s h e l l g l a n d .

T his b i r d a p p e a r e d p e r f e c t l y

norm al•
From t h e s e o b s e r v a t i o n s i t may seem r e a s o n a b l e to
c o n c lu d e t h a t t h i s f l u k e i s n o t , u n d e r o r d i n a r y c o n d i t i o n s ,
v e r y p a t h o g e n i c to wild ducks; a t l e a s t i n no w ise a p p r o a c h ­
ing th e c o n d itio n c re a te d in ch ick en s.

F u r t h e r , o f ov er a

dozen c a s e s examined i n which Prosthogonim us was p r e s e n t
i n t h e b u r s a , n o t one showed any n o t i c e a b l e g r o s s p a t h o l o g y .
However, a s i n o t h e r c a s e s o f p a r a s i t i s m , c o n d i t i o n s may
p r e v a i l to b r i n g a b o u t an e x c e p t i o n a l l y heavy i n f e s t a t i o n .
The c o n d i t i o n d e s c r i b e d by C handler (51) in th e c a s e o f th e
duck from Muskegon, M ic h ig a n , i n t h e a b s e n c e o f o t h e r
r e p o r t s o f p a t h o g e n i c i t y among d u c k s , i s p r o b a b l y the
e x c e p t i o n r a t h e r th a n t h e r u l e .

A lthough t h e p r e s e n c e o f

o t h e r d i s e a s e s , e i t h e r p a r a s i t i c o r b a c t e r i a l , may d e t e r m in e
t h e d e g r e e o f i n f e s t a t i o n a s w e ll a s p a t h o g e n i c i t y .
The f o r e g o i n g d i s c u s s i o n has d e a l t a l m o s t e x c l u s i v e l y
w i t h P rosth o g o n im u s m a c r o r c h i s (Macy, 1 9 3 4 ) .

This s p e c i e s

(18)

h a s b e e n t h e o n ly one r e c o v e r e d from f e e d i n g e x p e r i m e n ts
a t t h e S a n c t u a r y , and t h e o n ly one ta k e n from t h e o v i d u c t
o f b i r d s c a u g h t t h e r e d u r i n g t h e b r e e d i n g season*

J u d g in g

from M acy's work, and th e o t h e r p u b l i s h e d work on th e
s u b j e c t , t h i s i s by f a r th e most p r e v a l e n t s p e c i e s o f o v i ­
d u c t f l u k e s i n w a te r f o w l and c h i c k e n s i n t h e U nited S t a t e s *
P ro sth o g o n im u s r u d o l p h i i S k r j a b i n , 1919
This s p e c i e s h a s b e e n r e c o r d e d p r e v i o u s l y o n l y once
i n N o rth America ( 7 0 ) , h a v i n g b ee n t a k e n from t h e b u r s a
o f w ild m a l l a r d s i n Minnesota*

However, o f t h e m i g r a t i n g

f i r s t - y e a r ducks examined, a l a r g e p e r c e n t a g e have b e e n
i n f e o t e d w i t h t h i s s p e c i e s , t h e worm3 b e i n g found i n eaoh
c a s e i n the b u r s a *

On th e whole t h e y a v e r a g e o n ly a b o u t

one h a l f a s l a r g e a s P . m a c r o r o h i 3 . and may e a s i l y be t o l d
from t h a t s p e c i e s by the c o n f in e m e n t o f the u t e r u s w i t h i n
t h e i n t e s t i n a l c r u r a , and th e e x t e n s i o n o f t h e v i t e l l a r i a
p o s t e r i o r to th e t e s t e s *

E*

r u d o l p h i i h a s b ee n ta k e n from th e f o l l o w i n g d u c k s :

G l a u c i o u e t t a c l a n g u l a a m e r i c a n a . Nyroca a f f i n i s . Nyroca
o o l l a r i 3 , Anas p l a t y r h y n c h o s plat.yrhynchos „ and Anas
rubripes*

T hese, w i t h th e e x c e p t i o n o f A. p . p l a t y r h y n c h o s .

a l l r e p r e s e n t new h o s t r e c o r d s , and th e l o c a l i t y i s new*

(19)

So f a r no i n s t a n c e o f p a t h o g e n i c i t y h a s b een o b s e r v e d , and
w h eth e r or n o t t h i s s p e c i e s w i l l i n f e c t c h i c k e n s i s y e t to
be d e m o n s tr a te d *

P rosthogonim ps a n a t i n n s Msrkow, 190 8
Only one specimen o f t h i s s p e c i e s has b ee n t a k e n .

It

was found i n t h e b u r s a o f a f i r s t - y e a r b l a c k duck (Anas
r u b r l p e a ) , ta k e n a t t h e S a n c t u a r y ,
P . r u d o l p h i i were a l s o p r e s e n t .

Three specim ens o f

This s p e c i e s may be d i s ­

t i n g u i s h e d from th e two p r e v i o u s l y mentioned by the much
g r e a t e r l e n g t h o f t h e c i r r u s pouch, which e x t e n d s p o s t e r i o r
a l m o s t a s f a r a s t h e a c e ta b u lu m .

D iscussion
Macy s u g g e s t s t h a t P. a n a t i n u s , P. s k r j a b i n i , and
P . k a r a u s a i k i a r e so much a l i k e t h a t th e y may be found to
by synonymous, and a l s o s t a t e s t h a t t h e s e t h r e e s p e c i e s
may be b u r s a l m o d i f i c a t i o n s o f some o t h e r form r e c o r d e d
from t h e o v i d u c t .

Prom h i s work d e a l i n g w ith th e e f f e c t

o f the h o s t on th e anatomy o f t h e a d u l t , and from s i m i l a r
e x p e r i m e n t s and o b s e r v a t i o n s i n t h i s s t u d y , I b e l i e v e t h a t
when more work i s done w i t h h o s t m o d i f i c a t i o n on t h e s e form s,
t h e l a t t e r h y p o t h e s i s w i l l be v a r i f i e d .

Macy, w orking w ith

P . m a c r o r c h i s , found t h a t specim ens from the o v i d u c t o f the

( so)

d o m e s tic ducks and c h i c k e n s a r e more l i k e one a n o t h e r th a n
t h e y a r e l i k e specim ens from th e b u r s a o f e i t h e r h o s t *

In

h i s b u r s a l i n f e c t i o n e x p e r i m e n ts he was a b l e to i n f e c t
E n g l i s h s p a rr o w s a s w e l l a s cro w s.

There seems to be some

h o s t m o d i f i c a t i o n i n each o f t h e s e c a s e s .
P u r s u i n g t h i s m a t t e r f u r t h e r , 50 d r a g o n - f l y maiads
b e l o n g i n g to t h e genus T e t r a g o n e u r i a were c o l l e c t e d a l o n g a
p o r t i o n o f t h e l a k e s h o r e a t W i n te r g r e e n Lake between March
20 and 26, 1937.

Six o f t h e s e were d i s s e c t e d , and t h r e e

were found to be i n f e c t e d w ith o n e , t h r e e , and s i x c y s t s
respectively .

The o t h e r 42 n a i a d s were fed to two brown

l e g h o r n hens known to be l a y i n g , and to a m a l l a r d d u c k , each
a n i m a l r e c e i v i n g 14 n a i a d s .

Nine days a f t e r f e e d i n g the

n a i a d s , one o f t h e h ens was a u t o p s i e d and 26 l a r g e P r o a t h o gonimus m a c r o r c h i s were r e c o v e r e d from th e o v i d u c t .

A

r u p t u r e d o v u le was a b o u t h a l f way down t h e o v i d u c t , and
t h r e e o f th e f l u k e s were i n t h i s mass o f y o l k .

The o t h e r

h e n , a u t o p s i e d 14 days a f t e r i n f e c t i o n , had o n ly f o u r f l u k e s
p r e s e n t i n t h e o v i d u c t , t h e s e b e i n g o n ly s l i g h t l y l a r g e r
t h a n i n the form e r c a s e .

I n t h i s c a s e the hen was a b o u t

r e a d y to resume p r o d u c t i o n , f o r a p e r f e c t l y normal s o f t s h e l l e d egg was i n th e s h e l l g l a n d .

The duck was t i l l e d

t h r e e weeks a f t e r i n f e c t i o n and 14 f l u k e s were r e c o v e r e d

(21)

from t h e u p p e r h a l f o f t h e o v i d u c t .

The o v i d u c t o f t h i s

duck was becoming q u i e s c e n t , f o r i t was o n ly a b o u t h a l f a s
l a r g e a s i t should be i n a l a y i n g b i r d .

The f l u k e s from

th e duck were a b o u t t h e same s i z e a s th o s e from t h e h e n s ,
b u t when f i x e d and s t a i n e d , q u i t e a h i t o f m o d i f i c a t i o n
was e v i d e n t .

The f l u k e s from th e duck d i f f e r e d from th o s e

from t h e h e n s i n th e f o l l o w i n g p o i n t s :

t h e t e s t e s were

Only a b o u t one h a l f a s l a r g e a s i n th e l a t t e r , and th e
u t e r u s , w h ile showing th e 3ame g e n e r a l c o i l i n g , was v e r y much
s m a l l e r , and c o n t a i n e d o n ly a f r a c t i o n o f th e number o f eggs
found i n th e l a t t e r .
Prom t h e s e d a t a i t would a p p e a r t h a t th e p h y s i o l o g i c a l
c o n d i t i o n o f th e o r g a n , a s w e ll a s the organ and th e h o s t
may i n f l u e n c e th e morphology o f th e worm.

I t seem3 s a f e to

i n f e r t h a t when e x t e n s i v e s t u d i e s a r e made on th e h o s t p a r a s i t e r e l a t i o n s h i p s o f t h e genus Prosthogonim u3 s e v e r a l
o f th e r e c o g n i z e d s p e c i e s may be found to be h o s t o r organ
m o d ific a tio n s o f these s p e c ie s .

f a m i l y O p i s t h o r o h i i d a e Luhe, 1901
Qenus O p i s t h o r c h i s B l a n c h a r d , 1898
Of th e ducks examined a t th e W. K. K e llo g g B i r d S a n c t ­
u a r y a b o u t a dozen have been i n f e c t e d w i t h an o p i s t h o r c h i d
f l u k e i n th e p a n c r e a s and l i v e r .

These b i r d s were f o r the

(88)

moat p a r t c a p t i v e , w in g - p in io n e d b i r d s , b u t t h e p a r a s i t e s
have a l a o been observed i n w ild g o l d e n - e y e s t a k e n away
from th e S an ctu ary *

I n f e s t a t i o n has b ee n r e c o r d e d i n the

fo llo w in g s p e c ie s o f ducks:

m a l l a r d (Anas p l a t y r h y n c h o s

p l a t y r h y n c h o s ) . bl&ck duck (Anas r u b r i p e s ) . American g o l d e n ­
eye ( G rlauoionetta c l a n g u l a a m e r l c a n a ) , redhead (CTyrooa
a m e r i c a n a ) , and widgeon (Mareoa a m e r i o a n a ) .

P r e v i o u s to

t h i s s t u d y t h e s e f l u k e s have a l s o b ee n r e c o r d e d i n the
f o l l o w i n g c a p t i v e w a te r f o w l a t th e W. K. K ello g g B ird
Sanctuary:

r o s y - b i l l e d duck (P a e c i l o n i t t a baham ensis

r u b r l r o a t r i s ) , whooping swan ( Cygnus o y g n u s) , and b l a c k
swan ( Ohenopsis a t r a t a ) .

Prom t h i s l i s t o f s p e c i e s s e r v i n g

a s h o s t to t h i s p a r a s i t e i t i s r e a s o n a b l e to suppose t h a t
an y duck or swan may become i n f e c t e d i f th e i n f e c t i v e s t a g e
i s introduced in to the d i g e s ti v e t r a c t .

I t is h ig h ly prob­

a b l e t h a t o t h e r b i r d s such a s t h e k i n g f i s h e r , e t c . , m ight
a c t a s a h o s t f o r t h i s worm a s w e l l a s w a t e r f o w l .
S t u d i e s on th e morphology o f t h i s worm i n d i c a t e i t to
be a new s p e c i e s b e l o n g i n g to th e genus Amphimerus B a r k e r ,
1911, ( O p i s t h o r c h i i d a e ) .

The f o l l o w i n g d i a g n o s i s i s b as ed

on e x a m i n a t i o n o f a l a r g e number o f specim ens o b t a i n e d from
ducks.

S e r i a l s e c t i o n s o f p i e c e s o f p a n c r e a s have b e e n made,

a s w e l l a s a v e r y l a r g e number o f whole mounts o f f ra g m e n ts
o f worms.

Due to th e extrem e d i f f i c u l t y i n removing the

(23)

worms from t h e p a n c r e a s or l i v e r i n t a c t , o n l y t h r e e e n t i r e
worms h ave b ee n a v a i l a b l e f o r com plete m e a s u r e m e n ts , b u t
a v e r y l a r g e number o f f r a g m e n ts o f worms have been s t u d i e d ,
and th e d e t a i l s o f anatomy made o u t and compared.

A ll

measurements r e p o r t e d h e r e a r e from balsam m ounts.
$he genus Amphimerus was e r e c t e d by B a r k e r i n 1911,
(3) to c o n t a i n t h o s e s p e c i e s o f t h e genus O p i s t h o r c h i s i n
which t h e v i t e l l a r i a a r e d i v i d e d i n t o an a n t o v a r a l and
postovaral p o rtio n .
i n th e new g e n u s :

He p la c e d t h e f o l l o w i n g s i x s p e c i e s
Amphimerus o v a l i s B a r k e r . 1911; Amphimerus

i n t e r r u p t u s ( B r a u n , 1901) B a r k e r , 1911; A. l a n o e a ( D i e s i n g ,
1850) B a r k e r , 1911; A. n o v e r c a (Braun, 1901) B a r k e r , 1911;
A. p 3 e u d o f e l l n e u s (Ward, 1901) B a r k e r , 1911; A. s p e c i o s u s
( S t i l e s and H a s s a l l , 1894) B a r k e r , 1911.
Morgan (84) h a s shown t h a t 0 . n o v erca (B rau n , 1 9 0 1 ),
p l a c e d by B a r k e r i n th e genus Amphimerus. b e l o n g s to the
genus O p i s t h o r c h i s .

He a l s o d i s c u s s e d th e v a l i d i t y o f th e

new genus and c o n c l u d e s "

i t s r e t e n t i o n d e s i r a b l e in

the p r e s e n t s t a t e o f our knowledge o f t h i s g r o u p " .
In t h e p r e s e n t p a p e r a new s p e c i e s , Amphimerus e l o n g a t u s
i s added to t h e g e n u s , and Amphimerus (E e m a t o t r e p h i a ) f o d i e n s
L i n t o n , 1928 i s added a s " s p e c i e s i n q u i r e n d a e " .

(24)

Amphimerus e l o n g a t u s n . s p .
S pecific diagnosis:

O piathorohiinae.

Body v e r y narrow

and e l o n g a t e , t a p e r i n g from a b o u t the l a s t body t h i r d .
sm ooth.

T o t a l l e n g t h from 20 to 23 mm.

Skin

Body w id th g r e a t e s t

j u s t a n t e r i o r to o v a r y where i t measured 1 .0 9 mm to 1 . 1 2 mm.
Width a t t e s t e s .7 to .8 mm, a t ac e ta b u lu m .2 4 to .25 mm,
and a t

t h e a n t e r i o r end .028 to

e n tire ly absent.

.032 mm.

The o r a l s u c k e r i s

Mouth open in g sub t e r m i n a l , mouth

opening

(tube)

l e a d i n g i n t o a w e l l developed o v a l p h a r y n x , p la c e d

,06 to

t 08 mm from t h e a n t e r i o r

•032mra by .064 mm.

e n d ,and m e a su rin g .041 to

Acetabulum i n th e a n t e r i o r t h i r d o f

bo d y , measured .105 to .124 b y .093 to .099 mm.
a b o u t .1 to .15 mm l o n g .

Esophagus

I n t e s t i n a l c r u r a r e a c h i n g to or

n e a r l y to the p o s t e r i o r end, in some c a s e s t u r n i n g toward
t h e median l i n e .

T e s t e s i n t h e p o s t e r i o r e i g h t h o f body;

p la c e d one i n f r o n t o f th e o t h e r , o r s l i g h t l y o b l i q u e ;
lo n g o v a l i n s h a p e , b o r d e r s somewhat i r r e g u l a r i n o u t l i n e .
Average a b o u t 1 . 5 mm by . 3 mm.
l o b e d , .3 by .35 mm.

Ovary a n t e r i o r to t e s t e s ,

Reoeptaculum serainis p r o m i n e n t ,

l a r g e r t h a n o v a r y , *27 to .3 by .49 to 1 . 0 mm.
oanal p r e s e n t.

laurer*s

U t e r i n e c o i l s b e g i n j u s t a n t e r i o r to o v a r y ,

and a r e v e r y d e n s e , e x t e n d i n g over the i n t e s t i n a l c r u r a
to th e body w a l l a b o u t h a l f th e d i s t a n c e betw een the
o v a r y and a c e t a b u l u m , th e n t h i n n i n g o u t i n t o more or l e s s
r e g u l a r f o l d s c o n f in e d between t h e c r u r a .

V itellaria

(25)

d i v i d e d i n t o a d i s t i n c t a n t o v a r a l and p o s t o v a r a l p o r t i o n ,
e x t e n d i n g p o s t e r i o r beyond the c e n t e r o f th e p o s t e r i o r
t e s t i s , and a n t e r i o r a b o u t one t h i r d th e d i s t a n c e from the
o v a r y to t h e acetabulum *

Eggs lo n g o v a l , o p e r c u l a t e ,

m e a s u r in g *012 by .022 mm.
H a b i t a t : p a n c r e a t i c and b i l e p a s s a g e s o f v a r i o u s ducks
and swans.
D istribution:
Type:

M ic h ig a n , U. S. A.

U. S. U. M. Ho. 9051

P a r a t y p e Ho. 9052

A. e l o n g a t u s i s more l i k e A. s p e c i o s u s ( S t . and H . )
th a n an y o f th e o t h e r d e s c r i b e d s p e c i e s .

The s i z e , shape

o f b o d y , and e x t e n t o f c o i l i n g o f th e u t e r u s c l o s e l y
a p p r o x im a te t h a t s p e c i e s ,

I t may be d i s t i n g u i s h e d from

a l l the o t h e r s p e c i e s , however, by the a b s e n c e o f an o r a l
s u c k e r , and t h e shape and s i z e o f t h e t e s t e s , a c e ta b u lu m ,
p h a r y n x , and r e o e p t a c u l u m s e m i n i s .
The d e a t h o f a number o f b i r d s , p a r t i c u l a r l y th e
r o s y - b i l l e d d u c k , have b e e n a t t r i b u t e d e i t h e r d i r e c t l y o r
i n d i r e c t l y to t h i s f l u k e a t t h e W. K* K e llo g g B ird S a n c t u a r y .
As i n th e c a s e o f a l m o s t a l l h e l m i n t h i n f e s t a t i o n s , a v e r y
l a r g e number o f worms w i l l produce a h ar m f u l e f f e c t on the
h o s t , b u t a l t h o u g h 15 s p e c i e s o f f l u k e s have been re o o rd e d
from ducks a t th e S a n o t u a r y , A. e l o n g a t u s a p p e a r s to be the

(26)

one o f g r e a t e s t economic s i g n i f i c a n c e .
L i t t l e g r o s s p a t h o l o g y has b e e n o b s e r v e d .

On t h e

whole t h e p a n c r e a s a p p e a r s a l i t t l e a n e m ic , b u t /this i s
n o t c o n s t a n t and may be due to o t h e r c a u s e s .

S ections o f

i n f e c t e d p a n c r e a s show t h e t u b e s to be g r e a t l y e n l a r g e d
by t h e p r e s e n c e o f worms w ith a pronounced t h i c k e n i n g o f
t h e w a l l n e a r t h e a n t e r i o r r e g i o n o f t h e worm.

Since t h e r e

i s a l m o s t no m u s c u la r development i n t h e s e f l u k e s , i t i s
p r o b a b l e t h a t l i t t l e i f any movement t a k e s p l a c e .

More

o f t e n t h a n n o t , tv/o worms o c c u r s i d e by s i d e , and f r e ­
q u e n t l y e a c h worm i s doubled b ack on i t s e l f .

A sid e from a

t h i c k e n i n g o f the w a l l s t h e r e i s no d oubt some l e s s e n i n g
i n th e flo w o f th e p a n c r e a t i c f l u i d due to o c c l u s i o n o f
th e d u c t s .

This o f c o u r s e would be i n p r o p o r t i o n to the

extent of in fe c tio n .

The l i v e r i n most c a s e s i s l e s s

h e a v i l y i n f e c t e d th a n t h e p a n c r e a s , and i n many c a s e s n o t
at a ll,

No c a s e h a s b ee n found th u s f a r t h a t would s e r ­

i o u s l y h a n d i c a p t h e f u n c t i o n i n g o f the l i v e r .

A heavy

i n f e s t a t i o n r e s u l t s i n g r e a t l y lowered v i t a l i t y , poor
d i g e s t i o n , e m a c i a t i o n , a n d , e s p e c i a l l y i n young b i r d s , w i t h
th e a d v e n t o f some o t h e r f a c t o r such a s a d v e r s e w e a t h e r ,
other p a r a s ite s , e t c . , in death.

That th e worm i t s e l f

c o u ld c a u s e d e a t h i s n o t i m p o s s i b l e , b u t n o t p r o b a b l e ;
how ever, t h e end r e s u l t s a r e t h e same.

(27)

The e n t i r e l i f e h i s t o r y has n o t been d e te r m in e d a s
y e t , and w i l l be p u b l i s h e d a t a l a t e r d a t e .

That young

f i s h e s o a r r y th e i n f e c t i v e s t a g e has been e x p e r i m e n t a l l y
dem onstrated.

The f o l l o w i n g ex p e rim e n t was c o n d u c te d .

From a pen o f h a l f grown m a l l a r d s r a i s e d u nder s c r e e n
away from th e l a k e , two ducks were s e l e c t e d and plaoed i n
a s e p a r a te screened pen.

They were banded w i t h numbers

1444 and 1445, and d e s i g n a t e d a s Pen 1 .
of adult

A second

group

d u c k s , n o t u n d e r s c r e e n b u t n e v e r h a v i n g had

a c c e s s to the l a k e were d e s i g n a t e d by numbers 1441, 1442,
and 1443.

In eaeh c a s e b i r d s from th e o r i g i n a l pen were

used a s c o n t r o l s .
On August 6 , 1936, Pen 1 r e c e i v e d 40 s m a ll b a s s ,
p e r c h , and b l u e g i l l s , c u t up i n mash and g r a i n and
m oistened.

Pen 3 r e c e i v e d 24 s m a ll b a 3 S , p e r c h , and

b l u e g i l l s i n th e same manner*
t h r e e or f o u r p i e c e s .

The f i s h were c u t i n t o

In b o th c a s e s the ducks were

o bserved e a t i n g th e f i s h w i t h r e l i s h , even p i c k i n g i t
out o f the o th e r food.
75 f i s h e n t i r e .

On August 11, Pen 1 r e c e i v e d

They a t e th e e n t i r e l o t i n 12 h o u r s .

On th e same d a t e Pen 2 r e c e i v e d 7 minnows.
From Pen 1 duck 1445 was a u t o p s i e d on A ugust 15,
9 days a f t e r t h e b e g i n n i n g o f th e e x p e r i m e n t and found
i n f e c t e d w i t h a b o u t 50 f u l l y m a tu re Amphimerus e l o n g a t u s .

(28)

They had r e a c h e d f u l l s i z e , and were c o m p l e t e l y f i l l e d
w i t h eggs*

From Pen 2 duck 1442, an a d u l t female m a l l a r d

was a t u o p s i e d on th e same d a y , th e i n f e c t i o n b e i n g p r a c t i c ­
a l l y th e same a s i n th e above c a s e .

On August 19, duck

1444 was a u t o p s i e d and i n t h i s c a s e o n l y
noted.

5 worms were

On A ugust 85, duck 1441 was a u t o p s i e d and found

i n f e c t e d w i t h a b o u t 50 worms i n a b o u t the same c o n d i t i o n
a s the f i r s t two a n i m a l s .
A lthough time did n o t p e r m i t a v e r y e x t e n s i v e exam­
i n a t i o n , some o f th e f i s h were d i s s e c t e d , b u t no l a r v a l
f l u k e s were found e x c e p t s t r i g e i d f l u k e s o f the genus
Neascus.

A g r e a t d e a l more work i s to be done on t h i s

l if e h istory.

Buck 1443 was n o t a u t o p s i e d , and a l t h o u g h

t h i n f o r a b o u t two m onths, r e c o v e r e d co m pletely*

Fam ily M i e r o p h a l i d a e T r a v o s s o s , 1980
There h a s b e e n , and s t i l l

i s , a great deal of d is ­

agree m ent among p a r a s i t o l o g i s t s w ith r e g a r d s to the p r o p e r
r e l a t i o n s h i p s o f the group o f g e n e ra b e l o n g i n g to the
fam ily M ic ro p h a llid ae.

Ward (1901) e r e c t e d t h e s u b f a m i l y

M i o r o p h a l l i n a e w i t h i n t h e f a m i l y H e te r o p h y d i a e to c o n t a i n
h i s new genus M i o r o p h a l l u s .

S in ce t h a t time f i v e o th e r

g e n e r a have b een added to the group:

L ev in sen lella.

(29)

S p e l o t r e m a . S p e l l o p h a l l u a . Monocaeoum, and M a r it r e m a .

A

g r e a t d e a l has b e e n w r i t t e n by a number o f d i f f e r e n t work­
e r s w i t h i n th e l a s t few y e a r s w i t h r e g a r d to th e a p p a r e n t
i n c o m p a t i b i l i t y o f t h e s e s i x g e n e r a to form a n a t u r a l
grouping.

Some contend th e y do n o t b e l o n g t o g e t h e r a t

a l l , o t h e r s i n s i s t th e y should be c l a s s e d a s a s u b f a m i l y
u n d e r t h e H e t e r o p h y i d a e , and s t i l l o t h e r s t h a t th e y come
together n a tu r a lly
M ueller

i n t o a v a l i d f a m i l y . Tan Oleve

(151) g i v e a b r i e f d i s c u s s i o n o f

th e

and

controversy.

They a g r e e w i t h T r a v o s s o s , who f i r s t gave the group f a m i l y
rank in

1920, t h a t th e y do form a n a t u r a l f a m i l y .

w i l l be

considered o f fam ily rank h e r e .

They

Genus M a ritre m a N i c o l l , 1907
N i v o l l s e t up th e genus M a ritre m a i n 1907, naming
M. g r a t i o s u m a s t y p e .

The genus may be d i s t i n g u i s h e d

from th e o t h e r g e n e r a o f the f a m i l y by the s t r u c t u r e o f
t h e male g e n i t a l o r g a n s .

A w e l l developed s e m in a l v e s ­

i c l e , p r o s t a t i c c e l l s and a p r o t r u s i b l e c i r r u s , v/hich
opens i n a groove n e a r th e l e f t s i d e o f th e a c e t a b u l u m ,
is present.

They have a s p in e d s k i n , and t h e v i t e l l a r i a

a r e l a t e r a l , e x t e n d i n g mesiad a n t e r i o r to th e t e s t e s ,

(30)

m e e ti n g i n some o a s e s i n th e m i d - l i n e beh in d th e acetabulum *
J a g e r s k i o l d (39) h a s g i v e n a v e r y good d i s c u s s i o n o f
th e European forms*

He l i s t s f i v e s p e c i e s .

In a d d i t i o n

to t h e s e , t h r e e s p e c i e s have been d e s c r i b e d from N orth
America*

None o f the European forms were from ducks*

Swales (134) h a s d e s c r i b e d a s p e c i e s which he found im­
bedded i n th e i n t e s t i n a l w a l l o f b l a c k ducks a s S t r a p t o v i t e l l i a a c a d i a e n* g* , n* s p .

However, Dr* S w a les,

i n a p e r s o n a l communication inform s me t h a t h i s form r e a l l y
s h o u ld be p la c e d i n th e genus M a r i t r e m a *

Hence h i s form

i s th e f i r 3 t r e c o r d o f a member o f t h i s genus from North
A m e r ic a , a s w e l l a s the f i r s t r e c o r d from ducks*

In t h i s

c a s e a v e r y s e r i o u s p a t h o g e n i c d i s t u r b a n c e was n o t e d , th e
d e a t h o f a number o f ducks b e i n g caused by t h e s e f l u k e s *
They Were found i n g r e a t a b u n d a n c e , b o t h f r e e i n t h e lumen
and imbedded i n th e mucosa o f th e i n t e s t i n e , c a u s i n g a s e v e r e
inflam ation.
The o n l y o t h e r r e c o r d o f t h i s genus from North America
i s by f a n S ie v e and M u e l l e r (151) who, on a b a s i s o f a
s i n g l e immature specim en e a c h , d e s c r i b e d two new s p e c i e s ,
p l a c i n g them f i r s t i n t h e genus M i c r o p h a l l u s . b u t l a t e r
( M u e l l e r , 1934) removing them t o th e genus M a r i t r e m a *

These

worms were r e c o v e r e d from the i n t e s t i n e s o f f i s h and were

(SI)

n o t m a t u r e , b e c a u s e f i s h were u n d o u b te d ly a p u r e l y a c c i d e n t ­
a l d e f i n i t i v e h o s t to worms o f t h i s g e n u s .
I n a l a t e r s e c t i o n o f t h e same work ( p . 289) t h e s e
a u t h o r s s t a t e t h a t , s u b s e q u e n t to th e o r i g i n a l d e s c r i p t i o n ,
th e y o b t a i n e d o t h e r specim ens o f t h i s worm from A m b l o p l i t e s
ru p estris.

