/

THE UNIVERSITY

OF ILLINOIS

LIBRARY

NATURAL HISTORY SURVEY

ViTim

a

books. ^ overdue

U. of^. Library

ILLINOIS BIOLOGICAL
MONOGRAPHS

Vol. V July-October, 1919 Nos. 3 and 4

Editorial Committee

Stephen Alfred Forbes William Trelease

Henry Baldwin Ward

PUBUSHEO UNDER THE

Auspices or the Graduate School by
THE University or Illinois

Copyright, 1920 by the UNivERsrry o» Ilunots
Distributed December 31, 1920.

STUDIES ON MYXOSPORIDIA

A SYNOPSIS OF GENERA AND SPECIES
OF MYXOSPORIDIA

WITH 25 PLATES AND 2 TEXTFIGURES

BY

ROKUSABURO KUDO

Contributions from the

Zoological Laboratory of the University of Illinois

No. 158

TABLE OF CONTENTS

Introduction 7

General Remarks on Recent Observations 8

Myxosporidia Recorded in the Present Paper (List I) 9

Distribution of Myxosporidia 13

A. Geographical Distribution 13

B. Distribution of Myxosporidia in Animals 25

C. Distribution of Myxosporidia in the Organs of the Host 37

D. The Efifect of Environment on the Organal Distribution of Mj^osporidia in

Hosts 44

The Spore 47

Definition of Terms Used for Descriptions 50

Classification of Myxosporidia 52

Descriptions of Genera and Species 60

Suborder Eurysporea '. 60

Family Ceratomyxidae 60

Genus Leptotheca 60

Genus Ceratomyxa 65

Genus Myxoproteus 81

Genus Wardia 82

Genus Mitraspora 84

Suborder Sphaerosporea 86

Family Chloromyxidae 87

Genus Chloromyxum 87

Family Sphaerosporidae 99

Genus Sphaerospora 100

Genus Sinuolinea 104

Suborder Platysporea 106

Family Myxidiidae 106

Genus Myxidium 107

Genus Sphaeromyxa 118

Genus Zschokkella 122

Family Myxosomatidae 123

Genus Myxosoma 123

Genus Lentospora 125

Family Myicobolidae 128

Genus Myxobolus ^>!-^^. 128

Genus Henneguya 158

Genus Hoferellus 173

Myxosporidia Genera et Species Incertae 174

6 ILUNOJS BIOLOGICAL MONOGRAPHS [244

K^s to the Genera and Species of Myxosporidia 178

Key to the Genera of Myxosporidia 178

Key to the Species 179

Summary 196

Appendix: New Myxosporidia from Australia 197

Bibliography 200

General Explanation of Figures 210

Explanation of Plates 212

Index 261

245j STUDIES ON MYXOSPORIDIA—KUDO

INTRODUCTION

Ten years have elapsed since Auerbach (1910) published Die Cnido-
sporidien in which he gave a synopsis of the genera and species of Myxos-
poridia known up to that time. During this period new genera and a
number of new species have been added to the list of this particular group
of parasitic protozoa from the various parts of the world. It is, therefore,
desirable to have a complete monographic work including all the forms
reported up to the present time.

The main objects of the present paper are: 1) to describe a new genus
and a number of new species which have come under the observation of the
writer; 2) to collect all the genera and species recorded by various authors;
3) to propose a new classification by which some of the confusion now
existing may, probably, be avoided; 4) to show the geographical, zoological
and organal distribution in the light of more recent observations; and
5) to present a complete list of the names of the hosts in which Myxospori-
dia occur.

The writer believes himself to be in possession of as complete references
as possible under present conditions. However, he may be unaware of
some works which have not reached him owing to the war.

The Myxosporidia recorded by Labbe (1899) are arranged in almost the
same order as that author listed them, with some slight change such as
placing the type species at the front of each genus or removing a few species
to other genera, while those species which have been described since 1898
are arranged chronologically, no matter whether names are given the
species or not.

Some of the references are omitted, especially when they can be found
in Gurley (1894), Thelohan (1895), Labbe (1899), or Auerbach (1910).
The description of each species is given according to the first observer.
The observations of subsequent investigators are then mentioned in the
second place.

Each species is described according to the following scheme:

1) Specific name

2) Synonyms and literature

3) Habitat, including the locaHty and the date of observation

4) Vegetative form \^

5) Spore ^^"^

6) Remarks

I wish to express my appreciation to Professor Henry B. Ward whose
kindness has made the completion of this paper possible.

ILUNOJS BIOLOGICAL MONOGRAPHS (246

GENERAL REMARKS ON RECENT OBSERVATIONS

The total number of species of Myxosporidia reported up to date and
described in the following pages, excluding 12 ambiguous forms, reaches
237 of which 125 are species which have been observed since 1910,*
The distribution of these new forms is as follows:

Africa.. 6 species

Asia 23 species

Australia 1 species

Europe 31 species

North America 63 species

South America 1 species

Thus, the majority of the species were observed in other lands than
Europe, nearly half being recorded from North American waters. It is not
hard to anticipate from the observations made by Awerinzew, Davis,
Kudo, Mavor, Johnston and Bancroft, and others, that further investiga-
tions on the parasites in the localities where the study of the protozoa
under consideration was neglected, will bring out not only new and inter-
esting forms which will be quite different from the comparatively well
studied European species, but also many important facts that will clear
unknown or doubtful phases concerning the life history and structure of
Myxosporidia.

* Three species are included here which have been described (in Nipponese) by Miyairi'
in 1909.

247]

STUDIES ON MYXOSPORJDIA—KUDO

MYXOSPORIDIA RECORDED IN THE PRESENT PAPER

LIST I

Order MYXOSPORIDIA Butschli

I Suborder EURYSPOREA nom. nov. (see page 56)

I Family CERATOMYXIDAE Doflein

Genus 1 LEPTOTHECA Th61ohan
[15 species]

1) L. agilis Th^lohan (type species)

2) L. elongata Th61ohan

3) L. polymorpha (Th61.) Labbe

4) L. parva Th61ohan

5) L. renicola Th61ohan

6) L. hepseti Th61ohan

7) L. perlata (Gurley) Labb6

8) L. sp. Awerinz«w

9) L. macrospora Auerbach

10) L. informis Auerbach

11) i. longipes Auerbach

12) L. fusiformis Davis

13) L. scissura Davis

14) L. lobosa Davis

15) L. glomerosa Davis

Genus 2 CERATOMYXA Thelohan
[35 species]

1) C. arcuata Th6Iohan (type species)

2) C. sphaertdosa Thelohan

3) C. pallida Th61ohan

4) C. globurifera Thelohan

5) C. appendicidata Thelohan

6) C. truncata Thelohan

7) C. retictdaris Thelohan

8) C. inaequalis Doflein

9) C. linospora Doflein

10) C. ramosa Awerinzew

11) C. drepanopsettae Awerinzew

12) C. tylosuri Awerinzew

13) C. (?) spari Awerinzew

14) C. sp. (?) Awerinzew

15) C. sp. (?) Awerinzew

16) C. acadiensis Mavor

17)

C. sp. Georgevitch

18)

C. coris Georgevitch

19)

C. herouardi Georgevitch

20)

C. mesospora Davis

21)

C. sphairophora Davis

22)

C. taenia Davis

23)

C. atlenuata Davis

24)

C. recurvata Davis

25)

C. lunata Davis

26)

C. abbreviata Davis

27)

C. flagellifera Davis

28)

C. agglomerata Davis

29)

C. anwrpha Davis

30)

C. nwnospora Davis

31)

C. streptospora Davis

32)

C. aggregata Davis

33)

C. undtdata Davis

34)

C. navicularia Davis

35)

C. spinosa Davis

Genus 3 MYXOPROTEUS Doflein

[3 species]

1)

M. ambiguus (Thelohan) Doflein (ty

species)

2)

M. cordifortnis Davis

3)

M. cornutus Davis

Genus 4 WARDIA nov. gen.
[2 species]

1) W. ovinocua nov. spec, (type species)

2) W. ohlmacheri (Gurley) Kudo

Genus 5 MITRASPORA Fujita emend.
Kudo [3 species]

1) M. cyprini Fujita (type species)

2) M. caudata (Parisi) Kudo

3) M. elongata nov. spec.

10

ILLINOIS BIOLOGICAL MONOGRAPHS

[248

II Suborder SPHAEROSPOREA nom. nov. (see page 57)
I FamUy CHLOROMYXIDAE Thfilohan

Genus 1 CHLOROMYXUM Mingazzini

1) C.

2) C.

3) C.

4) C.

5) C.

6) C.

7) C.

8) C.

9) C.
10) C.

[22 species]
leydigi Mingazzini (type species)
caudatum Tli61ohan
quadratutn Th6Iohan
fluviatUe Th61ohan
mucronatutn Gurley
diploxys (Gurley) Th61ohan
protei Joseph
truttae lAgtr
cristatum L^ger
dubiutn Auerbach

11)
12)
13)
14)
15)
16)

17) C.

18) C.

19) C.

20) C.

21) C.

22) C.

sp. Awerinzew
thymaUi Lebzelter
koi Fujita

magnum Awerinzew
fundtdi Hahn
misgurni Kudo
fujitai Kudo
clupeidae Hahn
granulosum Davis
trijugum nov. spec.
catostomi nov. sp>ec.
joarJx nov. spec.

II Fanuly SPHAEROSPORIDAE Davis

Genus 1 SPHAEROSPORA Th^lohan
[10 species]

1) S. divergens Th61ohan (type species)

2) S. elegans Thfilohan

3) S. rostrata Th^lohan

4) 5. masovica Cohn

5) S. platessae Woodcock

6) S. angulata Fujita

7) S. sp. Davis

8) 5. polymorpha Davis

9) S. (?) sp. Southwell et Prashad
10) S. carassii nov. spec.

Genus 2 SINUOLINEA Davis
[5 species]

1) 5. dimorpha Davis (type species)

2) S. capsularis Davis

3) S. arborescens Davis

4) S. opacita Davis

5) S. brachiophora Davis

ni Suborder PLATYSPOREA nom. nov. (see page 57)
I FamUy MYXIDIIDAE Th6lohan

Genus 1 MYXIDIUM BUtschli
[26 species]

1) M. lieberkiihni Btitschli (type species)

2) M, incurvatum Th61ohan

3) M. sphaericum Th61ohan

4) M. histophUum Th^lohan

5) M. ^. Gurley

6) M. danilewskyi Laveran

7) M. giganteum Doflein

8) M. barbatulae C6pfede

9) M. giardi Cepede

10) M. pfdferi Auerbach

11) M. inflatum Auerbach

12) M. bergense Auerbach

13) M. procerum Auerbach

14) M. mackiei Bosanquet

15) M. macrocapstdare Auerbach

16) M. sp. Awerinzew

17) M. depressum Parisi

18) M. oviforme Parisi

19) M. anguillae Ishii

20) M. sp. Mavor

21) M. gadi Georg6vitch

22) M. glutinosum Davis

23) M. phyllium Davis

24) M. striatum Cunha et Fonseca

25) M. kagayamai nov. spec.

26) M. americanum nov, spec.

Genus 2 SPHAEROMYXA Thaohan

[7 species]

1) 5. balbianii Th61ohan (type species)

2) S. immersa (Lutz) Th61ohan

3) S. incurvata Doflein

4) S. sabrazesi Laveran et Mesnil

5) S. hellandi Auerbach

6) S. exneri Awerinzew

7) S. gasterostei Georgevitch

Genus 3 ZSCHOKKELLA Auerbach
[4 spedes]

1) Z. hildae Auerbach (type species)

2) Z. nova Klokacewa

3) Z. acheilognathi Kudo

4) Z. globulosa Davis

249]

STUDIES ON MYXOSPORIDIA—KUDO

If

n Fanuly MYXOSOMATIDAE Poche

Genus 1 MYXOSOMA Thdlohan
[3 species]

1) M. dujardini Th61ohan (type species)

2) M. (?) lobatum Nemeczek

3) M./unduli Kudo

1

2

3

4;

5

6

7

8

9;

10

11

12

13

14;

15

16

17

18

19

20:

21

22

23

24

2^:

26;
27
28
29
30;
31
32
33
34;
35
36
37
38

III Family MYXOBOL

Genus 1 MYXOBOLUS BUtschli
[63 species]
M. mulleri Biitschli (type species)
M. piriformis Thfilohan
M. unicapstdatus Gurley
M. fuhrmanni Auerbach
M. octdi-leucisci Trojan
M. toyamai Kudo
M. notattis Mavor
M. sp. Kudo

M. rohitae Southwell et Prashad
M. seni Southwell et Prashad
M. tnisgurni nov. spec.
M. pfeiferi Th61ohan
M. inaequalis Gurley
M. dispar Th61ohan
M. ellipsoides Th^Iohan
M. exiguus Thelohan
M. oviformis Th61ohan
M. lintoni Gurley
M. globosus Gurley
M. ohlongus Gurley
M. transovalis Gurley
M. obesus Gurley
M. cycloides Gurley
M. sphaeralis Gurley
M. anurus Cohn
M. sp. Gurley
M. sp. Gurley
M. sp. Gurley
M. cyprini Doflein
M. neurohius Schuberg et Schroder
M. aeglefini Auerbach
M. gigas Auerbach
M. volgensis Reuss
M. scardinii Reuss
M. physophilus Reuss
M. tnacrocapsularis Reuss
M. sandrae Reuss
M. bratnae Reuss

39;
40
41
42
43;
44
45
46
47
48
49
50
51
52
53
54

55;

56
57
58;
59
6O:
61
62
63

Genus 2 Ll^SfTOSPDRA Plehn
[6 species]
L. cerebrdis (Hofer) Plehn (type species)
L. midtiplicata Reuss
L. encephalina Mulsow
L. asymmetrica Parisi
L. acuta (Fujita) Kudo
L. dermatobia Ishii

DAE Thelohan

M. cyprinicola Reuss

M. balleri Reuss

M. squamae Keysselitz

M. cordis Keysselitz

M. musculi Keysselitz

M. sp. Miyairi

M. sp. Wegener

M. permagnus Wegener

M. roiundus Nemeczek

M. minutus Nemeczek

M. sp. Lebzelter

M. magnus Awerinzew

M. carassii Klokacewa

M. sp. Southwell

M. funduli Kudo

M. pleuronectidae (Hahn)

M. capsulatus Davis

M. nodularis Southwell et Prashad •

M. hylae Johnston et Bancroft

M. aureatus Ward

M. miyairii nov. spec.

M. lioi nov. spec.

M. orbiculatus nov. spec.

M. discrepans nov. spec.

M. mesentericus nov. spec.

Genus 2 HENNEGUYA Th61ohan
[32 species]

1) H. psorospermica Thelohan (type spe-

cies)

2) H. texta (Cohn) Labb6

3) H. minuta (Cohn) Labb6

4) H. oviperda (Cohn) Labb6

5) H. lobosa (Cohn) Labb6

6) H. peri-intestinalis C6p6de

7) H. media Thelohan

8) H. brevis Th6lohan

9) H. schizura (Gurley) Labb6
10) H. creplini (Gurley) Labb6

12

ILLINOIS BIOLOGICAL MONOGRAPHS

t250

11) H. linearis (Gurley) Labb6

12) H. gurleyi Kudo

13) H. strongylura (Gurley) Labb6

14) H. monura (Gurley) Labb6

15) H. kolesnikovi (Gurley) Labb6

16) H. macrura (Gurley) Th61ohan

17) H. zschokkei (Gurley) Doflein

18) H. sp. (Gurley) Labb6

19) H. sp. (Gurley) Labbe

20) H. tenuis Vaney et Conte

21) H. nusslini Schuberg et Schroder

22) H. legeri C6p6de

23) H. acerinae Schroder

24) H. gigantea Nemeczek

25) E. (?) sp. Nemeczek

26) H. gasterostei Parisi

27) H. neapolitana Parisi

28) H. wisconsinensis Mavor

29) E. brackyura "Ward

30) E. sdminicola Ward

31) H. miyairii nov. spec.

32) E. mictospora nov. spec.

Genus 3 HOFERELLUS Berg
[1 species]

I) H. cyprini Doflein

Appendix: Myxosporidia of unknown genera
and species [1 1 formsl

1) Gen. et spec, incert. Leydig

2) Gen. et spec, incert. Leydig

3) Gen. et spec, incert. Leydig

4) Gen. et spec, incert. Heckel et Kner

5) Gen. et spec, incert. Borne

6) Gen. incert. merlucii Perugia

7) Gen. incert. congri Perugia

8) Gen. et spec, incert. Linton

9) Gen. et spec, incert. Mingazzini
10) Gen. et spec, incert. Nufer

II) Gen. et spec, incert. Mavor
12) Gen. et spec, incert. Mavor

251] STUDIES ON MYXOSPORIDIA— KUDO 13

DISTRIBUTION OF MYXOSPORIDIA

A. GEOGRAPHICAL DISTRIBUTION

As will be seen from List III, Myxosporidia are common parasites of
fish in various parts of the world.

It is interesting to notice that the same species are found among fresh-
water or marine fish from waters in widely separated countries. It is
possible to think that Myxosporidia in marine fish may be carried into
remote waters by the migration of their hosts, while those infecting fresh-
water fish may be brought from one place to another by the transportation
of infected fish for breeding purpose, etc. It should be noted in this con-
nection that no intermediate host has yet been found in relation to myico-
sporidiosis.

The foUowings are the common species found in diflferent localities:

Leptotheca parva Th^l. Marseille, Bergen

Ceratomyxa sphaerulosa Th61. Monaco, Roscoff, Bergen

C. appendiculata Th61. RoscofiE, Marseille, Rovigno

C. drepanopsetlae Awerinzew Murman coast, Bergen, Woods Hole

Chloromyxum leydigi Ming. Roscoff, Monaco, Napoli, Rovigno, Beaufort

C. quadratum Th61. Roscoff, Marseille, Napoli, Beira

Sphaerospora elegans Th61. Bretagne, Karlsruhe, Lago di Garda

5. diver gens Th61. Napoli, Roscoff, Smalfjorden

Myxidium lieberkiihni Biitsch. Lago Maggiore, France, Germany, Lake

Mendota, Georgian Bay

M. incurvatum Th61. Napoli, Monaco, Roscoff, Bergen, Beaufort

M. bergense Auerbach. Bergen, St. Andrews

M. oviforme Parisi Napoli, Norwegian coast

Sphaeromyxa balbianii Th61. Roscoff, Napoli, Beaufort

Myxosoma dujardini Th61. France, Germany, Tokio(?)

On the other hand, some species are limited to certain localities. Five
species classified in the genus Sinuolinea by Davis are reported only from
Beaufort, N. C, U. S. A. The two species of the genus Wardia have been
found solely in the state of Illinois, U. S. A.

More detailed data are shown in the following list.

LIST II

ASIA
I Nippon
Myxosporidia of fresh water fish
1) Northern Part (Hokkaido)

Sapporo : Mitraspora cy print Fujita
Chloromyxum koi Fujita
Sphaerospora angulata Fujita
Lentospora acuta (Fujita) Kudo

14

ILLINOIS BIOLOGICAL MONOGRAPHS

[252

2) Central part (Hondo)

Tokio: MUraspora cy print Fujita

Chlorotnyxum misgurni Kudo
Chlorotnyxum fujitai Kudo
Spkaerospora carassii nov. spec.
Myxidiutn kagayamai nov. spec.
Zsckokkdla acheilognathi Kudo
Myxosoma dujardini (?) Th6lohan
Myxobolus toyamai Kudo
Myxobolus misgurni, nov. spec.
Myxoholus koi nov. spec.
Numazu: Myxidiutn anguiUae Ishii
Lentospora dertnatobia Ishii

3) Southern part (Kiushiu)
Fukuoka: Myxobolus sp. Miyairi

Myxobolus miyairii nov. spec
Henneguya miyairii nov. spec.

Katwan, Mirzapore (UJP.)
Mirpur, Decca district:

Bombay:

n India
A. Myxosporidia of fresh-water fish
Myxobolus sp. Southwell
Myxobolus rohitae Southwell et Prashad
Myxobolus sent Southwell et Prashad
Myxobolus nodularis Southwell et Prashad

B. Myxosporidian of reptiles
Myxidium mackiei Bosanquet

m BUSMA

In the vicinity of Ruby Mines: Spkaerospora sp. Southwell et Prashad

rV Kamtschatka
?Henneguya salminicola Ward

AUSTRALIA
Myxosporidian of amphibia
In the vicinity of Sidney: Myxobolus hylae Johnston et Bancroft

NUe:

1) Indian Ocean
Algoa Bay:
Beira:

East London:
Lorenfo Marques:

AFRICA

A. Myxosporidia of fresh-water fish
Myxobolus unicapsulatus Gurley
Henneguya strongylura Gurley

B. Myxosporidia of marine fish

Chioromyxum magnum Awerinzew
Chloromyxum quadraium Th61ohan
Chioromyxum magnum Awerinzew
Ceratomyxa tylosuri Awerinzew
Ceratomyxa spari Awerinzew
Ceratomyxa sp(?). Awerinzew
Ceratomyxa sp (?). Awerinzew
Sphaeromyxa exneri Awerinzew
2) South Atlantic Ocean

Luderitz Bay: Chioromyxum magnum Awerinzew

253]

STUDIES ON MYXOSPORIDIA—KUDO

15

NORTH AMERICA

Myxobolus globosus Gurley
Myxobolus globosus Gurley
Myxobolus sp. Kudo
Henneguya monura Gurley

Homer Park, HI.:
Salt Fork, Urbana, HI,:

Crystal Lake, Urbana, 111.

I United States
A. Myxosporidia of fresh-water fish

1) From Rivers emptying into Atlantic Ocean

Carlius, Va. (tribt. of Potomac River) : Myxobolus transovalis Gurley
Columbia, S. C. (Santee River) :
Kinston, N. C. (Neuse River) :
West Falmouth, Mass. :
Woodbury, N. J. (Delaware River) :

2) From Lakes and Rivers opening into the Gulf of Mexico

Fox River, trib. Mbsissippi: Myxobolus globosus Gurley
Lake Mendota, Wis.: Myxidium lieberkuhni Btitschli

Henneguya wisconsinensis Mavor et Strasser
NechesRiver, Palestin, Tex.: Henneguya macrura (Gurley) Th61ohan
Storm Lake, la. : Henneguya gurleyi Kudo

Stony Creek, 111. : Chloromyxum trijugutn nov. spec.

Myxobolus orbiculatus nov. spec.

Henneguya mictospora nov. spec.

Chloromyxum trijugum nov. spec.

Myxobolus orbiculatus nov. spec.

Wardia ovinocua nov. gen. nov. spec.

Chloromyxum catostomi nov. spec.

Myxobolus discrepans, nov. spec.

Mitraspora elongate nov. spec.

Myxidium americanum nov. spec.

Myxobolus mesentericus nov. spec.