These were f o r t h e most p a r t even younger than

th e form er specim ens some o f them n o t h a v i n g been l i b e r a t e d
from th e c y s t .

These l a t e r specim ens showed v a r i a t i o n i n

s i z e o f t h e a l i m e n t r y o r g a n s somewhat i n t e r m e d i a t e between
t h e i r two s p e c i e s , b u t the v i t e l l a r i a did n o t show any
v a r i a t i o n from th e type specim en.

They may be synonymous

a c c o r d i n g to the a u t h o r .
During th e c o u r s e o f t h i s i n v e s t i g a t i o n , sm all f l u k e s
b e l o n g i n g to t h i s genus have been r e c o v e r e d i n l a r g e numbers
from t h e i n t e s t i n e s o f f i v e American g o l d e n - e y e s ( Qlauci o n e t t a o l a n g u l a a m e r i o a n a ) . a s w e ll a s from a s i n g l e
American sc a u p duck (Nyrooa a f f i n i 3 ) .

In each c a s e th e

worms were f r e e i n th e i n t e s t i n e , none b e i n g found imbedded.
The b i r d s came from Kalamazoo, Calhoun and Roscommon Count­
i e s , M ic h ig a n .

No s p e c i f i c p a t h o l o g i c a l c o n d i t i o n could be

i d e n t i f i e d i n any o f th e d u c k s , a l t h o u g h two o f th e g o l d e n ­
eyes were p ic k e d up i n th e s t r e e t s o f B a t t l e C re e k , M ic h ig a n ,
a p p a r e n t l y u n a b l e to f l y , and i n somewhat o f a dazed c o n d i t ­
io n from h a v i n g flown i n t o e l e c t r i c w i r e s .

Both o f t h e s e

b i r d s were e m a c i a t e d , no ca u se f o r the c o n d i t i o n b e i n g

(32)

b e i n g found a s i d e from t h e l a r g e numbers o f s m a ll f l u k e s
in the i n t e s t i n e s .

They were b o th j u v e n i l e b i r d s , one a

m a l e , th e o t h e r a f e m a le .

That the em aciate d c o n d i t i o n o f

t h e s e b i r d s was due to t h e f l u k e s i s i m p r o b a b l e , f o r g o l d e n ­
eyes s h o t b e f o r e and a f t e r the time o f examining th e two
t h i n b i r d s , c o n t a i n e d an e q u a l number o f th e f l u k e s , and
were i n p e r f e c t f l e s h .
These f l u k e s a r e s u f f i c i e n t l y d i f f e r e n t to w a r r a n t
t h e naming them a s a new s p e c i e s .

The f o l l o w i n g d e s c r i p ­

t i o n i s b ased on th e s t u d y and measurements o f a v e r y
l a r g e number o f s p e c im e n s .

M a ritre m a n e t t a e n . s p .

Small f l u k e s , body ovid to somewhat tongue s h a p e d ,
o f t e n s h a r p l y t r u n c a t e a t th e p o s t e r i o r e n d .

Skin com­

p l e t e l y covered w i t h v e r y s m a ll 3 p i n e s , s l i g h t l y l a r g e r
and more t h i c k l y s e t a t t h e a n t e r i o r end.

T otal le n g th

v a r i e s from . 4 t o .45 mm, g r e a t e s t w id th from .24 to .29
mm.

G r e a t e s t w id th im m e d ia te ly beh in d a c e t a b u l u m .

The

r a t i o betw een body w id th and l e n g t h v a r i e s c o n s i d e r a b l y ,
d ep e n d in g upon th e s t a t e o f th e worm when f i x e d , b u t i n
general i t is about 1 :1 .8 .

The o r a l s u c k e r i s p la c e d

a t th e a n t e r i o r t i p o f t h e b o d y , t h e open in g b e i n g o n ly

(ss)

s l i g h t l y sub t e r m i n a l .

The s u c k e r i s s l i g h t l y w i d e r tha n

l o n g , m e a s u r i n g .0 3 5 to .054 i n w i d t h and .041 t o .054 mm
in le n g th .

The p r e p h a r y n x i s q u i t e c o n s p i c i o u s , v a r y i n g

from .0128 to .0 2 8 mm i n l e n g t h .

The pharynx i s s l i g h t l y

w i d e r t h a n l o n g , m e a s u r i n g .025 - .035 b y .032 - .044 mm
in w idth.

The esophagus i s q u i t e v a r i a b l e , r a n g i n g from

.0 064 to .0256 mm i n l e n g t h .

The e r u r a d i v e r g e r a p i d l y ,

th e y a r e r e l a t i v e l y l a r g e and t h i c k w a l l e d , r e a c h i n g n e a r ­
l y to t h e l a t e r a l m a r g in s o f th e b o d y , and n e v e r r e a c h to
the acetabulum .
len g th .

They m easure from .096 to .120 mm i n

The e n t i r e a l i m e n t r y s y s te m i s w i t h i n th e f i r s t

f o u r th o f the body.
.0 6 7 mm i n d i a m e t e r .

The a c e ta b u lu m v a r i e s from .057 to
I t i s f r e q u e n t l y somewhat p r o t r u d e d ,

and l i e s a t a p o i n t somewhat a n t e r i o r to th e m id d le o f th e
bo d y .

The r a t i o b e tw e e n t h e two s u c k e r s have proved to be

p r a c t i c a l l y c o n s t a n t and a v e r a g e s 1 : 1 . 4 , t h e a c e ta b u lu m
b e i n g a b o u t h a l f a g a i n a s b i g a s th e o r a l s u c k e r .
The t e s t e s a r e s u b - s p h e r i c a l , p la c e d a b o u t one h a l f
t h e d i s t a n c e from t h e a c e ta b u lu m to th e p o s t e r i o r t i p o f
th e body.

They l i e i n th e same p l a n e , and n o t f a r from

t h e l a t e r a l m a r g in s o f th e body.
from .038 t o .07 mm.

They r a n g e i n d i a m e t e r

The v e r y l a r g e o r e s c e n t shaped

c i r r u s pouch i s l o c a t e d i n f r o n t o f t h e a c e t a b u l u m , e x ­
t e n d i n g a c r o s s p r a c t i c a l l y t h e e n t i r e w i d t h o f th e body

(34)

a t th a t poin t.

I n t h e p ro x im a l two t h i r d s o f th e s a c i s

t h e s e m i n a l v s e i c l e , which i s f o llo w e d by a r e g i o n f i l l e d
w i t h l a r g e p r o a t a t i c c e l l s , and t h i s i n t u r n i s fo llo w e d
i n t h e d i s t a l p o r t i o n by t h e l o n g , n a r r o w , p r o t r u s i b l e
cirrus*

The c i r r u s i s s p i n e d , and i n many specim ens i s

e i t h e r e x t r u d e d or i n c o p u l a .

The fem ale g e n i t a l opening

i s c l o s e to t h e male o p e n i n g , u s u a l l y l y i n g j u s t beh in d i t .
Both o p e n i n g s l i e i n a g e n i t a l groove which i s a t th e l e f t
m a rg in o f t h e a c e t a b u l u m , t h e o p e n i n g s b e i n g i n th e f a c e
o f th e d e p r e s s i o n o p p o s i t e th e w a l l o f t h e a c e t a b u l u m .
The o v a r y i s d o r s a l and to t h e r i g h t o f th e a c e t a b u l u m ,
and m e a su r e s .041 - *051 by .064 - .07 mm.

Hear the

p o s t e r i o r b o r d e r o f t h e o v a r y i n some specim ens i s d i s ­
tin g u is h a b le a sm all s h e l l g la n d .

The u t e r u s i s q u i t e

v o l u m i n o u s , f i l l i n g most o f th e body b e h in d th e a c e t a b ­
ulum and e x t e n d i n g up to a b o u t t h e h e i g h t o f t h e p r e ­
pharynx i n a l o o p on e i t h e r s i d e o f th e b o d y .
t e n s i o n forw ard i s u n i q u e f o r th e g e n u s .

T his e x ­

The eggs a r e

r a t h e r l a r g e , y e l l o w i s h brown, and measure .009 - .018
by . 0 1 9 - .022 mm.
The v i t e l l a r i a a r e l o c a t e d l a t e r a l and v e n t r a l to
th e t e s t e s , t u r n i n g m e d i a l and d o r s a l i n f r o n t o f the
te ste s.

They do n o t h o w ev e r, meet i n th e m i d - l i n e o r

extend to th e p o s t e r i o r e x t r e m i t y .

The f o l l i c l e s a r e

n o t l a r g e , and a r e i n s m a l l c l u s t e r s .

(35)

The e x c r e t o r y b l a d d e r i s v e r y c o n s p i c u o u s , and i n
a s t a t e o f d i s t e n t i o n i s d i s t i n c t l y "V” s h a p e d , w hereas
when c o n t r a c t e d i t a p p e a r s "Y" s h a p e d .

The h o r n s o f t h e

b l a d d e r e x ten d a s f a r a n t e r i o r a s th e a n t e r i o r b o r d e r o f
t h e t e s t e s and p a s s j u s t m e s i a l to t h e i n n e r b o r d e r o f i t .
H ost:
a ffin is

G rla u c io n e tta c l a n g u l a a m e r ic a n a and Nyroca

(In testin e)

L ocality:
type:

M i c h i g a n , U. S. A.

U. S. H a t ' l Mus. No. 9122

P a r a t y p e Nos. 9123, 9124

M a r it r e m a n e t t a e i s p e r h a p s most l i k e th e two s p e c i e s
d e s c r i b e d by Van Oleve and M u e l l e r , b e i n g i n most r e s p e c t s
i n t e r m e d i a t e betw e en th e two.

However t h e r e was enough

d i f f e r e n c e to w a r r a n t i t s s e p a r a t i o n from th o s e s p e c i e s ,
f o r a l t h o u g h t h o s e a u t h o r s s t a t e t h a t t h e i r two s p e c i e s
may be synonymous, I b e l i e v e them to be d i s t i n c t .

In

shape M. n e t t a e i s much more t r u n c a t e a t t h e p o s t e r i o r
end th a n e i t h e r o f t h e Oneida Lake forms*

The g e n e r a l

r e l a t i o n s h i p s o f t h e o r g a n s i n M. n e t t a e i s more l i k e
M. o b s t i p u s th a n an y o f t h e o t h e r d e s c r i b e d s p e c i e s ,
b u t i t may be d i s t i n g u i s h e d from t h a t s p e c i e s by the
d i f f e r e n c e i n r a t i o betw e en the s u c k e r s , the g r e a t e r
e x t e n t o f t h e c r u r a and t h e c i r r u s pouch.

In no c a s e

was t h e c o n d i t i o n o f t h e s u c k e r s , c r u r a , and c i r r u s o f

(36)

M. meduia a p p r o x im a te d by M. n e t t a e .

I n M. n e t t a e

t h e r e i s aome v a r i a t i o n w i t h r e g a r d to t h e v i t e l l a r i a .
I n some c a s e s t h e y extend a b o u t a s f a r a s , or even s l i g h t l y
f u r t h e r t h a n , t h e c o n d i t i o n d e s c r i b e d f o r M. o b s t i p u s ,
w h i l e i n o t h e r i n d i v i d u a l s t h e y extend o n ly a b o u t a s f a r
p o s t e r i o r a s t h e p o s t e r i o r b o r d e r o f th e t e s t e s .

The

a c e t a b u l u m i n M. n e t t a e was n o t tu r n e d to th e l e f t a s
d e s c r i b e d f o r M. o b s t i p u s .

This p o i n t has p r o b a b l y been

o v e r s t r e s s e d by Van Gleve and M u e l l e r , f o r such an o r i e n t ­
a t i o n c ould e a s i l y be a n a r t i f a c t o f p r e s e r v a t i o n or
pressure.
A l l i n a l l t h e r e l a t i o n s h i p s w i t h i n t h i s genus a r e
v e r y p o o r l y u n d e r s t o o d , and i n f a c t t h e r e i s l i t t l e b a s i s
f o r s e t t i n g up s p e c i f i c c h a r a c t e r s a s y e t .

When th e l i f e

h i s t o r i e s o f t h e s e s p e c i e s a r e 7/orfced o u t i t may throw
some l i g h t on t h e s p e c i f i c r e l a t i o n s w i t h i n th e g e n u s .

(37)

Key t o t h e S p e c i e s o f the genus M aritrem a

1.

I n t e s t i n a l o r u r a n o t r e a c h i n g to c i r r u s pouch--M. medius
I n t e s t i n a l c r u r a r e a c h i n g to a c e ta b u lu m or b ey o n d
2

2.

I n t e s t i n a l c r u r a r e a c h i n g to o r n e a r l y to a n t e r i o r
m a rg in o f a c e t a b u l u m ...............................................................................3
I n t e s t i n a l c r u r a r e a c h i n g beyond a n t e r i o r b o r d e r o f
a c e t a b u l u m ....................
7

3*

Orura e q u a l t o o r l e s s th a n e s o p h a g u s ............. M. l l n g u i l l a
C ru ra l o n g e r th a n e s o p h a g u s .....................................T ...................... 4

4.

C e n t r a l s u c k e r e q u a l to o r s m a l l e r th a n o r a l s u c k e r . . . . 5
"Ventral s u c k e r l a r g e r th a n o r a l s u c k e r ......................................6

5.

V e n t r a l s u c k e r and o r a l s u c k e r e q u a l , l y i n g in middle
o f b o d y ...................................................................................M. obs t i p u s
V e n t r a l s u c k e r s m a l l e r th a n o r a l s u c k e r , l y i n g b e f o r e
m id d le o f b o d y ................................................................... .M. lepidum

6.

C i r r u s pouch much c o i l e d , pharynx l e s s th a n h a l f
d i a m e t e r o f o r a l s u c k e r .................
M. hum ile
C i r r u s pouch n o t c o i l e d n o t i c e a b l y , pharynx more tha n
h a l f diam eter o f o r a l su ck er, v i t e l l a r i a not reaching
p o s t e r i o r e n d . ...........................
.M. n e t t a e n . s p .

7.

V e n t r a l s u c k e r e q u a l to o r s m a l l e r tha n o r a l s u c k e r . . . . 8
V e n t r a l s u c k e r l a r g e r th a n o r a l s u c k e r ........... M. g r a t i o s u m

8.

Ovary i n m i d - l i n e , d o r s a l and p o s t e r i o r to a c e t a b u l u m . . .
......................... ..
M. a c a d i a e
Ovary to r i g h t o f a c e t a b u l u m ............... ....................M. subdolum

(38)

Genus 3 p e l o t r e m a L e v i n s e n , 1881

So f a r no r e c o r d o f t h i s genus from Worth America
h a s b e e n found i n t h e l i t e r a t u r e .

I t may be s e p a r a t e d

from t h e c l o s e s t genus ( S p e l l o p h a l m u s ) by t h e f a c t t h a t
t h e male c o p u l a t i o n o rg an c o n s i s t s o f a m usc ular p a p i l l a e ,
w herea s i n t h e l a t t e r s p e c i e s t h e r e a r e co m p lic a te d s e r i e s
o f p o c k e t s m e s i a l to t h e p a p i l l a e #

J e g e r s k i o l d has d i s ­

c u s se d t h e genus (.38) and g i v e n a good d e s c r i p t i o n o f th e
ty p e s p e c i e s ( 3 7 ) •

Four s p e c i e s have b ee n d e s c r i b e d i n

t h e g e n u s , one o f which h a s b een d e s c r i b e d from ducks#
S p e lo tr e m a pygmaem ( L e v i n s e n , 1881) h a s b ee n r e c o r d e d from
Onidema n i g r a and S o m a t e r i a m o l l i s e m i a i n Europe#
Among t h e many spe cim e ns o f M a ritre m a n e t t a e , r e c o v e r e d
from th e g o l d e n - e y e s m entioned i n the l a s t s e c t i o n , have
b e e n found i n ea ch c a s e a f l u k e s l i g h t l y l a r g e r th a n them,
and d i s t i n c t l y d i f f e r e n t i n a p p e a r a n c e ,

The l a t t e r f l u k e s

were d i s t i n c t l y tongue shaped and more e l o n g a t e th a n th e
form er.

They were n e v e r in v e r y l a r g e num bers, a b o u t 25

b e i n g t h e maximum number from any one h o s t .

These f l u k e s

have b ee n i d e n t i f i e d a s S p elo trem a pygmaem. and s i n c e th e y
h ave b e e n a d e q u a t e l y d e s c r i b e d by J a g e r s k i o l d , Cdhner and
N i c o l l , no d i s c u s s i o n o f th e anatomy o f t h e s e forms w i l l
be g i v e n h e r e .

They were n e v e r i n g r e a t enough numbers to

(39)

be o f s i g n i f i o a n . e e to t h e h o s t *

T h i s , ho w ev e r, c o n s t i t u t e s

a new h o s t r e c o r d a s w e l l a s the f i r s t r e p o r t o f th e form
from N o r t h M e r i c $ .

F am ily O y c l o o o e l i d a e K ossack, 1911
Genus Typhlocoelum S t o s s i o h , 1908
Typhlocoelum buoumerinum ( R u d o lp h i, 1809) and Typhlocoelum
oymbium( B1es i n g . 1850}
The genus Typhlocoelum was e r e c t e d by S t o s s i o h (126)
to c o n t a i n Typhlocoelum flavum ( M e h li s , 1931) a s t y p e ,
and

eucumerinum ( R u d o l p h i , 1809) a s a v a l i d s p e c i e s , and

a s d u b io u s f o r m s , Monostomum s p . (M agelhas, 1 8 8 8 ) , and
Monstomum s a r c i d i o r n i c o l a (H e gnin, 1 8 9 0 ) .

Kossack (49) r e c ­

o g n iz e d T. flavum and T. eucumerinum a s synon§®ous, th u s
making T. eucumerinum t h e ty p e o f t h e genus*

He r e t a i n e d

b o t h d u b io u s forms o f S t o s s i o h a s " s p e c i e s i n q u i r e n d a e " ,
and added Monostomum cymbium D i e s i n g , 1850*

S k r j a b i n (115)

e r e c t e d t h e genus T r a c h e o p h i l u s to c o n t a i n h i s new s p e c i e s
T*

3 iaow i,

and added to i t <T* (Monostomum) cymbium ( D i e s i n g ,

1 8 5 0 ) , T. s a r c i d i o r n i c o l a (Megnin, 1 8 9 0 ) , and a s a d o u b t f u l
s p e c i e s , T* oboval (Naumann, 1 9 0 9 ).

He d i f f e r e n t i a t e d th e

genus T r a c h e o p h i l u s from Typhlocoelum s o l e l y on the l o b i n g o f
t h e t e s t e s i n th e l a t t e r and th e e n t i r e c o n d i t i o n i n th e

(40)

form er.

W i te n b e rg (162) and S z i d a t (137) have p o in te d

o u t t h a t T. a i a o w i i a p r o b a b l y synonymous w ith T. cymbium.
bub t h e y r e t a i n e d t h e g e n u s .

In f a c t W itenberg e r e c t e d a

new genus Typhlultim um t o c o n t a i n s p e c i e s T. a a r c i d i o r n i o o l a (Megnin, 1 8 9 0 ) .

Joyeaux and Bear (46) have

shown t h a t T r a c h e o p h i l u s ( S k r j a b i n , 1913) and Typhlultimum
( W ite n b e r g ) a r e synonymous w i t h Typhlocoelum S t o s s i o h , 1902.
They r e c o g n i z e d b u t two v a l i d s p e c i e s .

Typhlocoelum am e ric a n a

d e s c r i b e d by M anter and W illiam s (71) i s w i t h o u t a doubt
synonymous w i t h T. eucumerinum.

The two v a l i d s p e c i e s w ith

t h e i r synonyms a r e :
Typhlocoelum eucumerinum ( R u d o lp h i, 1809)
Synonyms;

Monostomum flavum ( M e h l i s , 1 8 3 1 ) , Monostomum

a a r c i d i o r n i o o l a (Megnin, 1 8 9 0 ) , Mono3tomum cyoymerlnum (Rud
d o l p h i , 1 8 0 9 ) , Typhlocoelum flavum ( M e h l i s , 1 8 3 1 ) , Typhlooelum o b o v a le (Naumann, 1909) , Typhlocoelum r e t l c u l i a r e
( J o h n s o n , 1 9 1 9 ) , Typhlocoelum s a r c i d i o r n i c o l a (Megnin, 1890)
W i t e n b e r g , 1924, Typhlocoelum am e r ic a n a (Manter and W i llia m s ,
1928).
Typhlocoelum oymbium ( D i e s i n g , 1850)
Synonyms:

Monostomum cymbium D i e s i n g , 1850, T r a c h e o p h i l u s

a i s o w l S k r j a b i n , 1913.
Both o f th e above mentioned s p e c i e s have been r e p o r t e d
from R o r t h A m e r ic a , and a s Stunfcard p o i n t e d o u t a r e p r o b a b l y

(41)

v ery cosm opolitan.

M a n ter and W illiam s (71) have r e p o r t e d

them from N ebraska from ducks (M a r l l a a f f i n i s . D a f i l a a c u t a ,
and S p a t u l a o l y p e a t a ) . and S tunkard h as r e p o r t e d T. cymbium
from New York from th e p i e d - b i l l e d g r e b e ( Podilymbus p o d l c e p s ).
The w r i t e r h a s ta k e n b o t h s p e c i e s a t t h e W. K. K ello g g B ird
S a n c t u a r y , from a snow goose ( Chen h y p e r b o r e a h y p e r b o r e a )
a s w e l l a s ducks (A* jd. p l a t y r h y n o h o s . A* p l a t y r h y n c h o s
( d o m e s t i c ) and A. r u b r i p e s ).
S a i d a t ( 1 4 2 , 143) and Stunkard (132) w orking in d e p e n d ­
e n t l y , have com pleted t h e l i f e - c y c l e o f T_. oymbium.

Their

r e s u l t s a r e i n com p lete a g r e e m e n t , and b r i e f l y th e l i f e - c y c l e
o f t h i s sp ecies is as follow s.
b e i n g e x p e l l e d from t h e h o s t .

The eggs h a t c h a t t h e time o f
The m i r a c i d i u m , when l i b e r a t e d ,

a l r e a d y c o n t a i n s a f u l l y formed r e d i a .

The m ir a c id iu m swims

a round i n th e w a t e r u n t i l a s u i t a b l e s n a i l (Eeliosom a) i s
found.

The m i r a c i d i u m does n o t e n t e r the s n a i l , b u t a t t a c h e s

i t s e l f to i t , r e l e a s i n g t h e r e d i a which b o r e s i n t o th e s n a i l .
T h e re a r e no s p o r o e y s t or d a u g h t e r r e d i a s t a g e s .

T ailless

c e r c a r i a a r e produced i n th e r e d i a which e n c y s t i n th e lymph
spaces o f the s n a i l w ithout a fre e l i v i n g p erio d .
i n f e e t e d s n a i l i s i n g e s t e d by a

When an

w a t e r b i r d , th e worms e x c y s t

i n th e s m a l l i n t e s t i n e and f in d t h e i r way i n t o th e a i r
passages.
known.

By j u s t what r o u t e t h i s i s ac co m p lish ed i s n o t

Two p o s s i b l e r o u t e s have been s u g g e s t e d ; one by way

(42)

o f t h e b lo o d s y s t e m , t h e o t h e r by th e body c a v i t y and a i r
spaces.
S z i d a t was a b l e to r e c o v e r o n ly two A d u lt forms o f
T. cymbium from a duck a f t e r f e e d i n g 50-60 c y s t s , and
S t u n k a r d , u s i n g a duck a l s o , r e c o v e r e d none a t a l l a f t e r
feeding 6 c y s ts .

Most o f th e o b s e r v a t i o n s on t h i s genus

have i n d i c a t e d t h a t an i n f e c t e d b i r d seldom c a r r i e s more
th a n f i v e o r s i x worms.

My f i n d i n g s on 12 i n f e c t e d b i r d s

from W l n t e r g r e e n l a k e b e a r t h i s o u t to a s t r i k i n g d e g r e e ,
f o r I have found no more th a n t h r e e worms p r e s e n t i n any
b u t one o f t h e s e c a s e s .

The u s u a l p l a c e o f a t t a c h m e n t i s

i n t h e t r a c h e a j u s t i n s i d e th e g l o t t i s .

I t is h ig h ly prob­

a b l e t h a t an a d u l t b i r d seldom i s p a r a s i t i z e d by more than
f o u r or f i v e o f t h e worms.

Such l i m i t a t i o n on th e numbers

o f p a r a s i t e s i s due e i t h e r to o n ly a few worms r e a c h i n g
the d e f i n i t i v e p o s i t i o n in t h e h o s t or to the r a p id e x p u l ­
s i o n o f a l l b u t a few o f t h e p a r a s i t e s by th e h o s t .

This

l a t t e r h y p o t h e s i s seems th e more l i k e l y , i n view o f t h e
f a c t t h a t t h e p a r a s i t e s a r e p r a c t i c a l l y always n e a r th e
d i s t a l end o f the t r a c h e a or i n th e n a s a l s p a c e s .

C ertainly

a l a r g e number o f worms o f t h i s s i z e in the t r a c h e a o f a
b i r d would be c e r t a i n to produce a coughing and e x p e l l i n g
r e a c t i o n on t h e p a r t o f t h e h o s t .

The f a c t t h a t S z i d a t

r e c o v e r e d o n l y two worms from 5Q**60 c y s t s i s e a s i l y ex—

(43)

p l a i n e d on t h i a b a s i s , a s i s a l s o th e c a se o f Stun^ard
m e ntio ne d a b o v e , f o r t h e i r a n i m a l s were examined e i g h t
to t e n weeks a f t e r i n f e c t i o n ,

S z i d a t s t a t e s t h a t s i x days

a f t e r f e e d i n g c y s t s th e young worms had reac h ed th e t r a o h e a
and l u n g s , which f u r t h e r b e a r s o u t t h i s p o i n t .

I t takes

a b o u t s i x weeks f o r t h e worm to r e a c h m a t u r i t y .

C ertainly

any g r e a t number o f worms d e v e l o p i n g i n th e t r a c h e a and
l u n g s o f a b i r d o v e r a p e r i o d o f two o r t h r e e months would
pro d u ce e x p u l s i v e a c t i o n on th e p a r t o f t h e h o s t .
T h a t t h e d uck, i f u n a b le to e x p e l l a l l b u t a few o f
t h e p a r a s i t e s , may s u f f e r undue consequences i s shown by
th e f o l l o w i n g c a s e .

In J u l y , 1936, a h a l f - g r o w n m a l l a r d duck

was n o t i c e d on th e bank o f a sw ale a p p a r e n t l y d e a d .

It

s u d d e n l y j e r k e d up i t s h e a d , a l t h o u g h u n a b le to r i s e , and
gasped s p a s m o d i c a l l y s e v e r a l t i m e s , and the n s e t t l e d b ac k
a g a i n a s i f i n a coma.

I t was k i l l e d and a u t o p s i e d im m ediate­

l y and 19 f u l l y d ev e lo p ed and m a tu r e Typhlocoelum cuoumerinum
7/e r e

ta k e n from t h e n a s a l p a s s a g e s , t r a c h e a , l u n g s , a i r

s a c s , and coelom.

T his duck was n o t over two months old a t

t h e time o f a u t o p s y .

A lth o u g h t h e r e v/ere seven s p e c i e s o f

p a r a s i t e s i n th e i n t e s t i n e , the most numerous o f which
were C e s to d e s o f t h e genus F i m b r i a r i a . th e y were n o t i n
g r e a t enough abundance to have caused the c o n d i t i o n o f
the duck.

O th er ducks o f th e same age even more h e a v i l y

(44)

p a r a s i t i z e d w i t h i n t e s t i n a l p a r a s i t e s n e v e r showed such
symptoms.

A p p a r e n t l y , due to age or w eak ness, t h i s duck­

l i n g had b ee n u n a b l e to throw o f f th e p a r a s i t e s , and th e
heavy i n f e s t a t i o n r e s u l t e d .

The d a t a tend to s u p p o r t the

h y p o t h e s i s t h a t t h e worms r e a c h t h e i r d e f i n i t i v e p o s i t i o n
by m i g r a t i o n t h r o u g h th e blood s y s t e m , a l t h o u g h i t i s
q u i t e p o s s i b l e t h a t th e y m ight have m i g r a t e d from th e lu n g s
i n t o t h e a i r s p a c e s , t h e n c e i n t o t h e body c a v i t y .

S ince

Szid& t h as shown t h a t a t l e a s t s i x weeks i s r e q u i r e d f o r
t h e s e f l u k e s to m a t u r e , i t i s e v i d e n t t h a t t h i s duck must
have i n c u r r e d the i n f e s t a t i o n a t a v e r y e a r l y a g e , and t h i s
may e x p l a i n th e s e r i o u s r e s u l t .
Taxonomy o f t h e Senus Typhlocoelum
Morgan (84) i n h i s s t u d y on t h e f a m i l y O p i s t h o r c h i i d a e
h a s b r o u g h t o u t t h e f a c t t h a t th e shape o f the t e s t e s (lobed
or e n t i r e ) i s n o t , i n i t s e l f , a v a l i d g e n e r i c c h a r a c t e r
w ith in th a t fam ily.

Joyeaux and B ear (46) have a p p l i e d

t h i s r u l e in the case o f T ra o h eo p h ilu s. d e c la r in g i t
synonymous w i t h Ty p h lo c o e lu m .

The synonymy o f t h e s e forms

had b e e n s t a t e d a s a p r o b a b i l i t y by W itenberg (162) and
S zid a t (137).

S k r j a b i n (115) i n s e t t i n g up the genus

T r a o h e o p h i l u s , s t a t e d t h a t th e p r i n c i p a l d i f f e r e n c e b e ­
tween i t and Typhlocoelum was t h e unbranched b l i n d s a c s

^

A

Text f i g u r e 1 .

A a e r i e s o f Typhlocoelum showing th e

g r a d a t i o n o f t h e t e s t e s from an e n t i r e to
a lo b e d c o n d i t i o n *

The v a r i a t i o n i n a r r a n g e ­

ment o f t h e s e x g l a n d s , and th e p o s t e r i o r
e x t e n t o f t h e v i t e l l a r i a a r e a l s o shown*

o f t h e i n t e s t i n e and t h e e n t i r e t e s t e s *

The f i r s t c h a r a c t e r

i s , o f c o u r s e , d e p e n d e n t on t h e c o n d i t i o n o f th e p a r a s i t e a t
t h e tim e o f f i x i n g , w h i l e th e s e c o n d , a s s t a t e d above i s
c e r t a i n l y o f no more t h a n s p e c i f i c v a l u e .
A com p ariso n o f t h e anatomy o f th e two v a l i d s p e c i e s

(46)

—! • oymblum and T. cucumerinum--shows a t once t h a t in no
c h a r a c t e r o t h e r t h a n s h a p e o f t e s t e s do t h e s e two s p e c i e s
d i f f e r beyond th e e x t e n t o f i n d i v i d u a l v a r i a t i o n .

A care­

f u l co m p a riso n o f t h e two s p e c i e s shows t h e s i z e and shape
o f th e b o d y , e x t e n t and c h a r a c t e r o f v i t e l l a r i a , e x c r e t o r y
b l a d d e r , e x t e n t o f u t e r u s , p h a r y n x , mouth c a v i t y , p o s i t i o n
o f g e n i t a l p o r e , c i r r u s , and i n f a c t a l l o t h e r c h a r a c t e r s ,
to be e a s i l y w i t h i n th e r a n g e o f v a r i a t i o n found i n a s i n g l e
species.

C h a r a c t e r s f o r m e r l y c o n s i d e r e d to be o f s p e c i f i c ,

and even g e n e r i c v a l u e , such a s t h e p o s t e r i o r e x t e n t o f the
v i t e l l a r i a , l o b i n g o f t h e i n t e s t i n a l d i v e r t i c u l a e t c . , have
b ee n shown to be p u r e l y i h d i v i d u a l i s t i c (3ee t e x t f i g u r e 1 . ) *
A co m p a riso n o f o v e r 50 specim ens a s to the e x t e n t o f l o b i n g
o f t h e t e s t e s r a i s e s a q u e s t i o n a s to t h e v a l i d i t y o f t h i s
c h a r a c t e r , f o r a n a l m o s t p e r f e c t g r a d a t i o n has been n o t e d .
The p r e s e n c e o f a r u d i m e n t a r y a c e ta b u lu m i n T. cucumerinum
h a s b e e n t a k e n by some a u t h o r s a s s i g n i f i c a n t .

Stunkard

h a s shown t h a t T. cymbium h a s an a c e ta b u lu m i n th e l a r v a l
stag es.

The w r i t e r h a s found t h i s a p u r e l y i n d i v i d u a l c h a r ­

a c t e r i n specim ens o f b o t h s p e c i e 3 .

A s t u d y has been made o f

th e e g g s , m i r a c i d i a , and r e d i a o f T. cucumerinum, and the y
compare i n e v e r y r e s p e c t w i t h th e c o r r e s p o n d i n g s t a g e s d e s —
o r i b e d b y S s i d a t (1 4 8 , 143) and Stunkard (138) f o r T. cambium.

(4?)

When t h e l i f e h i s t o r y o f T. cucumerinum has been completed
i t w i l l be p o s s i b l e to d e f i n i t e l y sa y w hether t h e s e two
s p e c i e s a r e v a l i d o r synonymous.

At p r e s e n t a r e a s o n a b l e

doubt e x i s t s a s to t h e v a l i d i t y o f two s p e c i e s .

Fam ily N o t o c o t l y i d a e Luhe, 1909
Genus N o t o c o t y l u s D i e s i n g , 1839
N o t o o o t y l u s u r b a n e n a i 8 ( C o r t , 1914)
Harrah has d escrib ed the a d u l t o f t h i s s p e c ie s r a t h e r
b r i e f l y (34).

He has assumed t h a t he was d e a l i n g w ith the

a d u l t o f P e r c a r i a u r b a n e n s i s P o r t , 1914, w i t h o u t e x p e r i ­
m e n ta lly proving the r e l a t i o n s h i p .

Such a p r o c e e d u r e i s

o f c o u r s e , open to q u e s t i o n , and may o n ly be proved by
e x p e rim e n ta lly com pleting th e l i f e h i s t o r y .

The m a t e r i a l

upon which h i s d e s c r i p t i o n was b as ed was c o l l e c t e d by D r.
A l b e r t H a s s e l l i n Maryland from two ducks ( D a f i l a a c u t a
and Aix s p o n s a ) and a m u s k r a t ( F i b e r z i b e t h i c u s ) .

These

worms have b e e n ta k e n tw ic e i n t h i s s t u d y , i n one case
o n l y one specim en b e i n g p r e s e n t , i n the o t h e r 16 sp e c im e n s .
Both h o s t s were m a l l a r d s (Anas p l a t y r h y n c h o s p l a t y r h y n c h o s ) .
T h i s c o n s t i t u t e s a new h o s t and l o c a l i t y r e c o r d f o r t h e s e
p arasites.
S i n c e H a r r a h ' s d e s c r i p t i o n i s v e r y b r i e f , I have deemed

(48)

i t n e c e s s a r y to r e d e s c r i b e t h i s s p e c i e s from m a t e r i a l i n
my c o l l e c t i o n .

Medium s i z e d , e l o n g a t e f l u k e s , the a n t e r i o r p o r t i o n
o f t h e body t h i c k l y s e t w ith f i n e s p i n e s .

A n t e r i o r end

somewhat t a p e r i n g , p o s t e r i o r end more rounded.
2 . 5 to 4 . 2 mm, g r e a t e s t w i d t h .5 to 1 . 0 mm.
w e l l d e v e l o p e d , .1 4 to .2 mm i n d i a m e t e r .

Total le n g th

O ral s u c k e r
On th e v e n t r a l

s u r f a c e a r e t h r e e rows o f g l a n d s , the number b e i n g i n most
c a s e s t h i r t e e n i n ea ch row, b u t o c c a s i o n a l l y 14 a r e p r e s e n t .
The g l a n d s

i n th e median row a r e much l a r g e r tha n the l a t e r a l

rows.

g l a n d s a p p e a r more o r l e s s c i r c u l a r and show con­

The

s id e r a b le v a r i a t i o n in diam eter.

The esophagus b i f u r c a t e s

a t a point

.25 to .35 mm from th e a n t e r i o r end.

The c r u r a

p ro cee d to

n e a r t h e p o s t e r i o r end o f t h e worm, t u r n i n g

mesiad i n t h e r e g i o n o f th e t e s t e s and p a s s i n g i n t e r n a l to
those organs.

The c r u r a w a ll i s i r r e g u l a r , the c o n d i t i o n

c a l l e d by H a r r a h "numerous s h o r t d i v e r t i c u l a " .

The t e s t e s

l i e n e a r t h e p o s t e r i o r end o f th e body, and a r e l a t e r a l ,
ly in g a t the

3 ame

lev el.

u l a r l y on th e o u t e r s i d e .
from .4 8 t o .528 mm.

They a r e v e r y much l o b e d , p a r t i c ­
They measure i n th e l o n g a x i s

The ovary i s median between th e two

t e s t e s , and i s s e p a r a t e d from them by the i n t e s t i n a l c r u r a .
I t m e a s u r e s from .2 0 to .2 4 mm i n d i a m e t e r ana may be lobed

(49)

or e n t i r e .

The s h e l l g la n d l i e s j u s t a n t e r i o r to the o v a r y ,

a s does t h e s e m i n a l reeept& culum .

The u t e r u s i s w i t h i n t h e

o r u r a and winds v e r y r e g u l a r l y , e x t e n d i n g up to a b o u t the
p osterior p art

of

t h e second body f i f t h .

u r e s 1 . 0 to 1 . 3 mm i n l e n g t h .

The c i r r u s meas­

The s e m in a l v e s i c l e i n th e

p o s t e r i o r p a r t o f th e s a c i s v e r y s i n u o u s .

The g e n i t a l

p o r e i s l o c a t e d .27 - .5 2 mm beh in d t h e a n t e r i o r t i p o f th e
b o d y , b e i n g n e a r t h e b i f u r c a t i o n o f th e i n t e s t i n e .

The eggs

a r e y e l l o w , and have two p o l a r f i l a m e n t s .
H ost;

D a f i l a a c u t a , Aix s n o n s a . Anas

platyrhynchos

and F i b e r z l b e t h i c u s ( I n t e s t i n e and caecum)
L o cality :

M a r y la n d , M ic h ig a n , U. S. A.

S in c e t h e t h r e e r e c o r d s o f E a r r a h ( H a s s e l l ) a r e the
o n ly r e c o r d s a s i d e from th e p r e s e n t o n e , the i n d i c a t i o n i s
t h a t , e c o n o m i c a l l y , th e f l u k e i s o f l i t t l e s i g n i f i c a n c e .

Fam ily E chino3tom idae D i e t z , 1910
Genus Echinostoma D i e t z , 1910
Bohinostoma r e v o lu tu rn ( F r o e l i c h , 1902)
T his form i s p e r h a p s th e most common h e l m i n t h p a r a ­
s i t e o f ducks.

I t i s p r a c t i c a l l y c o s m o p o lita n i n d i s t r i ­

b u t i o n and h a s b e e n r e c o r d e d from a number o f o t h e r b i r d s

and mamraala a s i d e from d u c k s .

Johnson(43) f i r s t r e p o r t e d

on th e co m p le te l i f e h i s t o r y , showing t h a t Phy 3 a o c o i d e n t a l i s
s e r v e d b o t h a s f i r s t and second i n t e r m e d i a t e h o s t s .

The

m i r a e i d i u m h a t c h i n a b o u t t h r e e weeks, p e n e t r a t e th e s n a i l
and p ro d u ce m o th e r r e d i a which i n t u r n produce d a u g h t e r r e d l a
i n which r e l a t i v e l y l a r g e o e r o a r i a a r e d e v e l o p e d .

The c e r c a r i a

l e a v e t h e s n a i l f o r a w h i l e , p e n e t r a t e the same s n a i l , and
en cy st w ith in a s h o r t tim e.

They a r e in t r o d u c e d i n t o th e

f i n a l h o s t a l o n g w i t h t h e s n a i l , and d e v e lo p i n t o l a r g e
f l u k e s , found u s u a l l y i n t h e c o lon o f th e f i n a l h o s t , b u t
a l s o i n t h e s m a l l i n t e s t i n e , b u r s a and even the caecum and
s h e l l gland a t t i m e s .

Subsequent to Johnson*s work a number

o f s n a i l s h ave been found to be i n f e c t e d w ith t h i s s p e c i e s .
Limnea o b r u s s a (Say) i n W i n te r g r e e n Lake have been found to
be i n f e c t e d b o t h w ith r e d i a and en c y ste d c e r c a r i a , and ex­
p e r i m e n t a l i n f e c t i o n s have been induced by f e e d i n g th e s n a i l s
to d u c k s . Beaver

(6) has p u b l i s h e d a v e r y comprehensive ex­

p e r i m e n t a l s t u d y o f t h i s form.
As to t h e s e r i o u s n e s s o f t h i s p a r a s i t e , I b e l i e v e i t
to be n e g l i g a b l e among d u c k s .

No p a t h o l o g i c a l consequ ences

have b e e n n o te d i n any i n s t a n c e o f i n f e c t i o n w ith t h e s e
form s.

They have occured i n a b o u t 90^ o f a l l th e ducks

ex a m in e d , t h e r e f r e q u e n t l y b e i n g a s many a s t h r e e or f o u r

(51)

age g r o u p s o f th e p a r a s i t e p r e s e n t .

I have found sm all

f l u k e s , j u s t o u t o f t h e c y s t i n th e upper duodenum.

They

move down t h e i n t e s t i n e r a t h e r s l o w l y , u s u a l l y b e i n g n e a r
t h e maximum s i z e when r e a c h i n g t h e c o l o n .

A lthough t h e r e

i3 no a c t u a l d a t a to i n d i c a t e th e l e n g t h o f l i f e i n t h e s e
i n d i v i d u a l s , from t h e known f a c t s o f o t h e r l i f e h i s t o r i e s ,
i t i s p r o b a b l e t h a t , u n d e r n a t u r a l c o n d i t i o n s , th e y rem ain
in t h e h o s t a r e l a t i v e l y s h o r t time a f t e r r e a c h i n g maximum
s i z e , f o r no more th a n f o u r or f i v e worms o f maximum s i z e
u s u a l l y a r e found i n a s i n g l e h o s t .

This f a c t coupled

w i t h t h e p r e s e n c e i n a l m o s t e v e r y d u ck , o f f l u k e s o f the
same s p e c i e s i n v a r i o u s s t a g e s o f development seem to i n ­
d i c a t e a b r i e f s t a y i n th e f i n a l h o s t .

(52)

Genus S t e p h a n o p r o r a Odhner, 1910
S t e p h a n o p r o r a s p i n o s u a Odhner, 1910
These worms have b e e n ta k e n o n ly once d u r i n g t h i s
s t u d y , a b o u t 25 h a v i n g been r e c o v e r e d from a g o ld e n -e y e
p ic k e d up i n B a t t l e C r e e k , M ic h ig a n .

The s p e c i e s has

b ee n d e s c r i b e d by B i e t z (25) a s w e l l a s Odhner ( 9 7 ) .
T his c o n s t i t u t e s a new l o c a l i t y and h o s t r e c o r d f o r th e
s p e c i e s , h a v i n g p r e v i o u s l y been r e c o r d e d only from g r e b e s
in Europe.

They may be d i s t i n g u i s h e d from th e o t h e r E ch in -

ostomes by t h e d o r s a l b r e a k , i n the c o l l a r s p i n e s , th e s h o r t ,
wide c i r r u s pouch, and th e l i m i t a t i o n o f th e v i t e l l a r i a to
t h e r e g i o n b e h in d t h e o v a r y .

22 c o l l a r s p i n e s a r e p r e s e n t

i n S. s p i n u l o s u 3 .

F am ily P s i l o s t o m i d a e Odhner, 1913
Cenus S p a e r i d i o t r e m a O dhner, 1913
S p a e r i d i o t r e m a g l o b u l u s ( R u d o lp h i, 1914)
T h is s p e c i e s h a s b een e n c o u n te r e d i n l e s s e r scaup
ducks and g o l d e n - e y e s .

In the l a t t e r d u c k 3 , however,

o n ly v e r y s m a l l numbers have been f o u n d , u s u a l l y n o t
o v e r a dozen specim ens b e i n g p r e s e n t .

From t h r e e l e s s e r

s c a u p ducks c o l l e c t e d a t th e W. K. K e llo g g Bird S a n c t u a r y

(53)

i n M arch, 1937, l i t e r a l l y h u n d red s o f t h e s e s m a ll worms
were ta k e n *

They were found c h i e f l y i n the p o s t e r i o r

p a r t o f the sm all i n t e s t i n e *
P r i c e (106) h as d e s c r i b e d a s e r i o u s f a t a l c o n d i t i o n
i n l e s s e r s c a u p duck3 i n t h e v i c i n i t y o f Washington D. 0.
a s due t o t h i s f lu k e *

He d e s c r i b e d t h e c o n d i t i o n a s an

e n t i r i t i s o f t h e lo w er s m a ll i n t e s t i n e due to th e denud­
i n g o f t h e w a l l o f e p i t h e l i u m by t h e f l u k e s .

I t appears

from h i s d i s c u s s i o n t h a t th e d i s e a s e had developed r a p i d l y ,
p r o b a b l y k i l l i n g t h e b i r d s w i t h i n a s h o r t time a f t e r the
onset o f the d isease*

P r i c e has a l s o r e d e s c r i b e d th e s p e c i e s

from h i s own m a t e r i a l owing to t h e i n a d e q u a t e d e s c r i p t i o n s
of previous a u th o rs .

A lthough t h i s p a r a s i t e has b ee n r e ­

p o r t e d from e i g h t o t h e r s p e c i e s o f w a te r b i r d s , P r i c e ' s
i s th e o n l y r e c o r d o f a p a t h o g e n i c c o n d i t i o n b e i n g caused
by t h e s e f l u k e s .

F am ily S t r i g e i d a e R a i l l i e t , 1919
Senus O o t y l u r u s S z i d a t , 1928
0o t y l u r u s f l a b f e l l i f o r m l s ( ^ a u s t , 1917)
T his genus was d e s c r i b e d by S z i d a t i n 1929 ( 1 3 8 ) , and
may b e e a s i l y d i s t i n g u i s h e d from o t h e r S t r i g e i d a e by the

(54)

a b s e n c e o f v i t e l l i n e g l a n d s from the f o r e body and the m u sc u lar
s u c k e r l i k e b u l b i n th e b u r a a c o p u l a t o r i x .

Van H a i t 3 ma (153)

has r e p o r t e d t h e o n l y r e c o r d o f t h i s genus from ducks i n
North A m e r ic a .
1917).

This form i s Co t y l u r u s f l a b e l l i f o r m i s ( F a u s t ,

T h is a u t h o r has d e s c r i b e d th e a d u l t s o f th e s p e c i e s

and worked o u t th e l i f e - c y c l e e x p e r i m e n t a l l y .
a s w e l l a s t h e t e t r a c o t y l e , occ ur

The c e r c a r i a ,

i n s n a i l s , b o th forms b e i n g

known to i n f e c t a t l e a s t f i v e d i f f e r e n t s p e c i e s o f t h e s e h o s t s .
S in ce th e anatomy and l i f e h i s t o r y o f t h i s form a r e d i s c u s s e d
i n d e t a i l by Van H a i t s m a , th e y w i l l be o u i t t e d h e r e .
Of t h e a l m o s t 200 ducks examined i n t h i s s t u d y , p r o b ­
a b l y 6 0 - 7 0 $ have been i n f e c t e d , and o f th o s e examined d u r i n g
th e summer s e a s o n , i n f e c t i o n has b een n e a r 100$.

The worms

were o f t e n p r e s e n t i n l a r g e numbers, u s u a l l y somewhat more
numerous i n th e p o s t e r i o r h a l f o f the sm a ll i n t e s t i n e , b u t
f r e q u e n t l y found th r o u g h o u t t h e i n t e s t i n e from th e g i z z a r d
to th e i l i o e a e c a l v a l v e .

I n most o a s e s the y were f i r m l y

a t t a c h e d to t h e mucosa, and came l o o s e o n ly a f t e r f i v e to
t e n h o u r s im m ersion i n w a t e r .

A f t e r th e prolonged imm ersion,

by v i g e r o u s l y s h a k i n g the d i s h , many o f them were washed f r e e
and e a s i l y c o l l e c t e d from th e b o tto m o f the c o n t a i n e r *
Van H aitsm a h a s d i s c u s s e d b r i e f l y th e p a t h o lo g y o f
t h e s e p a r a s i t e s a s obse rv e d i n h i s e x p e r i m e n t a l b i r d s , b u t

(85)

s t a t e s t h a t g r e a t u n c e r t a i n l y e x i s t s i n t h i s r e g a r d due
to t h e c o n d i t i o n s u nder which th e o i r d s were m a in ta in e d *

It

i s h i g h l y p r o b a b l e t h a t , a l t h o u g h the p a t h o l o g y he d e s c r i b e d
may have been due to t h e p a r s s i t e s , i t was i n f l u e n c e d to
a l a r g e e x t e n t by t h e u n n a t u r a l c o n d i t i o n s under which the
b i r d s were k ept*

B i r d s r a i s e d and h e ld by the a u t h o r under

conditions q u ite sim ila r

to th o s e d e s c r i b e d by Van H aitsm a,

b u t n o t i n f e o t e d w i t h any p a r a s i t e s , have e x h i b i t e d q u i t e
s t r i k i n g l y , a l t h o u g h p r o b a b l y to a l e s s e r e x t e n t , th e symptoms
c i t e d by Van H aitsm a f o r b i r d s i n f e c t e d w ith _C. f l a b e l l l f o r m l s .
The denuded e p i t h e l i a l r e g i o n s caused by the a t ta c h m e n t
o f t h e s e worms have been s e e n i n a number o f c a s e s , b u t o n ly
two c a s e s o f extrem e p a t h o l o g y have been noted in t h i s s t u d y .
Two l e s s e r s c a u p d u c k s , c o l l e c t e d f o r mounting a t t h e W. K.
K e ll o g g B i r d S a n c t u a r y , were observed to be t h i n when t a k e n ,
and an e x a m i n a t i o n o f the i n t e s t i n e s o f t h e s e b i r d s r e v e a l e d
l a r g e numbers o f £ . f l a b e l l i f o r m i s a l o n g w ith much s m a l l e r
numbers o f two o t h e r s p e c i e s o f t r e m a t o d e s .

A lso a number

o f heromoragic a r e a s v/ere noted on the e p i t h e l i u m , i n some
c a s e s a blood c l o t b e i n g p r e s e n t .

When t h e s e c l o t s mere

b ro k e n open th e y were found to c o n t a i n , sometim es, a s many a s
f iv e 0. f l a b e l l i f o r m i s .

N e i t h e r o f th e o t h e r s p e c i e s o f

tr e m a t o d e s were noted n e a r or i n any o f t h e c l o t s .

(56)

A lth o u g h t h e work o f Van Haitsma h a s shown t h a t th e
i n d i v i d u a l o f t h i s s p e c i e s a r e v e r y s h o r t l i v e d , i n the
f i n a l h o s t ( d u c k s ) , o f t e n r e m a i n i n g l e s s th a n two weeks,
t h i s i n v e s t i g a t i o n h a s found th e s p e c i e s to be one o f the
most c o n s t a n t l y p r e s e n t o f a l l th e tr e m a to d e s o f ducks
e n c o u n te r e d , b e i n g second i n p r e v a l e n c e only to the common
Echinostome ( Echinostoma r e v o l u t u m ) .

Because o f t h i s

v e r y g r e a t abundance i n m i g r a t i n g ducks ( b o th s p r i n g and
f a l l ) , i t i s s a f e to s a y t h a t t h i s p a r a s i t e i s p r o b a b ly
one o f th e commonest o f a l l duck t r e m a t o d e s .

Its a b ility

to u s e a number o f s n a i l h o s t s f o r development o f i n t e r ­
m e d i a t e s t a g e s , and p r o b a b l e p a t h o l o g i c a l e f f e c t when
p r e s e n t i n l a r g e num bers, make i t i m p o r t a n t t h a t any one
r a i s i n g v e r y many ducks guard a g a i n s t i n f e c t i o n w ith t h i s
species.
C o t y l u r u s h as been ta k e n from the f o l l o w i n g s p e c i e s
o f ducks i n t h i s s t u d y :

Anas p la t y r h y n c h o s p l a t y r h y n c h o s ,

Anas r u b r i p e s . G l a u e i o n e t t a o l a n g u l a a m e r i c a n a . Querquedula
d i a c o r s , N e t t i o n c a r o l l n e n a i s . Mareca a m e r i c a n a , Nyroca
a f f i n i s , Ayroca c o l l a r i s . S p a t u l a c l y p e a t a , and Nyrooa
m arila.

(57)

F a m ily Earmoatomidae Odhner, 1912
S u b fa m ily Earmoatominae Loosa, 1900
W ite n b e rg (161) and Werby (158) have d i a c u s s e d th e
o l a a s i f i o a t i o n o f t h i s group i n d e t a i l .

There a r e f o u r

t r i b e s r e c o g n i z e d i n t h i s s u b f a m i l y , namely:
Harm os t o m e a . U r o t o c a e . and L e u o o o h l o r i d e a .

I t h y o g o n im a e .
This group i s

f o r t h e moat p a r t , l a r g e l y p a r a s i t i c i n b i r d s , b u t a c a r e f u l
a t u d y o f t h e l i t e r a t u r e h a s r e v e a l e d no r e p o r t o f t h e s e
p a r a s i t e s from d u c k s .
From th e b u r s a F a b r e c i i o f an immature b l a c k duck
(Anas r u b r i p e s ) , c o l l e c t e d a t W in te rg re e n Lake on O cto b e r
8 , 1936, two f l u k e s were o b t a i n e d , w hich, a t f i r s t , were
ta k e n to b e l o n g to t h e genus P ro s t h o g o n im u s .

Eowever,

when examined u n d e r a low power d i s s e c t i n g m ic r o s c o p e ,
i t became e v i d e n t t h a t t h e y were v e r y u n u s u a l fo rm s , a t
l e a s t f o r t h a t genus.

The f l u k e s were p i n k i s h - f l e s h in

c o l o r — t y p i c a l o f Prosthogonim us and assumed ab o u t the
same p e a r sh a p e i n w a t e r - - b u t th e e n t i r e v e n t r a l s u r f a c e
a p p e a re d to be a huge s u c k e r .

The s t r u c t u r e was even

more e v i d e n t v/hen the worms were s l i g h t l y f l a t t e n e d b e ­
tween two s l i d e s and examined a l i v e .

L ittle d e ta il of

the anatomy c o u ld be made o u t i n the l i v i n g worm due

(58)

to th e o p a c i t y o f t h e s k i n , so th e y were f ix e d and s t a i n e d .
U n f o r t u n a t e l y , one o f t h e specimens was l o s t i n th e s t a i n ,
and t h e f o l l o w i n g d e s c r i p t i o n i s n e c e s s a r i l y based on a
s i n g l e s p e c im e n .

A f t e r a v e r y c a r e f u l s t u d y o f the a n a t ­

omy o f t h i s s p e c im e n , i t has been found to go i n t o the
Tribe L e u c o c h lo rid ea , b u t sin c e

it

i s q u i t e d i f f e r e n t from

th e s i n g l e genus o f t h a t T r i b e ,

it

has been n e c e s s a r y to

e r e c t a new genus to c o n t a i n i t

as type.

L e u o o c h lo rid io m o rp h a n .
D iagnosis:
sim ilar

Leucochloridea.

g.

In g e n e ra l, q u ite

to t h e Genus L e u c o c h io r id iu r o .

I t d i f f e r s from

t h a t genus i n t h a t th e v i t e l l a r i a do n o t extend to th e
p o s t e r i o r p o r t i o n o f t h e worm, b e i n g l i m i t e d to t h e
a n t e r i o r p o r t i o n o f t h e worm.

Acetabulum v e r y l a r g e ,

much l a r g e r th a n t h e o r a l s u c k e r .

T e s t e s o b l i q u e , in

p o s t e r i o r f o u r t h o f b o d y , th e a n t e r i o r t e s t e s b e i n g to
th e l e f t o f th e m i d - l i n e .

The u t e r u s does n o t c r o s s i n

f r o n t o f the a c e t a b u l u m , b u t has an a s o e n d in g and d e s c e n d ­
i n g lo o p on e i t h e r s i d e o f t h e a c e ta b u lu m .

A w e ll d e v e l ­

oped c i r r u s pouch i s p r e s e n t , and c o n t a i n s a s p i n e d , p r o t r u s i b l e c i r r u s , th e pore o f which i s s u b t e r m i n a l and d o r sa l.

uv a r y s l i g h t l y l a t e r a l and between t h e two t e s t e s .

(59)

Type a p e e i e a :

M S ° o o h l p r I d l e m o r e h , ma 0r o c o t.v le

n . ep.