3) From the rivers opening into the Great Lakes

Black River, Ohio: Gen. et spec, incert. Linton
Put-In-Bay, Ohio: Myxobolus aureatus Ward
Henneguya brachyura Ward

B. Myxosporidia of marine fish (Atlantic Ocean)

Beaufort, N. C. : Leptotheca fusiformis T>a.v\&
Leptotheca scissura Davis
Leptotheca lobosa Davis
Leptotheca glomerosa Davis
Ceratomyxa mesospora Davis
Ceratomyxa sphairophora Davis
Ceratomyxa taenia Davis
Ceratomyxa attenuata Davis
Ceratomyxa recurvata Davis
Ceratomyxa lunata Davis
Ceratomyxa abbreviata Davis
Ceratomyxa flagellifera Davis
Ceratomyxa agglomerata Davfe^ \^^^
Ceratomyxa amorpha Davis
Ceratomyxa m^nospora Davis
Ceratomyxa streptospora Davis
Ceratomyxa aggregata Davis
Ceratomyxa undulata Davis >

16

ILLINOIS BIOLOGICAL MONOGRAPHS

[254

Woods Hole, Mass:

Locality imrecorded:
Sycamore, SL:

Ceratomyxa navictdaria Davis
Ceratomyxa spinosa Davis
Myxoproteus cordiformis Davis
Myxoproteus comutus Davis
Chloromyxum leydigi Mingazzini
Chloromyxutn granulosutn Davis
Sphaerospora polymorpha Davis
Sinudinea dimorpha Davis
Sinuolinea capsularis Davis
Sinuolinea arboresqens Davis
Sinuolinea opacita Davis
Sinuolinea brachiophora Davis
Myxidium incurvaium Th^lohan
Myxidium glutinosum Davis
Myxidium phyttium Davis
Sphaeromyxa balbianii Thfilohan
Zschokkella globulosa Davis
Myxoboliis capsulaius Davis
Ceratomyxa drepanopsettae Awerinzew
Chloromyxum funduli Hahn
Chloromyxum dupeidae Hahn
Myxosoma funduli Kudo
Myxobdus lintoni Gurley
Myxobolus funduli Kudo
Myxobolus pleuronectidae Hahn
Henneguya schizura (Gurley) LabM
C. Myxosporidian of Amphibia
Wardia ohlmacheri (Gurley) Kudo

II Canada
A. Myxosporidia of fresh-water fish
Georgian Bay (south, part) : Myxidium lieberkuhni Biitschli
Myxobolus notatus Mavor

Gen. et spec, incert. Mavor

i

B. Myxosporidia of marine fish (Atlantic Ocean)
Passamaquoddy Bay (at or
near the mouth of St. Croix

River), New Brunswick: Ceratomyxa acadiensis Mavor
Myxidium bergense Auerbach
M. sp. Mavor
Gen. et spec, incert. Mavor

m Alaska
Klutina Lake: Chloromyxum wardi nov. spec.
Stickeen River: Henneguya salminicola Ward

SOUTH AMERICA
A. Myxosporidia of fresh-water fish from the waters connected with Atlantic Ocean
Guiana: Myxobolus inaequalis Gurley
Surinam: Myxobolus inaequalis Gurley
Locality?: Henneguya linearis (Gurley) Labb6

255] STUDIES ON MYXOSPORIDIA—KUDO 17

B. Myxosporidian of marine fish (Atlantic Ocean)
Rio de Janeiro: Myxidium striatum Cunha et Fonseca

C. Myxosporidian of Amphibia
Brazil: Spkaeromyxa immersa (Lutz) Th61ohan

EUROPE
I Italy
A. Myxosporidia of fresh-water fish from lakes and rivers opening into Adriatic Sea
Lago di Como: MUraspora caudata (Parisi) Kudo

Myxidium lieberkiikni BUtschli
Lago di Garda: Sphaerospora degans Thfilohan

Henneguya gasterostei Parisi
Lago di Varamo : Henneguya minuta (Cohn)
Lago Maggiore: Myxidium lieberkiikni Biitschli
Milano: Myxidium lieberkiikni BUtschli

Myxobolus pfeifferi Thfilohan
Pa via: Myxobolus gigas AueThach.

Myxobolus ellipsoides Th^lohan

Henneguya peri-intestinalis C6p6de
Ticino River: Henneguya minuta fCohn)

B. Myxosporidia of marine fish

1) Ligurian Sea

Genova : Chloromyxum leydigi Mingazzini
Gen. incert. merluccii Perugia
Gen. incert. congri Perugia

2) Tyrrhenian Sea

Napoli: Leptotkeca agilis Th61ohan

Leptotkeca elongata Thflohan
Ceratomyxa arcuata Th^Iohan
Ceratomyxa appendiculata Th^lohan
Ceratomyxa truncata Thelohan
Ceratomyxa inaequalis Doflein
Ceratomyxa linospora Doflein
Myxoproteus ambiguus (Th61.) Doflein
Chloromyxum leydigi Mingazzini
Ckoromyxum quadraium Thelohan
Spkaerospora diver gens Thelohan
Myxidium incurvatum Th61ohan
Myxidium giganteum Doflein
Myxidium depressum Parisi
Myxidium oviforme Parisi
Spkaeromyxa balbianii Thdlohan
Spkaeromyxa incurvata Doflein
Spkaeromyxa sabrazesi Laveran et Mesnil
Lentospora asymmetrica Parisi
Myxobolus exiguus Th61ohan
Myxobolus miilleri BUtschli -^„_^^

Henneguya neapolitana Parisi

II Monaco
Mjrxosporidia of fish from Ligurian Sea
Leptotkeca elongata Th61ohan
Ceratomyxa sphaerulosa Thelohan

18 ILUNOJS BIOLOGICAL MONOGRAPHS [256

Ceratomyxa arcuata Th61oban
Ceratotnyota pallida Th61ohan
Ceratomyxa herouardi Georg^vitch
Ceratomyxa sp. Georg6vitch
Chloromyxum leydigi Mingazzini
Myxidium incurvatum Th61ohan
Sphaeromyxa sabrazesi Laveran et Mesnil

in France
A. Myxosporidia of fresh-water fish

1) From Rivers opening into Atlantic Ocean

Aigne: Myxobolus pfeiferi Th61ohan

Bretagne: Sphaerospora elegans Th61ohan

Lorraine: Myxobolus ovtformis Th61ohan

Nancy: Myxobolus pjeiferi Thelohan

Mame: Myxobolus pfeifferi Th61ohan

Seine: Myxobolus pfeijfferi Th61ohan

Paris: Chloromyxum fluviatUe Thilohan

Wimereux: Myxidium giardi C6p^de

2) From Rivers opening into Mediterranean Sea

Dauphin6: Myxobolus miilleri Butschli

Drac River: Myxobolus miilleri Butschli

Myxobolus pfeiferi Thelohan
Grenoble: Chloromyxum cristatum L^ger

Is&re River: Myxidium barbatulae C6pSde

Myxobolus oviformis Thelohan
Myxobolus miilleri Butschli
Myxobolus cycloides Gurley
Henneguya Ugeri C6pede
Lac d'Annecy: Myxobolus miilleri Biitschli
Lac de Paladru: Myxobolus cycloides Gurley
Lac du Bourget: Myxobolus obesus Gurley •

Henneguya peri-intestinalis C6p6de
Lyon?: Henneguya tenuis Vaney et Conte

Rhdne River: Myxobolus pfeiferi Th61ohan
Sa6ne River: Myxobolus pfeiferi Th61ohan

B. Myxosporidia of marine fish
1) From Atlantic Ocean

Arcachon Sphaeromyxa sabrazesi Laveran et Mesnil

Concameau: Ceratomyxa arcuata Th61ohan

Chloromyxum leydigi Mingazzini

Chloromyxum quadratum Th61ohan

Sphaerospora divergens Th61ohan

Myxidium incurvatum Thelohan

Sphaeromyxa balbianii Thelohan
Le Croisic: Leptotheca dongata Thelohan

Leptotheca parva Thelohan

Leptotheca renicola Thelohan

Ceratomyxa appendiculata Th61ohan

Myxoproteus ambiguus (Th61.) Doflein

Sphaerospora rostrata Thelohan

257]

STUDIES ON MYXOSPORIDJA—KUDO

19

CoDcameau: Ceratomyxa arcuataThiloha-n

Chloromyxum leydigi Mingazzini
CUoromyxum quadratum Thfilohan
Sphaerospora divergens Thdlohan
Myxidium incurvatum Th61ohan
Sphaeromyxa balbianii Th61ohan

Roscoff: Cer atomy xa sphaertdosa Th^lohan

Ceratomyxa arcuata Th6Iohan
Ceratomyxa appendiculata Th61ohan
Chloromyxum leydigi Mingazzini
Chloromyxum quadratum Th61ohan
Sphaerospora rostrata Thdlohan
Sphaerospora divergens Th61ohan
Myxidium incurvatum Thdlohan
Myxidium gadi Georg^vitch
Sphaeromyxa balbianii Th61ohan
Sphaeromyxa sabrazesi Laveran et Mesnil
Sphaeromyxa gasterostei Georg6vitch
Myxobolus miilleri Biitschli

Le-Vivier-sur-mer: Leptotheca parva Thdlohan

Myxidium sphaericum Thdlohan
Myxobolus exiguus Th^lohan

St.-Valery-en-caux: Ceratomyxa sphaerulosa Th6Iohan

2) From Mediterranean coast

Marseille: Leptotheca elongata Th^lohan

Leptotheca parva Th^lohan
Leptotheca renicola TWlohan
Leptotheca hepseti Thelohan
Ceratomyxa arcuata Thelohan
Ceratomyxa pallida Thelohan
Ceratomyxa globulifera Th6Iohan
Ceratomyxa appendiculata Th61ohan
Ceratomyxa reticularis Th61ohan
Chloromyxum leydigi Mingazzini
Sphaerospora rostrata Thelohan
Myxidium incurvatum Th61ohan
Myxidium sphaericum Thelohan
Sphaeromyxa balbianii Th61ohan
Myxobolus exiguus Thelohan

Banyuls: Leptotheca elongata Thelohan

Leptotheca polymorpha (Th61.) Labb6
Ceratomyxa arcuata Thelohan
Ceratomyxa globulifera Thelohan
Ceratomyxa appendiculata Thelohan
Ceratomyxa reticularis Th61ohan
Chloromyxum leydigi Mingazzini
Sphaerospora rostrata Thelohan
Myxidium incurvatum Thelohan
Myxidium sphaericum Thelohan
Sphaeromyxa balbianii Thelohan
Myxobolus exiguus Thelohan

20

ILLINOIS BIOLOGICAL MONOGRAPHS

1258

Villefranche: Ceratomyxa pallida Thfilohan

Ceratotnyxa truncata Th61ohan
Ceratomyxa coris Georg6vitch
Sphaeromyxa balbianii Th61ohan

Locality unknown: Leptotheca agilis Th61ohan

Leptotheca perlata (Gurley) Labb6
Myxidium lieberkUhni Biitschli
Myxidiutn histophilutn Th61ohan
Myxosoma dujardini Th61ohan
Myxobolus piriformis Th^lohan
Myxobolus dispar Th61ohan
Myxobolus obesus TWlohan
Henneguya psorospermica Thelohan
Henneguya media Thelohan
Henneguya brevis Thelohan
Hoferellus cyprini Doflein

C. Myxosporidian in a reptile
Myxidium danilewskyi Laveran

IV Germany
A. Myxosporidia of fresh-water fish
1) From Rivers opening into North Sea

Throughout country: Myxobolus cyprini Doflein

Berlin:
Bodensee:

Gutach:

Karlsruhe and its
vicinity:

Leipzig:
Mosel:

Neckar:

Rhine:

Henneguya ovipcrda (Cohn)
Chloromyxum dubium Auerbach
Myxobolus mulleri Biitschli
Myxobolus neurobitis Schuberg et Schroder
Henneguya niisslini Schub. et Schroder
Chloromyxum mucronatum Gurley
Sphaerospora elegans Thdlohan
Myxidium lieberkUhni Biitschli
Myxidium pfeijfferi Auerbach
Myxidium macrocapstdare Auerbach
Henneguya oviperda (Cohn)
Henneguya lobosa (Cohn)
Myxobolus gigas Auerbach
Myxobolus sp. Gurley
Myxobolus pfeiferi Thelohan
Myxobolus squamae KeysseUtz
Myxobolus cordis KeysseUtz
Myxobolus musculi KeysseUtz
Myxobolus exiguus Thelohan (Heidelberg)
Myxobolus mulleri ButschU
Myxobolus pfeiferi Th61ohan
Myxobolus squamae KeysseUtz
Myxobolus cordis KeysseUtz
Myxobolus musculi KeysseUtz
Henneguya psorospermica Th6lohan
Henneguya acerinae Schroder (Heidelberg)
Myxidium lieberkUhni Btitschli
Myxobolus mulleri ButschU
Henneguya psorospermica Th61ohan
Lentospora encephalina Mulsow

259]

STUDIES ON MYXOSPORIDIA—KUDO

21

2) From Rivers opening into Baltic Sea

AUe:
Frisches Haff:

Kurisches Haff :

Masurische Seen:

Pregel:

Weichsel:
3) Localities unknown:

Myxobolus midleri BUtschli
Myxidium lieberkuhni Butschli
Myxosoma dujardini Th61ohan
Myxobolus piriformis Thdlohan
Myxobolus dispar Th61ohan
Myxobolus exiguus Thdlohan
Myxobolus ovifortnis Th61ohan
Myxobolus tniilleri Butschli
Myxobolus cycloides Gurley
Myxobolus anurus Cohn
Myxobolus sp. Wegener
Myxobolus permagnus Wegener
Henneguya psorospermica Thelohan
Henneguya texta (Cohn)
Henneguya minuta (Cohn)
Henneguya lobosa (Cohn)
Henneguya crepUni (Gurley)
Myxosoma dujardini Thelohan
Myxobolus exiguus Thelohan
Myxobolus ovifortnis Thelohan
Myxobolus cycloides Gurley
Henneguya psorospermica Th61ohan
Henneguya texla (Cohn)
Henneguya creplini (Gurley) Labb6
Sphaerospora masovica Cohn
Myxobolus dispar Th61ohan
Myxobolus ellipsoides Thelohan
Myxobolus cycloides Gurley
Myxobolus anurus Cohn
Henneguya psorospermica Th61ohan
Henneguya texta (Cohn)
Myxosoma dujardini Thelohan
Myxobolus piriformis Thelohan
Myxobolus dispar Thelohan
Myxobolus exiguus Thelohan
Myxobolus oviformis Thdlohan
Myxobolus miilleri Butschli
Myxobolus cycloides Gurley
Myxobolus anurus Cohn
Myxobolus permagnus Wegener
Henneguya psorospermica Thelohan
Henneguya texta (Cohn)
Henneguya minuta (Cohn)
Henneguya lobosa (Cohn)
Henneguya creplini (Gurley) Labb6
Myxobolus cyprini DoSein

Chloromyxum leydigi Mingazzini
Myxidium lieberkiihni Butschli
Myxidium sp. Gurley
Lentospora cerebralis (Hofer) Plehn

22

ILUNOIS BIOLOGICAL MONOGRAPHS

(260

Helder:

Henneguya sckizura (Gurley) Labb6
HojereUus cyprini Doflein
Gen. et spec, incert. Leydig
Gen. et spec, incert. Leydig
Gen. et spec, incert. Leydig
Gen. et spec, incert. Bome

V Netheuland
Myxosporidian of marine fish
CMoromyxum quadratum Th61ohan

VI England
M3rxosporidia of marine fish
Firth of Qyde, More-

camb, etc.: Myxobolus aeglefini Auerbach

Liverpool (?) : Sphaerospora platessae Woodcock

Abelvaer:

Bergen:

Bergsfjord:

Boadsfjord:
Bodd:

Finkongkjeilen:

GronSy:

Hammerfest:

Harstad:

Honnigsvaag:

Kabelvaag:

Vn Norway
Mjrxosporidia of marine fish
Myxidium bergense Auerbach
Myxidium ovijorme Parisi
Zschokkelia hildae Auerbach
Myxobolus aeglefini Auerbach
Leptotheca parva Th^lohan
Leptotheca macrospora Auerbach
Leptotheca informis Auerbach
Leptotheca longipes Auerbach
Ceratomyxa sphaerulosa Thelohan
Myxidium incurvatum Thfilohan
Myxidium inflatum Auerbach
Myxidium bergense Auerbach
Myxidium procerum Auerbach
Sphaeromyxa heUandi Auerbach
Zschokkelia hildae Auerbach
Myxobolus aeglefini Auerbach
Myxidium bergense Auerbach
Zschokkelia hildae Auerbach
Zschokkelia hildae Auerbach
Myxidium bergense Auerbach
Zschokkelia hildae Auerbach
Myxidium bergense Auerbach
Zschokkelia hildae Auerbach
Myxidium bergense Auerbach
Zschokkelia hildae Auerbach
Myxidium bergense Auerbach
Myxidium ovijorme Parisi
Zschokkelia hildae Auerbach
Myxidium bergense Auerbach
Zschokkelia hildae Auerbach
Myocidium bergense Auerbach
Zschokkelia hildae Auerbach
Ceratomyxa drepanopsettae Awerinzew

261]

STUDIES ON MYXOSPORIDIA—KUDO

23

Myxidium bergense Auerbach
Zschokkella hildae Auerbach
Zschokkdla hildae Auerbach
Myxidium bergense Auerbach
Myxidium bergense Auerbach
LepMheca parva Thdlohan
Lepiotheca macrospora Auerbach
Myxidium oviforme Parisi
Zschokkella hildae Auerbach
Myxobolus aeglefini Auerbach
Myxidium bergense Auerbach
Zschokkella hildae Auerbach
Zschokkella hildae Auerbach
Myxidium bergense Auerbach
Myxidium bergense Auerbach
Zschokkella hildae Auerbach
Ceratomyxa drepanopsettae Awermzew
Myxidium bergense Auerbach
Myxidium oviform^ Parisi
Zschokkella hildae Auerbach
Zschokkella hildae Auerbach
Myxidium bergense Auerbach
Sphaerospora divergens Thdlohan
Zschokkella hildae Auerbach
Leptotheca parva Thelohan
Myxidium bergense Auerbach
Myxidium bergense Auerbach
Zschokkella hildae Auerbach
Leptotheca informis Th61ohan
Ceratomyxa drepanopsettae Awerinzew
Myxidium bergense Auerbach
Sphaeromyxa hellandi Auerbach
Myxidium bergense Auerbach
Myxidium oviforme Parisi
Zschokkella hildae Auerbach
Zschokkella hildae Auerbach
Myxidium bergense Auerbach
Myxidium oviforme Parisi
Zschokkella hildae Auerbach
Myxobolus aeglefini Auerbach

VIII Switzerland
Myxosporidia of fresh-water fish
1) From Lakes connected with North Sea

Neuchatel: Myxobolus fuhrmanni Auerbach

Myxobolus miUleri Biitschli
Henneguya oviperda (Cohn)
Henneguya zschokkei (Gurley) Doflein
Thun: Henneguya zschokkei (Gurley) Doflein

Zurich: Henneguya zschokkei (Gurley) Doflein

Lucerne: Myxosoma dujardini Th61ohan

Myxobolus ellipsoides Thelohan

Kiberg:
Kirkenes:
Eliistiana:
Kristiansand:

Lodingen:
Makur:

Mosjoen:

Nusfjord:

RSrvik:

Rossfjord:

SkjervS:

Skjotningsberg:

Smalfjorden:

Stavanger:

Svolvaer:

TjamS:

Torghatten:
Trondhjem:

Vikholmen:
Vardo:

24

ILUNOJS BIOLOGICAL MONOGRAPHS

[262

Myxobolus ovifortnis Th61ohan
Myxobolus midleri Biitschli
Henneguya psorospermka Th^lohan
Henneguya texta (Cohn)
Henneguya zschokkei (Gurley) Doflein
Gen. et spec, incert. Nufer

Wallen: Henneguya zschokkei (Gurley) Doflein

2) From Lake connected with Mediterranean Sea

Geneva: Myxobolus sphaeralis Gurley

Henneguya zschokkei (Gurley) Doflein

IX AtJSTKIA

A. Myxosporidia of fresh-water fish

1) From Wvers opening into Black Sea

Danube tributaries Chloromyxum thynwlli Lebzelter
and Neusiedler: Myxosotna (?) lobatum Nemeczek
Myxobolus aeglefini Auerbach
Myxobolus cyprini Doflein
Myxobolus rotundus Nemeczek
Myxobolus minutus Nemeczek
Myxobolus sp. Lebzelter
Henneguya acerinae Schroder
Henneguya gigantea Nemeczek

2) From Rivers opening into North Sea

Prag:

Krakau:

Rovigno:

Locality unknown:

Vienna:

Pergrad (Danube):

Volga (to Caspian
Sea):

Myxosoma dujardini Th61ohan
Myxobolus ellipsoides Th61ohan
Myxobolus oculi-leucisci Trojan
Myxobolus cyprini Doflein
Myxosporidia of marine fish (Adriatic Sea)
Leptotheca agilis Thelohan
Ceratontyxa pallida Th61ohan
Ceralomyxa appendiculata Th61ohan
Myxoproteus ambiguus (Th^l.) Doflein
Chloromyxum leydigi Mingazzini
Sphaeromyxa sabrazesi Laveran et Mesnil
Gen. et spec, incert. Heckel et Kner
C. Myxosporidian of Amphibia
Chloromyxum protei Joseph

X Serbia
Henneguya gigantea Nemeczek

XI Russia

A. Myxosporidia of fresh-water fish

Lentospora multiplicata Reuss
Myxobolus volgensis Reuss
Myxobolus scardinii Reuss
Myxobolus physophilus Reuss
Myxobolus macrocapsularis Reuss
Myxobolus sandrae Reuss
Myxobolus bramae Reuss
Myxobolus cyprinicola Reuss
Myxobolus balleri Reuss

263]

Don (to Black Sea) :
Locality unknown :

STUDIES ON MYXOSPORJDI A—KUDO

Myxobolus sp. Gurley
Zschokkella nova Klokacewa
Myxobolus tnagnus Awerinzew
Myxobolus carassii Klokacewa
Henneguya kolesnikovi (Gurley) Labb6

25

B. Myxosporidia of marine fish from Arctic Ocean
Murman coast: Ceratomyxa ramosa Awerinzew

Ceratomyxa drepatwpsettae Awerinzew
Myxidium sp. Awerinzew
Leptotheca sp. Awerinzew
Chloromyxum sp. Awerinzew

B. DISTRIBUTION OF MYXOSPORIDIA IN ANIMALS

The number of host species that harbor Myxosporidia is 237, as will be
seen from List III.

Tho two incompletely studied forms are found in Annelida and Insecta,
Myxosporidia are the parasites of Vertebrata, especially of Pisces, only few
being found infecting Amphibia and Reptilia. They are distributed among
these groups of animals as follows:

Number of host species

Annelida 1

Insecta 1

Pisces 223

Amphibia 8

Reptilia 4

Gurley (1894:101-105), Wasielewsky (1896:132-148), Labbe (1899:
133-161) and Auerbach (1910:36-45; 1911:471-494) gave lists in which
they recorded the names of host species. Wasielewsky arranged the names
alphabetically while others listed them according to their systematic order.
In the following pages, the writer followed Wasielewsky, i.e., the names of
the host species are arranged alphabetically as is supposed to be more
convenient in referring to the host than any form presented otherwise.

LIST III. LIST OF HOST SPECIES

Host

Organ Infected

Myxosporidian

Locality

Annelida
Nais lacustris {N. probo-
scidea)

Unknown

Abdominal cav-
ity

Branchiae

Myxobolus sp.

Chloromyxum diploxys

Myxobolus balleri
bramae

Germany

Insecta

Tortrix viridana L. (imago)

Pisces

Abramis batterus L

France
Russia

A. brama L

«

26

ILUNOIS BIOLOGICAL MONOGRAPHS

[264

Host

Organ Infected

Myxosporidian

Locality

Branchiae

Myxobolus cycloides

Germany

i<

ellipsoides

"(?)

«

exiguus

France

(1

ovifonnis

France

(C

rotundus

Austria

Gall-bladder

Sphaerospora masovica

Germany

Kidney

Myxobolus cyprini

«
t

Hungary

Subcut. conn.

gigas

Germany,

tissue of oper-

Italy

culum

A. vimba L

Branchiae

cycloides

Germany

«

ellipsoides

"(?)

«

oviformis

Germany

Acanthias acantkias L

Gall-bladder

Chloromyxum leydigi

France

A. blainvillei

(1

magnum
Henneguya acerinae

Africa

Acerina cernua L

Branchiae

Germany

" .Muscle

crepiini

((

Conn, tissue of

tenuis

France

aliment, canal

Eye

Myxobolus magnus

Russia

Muscle

Leptotheca perlata

France?