L e u c o o h lo ri d io m o r p h a m a c r o o o ty le n« s p .
Refer to P la te I I .

T o t a l l e n g t h 1.3 7 mm.

G reatest

w i d t h i n a p l a n e j u s t p o s t e r i o r to th e c e n t e r o f th e a c e ­
tabulu m , 0 . 6 2 mm.

The body n arro w s r a p i d l y in f r o n t o f the

a c e t a b u l u m , b u t i s somewhat rounded a t the a n t e r i o r end.
The p o s t e r i o r end i s v e r y b r o a d l y p o i n t e d .

The s k i n i s

e n t i r e l y sm ooth, w i t h t h e e x c e p t i o n o f the a c e ta b u lu m , the
s u r f a c e o f which a p p e a r s to be v e r y much roughened by s c a l e lik e stru c tu re s.

The a c e ta b u lu m measures .512 by .576 mm.

and i s e x t r e m e l y pow erfu l*

The o r a l s u c k e r i s p la c e d s l i g h t ­

l y s u b t e r m i n a l , i s w e ll developed and m easures .208 by .24 mm.
The p h ary n x m e a su r e s .067 mm i n d i a m e t e r .

I f an esophagus

i s p r e s e n t i t i s e x t r e m e l y s h o r t , none b e i n g d i s c e r n a b l e in
t h e ty p e s p e c im e n .

The i n t e s t i n a l c r u r a a r e l a r g e , r a t h e r

t h i n w a l l e d , and i n t h e t y p e , v e r y i r r e g u l a r .

They have no

a n t e r i o r l o o p a s i s o f t e n found i n th e genus L euooc hlorid um .
I t was i m p o s s i b l e to t r a c e th e c r u r a p o s t e r i o r to th e a n t e r ­
i o r b o r d e r o f t h e ac e ta b u lu m due to the o p a c i t y o f t h a t
s t r u c t u r e , a s w e l l as t h e d e n s i t y o f t h e eggs and v i t e l l a r i a .
A l l t h e s e x o r g a n s a r e l o c a t e d p o s t e r i o r to th e a c e ­
tabulum i n t h e p o s t e r i o r body f o u r t h .

The p o s t e r i o r t e s t i s

(60)

i s l o c a t e d s l i g h t l y to th e r i g h t o f t h e median l i n e , and
a l m o s t i n t h e extrem e t i p o f t h e body.
by *188 mm.

I t measures .181

The a n t e r i o r t e s t i s i s l o c a t e d to the l e f t

o f t h e m edian l i n e , t o u c h i n g th e p o s t e r i o - l a t e r & l b o r d e r
o f the acetabulum .

I t m e a su r e s .102 by .128 mm.

The

w e l l d e v e lo p e d c i r r u s p o u e h l i e s m edian, a s h o r t d i s t a n c e
from th e p o s t e r i o r end o f th e body.
by .064 mm i n d i a m e t e r .

I t i s s p h e r i c a l , .064

The c i r r u s e x te n d s r o u g h l y a n t e r ­

i o r and p o s t e r i o r t h r o u g h t h e s a c , t u r n i n g toward th e r i g h t
n e a r t h e p o s t e r i o r b o r d e r o f t h e s a c , and opening by means
o f a pore on t h e d o r s a l s u r f a c e j u s t a n t e r i o r to th e p o s t ­
e r i o r t i p o f the body.

The c i r r u s i s s p i n e d .

The ovary

i s to t h e l e f t o f t h e m i d - l i n e and between th e two t e s t e s .
I t m e asu res .134 by .057 mm.

J u s t to th e r i g h t o f th e

o v a r y , and i n a l i n e between th e two t e s t e s , l i e s th e
fecundarium .

The v i t e l l i n e d u c t meet between the f e c -

undarium and t h e o v a r y , forming a s m a ll v i t e l l i n e r e s ­
ervoir*

The v i t e l l a r i a a r e l o c a t e d on e i t h e r s i d e o f

t h e a c e t a b u l u m , f o l l o w i n g in g e n e r a l th e curve o f t h a t
o r g a n , and e x ten d p o s t e r i o r a b o u t one t h i r d th e l e n g t h
of th a t organ.
fo llic les.

They a r e composed o f i r r e g u l a r l y grouped

The u t e r u s h a s an a s c e n d i n g and a d e s c e n d i n g

(61)

lo o p on e i t h e r a i d e o f th e a c e t a b u l u m , e x t e n d in g o n ly a b o u t
a s f a r a n t e r i o r a s th e p o s t e r i o r l i m i t s o f the v i t e l l a r i a .
The u t e r u s o f th e ty p e specim en i s f i l l e d w ith r a t h e r
l a r g e , y e l l o w i s h eggs which measure .032 by .013 mm.
H ost;

Anas r u b r l p e s ( b u r s a )

L ocality:

W i n te r g r e e n Lake, A u g u s ta , Michigan

Type sp e c im e n :

U. S. N a t ’l . Mus. No. 9121

That t h i s d e s c r i p t i o n i s n e c e s s a r i l y based on a
s i n g l e whole mount specimen i s u n f o r t u n a t e .

However,

th e c h a r a c t e r s d e s c r i b e d a r e v e r y c l e a r , and t h i s f a c t
coupled w i t h t h e f a c t t h a t th e a d d i t i o n a l specimen which
was l o s t , showed th e same e x t e r n a l c h a r a c t e r s above d e s ­
c r i b e d , have made i t seem d e s i r a b l e to name t h i s worm a s
a new s p e c i e s and g e n u s .

Part II

Key to th e g e n e r a and d i a g n o s i s o f t h e f a m i l i e s
and g e n e r a and l i s t s o f the s p e c i e s o f tr e m a to d e s
r e p o r t e d from North American Ducks.

(62)

Key to t h e Trem atodes r e p o r t e d
from N o r t h American Ducks

1.

Sexes s e p a r a t e , b lo o d f l u k e s ..............................................................
Hermaphrod i t i c f l u k e s ............................................ , .............................. 6

2.

Females s l e n d e r , m a les l a r g e r t h a n fe m a le s and somewhat
f l a t t e n e d to form a g y n a e e o p h o r i c c a n a l in which the
fem a le i s c o n t a i n e d , G ynaeeophoric c a n a l e x t e n d i n g
a n t e r i o r to a c e t a b u l u m ............................................ M i c r o b i l h a r z i a
Females s i m i l a r
to m a les i n s h a p e ; m ales w i t h o u t a
w e l l d e v e lo p e d g y n a e e o p h o r ic c a n a l .............................................. 3

3.

Body c y l i n d r i c a l o r n e a r l y s o ...................................................... . . 4
Body f l a t t e n e d ........................................................................................... 5

4.

Female unknown; p o s t e r i o r end o f body t h r e a d - l i k e ,
m id d le p o r t i o n w id e r th a n e i t h e r end; no g y n a e e o p h o ric
c a n a l ; s u c k e r s p r e s e n t ..........................................T r i c h o b i l h a r z i a
Male and fem a le l o n g and s l e n d e r ; g y n a e e o p h o r ic c a n a l
re d u c e d to a s h o r t groove i n a n t e r i o r p a r t o f body; one
s u c k e r p r e s e n t , or e n t i r e l y a b s e n t
G igantobilharzia

5.

S u ck ers p r e s e n t ; common caecum w i t h o u t l a t e r a l den­
d r i t i c b r a n c h e s .................................................................B i l h a r z i e l l a
S u c k e rs a b s e n t ; common caecum w i t h s h o r t l a t e r a l den­
d r i t i c b r a n c h e s ...................................................... D endri t o b i l h a r z i a

6.

Body d i v i d e d i n t o a f o r e body and a hind b o d y , the
o r g a n s o f a t t a c h m e n t on t h e f o r e body and t h e se x
o r g a n s i n t h e h in d b o d y ....................................................................... 7
Body n o t so d i v i d e d ............................................ ....................................9

7.

V i t e l l a r i a e x t e n d i n g i n t o a n t e r i o r body, two appe nd­
a g e s on f o r e b o d y . . . . . ...................................................P a r a s t r i g e a
V i t e l l a r i a c o n f i n e d to p o s t e r i o r b o d y ....................................... 8

8.

B u r s a c o p u l a t o r i x w i t h a m u s c u la r s u c k e r - l i k e r i n g .............
...................................C o ty lu ru 3
B u r s a c o p u l a t o r i x a s im p le s a c ...................................... Apatemon

9.

S t r u c t u r e s t r i g e i d - l i k e , b u t body n o t d i v i d e d ; a c c e s s ­
o r y o r g a n s o f a t t a c h m e n t p r e s e n t ..........................C y a t h o c o t y l e
Organs o f a t t a c h m e n t c o n s i s t i n g o f one or two s u c k e r s . . .
10

(63)

10*

With o n l y one s u c k e r p r e s e n t , and t h a t a t t h e a n t e r i o r
end ( i n ty p h l o c o e l u m t h e r e i s no s u c k e r p r e s e n t ) ............... 11
With two s u c k e r s p r e s e n t ...................................................................... 17

11.

With a w e l l d e v e lo p e d o r a l s u c k e r , i n t e s t i n e s end
b l i n d l y ..............................................
12
O r a l s u c k e r o n l y s l i g h t l y deve lop ed or a b s e n t , i n t e s ­
t i n e s c o n t i n u o u s , making a com plete c i r c l e a t the
p o s t e r i o r e n d ............................................................................................... 15

12.

A p harynx p r e s e n t , p a r a s i t i c i n th e k i d n e y .............. E u o o ty le
Ph ary n x a b s e n t , p a r a s i t e s o f th e i n t e s t i n e and c a e c a ..........
................................................. 13

13.

On t h e v e n t r a l s u r f a c e one o r more rows o f g l a n d s
p r e s e n t .......................................................
14
No g l a n d s p r e s e n t on v e n t r a l s u r f a c e .
Paramonostomum

1 4.

Glands d i s t i n c t and p r o t r u s i b l e ; in 3 -5 r o w s . . N o t o c o t y l u s
G lands n o t d i s t i n c t ; n o t p r o t r u s i b l e ; th e c e n t r a l row
fo rm in g a r i d g e ..................................................................... C a t a t r o p h i s

15.

I n t e s t i n e s u n b r a n c h e d , body t a p e r i n g toward th e a n t e r ­
i o r e n d , a weak s u c k e r p r e s e n t ........................................................ 16
I n t e s t i n e s b e a r i n g d i v e r t i c u l a on the i n n e r s i d e , body
o v a l i n s h a p e , e q u a l l y rounded a t b o t h e n d s . . Typhlocoelum

1 6.

The s e x g l a n d s
the i n t e s t i n e ,
ly in g a t about
Sex g l a n d s l i e
o v a r y b etw e en

form ing a t r i a n g l e i n p o s t e r i o r lo o p o f
th e o v a r y and a n t e r i o r t e s t i s u s u a l l y
t h e same h e i g h t . . ...............
Cyclocoelum
i n more or l e s s o f a s t r a i g h t l i n e , the
th e two t e s t e s ................................. Opthalmophagus

17.

With th e a c e ta b u l u m a t t h e p o s t e r i o r end o f th e body.
Acetabulum b e a r i n g two l i p s .............................................Z y g o co ty le
With a c e t a b u l u m on th e v e n t r a l s u r f a c e o f th e b o d y
18

18.

With a c o l l a r around o r a l s u c k e r in which a r e one or
two rows o f l a r g e s p i n e s ......................................................................19
W ithout c o l l a r or s p i n e s around o r a l s u c k e r ............................ 23

19.

Head s p i n e s i n a double row ................................................................ 20
Head s p i n e s i n a s i n g l e r o w ................................................................ 22

80.

V i t e l l a r i a b a r e l y r e a c h i n g a c e t a b u l u m ........................................ 21
V i t e l l a r i a r e a c h i n g to or a n t e r i o r to a c e t a b u l u m ...................
.............................Hypodereum

(64)

21* Medium to l a r g e f o r m s , u t e r u s l o n g ........................ Echlnostoma
Small E c h i n o s t o m e s , u t e r u s s h o r t , eggs r e l a t i v e l y l a r g e
................................................. Eohinoparyphium
22

23

.
*

C i r r u s pouch s h o r t , 22-26 head s p i n e s , P a r s p r o s t a t i c a
a b s e n t, v i t e l l a r i a not reaching ovary
S te p h a n o p r o r a
29-31 head s p i n e s , v i t e l l a r i a r e a c h i n g a n t e r i o r to
a c e t a b u l u m ................................................................
H im a s th la
G e n i t a l p o r e a t p o s t e r i o r end o f body . .L e u c o e h lo r id io m o rp h a
G e n ita l pore n o t a t the p o s t e r i o r end
........... . . ... ...34

24.

A w e l l d e f i n e d c i r r u s pouch p r e s e n t ...........................
33
C i r r u s pouch a b s e n t ............................................................................... 25

25.

Numerous t e s t e s p r e s e n t i n m a tu re s p e c i m e n s . . . Orchipedum
Only two t e s t e s p r e s e n t ...................................................................... 26

26.

V e n tr a l su ck er com pletely modified in to a g e n i t a l sucker
o f gonytol
.......................
Tocotrema
V e n t r a l s u c k e r and g e n i t a l p o r e s e p a r a t e ............................... 27

27.

A g e n i t a l a t r i u m p r e s e n t i n which a d i s t i n c t p a p i l l a e
i s p r e s e n t ..................................................
28
No p a p i l l a e p r e s e n t ..................................
30

28.

P a p i l l a e n a r r o w .......................................................
. L evinseniella
P a p i l l a e more or l e s s cone s h a p e d ......................
.29

29.

Vagina o p e n i n g n e a r b a s e o f th e p a p i l l a e
S p e lo tre m a
Vagina o p e n i n g n e a r t h e mouth o f t h e g e n i t a l a t r i u m ..........
...........................S p e l o p h a l l u 3

30.

Small f o r m s , r e c e p t a c u l u m s e m i n i s a b s e n t , v i t e l l a r i a l i e
d o r s a l i n r e g i o n o f a c e t a b u l u m ..............................Gymnophallus
L a r g e r , u s u a l l y more e l o n g a t e form§, v i t e l l a r i a l a t e r a l
to i n t e s t i n a l c r u r a , a s e m in a l r e c e p ta b u l u m p r e s e n t . . . 31

3 1.

V i t e l l a r i a d i v i d e d on e i t h e r s i d e i n t o an a n t o v a r a l and
p o s t o v a r a l p o r t i o n , the p o s t o v a r a l p o r t i o n e x t e n d i n g to
n e a r p o s t e r i o r end o f th e b o d y ................................... Amphimerus
V i t e l l a r i a n o t d i v i d e d , c o n t i n u o u s i n l a t e r a l f i e l d s . . 32

32.

V i t e l l a r i a e x t e n d i n g a n t e r i o r to a c e t a b u l u m . . . . M e t o r c h i s
V i t e l l a r i a n o t r e a c h i n g a c e t a b u l u m ......................O p i s t h o r c h l a

(65)

33.

G e n i t a l o p e n in g v e r y n e a r , and u s u a l l y l a t e r a l to a c e t a b ­
u lum , o p e n in g i n t o a d e p r e s s i o n .................................... .M a ritre m a
G e n i t a l o p e n in g a n t e r i o r to a c e t a b u l u m ......................................34

34.

U t e r u s l y i n g f o r t h e most p a r t a n t e r i o r to o v a r y ............... 35
U t e r u s w i t h w e l l d e f i n e d c o i l s p o s t e r i o r to o v a r y which
u s u a l l y r e a c h to n e a r p o s t e r i o r end o f b o d y .......................... 39

35.

V i t e l l a r i a n o t r e a c h i n g p o s t e r i o r any f u r t h e r th a n the
a n t e r i o r t e s t i s . V agina and c i r r u s v e r y l o n g , e x t e n d ­
................... P h i l o p h t h a l m u s
i n g p o s t e r i o r to a c e t a b u l u m
V i t e l l a r i a c o n s i d e r a b l y o v e r l a p p i n g i n t e s t i n e , and ex­
t e n d i n g to th e p o s t e r i o r body e x t r e m i t y . C i r r u s and
v a g i n a n o t e x t e n d i n g p o s t e r i o r to a c e ta b u lu m n o t i c e a b l y . .

3 6.

Body e l o n g a t e , n a r r o w ..............................................................................37
Body n o t e l o n g a t e , r o u n d e d ................................................................. 38

37.

T e s t e s o v a l , e n t i r e , p o s t e r i o r end r o u n d e d . . . . Psilostomum
T e s t e s e l o n g a t e , s l i g h t l y n o t c h e d , p o s t e r i o r end more
o r l e s s p o i n t e d ....................................................................P silo c h a sm u s

38.

A cetabulum i n second body f o u r t h .................................. P s i l o t r e m a
A cetabulum n e a r c e n t e r o f b o d y .............................S p a e r i d i o t r e m a

39.

T e s t e s one i n f r o n t o f th e o t h e r or s l i g h t l y o b l i q u e
g e n i t a l p o r e i n f r o n t o f a c e t a b u l u m ........................ P l a g i o r c h i s
T e s t e s a t th e same l e v e l , g e n i t a l o p e n in g s n e a r a n t e r ­
i o r t i p o f b o d y ...........................................................................................40

40.

Male and fem ale o p e n in g s v e r y c l o s e
t o g e t h e r ..
.............................................P ro s thogonimus
Male and fem ale o p e n in g s r a t h e r w id e l y s e p a r a t e d ................ ..
....................................... Schistogonim us

f 66)

D i a g n o s i s o f F a m i l i e s and G enera, and l i s t s
o f s p e c i e s o c c u r r i n g i n Worth American Ducks

S u b c la s s Digenea

T re m ato d a.

I n t e r n a l l y p a r a s i t i c f l u k e s , w i t h one

o r more s u c k e r s which a c t a s h o l d - f a s t o r g a n s .

The l i f e

c y c l e i s i n d i r e c t , and alw a ys i n c l u d e s a m o l l u s c a n h o s t
a t one s t a g e .

U su ally u t i l i z i n g a v e r t e b r a t e as h o st in

the a d u l t or sexual s t a g e .

Of the s i x t y odd f a m i l i e s the

f o l l o w i n g have b ee n r e c o r d e d from Worth American d ucks:
Param phistom idae, A canthostom idae, P l a g i o r o h i d a e ,
O p is th o rc h iid a e , E etero p h yid ae, M ic ro p h a llid ae, O rchipedidae,
P h ilo p h f c h a l m id a e , C y o l o c o e l i d a e , E u c o t y l i d a e , W o t o c o t y l i d a e ,
E chinostom idae, P silo sto m id a e , O yathooolyoidae, S t r i g e i d a e ,
H arm o sto m id a e , and S c h i s t o s o m i d a e •

F am ily PARAMPHISTOMIDAE F i s c h o e d e r , 1901
Medium s i z e d f l u k e s , w ith c o n i c a l or c y l i n d r i c a l
shaped b o d i e s , and w i t h t h e s k i n unarmed.

P o s te r io r sucker

l o c a t e d a t , or s l i g h t l y a n t e r i o r and v e n t r a l t o , t h e p o s ­
t e r i o r end o f th e body.

O ral s u c k e r w e ll d e v e l o p e d , o f t e n

w i t h two d i v e r t i c u l a o r a r i n g o f d i v e r t i c u l a from i t .

(67)

Pharynx a b s e n t , i n t e s t i n a l c r u r a s i m p l e .

Excretory bladder

b a g - f o r m , t h e e x c r e t o r y p o r e b e i n g l o c a t e d d o r s a l l y and
s l i g h t l y a n t e r i o r to t h e p o s t e r i o r end.
o p e d , e n t i r e , l o b e d , or b r a n c h e d .

Testes w ell d e v e l­

C i r r u s p r e s e n t or a b s e n t .

Ovary s m a l l e r t h a n t e s t e s , l y i n g p o s t e r i o r to them.
U t e r u s d o r s a l , few c o i l s .

V i t e l l a r i a u s u a l l y vvell d e v e l o p e d .

G e n i t a l o p e n i n g d o r s a l , m e d ia n , i n t h e a n t e r i o r body t h i r d .
Eggs n um e rous, l a r g e , w i t h or w i t h o u t f i l a m e n t s .

P arasites

o f f i s h , a m p h i b i a , r e p t i l e s , b i r d s , and mammals.

Genus Z y g o o o ty le S t u n k a r d , 1916
Param phistom idae.
shaped f l u k e s .
d iv erticu la.

Medium s i z e d , more or l e s s c o n i c a l

O ral s u c k e r w e ll d e v e l o p e d , w ith a p a i r o f
O ra l s u c k e r p la c e d v e n t r a l and s u b - t e r m i n a l .

A cetabulum p o s t e r i o r , t e r m i n a l , and w i t h tv/o p r o j e c t i n g
lip s.

D iv id e d i n t o an a n t e r i o r and p o s t e r i o r p o r t i o n by a

partitio n .

T estes w ell developed, s l i g h t l y lobed.

s m a l l e r th a n t e s t e s , p o s t e r i o r to them.

F olds o f u t e r u s

few, c o n f i n e d b e tw e e n t h e i n t e s t i n a l c a e c a .
7/ e l l

developed.

Ovary

V itellaria

C i r r u s a b s e n t , g e n i t a l pore l o c a t e d n e a r

th e b i f u r c a t i o n o f th e i n t e s t i n e s .
Type and o n ly s p e c i e s :
1836) .

Z ygoootyle lunatum ( D i e s i n g ,

(68)

Z y g o o o ty le lu n a tu m ( B i e s i n g , 1836)
H ost:

Anas m e l a n o t u s , A. i p e o u t i r i . A. m o s o h a t a ,

A. jo. p l a t y r h y n o h o s . A. jo. d o m e s t l o u s . A. r u b r i p e s .
Mar11a a m e r i c a n a , N e t t l o n c a r o l i n e n s e . B r a n t a c a n a d e n s i s .
A nser a n s e r d o m e s t l o u s . O a l l i n a g o d e l i c a t a , Eimantopus
w i l s o n i i , Cervus d i o h a t o m u a . Bos t a u r u s .
L o cality :

N o r t h , C e n t r a l , and South A m erica.

R eference:

102, 129.

Jfemily ACANTHOSTOMIBAE P oche, 1925
Skin s p i n e d .

I n t e s t i n a l cru ra sim ple.

b l a d d e r "Y" s h a p e d .
th e a c e t a b u l u m .
developed.

Excretory

G e n i t a l o p eh in g m e d ian , l y i n g b e f o r e

C i r r u s pouch a b s e n t , c i r r u s s l i g h t l y

Sex g l a n d s l y i n g b e h in d the a c e t a b u l u m , the

ovary in f r o n t o f the t e s t e s .

V i t e l l a r i a w e ll d e v e l o p e d .

Only one genus r e p o r t e d from d u c k s .

Genus Gymnophallus Odhner, 1900
A canthostom idae.
skin.

S m all, t h i c k , ov al worms w i t h s p in e d

O r a l s u c k e r sub t e r m i n a l , a c e ta b u lu m a b o u t th e m id d le

of body.

Pharynx s m a l l , esophagus medium l e n g t h .

o r u r a s h o r t , n o t r e a c h i n g beyond a c e ta b u l u m .

In testin al

The t e s t e s l i e

l a t e r a l l y beh in d the a c e t a b u l u m , t h e o vary j u s t a n t e r i o r to
them.

A receptaculum sem inis i s a b s e n t, L a u r e r 's canal

(69)

p resen t.

The v i t e l l a r i a a r e d o r s a l , n e a r the a c e ta b u lu m and

a r e composed o f o n ly a s m a l l number o f f o l l i c l e s on e i t h e r
sid e.

The u t e r u s i s l o c a t e d i n the p o s t e r i o r h a l f o f th e body.

Gymnophallus b u r s i c o l a Odhner, 1900
H ost:

S o m a te ri a m o l l i s s i m a

L o cality :
R eference:

Europe
95, 96

Gynmophallus c h o l e d o c u s Odhner, 1900
H o s t:

S o m a te r i a m o l l i s s i m a . Todorna to d o r n a ( G a l l - b l a d d e r )

L o cality :
Reference:

Europe
65, 95, 97

Gymnophallus d i p s l l i s N i c o l l , 1907
H o s t:

Oidemia f u s o a . 0 . n i g r a (Bursa F a b r i c i i )

L ocality:
R eference:

Europe
M c I n to s h , 1927; N i c o l l , 1906, 1907, 1909

Gymnophallus s o m a t e r i a e ( L e v i n s e n , 1881)
H o s t:

Somateria m o llissim a

L o cality :
Reference:

Europe
95, 96

Fam ily PLAGIOROHIDAE L u h e, 1899
Synonyms:

L e p o d e rm a tid a e L o s s , 1901; P l a g i o r c h i i d a e

Ward, 1917; P r o s t h o g o n i m i d a e N i c o l l , 1924; R e n i f e r i d a e

(70)

B a e r , 1924.

Body e l o n g a t e to o v a l , m o d e r a t e l y f l a t t e n e d to c y l i n d ­
rica l.

S kin s p i n e d , a s a r u l e , o v er e n t i r e s u r f a c e .

Suckers

w e l l d e v e l o p e d , a c e ta b u l u m b e f o r e t h e c e n t e r o f th e body.
P re p h a r y n x s h o r t , p harynx p r e s e n t , esophagus s h o r t to medium.
I n t e s t i n a l c r u r a v a r i a b l e , b u t u s u a l l y r e a c h to p o s t e r i o r
p a r t o f body.

E x c r e t o r y b l a d d e r "Y" s h a p e d , th e b r a n c h e s

r e a c h i n g o f t e n beyond th e m id d le o f th e body.

The t e s t e s a r e

e i t h e r e n t i r e or lo b e d and may l i e a t the same l e v e l , o b l i q u e ,
or one i n f r o n t o f th e o t h e r .

The c i r r u s pouch i s s t r o n g l y

d e v e l o p e d , and u s u a l l y c o n t a i n s a s e m in a l r e s e r v o i r , a p a r s
p r o s t a t i c a and a p r o t r u s i b l e c i r r u s .

The o v ary l i e s b e f o r e

t h e t e s t e s , u s u a l l y b e h in d t h e a c e t a b u l u m .
s e m i n i s may be p r e s e n t o r a b s e n t .
uterus l i e

A r e c e p ta c u lu m

The l o o s e f o l d s o f the

f o r t h e most p a r t between th e t e s t e s and a c e t a b ­

ulum, or l o o s e f o l d s may ex tend to the p o s t e r i o r end o f the
body.

The v i t e l l a r i a a r e l a t e r a l .

The g e n i t a l pore l i e s

median to l a t e r a l , and u s u a l l y a n t e r i o r to t h e a c e t a b u l u m .
The numerous eggs a r e y e l l o w .

Of th e f o r t y odd g en e ra

r e c o g n i z e d i n t h i s f a m i l y o n l y t h r e e have been r e c o r d e d
from d u c k s .

(

71)

Genua P l a g i o r o h l s Luhe, 1899
P lagiorchidae.

Body l o n g o v a l to e l o n g a t e , f r e q u e n t l y

more o r l e a s p o i n t e d a t th e e n d s .

Skin a p i n e d .

n e a r l y e q u a l i n s i z e , esophagus s h o r t .

S u ck ers

T e s t e s e n t i r e , one

behind th e o th e r or s l i g h t l y o b liq u e in p o s t e r i o r h a l f of
the body.

Ovary r o u n d , r e c e p t a c u l u m s e m i n i s a b s e n t .

The

u t e r u s may or may n o t r e a c h p o s t e r i o r to the t e s t e s .

The

v i t e l l a r i a a r e composed o f s m a l l , numerous f o l l i c l e s i n
t h e l a t e r a l p o r t i o n o f th e b o d y , r e a c h i n g from th e pharynx
to t h e p o s t e r i o r end o f the body.

The g e n i t a l pore l i e s a

s h o r t d i s t a n c e a n t e r i o r to th e a c e t a b u l u m .
Type s p e c i e s :

P lag io rch is v e s p e rtilio n is

( M u e l l e r , 1784)

P l a g l o r c h i s p o t a n l n i S k r j a b i n , 1928
H o st:

Anas p l a t y r h y n o h o s ( I n t e s t i n e )

L ocality:
Reference:

S ib eria
117
Genus P ro sth o g o n im u s Luhe, 1899

Synonym:

Prymnopryon L o o ss, 1899

Small to medium s i z e d f l u k e s , body f l a t t e n e d , a n t e r i o r
end somewhat p o i n t e d , p o s t e r i o r u s u a l l y b r o a d l y ro u n d e d ,
g r e a t e s t w id th p o s t e r i o r to m id d le o f body.
P re p h a ry n x very s h o r t,

esophagus medium.

t h e same l e v e l , b e h i n d a c e t a b u l u m .

Skin s p i n o u s .

T e3te s l y i n g a t

C i r r u s pouch c o n t a i n s

(78)

a l a r g e s e m in a l v e s i c l e .