Acheilognathus lanceolatum

Gall-bladder,

Temm. et Schl

Gall-duct

Zschokkella acheilognathi
Myxobolus cycloides

Nippon
Germany

Alburnus alburnus L

Branchiae

(A. lucidus Heck)

((

dispar

«

«

ellipsoides

Germany ?

It

oviformis

France

" , kidney

obesus

(1

Muscle and

dispar

f(

spleen

Eye

miilleri

Switzerland

Alosafinta Cuv. var. lacus-

tris Fatio

Kidney

Base of spines of
2nd dorsal fin

Mitraspora caudata

Italy

Ameiurus tnelas Raf

Henneguya gurleyi

U. S. A.

Ancyhpsetta quadrocdlata

GUI

Gall-bladder

Ceratomyxa undulata
Myxidium anguiUae

«

AnguUla japonica Temm.

Integument

Nippon

etSchL

«

Lentospora dermatobia
Myxidium giardi

«

A. vulgaris Flemm

Kidney
Subcutaneous

France

Apkredoderus sayanus Gill.

intermusc. tiss.

Henneguya monura

U. S. A.

Apogon rex-mtdlorum Cuv.

GaU-bladder

Myxidium oviforme

Italy

Argentina situs Nilss

«

procerum
Chloromyxum quadratum

Norway

Ariodes polystaphylodon

Muscle

Africa

Aspius rapax Ag

Branchiae

Myxosoma{7) lobatum
Myxobolus miilleri
Henneguya acerinae

Austria

Aspro asper L

((

France

A. zingd Cuv

«

«

2651

STUDIES ON MYXOSPORIDIA—KUDO

27

Host

Organ Infected

Myxosporidian

LocaUty

Atherina hepsetus L

Gall-bladder

Leptotheca hepseH
Myxoproteus cornutus
Myxidium striatum

France

Bairdiella chrysura

Urin. bladder

U. S. A.

B. rouchus Cuv. et Val

GaU-bladder

Brazil

Barbus barbus L

Kidney, spleen,
intestine, ovary,

(5. fluviatUis)

Myxobolus pfeifferi

France,

etc.

Germany

Inner surface of

squamae

Germany

scale

Muscle of ven-

cordis

<f

tricle

Muscle, kidney

muscuH

«

B. plebejus Val

((

pfeifferi
miiUeri

Italy

Germany

Burma

B. vulgaris Flem

Branchiae

Barilius barnc ;..

Under scales

Sphaerospora sp.
Myxidium sphaericum
Myxidium sphaericum
Sphaeromyxa incurvata
Chloromyxum quadratum
Myxidium incurvatum
Sphaerospora divergens

Belone acus Risso

GaU-bladder

it

France

B. belone L

«

Blennius ocellatus

«

Italy
«

B. gattorugine Brunn

Kidney
Gall-bladder

B. pholis L

France

Renal tubules

«

Blicca bjdrkna L

Branchiae

Myxobolus cydoides
ellipsoides

Germany
"?

«

«

macrocapsularis

Russia

«

oviformis

Germany

Box boops L

Gall-bladder .

Ceratomyxa pallida

France,

Italy

B. salpa L

Gall-bladder

Ceratomyxa herouardi
pallida

Monaco

((

France,

Italy

Kidney

Henneguya neapolitana

Italy

BrevoorUa tyrannus Latr....

Muscle

Chloromyxum clupeidae

U. S. A.

Brosmius brosme Ascanius..

GaU-bladder

Leptotheca longipes

Norway

«

Sphaeromyxa hellandi

({

CaUionymus lyra L

«

Myxidium incurvatum

France,

Norway

Muscle

Chloromyxum quadratum

France

Carassius auratus L

Branchiae

Lentospora acuta
Sphaerospora angulata

Nippon
«

Kidney

«

Mitraspora cyprini

<(

Subcutaneous

tiss. of head

Henneguya miyairii

«

(C. carassius L.)

Body cavity
Branchiae

Myxobolus sp.

dispar

Germany
«

u

Sphaerospora carassii

Nippon

(C. vulgaris L.)

Body cavity,
Uver, intestine

Myxobolus carassii

Russia

GaU-bladder

Zschokkella nova

((

Carcharhinus limbatus

(<

Chloromyxum leydigi

U.S.A.

C. sp

«

Ceratomyxa flagellifera

«

28

ILLINOIS BIOLOGICAL MONOGRAPHS

[266

Host

Organ Infected

Myxosporidian

Locality

Carpiodes difformis

Branchiae

Myxobolus discrepans

U. S. A.

Catostomus comtnersonii

Lac

Gall-bladder

Chloromyxum catostomi
Sphaeromyxa hellandi

«

Centronotus gundlus

((

Norway

Cepola rubescens L

((

balbianii

France

Cestracion tiburo

«

Chloromyxum leydigi

U. S. A.

(1

Ceratomyxa mesospora

a

C. zygaena

((

Ceratomyxa mesospora

it

((

recurvata

it

«

Chloromyxum leydigi

ti

(i

Leptoiheca fusiformis

it

Chaetodipterus faber

Gall-bladder

Ceratomyxa streptospora

It

Urin. bladder

Myxoproteus cordiformis

It

Chondrostoma nasus L

Branchiae

Myxobolus exiguus

Germany

"

Gen. et spec, incert.

Switzerland

Tongue

Gen. et spec, incert.

?

Citharus linguataGthi

Gall-bladder

Myxidium depressum

Italy

Clupea harengus Young

((

Ceratomyxa sphaerulosa

Norway

Muscle

Chloromyxum dupeidae

U. S. A.

C. pilchardus Walb.

{Alosa sardina)

Gall-bladder

Ceratomyxa truncata
Sphaeromyxa balbianii

France, Italy
Italy

it

Cohitis barbatula L

Kidney
Urin. bladder

Myxidium barbatulae
Henneguya legeri

France

«

C. fossilis L

Branchiae,

Conger conger L.

kidney, spleen

Myxobolus piriformis

Germany

{Leptocepkalus c.)

Gall-bladder

Gen. incert. congri

Italy

Coregonus lavaretus L.

(C.fera)

Branchiae

Henneguya sp.
Myxobolus sphaeralis

France?

\^^ 'J ^' "/

" (mucosa)

Switzerland

Muscle

Henneguya kolesnikovi

Russia

«

zschokkei

Switzerland

C. exiguus albellus

" , branchia

it

«

C. wartmanni nobilis

« «

tt

«

Coris giofredi Risso

Gall-bladder

Ceratomyxa coris
it

France

C.julisL

((

tt

Muscle

Chloromyxum quadratum

ti

Gall-bladder

Myxidium oviforme

Italy

CoUus gobis L

Branchiae

Myxobolus mulleri
Myxidium sp.
Ceratomyxa inaequaiis

France

C. scorpitis

«

Russia

Crenilabrus mediterraneus..

"

Italy

C. tnelops L '

Eye

Myxobolus miiUeri

Germany

J w

France

Gall-bladder

Ceratomyxa arcuata

France

Kidney

Sphaerospora diver gens

K

C. pavo Cuv. et Val

Gall-bladder

Ceratomyxa inaequaiis
Lentospora asymmetrica

Italy

Kidney

Italy

«

Sphaerospora diver gens

France, Italy

Cydopterus lutnpus L

Gall-bladder

Myxidium infiatum

Norway

267]

STUDIES ON MYXOSPORIDIA—KUDO

29

Host

Organ Infected

Myxosporidian

Locality

Cyitoscion regalis

Gall-bladder

Myxidium glutinosum

U. S. A.

Urin. bladder,

ureters

Sinuolinea dimorpha

«

CypHnodon variegatus

Subcutaneous

tissue

Myxobolus lintoni

((

Visceral conn.

tissue

capsulatus

«

CypHnus carpioli

Branchiae

cyprinicola

Russia

((

dispar

France,
Germany

<(

oviformis

France ?

<c

toyamai

Nippon

tl

Myxosoma dujardini

France

((

Myxobolus koi

Nippon

Brain

Lentospora encephalina

Germany

Gall-bladder

Chloromyxum koi

Nippon

Kidney

Hoferellus cyprini

Germany,
France

" liver, spl.

Myxobolus cyprini

Germany,
Himgary

Kidney

Milraspora cyprini

Nippon

«

Spkaerospora angulata

«

C. (Rasbora) daniconius

Subcutaneous

Day

intermuscular

tissue

Myxobolus sp.

India

Muscles

Myxobolus nodularis

India

Dasybatis hastatus

Gall-bladder

Chloromyxum leydigi
Leptotheca scissura

U. S. A.

((

«

D. sabina

«

Chloromyxum leydigi

«

Drepanopsetta platessoides

Fabr

(1

Ceratomyxa drepanopsettae

Russia

Erimyzon sucetta oblongus

Lac {Catostomus

Branchiae

Myxobolus globosus

U. S. A.

tuberctdatus)

Integument

oblongus

i(

Esox lucius L....

Branchiae

anurus

Germany

((

Henneguya lobosa

«

<(

Henneguya psorospermica

France,
Germany

Intestinal wall

Henneguya peri-inkstinalis

France,
Italy

Eye muscle, etc.

Henneguya sckizura

U. S. A.

Ovary

Henneguya oviperda

Germany,
Switzerland

Urin. bladder

Myxidium lieberkUhni

France, Ita-
ly, Canada,
U. S. A.,
Germany

Fundidus sp

Muscle

Chloromyxum funduli

U. S. A.

30

ILLINOIS BIOLOGICAL MONOGRAPHS

[268

Host

Organ Infected

Myxosporidian

Locality

P. diaphanus..
F. keteroditus..

F. majalis..

Gadus aeglefinis L.,
G. caUarias L

G. tsmarkii Nilss..

G.merlangus'L..

G. morrhuaJj....
G. poUackius L..
G. virens L

Galeocerda tigrinus

Galois galeus L. (G. canis).
Gambusia affinis

GasterosUus aculeatus.

G. pungitius L..

G. spinachia

Gobio gobio'L.
(G. fiuviatUis).

Gobius fluviatUis L

Gobius paganeUus L

Hdiases ckromis Gthr..

Hippocampus brevirostris
Cuv.

Muscle

Branchiae,

Muscle

Branchiae
«

" , muscle
Gall-bladder
Cartilage
Urin. bladder
Cartilage
Gall-bladder
Urin. bladder
Cartilage, bone

of cranium,

eye

Gall-bladder

Cartilage

Cartilage

Gall-bladder
«

Urin. bladder
Gall-bladder

Kidney (r. t.),
ovary

Kidney
" (r. t), ovary

Gall-bladder

Fin

Fin, spleen,
kidney, liver
Branchiae
Body-cavity
Gall-bladder

Myxoholus funduli

Myxobolus funduli
Myxosoma funduli
Myxosoma funduli
Myxobolus funduli
Myxidium incurvatum
Myxobolus aeglefini
Zschokkella kildae
Myxobolus aeglefini
Myxidium oviforme
Zschokkella hUdae

Myxobolus aeglefini

Leptotheca informis
Myxobolus aeglefini
Myxobolus aeglefini
Myxidium gadi

bergense
Zschokkella hildae
Ceratomyxa lunata

sphaerulosa
Myxidium incurvatum

phyUium

Henneguya media

brevis
Sphaerospora elegans
Henneguya gasterostei

brevis

media
Sphaerospora elegans
Sphaeromyxa gasterostei

Myxobolus miilleri

oviformis

cycloides

Gen. et. spec, incert.

Ceratomyxa arcuata

Ceratomyxa arcuata

Myxidium incurvatum
Sphaeromyxa sabrazesi

U.S. A.

Germany

Norway

Germany

Norway

Norway

Germany,

England

Norway

Germany

Germany

France

Norway
«

U. S. A.
France
U. S. A.

France

«

" , Italy
Italy
France

Germany

France

Germany
((

Italy
Italy,
Monaco

Italy
France,

Italy,

Himgary

269]

STUDIES ON MYXOSPORIDJA—KUDO

31

Host

Organ Infected

Myxosf>oridian

Locality

H. guUulatus Cuv

Gall-bladder

Sphaeromyxa sabrazesi

Fr., Hun.,

Hippoglossoides litnan-

Monaco

doides

Urin. bladder

Sphaerospora diver gens

Norway

Hippoglossus vulgaris

Flenun

Gall-bladder

Ceratomyxa drepanopsettae

Norway
Russia

((

ramosa

Hybognathus nuchalis Ag....

Conn, tissue of

the head

Henneguya macrura

U. S. A.

Idus tndanotus Jleck

Branchiae

Myxobolus eUipsoides

Germany?

Muscle

Leniospora multiplicata

Russia

Labeo rohita

Branchiae

Myxobolus rohilae

India

Fins

Myxobolus seni

«

Ldbeo niloticus Forsk

?

Myxobolus unicapsulatus

Egypt

Lahrus turdus

GaU-bladder

Ceratomyxa linospora

Italy
U. S. A.

Leiostomus xanthurus

i<

aggregata

Lepisosleus platyslomiis

Urinary bladder

Sphaerospora sp.

«

Lepomts cyanellus Raf

Mesentery

Myxobolus mesentericus

«

Urin. bladder

Henneguya mictospora

M

Kidney

Mitraspora elongata

<(

L. humilis Girard

Ovary

Wardia ovinocua

l<

Urinary bladd.

Henneguya mictospora

«

L. tnegalotis Raf

Gall-bladder

Chloromyxum trijugum

«

Leuciscus cephdus {Squalis

Air-bladder,

France,

cephalus) ,

branchiae

Myxobolus miilleri

Germany

Branchiae

ellipsoides

« «

»

Gall-bladder

Chloromyxum fluviatile

France

L. lucius L

Branchiae

Myxobolus sp.

Germany?

L. phoxinus L. {Phoxinus

Conn. tiss. of

laevis Ag.)

kidney; ovary

Myxidium histophilum
Myxobolus miilleri

France

<<

«

L. rutilus L

Branchiae

ellipsoides

Germany?

<i

miilleri

c<

Myxosoma dujardini

France

Conn. tiss. un-

der the mouth

muc. mem-

brane

Myxobolus fukrmanni

Switzerland

Opercle, pseu-

do branchiae,

France,

kidney

cydoides

Germany

Vitreous body

of eye

oculi-leucisci

Austria

?

Henneguya sp.

Germany

Heart

Gen. et spec, incert.

((

Leuciscus sp

Branchiae

Myxobolus minutus

Austria

<(

Myxosoma (?) lobatum

«

Lophius budegassa Spin

GaU-bladder

Ceratomyxa appendiculata

Italy,
France

L. piscatorius Li

((

Ceratomyxa appendiculata

<i

32

ILUNOJS BIOLOGICAL MONOGRAPHS

(270

Host

Organ Infected

Mjrxosporidian

Locality

L. piscatorius L

Urin. bladder

Myxoproteus atnbiguus

France,

Italy,

'

Hungary

Lota lota L. (L. vulgaris)

Branchiae

Myxobdus midUri

Germany

«

Myxoholus cydoides

Germany

Gall-bladder

CUoromyxum dubium

" , Austria

Urin. bladder,

kidney

mucronatum

France

Urin. bladder

Myxidium lieberkiihni

France,
Germany

((

Sphaerospora degans

Germany

Lucioperca lucioperca L

Branchiae

Henneguya acerinae

" , Austria

(L. Sandra Cuv.)

" , head, fin,

circle

Myxoholus sp.

?

<i

Henneguya gigantea

Aus., Servia

«

Gen. et sp. incert.

Austria

Muscle

Myxoholus sandrae

Russia

L. vdgensis Pall

Branchiae, cor-

nea, dorsal fin

volgensis

«

Mdanogrammus aeglefinis..

GaU-bladder

Myxidium hergense

Canada

Menticirrkus americanus L.

<(

striatum

BrazU

Merluccius merluccius L.

(M. vulgaris)

«

Ceratomyxa glohulifera
Leptotheca dongata

France

V'*** • ■'^^^i*' »*•/• •••••••••••••••••••

«

" , Italy

i<

Gen. incert. merluccii

Italy

Micropogon undulatus

«

Ceratomyxa aggregata

U.S.A.

Micropterus salmoides Lac .

Urin. Bladder

Henneguya mictospora

U. S. A.

Misgurnus anguiUicau-

datus Cantor

Branchiae

Myxoholus ^.
CUoromyxum fujitai

Nippon

«

GaU-bladder

«

misgumi

«

«

Myxidium kagayamai

«

«

Myxoholus misgumi

c<

Molva vulgaris Flem

Bone

aeglefini
Leptotheca informis

Austria

GaU-bladder

Norway

"

Sphaeromyxa hellandi

«

MottUa maculata Risso

«

baJbianii

France

M. tricirrata BL

«

Ceratomyxa arcuata
Leptotheca dongata

<<

<i

", Monaco

«

Sphaeromyxa halbianii

France

«

sabrasesi

Monaco

Mugil auratus Risso

Intestine, stom-

ach, spleen,

pyloric coeciun

Myxoholus miiUeri

Italy

Kidney

exiguus

" , France

M. capita Cuv „

«

exiguus

<(

M. cephalus L

GaU-bladder

Myxidium incurvatum

U. S. A.

M. cheh Cuv„

Stomach,spleen,

Italy,

kidney, etc.

Myxoholus exiguus

France

2711

STUDIES ON MYXOSPORIDIA—KUDO

33

Host

Organ Infected

M3Ttosporidian

Locality

M.sp

Kidney

Sphaerospora rostrata

Italy, France
Monaco

• i3p

Muraena sp

Gall-bladder

Ceratomyxa sp.

Mustelus cams Mitch.

{M. vulgaris)

Gall-bladder

Ceratomyxa sphaerulosa
Myxidium incurvalum
Ckloromyxutn quadratum

France

Nerophis aequoreus L

Gall-bladder

France

Muscle

«

N. annulatus

Gall-bladder

Myxidium incurvalum

«

«

Sphaeromyxa sabrazesi

Monaco

N. lumbricifortnis

«

Myxidium incurvalum
Gen. et spec, incert.

France

Notropis megalops Raf

Subcut. tissue

U. S. A.

N. gilberti J. et M

Muscle

Myxobolus orbicttlatus
aurealus

U.S.A.

N, blennius

«
Fins

«

N. anogenus

C(

«

Henneguya brachyura

((

Oncorkynchus keta

Under the skin

Henneguya salminicola
«

Kamtschatk a

0, kisulch

Connective tiss.

«

of body muscle

«

Alaska

0. nerka

Gall-bladder

Chloromyxum wardi

Ceratomyxa arcuata
Sphaerospora polymorpha
Ceratomyxa arcuata

«

Ophidium vasalli

Gall-bladder

Monaco

Opsanus tau

Urin. bladder

U. S. A.

Pagellus centrodontus Del....

Gall-bladder

France,

Italy

Paralichthys albiguUus

J. et G

Urin. bladder

navicularia

U. S. A.

^ ^

«

spinosa

«

«

Leptotheca glomerosa

4(

«

Sinuolinea brachiophora

((

K

capsularis

«

(1

opacita

(1

P. dentatus

Gall-bladder

Ceratomyxa drepanopsettae
navicularia

«

Urin. bladder

(1

<<

Leptotheca lobosa

C<

et

Sinuolinea capsularis

«

Parasilurus asotus L

Intestinal wall

Myxobolus miyairii
Ceratomyxa monospora
Myxobolus sp.
Henneguya wisconsinensis

Nippon
U. S. A.

Peprilus alepidotus

Gall-bladder

Perca flavescens

Spleen
Urin. bladder

(i

«

P. fluviatilis

Branchiae

■ texta

Italy,
Germany

«

Henneguya minuta

«

«

Myxobolus sp.

Germany

" , operculum

permagnus

«

((

Myxosoma dujardini

Switzerland

«

Henneguya psorospermica

«

Phoxinus (Clinostomus)

Under scales on

fundidoides Girard

ext. surf.

Myxobolus transovalis

U. S. A.

P.laems

Branchiae

muJleri

France

34

ILUNOIS BIOLOGICAL MONOGRAPHS

[272

Host

Organ Infected

Myxosporidian

Locality

P. laevis

Kidney, ovary
Urin. bladder
Urin. bladder

Gall-bladder

Membrane lin-
ing branchial
cavity

Gall-bladder
?

Branchiae

Gall-bladder
<(

Otic-capsule
Muscle

i(

GaU-bladder

Subcutaneous
muscL tissue

Gall-bladder

«

«
«

Gall-duct
Gall-bladder

«

«

«

Branchiae
Cartilage, pe-
richondrium

Branchiae
«

«

GaU-bladder

Muscle, spleen

Air-bladder
«

Gall-bladder

Myxidium histophUum
Sphaerospora degans
ZschokkeUa hildae

Leptotheca polymorpha

Henneguya linearis

Myxobolus notaius

inaequalis

Henneguya linearis

? Ceraiomyxa drepanopsettae
«

Sphaerospora platessae
Chloromyxum dupeidae
dupeidae
dupeidae

Ceraiomyxa acadiensis
Myxidium sp.

Myxobolus pleuroneciidae

Chloromyxum leydigi
Leptotheca scissura
Myxidium giganteum
Chloromyxum leydigi
Myxidium sp.
Chloromyxum sp.

Chloromyxum leydigi
Chloromyxum leydigi

Chloromyxum leydigi

Ceratomyxa sp. (?)
Myxobolus cydoides

Lentospora cerebralis

Myxobolus cydoides
scardinii
Myxosoma dujardini

Myxidium macrocapsulare

Myxobolus dispar

permagnus
physophilus

Ceratomyxa sp. (?)

France

Phycis blennioides Br

P. mediterraneus (P.
i>hvcis L.)

Germany
Norway

France

Pimelodus sebae Cuv. et
Val

Pimepkales notatus Raf

Piramutana blochi C. et V.

Platystoma fasciatum L

Pleuronectes flesus L

S. America
Canada

S. America

«

Norway
«

P. platessa L

Pomolobus aestivalis

England
U. S. A.

P. mediocris Mitch

II

P. pseudoharengus Yoxmg..
Pseudopleuronedes amer-
icanus

II

Canada

Pteroplatea madura
Le Sueur

(1

U.S.A.
<i

Raja asterias

II

Italy
France

R. bails L

R. radiaia

Germany ?
Murman

R, davata L

Coast
France

R. undulaia Lac

II

Rkina squatina L

France,

RMnobaihus sp. (?) Awer....
Rhodeus amarus BL

Germany
Africa
Germany

Salmo foniinalis Mitch

Scardinius erythrophihal-
mus

II
II

Scatophagus argus

Russia

France,

Germany
<i

France
Germany
Russia
Africa

2731

STUDIES ON MYXOSPORIDJA—KUDO

35

Host

Organ Infected

Myxosporidian

Locality

Scoliodon terrae-novae

Gall-bladder

Ceratomyxa abbreviata

U. S. A.

((

attenuata

((

«

sphairophora

((

«

taenia

((

«

Chloromyxutn leydigi

«

Sconiber scotnbrus L

«

Leptotheca parva

France,

Norway

Kidney

renicola

France

?

Gen. et sp. inc.

Germany?

Scorpaena porcus L

Gall-bladder

Ceratomyxa arcuata

France

S. scrofd L

«

Ceratomyxa arcuata

«

«

Myxidium incurvatum

«

S. sp

«

Leptotheca agilis

", Germany
Monaco

Scylliwn canicula

((

Ceratomyxa sphaerulosa

<(

Chloromyxum leydigi

" , France
Germany

S. dsterids

«

a

Italy

Norway

Fastem

Sebastes dactylopterus

((

Leptotheca macrospora
Leptotheca sp.