Ovary u s u a l l y s l i g h t l y l a t e r a l

t o median l i n e , a n t e r i o r to t e s t e s , and p o s t e r i o r or m o s t l y
p o s t e r i o r to a c e t a b u l u m .

Beceptaculum s e m i n i s p r e s e n t .

U t e r u s e x t e n d s to th e l a t e r a l p o r t i o n s o f th e body, and
r e a c h e s to th e p o s t e r i o r end.

G e n i t a l p o re l a t e r a l to

t h e o r a l s u c k e r , th e male and female o p e n in g s c l o s e t o g e t h e r .
L a u r e r ’s c a n a l p r e s e n t .

V itellaria la te ra l.

Eggs numerous

and s m a l l .
Type s p e c i e s :

P ro sth o g o n im u s o v a t u s ( R u d o lp h i, 1803)

P ro sth o g o n im u s a n a t i n u s Markow, 1902
H ost:

Anas p l a t y r h y n c h o s d o m e s tic u s (Bursa o f F a b r e c i u s )

L o cality:
Reference:

Russia
6 5 , 69

P ro sth o g o n im u s c u n e a t u s ( R u d o l p h i , 1809)
H o st:

A rdea c i n e r e a , C a s a r c a c a s a r e a , Oolymbus

n i g r i c a n s . Oorvua c o r n l x , £ . c o r o n e . Cygnus o y g n u s , ffullca
a t r a , G a r r u l u s g l a n d a r i u s , Grus c i n e r e a , N e t t i o n o r e c a ,
Nyroca c l a n g u s . P a s s e r d o m e s t i c u s . G a llu s d o m e s t i c u s . 1
L o c a l i t y : E u ro p e , A s i a , North America
R eference:

1 5 , 18 , 69

1. T h is s p e c i e s has a l s o been ta k en from the red-w inged b l a c k
b i r d ( A g e l a i u s p h o e n i c e u s p h o e n i c e u s ) and th e bronzed g r a c k l e
( Q u i s c a l u s q u i s c u l a a e n e u s ) a t th e W. K, Kellogg Bird S a n c t u a r y .

(73)

P ro sth o g o n im u s h o r i u o h i l M o r i s h i t a and Tsuchim ochi, 1925
B o a t:

Ana a jc>, ( domes t i c u s ) and Anaer a* ( domes t i o u a )

L ocality:

Formosa

Reference:

69

P ro sth o g o n im u s m a o r o r c h i a Maey, 1934
Host:

Anas jd. p l a t y r h y n o h o s . A. jd. d o m e s t i c u s , A.

r u b r i p e s . G a llu a d o m e s t i c u s (Egg, O v i d u c t , Bursa)
L ocality:

N orth America ( G r e a t Lakes Region)

Reference:

6 8 , 69, 50, 51,

P ro sthogonim us o v a t u s ( R u d o l p h i , 1803)
H ost:

S p a t u l a c l y p e a t a . Anaa f e r i n a . Anas g l a c i a l i a ,

Nyrooa m a r i l a , Anas m u s i o a , Nyroca h y e m a i l s . and 35 o t h e r
species of b ird s .
L ocality:
R eference:

E u ro p e , A s i a , South America
15 , 18, 64, 61, 69

P ro sth o g o n im u s r u d o l p h i S k r j a b i n , 1919
H o s t:

Anas p l a t y r h y n o h o s d o m e s t i c u s , A. jo. p l a t y r h y n o h o s ,

A. r u b r i p e s , Nyroca c o l l a r l s , Nyroca m a r i l a , Nyroca a f f i n i s ,
G lau cio n etta clangula am ericana.
L ocality:
R eference:

E u ro p e , N orth America
69

P r o s thogonimus s k r jab i n i Zakharov/, 1920
H ost:

Anas p l a t y r h y n o h o s dom e stic u s

(74)

L ocality:

Russia

R eference:

6 , 69

P ro s th o g o n im u s p e l l u o i d u s (v L ins tow, 1873)
H o s t:

Anas p l a t y r h y n c h o s d o m e s t i c u s , Numenius a r q u a t u s

and G a ll u s d o m e s t i c u s .
L ocality:

Europe

R e f e r e n o e : 69
Genus S c h isto g o n im u s Luhe, 1909
The d i a g n o s i s f o r t h i s genus i s p r a c t i c a l l y the same
a s f o r P ro sthogo nim us . w i t h th e e x c e p t i o n t h a t the g e n i t a l
o p e n i n g s o f th e male and female system a r e r a t h e r w id e l y
s e p a r a t e d , and do n o t open c l o s e t o g e t h e r as in the l a t t e r
genus.

The u t e r u 3 i s c o n f i n e d w i t h i n the i n t e s t i n a l c r u r a .

Type and o n ly s p e c i e s :

S. r a r u s (B rau n , 1901)

S ch isto g o n im u s r a r u s (B raun, 1901)'
H o st:

Anas p l a t y r h y n o h o s , S p a t u l a c l y p e a t a , F u l i c a a t r a

L ocality:
R eference:

Europe
1 5 , 1 8 , 65

F am ily OPISTHOPCHIIDAE Luhe, 1901
Body u s u a l l y somewhat f l a t t e n e d , n o t v e r y m u s c u l a r ,
e l o n g a t e , th e a n t e r i o r end somewhat p o i n t e d .
or s p i n e d .

Skin smooth

The s u c k e r s u s u a l l y c l o s e t o g e t h e r , and n o t

s tr o n g ly developed.

Pharynx and esophagus p r e s e n t .

In tes­

(

75)

t i n e s u s u a l l y r e a c h i n g to n e a r the p o s t e r i o r end.

Excretory

b l a d d e r ”Yn s h a p e d , t h e stem v e r y l o n g , and u s u a l l y s ig m o id .
E x c r e t o r y p o r e a t th e p o s t e r i o r e x t r e m i t y or s l i g h t l y su b ­
t e r m i n a l and v e n t r a l .

T e s t e s n e a r p o s t e r i o r end o f the

body, u s u a l l y one beh in d th e o t h e r , b u t sometimes s l i g h t l y
o b l i q u e , e n t i r e or l o b e d .

C i r r u s pouch a b s e n t .

r e c e p t a c l e w e l l d e v e l o p e d , n e a r th e o v a r y .
and i n f r o n t o f th e t e s t e s .

Seminal

Ovary s m a l l e r

U te r u s f i l l i n g most o f the

body s p a c e betw e en th e o v ary and a c e ta b u l u m .
w e l l d e v e l o p e d , l a t e r a l to the c r u r a .

V itellaria

G e n i t a l pore i n

f r o n t o f t h e a c e t a b u l u m , u s u a l l y v e r y c l o s e to t h a t o r g a n .
L a u re r's canal p re se n t.

E g g s , s m a l l , numerous, y e l lo w in

color •
In r e p t i l e s , f i s h , b i r d s , and mammals.

In the l i v e r ,

g a l l - b l a d d e r , or p a n c r e a s .
Genus O p i s t h o r c h i s B l a n c h a r d , 1895
Body d i s t i n c t l y , o f t e n v e r y g r e a t l y , e l o n g a t e d , a n t e r i o r
end a t t e n u a t e d ; p o s t e r i o r end b r o a d e r .
w ithout s p in e s .

Skin g e n e r a l l y smooth,

E x c r e t o r y v e s i c l e u s u a l l y "Y" shaped w ith

lo n g sigm oid stem w inding between th e t e s t e s or p a s s i n g
d o r s a l to th e b r a n c h e s o f th e l a t t e r .
absent.

C o p u l a to r y organs

T e s t e s i n the p o s t e r i o r p o r t i o n o f the b o d y , and

p la c e d e i t h e r o b l i q u e l y or d i r e c t l y behind one a n o t h e r ; more

(76)

or l e a s lo b e d or d e n d r i t i c ,
canal p re s e n t.

^ v a r y s im p le or l o b e d .

Receptaculum se m in is p r o m in e n t .

L & u r e r 's

U terine

o o i l s extend from the o v a r y to th e v e n t r a l sticker and do
n o t o v e r l a p t h e i n t e s t i n a l ca ec a to any marked e x t e n t .
V i t e l l a r i a m o d e r a t e l y d e v e l o p e d , l a t e r a l o f the i n t e s t i n a l
c a e c a and n o t e x t e n d i n g a n t e r i o r o f th e v e n t r a l s u c k e r ,
e n d i n g p o s t e r i o r l y a t the l e v e l o f the o v a r y .

V itellaria

form one r e g i o n .
Type s p e c i e s :

0 . f e l l n e u s ( R i v o l t a , 1884)

O p i s t h o r c h i s geminus (L o o s s , 1896)
H o s t:

Anas p l a t y r h y n o h o s , C iro u s a e r u g i n o s u s ( L i v e r )

L o cality :

E g y p t, A sia

R eference:

3 , 60, 61, 62, 84

O p i s t h o r c h i s a i m u la n s (L o o s s, 1896)
H ost:

Anas p l a t y r h y n o h o s , P. p e n e l o p e . P e r n i s a p i v o r u s ,

Circus a e ru g in o su s♦
L ocality:

E g y p t, Europe

Reference:

3 , 60, 61, 62, 84

Genus Amphimerus B a r k e r , 1911
Synonym:

O p i s t h o r c h i s B l a n c h a r d , 1895

Body f l a t t e n e d , e l o n g a t e d , a n t e r i o r end t a p e r i n g .

Skin

f r e q u e n t l y covered w h o lly or in p a r t by s m a l l r e t r o s e s p i n e le ts.

E x c r e t o r y system "Y" shaped w ith sigmoid stem w inding

(77)

b etw e en t e s t e s .

Copulatory organs a b s e n t.

Testes in p o s te r ­

i o r p o r t i o n o f b o d y , s i m p le o r l o b a t e ; the one o o l i q u e l y
p o s t e r i o r to th e o t h e r , o r i n some c a s e s one behind the
other.

Ovary a n t e r i o r to t e s t e s , s im p le or l o b a t e .

L a u r e r ’s

c a n a l p r e s e n t ; r e c e p t a c u l u m s e m in is w e l l d e v e lo p e d ; u t e r i n e
co ils an terio r

to o v a r y and may extend l a t e r a l l y over the

i n t e s t i n a l c a e c a ; v i t e l l a r i a w e l l d e v e l o p e d , l a t e r a l to the
i n t e s t i n a l oaeca and d i v i d e d i n t o two d i s t i n c t r e g i o n s by
a b r e a k o p p o s i t e t h e o v a r y ; n o t e x t e n d i n g a n t e r i o r l y beyond
the a c e ta b u lu m b u t f r e q u e n t l y e x t e n d i n g p o s t e r i o r l y to or
beyond th e p o s t e r i o r t e s t e s .
Type s p e c i e s :

Amphimerus o v a l i s B a r k e r , 1911

Amphimerus e l o n g a t u s n . s p .
H o s t:

G l a c u i o n e t t a c l a n g u l a a m e r i c a n a . Anas jo. p l a t y -

r h y n c h o s , A. ja. d o m e s tic u s , Mareca a m e r i c a n a , Anas r u b r i p e s ,
Nyroca a m e r i c a n a , Nyroca a f f i n i s . Nyroca v a l i s i n a r i a .
L ocality:
Reference:

N orth America (M ichigan)
p r e s e n t paper
Genus M e to r c h i s L o o ss, 1899

Body somewhat com pre ssed , more or l e s s p o in te d a t
a n t e r i o r end.

Skin s p in e d i n most c a s e s .

The i n t e s t i n a l

•crura r e a c h i n g o n ly to the r e g i o n o f the t e s t e s in most
species.

The t e s t e s a r e e n t i r e to l o b e d , and a r e more o f t e n

(78)

a t t h e same l e v e l th a n one behind t h e o t h e r .

The c o i l s o f

th e u t e r u s a r e com pact, and o v e r l a p the i n t e s t i n a l c r u r a .
The v i t e l l a r i a a r e l a t e r a l to th e i n t e s t i n e s , and b e g i n
b e fo re the acetabulum .
Type s p e c i e s :

M. a l b i d u s (B raun, 1893)

M e t o r c h i s c r a s s i u s c u l u s ( B u d o lp h i, 1809)
H o s t:

Mergus s e r r a t o r , Anas q u e r q u e d u l a . and a number

o f hawks and e a g l e s .
L ocality:
R eference:

Europe
5 7 , 61

M e t o r c h i s o r i e n t a l i s Tanabe, 19 21
Host:

"Bucks"

L o cality :
R eference:

Ja p an
6

M e t o r c h i s t e n e r Kowalewski, 1903
H o s t:

Mergus m e rg a n se r ( L i v e r )

L ocality:
R eference:

Europe
57

M e t o r c h i s zanthosomus ( C r e p l i n , 1846)
Host:

Oidemia n i g r a , P odiceps r u f l c o l l i s . Mergus

s e r r a t o r , Anas p l a t y r h y n o h o s , Anas q u e r q u e d u l a .
L ocality:
R eference:

Europe
18

(79)

F a m ily EETEROFHYIDAE Odhner, 1914
Synonyms:

Coenogonimidae N i c o l l , 1907; C otylogonim idae

N i o o l l , 1907; E i c r o p h a l l i d a e D r a v a s s o s , 1920; A p l o r c h i d a e
Vi&na, 1924; E a p l o r c h i d a e , T r a v a s s o s , 1924.
S n a i l t o v e r y s m a ll t r e m a t o d e s w ith body covered w i t h
s c a l e - l i k e s p i n e s and f r e q u e n t l y w i t h a crown o f c i r c u m o r a l
spines.

Pharynx alw ays p r e s e n t .

Body u s u a l l y d i v i d e d i n t o

a m o t i l e a n t e r i o r , f l a t t e n e d r e g i o n devoid o f g e n i t a l i a ,
and a p o s t e r i o r p a r t c o n t a i n i n g the g e n i t a l o r g a n s .

V entral

s u c k e r u s u a l l y r ed u ce d and i n t i m a t e l y a s s o c i a t e d w ith the
g e n ita l pore.

G e n i t a l d u c t s u s u a l l y opening i n t o a common

g e n i t a l s i n u s which f r e q u e n t l y c o n t a i n s a c o p u l a t o r y organ
known a s a g o n o t y l .
in p o s itio n .

G e n i t a l p o re e i t h e r median or l a t e r a l

Ovary and t e s t e s h i g h l y v a r i a b l e i n s h a p e ,

t h e o v a r y a l m o s t alw ays a n t e r i o r to t h e t e s t e s .
pouch l a c k i n g .

Seminal r e c e p t a c l e v o lu m in o u s.

C irrus
U te r u s

u s u a l l y n o t e x t e n d i n g a n t e r i o r to th e g e n i t a l p o r e .

Para­

s i t e s o f mammals, b i r d s , and f i s h e s .
Genus Tocotrema L o o s s , 1899
Synonyms:

G r y p t o c o t y l e Luhe, 1899; B e rm o c y stls S t a f f o r d ,

1905; E allum Wigdor, 1916.
Pharynx p r e s e n t ; esophagus s h o r t , b i f u r c a t i o n n e a r e r
t h e o r a l s u c k e r th a n t h e a c e t a b u l u m .

I n t e s t i n a l ca ec a extend

(80)

to t h e p o s t e r i o r end o f t h e b o d y , t e r m i n a t i n g beh in d t e s t e s *
V e n t r a l s u c k e r , m e d ia n , more or l e s s r u d i m e n t a r y , in r e ­
l a t i o n w ith th e a n t e r i o r p o r t i o n o f th e g e n i t a l s u c k e r ,
and com m unicating w i t h t h e e x t e r i o r th ro u g h the g e n i t a l
p o re which i s l o c a t e d i n th e c e n t e r o f the g e n i t a l s u c k e r .
G e n i t a l s u c k e r w e l l developed , s i t u a t e d i n the median l i n e
a b o u t midway o f th e bod y .

The g e n i t a l s i n u s which opens

t o th e e x t e r i o r th r o u g h th e g e n i t a l p o r e , and i n t o which
th e vas d e f e r e n s and v a g i n a o pen , i s p o s t e r i o r o f the
v e n tra l sucker.

A g e n i t a l p a p i l l a e p r o j e c t s i n t o th e g e n i t a l

s i n u s and i s p r o t r u s i b l e through th e g e n i t a l p o r e .

P ro static

p o r t i o n o f v a s d e f e r e n s w e l l d e v e l o p e d , d o r s a l to and behind
t h e g e n t i a l su c k e r *

Seminal v e s i c l e v/ell d e v e lo p e d , s i t u a t e d

behind th e g e n i t a l s u c k e r, arranged in tra n s v e rs e lo o p s,
d o r s a l o f th e c o i l s o f t h e u t e r u s .

T estes near p o s te r io r

end o f b o d y , i r r e g u l a r l y o v a l or g l o b u l a r , and u s u a l l y
s l i g h t l y l o b e d , s i d e by s i d e , o r r i g h t t e s t i s o b l i q u e l y
b eh in d t h e l e f t .

Seminal r e c e p t a c l e i n f r o n t o f and to the

rig h t of l e f t t e s t i s .

Ovary i r r e g u l a r l y o v a l , or u s u a l l y

l o b e d , commonly l i k e a c l o v e r l e a f , s i t u a t e d on the r i g h t
o f th e median l i n e , i n f r o n t o f th e se m in al r e c e p t a b l e .
V i t e l l a r i a extend a c r o s s i n f r o n t o f g e n i t a l p o r e , on the
d o r s a l s i d e o f the b r a n c h e s o f the i n t e s t i n e and o f t e n

(81)

meet i n t h e median l i n e .

A n t e r i o r l i m i t s o f th e v i t e l l a r i a

u s u a l l y a t a c o n s i d e r a b l e d i s t a n c e p o s t e r i o r o f th e b i f u r ­
c a t i o n o f t h e i n t e s t i n e ; p o s t e r i o r l y th e y extend behind
t h e t e s t e s and u s u a l l y c r o s s the median l i n e .

Transverse

v i t e l l i n e d u c t s l o c a t e d i n th e neighborhoo d o f the boundary
betw een t h e o v a r i a n and t e s t i c u l a r z o n e s .

U teru s d is p o s e d

i n a few l o o p s i n t h e median l i n e ; none i n f r o n t o f the
g e n ita l pore.
Type s p e c i e s :

T. l i n g u a ( C r e p l i n , 1 8 2 5 ) , F i s c h o e d e r ,

1903.
Tocotrema ooncavum ( C r e p l i n , 1825) F i s c h o e d e r , 1903
H o s t:

Mergus m e r g a n s e r . Mergus s e r r a t a , Anas h o r n s -

c u c h i i . Nyroca m a r i l a . Nyroca c l a n g u l a , Nyroca h y e m a l i s ,
Oidemia f u s c a . and a number o f o t h e r w a te r b i r d s ,
lo cality :

Europe

Reference:

86, 110

F am ily MICR03PHALLLDAE T r a v a s s o s , 1920
Synonyms:

H e t e r o p h y i d a e , Odhner, 1914; M i c r o p h a l l i n a e

( H e t e r o p h y i d a e ) Ward, 1901.
Small to v e r y s m a ll t r e m a t o d e s , s u p e r f i c i a l l y r ese m ­
b l i n g t h e l a s t f a m i l y , and f r e q u e n t l y c l a s s i f i e d as a s u b ­
f a m i l y o f th e H e t e r o p h y i d a e .

Body s p in e d or smooth.

A cetab­

(82)

ulum and g e n i t a l o p e n i n g s s e p a r a t e d .

I n t e s t i n a l c ru ra not

r e a c h i n g beyond t h e a n t e r i o r b o r d e r of the t e a t e a , and o f t e n
n o t beyond t h e a c e t a b u l u m .
o f the acetabulum .

G e n i t a l pore u s u a l l y to the l e f t

The g e n i t a l a t r i u m c o n t a i n s a m u sc u la r

p a p i l l a e , or a p r o t r u s i b l e c i r r u s .
is p rese n t.

A l a r g e s e m in a l v e s i c l e

The t e s t e s u s u a l l y l i e a t the same l e v e l , behind

the a c e t a b u l u m , th e o v a r y b e i n g a n t e r i o r , and u s u a l l y to
th e r i g h t o f th e m i d - l i n e .

V ite lla r ia post-acetabular.

P a r a s i t e s o f f i s h , r e p t i l e s , b i r d s and mammals.
Genus S p e lo tr e m a J a g e r s k i o l d , 1901
M icrophallidae.

Small f l u k e s , w ith s p in e d s k i n .

I n t e s t i n e s n o t r e a c h i n g beyond a n t e r i o r b o r d e r o f t e s t e s .
E x c r e t o r y b l a d d e r s m a l l , n o t e x t e n d i n g to the t e s t e s .
T e s t e s l i e a t th e same l e v e l .
o f 6-8 s e p a r a t e f o l l i c l e s .

V i t e l l a r i a l a t e r a l , composed

G e n i t a l a t r i u m always opening

th r o u g h a m u s c u la r p a p i l l a e .

No c o m p lic ate d d i v e r t i c u l a a s

in the case o f S p e lo p h a llu s , p r e s e n t.
th e b a s e o f th e male p a p i l l a e .

The v a g in a opens in

P a rs p r o s t a t i c a and se m in a l

v e s i c l e w ell developed.
Type s p e c i e s :

S p elotrem a pygmaeum (Le v inse n 1981)

S p e lo tr e m a pygmaeum (L e v in se n 1881)
H ost:

S o m a te ri a m o l l i s s i m a , Oidemia n i g r a , G l a u o i o n e t t a

olangula am ericana.

(83)

L ocality:
R eference:

E u ro p e , North America
37 , 40

Genus M a ritre m a N i c o l l , 1907
Synonym:

S t r e p t o v i t e l l a S w ales, 1933

Small f l u k e s w i t h a s p in e d s k i n .
developed.

Suckers m o d e r a t e l y

P r e p h a r y n x , pharynx and esophagus p r e s e n t ,

c r u r a n e v e r r e a c h i n g heyond a n t e r i o r b o r d e r o f t e s t e s .
E x c r e t o r y b l a d d e r "V" s h a p e d , l a r g e , o f t e n e x t e n d i n g
a n t e r i o r to th e t e s t e s .

G e n i t a l p o re a t l e f t o f a c e t a b ­

ulum w i t h a lo n g c i r r u s pouch l y i n g t r a n s v e r 3 l y between
the a c e ta b u lu m and t h e i n t e s t i n a l c r u r a .

Ovary r i g h t of

m i d - l i n e , f r e q u e n t l y o v e r l a p p i n g a c e ta b u lu m .
t h e same l e v e l , beh in d a c e t a b u l u m .

Testes a t

V itellaria la te ra l

e x t e n d i n g mesiad a n t e r i o r to t e s t e s , f r e q u e n t l y forming
a c o m p lete c i r c l e i n p o s t e r i o r h a l f o f body.
f i l l i n g p o s t e r i o r h a l f o f body.
Type s p e c i e s :

M. g r a t i o s u m N i c o l l , 1907

M a ritre m a a c a d i a e S w a l e s , 1933
Host:

Anas r u b r i p e s

L ocality:
R eference:

North America (Canada)
133

M a r it r e m a n e t t a e n . s p .

U te r u s

(84)

H ost:

G l a u c i o n e t t a c l a n g u l a a m e r i c a n a . Nyroca a f f i n i s ,

lo c ality :

N orth America (Michigan)

R eference:

p r e s e n t paper

Genus L e v i n s e n i e l l a S t i l e s and H a s s e l l , 1901
Synonym:

L e v i n s e n i a J a g e r 3 k i o l d . 1900

Body s m a l l to v e r y s m a l l , s h o r t , f o r the most p a r t
pear shaped.

S uckers r a t h e r s m a l l .

Esophagus l o n g .

Pharynx

l y i n g n e a r th e o r a l s u c k e r , a l t h o u g h a p rep h ary n x i s always
present.

I n te s t in a l crura s h o rt.

G e n i t a l pore l y i n g a t a b o u t

th e same l e v e l a s a c e t a b u l u m , w ith a more or l e s s m usc ular
g en ital sin u s.

P e n is a b s e n t .

V e s i c u l a s e m i n a l i s l y i n g some­

what i n f r o n t o f th e a c e t a b u l u m , the two t e s t e s s l i g h t l y
p o s t e r i o r to i t , and a t the same l e v e l .
to t h e r i g h t o f th e a c e t a b u l u m .
L a u re r's canal p re s e n t.

The ovary l i e s j u s t

Receptaculum s e m in is a b s e n t ,

The s m a l l , compact, r o s e t t e l i k e

v i t e l l i n e r e s e r v o i r l i e s im m ed ia tely behind th e a c e ta b u lu m .
The u t e r u s i s l i m i t e d to th e p o s t e r i o r p a r t o f the body.
The e x c r e t o r y b l a d d e r i s r e l a t i v e l y l a r g e and rtV,t s h a p e d .
Type s p e c i e s :

L. brachysoma ( G r e p l i n , 1957)

L e v i n s e n i e l l a brachysoma ( C r e p l i n , 1937)
H ost:

C la n g u la h y e m a l i s , Oidemia n i g r a . Bucephala

c l a n g u l a , and a number o f o t h e r w a te r b i r d s .
L ocality:

Europe

(85)

R eference:

3 8 , 65

L e v i n s e n i e l l a m in u t a P r i c e , 1934
H ost:

Nyroca a f f i n i a

L o cality :

West I n d i e s

R e f e r e n c e : 107
L e v i n s e n i e l l a p e l l u o i d a J a g e r s k i o l d , 1907
H ost:

Anas p l a t y r h y n o h o s , Nyroca f u l i g u l a ( I n t e s t i n e )

L ocality:
R eference:

Europe
3 8 , 65

Genus S p e l o p h a l l u s J a g e r s k i o l d
Body s m a l l , s h o r t , b i s c u i t or p e a r s h a p e d .
ra th e r sm all.

Esophagus l o n g .

prepharynx p r e s e n t .

Pharynx n e a r th e o r a l s u c k e r ,

Crura s h o r t and w id e .

Acetabulum l o c a t e d

a t th e b e g i n n i n g o f the p o s t e r i o r body t h i r d .
i n g l y i n g n e a r th e a c e ta b u lu m .

Suckers

G e n i t a l open­

The v a g i n a opens n e a r the

g e n i t a l p o r e i n the g e n i t a l s i n u s , r a t h e r th a n a t the ba3e
o f th e g e n i t a l s i n u s a s i n S p e l o t r e m a .
c o n t a i n s a "K eg e lfo rm ig en K o r p e r " .
w e ll d e v e l o p e d .
ulum.

The g e n i t a l s i n u s

The p r o s t a t i c c e l l s a r e

V e s i c u l a s e r a i n a l i s l a r g e , b e f o r e th e a c e t a b ­

T e s t e s a t th e same l e v e l behind th e v e n t r a l s u c k e r .

C i r r u s pouch a b s e n t .

Ovary to th e r i g h t o f th e a c e ta b u lu m .

R ece p tac u lu m s e m in is a b s e n t .

The l a r g e , com pact, r o s e t t e

v i t e l l a r i a l i e sy m m e tr ic a l behind th e t e s t e s , th e \ i t e l l i n e
r e s e r v o i r im m e d ia te ly behind the a c e ta b u lu m .

U t e r u s c o n f in e d

(86)

to th e p o s t e r i o r p a r t o f th e body.

E x c r e t o r y b l a d d e r "V"

shaped.
Type s p e c i e s :

S. primus J a g e r s k i o l d , 1908

S p e l o p h a l l u s prim us J a g e r s k i o l d , 1908
H o s t:

S o m a te r i a m o l l i s s i m a . Haematopus o s t r a l e g u s

lo cality :

Europe

Reference:

3 9 , 65

Family ORCHIPEDIME S k r j a b i n , 1924
More or l e s s e l o n g a t e worms w i t h a smooth s k i n .
t e r i o r end somewhat s e t o f f from th e p o s t e r i o r end.
ulum somewhat l a r g e r th a n th e o r a l s u c k e r .
esophagus a b s e n t .

An­
A c e ta b ­

Pharynx p r e s e n t ,

I n t e s t i n e s p a r a l l e l to s i d e s o f bo d y ,

r e a c h i n g to n e a r th e p o s t e r i o r end o f the body.

Testes

num erou s, l y i n g betw e en the i n t e s t i n a l c r u r a , and f i l l i n g
most o f t h e s p a c e from n e a r the o vary to n e a r the p o s t e r i o r
end.

C i r r u s pouch a b s e n t .

short.

Seminal v e s i c l e l a r g e , u t e r u s

V i t e l l a r i a v/ell d e v e l o p e d , composed o f numerous

f o l l i c l e s , e x t e n d i n g from th e p o s t e r i o r end to th e a c e t a b ­
ulu m , l y i n g d o r s a l , l a t e r a l , and v e n t r a l to the c r u r a .
G e n i t a l p o r e m e d ia n , n e a r the p h a ry n x .
p resen t.

Eggs l a r g e , b u t few i n number.

P arasites of b ird s.