5. norvegicus

«

Finmark

5. viviparus H. Kr

«

Leptotheca macrospora
Spkaeromyxa sabrazesi

Norway
Monaco

Siphonostoma rondeletii

«

Siphostoma floridae

«

balbianii

U. S. A.

Urin. bladder

Sinuolinea arborescens

«

S. louisianae

Gall-bladder

Spkaeromyxa balbianii
Myxidium gadi
Ceratomyxa (?) spari
navicularia

«

Solea vulgaris

«

France

Spcrus berda

i(

Africa

Sphaeroides tnaculatus

Urin. bladder

U. S. A.

((

Sinuolinea capsularis

((

<(

Zschokkella globulosa

«

Spinax spinax L

GaU-bladder

Chloromyxum leydigi

Italy,
France

Squalis agassizii Heck

Branchiae

Myxobolus miilleri

France

S. a. Savigny Bona-

parte

«

Myxobolus miilleri
Chloromyxum clupeidae

((

Stenotomus ckrysops L

Muscle

U. S. A.

Urin. bladder

Gen. et sp. incert.

Canada

Syngnathus acus L

Gall-bladder

Myxidium incurvatum
Spkaeromyxa sabrazesi

France

«

Italy,

Monaco

Muscle

Chloromyxum quadratum

France

S.typMe

GaU-bladder

Myxidium incurvatum

France

Synodontis schaU Bl. Schn..

Integum. of

cephalic reg.

Henneguya strongylura

Egypt

?

Myxobolus inaequalis

S. America

Synodus faetans

Gall-bladder

Ceratomyxa agglomerata
amorpha

U. S. A.

((

«

Tautogolabrus adspersus

Walb

Muscle

Chloromyxum clupeidae

((

36

JLUNOJS BIOLOGICAL MONOGRAPHS

(274

Host

Organ Infected

Myxosporidian

Locality

ThymaUus thytnallus L

Gall-bladder

Chloromyxum thymaUi

Austria

((

Myxobolus sp.

«

Neurilemma (?)

pfeifferi

Germany ?

Tkysanophris japonicus

Gall-bladder

Sphaeromyxa exneri

Africa

Tinea tinea L. {T. vulgaris).

Branchiae
Air-bladder,

Myxobolus piriformis

France,
Germany

kidney, etc.

eUipsoidcs

(1

GaU-bladder

Chloromyxum cristatum

France

<(

Myxidium pfeiferi

Germany

Kidney

Myxobolus cy print

Germany,
Hungary

Torpedo narce Risso

Gall-bladder

Chloromyxum leydigi
Chloromyxum leydigi
Chloromyxum leydigi
"Ceratomyxa reticularis
Myxidium incurvatum

France

T. oceUata

((

Germany

T. torpedo L

((

Trachinus draco IL

«

«

((

«

Trackurus trachurus L

Muscle

Chlordmyxum quadrcUum

France,
Germany

Trutta fario L

Gall-bladder,

Gall-duct

Chloromyxum truttae

France

Nervous syst.

Myxobolus neurobius

Germany

Subcutaneous

conn. tiss. at

"•

base of fin

Henneguya niisslini

«

T. iridea Gibb

Cartilage, peri-
chondrium

Lentospora eerebralis

«

T. solar L

«
Gall-bladder

' cc

Myxidium oviforme

«

Norway

Trygon pastinaca L

Gall-bladder

Chloromyxum leydigi
Leptotheca agilis

France

" , Italy

Tylosurus inarianus

Urin. bladder

Chloromyxum granulosum
Ceratomyxa tylosuri

U. S. A.

T. scMsmatorhynckus

Gall-bladder

Africa

Urophycis ckuss

«

acadiensis

Canada

Zoarees angularis

«

acadiensis

«

Amohibia -•

Kidney
Gall-bladder

Wardia ohlmacheri

Bufo lentiginosus

U. S. A.

B. marinus L

Sphaeromyxa immersa
Myxobolus hylae
Sphaeromyxa immersa

Brazil

Hyla aurea

Testis, ovary
GaU-bladder

Australia

Leptodactylus oceUaius

Brazil

Molge cristata Laur.

(Tritonc.) ;

«

Chloromyxum caudatum

France

Proteus anguineus L

Kidney
«

Chloromyxum protei
?Wardia ohlmacheri
?Wardia ohlmacheri

Austria

Rana esctdenta

France?

R. temporaria (R.fusca)

*

Reptilia

Emys orbicularis L.

{Cistudo europaea)

Kidney

Myxidium danilewskyi

Russia,

France

2751

STUDIES ON MYXOSPORJDIA—KUDO

37

Host

Organ Infected

Myxosporidian

Locality

Lacerta sp

Ovarian egg
Kidney

Gen. et spec, incert.

Myxidiutn mackiei

americanum

Italy
India

Trionyx (Amyda) ganzeti-
cus

T. spinifera

U. S. A.

C. DISTRIBUTION OF MYXOSPORIDIA IN THE ORGANS OF THE HOST

Altho some species are found in various organs of the host animal, the
majority has one or two particular seats of infection. Among the various
organs which become infected, the gall-bladder is most frequently infected.
The kidney, branchia and urinary bladder have less chances of being
parasitized. As to the infection of the reproductive organs of the host,
little is known. The male reproductive organ becomes rarely infected,
being reported only twice. The female reproductive organ, however, is
more frequently infected. The infection of the next generation of the host
animal thru the infected ovum which is known to occur in some Micro-
sporidian parasites, has not been reported in Myxosporidia as yet.

LIST IV. ORGANS OF HOST INFECTED BY MYXOSPORIDIA

I. Pisces

1) Integument. — Sphaerospora sp. Southwell et Prashad (under the scales)

Myxobolus seni Southwell et Prashad (fin)

Myxobolus transovdis Gurley (under the scales)

Myxobolus unicapsulatus Gurley (head)

Myxobolus cycloides Gurley (opercle)

Myxobolus inaequalis Gurley (head)

Myxobolus sp. Gurley (opercle, head, fin)

Myxobolus squamae Keysselitz (inner surface of the scales)

Myxobolus volgensis Reuss (fin)

Myxobolus pertnagnus Wegener (operculum)

Myxobolus aureatus Ward (fin)

Henneguya brachyura Ward (fin-ray)

Henneguya linearis (Gurley) Labb^ (membrane lining branchial cavity)

Henneguya gurleyi Kudo (base of spines of dorsal fin)

Henneguya strongylura (Gurley) Labb6 (cephalic region)

Lentospora dermatobia Ishii

2) Connective tissue. — Myxidiutn anguillae Ishii (subcutaneous)

Myxobolus fuhrmanni Auerbach (under the oral mucous membrane)

Myxobolus oviformis Th61ohan (subcutaneous)

Myxobolus lintoni Gurley (subcutaneous)

Myxobolus oblongus Gurley (chiefly of the head)

Myxobolus gigas Auerbach (of operculum, sides and fins)

Myxobolus capsulatus Davis (visceral)

Henneguya kolesnikovi (Gurley) Labb6 (interstitial)

Henneguya niisslini Schuberg et Schroder (at the base of dorsal fin)

Henneguya miyairii Kudo (of the head)

Gen. et sp. incert. Linton (subcutaneous)

38 ILLINOIS BIOLOGICAL MONOGRAPHS [276

3) Muscle. — Leptolheca perlata (Gurley) Labb£

Chloromyxum quadratum Th^lohan

Chloromyxum fttnduli Hahn

Chloromyxum clupeidae Hahn

Lentospora multiplicata Reuss

Myxoboltis notatus Mavor (connective tissue of voluntary muscle)

Myxoholus pfeifferi Tli6lohan

Myxobolus sandrae Reuss

Myxoholus musculi Keysselitz

Myxobolus funduli Kudo

Myxobolus pleuronectidae Hahn

Myxobolus sp. Southwell (subcutaneous intermuscular tissue)

Myxobolus nodularis Southwell et Prashad

Myxobolus orbiculatus Kudo

Henneguya creplini (Gurley) Labb€

Henneguya monura (Gurley) Labbfi

Henneguya zschokkei (Gurley) Doflein

Henneguya salminicola Ward

Gen. et spec, incert. Leydig

4) Eye. — Sphaerospora platessae Woodcock (optic capsule)

Myxobolus oculi4eucisci Trojan (vitreous body)

Myxobolus ellipsoides Thflohan

Myxobolus miiUeri Biitschli

Myxobolus aeglefini Auerbach

Myxobolus volgensis Reuss

Myxobolus magnus Awerinzew

Henneguya sckizura (Gurley) Labb€ (intercellular tissue of eye muscle)

5) Branchiae. — Sphaerospora carassii Kudo

Myxosoma dujardini Th61ohan

Myxosoma (?) lobatum Nemeczek

Myxosoma funduli Kudo

Lentospora acuta (Fujita)

Myxobolus piriformis Th^lohan

Myxobolus toyamai Kudo

Myxobolus rohitae Southwell et Prashad

Myxobolus dispar Thfilohan

Myxobolus ellipsoides Th^lohan

Myxobolus exiguus Thdlohan

Myxobolus oviformis Thfilohan

Myxobolus miilleri Biitschli

Myxobolus globosuS Gurley

Myxobolus cycloides Gurley (also pseudobranchiae)

Myxobolus sphaeralis Gurley

Myxobolus anurus Cohn

Myxobolus sp. Gurley

Myxobolus gigas Auerbach

Myxoboltis volgensis Reuss

Myxobolus scardinii Reuss

Myxobolus macrocapsidaris Reuss

Myxobolus bramae Reuss

Myxobolus cyprinicola Reuss

277] STUDIES ON MYXOSPORIDIA— KUDO 39

Myxobolus balleri Reuss
Myxobolus sp. Miyairi
Myxobolus sp. Wegener •

Myxobolus perntagnus Wegener
Myxobolus rotundus Nemeczek
Myxobolus minutus Nemeczek
Myxobolus funduli Kudo
Myxobolus koi Kudo
Myxobolus discrepans Kudo
Henneguya psorospermica Th6lohan
Henneguya texla (Cohn)
Henneguya minuta (Cohn)
Henneguya lobosa (Cohn)
Henneguya creplini (Gurley) Labb6
Henneguya linearis (Gurley) Labb6
Henneguya acerinae Schroder
Henneguya gigantea Nemeczek
Gen. et spec, incert. Heckel et Kner

6) Heart. — Myxobolus cordis Keysselitz (muscle of ventricle and bulbus arteriosus)

Gen. et spec, incert. Leydig (auriculo-ventricular valve)

7) Air bladder. — Myxobolus ellipsoides Th^lohan (conn, tiss.)

Myxobolus miilleri Butschli (conn, tiss.)
Myxobolus physophilus Reuss (sxirface)
Myxobolus perntagnus Wegener

8) Body-cavity (cyst). — Myxobolus sp. Gurley

Myxobolus carassii Klokacewa
Gen. et spec, incert. Leydig

9) Nervous tissue. — Myxobolus neurobius Schuberg et Schroder

Lentospora encephalina Mulsow (blood vessel of brain)

10) Bone, cartilage, perichondrium. — Lentospora cerebralis (Hofer) Plehn

Myxobolus aeglegini Auerbach
Henneguya brachyura Ward (of fin)

11) Stomach, pyloric cecum. — Myxobolus exiguus Thfilohan

Myxobolus mesentericus Kudo
Henneguya tenuis Vaney et Contc

12) Liver. — Myxobolus ellipsoides Th61ohan

Myxobolus oviformis Th^lohan
Myxobolus cyprini Doflein
Myxobolus cordis Keysselitz
Myxobolus musculi Keysselitz
Myxobolus carassii Ellokacewa
Myxobolus mesentericus Kudo

13) Gall-bladder. — Leptoiheca agilis Th^lohan

Leptotheca elongata Thfilohan
Leptotheca polymorpha (Th61ohan) Labb6
Leptotheca parva Th61ohan
Leptotheca hepseti Th61ohan
Leptotheca sp. Awerinzew
Leptotheca macrospora Auerbach
Leptotheca informis Auerbach
Leptotheca longipes Auerbach

40 ILUNOIS BIOLOGICAL MONOGRAPHS 1278

Leptotheca fusifonnis Davis
LepMheca scissura Davis
Caratomyxa arcuaia Thflohan
Ceratomyxa spkaerulosa Th^lohan
Ceratomyxa paUida Th61ohan
Ceratomyxa globurijera Thelohan
Ceratomyxa appendiadata Th61ohan
Ceratomyxa truncata Thelohan
Ceratomyxa reticularis Th6lohan
Ceratomyxa inaequalis Doflein
Ceratomyxa linospora Doflein
Ceratomyxa ramosa Awerinzew
Ceratomyxa drepanopsettae Awerinzew
Ceratomyxa tylosuri Awerinzew
Ceratomyxa (?) spari Awerinzew
Ceratomyxa sp. (?). Awerinzew
Ceratomyxa sp (?). Awerinzew
Ceratomyxa acadiensis Mavor
Ceratomyxa sp. Georgdvitch
Ceratomyxa coris Georgfivitch
Ceratomyxa herouardi Georg6vitch
Ceratomyxa mesospora Davis
Ceratomyxa sphairophora Davis
Ceratomyxa taenia Davis
Ceratomyxa attenuata Davis
Ceratomyxa recurvata Davis
Ceratomyxa lunaia Davis
Ceratomyxa abbreviata Davis
Ceratomyxa flagellif era Davis
Ceratomyxa agglomerata Davis
Ceratomyxa amorpha Davis
Ceratomyxa monospora Davis
Ceratomyxa streptospora Da\'is
Ceratomyxa aggregata Davis
Ceratomyxa undidata Davis
CUoromyxum leydigi Mingazzini
CUoromyxum fluviatile Th61ohan
CUoromyxum truttae L6ger (also gall-duct)
CUoromyxum cristatum Ldger
CUoromyxum dubium Auerbach
CUoromyxum sp. Awerinzew
CUoromyxum thymaUi Lebzelter
CUoromyxum koi Fujita
CUoromyxum magnum Awerinzew
CUoromyxum misgumi Kudo
CUoromyxum fujiiai Kudo
CUoromyxum trijugum Kudo
CUoromyxum catostomi Kudo
CUoromyxum wardi Kudo
Sphaerospora masovica Cohn
Myxidium incurvatum Thflohan
Myxidium sphaericum Thdohan

2791 STUDIES ON MYXOSPORIDJA—KUDO 41

Myxidiutn sp. Guriey (only in gall-duct)

Myxidium giganteum Doflein

Myxidium pfei£eri Auerbach

Myxidiutn inflatum Auerbach

Myxidium bergense Auerbach

Myxidiutn procerutn Auerbach

Myxidiutn tnacrocapsulare Auerbach

Myxidiutn sp. Awerinzew

Myxidiutn ovifortne Parisi

Myxidium sp. Mavor

Myxidium kagayamai Kudo

Myxidium gadi Georg6vitch

Myxidium glutinosum Davis

Myxidium phyllium Davis

Myxidium striatum Cunha et Fonseca

Myxoholus misgurtii Kudo (few spores only)

Myxobolus sp. Lebzelter (spores only)

Sphaeromyxa balbiatiii Th61ohan

Sphaeromyxa incurvata Doflein

Sphaeromyxa sabrazesi Laveran et Mesnil

Sphaeromyxa hellandi Auerbach

Sphaeromyxa exneri Awerinzew

Sphaeromyxa gasterostei Georgdvitch

Zschokkella twva Klokacewa

Zschokkella acheilognathi Kudo (also in gall-duct)

Gen. incert. congri Perugia

Gen. incer. merlucii Perugia

Gen. et spec, incert. Mavor

14) Spleen. — Myxobolus piriformis Th^lohan

Myxobolus sp. Kudo
Myxobolus ellipsoides Th61ohan
Myxobolus exiguus Thdohan
Myxobolus omformis Th^lohan
Myxobolus pfeifferi Th^lohan
Myxobolus cyprini Doflein
Myxobolus cordis Keysselitz
Myxobolus musculi Keysselitz
Myxobolus mesenlericus Kudo

15) Intestine. — Myxobolus exiguus Thelohan

Myxobolus miilleri Biitschli
Myxobolus pfeifferi Th61ohan
Myxobolus miyairii Kudo
Myxobolus carassii Klokacewa
Myxobolus mesenlericus Kudo
Hentieguya peri-intestinalis C6pSde
Henneguya tenuis Vaney et Conte

16) Ovary.^Wardia ovinocua Kudo

Sphaerospora elegans Th61ohan
Myxidium histophilum Th61ohan
Myxobolus pfeifferi Thelohan
Myxobolus miilleri Biitschli
Myxobolus musculi Keysselitz

42 ILLINOIS BIOLOGICAL MONOGRAPHS [280

Eenneguya oviperda (Cohn)
Henneguya media Thflohan
Eenneguya brevis TMlohan

17) Kidney. — a) Urinary tubules. — Leptotheca renicola Thdlohan

Mitraspora caudata (Parisi) Kudo
Mitraspora cyprini Fujita (also in ureter)
Mitraspora elongata Kudo
Sphaerospora elegans Thfilohan
Sphaerospora divergens Th^ohan
Eenneguya media Th61ohan
Eenneguya brevis Th61ohan
Eenneguya gasterostei Parisi
Eoferellus cyprini Doflein

b) Tissue. — Mitraspora elongala Kudo

CUoromyxum quadratum Thelohan

Sphaerospora rostrata Thelohan (^lalpighian bodies)

Myxidium kistophilum Th61ohan

Lentospora asymmetrica Parisi

Myxobolus pfeifferi Tli61ohan

Myxobolus cyprini Doflein

Eenneguya neapolitana Parisi (conn. tiss. of ren. tubules)

Eoferellus cyprini Doflein

c) Seat, unstated. — CUoromyxum mucronaium Gurley

Sphaerospora angtdata Fujita
Myxidium barbatulae CdpMe
Myxidium giardi C^pede
Myxobolus piriformis Thelohan
Myxobolus ellipsoides Thdlohan
Myxobolus exiguus Thelohan
Myxobolus oviformis Thelohan
Myxobolus mOUeri Btitschli
Myxobolus obesus Gurley
Myxobolus cycloides Gurley
Myxobolus cordis Keysselitz
Myxobolus musculi Keysselitz

18) Urinary bladder. — Leptotheca lobosa Davis

Leptotheca glomerosa Davis
Ceratomyxa naricularia Davis
Ceratomyxa spinosa Davis
CUoromyxum mucronaium Gurley
CUoromyxum granulosum Davis
Sphaerospora elegans Thelohan
Sphaerospora divergens Th61ohan
Sphaerospora polymorpha Davis
Sphaerospora sp. Davis
Sinuolinea dimorpha Davis (also in ureter)
Sinuolinea capsularis Davis
Sinuolinea arborescens Davis
Sinuolinea opacHa Davis
Sinuolinea brachiophora Davis
Myxidium libber kiihni Biitschli
Zschokkella hildae Auerbach
Zschokkella globulosa Davis

281]

STUDIES ON MYXOSPORIDIA—KUDO

43

Myxoproteus atnbiguus (Th^lohan) Doflein

Myxoproteus cordiformis Davis

Myxoproteus cornutus Davis

Henneguya legeri C6pSde

Henneguya wisconsinensis Mavor et Stxasser

Henneguya mictospora Kudo

Gen. et spec, incert. Mavor

19) Testis. — Myxobolus pfeifferi Th61ohan (only spores)

20) Mesentery. — Myxobolus mesentericus Kudo

21) Seat unknown. — Henneguya sp. Gurley (integument?)

Gen. et. spec, incert. Borne

II Amphibia

1) Gall-bladder. — CMoromyxum caudatutn Th^Iohan

Sphaeromyxa immersa (Lutz) Thfilohan

2) Urinary tubules of kidney. — Wardia oUmacheri (Gurley) Kudo

CMoromyxum protei Joseph

TABLE I

4-1

a

a

Si

1

o

3

<

i

'•3

i

1

M

O
CO

t3

O
11

33

3,

C/5

=3

1

5

.2

O

1

1(a)*

1

rtT3
t33

2
2
3

b

Si

a

o
w 9

Leptotheca

Ceratomyxa

Myxoproteus....

Wardia

1

1(a)

3(a)

Kb)*

1(a)

Kb)

1(c)*

2(a)

Kb)

1(c)

2(a)

Kb)

2(c)

3(a)

Mitraspora

3

14

1

Chlorom3rxum ..

1

2

4
5

1

1

1

1

1

Sphaerospora....

Sinuolinea

Myxidium

1

18

7
2

1

Sphaeromyxa....

Zschokkella

2

Myxosoma

3
1

28

8

Lentospora

1

10
4

6

3

1
9

4

6
1

1

4

1
1

1
1

1

Kb)
2(b)
9(c)
3(a)
Kb)
1(a)
Kb)

1

1

Myxobolus

Henneguya

2

1

7

2

10

6
2

2

4
3

3

1

.... 1
1

Hoferellus

Total

16

10

18

8

41

1

4

9

^

^

7

88

10

s

2

10

39

3

24

1

7

' For (a), (b) and (c), see page 42.

44

ILLINOIS BIOLOGICAL MONOGRAPHS

[282

3) Testis, oviduct. — Myxobcius hylat Johnston ct Bancroft

m RZPTILIA

1) Kidney (ren. tub.). — Myxidium danilewskyi Laveran

Myxidium mackiti Bosanquet
Myxidium atnericanum Kudo

2) Ovary. — Gen. et ^)ec. incert. Mingazzini

rV Insecta
1) Abdominal cavity. — Chhromyxum diploxys Guriey

V Annelida
Myxoboius ^. Guriey

The data in this section are summarized on the preceding page (Table I).

D. THE EFFECT OF ENVIRONMENT ON THE ORGANAL DISTRIBUTION
OF MYXOSPORmL\ IN HOSTS

Myxosporidia are almost equally distributed among marine and fresh-
water fishes in regard to the number of species. This is shown in the follow-
ing table.

T.\BLE II

Number of species

Number of species

Other hosts

Genus

found in marme

found in fresh-

fi»h

water fish

RepL

Amph.

In.sect

Annelida

Leptotheca (15)*.„.

15

Ceratomyxa (35)....

35

Myxoproteus (3)....

3

Wardia (2)

1

1

Mitraspora (3)

3

....

....

Ghloromjnum (22)

7

12

2

....

Sphaerospora (10).

5

5

....

Sinuolinea (5)

5

....

Myxidium (26)

17
(2 common)

8
(2 common)

3

....

Sphaeromyxa (7)....

6

....

....

1

....

ZschokkeUa (4)

2

2

....

....

Mjniosoma (3)

1

2

....

....

Lentospora (6)

2

(2 common)

6

(2 common)

....

Mjrxobolus (63)

5

56

1

1

Henneguya (32)

1

31

HofereUus (1)

....

1

....

....

Gen. et spec

incert (12)

4

7

1

....

....

Total 237+12

104+4

134+7

4

5

1

1

* The number in parenthesis denotes the number of species in the corresponding genus.

2831

STUDIES ON MYXOSPORJDJA—KUDO

45

These genera have certain relations to the organal distribution in the
body of the host, which are shown in List IV (page 37) and Table I (on page
43) and which can be put together as follows :

TABLE III

Genus

Number of species
found in body-
cavity

Number of species
found in tissue

Number of

species found

in both

Seat
unknown

Leptotheca (15)

14

35
3
1
2

18

' 4

5

22
6
4

1
2
4

4

1

1

4
4

4

3

5

59

28

5

1

1

1

Ceratomyxa (35)

Myxoproteus (3)

Wardia (2)

Mitraspora (3)

Chloromyxum (22)

Sphaerospora (10)

Sinuolinea (5)

Myxidium (26)

Sphaeromyxa (7)

Zschokkella (4)

Myxosoma (3)

Lentospora (6)

Myxobolus (63)

Henneeuva (32)

Hoferellus (1)

Gen. et spec, inct (12)....

3

Total 237+12

121+4

109+5

3

4+3

From the facts shown in the above tables, the following conclusions can
be drawn.

1) The genera Leptotheca (one species in tissue), Ceratomyxa, Myxo-
proteus, Sinuolinea and Sphaeromyxa (one species in Amphibia) include
parasites from the body cavity of marine fish.

2) The majority of the genera Lentospora, Myxosoma, Myxobolus (one
species in Amphibia), Henneguya and Hoferellus include parasites in tissues
of fresh-water fish.

3) The genera Chloromyxum, Sphaerospora, Myxidium and Zschok-
kella include forms that infect the body-cavity as well as the tissue of
marine and fresh-water fishes.

4) The genus Mitraspora includes three species that parasitize the
fresh-water fish.

5) The genus Myxidium has three species found in the kidney of
reptiles.

46 ILLINOIS BIOLOGICAL MONOGRAPHS [284

6) The genera Wardia, Chloromyxum, Sphaeromyxa and Myxobolus
include species parasitic in Amphibia.

7) The genus Chloromyxum has one species found in an insect.

8) The genus Myxobolus has one species found in an annelid, which
was not normally recorded.

285]

STUDIES ON MYXOSPORIDIA—KUDO

47

THE SPORE

As will be shown later (page 52-55), the spore stage is still the only
constant character by which various forms of Myxosporidia are identified
from each other. For this reason it is necessary to have a clear conception
of the form and structure of the spore and at the same time to define the
terms used in the present paper, even tho they have commonly been
used heretofore.

Anterior end
Foramen or polar capsule

Shell
sutural ridge

Polar capsule
Coiled polar filament

Nuclei 07 sporoplasm
Sporoplasm

lODINOPHlLOUS vacuole
POSTXKIOX END

A B

Textfigure
Diagrammatical front [A] and side p] views of a Mvxoboltjs spore.

(For further explanation see following pages.)

The spore of Myxosporidia is covered by a shell, which is composed
of two valveSy usually symmetrical in form and size that come in contact
in the sutural plane. The sutural line is straight in most cases, tho some-
times curved like an S. It is more or less thickened, forming the sutural
ridge. The sutural ridge is to be made out clearly in fresh as well as in
stained preparations and furnishes important data in regard to the classi-
fication of the parasite. The thickness of the shell- valve is usually uniform;
in some species (Myxobolus), however, it may differ slightly in different
parts of the shell. Besides, in many species of Myxobolus, the shell
differentiates a small triangular intercapsular appendix on the inside at the
anterior end directed posteriad between two polar capsules.

The form of the spore varies greatly owing to the shape of the shell
together with its variously developed appendages; 1) lateral appendages
as in Ceratomyxa, 2) anterior processes as in Myxoproteus, 3) posterior
processes as in Wardia (fringe-like), Mitraspora (filiform), Hoferellus
(spinous), Henneguya (tail-form), etc.

48 ILLINOIS BIOLOGICAL MONOGRAPHS [286

The surface of the shell may be smooth or exhibit various markings.
More or less conspicuous ridges varying in form and number in different
species, may run parallel to the sutural line, may show a network-like
structure or may exhibit short tooth-like processes arising from the sutural
ridge and radiating toward the center of each valve. When the ridges are
fine, they form delicate striations, arranged usually parallel to the sutural
line. Tho these markings are usually easily seen in vivo, they are very
often more readily studied in stained preparations.

Inside of the shell are present the polar capsules and sporoplasm.
Gurley (1894:120) and Davis (1917:210) used the term "capsules" instead
of polar capsules because of the facts that "the situation implied by the
the latter (polar capsule) is not constant" (Gurley) and that "they are
often not in the position indicated by the term polar capsule" (Davis).
The present writer, however, does not agree with these authors and retains
the commonly used term, polar capsule, thruout the present paper on
the basis of the fact that these polar capsules are situated at or near the
more or less attenuated anterior end in the great majority of species or at
each end (in Myxidiidae) of the spore, except in the few cases as in Wardia
in which they are situated in the central portion and have the foramina at
the anterior end of the spore.

The polar capsules may be pyriform or spherical. They are located
at or near one end (anterior end) of the spore. In Myxidiidae, one polar
capsule is situated at each end, in which case no distinction can be made
between the anterior and posterior ends. The end or side opposite to the
anterior, is the posterior end of the spore. The number of polar capsules
in a spore varies according to the dififerent genera. There is only one polar
capsule in the spore of unicapsular Myxobolus, four in Chloromyxum, two
in all the other genera. They may be equal or unequal in form and size.
When two polar capsules are located at the anterior end, they may be
convergent or divergent. Each has a foramen to the outside of the spore
thru the shell in or near the sutural line, thru which the polar filament is
extruded. The foramen is observable in the fresh condition. Staining
will very often show clearly the canals thru the shell. Each polar capsule
has an independent foramen.

In the polar capsule exists a coiled polar filament, which in most
cases can be recognized without diflficulty in the fresh condition. The
polar filament is as a rule a more or less extended, probably hollow thread
connected with the polar capsule, which is extruded from the spore thru
the foramen under the action of the stimulants such as the digestive fluid of
the host or certain chemicals. In Sphaeromyxa it is rather short and thick,
tapering to a point. The polar filament is coiled around the longest
axis of the polar capsule, except in Sphaeromyxa in which it is coiled
around an axis perpendicular to the longest axis of the polar capsule.

287] STUDIES ON MYXOSPORIDIA— KUDO 49

The sporoplasm occupies the extracapsular cavity at the posterior
region of the spore. It is of granular structure with almost always two
nuclei. Besides, it has an iodinophilous vacuole mostly round or oval in
the spores of the family Myxobolidae. It occurs thruout the spore stage
and is the important character of the said family. The contents of the
vacuole is probably of glycogenous nature and is stained deeply with
iodine. Small refringent fat globules have also been observed in the spore.

Davis (1917:212) proposed to use capsular and postcapsular sides in
place of anterior and posterior ends which have most frequently been
used and are also used in the present paper. The latter terms can be
employed as properly as Davis' terms except in the case of the Myxidiidae,
where both terms, strictly speaking, are inapplicable.

Tho various abnormal spores are very often encountered in several
species, the majority of the spores are of typical form, structure and size.
In Myxosoma and Myxobolus, the spore sometimes develops a short
posterior process, which is highly developed in the spore of the genus
Henneguya.

Young spores, generally speaking, are more rounded in form than the
mature form, while the mature spores, as a rule, are of definite form,
structure and size characteristic to the species. It should, however, be
kept in mind that there is a certain amount of variation among these
characters.

As is generally recognized, one must mention whether the spores were
measured in fresh condition or in fixed and stained state. The fresh spore
is generally more or less larger than the mounted one.

so LUNOIS BIOLOGICAL MONOGRAPHS [288

DEFINITION OF TERMS USED FOR DESCRIPTIONS

Anterior end. — The end of the spore where the polar capsules open; in most

cases the polar capsules are situated at this end.
Anterior process. — The spinous process of the shell at the anterior end of

the spore of the genus Myxoproteus.
Breadth. — The larger diameter of the spore measured at right angles to the

length or sutural diameter; the shorter diameter thus measured being

the thickness.
Capsulogenous cell. — A small island of protoplasm with a nucleus, in which

polar capsule becomes differentiated.
Cyst. — The vegetative form of more or less conspicuous size in tissues of the

host, surrounded usually by a membranous structure composed of the

host issue.
Disporous. — The character of a trophozoite of forming only two spores.
Foramen. — Opening of the polar capsule* thru which the polar filament is

extruded.
Front view. — The view in which length and breadth of the spore are laid

horizontally.
Gemmules. — A small mass of trophozoite separated from the mother body

by plasmotomy. Used by Davis (1917). (See page 105.)
lodinophilous vacuole. — The vacuole in the sporoplasm of the spore of the

family Myxobolidae, the contents of which are stained brownish with

iodine.
Lateral process. — The lateral prolongation of the shell-valve at right angles

to the sutural plane.
Length. — Antero-posterior diameter of the spore in the sutural plane;

equivalent to sutural diameter. ♦

Longitudinal striations. — Fine ridges or thickenings marked longitudinally

on the shell of the spore.
Mesoplasm. — An intermediate layer between ectoplasm and endoplasm,

coined by Cohn in the case of Myxidium lieherkiihni (see page 107).
Mictosporous. — The character of the trophozoite of forming a variable

number of spores in an individual.
Monosporous. — The character of the trophozoite of forming a single spore.
Pansporoblast. — Coined by Gurley (1893:408) used here in the same mean-
ing, an enclosed area in the endoplasm of the vegetative form, in which

two sporoblasts become dififerentiated.

289] STUDIES ON MYXOSPORIDIA—KUDO 51

Plasmogamy. Fusion of two trophozoites, coined by Doflein (1898).
Plasmotomy. — Division of trophozoite into daughter individuals, coined by

Doflein (1898).
Polar capsule. — The pyriform or spherical, hollow body in the spore which

forms a polar filament.
Polar filament. — The filament which is coiled inside the polar capsule.
Polysporous. — The character of the trophozoite of forming spores, more

than two.
Posterior filament. — Fine posterior appendage of the spore.
Posterior processes. — Posterior differentiations of the shell.
Ridge. — Linear or network-like elevation of the shell of the spore.
Shell. — The envelope of the spore.

Shell-valves. — Two valves which compose the shell of the spore.
Sporoplasm. — The protoplasmic mass found inside of the spore (amebula

or sporozoite) , usually situated in the posterior portion of the spore.
Sutural diameter. — Same as length.

Sutural edge. — The edge of the shell-valves cut by the sutural plane.
Sutural line. — The line on the shell of the spore marked by the sutural plane.
Sutural plane. — The plane on which two shell- valves meet together.
Sutural ridge. — The ridge marking the sutural line.
Tail. — The posterior prolongation of the valves from the median posterior

end; it may be a single process or bifurcated.
Thickness. — See breadth.

Trophozoite. — The vegetative or multiplicative stage of a Myxosporidian.
Vegetative form. — Same as trophozoite.

52

ILLINOIS BIOLOGICAL MONOGRAPHS

[290

CLASSIFICATION OF MYXOSPORIDIA

The classification of Myxosporidia, was first carried out by Thelohan
as early as 1892, who considered rightly that the spore was the only reliable
means for the purpose. In 1899 and 1901, Doflein introduced into the
classification two Legions, Disporea and Polysporea, and a new family.
This plan has generally been followed by various authors in dealing with
these protozoa.*

The classification of the said author, however, no longer agrees with
our present knowledge of the animals. In the first place, as was pointed
out by some authors, for instance Davis (1917:217), it is far from being
correct to divide the Myxosporidia into two Legions, Disporea and Poly-
sporea, on the basis of the number of spores formed in each vegetative
form, since this differs even in one and the same species as was observed
by Leger, Auerbach, Awerinzew, Parisi, Georgevitch, Davis, Kudo and
others (see Table IV on page 53).

Auerbach who had observed numerous interesting facts in this group,
had adopted Doflein's classification in his splendid work (1910) by simply
adding two genera, Zschokkella and Lentospora, to the family Myxidiidae.
In the following year (1911), he tried a new classification, on the same
basis as Doflein did, by introducing two new Legions besides these two
already existing, and by discarding all the families. Thus:

I Monosporea
II Mictosporea

III Disporea
rv Polysporea

a) Genus

a) Genus

b) Genus

c) Genus

d) Genus

e) Genus

f) Genus

a) Genus

b) Genus

a) Genus

b) Genus

c) Genus

d) Genus

e) Genus

Coccomyxa

Zschokkella

Myxoproteus

Myxidium

Sphaeromyxa

Chloromyxum

Sphaerospora

Ceratomyxa

Leptotheca

Myxosoma

Lentospora

Myxobolus

Henneguya

Hoferellus

As will be distinctly seen from Table IV, the classification not only fails
to improve Doflein's classification in bringing together the genera, Myxo-

* Doflein still uses the same classification in his recent work (1916).

291)

STUDIES ON MYXOSPORIDIA—KUDO

S3

proteus, Myxidium and Sphaeromyxa into Mictosporea, and Lentospora
and Henneguya into Polysporea, but increases the confusion concerning
relationship among the genera.

TABLE IV

Genus

Mono-

Mono-

Mono-,

and

and

Di-

di- and

di-

poly-

sporous

poly-

sporous

sporous

sporous

8

3

23
2

■■

1

2

2

1

2
2

1

1

2

2

1

2

1
1

Di- and

poly-
sporous

Poly-
sporous

Unknown

Leptotheca
15 species....

Ceratomyxa
35 species....

Myxoproteus
3 species

Wardia*

2 species

Mitraspora*

3 species

Chloromyxum

22 species...
Sphaerospora

10 species

Sinuolinea*

5 species

Myxidium

26 species

Sphaeromyxa

7 species ,

Zschokkella

4 species

M5Tcosoma

3 species

Lentosp)ora

6 species

Myxobolus

63 species

Henneguya

32 species

Hoferellus

1 species

1
1
4

2
1
9

5
1

3

2

54

25

1

7
5
1
1

7
2

10
2
1

2
9
5

* These three genera are unknown to Auerbach, except two species which were formerly
placed in Leptotheca and Sphaerospora.

54 ILLINOIS BIOLOGICAL MONOGRAPHS [292

Parisi (1912) followed Auerbach in his paper dealing with Myxosporidia
from Italian waters. Poche (1913) put Auerbach's classification in better
form as follows:

Order: MjTcosporidia

2 Superfamily Mictosporea

2 Family Myxidiidae

2 Genus Zschokkella

3 Genite Myxoproteus

4 Genus Myxidium

5 Genus Sphaeromyxa

6 Genus Sphaerospora

3 Family Chloromyxidae

7 Genus Chloromyxum

3 Superfamily Disporea

4 Family Ceratomyxidae

8 Genus Ceratomyxa

9 Genus Leptotheca

4 Superfamily Polysporea

5 Family Myzosomatidae (Poche)

10 Genus Myxosoma

11 Genus Lentospora

6 Family Myxobolidae

12 Genus Myxobolus

13 Genus Henneguya

14 Genus Hoferellus

For the same reason given in discussing Auerbach, this, however, is not
conformable with the present state of knowledge regarding these protozoa.

It was not until 1917 that the classification of the Myxosporidia
approached to a more natural state in the valuable work by Davis (1917:
219-221). He pointed out sharply the unsatisfactory features in Doflein's
classification and proposed a different system as follows:

Order: Myxosporidia.

Suborder I Myxosporea Davis
Family 1 Ceratomyxidae

Genus 1 Leptotheca

Genus 2 CeratomjTca
Family 2 Sphaerosporidae Davis

Genus 1 Myxoproteus

Genus 2 Sphaerospora

Genus 3 Sinuolinea
Family 3 Myxidiidae

Genus 1 Myxidiima

Genus 2 Sphaeromyxa

Genus 3 Zschokkella
Family 4 Chloromyxidae

Genus 1 Chloromyxum

2931

STUDIES ON MYXOSPORJDIA—KUDO

55

Suborder n Cystosporea Davis
Family 1 Myxosomidae* Davis
• Genus 1 Myxosoma

Genus 2 Lentospora
Family 2 Myxobolidae

Genus 1 Myxobolus •

Genus 2 Henneguya
Genus 3 Hoferellus

Thus, Davis selected the form of the spore for the establishment of two
suborders and further rearranged the genera into closer positions to show
relationship to each other better than any one of the previous authors.
He, however, named the suborders according to a secondary character,
i.e., the seat of the parasites in the host. According to his definition the
trophozoites of the species belonging to Myxosporea are "with few excep-
tions free living in the body-cavity," while those of Cystosporea "with
few exceptions" are tissue parasites.

From TABLE III on page 45, are taken the following data regarding
this point:

Total nimiber

of species

known

Number of

species found in

body cavity

Number of

species found in

tissue

Number of

species found in

both places

Seat
unknown

M)ntosporea.
Cystosporea.

132
105

114

7

14

95

Thus it appears that the terms Myxosporea and Cystosporea do not
seem to be properly used. These may be replaced by terms that denote
the first and common character of the suborders.

The suggestions as to the adoption of other characters than the spore
for the divisions of Myxosporidia, proposed by Awerinzew (1907:831;
1908:64), Auerbach (1910:161) and Davis (1917:217) can only be applied
in the future. At the present time, the characters concerning the vegetative
form do not appear to afford a better and more natural basis for the
classification of Myxosporidia than those of the spore. Thus from the
taxonomic point of view the present situation does not seem to be much
improved as compared with that at the end of the last century.

The writer proposes in the following pages a new classification based on
the characters of the spore.

* Davis did not notice the establishment of the family Myxosomatidae by F. Poche (1913),
including exactly the same genera. See page 54.

56 ILLINOIS BIOLOGICAL MONOGRAPHS [294

Order MYXOSPORIDIA Butschli 1881
Suborder EURYSPOREA nom. nov.
Largest diameter of the spore at right angles to the sutural plane.
One polar capsule on each side of the plane. Sporoplasm with no iodin-
ophilous vacuole. Vegetative form found in body cavity (except 2 species) .
Great majority parasites of marine fish. Monosporous, disporous and
polysporous.

Family CERATOMYXIDAE Dofiein 1899
With the characters of the suborder.

Genus LEPTOTHECA Thelohan 1895
Shell-valves of spore hemispherical or shortly rounded, 15 species.
Disporous (7 unknown). 14 species in body-cavity; 1 in tissue; all in marine
fish. Type species: Leptotheca agilis Thelohan.

Genus CERATOMYXA Thelohan 1892
Shell- valves, conical and hollow, attached on the bases; free ends
extended, tapering to more or less sharply pointed or rounded ends.
Sporoplasm usually does not fill the cavity, but is located asymmetrically
in it. 35 species. Disporous (23 species), monosporous and disporous
(3 species), disporous and polysporous (4 species) and unknown (5 species).
All (except 2 species in urinary bladder) in the gall-bladder of marine fish.
Type species: Ceratomyxa arcuata Thelohan.

Genus MYXOPROTEUS Doflein 1898 emend. Davis 1917
Spores roughly pyramidal; with or without distinct processes from the
base of the pyramid. 3 species. Disporous (one species unknown). All
in urinary bladder of marine fish. Type species: Myxoproteus atnhiguus
(Thelohan) Doflein.

Genus WARDIA nov. gen.
Spore form of isosceles triangle with two convex sides. Oval in profile.
Surface of shell with fine ridges which turn into fringe-like processes at the
posterior end. The polar capsules, large and perfectly spherical, situated
at the central portion of the spore, opening at the anterior tip. Two spe-
cies. Polysporous (one species unknown). Tissue parasite (one species)
of fresh-water fish and amphibia, both found in Illinois, U. S. A. Type
species: Wardia ovinocua nov. spec.

Genus MITRASPORA Fujita 1912 emend. Kudo
Spores spherical or ovoidal. Two polar capsules pyriform, one situated
on each side of the sutural plane. Shell longitudinally striated; with or
without long and fine filaments projecting posteriorly in a row at right

295] STUDIES ON MYXOSPORJDIA— KUDO 57

angles to the sutural plane at the posterior side. 3 species. Disporous
and polysporous. All found in kidney of fresh- water fish. Type species:
Mitraspora cyprini Fujita.

Suborder SPHAEROSPOREA nom. nov.
Spores spherical or subspherical, with two 'to four polar capsules.
Sporoplasm without iodinophilous vacuole. Vegetative form found in
body-cavity and tissue. Monosporous, disporous and polysporous.
Parasites of marine and fresh-water fish and amphibia.

Family CHLOROMYXIDAE Thelohan 1892*

Spores with four polar capsules. Monosporous, disporous and poly-
sporous.

Genus CHLOROMYXUM Mingazzini 1890

With the characters of the family. 22 species. 18 in body cavity;
4 in tissue. 7 from marine and 12 from fresh-water fish, 2 in amphibia,
1 in insect. Type species: Chloromyxutn leydigi Mingazzini.

Family SPHAEROSPORIDAE Davis 1917
Spores with two polar capsules. Monosporous, disporous and polyspo-
rous.

Genus SPHAEROSPORA Thelohan 1892
Spores with two polar capsules. Monosporous, disporous and poly-
sporous. 10 species. Body-cavity and tissue. 5 from fresh-water and 5
marine fish. Type species: Sphaerospora divergens Thelohan.

Genus SINUOLINEA Davis 1917
Spores with or without lateral processes. Two polar capsules spherical.
Sutural line sinuous. 5 species. Disporous and polysporous. In the
urinary bladder of marine fish. Type species: Sinuolinea dimorpha Davis.

Suborder PLATYSPOREA nom. nov.
Sutural plane of the spore coincides with or at an acute angle to the
longest diameter. One or two polar capsules. Sporoplasm with or without
an iodinophilous vacuole.

Family MYXIDIIDAE Thelohan 1892
Two polar capsules, one at each end. Sporoplasm without any
iodinophilous vacuole. Spores fusiform.

* Th61ohan (1892) used the terms: Chloromyxidees, Myxidid^es, Myxobolid^es, which
Gurley (1893) made over into Chloromyxidae, Myxidiidae, Mj^obolidae, so that the credit
of recognizing and establishing these families belongs to Thelohan.

58 ILLINOIS BIOLOGICAL MONOGRAPHS [296

Genus MYXIDIUM ButschU 1882
Spores more or less regularly fusiform, with pointed or rounded ends.
Polar filaments long and fine. 26 species. Monosporous, disporous and
polysporous. 22 in body-cavity; 4 in tissue. 15 in marine and 6 in fresh-
water fish, 2 in fishes from both waters and 3 in reptilia. Type species:
Myxidium Ueberkuhni Biitschli.

Genus SPHAEROMYXA Thelohan 1892
Spores fusiform, with truncated ends. Polar filament short and thick.
Trophozoites large and disc shaped. 7 species. Polysporous (2 unknown).
6 in body-cavity; 1 unknown. 6 in marine fish; 1 in amphibia. Type
species: Sphaeromyxa halhianii Thelohan.

Genus ZSCHOKKELLA Auerbach 1910
Spores, semicircular in front view; pointed at ends. Polar capsules
large and spherical, opening on the flat edge near the tips. Sutural line
usually curved in S-form. 4 species. Monosporous, disporous and poly-
sporous. Body-cavity. 2 from marine and 2 from fresh-water fish. Type
species: Zschokkella hildae Auerbach.

Family MYXOSOMATIDAE Poche 1913
Two polar capsules at the anterior end. Sporoplasm without iodino-
philous vacuole.

Genus MYXOSOMA Thelohan 1892
Spores ovoidal, flattened and more or less elongated. 3 species. Poly-
sporous. Tissue parasites. 2 in fresh-water and 1 in marine fish. Type
species: Myxosoma dujardini Thelohan.

Genus LENTOSPORA Plehn 1905
Spores similar to Myxobolus in form. Sporoplasm without any iodino-
philous vacuole. 6 species. Disporous and polysporous (2 unknown). 1
in marine and 3 in fresh-water fish, 2 from fishes in both waters. Type
species: Lentospora cerebralis (Hofer) Plehn.

Family MYXOBOLIDAE Thelohan 1892
Spores with one or two polar capsules at the anterior end, with or
without posterior processes. Sporoplasm with an iodinophilous vacuole.
Majority polysporous in fresh-water fishes.

Genus MYXOBOLUS Biitschli 1882
Spores ovoidal or ellipsoidal; flattened. One or two polar capsules at
the anterior end. Shell without posterior process. 63 species. Poly-
sporous (9 species unknown). 59 species in tissue; 4 unknown. 5 in
marine and 56 in fresh-water fish, 1 in annelid and 1 in amphibia. Type
species: Myxobolus mUlleri Biitschli.

2971 STUDIES ON MYXOSPORIDIA—KUDO 59

Genus HENNEGUYA Thelohan 1892
Spores more or less globular or ovoidal. Two polar capsules at the
anterior end. Posterior end of the shell-valves prolonged into more or less
extended processes, which unite and form a tail in the median line. 32
species. Polysporous, disporous and monosporous. 28 species in tissue
and 4 in body-cavity. In fresh- water fish, except one. Type species:
Henneguya psorospertnica Thelohan.