L a u re r's canal

(87)

Genua Orchipedum B ra u n , 1908
S in ce t h i s i s t h e o n ly genus i n th e f a m i l y , i t w i l l
n o t be n e c e s s a r y to r e p e a t the d e s c r i p t i o n .
Type s p e c i e s :

()• t r a c h e i c o l a . B ra u n , 1901

Orchipedum t r a c h e i c o l a B r a u n , 1901
Host:

Oidemia f u s c a (T rachea)

L ocality:

Europe

R eference:

18

Fam ily PHILOPHTHALMIBAE L ooss, 1899
Medium s i z e d , m u s c u la r f l u k e s , w i t h unspined skin^
S u ckers s t r o n g l y d e v e l o p e d .
sucker.

Pharynx l a r g e r than th e o r a l

Esophagus v e r y s h o r t .

T e s t e s c l o s e t o g e t h e r , and

one beh in d t h e o t h e r i n th e p o s t e r i o r end o f the body.
C i r r u s pouch w e l l d e v e l o p e d , e l o n g a t e .
f r o n t o f the t e s t e s .

C o i l s o f th e u t e r u s numerous, r e a c h i n g

a n t e r i o r to the a c e t a b u l u m .

V itellaria la te ra l.

p ore l y i n g i n f r o n t o f the a c e ta b u l u m .
ent.

Ovary l y i n g i n

G enital

L a u re r's canal pre-

Eggs l a r g e , c o n t a i n i n g a developed m ir a c id iu m .
These forms a r e e c t o p a r a s i t i c , and a r e found i n the

e y e , b o r d e r o f the c l o a c a e t c . , o f b i r d s .
Genus P h i lo p h th a lm u s Looss, 1899
With t h e c h a r a c t e r s o f the f a m i l y .

C i r r u s pouch and

(88)

v a g i n a v e r y l o n g , r e a c h i n g p o s t e r i o r to th e a c e ta b u lu m .
V i t e l l a r i a always i n th e shape o f a sim p le t u b e , l y i n g
l a t e r a l to t h e c r u r a , and p a s s i n g mesiad i n the r e g i o n o f
th e o v a r y .
Type s p e c i e s :

P. p a lp e b r a r u m L o o s s , 1899

P h i l o p h t h a l m u s a n a t i n u s Sugimoto, 1928
H ost:

Anas p l a t y r h y n o h o s

L ocality:
Reference:

Formosa
123

Fam ily CYCLOCOELIDAE K ossack, 1911
E n d o p a r a s i t i c tr e m a t o d e s w ith l a r g e to m iddle s i z e d ,
m u s c u la r b o d i e s .

Kouth opening t e r m i n a l or sub t e r m i n a l ,

s u r ro u n d e d by a m u s c u la r s u c k e r , u s u a l l y much r e d u c e d ,
and e n t i r e l y w a n t i n g in some c a s e s .

Acetabulum r a r e l y

p r e s e n t , pharynx l a r g e , m u s c u l a r ; esophagus lo n g .

In testin ­

a l c r u r a s i m p l e or p o s s e s s i n g i n t e r n a l c a e c a , an a sto m o s in g
i n t h e p o s t e r i o r end o f th e body.

E x c r e t o r y b l a d d e r between

p o s t e r i o r i n t e s t i n a l a r c h and end o f body w ith median d o r s o term inal pore.
ynx.

G e n i t a l pore m ed ian, u s u a l l y v e n t r a l to p h a r ­

C o p u l a t i o n o r g a n s p r e s e n t , w ell d ev e lo p e d ; se m in a l

v e s i c l e i n c i r r u s pouch.

V i t e l l a r i a i n g e n e r a l l y i n g between

body w a l l and i n t e s t i n a l c r u r a , sometimes s u r r o u n d i n g the
la tte r.

G e n i t a l g l a n d s between i n t e s t i n a l c r u r a , sim p le or

l o b e d , form ing the p o i n t s o f a t r i a n g l e , or i n a s t r a i g h t

(89)

lin e*

L a u r e r ' s c a n a l w a n ti n g ; re c e p ta b u lu m se m in is p r e s e n t .

U te r u s s t r o n g l y d e v e l o p e d , l y i n g i n more or l e s s r e g u l a r
f o l d s betw een i n t e s t i n a l c r u r a over ?/hich the y sometimes
e x t e n d ; u s u a l l y f i l l i n g e n t i r e sp a ce between c r u r a .

Eggs

numerous, w i t h o u t p o l a r f i l a m e n t , u s u a l l y c o n t a i n i n g w ell
developed m i r a c i d i a w i t h c h a r a c t e r i s t i c double eye s p o t s .
P a r a s i t i c i n body c a v i t y , l u n g s , t r a c h e a , and n a s a l
c a v i t i e s o f w ater b i r d s .
Genus Cyclocoelum B r a n d e s , 1892
Medium to l a r g e tr e m a t o d e s w ith a m u s c u l a r , somewhat
f l a t t e n e d b o d y , somewhat t a p e r i n g a t th e a n t e r i o r end, and
b r o a d l y rounded a t t h e p o s t e r i o r end.

The mouth opening i s

t e r m i n a l , and i s su rro u n d e d by a weakly m uscular s u c k e r .
The i n t e s t i n a l c r u r a a r e s i m p l e , a n a sto m o s in g a t t h e p o s ­
t e r i o r end.

The g e n i t a l pore u s u a l l y l i e s a t the h e i g h t o f

the a n t e r i o r b o r d e r o f the ph ary n x .

The c i r r u s pouch i s

r e l a t i v e l y s m a l l , l y i n g to th e s i d e o f the esophagus, and
sometimes r e a c h i n g p o s t e r i o r to the i n t e s t i n a l c r u r a .
c i r r u s i s sm a ll and c y l i n d r i c a l .

The

The v i t e l l a r i a l i e l a t e r a l

to t h e i n t e s t i n a l c r u r a , th e y a r e composed o f numerous s m a ll
f o l l i c l e s , e x t e n d i n g from th e r e g i o n o f the b i f u r c a t i o n o f
the i n t e s t i n e to th e e x c r e t o r y b l a d d e r .

The e x c r e t o r y

(90)

b l a d d e r i s t y p i c a l o f th e f a m i l y .

The sex g la n d s form a

t r i a n g l e i n th e p o s t e r i o r l o c p o f th e i n t e s t i n e , th e t e s t e s
u s u a l l y l a r g e r th a n th e o v a r y .
o n ly s l i g h t l y l o b e d .

The t e s t e s a r e e n t i r e or

The u t e r u s f i l l s p r a c t i c a l l y the e n t i r e

s p a c e bounded by t h e i n t e s t i n e , th e c o i l s b e i n g more or l e s s
reg u lar.

The s e x g l a n d s a r e i n some c a s e s s e p a r a t e d by f o l d s

o f the u t e r u s .

The eggs a r e t h i c k s h e l l e d , and c o n t a i n

m i r a c i d i a when d e p o s i t e d .
Type s p e c i e s :

£ . m u t a b i l e ( Z e d e r , 1800)

Cyolocoelum a r o u a t u s (B r a n d e s , 1892)
H o s t:

B u c ep h a la o l a n g u l a , Kergua a l b e l l u s

L o cality:
R eference:

Europe
46 , 49, 161

Cyolocoelum c o e l o n o t u s W ite n b e rg , 1926
H o st:

Nyroca f e r i n a

L ocality:
R eference:

Europe
46, 161

C y c lo clelu m l a e v i g a t u s K ossack , 1911
H o s t:

Nyroca f u l i g u l a , Oidemia f u a e a , 0. n i g r a ,

S o m a te ria m o l l i s s i m a , C lan g u la h yem alis
L o cality:
R eference:

Europe
46, 49, 161

Cyolocoelum pseudomicrostomum H a r r a h , 19 22

(91)

H ost:

"w ild duck"

L o cality :

North America and Europe (Th is s p e c i e s has

been ta k e n from the body c a v i t y o f the c o o t a t V /intergreen
Lake)
Reference:

34, 46, 125

Oyclocoelum r o b u s t u 3 ( S t o s s i o h , 1902)
Hosti

Nyroca f u l i g u l a

L o cality:
Reference:

Europe
4 6 , 4 9 , 126, 161

Genus Typhlocoelum S t o s s i c h , 1902
Synonym:

Typhlultimum W ite n b e rg , 1924; T r a c h e o p h ilu s

S k r j a b i n , 1913.
With th e c h a r a c t e r s o f th e f a m i l y .

Medium to l a r g e

t r e m a t o d e s w i t h a v e r y m u s c u la r body, f l a t t e n e d , and b r o a d ­
l y rounded a t e i t h e r end.

The mouth opening i s s u b t e r m i n a l ,

may o r may n o t be surro u n d e d by a weak s u c k e r .

There i s

o f t e n a s m a ll t o n g u e - l i k e p r o j e c t i o n a n t e r i o r to th e mouth
opening.

A s m a ll v e n t r a l s u c k e r may be p r e s e n t .

i s m u s c u l a r , t h e esophagus b e i n g v e r y s h o r t .

The pharynx

The i n t e s t i n a l

c r u r a have d i v e r t i c u l a e from t h e i n n e r b o r d e r , th e y a n a s t o ­
mose a t th e p o s t e r i o r end.
or e n t i r e .

The t e s t e s may be s t r o n g l y lobed

V i t e l l a r i a l a t e r a l , e x t e n d i n g d o r s a l and v e n t r a l

(92)

o f th e c r u r a .
Cycloooelum.

The e x c r e t o r y b l a d d e r i s s i m i l a r to
The c o i l s o f t h e u t e r u s a r e v e r y h e a v y ,

e n t i r e l y f i l l i n g th e a v a i l a b l e sp a ce w i t h i n the c r u r a .
Eggs y e l l o w , c o n t a i n i n g a m a ture m i r a c i d i u m .
Type s p e c i e s :

Typlocoelum cucumerinum( R u d o l p h i . 1®09)

Typhlocoelum cucumerinum ( R u d o lp h i, 1809)
H ost:

Anas p l a t y r h y n c h o s . Oidemia f u s c a , Nyroca f u l i g u l a ,

may i T a . S o m a te r i a m o l l l s s i m a . Olangula hyem alis , Mergus
a l b e l l u s . M. s e r r a t o r .
L o cality :
Reference:

Worth A m e ric a , South A m erica, Europe
46, 49, 71, 126, 161

T yphlocoelum cymbium B i e s i n g , 1850
H ost:

Anas p l a t y r h y n c h o s , Wyroca a f f l n i s , S p a t u l a

o l y p e a t a . Anas r u b r i p e s , Podylymbus p o d i c e p s , L a f i l a a c u t a ,
L o cality:
Reference:

Worth A m erica, South A merica, J a p a n , Europe.
4 6 , 4 9 , 71, 126, 131, 161.
Genus Ophthalmophagua S t o s s i c h , 1902

Synonyms:

Hyptiasmus Z o s s a c k , 1911; Spanometra Z o ssa c k ,

1911; Transcoelum W i te n b e r g , 1926
Body f l a t t e n e d , m u s c u l a r , the ends r o u n d e d .
w e l l d e v e l o p e d , the i n t e s t i n e s s i m p l e , u n b r a n c h e d .

Pharynx
The

g e n i t a l pore l i e s v e n t r a l a b o u t mid-way o f th e p harynx.

The

c i r r u s pouch r e a c h e s p o s t e r i o r a s h o r t d i s t a n c e behind the
crura.

The v i t e l l a r i a l i e l a t e r a l to the i n t e s t i n e s , and

(93)

s u r ro u n d them d o r s a l and v e n t r a l , th e y r e a c h from the b i ­
f u r c a t i o n t o th e p o s t e r i o r end o f the body.
g la n d s l i e

The t h r e e sex

i n a more or l e s s s t r a i g h t l i n e , the p o s t e r i o r

t e s t e s b e i n g n e a r th e p o s t e r i o r l i m i t o f th e i n t e s t i n a l
loop.

The o v a r y i s somewhat s m a l l e r , and between the t e s t e s .

The c o i l s o f th e u t e r u s extend l a t e r a l to th e i n t e s t i n a l
crura.

The eggs a r e t h i c k - s h e l l e d , y e l l o w , and oval i n s h a p e .

Type s p e c i e s :

0. s i n g u l a r i s S t o s s i c h , 1902

Ophthalmophagus m a s s i n o i ( W ite n b e r g , 1926)
H ost:

"wild duck"

L ocality:

R ussia

Reference:

161

F am ily EUCOTYLIDAE S k r j a b i n , 1924
Monostomes.

Mouth opening s u b t e r m i n a l .

esophagus p r e s e n t or a b s e n t .

Pharynx p r e s e n t ,

I n t e s t i n a l c r u r a r e a c h i n g to

n e a r t h e p o s t e r i o r b o r d e r o f the body.

Testes behind,

t r a n s v e r s e t o , o r a t th e same h e i g h t o f the o v a r y , l y i n g
l a t e r a l to th e i n t e s t i n a l c r u r a n e a r the s i d e s o f the body.
C i r r u s pouch a b s e n t .

U teru s i n numerous f o l d s , w inding

t r a n s v e r s l y to n e a r th e p o s t e r i o r end o f the bod y , a few
c o i l s a n t e r i o r to t h e g e n i t a l p o r e .
the i n t e s t i n e s .

V i t e l l a r i a l a t e r a l to

G e n i t a l pore behind th e b i f u r c a t i o n o f the

(94)

in testin es*

Eggs s m a l l , numerous, w i t h o u t f i l a m e n t s .

P a r a s i t i c i n th e k id n e y s o f b i r d s .
Genus E u c o t y l e Cohn, 1904
E l o n g a t e d , f l a t t e n e d , medium s i z e d monos tomes; a n t e r i o r
end t r i a n g u l a r and s e t o f f from th e re m a in d e r o f body by a
d o r s a l and v e n t r a l t r a n s v e r s e m u s c u la r r i d g e ; p o s t e r i o r end
rounded*

T e s t e s o p p o s i t e each o t h e r , e i t h e r e n t i r e l y e x t r a -

c a e c a l , p a r t l y o v e r l y i n g th e c a e c a , or occupying th e e n t i r e
w i d t h o f t h e body v/ith th e median b o r d e r s to u c h i n g .

Ovary

l o b a t e , p r e t e s t i c u l a r , and to th e s i d e o f the median l i n e .
Y i t e l l a r i a e x t r a c a e c a l , commencing im m edia tely p o s t e r i o r to
th e t r a n s v e r s e m u s c u la r r i d g e and e x t e n d i n g backward a s f a r
a s t h e t e s t e s or beyond.
Type s p e c i e s ;

E. n e p h r i t i c a ( M e h li s , 1846) Cohn, 1904

E u c o t y l e zakharow i S k r j a b i n , 1920
H o s t:

Nyrooa f u l i g u l a

L ocality:

Europe

Reference:

116

E u c o t y l e w e h r i P r i c e , 1930
H ost:

Q uequerdula d i s o o r s

L o cality:
R eference:

N o rth America
104

(95)

Fam ily N0TOCOTYLIBAE Luhe, 1909
Small monostomes t a p e r i n g a t b o th e n d s , p o s t e r i o r end
b r o a d l y r o u n d e d , a n t e r i o r s l i g h t l y more a t t e n u a t e d .

Gener­

a l l y w i t h rows o f p a p i l l a e on the v e n t r a l s u r f a c e formed o f
u n i c e l l u l a r dermal g l a n d s .

Esophagus s h o r t , w i t h o u t pharynx;

i n t e s t i n a l caeca w ith s h o r t d i v e r t i c u l a , extending e n t i r e
l e n g t h o f bo d y .
where i t

G e n i t a l po re m e d ian , e x c e p t i n Hudoootyle

is d i s t i n c t l y l a t e r a l , u s u a lly near o ral sucker.

C irru s sac e lo n g a te .
p o s t e r i o r en d .

T estes sym m etrical, e x t r a c a e o a l , near

Ovary between t e s t e s .

a n t e r i o r to t e s t e s .

V itellaria la te ra l,

U t e r i n e c o i l s between c i r r u s sac and

g e n i t a l g la n d s , t r a n s v e r s e , r e g u l a r , u s u a lly not extending
outside i n t e s t i n a l c ru ra .

Eggs sm all w ith lo n g p o l a r f i l a ­

ment on each end.
P a r a s i t e s o f th e i n t e s t i n e and caeca o f b i r d s and
mammals.
Genus H o to c o t y l u s B i e s i n g , 1839
Body e l o n g a t e , b r o a d l y rounded a t the p o s t e r i o r end,
t a p e r i n g more a t t h e a n t e r i o r end.
l y s e t w ith f i n e s p i n e s .

The v e n t r a l s u r f a c e t h i c k ­

Three to f i v e rows o f v e n t r a l g la n d s

p r e s e n t , which a r e p r o t r u s i b l e .

The v a g i n a i s v e r y m u s c u l a r ,

and i s a b o u t h a l f a s lo n g a s th e c i r r u s pouch.
Type s p e c i e s :

N o to c o t y lu s a t t e n u a t u s ( R u d o lp h i, 1809)

(96)

N o t o c o t y l u a a t t e r m a t u s ( R u d o lp h i, 1809)
H0 3 1 :

Anas p l a t y r h y n c h o s . A. p e n e l o p e . A. o r e c c a ,

A. a c u t a , C l a n g u la h y e m a i l s , S p a t u l a c l y p e a t a . Mergus
m e r g a n s e r . S o m a te r i a m o l l i 3 3lm a. and a number o f o t h e r
b i r d s i n c l u d i n g c h i c k e n s and g e e s e .
L ocality:

Europe

R eference:

45, 49 , 65, 96

N o t o c o t y l u s a e g y p t i a c u s (Odhner, 1905)
H o s t:

Anas p l a t y r h y n c h o s

L ocality:

Egypt

Reference:

95

N o t o c o t y l u s s e i n i t a Fuhrmann, 1919
Host:

Anas p l a t y r h y n c h o s , A. querq u ed u la

L o cality :

Europe

R eference:

31

No t o c o t y l u s u r b a n e n s i s ( C o r t , 1914)
H o s t:

D a f i l a a c u t a , Aix s p o n s a , F i b e r z i b e t h i c u s ,

Anas p l a t y r h y n c h o s
L ocality:

North America

Reference:

34, p r e s e n t paper
Genus C a t a t r o p i s Cdhner, 1905

Body e l o n g a t e , a n t e r i o r and p o s t e r i o r ends u s u a l l y
eq u a lly rounded.

A n t e r i o r h a l f o f v e n t r a l s u r f a c e covered

(97)

w i t h t h r e e rows o f n o n - p r o t r u s i b l e p a p i l l a e ; median row
s e t on a r i d g e or k e e l ; l a t e r a l rows each c o n t a i n i n g e i s ’h t
to t w e l v e g l a n d s .

Vagina s t r o n g l y d e v e lo p e d , u s u a l l y a s

l o n g a s c i r r u s pouch.
Type s p e c i e s :

C a t a t r o p i s v e r r u c o s a ( F r o e l i c h , 1789)

O a t a t r o p i s v e r r u c o s a ( P r o e l i c h , 1789)
Host:

Anas p l a t y r h y n c h o s , S o m a te ria m o l l i s s i m a , Mergus

s e r r a t o r , B u c ep h a la c l a n g u l a , C la n g u la h y e m a i l s . Oidemia
n i g r a . a s w e ll a s c h i c k e n s and g e e 3 e .
L ocality:
Reference:

Europe
49, 65

Genus Paramonos tomum Luhe , 1909
Body c o m p re ssed , egg s h a p e d , g r e a t e s t b r e a d t h a l i t t l e
caudad from m id d le o f bod y, p o s t e r i o r end b r o a d l y ro unded,
an terio r

t a p e r i n g and p o i n t e d ; a n t e r i o r h a l f o f v e n t r a l

s u r f a c e t h i c k s e t w i t h s h o r t heavy s p i n e s .
absent.

C i r r u s pouoh weakly m u s c u l a r .

V e n t r a l g la n d s

Vagina u s u a l l y one-

h a l f l e n g t h o f c i r r u s pouch.
Type s p e c i e s :

P.

a l v e a tu m ( M e h li s , 1846)

Paramonostomum a l v e a t u m ( M e h li s , 1846)
H ost:

Oidemia f u s c a , Nyroca m a r i l a , C lang ula h y e m a ils

S o m a te ri a m o l l i s s i m a , Anser a n s e r , B r a n ta b e r n i c l a , Gjrgnua,

(98)

oygnus.
L o cality :

Europe

R eference:

6, 49

Family ECHINOSTOUIBA.E D i e t z , 1910
Body more or l e s s e l o n g a t e .

A n t e r i o r end w ith a c o l l a r ,

i n th e c e n t e r o f which i s th e mouth opening and o r a l s u c k e r ,
and on which a r e th e l a r g e c o l l a r s p i n e s , the number o f which
are constant for a sp e cie s.

The s p i n e s may be i n a double or

s i n g l e row , or may be d iv id e d i n t o two l a t e r a l f i e l d s .
u s u a l l y s p in e d over a n t e r i o r h a l f o f body.
close to g e th er.

Skin

Suckers r a t h e r

Acetabulum u s u a l l y l a r g e r than o r a l s u c k e r ,

and v e r y m u s c u l a r .

Pharynx p r e s e n t .

In te s tin a l bifu rcatio n

a n t e r i o r to v e n t r a l s u c k e r .

I n t e s t i n a l c r u r a r e a c h to or

n e a r p o s t e r i o r end o f body.

E x c r e t o r y b l a d d e r HYrt sh a p ed .

The t e s t e s l i e one beh in d th e o t h e r , u s u a l l y in th e p o s t e r i o r
p o r t i o n o f t h e body.

A c i r r u s pouch u s u a l l y w e ll d e v e lo p e d ,

p l a c e d d o r s a l to th e a c e ta b u lu m , g e n i t a l pore u s u a l l y j u s t
a n t e r i o r to a c e t a b u l u m .

Ovary on or to the r i g h t o f the mid­

l i n e , im m e d ia te l y in f r o n t o f the t e s t e s .
in is absent.

Receptaculum sem-

U te r u s v a r y i n g i n e x t e n t , u s u a l l y a n t e r i o r to

o v a r y , i n t r a n s v e r s e c o i l s r e a c h i n g j u s t a n t e r i o r to a c e ta b u lu m .

(99)

V i t e l l a r i a l a t e r a l , s u r r o u n d i n g c r u r a , and o f t e n r e a c h i n g
m i d - l i n e p o s t e r i o r to th e t e s t e s .

L a u re r's canal p re s e n t.

Eggs i n g e n e r a l , r a t h e r l a r g e .
P a r a s i t e s o f b i r d s and mammals.
Genus Schinostoma D i e t z , 1910
Medium s i z e to l a r g e f l u k e s , w ith body e l o n g a t e .

Head

s p i n e s k i d n e y - s h a p e d , a r r a n g e d i n a doub le row, th e a n t e r i o r
row n o t m e e t i n g d o r s a l l y .
spines.

A n t e r i o r end w ith numerous body

B i f u r c a t i o n o f th e i n t e s t i n e im m edia tely in f r o n t

o f acetabulum .

C i r r u s pouch s m a l l , l y i n g f o r th e most p a r t

a n t e r i o r to t h e a c e t a b u l u m .

T e s t e s v a r i a b l e in s h a p e , l o n g e r

th a n w id e , and p la c e d m e d ia n , one behind the o t h e r in the
p o s t e r i o r h a l f o f body.

Ovary s p h e r i c a l or o v a l , median to

somewhat l a t e r a l , a s h o r t d i s t a n c e a n t e r i o r to t e s t e s .
w e l l d e v e l o p e d , numerous c o i l s .

U te r u s

V i t e l l a r i a l a t e r a l , reaching

a n t e r i o r t o , or a l m o s t t o , a c e t a b u l u m , more e x t e n s i v e p o s ­
t e r i o r to t e s t e s , r e a c h i n g p o s t e r i o r end.

Eggs l a r g e , t h i c k

sh elled •
Type s p e c i e s :

Echinostoma revolutum ( F r o e l i c h , 1802)

E c h i n o s toma r e v o lu tu m ( E r o e l i c h , ' 1802)
H o s t:

Anas p l a t y r h y n c h o s , Anas p e n e l o p e , A. q u erq u e-

d u l a , Nyroca f e r i n a , Oidemia n i g r a , and a number o f o t h e r
b i r d s and mammals, i n c l u d i n g man.

(100)

lo cality :

Cosm opolitan

Reference:

7 , 25, 42
Genus Ech ino paryphium D i e t z , 1910

Body s m a l l .

Head s p i n e s k i d n e y - s h a p e d , in a double row,

a n t e r i o r row n o t m e e ti n g d o r s a l l y .
spined.

A n t e r i o r end o f body

Acetabulum on a n t e r i o r b o r d e r o f second body t h i r d .

B i f u r c a t i o n o f i n t e s t i n e s im m edia tely i n f r o n t o f a c e ta b u lu m .
T e s t e s l o n g o v a l , one beh in d th e o t h e r in p o s t e r i o r body
h alf.

C i r r u s pouch d o r s a l to a c e ta b u l u m .

Ovary s p h e r i c a l ,

m edian o r s l i g h t l y l a t e r a l , im m ed ia tely a n t e r i o r to t e s t e s .
U terus s h o r t .

V i t e l l a r i a l a t e r a l , not reaching a n t e r i o r as

f a r as acetabulum .
Type s p e c i e s :

Eggs r e l a t i v e l y l a r g e .
E. e l e g a n s (Looss, 1850)

E ohlnoparyphlum b a c u l u s ( D i e s i n g , 1850)
H o s t:

Mergus a l b e l l u s , Nyroca m a r i l a , Bucephala c l a n g u l a .

S p a t u l a c l y p e a t a , Oidemia f u s c a , Colymbus a r c t i c u s .
L ocality:
Reference:

Europe
25

Ech inoparyphium paraulum ( D i e t z , 1909)
Host:

Anas p l a t y r h y n c h o s , Anas p e n e l o p e , and a l s o

g e e s e , sw ans, and g r e b e s .
L ocality:
R eference:

Europe
1 1 , 25, 121, 122

(101)

Eohinoparyphium r e o u r v a tu m (v L in sto w , 1873)
H ost:

Anas p l a t y r h y n c h o s . Nyroca m a r i l a , N. f u l i g u l a ,

c h i c k e n s and g r e b e s .
L o cality :

Europe

Reference:

9, 33, 76, 77
Genus S te p h a n o p ro r a Odhner, 1910

Synonyms:

M o l i f e r D i e t z . 1909; M e sorchis D i e t z . 1909;

C i r r u s pouch p r e s e n t , n o t r e a c h i n g p o s t e r i o r l y to middle
o f a c e t a b u l u m , c o n t a i n i n g a s h o r t , b u t m uscular c i r r u s .
26 head s p i n e s p r e s e n t .
Type s p e c i e s :

22 -

Pars p r o s t a t i c a p r e s e n t.

_S. d e n t i c u l a t a (H ud o lp h i, 1802)

S t e p h a n o p r o r a s p i n o s a Odhner, 1910
H o s t:

G l a u c i o n e t t a c l a n g u l a a m e r i c a n a , and a number

o f grebes.
L ocality:

E u ro p e , North America

R eference:

25, 97
Genus E i m a s t h l a B i e t z , 1909

Head s p i n e s i n a c o n t i n u o u s row, w ith 29-31 s p i n e s .
V i t e l l a r i a r e a c h i n g from n e a r the p o s t e r i o r end to n e a r
t h e c i r r u s pouch.

A n t e r i o r p o r t i o n o f body o f normal l e n g t h .

Eggs w i t h o u t f i l a m e n t s .
Type s p e c i e s :

H. r h i g e d a n a D i e t z , 1909

(102)

H i m a s t h a l a e l o n g a t a L i n t o n , 1928
H o st:

Oidemia d e g l a n d i

L o cality :

N orth America

Reference:

59
Genua Heteroechinoatomum Odhner, 1910

S i m i l a r to th e a b o v e .
oped.

20

or

22

C i r r u s pouch m o d e r a t e l y d e v e l ­

head s p i n e s .

Type s p e c i e s :

H. mordax (L o o s s, 1899)

Odhner h a s ta k e n a s p e c i e s from Mergus m erganser i n
Sweden f o r which he h a s g i v e n no s p e c i f i c name.
Genus Hypoderaeum D i e t z , 1909
M iddle s i z e d , e l o n g a te d f l u k e s .

Head s p i n e s w e ll

d e v e l o p e d , a r r a n g e d i n a double row, the o u t e r one n o t
m eeting d o r s a l l y .

A n t e r i o r end o f body s p i n e d .

la r g e , near o ral sucker.

Testes elongate, e n t i r e , lying

j u s t b ack o f th e m iddle o f th e body.
p o s t e r i o r to a c e t a b u l u m .

Acetabulum

U teru s lo n g .

C i r r u s pouch r e a c h i n g
V itellaria la te ra l,

r e a c h i n g a n t e r i o r to a c e ta b u lu m .
Type s p e c i e s :

H. conoideum (B loch, 1782)

Hypoderaeum conoideum (B loch, 1782)
H ost:

Anas p l a t y r h y n c h o s , A. q u e r q u e d u l a , Nyroca

m a r i l a , S p a t u l a o l y p e a t a , Mergus m e r g a n s e r , and a l s o from
c h i c k e n s , g e e s e , and g r e b e s .