Genus HOFERELLUS Berg 1898
Spores pyramidal, with two posterior processes from the lateral faces.
1 species. Polysporous. Tissue and body-cavity of fresh-water fish.
Type and only species: Hoferellus cy print Doflein.

eO ' ILLINOIS BIOLOGICAL MONOGRAPHS ' [298

DESCRIPTIONS OF GENERA AND SPECIES
Suborder EURYSPOREA nom. nov.
The definition of the suborder is recorded on page 56.

Family CERATOMYXIDAE Doflein
1899 Ceratomyxidea Doflein 1899 : 378

1901 Ceratomyxidae Doflein 1901 : 182

The characters of the family are described on page 56,

Genus LEPTOTHECA Thelohan
1895 Leptotheca Thelohan 1895 : 331

The characters of the genus are described on page 56.
Type species: Leptotheca agilis Thelohan.

LEPTOTHECA AGILIS Thelohan
[Figs. 1 to 5]
1892 Ceratomyxa agilis Th61ohan 1892 : 962

1895 Leptotheca agilis Thflohan 1895 : 332

1898 Leptotheca agilis Doflein 1898 : 294, 297

Habitat: Gall-bladder of Trygon pastinaca L. and Scorpaeona sp.;
France, Rovigno, Napoli.

Vegetative form: Form generally elongated. Anterior end rounded
where a mass of fat globules is found, while the posterior end terminates
in a point. Size not exceeding 85^1 by 20 to 25/n. The posterior part is
sometimes divided into a certain number of lobes. In the protoplasm,
the globules are clearly seen. Pseudopodia are localized at the anterior
portion of the body. They are long, 40 to 50/i in length, filiform and very
active in moving from back toward front, just like the motion of oars.
Disporous.

Spore: Slightly elongated. Dimensions: sutural diameter 6 to 7n,
breadth 11 to 12/t.

LEPTOTHECA ELONGATA Thelohan
[Fig. 6]

1895 Leptotheca dongata Thflohan 1895 : 332

1898 Leptotheca dongata Doflein 1898 : 312

1917 Leptotheca dongata Georgevitch 1917b : 99-106

Habitat: Gallbladder of Merluccius merluccius L. (M. vulgaris) and
Motdla tricirrata; Marseille, Banyuls, Le Croisic, Napoli, Monaco.

2991 STUDIES ON MYXOSPORIDIA—KUDO 61

Vegetative form: Form variable. Many individuals show, however,
a very characteristic form. It is elongated and has the length of about
120/i. The anterior end is enlarged into a disc-shaped depression, on the
edge of which, the branched pseudopodia are formed. The body gradually
narrows itself toward the posterior end. Also club-shaped, etc. The short
lobose pseudopodia show no movement like that of oars.

Georgevitch's form: Young forms, oval or rounded, are attached to
the epithelial cells of the bladder with a long filiform pseudopodium at the
free end. Such forms often agglomerate in great number.

Spore: Form similar to the spore of Leptotheca agilis. Dimensions on
the average: Sutural diameter 12 to IS/x, breadth 18 to 20/x.

LEPTOTHECA POLYMORPHA (Thelohan) Labbe
1895 Leptotheca elongata Thelohan 1895 : 332-333

1899 Leptotheca polymorpJia Labb6 1899 : 88

Habitat: Gall-bladder of Phycis mediterraneus {P. phycis L.); Banyuls.

Vegetative form: Form extremely polymorphous, with three main
types. 1) Somewhat regularly club-shaped, with lobose pseudopodi^,, some-
times filiform at one end. 2) Irregular as is the case with Ceratomyxa
truncata, with long (ISfx) ectoplasmic processes, which are motionless or
very slow in motion. Lobose pseudopodia are formed actively. 3) More
or less rounded with bristle-like filose pseudopodia. Intermediary forms
are also found. Often many individuals unite together. The protoplasm
is much different from other forms, i.e., more homogeneous and compact.
Granules are hardly visible on account of vacuolar appearance.

Spore: Dimensions: sutural diameter 10 to 12)ti, breadth 18 to 20jti,
length of polar filament 40m.

LEPTOTHECA PARVA Thelohan
[Fig. 7]
1895 Leptotheca parva Thelohan 1895 : 333

1912 Leptotheca parva Auerbach 1912 : 42-43

Habitat: Gall-bladder of Scomber scombrus L.; Marseille, Le Croisic,
Le Vivier-sur-mer, Kristiansund, Stavanger, Bergen.

Vegetative form: Form ordinarily rounded, spherical or subspherical.
Often club-shaped. Size not larger than 12 to 15/x in diameter. Proto-
plasm finely granular. Pseudopodia lobose.

Spore: Small, more or less elongated, curved in arch-form. Dimen-
sions: sutural diameter 3 to 4jLt, breadth 8 to IOai.

LEPTOTHECA RENICOLA Thelohan
1895 Leptotheca renicola Thelohan 1895:333

Habitat: Urinary tubules of the kidney of Scomber scombrus L.; Mar-
seille, Le Croisic.

62

ILLINOIS BIOLOGICAL MONOGRAPHS

[300

Vegetative form: Small. No marked character.

Spore: Globular. Form similar to the spore of Sphaerospora. Dimen-
sions: sutural diameter Sn, breadth 10/x.

LEPTOTHECA HEPSETI Thelohan
1895 LepMkeca hepseti Th€lohan 1895 : 334

Habitat: Gall-bladder of Atherina hepsetus L.; Marseille. Of rare
occurrence; Thelohan observed it but once.

Vegetative form: Not described.

Spore: Form triangular with rounded angles. Dimensions: sutural
diameter 7 to Sju, breadth 12 to 15/i.

LEPTOTHECA PERLATA (Gurley) Labbe
[Fig. 8]
1883 Balbiani 1883 : 201, 204

1894 CUoromyxum {Sphaerospora) perlatutn Gurley 1894 : 272
1899 LepMheca perlata Labb6 1899 : 88

Habitat: Acerina cernua L.; France (?).
Vegetative form: Not described.

Spore: Elliptic. Two small polar capsules converging. Dimensions
not given.

LEPTOTHECA sp. Awerinzew
[Figs. 16, 17]

1908 LepMheca sp. Awerinzew 1908 : 51, 52

Habitat: Gall-bladder of Sehastes norvegicus; Eastern Finmark?
Vegetative form: Rounded form with clear differentiation of proto-
plasm into ectoplasm and endoplasm. Plasmotomy occurs.
Spore: Undescribed. No figure.

LEPTOTHECA MACROSPORA Auerbach
[Fig. 9]

1909 LepMheca macrospora Auerbach 1909 : 70-71

1910 LepMheca macrospora Auerbach 1910b : 768-769
1910 LepMheca macrospora Auerbach 1910c : 167
1912 LepMheca macrospora Auerbach 1912 : 42-43

Habitat: Gall-bladder of Sehastes viviparus H. KT.ai,ndS.dactylopterus;
Bergen, Kristiansund (May, September).

Vegetative form: Trophozoites spherical with the average diameter
of 26 to 30/1. Homogeneous ectoplasm layer exhibits somewhat active
ameboid movements. Endoplasm, granular in living specimen, is rather
sharply distinguishable from the ectoplasm and contains large nuclei.

301] STUDIES ON MYXOSPORIDIA—KUDO 63

Spore : Size large. Form resembles to that of Leptotheca parva. Dimen-
sions: sutural diameter and tliickness I3n, breadth 26/i. Polar capsules
short oval, with a length of 5.2/i, length of polar filament about 130m
(KOH). In the second host, a few normal and numerous abnormal spores
with three or four polar capsules were observed.

LEPTOTHECA INFORMIS Auerbach
[Fig. 10]

1910 Leptotheca informis Auerbach 1910b : 770-771

1912 Leptotheca informis Auerbach 1912 : 42-44

Habitat: Gall-bladder of Molva vulgaris Flem. and Gadus merlangus;
Bergen, Tjomo.

Vegetative form: Young trophozoites with somewhat long and narrow
pseudopodia formed of hyaline ectoplasm; movements active. The proto-
plasm is differentiated into ectoplasm and endoplasm. When stained, two
large (7 to 9/i) and two small (3 to 4n) nuclei were observed in an individual,
27m long excluding the pseudopodia. Sporulating trophozoites are gen-
erally round and each forms two spores, which are developed independently
to each other (i.e., not in ordinary pansporoblast). Auerbach observed
centrosomes in the nuclei of larger type in division. Disporous.

Spore: Large and heavily built. Greatly curved. Sutural line fairly
well marked. Polar capsules round. Dimensions: sutural diameter lO/t,
breadth 18 to 20m, thickness 9m, diameter of polar capsules 3 to 4m. Sporo-
plasm contains two nuclei, 3.5 to 4m in diameter.

LEPTOTHECA LONGIPES Auerbach
[Fig. 11]
1910 Leptotheca longipes Auerbach 1910b : 771

1912 Leptotheca longipes Auerbach 1912 : 42^3

Habitat: Gall-bladder of Brosmius brosme Asc; Bergen (May).

Vegetative form: Trophozoites elongated or rounded. Only few
pseudopodia which are very long. Small forms with a very long process,
were observed in large numbers; length of the body being 10m, while the
process was 60m long. Endoplasm contains nuclei of various sizes. Disporous.

Spore: Form similar to that of Leptotheca informis, though smaller.
Dimensions: sutural diameter 8 to 9m, breadth 12 to 14m, thickness 8m,
diameter of polar capsule 2.5m.

LEPTOTHECA FUSIFORMIS Davis
[Fig. 12]
1917 Leptotheca fiisiformis Davis 1917:222

Habitat: Gall-bladder of Cestracion zygaena; Beaufort (July).

64 ILUNOIS BIOLOGICAL MONOGRAPHS [302

Vegetative form: Pyriform, tapering gradually toward the posterior
end, which usually terminates in a long, slender process; colorless and
transparent. Progressive movements rapid. Endoplasm granular, the
granules being more abundant at the anterior end. The average size of
full-grown individuals: 50/i by 13/i. Disporous.

Spore: Elliptical in front view; fusiform in side view. Sutural plane
slightly oblique to the longest diameter, the line forming a marked ridge.
Polar capsules open on opposite sides of the spore. Sporoplasm finely
granular, confined to the central part of spore. Dimensions : sutural diame-
ter 9/i, breadth 16)u, polar capsule 4.5m long, length of polar filament 30/i.

LEPTOTHECA SCISSURA Davis
[Fig. 13]
1917 Leptotheca scissura Davis 1917 : 222

Habitat: Gall-bladder of Dasybatis hastatus and Pteroplatea maclura
Le Sue; Beaufort (July, August).

Vegetative form: Young form elongated, with long attenuated posterior
process; usually sUghtly constricted just posterior to rounded anterior end,
which bears numerous long, filiform pseudopodia. Progressive movement
rapid. Ectoplasm distinguishable at the anterior end. Endoplasm usually
filled with small, clear, colorless spherules, which become larger and
yellowish as the body increases in size. Each spherule contains one to
several dark-brown granules, which increase in size and number and finally
collect in an irregular clump at the centre of spherule. The larger indi-
viduals are usually flattened dorso-ventrally. The posterior end is divided
into long slender processes, presenting sometimes a network caused by the
fusion of two or more adjacent processes. Full-grown forms: length 125 to
150/i, breadth 20 to 25^. The longest observed, 195^1 by 16jLt. Disporous.

Spore: Elliptical in front view; somewhat flattened along the posterior
side. Sutural line distinct and at right angles to the longest diameter.
Polar capsules have foramina at some distance from the capsular margin.
Sporoplasm finely granular, nearly filling both valves. Dimensions:
sutural diameter ll/x, breadth 22/li, diameter of polar capsule 4^.

LEPTOTHECA LOBOSA Davis
[Fig. 14]
1917 Leptotheca lobosa Davis 1917 : 223

Habitat: Urinary bladder of Paralichthys dentatus L.; Beaufort (July).

Vegetative form: Usually spherical which may form a large rounded
pseudopodium composed of ectoplasm. Body colorless and transparent
to translucent. Ameboid movements very slow. Ectoplasm contains
coarse granules, which are of uniform size and very distinct. Endoplasm

303] STUDIES ON MYXOSPORIDIA— KUDO 65

less granular and more transparent than ectoplasm, containing numerous
large, yellow, fat globules, which are abundant in large forms. Diameter
up to 24ai. Disporous.

Spore: Elliptical in front view; valves slightly tapering but rarely
alike. Sutural line forming a sinuous ridge. Polar capsules open at some
distance from the anterior margin. Sporoplasm nearly filling both valves.
Free spores are often seen to remain united at the sutural line. Dimensions:
sutural diameter 9 to 10/i, breadth 16 to IS/u; diameter of polar capsule 3/t.

LEPTOTHECA GLOMEROSA Davis
[Fig. 15]
1917 Leptotheca glomerosa Davis 1917 : 223

Habitat: Urinary bladder of Faralichthys albiguttus] Beaufort.

Vegetative form: Rounded or somewhat irregular in shape, with short
iobose pseudopodia. Body transparent and colorless. Ameboid move-
ments slow. Ectoplasm hyaline, forming a distinct outer layer. Endo-
plasm finely granular, with numerous small fat globules varying in size.
Almost entire body is used for spore formation. Diameter of rounded
sporulating trophozoite about llfx. Disporous.

Spore: Approximately cylindrical; valves rounded at ends. The coiled
polar filament not visible in the polar capsule. Sutural line at right angles
to the longest diameter. Sporoplasm finely granular, fills the extracapsular
cavity of spore. Dimensions: sutural diameter 4.5/i, breadth 9/u, diameter
of polar capsule 2/t.

Genus CERATOMYXA Thelohan

1892 Ceratomyxa Thdohan 1892 : 169, 171, 175

1895 Ceratomyxa Thelohan 1895 : 334

The characters of the genus described on page 56.
Type species: Ceratomyxa arcuata Thelohan.

CERATOMYXA ARCUATA Thelohan

[Figs.

18 to 22]

1892

Ceratomyxa arctuita

Thelohan

1892a

:1091

1895

Ceratomyxa arcuata

Thelohan

1895 :

335-336

1899

Ceratomyxa arcuata

Labb6

1899 :

90

1912

Ceratomyxa arcuata

Parisi

1912 :

290-291

1913

Ceratomyxa arcuata

Jameson

1913 :

2

1916

Ceratomyxa arcuata

Georgevitch

1916a

:3

Habitat: Gall-bladder of Pagellus centrodontus C. et V., Crenilabrus
melops L., Motella tricirrata Bl., Ophidium vasalli, Gohius paganellus L.,
Heliases chromis Gthr.; Scot paena scrofa L., S. porcus L.; France (Marseille,
Banyuls, Concarneau, Roscoff), Italy (Napoli, summer), Monaco (May).

66 ILLINOIS BIOLOGICAL MONOGRAPHS [304

Vegetative form: Polymorphous; generally club-shape, pseudopodia
localized at the broad end; the other end cylindrical or terminating in a
sharp point. Some other different forms. Pseudopodia, always localized,
lobose pointed at the extremities. Ectoplasm hyaline and thin. Endo-
plasm contains fat globules and particular elements, mostly large refractive
globules, which seem to disappear in the sporulating individuals. Dimen-
sions (maximum): length 35 to 40/x, breadth 12 to IS/i, pseudopodia about
10m loiig- Disporous.

Spore: Arch form. Shell valves equal. Sporoplasm occupies the
extracapsular cavity of the spore. The length varies rather considerably
according to the development of the lateral processes, which are occasionally
acuminated or very short. Often extremities are rounded. Dimensions
(Thelohan) ; breadth 20 to 30/i, sutural diameter 5 to 8/i. Parisi's measure-
ments: breadth 25 to 31/i, sutural diameter 5.5 to 6/i, length of polar
capsules 3.5 to An, length of polar filaments 25/x.

Remarks: The writer agrees with Parisi in eliminating Labbe's two
subspecies (1899:90), as they are too arbitrary.

CERATOMYXA SPHAERULOSA Thelohan

[Figs.

23 to 24]

1892

Ceratomyxa sphaerulosa

Thelohan

1892

171

1895

Ceratomyxa sphaerulosa

Thelohan

1895

334-335

1909

? Ceratomyxa sphaerulosa

Auerbach

1909

80

1912

Ceratomyxa sphaerulosa

Auerbach

1912

4,45

1916

Ceratomyxa sphaerulosa

Georg^vitch

1916a

:3

Habitat: Gall-bladder of Mustelus canis Mitch. (M. vulgaris), Galeus
galeus L. (G. canis), Clupea harengus, Scullium canicula Cuv.; St-Valery-
en-Caux, Roscoff, Bergen, Monaco (May).

Vegetative form: Form more or less definite among the adults. Gen-
erally elongated. Both ends slightly attenuated. Wide in the middle part
of the body. Lobose pseudopodia at one of the extremities. Others
more massive or more or less regularly spherical, in which case the pseu-
dopodia are formed from the whole surface. Spherical form does not
exceed 50 to 60iu in diameter. Other forms 90 to lOO/x by 30 to 40/i (largest).
Young forms colorless and are more variable than the adults. Protoplasm
homogeneous and finely granular. Adult form, on the contrary, yellowish
or greenish yellow. The endoplasm is filled with small (5ju in diameter)
spheres, in the centre of which 5 to 6 small granules, yellowish brown or
greenish in color, are present. Disporous.

Spore: Remarkably large. Polar filament can be seen in vivo (easily
extruded by KOH, ether, etc.) Sporoplasm occupies one of the shell- valves,
while a small mass of very pale looking substance is seen in the other.
Dimensions: sutural diameter 10 to 12/i, breadth 90 to 100/x, subspherical
polar capsule 6 to 7 by 5/i, sporoplasm 12 to 15 by 8 to 9/i.

305] STUDIES ON MYXOSPORIDIA—KUDO 67

CERATOMYXA PALLIDA Thelohan

1895

Ceratomyxa pallida

TWlohan

1895 : 336-337

1898

Ceratomyxa pallida

Doflein

1898 : 341

1916

Ceratomyxa pallida

Georg6vitch

1916b : 2, 3

Habitat: Gall-bladder of Box boops L. and B, salpa L.; Marseille,
Villefranche, Rovigno, Monaco (May).

Vegetative form: Ordinarily spherical not exceeding 16 to 20/i in
diameter. Many individuals often found in massive groups. Pseudopodia
lobose and mostly short. Protoplasm extremely pale with fine granules.

Spore: Dimensions: sutural diameter 5^, breadth 25 to 30)u.

CERATOMYXA GLOBULIFERA Thelohan
[Fig. 25]
1895 Ceratomyxa globulifera Th61ohan 1895 : 338

Habitat: Gall-bladder of Merluccius merluccius L. {M. vulgaris) \
Marseille, Banyuls.

Vegetative form: Polymorphous. Elongated into long branches,
including endoplasm. Endoplasm contains small refractive globules.

Spore: Elongated. Shell- valves unequal, one being longer and finer
than the other. Dimensions: sutural diameter 10/i, breadth 50/x.

CERATOMYXA APPENDICULATA Thelohan

[Fig. 26]

1892 Ceratomyxa appendiculata Th61ohan 1892a : 963-964

1895 Ceratomyxa appendiculata Thelohan 1895 : 337

1898 Ceratomyxa appendiculata Doflein 1898 : 300, 311

Habitat: Gall-bladder of Lophius piscatorius L., L. budegassa Spin.;
RoscoflF, Le Croisic, Marseille, Banyuls, Napoli, Rovigno.

Vegetative form: Extremely polymorphous. Young form spherical,
spatulaform, club-shape, etc. In adult form, the main thick part of the
body, in which spore formation takes place, forms 1 to 6 long prolongations,
twice or three times longer than the main part of the body. Pseudopodia
lobose, filiform or elongated with enlargements. Disporous.

Spore: Lateral prolongations of shell- valves well developed. Dimen-
sions: sutural diameter 5 to J/x, breadth 50/i.

CERATOMYXA TRUNCATA Thelohan
[Fig. 27]

1895 Ceratomyxa iruncata Th61ohan 1895 : 336

1912 Ceratomyxa truncata Parisi 1912 : 289-290

Habitat: Gall-bladder of Clupea pilchardus Walb. {Alosa sardina);
Marseille, Villefranche, Napoli (August, September).

68 ILLINOIS BIOLOGICAL MONOGRAPHS [306

Vegetative form: Polymorphous. Ordinarily more or less rounded,
with lobose pseudopodia. Pseudopodia long and often shows very active
movements. Endoplasm very finely granular, contains small fat globules
which are found in irregular mass or in a circular form. Disporous.

Spore: Valves are short and truncate. Sporoplasm occupies the whole
cavity. Spores with three valves are frequently encountered. Dimensions :
breadth 25jLt, 5^ in sutural diameter. According to Parisi, spores with two
shell- valves are rather few (10%), while those with three (70%) and four
shell- valves (20%) prevail in number! Dimensions: breadth 20 to 30/*,
length of the polar filament 45ac.

CERATOMYXA RETICULARIS Thelohan

[Fig. 28]

1895 Ceratomyxa reticularis Th61ohan 1895 : 337-338

Habitat: Gall-bladder of Trachinus draco L.; Banyuls.

Vegetative form: Extremely polymorphous. Generally spherical or
club-shaped. Well developed trophozoites have the similar form as in
C. appendiculata. Endoplasm highly reticular, with refringent fluid.

Spore: Shell valves are short and truncate, one of which is curved to the
rear. Dimensions: sutural diameter 12 to 15/x, breadth 45 to 50/i.

CERATOMYXA INAEQUALIS Doflein

[Fig. 29]

1898 Ceratomyxa inaequalis Doflein 1898 : 284-285

Habitat: Gall-bladder of Crenilabrus mediterraneus and C. pavo;
Napoli.

Vegetative form: Form usually club-shaped. Protoplasm in active
motion, is differentiated distinctly into ectoplasm and endoplasm. Body
yellowish brown by the presence of granules in endoplasm. Inactive forma-
tion of pseudopodia. Ameboid movements or progressive movements by
means of the posterior process. Size: 20 to 40m by 5 to 10/x in average.
Length of the posterior process up to 30/i. After spore formation, only
two nuclei remain in protoplasm, which apparently degenerate later.
Disporous.

Spore: Elliptical, somewhat flattened. Massive. Very transparent.
Both ends round, but unequally built, i.e., one end is club-shaped. Polar
capsules are somewhat round and are bound to the shell by protoplasmic
bridges. The polar filament is not seen in fresh spores. Dimensions:
sutural diameter 6ju, breadth 31/i, diameter of the polar capsule 2.5 to
3fi, length of polar filament is half breadth of the spore (diluted nitric acid).

307] STUDIES ON MYXOSPORJDIA— KUDO 69

CERATOMYXA LINOSPORA Doflein
[Figs. 30 to 31]
1898 Ceralomyxa Unospora Doflein 1898 : 285

Habitat: Gall-bladder of Labrus turdus; Napoli.

Vegetative form: Club- or spindleshape. Protoplasm highly granu-
lated. Body whitish grey, though very transparent. Pseudopodia very
fine and only formed at the anterior end of the body. Size: 30 to 35jw by
16 to 18/x. Disporous.

Spore: Form symmetrical with long thread-like lateral processes. In
sporoblast, the processes are wound around the spore. It is twice as long as
the breadth of the spore. Polar capsules large and spherical pyriform.
Dimensions: total breadth 50^, breadth of the main part of the spore 10
to 12fi, sutural diameter 5ju, length of lateral process 20/i. "Polar filament
was too fine to be measured."

CERATOMYXA RAMOSA Awerinzew

[Figs. 32 and 33]

1907 Ceralomyxa ramosa Awerinzew 1907 : 831-834

*1908 Ceralomyxa ramosa Awerinzew 1908 : 60-66

Habitat: Gall-bladder oi Hippoglossus vulgaris Flemm.; Kjellebjord,
Murman coast.

Vegetative form: Form irregular ameboid, owing to the presence of
peculiar pseudopodia. The middle part of the body is enlarged into an
ellipsoidal form, where nuclei and sporoblasts are present. From this part
two, rarely one or three processes are formed, which branch out several
pseudopodia of different length. The finer portions of pseudopodia anasto-
mose each other and form a characteristic and remarkable network.
Differentiation of protoplasm is not very distinct. Ectoplasm is not well
developed, tho covering the entire surface of the body as a thin la)'er.
Endoplasm slightly vacuolated and granular, forms the greater part of the
body. Disporous and polysporous.

Spore: Form and size (?) resemble C. arcuata. Slightly curved toward
the posterior side. Valves usually unequally built, one being longer than
the other. Sporoplasm almost always asymmetrically situated in the shell.
Polar capsules on each side of the sutural plane and of the plane perpen-
dicular to the sutural plane, cutting the spore into two equal parts. Young
spores in development ellipsoidal to kidne)^ bean shape. Dimensions:
sutural diameter 12 to 20/i, breadth 50 to 80/x.

* Professor J. Zeitlin has kindly translated some part of the paper, for which the writer
expresses his thanks.

70 ILLINOIS BIOLOGICAL MONOGRAPHS [308

CERATOMYXA DREPANOPSETTAE Awerinzew
[Figs. 34 to 39]

1907 Ceratomyxa sp. Awerinzew 1907 : 832-833

1908 Ceratomyxa drepanopsettae Awerinzew 1908 : 1-41, 45-47

1909 Ceratomyxa drepanopsettae Awerinzew 1909:74-112

1912 Ceratomyxa drepanopsettae Auerbach 1912 : 44-45
1918 Ceratomyxa drepanopsettae Kudo 1918 : 14-15

Habitat: Gall-bladder of Pleuronectes platessa, P.flesus, Drepanopsetta
plaUessoides, Hippoglossus vulgaris, Hippoglossoides limandoides and
Paralichihys deniatus; Murmankuste, Kabelvaag, Rorvik, Tjomo, Woods
Hole (August, September).

Vegetative form: Polymorphous. Usually very much elongated and
slender forms. Protoplasm differentiated. Endoplasm coarsely granular.
Pseudopodia lobose and filiform (2 to 3n), with which the trophoz9ites
attach themselves to the epithelium of the bladder. Disporous.

Spore: Curved toward the posterior side. Shell with rounded ends.
Valves almost always unequally built. Dimensions: breadth 50 to 80/i.
Auerbach's form: Form variable. Size: sutural diameter about 12 to
14/*, breadth about 56/i, diameter of polar capsule about 4 to 6/x, length of
the cavity in which the sporoplasm is located about 34n. Kudo's form:
Variable. Sutural diameter 8 to 10m, average breadth 64^, diameter of
polar capsule 6/i.

CERATOMYXA TYLOSURI Awerinzew
[Figs. 40 and 41]

1913 Ceratomyxa tylosuri Awerinzew 1913a : 153

Habitat: Gall-bladder of Tylosurus schismaiorhynchus; Lorengo
Marques, Delagoa Bay (Africa).

Vegetative form: Large, irregular, disc-like or large ameboid, with
blunt lobose pseudopodia and highly granular protoplasm.

Spore: Large. The anterior edge arch-shape, while the posterior edge
has two small horns which are located symmetrically to the sutural line.
Polar capsules elongated and are separated from binuclear sporoplasm by a
special membrane. Rarely spore with three polar capsules. Dimensions
breadth 124 to 140^, sutural diameter 40 to 45/*, thickness 25 to 30/«.

CER.\TOMYXA (?) SPARI Awerinzew

[Figs. 42 and 43]

1913 Ceratomyxa (?) spari Awerinzew 1913a : 153-154

Habitat: Gall-bladder of Sparus berda; Lorenfo Marques, Delagoa
Bay (Africa).

309] STUDIES ON MYXOSPORIDIA—KUDO 71

Vegetative form: Large (100 to 120/*), disc-form ameboid, containing
a large number of enclosures and granules of different size. In one case, a
number of this form, carrying no spore, underwent budding, which resulted
in forming spherical forms of various size, some of which divided again into
2 to 6 parts (Plasmotomy?). Monosporous and disporous.

Spore: More or less curved. Two polar capsules lie closely together
on each side of the sutural plane. Ends of shell- valves are rounded.
Dimensions: breadth 50 to 60/x, sutural diameter 12 to IS/it, thickness
12 to 15ai, polar filament very long (length not given).

Remarks: Awerinzew thinks this is the intermediate form between
Leptotheca and Ceratomyxa.

CERATOMYXA sp. (?) Awerinzew
1913 Ceratomyxa sp. (?) Awerinzew 1913a : 154

Habitat: Gall-bladder of Scatophagus argus; Delagoa Bay (Africa).

Vegetative form: Small, disc-form ameboid (25 to 35/i), containing
two spores of indistinct contour, on account of incomplete formation of the
shell. Two spores, apparently, developed in one pansporoblast. Dis-
porous.

Spore: Form could not exactly be made out. Polar capsules were
arranged like those of other Ceratomyxa.

CERATOMYXA sp. (?) Awerinzew
1913 Ceratomyxa sp. (?) Awerinzew 1913a : 154-155

Habitat: Gall-bladder of Rhinobathus Awer. (?); Lorenzo Marques
(Africa).

Vegetative form: Irregular shape. Endoplasm highly granular. In the
epithelial layer of the gall-bladder numerous, spherical cysts (30 to 35ju)
were found. Two spores are formed in one pansporoblast. Disporous.

Spore: Cylindrical with broad and slightly rounded ends. Dimensions:
sutural diameter 16 to 19/x, breadth 70 to 80/*, thickness 16 to 19ju.

CERATOMYXA ACADIENSIS Mavor
[Figs. 44 to 47]

1915 Ceratomyxa acadiensis Mavor 1915 : 27-30

1916 Ceratomyxa acadiensis Mavor 1916 : 551-574

Habitat: Gall-bladder of Urophycis chuss (trophozoites are attached to
undetermined Myxosporidia, see p. 176), Zoarces angularis, Pseudo-
pleuronectes americanus ; New Brunswick (Canada) (July to September).

Vegetative form: Polymorphous. Typically club-shaped with very
long tail, or irregularly stellate. Pseudopodia show rigidity. Sometimes

72 ILLINOIS BIOLOGICAL MONOGRAPHS [310

clumps of protoplasm along their length, which are connected by thin
hyaline filaments of ectoplasm. Differentiation of protoplasm is usually
observable at the anterior region. Dimensions: length, excluding tail,
12 to 15^1, breadth 10 to 20/x, tail up to 60/i. Disporous.

Spore: Wide, short and slightly compressed dorso-ventrally, with very
long fine lateral filaments. Polar capsules spherical. Polar filament
invisible in vivo. Dimensions: breadth 40 to 50/i, sutural diameter
7 to 8m, diameter of polar capsule 3 to 4^, length of polar filament 70^,
length of lateral filaments 250 to 300/x.

CERATOMYXA sp. Georgevitch
1916 Ceratomyxa sp. Georgevitch 1916a : 3

Habitat: Gall-bladder of Muraena sp.; Monaco (May).
Vegetative form: No description.
Spore: No description. No figure.

CERATOMYXA CORIS Georgevitch
[Fig. 48]

1916 Ceratomyxa coris Georgevitch 1916a : 4-5

1917 Ceratomyxa coris Georgevitch 1917a : 1-20

Habitat: Gall-bladder of Coris julius, C. giofredi; Villefranche (March,
June).

Vegetative form: Various forms, club-shape, spherical or elongated,
with lobose or filiform pseudopodia. Disporous and rarely Polysporous.

Spore: More or less ellipsoidal. Lateral prolongations of the shell-
valves short and truncate. Sutural line straight. Sporoplasm, elongate,
rounded, elliptical, fills a part of the extracapsular cavity of the spore.
Polar capsules rounded, almost spherical, not converging. Dimensions
not given.

Remarks: Georgevitch observed (1917: Fig. 30) that spores of Glugea
marionis occurred in disporous trophozoite of Ceratomyxa coris, which he
thought to have happened accidentally by plasmogamy of these two Cnido-
sporidia. The above mentioned figure, however, strongly suggests that
G. marionis may be leading parasitic life in the trophozoite of C. coris.

CERATOMYXA HEROUARDI Georgevitch
[Fig. 49]

1913 Leptotheca (?) sp. Jameson 1913 : 2

1916 Ceratomyxa herouardi Georgevitch 1916a : 5-8

1916 Ceratomyxa herouardi Georgevitch 1916b : 717-19, 983-985

1917 Ceratomyxa herouardi Georgevitch 1917 : 375-399

Habitat: Gall-bladder of Box Salpa L.; Monaco (May).

3111 STUDIES ON MYXOSPORIDIA— KUDO 73

Vegetative form: Polymorphous. Elongated with same breadth or
tapering to one end; club-shaped with roundish enlargements. Young
trophozoites spherical or pyriform. Pseudopodia long and narrow or
broad and bi- or multi-lobate. Body colorless both in the young and the
adult. Protoplasm homogeneous and finely granular. Disporous and
polysporous. Spores are found inside of the endoplasm and in the roundish
buds, ordinarily two spores being formed in each bud. Number of buds
on one trophozoite varies. Plasmotomy by budding and division.

Spore: Elongated elliptic. Polar capsules spherical and large. Sutural
plane cuts the spore into exactly equal two parts. Two nuclei in sporoplasm
are rather small and are always in one of the shell-valves. Dimensions
not given.

Remark: The form, mentioned by Jameson in the same seat, host and
locality, that "has something of the appearance of a Leptotheca" and that
is also "almost certainly neither of the two Myxosporidia — Ceratomyxa
pallida Sind Henneguy a neapolitana . . . ," is probably identical with the
present form.

CERATOMYXA MESOSPORA Davis

[Fig. 50]

1917 Ceratomyxa mesospora Davis 1917 : 223-224

Habitat: Ga,\\-h\a.dder oi Cestracion zygaena, C . tiburo; Beaufort (July).

Vegetative form: Pyriform, elongate, with long, slender posterior
process. Numerous filiform pseudopodia at anterior end. Progressive
movements rapid. Body colorless. No sharp demarcation between
ectoplasm and endoplasm. Endoplasm finely granular and filled with
small, colorless, homogeneous spherules. Spherules absent at anterior end.
Size: total length 70 to 85^, length exclusive of posterior process 50 to 75ju,
breadth 20 to 25/1. Disporous.

Spore: Greatly elongate, each valve forming a slightly tapering cone,
rounded at the apex. Valves not compressed. Sutural plane forming an
acute angle with the longest diameter. Polar capsules conspicuous. Coiled
polar filaments very distinct. Polar capsules are remarkable in that they
are asymmetrically situated, one being always located in the widest part
of the spore, while the other being a little to one side. Sporoplasm asym-
metrically situated, sometimes being entirely confined to the larger valve.
Dimensions: breadth 50 to 65/i, sutural diameter about 8^, diameter of
polar capsule 4.5/x, length of polar filament 90jLt.

Remarks: Similar to C. sphaerulosa Thel. and occurs with C. recurvata
Davis in the same organ.

CERATOMYXA SPHAIROPHORA Davis

[Fig. 51]
1917 Ceratomyxa sphairophora Davis 1917 : 224

Habitat: Gall-bladder of Scoliodon terrae-novae; Beaufort.

74 ILLINOIS BIOLOGICAL MONOGRAPHS [312

Vegetative form rPyriform, elongate. Numerous fine filiform pseudo-
podia at anterior end. Progressive movements rapid. Body colorless and
transparent. Ectoplasm clear and homogeneous. Structure of endoplasm
highly variable, in majority of trophozoites filled with transparent homo-
geneous spherules. Small fat globules at the anterior end. In some sporu-
lating individuals, the endoplasm shows vacuolated structure without any
spherules, usually, however, sporulating trophozoites exhibit well-defined
spherules. The spherules or vacuoles, as the case may be, are separated by
a thin layer of distinctly granular endoplasm containing numerous rod-
shaped or rounded, colorless bodies, which in their appearance are strikingly
like small bacteria tho they are not bacteria, as they fail to take Giemsa
stain. Size of sporulating trophozoites 100 to llO/i by 25/i. Disporous.

Spore: Shell- valves greatly elongated, tapering gradually toward the
ends. Long, attenuated ends of valves hollow and so fragile that it is
almost impossible to find an example in which they are not more or less
distorted. Sutural plane perpendicular or only slightly oblique to the
longest diameter. Polar capsules are spherical and large; slightly conver-
gent, opening some distance apart on the anterior side. Coiled polar
filament distinct. Sporoplasm confined to large, central part of spores,
but extending farther into one valve than the other. Dimensions: total
breadth 115 to 140/:, sutural diameter about 12/i, diameter of polar capsules
(>H, length of polar filament 75/z.

CERATOMYXA TAENIA Davis

[Figs. 52 and 53]
1917 CeraUmyxa taenia Davis 1917 : 224-225

Habitat: Gall-bladder of ScoUodon terrae-novae; Beaufort.

Vegetative form: Similar to those of C. sphairophora Davis, and no
character has been found by which they may be distinguished. Sporulating
trophozoites can be easily distinguished on account of the very different
appearance of the spore and their different arrangement within the tropho-
zoites. The spores of this species are situated, as is usually the case in
Ceratomyxa, with the greater part of the spore parallel to the long axis of
the trophozoite, only a part of one valve being bent back along the rest
of the spore. Size: sporulating trophozoites length SO/x, breadth 25/i.
Disporous.

Spore: Valves greatly elongated. Shell very thin, the membrance on
opposite sides of each valve being in contact for about two-thirds of its
length, forming a thin ribbonlike structure; basal third of each valve only
slightly compressed; terminal ribbonlike portion of each valve usually
twisted so that plane of ribbon is at right angles to the main part of Jjje
spore. Polar capsules small, pyriform to spherical and convergent. Coiled
polar filament indistinct. Sutural plane perpendicular to the longest

3131 STUDIES ON MYXOSPORIDIA—KUDO 75

diameter. Sporoplasm finely granular, filling the basal third of each valve,
sometimes extending farther into one valve than the other. Dimensions:
breadth 140 to 150/i, breadth of central portion 45jli, sutural diameter 6/Lt,
diameter of the polar capsules 3/t.

CERATOMYXA ATTENUATA Davis

[Fig. 54]

1917 Ceratomyxa attenuata Davis 1917 : 225

Habitat: Gall-bladder of ScoUodon terrae-novae; Beaufort (July).

Vegetative form: Elongate, pyriform, with long, tapering posterior
process; at anterior end numerous long filiform pseudopodia. Progressive
movements rapid. Ectoplasm distinct only at anterior end. Endoplasm
filled with small, refractive, yellowish or brownish granules, which are
uniformly distributed throughout the trophozoite. Between the brownish
granules, the endoplasm is clear and colorless, not granular, except at
extreme anterior end where it contains a clump of small fat globules. Size
of full-grown trophozoites 100 to 120 by 27 /u. Disporous.

Spore: Valves greatly elongated; a symmetrical, one valve being about
15/i shorter than the other and ending abruptly; the longer valve tapering
gradually to a point. About midway of each valve, is a thin septum;
external to the septum the valves are empty. Polar capsules are large,
opening on the anterior margin. Coiled polar filaments distinct. Sutural
plane oblique to longitudinal axis, usually forming a ridge. Sporoplasm
asymmetrically situated in central part of the spore. Dimensions: breadth
llSju, sutural diameter 9n, diameter of polar capsules 4.5/1, length of polar
filament 60jLt.

CERATOMYXA RECURVATA Davis

[Figs. 55 and 56]

1917 Ceratomyxa recurvata Davis 1917 : 225-226

Habitat: Gall-bladder of Cestracion zygaena; Beaufort (July).

Vegetative form: Pyriform with long, slender posterior process.
Body colorless. Actively motile, forming filiform pseudopodia of ectoplasm
at anterior end. Endoplasm colorless and granular, filled with large,
homogeneous spherules. Full-grown trophozoites 130 to 175)u, length of
the main body about lOOfi. Spores are developed singly from distinct
sporoplasts and not necessarily in pairs. Disporous and polysporous (up
to 10 spores, 6 and 8 are common numbers).

Spore: Valves greatly curved toward the posterior side, usually sym-
metrical, but occasionally one may be much more incurved than the other.
Valves circular in cross section at the base but toward the ends greatly

76 ILLINOIS BIOLOGICAL MONOGRAPHS [314

flattened. Ends of valves sharply pointed. Polar capsules large, opening
at some distance from the anterior margin. Coiled polar filaments dis-
tinct. Sporoplasm finely granular usually extending farther into one valve
than the other. Dimensions: breadth between points of greatest curva-
ture about 16/11, sutural diameter 8 to 9/i, diameter of polar capsules 4.5/*.

CERATOMYXA LUNATA Davis

[Figs. 57 to 60]

1917 CeraUmyxa lunata Davis 1917 : 226-227

Habitat: Gall-bladder of Galeocerda tigrinus; Beaufort (August).

Vegetative form: Pyriform, rounded after being on the slide for some
time. Progressive movements slow Endoplasm filled with large, homo-
geneous spherules, which are usually colorless, sometimes light yellow.
At extreme anterior end, the endoplasm contains numerous small fat
globules. Disporous.

Spore: Considerably variable in size and form. The larger and more
typical are more or less crescent-shaped; symmetrical; valves curved
toward rear, terminating in more or less rounded ends. Polar capsules
large and open on opposite sides of spore. Coiled polar filament distinct.
Sporoplasm finely granular, symmetrically situated in spore. Smaller
spores differ from large ones chiefly in size; valves are much shortened
and have a greater curvature, with more distinctly rounded ends. Dimen-
sions: breadth 30m (longest 38^), sutural diameter 9/i, diameter of polar
capsules 4/*, length of polar filament 37/*. Small forms: breadth 15/*,
sutural diameter 7/*, diameter of polar capsules ^n.

CERATOMYXA ABBREVIATA Davis
[Fig. 61]
1917 Ceratomyxa abbreviala Davis 1917 : 227

Habitat: Gall-bladder of Scoliodon terrae-novae; Beaufort (August).

Vegetative form: Elongate, pyriform, with usually a very long, slender
posterior process. Body colorless. Progressive movements rapid. Dis-
tinct differentiation of protoplasm, posterior process usually composed of
ectoplasm (rarely endoplasm may extend into it for a short distance).
Pseudopodia, short, tapering or filiform at anterior end. Dimensions:
length up to 90/*, breadth 10 to 12/*, diameter of rounded sporulating
trophozoites about 27/t. Disporous.

Spore: Roughly crescent-shaped; sutural diameter exceptionally
great in comparison with the breadth. Ends of valves ^rounded, slightly
asymmetrical. Shell exceptionally tough and resistant to reagents. Polar
capsules large, prominent and open on opposite sides of spore. Sporoplasm
finely granular, confined entirely to one valve. Dimensions: breadth 17/*,
sutural diameter 14/*, diameter of polar capsules 4.5/*.

315] STUDIES ON MYXOSPORIDI A— KUDO 77

CER\TOMYXA FLAGELLIFERA Davis
[Fig. 62]
1917 Ceratomyxa flageUifera Davis 1917:227

Habitat: Gall-bladder of Carcharhinus sp?; Beaufort (July).

Vegetative form: Pyriform, short, tapering toward the posterior end,
sometimes dividing into a number of long, slender, transparent processes.
Extremely long filiform pseudopodia, developed at anterior end, can be
seen to sweep slowly back like a whiplash until they come to lie by the side
of the body. Progressive movements slow. Ectoplasm clear, transparent,
forming a distinct layer at anterior end. Endoplasm in large trophozoites
filled with large numbers of rod-shaped, bacteria-like bodies, which are more
abundant in the anterior half than in the posterior. Endoplasm in younger
trophozoites, with much less or without any bacteria-like bodies, shows a
vacuolated structure. Size up to 115 to I20fi in length and 40 to 45/* in
breadth. Disporous.

Spore: Valves greatly elongated, conical, with rounded ends. Sutural
ridge well marked. Polar capsules large, opening on opposite sides* of
spore. Coiled polar filament very distinct. Sporoplasm granular, symmet-
rically situated, but extending only a short distance into each valve.
Dimensions: breadth II8/1, sutural diameter 12/x, diameter of polar cap-
sules 6/i.

CERATOMYXA AGGLOMERATA Davis

[Fig. 63]

1917 Ceratomyxa agglomerata Davis 1917 : 228

Habitat: Gall-bladder of Synodus foetans; Beaufort.

Vegetative form: Pyriform, usually with long, slender, posterior
process. Body colorless and transparent. Actively motile, moving by
means of characteristic wavelike movements of the ectoplasm, from which
are projected numerous short, conical to filiform pseudopodia. Pseudo-
podia travel back along sides of body for about one-third its length and
then disappear, new ones being continually formed at the anterior end.
Ectoplasm distinguishable at anterior end. Endoplasm clear, very trans-
parent, usually homogeneous, sometimes finely granular. Large numbers
of fat globules usually present. Size of sporulating trophozoites 38/i by
12/i. Disporous.

Spore: Asymmetrical, one valve being smaller and more attenuated
than the other; larger valve compressed. Polar capsules spherical. Coiled
polar filaments indistinct. Sporoplasm filling nearly entire smaller valve,
but only extending a short distance into the larger one. Dimensions:
breadth 24 to 28ju, sutural diameter 5ju, diameter of polar capsules 3ju.

78 ILLINOIS BIOLOGICAL MONOGRAPHS [316

CERATOMYXA AMORPHA Davis
[Fig. 64]
1917 Ceratomyxa atnorpha Davis 1917 : 228

Habitat: Gall-bladder of Synodus foetans; Beaufort.

Vegetative form: Rounded or irregular in shape, with short lobose
pseudopodia; not pyriform; slowly ameboid. Body colorless. Ectoplasm
well developed, forming a distinct layer; transparent, finely granular.
Endoplasm granular, with large numbers of small fat globules scattered
through it or aggregated into one or two large clumps (difference between
the present form and C. agglomerata). Disporous.

Spore: Asymmetrical; crescent-shaped; valves short, conical, some-
what compressed. One valve distinctly smaller and more conical than the
other. Sutural ridge perpendicular to longitudinal axis. Polar capsules
large, opening at some distance from the anterior side. Coiled polar
filaments distinct. Sporoplasm granular, asymmetrically situated, being
chiefly confined to smaller valve. Dimensions: breadth 27/z, sutural
diameter 11/i, diameter of polar capsules 4ju.

CERATOMYXA MONOSPORA Davis

[Figs. 65 to 67]

1917 Ceratomyxa monospora Davis 1917 : 228-229

Habitat: GdJi[-\Adididitr oi Peprilus alepidotus; Beaufort. Abundantly
present in June, much less in July, being entirely absent in the bladder at
the end of the month.

Vegetative form: Pyriform, with a slender posterior process and one to
several filiform pseudopodia at anterior end. Body colorless and trans-
parent. Movements very slow. No clear differentiation between ecto-
plasm and endoplasm, the entire body being composed of a clear, finely
granular protoplasm. Fat globules more abundant in larger individual,
which are aggregated into small clumps. Size of vegetative trophozoites
up to 24/x in length and 15m in width. Monosporous form much smaller
than disporous. Monosporous and disporous. Nearly entire substance
of trophozoite is used up in spore formation.

Spore: Crescent-shaped. Valves cylindrical, tapering toward the end,
which is rounded and compressed. Curvature of valves varies. One
valve is more attenuated than the other. Sutural ridge perpendicular to
the longest diameter. Polar capsules large. Sporoplasm usually asym-
metrically situated. Dimensions: breadth 18 to 25/x, sutural diameter
5 to 6jLi, diameter of polar capsules 3/i.

Remarks: This species is evidently very close to C. pallida Th61.
Similar form was found in Prionotus evolans (gall-bladder), which showed
somewhat larger trophozoites and spores than C. monospora.