(103)

L o cality :

Europe

R eference:

25, 75

Family PSIL0ST0MIBAE Odhner, 1913
Body n o t e s p e c i a l l y f l a t t e n e d , and in r e l a t i o n to i t s
shape and d i a m e t e r , v e r y v a r i a b l e i n l e n g t h , v a r y i n g from
.5 to

8

mm.

Skin smooth or o n ly s l i g h t l y s p i n e d .

p h a r y n x , and esophagus p r e s e n t .

B i f u r c a t i o n o f th e i n t e s t i n e

a l i t t l e a n t e r i o r to th e a c e ta b u lu m .
s l i g h t l y subterm inal.

P re p h a r y n x ,

Oral s u c k e r u s u a l l y

The ac eta bulum l a r g e r and more power­

f u l tha n the o r a l s u c k e r , and r a t h e r n e a r i t .

In testin al

c r u r a r e a c h i n g to or n e a r l y to the p o s t e r i o r end.

Excretory

b l a d d e r nY" s h a p e d , th e e x c r e t o r y system c o n s i s t i n g o f a
s u b c u t a n e o u s netw ork of c o l l e c t i n g t u b u l e s and a l a r g e s i n u s
i n the r e g i o n o f th e a c e ta b u lu m .

T e s t e s in p o s t e r i o r h a l f o f

b o d y , one b e h in d th e o t h e r or s l i g h t l y t r a n s v e r s e .

The ovary

l i e s j u s t a n t e r i o r to the t e s t e s , and i s e i t i e r median or
s l i g h t l y to the r i g h t o f th e m i d - l i n e .
P a r a s i t e i n th e i n t e s t i n e o f b i r d s .
G en u s P s i l o s t o m u m L o o s s , 1899
M o re o r l e s s
sm ooth.

elongated,

A cetabulum n e a r

Esophagus s h o r t .

rounded

the a n t e r i o r

p o sterio rly .

Skin

end, very pow erful.

Subcutaneous netw ork o f e x c r e to r y system

(104)

v e r y w e l l deve loped i n a n t e r i o r r e g i o n .

C i r r u s pouch d o r s a l

to a c e t a b u l u m , s e m in a l v e s i c l e l y i n g behind the a c e ta b u lu m ,
and d i v i d e d i n t o two p a r t s .
lo n g and w e l l d e v e l o p e d .
to th e a c e t a b u l u m .

Pars p r o s t a t i c a a b s e n t.

C irrus

Ovary m edian, l y i n g j u s t p o s t e r i o r

U te r u s m o d e r a t e l y l o n g . Female g e n i t a l

pore p r o v id e d w i t h s p h i n c t e r .
v e n t r a l to the c r u r a .

V i t e l l a r i a l y i n g l a t e r a l and

G e n i t a l pore somewhat a n t e r i o r to

acetabulum .
Type s p e c i e s :

Psilostomum b r e v i o o l l e ( C r e p l i n , 182S)

P silostom um b r e v i c o l l e ( C r e p l i n , 1829)
H o s t:

Kaematopus o s t r a l e g u s . Oidemia n i g r a . Clangula

h y e m a li s ( I n t e s t i n e )
L o cality:

Europe

R eference:

1 8 , 99
Genus P silo c h asm u s Luhe, 1909

Very s i m i l a r to th e l a s t g en u s .

M o d e ra tely f l a t t e n e d ,

a n t e r i o r body and esophagus l o n g , p o s t e r i o r end p o i n t e d .
C r u r a and v i t e l l a r i a endin g s h o r t l y b e f o r e the end o f the
bo d y .

E x c r e t o r y system b e g i n n i n g from a s i n u s n e a r the

a c e t a b u l u m , and a cu tan eo u s n e tw o r k .

C i r r u s pouch m u s c u l a r .

Seminal v e s i c l e r e a c h i n g to the p o s t e r i o r end o f the a c e t a b ­
ulum; two p a r t e d .

C i r r u s lo n g and c u r v e d .

V i t e l l a r i a composed

o f numerous s m a ll f o l l i c l e s , s u r r o u n d i n g i n t e s t i n a l c r u r a .

(105)

Type s p e c i e s :

P. oxyurus ( C r e p l i n , 1825)

P s ilo c h a s m u s oxyurus ( C r e p l i n , 1825)
H ost:

Nyroca m a r i l a . N. f u l i g u l a . Oidemia n i g r a .

Tadorna t a d o r n a . B uceph ala c l a n g u l a . C langula hyem ails .
L o cality:

Europe

R eference:

18, 99

P s ilo c h a s m u s l e c i t h o s u s O t t e , 1926
H o s t:

Anas p l a t y r h y n c h o s

L ocality:

L ettland

Reference:

123

P s i l o c h a s m u s l o n g i c i r r a t u s S k r j a b i n , 1913
H ost:

Nyroca ny ro ca

L o cality:

Europe

Reference:

114
Genus P s i l o t r e m a Odhner, 1913

Body r o u n d e d .
fourth.

Acetabulum l y i n g in the second body

S p in e s p r e s e n t , b u t r e a c h i n g a n t e r i o r end o n ly on

th e d o r s a l s i d e .
the l a s t genus.

Esophagus s h o r t .

The c i r r u s pouch s e p a r a t e s the s in u s i n

f r o n t o f th e a c e ta b u lu m .
w all.

C irrus s h o rt.

behind a c e t a b u l u m .
eggs.

E x c r e t o r y system a s in

Seminal v e s i c l e w i t h o u t a thick:

Ovary to r i g h t o f m i d - l i n e , im m ed ia tely
U te r u s v e r y s h o r t w ith no more than 4-5

G e n i t a l pore a t the l e v e l o f the middle to p o s t e r i o r

b o rd e r of pharynx.

(106)

Type s p e c i e s :

P. sim lllim u m (Muhling, 1898)

P s i l o t r e m a sim illi m u m (Muhling, 1898)
H ost:

Nyroca n y ro c a

L ocality:

Europe

Reference:

74,

88,

99

P s i l o t r e m a s p i c u l i g e r u m (Muhling, 1898)
H ost:

Nyroca n y r o c a

L o cality :

Europe

Reference:

88,

99

Genus S p a e r i d i o t r e m a Odhner, 1913
Body s t r o n g l y ro u n d e d .

Skin smooth, o r a l s u c k e r su b ­

t e r m i n a l , pharynx m u s c u l a r .

Acetabulum i n or s l i g h t l y b e ­

hind t h e c e n t e r o f the body, v e r y w e ll d e v e lo p e d , p r o t r u d i n g .
Esophagus a s l o n g a s the ph ary n x .
e r i o r l i m i t s o f the v i t e l l a r i a .

Crura extend to th e p o s t ­
G e n i t a l pore to th e l e f t

o f the m i d - l i n e a t the l e v e l o f the p o s t e r i o r margin o f the
oral

3u c k e r .

C i r r u s pouch c o n t a i n s a se m in a l r e c e p t a c l e ,

p r o s t a t i c c e l l s and a p r o t r u s i b l e c i r r u s .

T estes behind the

a c e t a b u l u m , one beh in d the o t h e r , o r s l i g h t l y o b l i q u e , the
a n t e r i o r one b e i n g s l i g h t l y more d o r s a l .
o v i d , m e d ia n , i n f r o n t o f t e s t e s .

Ovary s p h e r i c a l to

U teru s c o n s i s t s o f a few

(107)

c o i l s , m ostly in f r o n t o f acetabulum .

V i t e l l a r i a surround

i n t e s t i n a l c r u r a , m e e t i n g i n t h e m i d - l i n e d o r s a l l y beh in d
th e t e s t e s .
Type s p e c i e s :

S. g l o b u l u s ( R u d o lp h i, 1819)

S p a e r i d i o t r e m a g l o b u l u s ( R u d o lp h i, 1819)
H o s t:

Nyroca a f f i n i s . N. f u l i g u l a . N. m a r i l a . Anas

a c u t a , Mergus m e r g a n s e r . M.

3e

r r a t o r . C langula h y e m a i l s ,

Cygnus c y g n u s . A lca t o r d a . G l a u c i o n e t t a c l a n g u l a a m e r i c a n a .
L ocality:
Reference:

E urope, E g y p t, North America
1 8 , 65, 99, 106

Family CYATEOCOTYLIDAE Poche, 1925
B ig en ea w i t h d is to m e l i k e body a r ran g m e n t and s t r i g e i d
lik e stru ctu re.

I t d i f f e r s from th e S t r i g e i d a e i n t h a t a

l a r g e w e l l deve loped c i r r u s pouch i s p r e s e n t , which c o n t a i n s
a s e m in a l v e s i c l e , p a r s p r o s t a t i c a and c i r r u s .

The g e n i t a l

p o re i s p o s t e r i o r .
P a r a s i t e s o f r e p t i l e s , b i r d s , and mammals.
Genus C y a t h o c o t y l e Muhling, 1896
C h a r a c t e r s o f the f a m i l y .
and somewhat f l a t t e n e d .
V itellaria la te ra l.

Somewhat l o n g e r than wide

H o l d f a s t organs w ell developed

E x c r e t o r y p o re somewhat s u b - t e r m i n a l ,

o p e n in g on th e d o r s a l s u r f a c e .
Typa s p e c i e s :

p r u s s i c a M uhling, 1896

(108)

C y a t h o c o t y l e p r u s s i c a M u h lin g , 1896
Host:

C l a n g u la h y e m a li s ( I n t e s t i n e )

L o cality :
Reference:

Europe
87

Fam ily STRIGEIDAE R a i l l i e t , 1919
Body u s u a l l y d i v i d e d by a c o n s t r i c t i o n i n t o two
r e g i o n s , an a n t e r i o r b e a r i n g o h i e f l y t h e s p e c i a l o rg an s
f o r a t t a c h m e n t and a p o s t e r i o r u s u a l l y c o n t a i n i n g the
ma^or p o r t i o n o f t h e g e n i t a l i a .

Forebody r e g i o n f l a t t e n e d

and I n c u r v e d v e n t r a l l y or c u p - s h a p e d .

Hind body r e g i o n

more or l e s s c y l i n d r i c a l , o v i d , or c o n i c a l .
t e r m i n a l or s u b t e r m i n a l .
weak.

Oral s u c k e r

Acetabulum u s u a l l y p r e s e n t b u t

S p e c i a l o rg ans o f a t t a c h m e n t p o s t e r i o r to a c e t a b ­

ulum and sometimes c o v e r i n g the l a t t e r .

I n t e s t i n a l crura

w i t h o u t d i v e r t i c u l a , u s u a l l y e x t e n d i n g to p o s t e r i o r end
o f bod y.

G e n i t a l pore u s u a l l y a t p o s t e r i o r end and more

or l e s s d o r s a l .

C i r r u s and c i r r u s pouch u s u a l l y a b s e n t .

U t e r u s w i t h a few c o i l s and c o n t a i n i n g only a few eggs.
P a r a s i t e s o f f i s h , r e p t i l e s , b i r d s and mammals.
Genus Apatemon S z i d a t , 1929
Small S t r i g e i n a e .

V i t e l l a r i a always co n f in e d to

p o s t e r i o r bo dy, where i t s g r e a t e s t l e n g t h l i e s v e n t r a l .

(109)

A t y p i c a l neck b r a n c h a b s e n t .

G e n i t a l cone l i t t l e d i f f e r ­

e n t i a t e d , and composed o f parenchymemous body.

Bursa a

s im p le d e p r e s s i o n w i t h o u t a m u sc u lar s u c k e r - l i k e s t r u c t u r e .
Type s p e c i e s :
Apatemon g r a c i l i s
H ost:

A. g r a c i l i s ( R u d o lp h i, 1819)
( R u d o l p h i , 1819)

Mergus m e r g a n s e r . M. a l b e l l u s . M. s e n a t o r .

Oidemia f u s c a . B uc epha la c l a n g u l a . Oidemia n i g r a . Anas
p l a t y r h y n c h o s . Anser a l b i f r o n s . Oolumba l i v i a
L ocality:
R eference:

(Intestine)

Europe
12, 149, 137
Genu3 O o ty lu r u s S z i d a t , 1929

From s m a l l e r tha n to medium s i z e d S t r i g e i n a e .
c o n f i n e d t o p o s t e r i o r body , which i t f i l l s
end.

B u r s a a sim p le s a c .

V itellaria

to the p o s t e r i o r

A t y p i c a l g e n i t a l cone a o s e n t .

b u l b l i k e s u c k e r around th e o p e n in g .

In a number o f w a te r

b ird s.
Type s p e c i e s :

0. c o r n u t u s ( R u d o lp h i, 1809)

O o t y l u r u s c o r n u t u s ( R u d o lp h i, 1809)
H o s t:

Anas p l a t y r h y n c h o s , Nyroca m a r i l a , geese and

g r e b e s , and o t h e r w a t e r b i r d s .
L o cality :
R eference:
O otylurus

Europe
1 2 , 1«37

erraticu s

( R u d o l p h i , 1809}

A

(110)

H ost:

Mergus a l b e l l u s . Colymbus, a r c t l c u s , Uriga a a l g e .

L o cality :
R eference:

Europe
6 5 , 137

O o t y l u r u s f l a b e l l i f o r m i s ( P a u s t , 1917)
H ost:

Anas p l a t y r h y n c h o s . A. r u b r i p e s . Mareca a m e r ic a n a ,

Querquedula d i s c o r a . N e t t i o n c a r o l i n e n s e . S p a tu la o l y p e a t a .
G l a u c i o n e t t a c l a n g u l a a m e r i c a n a . Nyroca a f f i n l s , N. m a r i l a ,
N. c o l l a r i s . N. a m e r i c a n a , N. v a l i s e n i r a .
L ocality:
R eference:

America (North)
151, p r e s e n t paper
Genus P a r a s t r i g e a S z i d a t , 1927

M idd le s i z e d and l a r g e r S t r i g e i n a e .
i n t o t h e f o r e body.

V i t e l l a r i a extend

I t i s v e r y much l i k e the genus S t r i g e a

b u t has two v e r y pronounced appendages on the f o re b o d y .
Type s p e c i e s :

P , c i n c i n c t a {Brandes, 1888)

P a r a s t r i g e a r o b u s t a S z i d a t , 1929
H o st:

Anas p l a t y r h y n c h o s

L ocality:
R eference:

Europe
137
Family HARMOSfOMIDAE Odhner, 1912
S u bfam ily Harmostominae Loos 3 , 1900

Small to medium s i z e d d i s t o m e s .

Body o v a l , e l o n g a t e .

(Ill)

or s p i n d l e s h a p e , o v a l i n c r o s s s e c t i o n *
smooth*
other.

Skin spine d or

Suckers may be e q u a l , or e i t h e r l a r g e r than the
Acetabulum e i t h e r i n or s l i g h t l y b e f o r e or behind

the m id d le o f the body.

A lim e n tr y t r a c t c o n s i s t i n g o f an

o r a l s u c k e r , p r e p h a r y n x , powerful p harynx, v e r y s h o r t e s o ­
p h ag u s , and i n t e s t i n a l c r u r a which u s u a l l y r e a c h to nea r
the p o s t e r i o r end.

E x c r e t o r y pore nea r th e p o s t e r i o r end

T e s t e s i n p o s t e r i o r body h a l f .
p resent.

O i r r u s pouch w i t h winding c i r r u s p r e s e n t .

l y i n g between th e t e s t e s .
or a b s e n t .

V esicula sem inalis u su a lly
Ovary

Receptaculum s e m in is p r e s e n t

U te r u s lo n g and w inding.

V itellaria la te ra l.

G e n i t a l p o r e m edian, e i t h e r v e n t r a l or d o r s a l a t or nea r
the end o f th e body.

Eggs numerous and

3m a l l .

In b i r d s and mammals.
Genus L euoochloridiom orpha n. g.
See page 58 f o r d e s c r i p t i o n .
Type s p e c i e s :

L euoochloridiom orpha m a c r o c o ty le n. s p .

L e u c o c h lo r id io m o rp h a m a c r o c o ty le n. sp.
H o st:

Anas r u b r i p e s

L ocality:

North America (Michigan)

Family SCHISTOSO&IDAE L o o ss, 1899
Synonyms:

S o h i s to s o m a tid a e

Poche, 1907; B l l h a r z i i d a e

(112)

Odhner, 1918.
Sexes s e p a r a t e , pharynx a b s e n t , esophagus s h o r t , term ­
i n a t i n g p o s t e r i o r l y i n a b i f u r c a t i o n to form i n t e s t i n a l
b r a n c h e s or ca ec a which j o i n c a u d a l l y a t the c a e c a l union
to form a s i n g l e , s l e n d e r i n t e s t i n a l caecum t e r m i n a t i n g
n e a r th e p o s t e r i o r end o f th e body.

Suckers p r e s e n t or

a b s e n t ; a c e t a b u l u m , when p r e s e n t , cephalad o f th e g e n i t a l
pore,

^ody o f male may be widened caudad o f acetabulum and

have th e s i d e s in c u rv e d v e n t r a l l y , form ing a g y n ae co p h o ric
c a n a l i n which th e female l i e s .
more f o l l i c l e s .

T e s t e s c o n s i s t o f f o u r or

O i r r u s pouch p r e s e n t or a b s e n t .

more s l e n d e r th a n the m a le .

Female

Ovary e l o n g a t e , sometimes

s p i r a l l y curved , and l y i n g cephalad o f the c a e c a l u n io n .
L a u r e r ' s c a n a l p r e s e n t or a b s e n t .

V i t e l l a r i a extensive,

e x t e n d i n g from th e d i s t a l p o le o f the ovary to the p o s t e r i o r
end o f t h e body.

P a r a s i t i c in the blood v e s s e l s o f b i r d s

and mammals.
Genus M i o r o b i l h a r z i a P r i c e , 1929
S chistosom inae.

Male l a r g e r than f e m a le .

Gynaecophoric

c a n a l w e l l d e v e lo p e d , commencing i n f r o n t o f the a c e ta b u lu m .
S u ckers p r e s e n t i n b o th s e x e s .
t h a t o f S c h is t o s o m a .

D i g e s t i v e t r a c t s i m i l a r to

T e s t e s 18-20 i n number, a r r a n g e d in

two i r r e g u l a r rows i n a n t e r i o r h a l f o f body.

G e n i t a l pore

(113)

s i t u a t e d a b o u t midway betw een a c e ta b u lu m and the a n t e r i o r
testis.

Female s l e n d e r , a l m o s t c y l i n d r i c a l a n t e r i o r l y ,

flatten ed p o s te rio rly .

Ovary l o o s e l y s p i r a l , s l i g h t l y p r e -

e q u a to r i a l in p o s i t i o n .
egg.

U te r u s l o n g and c o n t a i n i n g a s i n g l e

V i t e l l a r i a occupy a b o u t o n e - h a l f o f body l e n g t h .

Larva

unknown or u n r e c o g n i z e d .
Type s p e c i e s :

M. c h a p i n i P r i c e , 1929

M i o r o b i l h a r z i a c h a p i n i P r i c e , 1929
H o s t:

M arila a f f i n i s

L o cality:
Reference:

N orth America
103
Genus B l l h a r z i e l l a L oo ss, 1899'1'

B ilh arziellin ae.

Both s e x e s w ith the p o s t e r i o r p a r t o f

t h e body d i s t i n c t l y f l a t t e n e d .

Female s h o r t e r tha n the m ale.

I n t e s t i n a l ca ec a u n i t e d p o s t e r i o r l y a t or n e a r e q u a t o r o f
bod y .

Common caecum l o n g , w i t h o u t l a t e r a l b r a n c h e s , and ex­

t e n d i n g i n a z i g z a g manner to p o s t e r i o r end o f body.

Male

g e n i t a l opening s i t u a t e d a t l e f t s i d e o f body a c o n s i d e r a b l e
d i s t a n c e caudad o f a c e ta b u lu m .

C i r r u s pouch p r e s e n t , oon-

1 . S in ce t h i s m a n u s c r i p t was prep ared McLeod ( J o u r . P a r a s i t o l .
2 3 :4 5 6 -4 6 6 ) has d e s c r i b e d a new s p e c i e s o f Schistosome
( P a e u d o b i l h a r z i e l l a q u e r q u e d u la e n . s p . ) from Tuerquedula
d i s c o r s ” i n Ganada'7“ This genus is q u i t e s i m i l a r to B l l h a r z i e l l a .
d i f f e r i n g c h i e f l y i n the r e l a t i v e l e n g t h o f the a n t e r i o r and
p o s t e r i o r body r e g i o n . McLeod would t r a n s f e r B. yokogawaj to
P s e u d o b i l h a r z i e l l a . The genus P s e u d o b l l h a r z i e T l a was d e s c r i b e d
b y Ejsmont in 1929.

(114)

t a i n i n g t h e p r o s t a t e and th e e j a c u l a t o r y d u c t .
v e s i c l e lo n g and f r e e i n t h e parenchyma.

Seminal

T e ste s ab o u t 110

i n number, in p o s t e r i o r p a r t o f body on eaoh s i d e o f the
common caecum.

Female g e n i t a l opening im m ediately p o s t e r i o r

to a c e t a b u l u m .

U t e r u s s h o r t c o n t a i n i n g a s i n g l e egg.

e l l a r i a s i t u a t e d on each s i d e o f th e common caecum.

V it­
Egg

e l o n g a t e d a n t e r i o r l y , e n l a r g e d and provided w ith a sm a ll
spine p o s t e r i o r l y .
Type s p e c i e s :

B. p o l o n l c a (Kowalewski, 1895)

B l l h a r z i e l l a p o l o n i c a (Kowalewski, 1895)
H o st:

Anas p l a t y r h y n c h o s . Querquedula q u e r q u e d u l a .

N e t t i o n c r e c c a . D a f i l a a c u t a , F u l i g u l a f u l i g u l a . Nyroca
l e u c o p h t h a l m a . Ardea c i n e r e a and Cygnus o l o r .
L ocality:
R eference:

Europe and North America
18, 10.3, 136, 137

B i l h a r z i e l l a yokogawal O is o , 1927
H o s t:

Anas p la t y r h y n c h o s

L ocality:
Reference:

Formosa
6

, 103

Genus D e n d r l t o b i l h a r z i e S k r j a b i n and Zakharow, 1920
B ilh arziellin ae.

Body o f b o th sexes e l o n g a t e d .

w i t h o u t s p i n e s or t u b e r c u l e s .

Suckers a b s e n t .

C uticle

D igestive

(115)

system s i m i l a r to t h a t i n B i l h a r z i e l l a , common caecum I o n s ,
z i g z a g , and p r o v id e d w i t h s h o r t , c lu b -s h a p e d or branched
l a t e r a l caeca.

G e n i t a l pore o f male i n a n t e r i o r p a r t of

body and to the l e f t o f th e median l i n e .

T e s t e s numerous

s i t u a t e d on each s i d e o f t h e common caecum and e x t e n d in g
from t h e c a e c a l u n io n to th e p o s t e r i o r end o f body.
s p i r a l and s i t u a t e d between the c a e c a l b r a n c h e s .

Ovary

V itellin e

f o l l i c l e s numerous, s i t u a t e d a l o n g the co u rse o f th e common
caecum.
Type s p e c i e s :

D. p u l v e r u l e n t a (B raun, 1901)

D e n d r i t o b i l h a r z i a p u l v e r u l e n t a (Braun, 1901)
H o s t:

Anas p l a t y r h y n c h o s . Querquedula q u e r q u e d u l a .

L ocality:

A f r i c a , and Europe

Reference:

18, 65, 103
Genus T r i c h o b i l h a r z i a S k r ja b in and Zakarow, 1920

B ilh arziellin ae.
tions;

Body s l e n d e r and d iv id e d i n t o two p o r ­

th e a n t e r i o r wider p o r t i o n s e p a r a t e d from the p o s t e r i o r

t h r e a d l i k e p o r t i o n by a s l i g h t d i l a t i o n .
than acetabulum .

Gynaecophoric c a n a l a b s e n t .

and s e m i n a l v e s i c l e p r e s e n t .
in p o s t e r i o r p o r t i o n o f body.
Type s p e c i e s :

Oral suc ker s m a l l e r
C i r r u s pouch

T e s t e s numerous and s i t u a t e d
Female unknown.

T. k o ssa re w i S k r j a b i n and Zakarow, 1920

(116)

T r l c h o b i l h a r z i a k o s s a r e w i S k r j a b i n and Zakarow, 1920
Host:

Querquedula querq u ed u la

L ocality:

Europe

R eference:

103
Genus G i g a n t o b i l h a r z i a Odhner, 1910

B ilh arziellin ae.
th e m a l e .

Female c y l i n d r i c a l and s h o r t e r than

P o s t e r i o r e x t r e m i t y o f b o th se x es provided w ith

l a t e r a l lo belike p ro jec tio n s.
tu b ercu les.

Suckers a b s e n t .

C u t i c l e w ith o u t s p i n e s or
Gynaecophoric c e n a l reduced to

a s h o r t g r o o v e , s i t u a t e d in a n t e r i o r p a r t o f body.
system s i m i l a r to t h a t o f B i l h a r z i e l l a .

D igestive

T e ste s o r i g i n a t e

caudad o f g y n a e c o p h o r ic c a n a l and extend to p o s t e r i o r end
o f body.

C i r r u s pouch a b s e n t .

G e n i t a l pore s i t u a t e d a t

a n t e r i o r end of g y n a e c o p h o r ic c a n a l and s l i g h t l y to the l e f t
o f the median l i n e .

Ovary m o d e r a te ly long and s p i r a l .

V it­

e l l i n e f o l l i c l e s occupy about n in e t e n t h s o f body l e n g t h .
U t e r u s s h o r t and c o n t a i n i n g a s i n g l e egg.
Type s p e c i e s :

_G. a c o t y l e a Odhner, 1910

G igantob i l h a r z i a monocotylea S z i d a t , 1930
H ost:

Larus r i d l b u n d u s , Anas p la ty rh y n c h o s

L ocality:

Europe

R eference:

139

(117)

I n t h e f o l l o w i n g l i s t s , the tr e m a to d e s t h a t have been
r e c o r d e d from ducks o c c u r r i n g i n North America a r e a rra n g e d
by h o s t .
Anas p l a t y r h y n c h o s p l a ty r h y n c h o s and A. p l a ty r h y n c h o s
( dom estic)
Z y g o c o ty le l unatum

p.

9

, 67

P l a g i o r c h i s p o t a n i n i p. 71
P rosthogonim us a n a t i n u a p . 19, 72
P rosthogonim us h o r i u c h i i p . 73
P ro sthogonim us m a c r o r c h is

p. 14, 73

P rosthogonim us r u d o l p h i i

p. l a ,

Pros thogonimus s k r j a b i n i

p.

Prosthogonim us p e l l u c i d u s
p.

74

O p i s t h o r c h i s geminus

p.

7$

p . 76

O p is th o rc h is sim u lan s

24, 77

p.

M e to r c h is zanthosom us
h ev in se n ie lla

73

p. 84

S chistogonim us r a r u s

Amphimerus e l o n g a t u s

p.

78

pellucida

p.

35

P hilophthalm us a n a tin u s

p.

87,

T y p h l o c o e l u m cymbium

88

p . 39, 92
p . 39, 92

T y p h lo co e lu m cucumerinum
N otoootylus a tte n u a tu s

73

p.

96

N o t o c o t y l u s a e g y p t i a c u s p . 96

(118)

N o t o o o t y l u s s e i n l t a p. 96
N otocotylus urb an en sls
O atatro p is verrucosa

p.
p.

Eohlnoatoma revolutum

47

49,

Eohinoparyphlum paraulum

p.

Echinoparyphium recurvaturn
p.

P silo c h asm u s l e c l t h o s u s
Apatemon g r a c i l i s

96

97

p.

Hypoderaeum conoideum

t

99
100

p.

101

102

p. 105

p. 109

Cotylurus f l a b e l l i f o r m i s
P arastrigea robrsta

53,

p.

110

p. 110

B i l h a r z i e l l a polonica

p.

114

D e n d rito b ilh arzia pulverulenta
G r i g a n t o b i l h a r z i a monocotylea

p. 115
p. 116

Anas r u b r i p e s
Z y g o c o ty le lunatum

p . 9 , 67

Prosthogonim us r u d o lp h ii
Amphim erus e l o n g a t u s
M aritrem a a c a d ia e

P* 2 4 ,

73

77

p . 83

T y p h l o c o e l u m cymblum
E chinostom a re v o lu tu m
O otylurus

p. 18,

p . 3 9 , 92
p.

fla b ellifo rm is

4 9

, 99

p-

53

§ n 0

(119)

L eu c o c h lo r id io m o rp h a maoroco t y l e

p.

59

Mareca arnerioana
Amphimerus e l o n g a t u s

p . 24,

Echinostoma revolutum

p.

C otylurus f l a b e l l i f o r m i s

77

49,

p.

99
53

i 10

Mareca pene lope*
O p i s t h o r o h i s sim ulans

p* 76

N o to c o t y l u s a t t e n u a t u s

p. 96

Echinostoma rev o lu tu m

p. 49, 99

Echinoparyphium paraulum

p. 100

D afila acuta
Typhlocoelum cymbium

p. 39, 92

N o to c o ty lu s a t t e n u a t u s

p. 96

N o to c o t y lu s u r b a n e n s l s

p . 47, 96

S p a e r i d i o t r e m a g lo b u lu s

p. 52, 107

B i l h a r z i e l l a polonica

p. 114

N ettio n carolinense
E chinostom a re v o lu tu m

p.

4 9

t

99

* H o s t s marked th u s a r e n o t n a t i v e North American forms
b u t a r e in c lu d e d h e re b ec au se th e y a r e e i t h e r ta k en
o c c a s i o n a l l y in North America or a r e s u b s p e c i e s o f
th e n a t i v e form.

t m

(120)

Qotylurus f l a b e l l i f o r m l a

p.

53,

110

Querquedula q u e r q u e d u l a *
M e to r c h i s c r a s s i u s o u l u s

p. 78

M e t o r c h i s zanthosomus
N otocotylus s e i n i t a

p. 78
p. 96

Echinostoma rev o lu tu m

p.

49,

Eypoderaeum conoideum

p.

102

B l l h a r z i e l l a polonica

p.

114

99

B e n d rito b ilh a rz ia pulverulenta
T r i c h o b i l h a r z i a Jfcoss a r e w i

p. 115

p . 116

Querquedula d i s o o r s
E u o o t y l e w e h r i p . 94
E chinostom a rev o lu tu m
0otyluru3

p . 49, 99

p.

fla b ellifo rm is

T y p h l o c o e l u m ou cumerimum

531

p . 39, 92

P s e u d O b ilh a r z ie lla querquedulae

Spatula cly p eata
Prosthogonimus

ova tus

Schlstogonimus

rarus

Typhlocoelum oymbium

no

P« 73
p . 74
p . 39 ^ 93

p . 113

(121)

N otocotylus a tte n u a tu s

p . 96

Echinoparyphium ba c u lu a
Kypoderaeum conoideum

p. 100
p. 102

Q o ty lu r u s f l a b e l l i f o r m i s

p« 53, 110

A ix sponsa
N o to c o t y lu s u r b a n e n s l s

p. 47, 96

Nyroca a m erica n a
Typhlocoelum cymbium

p,

Echinoatomum revolutum

p,

Q o ty lu r u s f l a b e l l i f o r m i s
Ampherimus e l o n g a t u s
Z ygocotyle lunatum

39

96
49,

99

p. 53, 110

p. 24, 77
p. 9, 68

Nyroca c o l l a r i s
P r o s thogonimus r u d o l p h i i
Echinostomum revolutum

p . 18, 73
p. 49, 99

Q o ty lu r u s f l a b e l l i f o r m i s

p. 53, 110

Nyroca v a l l s i n e r i a
Ampherimus e l o n g a t u s
E c h i n o s tomum r evolutum

p . 24, 77
p. 49, 99

(122)

Nyroca mar11a
Prosthogonimua r u d o l p h i i
Prosthogonim us ovatua
Tocotrema concavum

p. 18, 73

p. 73

p. 81

Typhlocoelum oucumerinum
Paramonoatomum a lv eatu m

p.

p.

3 9
97

Echinoparyphium r e cu rv atu m

p. 101

Hypoderaeum oonoldeum

p. 102

Psiloohasmua oxyurus

p.

Spaeridiotrem a globulus
O o ty lu ru a c o r n u tu 3

, 92

105

p . 5 2 , 107

p. 109

Q o ty lu r u s f l a b e l l i f o r m i s

p.

55,

110

Nyroca a f f i n i s
Prosthogonim us r u d o lp h ii
Ampherimus e l o n g a t u a
M aritrem a n e t t a e

p* 18, 73

p. 24,

77

p . 3 2 , 84

L e v in a e n ie lla m inuta

p . 85

T y p h l o c o e l u m cymbium

p . 5 9 , 92

S p aerid io trem a glo b u lu s
Q otylurus

fla b ellifo rm is

M io ro b ilh a rz ia ch apini

Nyrooa n y r o c a *

p.

5 3

, 107

p . 5 3 , 110

nz

(123)

P sllo c h a sm p a l o n g i c i r r a t u s
P s i l o t r e m a sim illlm um

p* 105

p. 106

P sllo trem a spiculigerum

p. 106

Nyroca f u l i g u l a *
L ev in aen lella pelluoida

p. 85

Gyoloooelum l a e v i g a t u s

P» 90

Oyolocoelum r o b u a t u a

p. 91

Typhlocoelum oucumerinum
Eohinoparyphium reourvaturn
P silochasm ua oxyurua

3 9 # 92

p.

p. }oi

p. 105

S p a e r i d i o t r e m a g lo b u l u s
B i l h a r a i e l l a p o lo n l c a

p#

53

^ iq7

p. 114

Nyroca f e r i n a *
Gyoloooelum o o e l o n o tu a

p. 90

Echinostoma r e v o lu tu m

p. 49, 99

P rosthogonim us ovatua

p. 73

S la u c io n e tta olangula c la n g u la *
L e v i n a e n i e l l a brachyaoma
Gyoloooelum a r c u a t u a
G atatro p is verrucosa

p. 84

P» 90
p . 97

(124)

gohinoparyphium baoulu3
Psiloohasmua oxyurua
Apatemon g r a c i l i s

p. 100
p.

105

p . iq9

S l a u c i o n e t t a c l a n g u l a am erlca na
Prosthogonim ua r u d o l p h i i

p,

Ampherimus e l o n g a tu a

p . 24,

S p elo trem a pygmaeum

p,

M aritrem a n e t t a e

73
77

3 9 ^ 93

P* 32, 84

Echinostoma r e v o lu tu m

p« 49 , 99

S te p h a n o p ro r a a p in o a a

p. 52,

S p a e r i d i o t r e m a g lo b u l u s

101

p . 52, 107

C lan g u la hy em alis
Prosthogonim us o vatua

p. 73

Toco trema ooncavum

31

Cyclocoelum l a e v i g a t u a

p* 90

Typhlocoelum oucumerlnum
N o to c o t y lu s a t t e n u a t u s
C atatro p is verrucosa

p. 39, 92
p. 96

p. 97

Paramonostomum alveaturn

p. 97

Psilostomum b r e v i c o l l e

p, 104

P siloc hasm us oxyurus

p. 104

(125)

S p a e r i d i o t r e m a g lo b u l u s p. 52, 107
O y ath o o o ty le p r u s s l c a

p . 108

Oidemia d e g l a n d i
H im a s th a la e l o n g a t a

p . 102

Oidemia f u s c a *
Oymnophallus d i p s i l i s p. 69
Tocotrema ooncavum

p. 81

Orohipedum t r a c h e i o o l a

p. 87

Cyclocoelum l a e v i g a t u s

p, 90

Typhlocoelum cuoumerinum

P» 39, 92

Paramonos tomum alveatum
Echinoparyphium b a c u l u s
Apatemon g r a c i l l s

p . 100

p. 109

O i d e m i a n i g r a "'
Qym nophallus d i p s i l i s

p . 69

E e to rch is

p.

zanthosom us

S p e l o t r e m a pygmaeum

p.

L e v l n s e n i e l l a brachy30ma
Cyolocoelum l a e v i g a t u s

78

33,

82

p>

34

p.

90

(126 )

G atatro p is verrucosa

p,

Echinostoma rev olutum

p.

P silo c h a sm u s oxyurus
Apatemon g r a c i l i s

97
49,

p,

99

105

p . 109

Somater l a m o l l i s s l m a
Grymnophallua b u r s i c o l a

p . 69

Gymnophallus c holedocu s

p . 69

O.ymnophallu 3 s o m a t e r i a e

p , 69

Spelotrem a pygmaeum

p . Z 8 t 92

S p e l o p h a l l u s primus

p> gg

Gyclocoelum l a e v i g a t u s

p#

Typhlocoelum cucumerinum
N o to c o t y l u s a t t e n u a t u s
C atatropis verrucosa

90

p. 39, 92
p. 96

p. 97

Paramonostoinum alveatum

p. 97

Mergus merganser
k e to rc h is tener

p . 78

N o to c o t y lu s a t t e n u a t u s

p.

96

Hypoderaeum concoid eum

p#

102

Heteroechlnostomum s p «

p,

102

S p a e r i d i o t r e m a g lo b u lu s
Apatemon g r a c i l i s

p* 109

p# ^2 t 107

(127)

Tocotrema conoavum

p. 81

Mergus s e r r a t o r
M e to rc h is c r a s s i u s c u l u s

p . 78

M e to rc h is zanthosomus p. 78
Tocotrema conoavum

p. 81

Typhlocoelum cucumerinum
O atatropis verrucosa

p. 39, 92

p. 97

S p a e r i d i o t r e m a g lo b u lu s
Apatemon g r a c i l i s

p. 52, 107

p . 109

N e t t i o n oreoca
Prosthogonimus cu n e a tu a
N o t o c o t y lu s a t t e n u a t u s
B l l h a r z i e l l a p o lo n lo a

p. 72
p#

95

p. 114

(128)

SUMMARY and CONCLUSIONS

F i f t e e n s p e c i e s of trem a todes b e l o n g in g to te n d i f ­
f e r e n t f a m i l i e s , a r e r e p o r t e d from ducks in Michigan*
Three new s p e c i e s and one new genus a r e d e s c r i b e d .
Such d a t a a s i t has been p o s s i b l e to o b t a i n w ith
r e g a r d to th e l i f e h i s t o r y , p r e v a l e n c e , p a t h o g e n i c i t y
and h o s t - p a r a s i t e r e l a t i o n s h i p s a r e p r e s e n te d under
each s p e c i e s .
From t h e s e s t u d i e s and the l i t e r a t u r e , i t has been
found t h a t f i v e o f t h e s e s p e c i e s a r e p a t h o g e n ic in ducks
and o n e , n o r m a lly p a r a s i t i c in ducks, i s v e r y s e r i o u s
i f p r e s e n t in c h i c k e n s .
Evidence so f a r i n d i c a t e s l i t t l e or no h o s t s p e c i f i c
i t y among duck tr e m a t o d e s .
By means o f the l i t e r a t u r e , a key to the g e n e r a ,
t h e f a m i l y and g e n e r i c d i a g n o s i s , and l i s t s of the
s p e c i e s o f tr e m a to d e s t h a t have been recorded from
North American d u c k s , a r e p r e s e n t e d . 76 s p e c i e s ,
b e l o n g i n g to 42 gene ra and 17 f a m i l i e s a r e r e p r e s e n t e d .

LITERATURE

A r n s d o r f f , A.

1908. Monostomurn v ic a r iu m n. s p .
Z b l . f . B a k t e r . I . 0 . 4V:362-366
B a r k e r , F. D.

1907. V a r i a t i o n s in the V i t e l l i n e and V i t e l l i n e
D ucts o f Three Distomes o f the Genus O p i s t h o r c h i s .
T r a n s . Am. M ie r. Soc. 27:99-100
1911. The Trematode Genus O p i s t h o r c h i s .
A rc h , de P a r a s i t o l . 14:513-561
1915. P a r a s i t e s o f the American Muskrat ( F ib e r
zibethicus)
J o u r . P a r a s i t o l . 1:184-197
1916. A New Monostome Trematode p a r a s i t i c in the
M uskrat w ith a key to the p a r a s i t e s o f the M u sk rat.
T r a n s . Am. M ic r. Soc. 35:175-184
B a y l i s , H. A.
1929. A Manual o f H elm inthology.
Lond on.

New York and

B e a v e r , P. C.
1937. E x p e r im e n ta l S t u d i e s on Echinostomum
r e v o lu tu m (Froelich)^ a f l u k e from b i r d s and
mammals. 111. B i o l . Monogr. 15 (1) 1-96
B e n n e t t , H. J .
1936. The l i f e h i s t o r y o f Cotylophoron c o ty lo p h o r u m ,
a tr e m a to d e from r u m i n a n t s .
111. B i o l . Monogr. 14 (4) 1-119
B i t t n e r , H.
1925. Ein B e i t r a g zur U bertragung und sur
M orpholog ie von Echinoparyphium r e c u r v a t u m .
B erl.
t i e r a r t z e l . Wschr. 41:82-86
1927. P a r a s i t o l o g i s c h e Beobachtungen.
Z. I n f . k r k h . H a u s t i e r e 30:213-227
B olle

1925. Uber e i n e n Taubentrematoden aus d er
G attung Echinostomum. B t s c h . T i e r a r z t l . Wschr.
3 3 :529-531

(130)

12.

13.
14.

B r a n d e s , G.
1890. Hie f a m il e d e r H olostom id ae.
Z o o l . J b . S y s t . 5:5 49-5 04
1892. R e v is io n d er Monostomiden.
Z b l . f . B a k t e r . I . 0 . 12:504-511
B r a u n , M.
1900. Trematoden d e r C h i r o p t e r a .
Ann. k . k . Hofmus. Wien. 15:217-236

15.

1901. Trematoden der Bursa F a b r i c i i , des
E i l e i t e r s und der E i e r der VBgel. Z b l . f .
B a k t e r . I . 0 . 29:12-19

15.

1901.
E itt.

17.

1901. Zur R e v i s i o n der Trematoden der VBgel
Z b l . f . B a k t e r . I . 0. 29:5 60-568, 895-897, 941-948

18.

1902.
Z ool.

Trematoden d e r C h e l o n i e r .
Z o o l. Ivlus. B e r l i n . 2:1-158

F a s c i o l i d e n d er VBgel
J b . S y s t . 16:1 -1 6 3

19.

C&navan, W. P.
1934. On a trem a tode A llopyge u n d u l a t u s n. s p . ,
p a r a s i t i c in L i l f o r d ' s Crane. P a r a s i t o l .
26 (1) 117-121

20.

C iurea, J .
1924. H e te r o p h y i d e s de l a Feune P a r a s i t a i r e de
Roumanie. P a r a s i t o l . 16 f l ) 1-22

21.

Cobbald , T. S.
1860. Synopsis o f th e B is to m id a e .
P r o c . L in n . Soc. London. 22:363-366

22

.

D a v i e s , E.
1934. On the anatomy o f the Trematode P t a s i g e r
e x a e r e t u s ( E c h in o s to m id a e ) D i e t z , 1909, from the
i n t e s t i n e o f P h a l o c r o c o r a x carbo (C orm ora nt).
P a r a s i t o l . 26 (1) 133-138

23.

Dawes, B.
1936. On a c o l l e c t i o n o f Paramphistomidae from
E a l a y a , w ith a r e v i s i o n o f the genus Paramphistomum.
P arasito l.
28 (3) 330-349

24.

B i e s i n g , C. M.
1836. Monographie der g a t t u n g e n Amphistoma und
B e p l o d i s c u s Ann. Wiener E us. N a tu rg e s c h . 1:235-260

(131)

25.

D i e t z , E.
1910. Die Echinostomiden der Vtfgel.
Z o o l. J b . S uppl. 12:265-512

26.

Edwards, E. E.
1927. On the Anatomy o f the Trematode Paryphostomum
r a d i a t u m , D i e t z , 1909. P a r a s i t o l . 19 (2) 245-260

27.

F a l l i a , A. M.
1934. A Note on some i n t e r m e d i a t e h o s t s o f
Eehinostoma r e v o l u t u m . P ro e . H e l . Soc. Wash. 1 (1) 4-5

28.

F a u s t , E. 0 .
1924. Notes on O r n i t h o b i l h a r z i a odhneri n. s p .
from t h e A s i a t i c Curlew. J o u r . P a r a s i t o l . 11 ( 1 ) 50-55

29.

30.

and N. N i s h i g o r i .
1926. The l i f e c y c l e o f two s p e c i e s o f E e te r o p h y i d a e
p a r a s i t i c in Mammals and B i r d s .
J o u r . P a r a s i t o l . 13 (2) 91-129
Fuhrmann, 0.
1904. Neue Trematoden. Z b l. f . B a k t e r . I .
3 7 :5 8 - 6 4

0.

31 .

1919. Notes h e l m i n t h o l o g i q u e s s u i s s e s .
Rev. s u i s s e de Z o o l. 27:354-376

32.

1928. Trematoden in K u k e n t h a l ’s Handbuck der
Z o o l o g i e . B e r l i n and L e i p z i g .

33.

H a r p e r , W. F.
1929. On th e s t r u c t u r e and l i f e - h i s t o r y of
B r i t i s h F r e s h - w a t e r l a r v a l t r e m a to d e s .
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34.

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S t r i g e a . Z e i t . P a r a s i t e n k d e . 2:583-588

139.

1929. Zur E n t w i c h l u n g s g e s c h i c h t e des B l u t t r e m a t o d e n
der E nten, B i l h a r z i e l l a p o lo n ic a.
Zbl. f . B a k ter.
I l l ; 461-470

140.

1930. G i g a n t o b i l h a r z i e monocotylea n. sp
e i n n e u e r B l u t p a r a s i t aus O s t p r u s s i s c h e n
W a a s e rv o g e l. Z e i t . P a r a s i t e n k d e . 2:583-588

141.

1931. B e i t r a g e zur E n t w ic k l u n g s g e s c h ic h t e
d e r H o lo sto m id en I v . Die c e r c a r i a des E ntenp a r a s i t e n Apatemon g r a c i l i s R u d o lp h i. Z e i t .
P a r a s i t e n k d e . 3:1 60-1 73

(141)

142.

1932. Zur S n t w i e k lu n g s g e s c h ic h t e der C y c lo c o e lid e n
d e r L egenszyklus von T ra c h e o p h ilu s s i s o w i .
Z o o l . Anz. 100-205-213

143.

1933. Uber d i e Entw icklung und den I n f e k t i o n 3modus
o f T r a e h e o p h i l u s s i s o w i , e i n e s L u f t r o h r e n 3 e h m a r o tz e rs
d e r E n ten aus d e r T re m ato d en fam ilie d er Z y k l o z o l i d e n .
T i e r a r t z l . R u n d s ., 3 9 : 9 5 -9 9

144.

T a l b o t , S. B.
1934.

T r a n s . Am, M ic r. Soc.

53 (1) 41.

145.

T a b a n g u i, M. A.
1928. Trematode P a r a s i t e s o f P h i l i p p i n e
V e r t e b r a t e s P h i l . J o u r . S c i . 36 (3) 351-369

146.

V e v e r s , G. M.
1923. O b s e r v a t i o n s on the l i f e - h i s t o r y o f
Eypoderaeum oonoideum (Bloch) and Echinostomum
r e v o lu tu m ( F r o e l i c h ) Trematode p a r a s i t e s o f the
do m e stic duck. Ann. A p l . B i o l . 10:134-136

147.

ViJgel, Hans
1937, Beobachten uber d i e L i e b e n g e s c h i c h t e von
O p i s t h o r c h i s f e l i n e u w i n O s t p r u s s e n . Z b l. f .
B akter. I . 0 .
38:250-254

148.

v . l i n 3 t o w . 0.
1873. E i n i g e neue Bistomum und Berraerkingen
u b e r d i e w e i b l i c h e n Semalorgane der Trematoden.
A r c h , N a t u r g . 39 ( l ) 95-108
1875. Beobachtungen an neuen und bekannten
H e lm in th e n . Arch* N a tu r g . 41 (1) 183-207

149.
150.

1877.

H e lm ln th o lo g i a Arch. N a tu rg ,

43 (1) 1-18

151.

Van C l e v e , K. J . and J . P. M u e lle r
193 2-1934. P a r a s i t e s o f Oneida Lake F i s h e s ,
A n a ls R o o s e v e l t Wild L i f e S t a t i o n Vol. 3,
n o s . 1 , 2, 3 , 4 ,

152.

Van H a i t s m a , J . P.
1930. S t u d i e s on the Trematode Family S t r i g e i d a e
( H o lo s to m id a e ) XXI. L i f e - c y c l e and D e s c r i p t i o n o f
th e C e r c a r i a o f Q o ty lu r u s r a i c h i g a n e n s i s ♦
J o u r . P a r a s i t o l . 16 (4) 230

<142)

153.

154.

1931. S t u d i e s on the Trematode Family S t r i g e i d a e
( H o l o s t o m i d a e ) XXII. C o ty lu ru s f l a b e l l i f o r m i s
( F a u s t ) and i t s l i f e h i s t o r y .
P a p e r s M ic higan Acad. S c i . 8:447-482
W ar|,H . B.
1894. On th e p a r a s i t e s o f Lake f i s h I . Notes
on th e s t r u c t u r e and l i f e h i s t o r y o f Bistoma
opacum n . s p . P r o c . Am. M ic r. Soc. 15:173-182

155.

1895. H e lm in th o lo g i s c h e N otizen
Z b l . f . B a k t e r . I . 0 . 17:304-309

156.

1901. Notes on the P a r a s i t e s o f Lake f i s h .
t h e s t r u c t u r e o f th e c o p u l a t o r y organs i n
M i c r o p h a l l u s n . g. T r a n s . Am. M ic r. Soc.
2 2:175-187

157.

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1918. Trematoda i n F resh-W ater B io lo g y .
John Wiley and Sons. New York.

158.

Werby, E. J .
1928. On th e Trematode Genus Harmostomum w ith
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T r a n s . Am. M ic r . Soc. 47 (1) 68-82

159.

Weseriburg-Lund , C.
1931. C o n t r i b u t i o n s to the development o f the
Trematoda D ig en e a. P a r t I . The b i o l o g y o f
L e u c o c h lo r id iu m paradoxum. 1Viem. Acad. S c i.
L e t t . Denmark. Copenhagen, s e c t . S c i . 9m
s e r . , t . i v , no. 3 , pp. 90-1^2

160.

W i l l e y , C. H.
1933. The Lymph system o f Sygoco^yle lunatum .
P a r a s i t o l . 25 (2) 242-248

161.

W i te n b e r g , G.
1925. V ersuch e i n e r Monographie der Trematoden
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Z o o l . J b . S y s t . 51:167-254

162.

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K o s s a c k , 1911. Z o o l. J b . S y s t. 52:103-186

163 •

1929. S t u d i e s on the Trematode fa m ily F e t e r o p h y i d a e .
Ann. Trop. Med. and P a r a s i t o l . 33 (2) 131-239

On

(143)

164 •

1930. C o r r e c t i o n s to my paper ’’S tu d ie s on the
Trematode f a m ily K e t e r o p h y i d a e ” .
Ann. Mag. N at. H i s t . s e r . 10, 5:412-414

165.

W o l f f h u g e l , K.
1900. B e i t r a g aur Kenntnis der V o g e lh e lm in th e n .
Inaug. B is s . B asel.

166.

Woodhead , A. S.
1935. The Mother ^ p o r o c y s ts o f L eu c o c h lo rid iu m .
J o u r . P a r a s i t o l . 21 (5)

167.

Woodhead , A. E. and H. M a le q itz
1936. Mediogonimus o v i l a c u s n . g. n. s p .
J o u r . P a r a s i t o l . 22 (3) 273-275

168.

W r i g h t , S.
1912. Notes on the anatomy o f the Trematode
M i c r o p h a l l u s o p ac u s, T r a n s . Am. M icro. Soc.
31:167-175

(144)

Index

A m phim erus.. . . . . . . . . . . 2 4 ,

76

e l o n g a t u s ............... ,24,

77

Apa temon

........................

g racilis

100

b a c u l u s ............

108

p a r a u l u m ..................................... 1Q0

............ 109

r e c u r v a t u m ................................ 101

B i l h a r z i e l l a ............................ 113
polonioa.

E c h i n o p a r y p h i u m .................................100

................... 114

y o k a g a w a i........................114
C a t a t r o p i s ................................

96

E c h i n o s t o m a .........................................

99

r e v o l u t u m ......................... . 4 9 , 99

94

E u c o t y l e ......................

94

z a k h a r o v / i .................................

v e r r u c o s a . . . . . ............ 97

v / e h r i ............................................ 94

C o t y l u r u a ................................... 109

G i g a n t o b i l h a r z i a .............................. 116

cornu t u s .......................... 109

monoco t y l e a ..............................116

e r r a t i c u s ....................

109

f l a b e l l i f o r m i s . . . 5 3 ,110
C y a t h o c o t y l e .............................107
p ru ssica.

................... ^ 8

C y c lo c o e l u m ............................. 89

G y m n o p h a l l u 3 .......................................

68

................

69

c h o l e d o c u s ...............................

69

d i p s i l i s . . . . . . ............

69

s o m a t e r i a e ...............................

69

b u rsico la.

*102

a r c u a t u s .............................90

E e t e r o e c h i n o s t^firam a p

c o e l o n o t u s ....................

90

Him as t h a l a .............................................1 0 1

l a e v i g e t u a . . ................ 90

e l o n g a t e ..................................... 102

pseudoraicrostomurn.. . 90
r e b u s t u s . . . ......................91
D en d rito b ilh arzia.

.......... 114

p u l v e r u l e n t a ................. 115

Eypodereum .

.................................. 102

c o n o i d e u m .................................. 1 0 2
L e u c o c h l o r d i o m o r p h a ......................I l l
m a c r o c o t y l e ................. * *59

,-1 1 1

(145)

L e v i n s e n i e l l a ............................ 84

Orohipedum................................. 87

brachya om a....................... 84

t r a o h e i c o l a ....................87

r a i n u t a ..................................... 85

Paramonostomum........................ 97

p e l l u e i d a ............................. 85

a l v e a t u m ...........................97

M a r i t r e m a .................................. 32 ^83

P a r a a t r i g e a .............................110

acad iae.

................... 3 3

r o b u s t a ..........................110

n e t t a e ......................... . 3 2 , 84

P h i l o p h t h a l m u s ........................ 87

M e t o r c h i s ...........................................77

a n a t i n u s ...........................88

c r a s s i n s c u l u s ................... 78

P l a g i o r c h i s ............................... 71

o r i e n t a l i s ...........................78

p o t a n i n i ......................

71

t e n e r ........................................78'

P s e u d o b i l h a r z i e l l a .............113

z a n t h o s o m u s ........................ 78

q u e r q u e d u l a e ............... 113

M i c r o b i l h a r z i a ....................... 112

P rosthogonim us............... 13.. 71

.113

a n a t i n u s ................ 1 9 , 72

.....................95

cunea t u s ...........................72

c h a p i n i .....................
N otocotylus

a e g y p t i a o u s ........................96
atten u atu s

...............*96

hor i u o h i i

...........73

m a c r o r c h i s ............... 14,73

^6

ova t u s ...............................73

u r b a n e n s i s ................* 4 7 , 9 6

r u d o l p h i i ............... ..18,73

s e i n i t a .........................

s k r j a b i n i .........................73

Opia t h o r c h i a ..............................
g e m i n u s ...................................76

p e l l u c i d u s .......................74

76

P s i l o c h a s m u s .......................... 104

Opthalmophagua.........................*92

o x y u r u s .......................... 105

m a s s i n o i ................................93

l e c i t h o s u s ................... 105

a i m u l a n s .............

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lo n g ic irra tu s
P a ilo s to m u m

105

Z y g o o o ty le ..................................... 9, 67

103

lu n a tu m ..................................9, 68

b r e v i o o l l e ................. 104
P s i l o t r e m a .............................105
aim i l l i n i u m . . . . . . . . 1 0 6
s p i c u l i g e r u m .............106
S c h i a t o g o n i m u a ...................... 74
r a r u s ............................... 74
S paer id io t r ema....................106
g l o b u l u s ..............52 , 107 ^
S p e l o p h a l l u s .......................... 85
p r i m u s .............................86
S p e l o t r e m a ............................... 82
pygmaeum................ 58, 82
S t e p h a n o p r o r a ......................101
s p i n o s a ............... 5 2 , 101
T o e o tr e m a ...........................

79

concavum........................ 81
T rich o b ilh arzia. . . . . . . . 1 1 5
k o s s a r e w i ....................116
Typhlocoelura

39 , 91

cuo um erinu m .. . . 3 9 ,92
cymbium

39, 92

P late I

Amphimerus e l o n g a t u a n . a p .
See page 24 f o r d e s c r i p t i o n

E n t i r e worm w i t h v i t e l l a r i a o m itte d *
E x c r e t o r y b l a d d e r added from a e r i a l
section*

jfigure

fig u r e

•

A n t e r i o r t i p o f worm showing mouth
op en in g and p h a r y n x .

f ig u r e

P o s t e r io r t i p o f worm 3howing pos**
t e r i o r v i t e l l a r i a , t e s t e a , o vary,
and 3em inal r e c e p t a c l e .

figure 4.

Acetabulum and g e n i t a l p o r e .

Plate II

f i g u r e 1*

L e u o o o h lo r id io m o rp h a m a o r o o o t y l e
n . g . , n . s p . v e n t r a l v ie w . See
page 59 f o r d e s c r i p t i o n .

f i g u r e S.

M a ritre m a n e t t a e n . s p . v e n t r a l
v ie w .
See page 552 f o r d e s c r i p t ­
ion.