3171 STUDIES ON MYXOSPORIDJA—KUDO 79

CERATOMYXA STREPTOSPORA Davis
[Figs. 68 and 69]
1917 CeraUmyxa streptospora Davis 1917 : 229

Habitat: Gall-bladder of Chaetodipterus faber; Beaufort (June, but
not in July).

Vegetative form: Pyriform, colorless and transparent. A few conical,
filiform, wavelike pseudopodia at anterior end. Ectoplasm recognizable at
anterior end. Endoplasm finely granular, with a few, small, fat globules,
filled with transparent, homogeneous spherules. Size: 48 by 12/x to 60
by 9iJL. Disporous.

Spore: Compressed valves greatly elongated, with rounded ends.
Sutural ridge. Polar capsules spherical. Coiled polar filament indistinct.
Sporoplasm finely granular, entirely filling both valves. Dimensions:
breadth 34 to 39/x, sutural diameter 4)u, diameter of polar capsules 3/*.

CERATOMYXA AGGREGATA Davis
[Fig. 70]
• 1917 Ceratomyxa aggregata Davis 1917 : 229

Habitat: Gall-bladder of Leostomus xanthurus, Micropogon undulatus;
Beaufort (July).

Vegetative form: Form rounded to somewhat irregular in shape, rarely
pyriform; slowly ameboid. Body colorless and transparent. No clear
differentiation of protoplasm. Endoplasm finely granular, containing
numbers of small fat globules. Sporulating trophozoites show a tendency
to collect in groups composed of a large number of individuals so closely
associated that it is often impossible to make out the individual outlines.
Size of full-grown form 18/i by 14/i. Disporous.

Spore: Crescent-shaped; valves much elongated, tapering toward the
ends, which are compressed. Polar capsules spherical and opaque. Sporo-
plasm granular, situated symmetrically in the spore cavity. Dimensions:
breadth about 50jtt, sutural diameter 6 to 7/x, diameter of polar capsule
3.5ai.

CERATOMYXA UNDUE ATA Davis

[Fig. 71]
1917 Ceratomyxa undulata Davis 1917 : 230

Habitat: Gall-bladder of Ancylopsetta quadrocellata Gill.; Beaufort
(June to August).

Vegetative form: Pyriform, sometimes fusiform, tapering toward
posterior end. Movements rapid. Body colorless. Ectoplasm observable
at anterior part, constantly undergoes rapid, wavelike undulating move-
ments and extrudes fine conical or filiform pseudopodia. Pseudopodia

80 JLUNOIS BIOLOGICAL MONOGRAPHS [318

are formed very rapidly and vary in length. After reaching a considerable
length the pseudopodia usually travel posteriorly along sides of body
and then disappear. Endoplasm very transparent, often vacuolated,
containing numerous small fat globules. Size of full-grown trophozoite:
25/1 by 10 to 12/i in average. Disporous.

Spore: Crescent-shaped. Valves cylindrical, not compressed, ends
rounded, one valve being somewhat longer and more conical than the other.
Polar capsules convergent. Coiled polar filaments distinct. Sporoplasm i

granular, asymmetrically situated, sometimes being almost confined to
more conical valve. Dimensions: breadth 22 to 24m, sutural diameter 6/i,
diameter of polar capsules 3/x.

CERATOMYXA NAVICULARIA Davis

[Figs. 72 and 73]
1917 Ceratomyxa navicularia Davis 1917 : 230

Habitat: Urinary bladder of Paralichthys dentatus, P. albiguttus,
Sphaeroides maculatus; Beaufort (June to August).

Vegetative form: Rounded or slightly irregular in shape, never pyri-
form. Body colorless. Very slow ameboid. No distinct ectoplasm.
Entire trophozoite finely granular, containing a few small fat globules.
Nearly entire body is used up in the formation of spores. Diameter about
17/1. Disporous.

Spore: Variable in shape and size. Symmetrical or asymmetrical,
often boat-shaped, slightly compressed dorso-ventrally, with rounded
ends. Polar capsules convergent, shows polar filament indistinctly.
Sporoplasm finely granular, extending into both valves, but usually
somewhat farther into one than the other. Dimensions: breadth 14 to 22/x
(average 16/Lt), sutural diameter 5 to 7.5m (average 6m), diameter of polar
capsules 2m.

CERATOMYXA SPINOSA Davis

[Fig. 74]

1917 Ceratomyxa spinosa Davis 1917 : 230

Habitat: Urinary bladder of Paralichthys albiguttus; Beaufort.

Vegetative form: Rounded or slightly irregular in shape, with short,
lobose pseudopodia; slowly ameboid. Body colorless and transparent.
Distinct differentiation of protoplasm along the entire surface, ectoplasm
forming outer layer. Endoplasm faintly granular, with numerous small
fat globules. Monosporous and disporous.

Spore: Central portion greatly enlarged; ovoid, with very long tapering
process extending out from each end. Sutural line perpendicular to the
longest diameter. Polar capsules large and spherical. Sporoplasm finely

319] STUDIES ON MYXOSPORIDIA—KUDO 81

granular, chiefly located in one valve, extending into the other only a short
distance beyond the capsule. Dimensions: breadth 80/*, breadth of
enlarged central portion IS^t, sutural diameter 7/*, diameter of polar cap-
sules 4)Lt.

Genus MYXOPROTEUS Doflein emend. Davis

1898 Myxoproteus Doflein 1898 : 287

1917 Myxoproteus Davis 1917 : 219

The characters of the genus are described on page 56.
Type species: Myxoproteus ambiguus (Thelohan) Doflein.

MYXOPROTEUS AMBIGUUS (Thelohan) Doflein
[Figs. 75 to 80]

1895 Myxosoma atnbiguum Th61ohan 1895 : 344

1898 Myxoproteus ambiguus Doflein 1898 : 287-288

Habitat: Urinary bladder of Lophius piscatorius; Le Croisic, Rovigno,
Napoli.

Vegetative form: Polymorphous. Color milky white. Protoplasm is
filled with numerous granules and fat globules. Pseudopodia, short,
pointed lobose. Plasmogamy and plasmotomy take place. Many small
individuals formed apparently by plasmotomy, often, make up groups.
Disporous, polysporous?

Spore: Almost pyramidal, with anterior processes. Two very large
polar capsules at the anterior end. The distance between these capsules
is equal to or greater than the diameter of the capsules. Sporoplasm with
two nuclei. Dimensions: length 25/i, breadth 18 to 20/x, diameter of polar
capsules 7/i.

MYXOPROTEUS CORDIFORMIS Davis

[Figs. 81 to 83]

1917 Myxoproteus cordiformis Davis 1917 : 231

Habitat: Urinary bladder of Chaetodipterus faber; Beaufort (June,
July).

Vegetative form: Rounded; very slowly ameboid, usually forming a
single, short, lobose pseudopodium. Body colorless and transparent.
Ectoplasm not distinct. Entire trophozoite finely granular, with a few
fat globules. Rarely vacuolar. Small trophozoites often show a single
large, central vacuole. Rounded sporulating trophozoites 18/t in diameter.
Disporous.

82 ILLINOIS BIOLOGICAL MONOGRAPHS [320

Spore: Heart-shaped in front view, with peculiar wing-like expansions
on each side which contain remains of parietal cells. Sutural plane oblique
in position. Capsulogenous cells distinct. Sporoplasm finely granular,
fills the extracapsular cavity of the spore. Dimensions: length \2n,
breadth 10 to ll/x, thickness 6/1, polar capsules 3 to 4/i in diameter.

MYXOPROTEUS CORNUTUS Davis
[Fig. 84]

1917 Myxoproteus corntdus Davis 1917 : 231

Habitat: Urinary bladder of Bairdiella chrysura; Beaufort.

Vegetative form: Somewhat elongated or irregular in shape, with
short lobose pseudopodia; slowly ameboid. Differentiation of protoplasm
clear. Ectoplasm well developed, hyaline; in rounded individuals forming
a distinct layer around the body. Endoplasm opaque, contains coarse
refringent granules varying in shape, and a few fat globules. In contracted
rounded resting condition, endoplasm becomes condensed, while ectoplasm
appears more abundant. Rounded trophozoites up to 27^ in diameter.
Disporous.

Spore: Heart-shaped, with two anterior processes. Shell very thick.
Polar capsules large, opening some distance apart. Coiled polar filament
distinct. Sporoplasm finely granular, with a few small fat globules, fills
the extracapsular cavity of the spore. Dimensions: sutural diameter
exclusive of the processes 9/u, breadth 12)u, length of processes 5n, diameter
of polar capsules 3^1.

Genus WARDIA nov. gen.

The characters of the genus are described on page 56.
Type species: Wardia ovinocua nov. spec.

WARDIA OVINOCUA nov. spec.

[Figs. 85 to 95]

Habitat: Ovum and connective tissue of ovary of Lepomis humilis
Girard;* Salt Fork, 111. (November). Only one fish, 6.5 cm. long with
normal appearance, was found to be infected.

Vegetative form: Trophozoites form cysts visible to the naked eyes
as white spherical spots in the pink-colored ovary. Four cysts present.
The cyst (Figs. 85 and 86), in section, shows a circular form surrounded by
several layers of hypertrophied nurse cells and connective tissue, in which
many large blood vessels are present. Protoplasm is not clearly differ-

*Professor F. Smith of the Department kindly identified all the fish that were collected
in the vicinity of Urbana and mentioned in this paper as hosts, for which the writer wishes to
express his appreciation.

321] ■ STUDIES ON MYXOSPORIDIA— KUDO 83

entiated into ectoplasm and endoplasm, the whole protoplasm showing
granulated reticular structure. Cysts contained numerous fully developed
spores and a small number of spores in developmental stages, which sug-
gested the fact that two spores rise from each pansporoblast. The parasite
is also found in the state of diffuse infiltration in the connective tissue
around the cyst. Diameter of cysts 316 to 445/i in sections. Polysporous.
Spore: In front view, isosceles triangular form, two sides of which
usually convex, with more or less attenuated anterior end (Figs. 87, 90,
92); in profile, ellipsoidal (Fig. 88); and oval viewed from the anterior
end (Fig. 89). Sutural plane at right angles to the longest diameter (Figs.
87 and 89), Shell comparatively thin except at the anterior end and has
many fine network-like ridges on the surface. These ridges are hardly
observable on fresh spores on account of their fine form and the conspicu-
ously large polar capsules lying in the spore. When stained, however, they
are not only made distinctly visible, but the prolongation of each ridge
from the posterior edge wbiph forms about l/x long fringe-like structure is
also more clearly recognized (Figs. 90-95). Two large and spherical polar
capsules located in the central portion of the spore. Coiled (5 to 6 times)
polar filament extremely distinct. The openings of polar capsules at the
anterior end. Sporoplasm comparatively small, finely granular, without
any vacuole, contains two small nuclei, when stained. Dimensions in vivo:
sutural diameter 9 to lO/x, breadth 10 to 12/i, thickness 6ju, diameter of the
polar capsule 4/i, length of polar filament 35 to 45/x.

WARDIA OHLMACHERI (Gurley) Kudo

[Figs. 96 and 97]

1893 Myxosporidian Ohlmacher 1893 : 561-567

1893 Chloromyxum ohlmacheri Whinery 1893 : 660-662

1894 Chloromyxum {Spkaerospora)

ohlmacheri Gurley 1894 : 267-272

1895 ? Leptotheca ranae Th61ohan 1895 : 383
1899 Leptotheca ohlmacheri Labh€ 1899 : 87

Habitat: Urinary tubules of kidney of Bufo lentiginosus Shaw and
kidney of Rana esculenta and R. temporaria {R. fusca) ; Sycamore, De Kalb
county. 111.

Vegetative form: Not found.

Spore: Transversely elliptic. Sutural plane perpendicular to the longer
axis of the spore. A well defined undulate-parallel longitudinal striation
on the shell. Sutural ridge comparatively well marked. Two polar capsules
lying side by side, occasionally only one. Dimensions: sutural diameter
6ju, breadth 8/i, diameter of polar capsule 3 to 3.5/:, length of polar filament
6 to 8 times the breadth of spore (48 to 64/i).

Remarks: This form is apparently very much different from any
species of genus Leptotheca, in the general form, form of polar capsules,

84 ILLINOIS BIOLOGICAL MONOGRAPHS [322

striations on the shell and the habitat. Tho the form of the spore is differ-
ent from the type species of the genus Wardia and nothing is known about
the vegetative form, the presence of large spherical polar capsules in the
central portion of the spore, the striations on the shell and the occurrence
of the same nature, i.e., from fresh waters in the close-by localities, show
its nearer relationship to the genus Wardia than to the genus Leptotheca.
Hence it is placed here provisionally.

Genus MITRASPORA Fujita emend. Kudo
1912 Mitraspora Fujita 1912 : 259-260

The characters of the genus are described on page 56.
Type species: Mitraspora cyprini Fujita.

MITRASPORA CYPRINI Fujita

[Figs. 98 to 104]

1912 Mitraspora cyprini Fujita 1912 : 259-260

Habitat: Renal tubules of kidney and ureters of Cyprinus carpio L.
and Carassius auratus L. ; Sapporo (winter), Tokio (March).

Vegetative form: Fujita's only description is as follows: "The sporo-
blast contains generally three or four spores." The present writer observed
a similar form in the ureter and the renal tubule of the kidney of Cyprinus
carpio L., in Tokio, The observations are as follows: Trophozoites small
ameboid (Fig. 98). Body colorless. Movements tardy. Differentiation
of protoplasm imperfect. The hyaline ectoplasm recognizable at one side
of the body, where lobose pseudopodia are formed (Figs. 98-99). Endo-
plasm granular with vacuoles and brownish granules, which become
larger as the body grows. Size 10 to 40/i. Disporous (Kudo) and poly-
sporous (?, Fujita).

Spore: Fujita's descriptions are as follows: Form resembles monk's
hood, slightly attenuated at its anterior end. Shell uniformly thin, except
at two points of the truncated posterior end. Each shell valve has eight
distinct striations which run longitudinally and turn into long cilia up to
5.8jLi long, planted in a single row at the posterior end of the spore. Two
polar capsules at the anterior end. The nucleus is obscure and no vacuole
is present. Dimensions: length 10 to 13/x, breadth Sn, polar capsules
3.8/x by 2n, length of polar filament 15/i (weak glycerine). The writer
observed the following facts: More rounded with rounded anterior end in
front and side views. Shell more or less thick along the entire posterior
margin. Striations on shell, variable in number. Sporoplasm granular,
without any vacuole, exhibits two nuclei when stained. Posterior filaments
5 to 6 in number and 5 to 6/i long, being absent in some spores. Dimen-
sions in vivo: length lO/x, breadth 8 to 9/i, thickness 6 to S/x, polar capsule
4n by 1.5 to 2/i, length of polar filament 35 to 40/i.

323] STUDIES ON MYXOSPORIDIA—KUDO 85

Remarks: Tho Fujita does not describe the vegetative form and there
are some differences in the form and size of the spore between the forms,
the writer does not find out any objection against the union of the above
mentioned two forms.

f

MITRASPORA CAUDATA (Parisi) Kudo

[Figs. 105 to 107]

1910

Sphaerospora caudata

Parisi

1910

: 253-254

1912

Sphaerospora caudata

Parisi

1912

:289

1913

Sphaerospora caudata

Parisi

1913

: 396-402

Habitat: Renal tubules of kidney of Alosa finta Cuv. var. lacustris
Fatio; Lake Como.

Vegetative form: Rounded or variously elongated owing to the move-
ments. Protoplasm is distinctly differentiated into ectoplasm and endo-
plasm. Ectoplasm, hyaline and homogeneous, forms slowly moving lobose
pseudopodia. Endoplasm granular, contains yellow globules and fat
granules. Disporous and polysporous.

Spore: Subspherical in front view; oval in profile; anterior end being
more rounded than the posterior end. Shell rather thick, longitudinally
striated. In front view, the posterior end enlarged into a quadrangular
form, which appears as a small spine in side-view and which projects
backward long and fine filaments, usually six in number. Two well devel-
oped polar capsules open on each side of the sutural plane. Polar filament
coiled 5 to 6 times. Sporoplasm without any iodinophilous vacuole.
Dimensions: external length 10 to llju, internal length 7 to 9n, length of
polar capsules 4 to 4.5)u, length of polar filament up to 48/x, length of pos-
terior filaments up to 28/i.

MITRASPORA ELONGATA nov. spec.
[Figs. 602 to 621]

Habitat: In the urinary tubules and tissue of kidney of Lepomis
cyanellus; Crystal Lake, Urbana, 111. From June to July, all the fish
examined, 36 in number and 10 cm. in average length, were found to be
infected. Other fish such as Lepomis pallidus and Lepomis humilis, caught
at the same time, were free from the infection. Early in June, the number
and size of the parasites in a host body were rather small and only a small
number of spores could be recognized in the fresh state with the addition
of potassium hydrate solution. The growth of the parasite was rather
remarkable during the hot weeks in the latter part of June and July so
that every fish caught on July 17th showed a heavy infection, exhibiting
small whitish pustules over the surface of the organ. During June,
vegetative forms and spores were found in the lumen of the urinary tubules.

86 - ILUNOIS BIOLOGICAL MONOGRAPHS [324

altho some contained the parasitic masses in the tissue. About the middle
of July, the parasite forms conspicuous cysts in the tissue thruout the
organ. The c)^t may or may not be covered by a thick layer of connective
tissue from the host. Aside from this hypertrophy, the host did not show
any pathological change which covdd be recognized. »

Vegetative form: Youngest trophozoite found in the urinary tubule
is multinucleate, rounded, and of from 20 to 50ai in diameter. The proto-
plasm is not differentiated, the entire body is finely granular or coarsely
reticular in structure. In the protoplasm are to be seen nuclei and sporo-
blasts at different stages of development. The union of two propagative
ceUs similar to that of Myxoholus toyamai produces a small body which
developes into a single sporoblast and ultimately into a single spore
(Figs. 605-613). In later stages, the trophozoite reaches a size of 200/1
in diameter showing many stages of spore formation and mature spores,
surrounded by thick layers of connective tissue from the host. Poly-
sporous.

Spore: Elongated oblong with pointed anterior and truncated posterior
extremities. The width is often greatest at the middle of the polar cap-
sules, the posterior portion is much narrower than the anterior. Nearly
circular in the cross-section thru the polar capsules. The shell is thin,
the sutural line being faintly marked in fresh state. It generally is
obliquely located in relation to the capsules. The shell also shows fine
longitudinal striations, 14 to 16 in number, on each valve. The sutural
line as well as the striations are best seen in spores stained with Heiden-
hain's iron hematoxylin. Two polar capsules elongated pyriform, mostly
equal in size, occupy the anterior half of the spore. Abnormal situations
of the polar capsules are sometimes observed (Fig. 619). The coiled polar
filament is faintly visible in fresh spores. It is spirally coiled along the
wall of the polar capsule without any central axis. This fact was clearly
observed in stained section as is shown in Figs. 620 and 621. The fila-
ment has seven or eight windings, thus agreeing with the actual length
of the extruded polar filament. The polar filament was extruded under
the action of potassium hydrate solution. The extrusion takes place
even in some spores which were treated with Schaudinn's fixative and
kept in 95 per cent alcohol for three months (see the similar observations
on Myxoholus discrepans on page 157). The sporoplasm is finely granular
and transparent. When stained, it shows two nuclei in the center or near
the posterior part of the body. Dimensions of preserved spores: length
15 to 17/ii, breadth 5 to 6/i, thickness 4.5 to 5.5/z, polar capsule 7.5^ by 2/x,
length of polar filament 40 to 50/Lt.

Suborder SPHAEROSPOREA nom. nov.
The definition of the suborder is recorded on page 57.

325] STUDIES ON MYXOSPORIDIA—KUDO

Family CHLOROMYXIDAE Thelohan

1892 Chloromyxidees Thfilohan 1892 : 173

1895 ChloromyxidSes Thaohan 1895 : 344

The characters of the family are described on page 57.

Genus CHLOROMYXUM Mingazzini

1890 Chloromyxum Mingazzini 1890 : 160

1892 Chloromyxum Thdlohan 1892 : 173-176

1895 Chloromyxum Thflohan 1895 : 344

The characters of the genus are described on page 57.
Type species: Chloromyxum leydigi Mingazzini.

CHLOROMYXUM LEYDIGI Mingazzini
[Figs. 108 to 113]

87

1851

Leydig

1851

: 225-234

1852

Leuckart

1852

:435

1854

Lieberkiihn

1854

:352

1890

Chloromyxum leydigi

Mingazzini

1890 :

: 160-164

1892

Chloromyxum leydigi

Thelohan

1892 ;

: 166, 169-170

1894

Chloromyxum leydigi

Chloromyxum incisum

Gurley

1894;

: 25^260

1895

Chloromyxum leydigi

Chloromyxum incisum

Thelohan

1895 :

: 345-346

1898

Chloromyxum leydigi

Doflein

1898 :

: 292, 310, etc.

1912

Chloromyxum leydigi

Erdmann

1912

: 149-162

1916

Chloromyxum leydigi

Georg6vitch

1916a

:3

1917

Chloromyxum leydigi

Davis

1917

: 236-237

1917

Chloromyxum leydigi

Erdmann

1917

: 276-321

1918

Chloromyxum leydigi

Georgfivitch

1918

: 182-189

Habitat: Gall-bladder of Rhina squatina L., Spinax spinax L., ScylUum
canicula, S.asterias,Raja batis L., i?. clavata L., R. undulata Lac, Torpedo narce
Ris., T.marmorata, T.ocellata, T. torpedo 'L.,Acanthias acanthias L,.,Trygon
pastinaca L., Dasybatis hastatus, D. sabina, Pteroplatea machira Le Sueur,
Scoliodon terrae-novae, Cestracion zygaena, C. tiburo, Carcharhinus limbatus;
Roscoflf, Concarneau, Marseille, Banyuls, Rovigno, Heligoland, Beaufort,
Monaco (May), Napoli, Genova. Erdmann observed the species at
Naples from March to August. She noticed mixed infection with Cerato-
myxa reticulata and especially with Leptotheca parva. Georgevitch studied
the parasite at Monaco from February to April.

Vegetative form: Polymorphous, being spherical, oval or irregular.
The change of the form rather rapid under favourable conditions. Differ-
entiation of protoplasm distinct. Ectoplasm with pseudopodia of various
form, i.e., lobose, filiform or intermediary; short, pointed or branched.
Endoplasm alveolar, filled with yellowish granules. Doflein observed the
plasmotomic multiplication of young trophozoites. Polysporous. Erd-
mann's observations (1917) may be summarized as follows: Ameboid.

88 II.UNOJS BIOLOGICAL MONOGRAPHS [326

Color of the body greenish to dark green. The protoplasm is clearly dif-
ferentiated into ectoplasm and endoplasm. The ectoplasm is hyaline and
covers the entire surface of the body. It appears as a fine fibrous structure
when fixed with Bouin's solution. The endoplasm contains besides nuclei,
two kinds of spherules; one smaller and yellowish "color-carriers" and the
other larger and light to dark greenish reserve bodies. The color-carrier
is in part composed of myelin, while the reserve body is of glycogenous
nature. The infection was studied experimentally per os: young tropho-
zoites appeared in 3 to 5 days which continued to 10th day, various tropho-
zoites were seen in 13 to 19 da}^, and sporulating individuals were first
recognized in 39 days after the infection. The trophozoite multiplies in
number either by fission or by budding. It usually contains enclosures
which seem to be degenerating erythrocytes. Mictosporous.

Spore: Ovoidal. Shell- valves show wide edge at sutural plane, which
is attenuated at the anterior end and forms a quadrilateral process at the
posterior end, from which a row of ciUa grows. Shell-valves have ridges
(6 to 7), which run parallel to the posterior margin. The striations may
vary considerably. Four polar capsules at the anterior end. Dimensions:
length 8/i. Erdmann gave the following dimensions: Spores from Torpedo
marmorata and T. ocellata: length 6 to 9yL, breadth 5/i, polar capsule 3^
by 2/i. Those from Scyllium asterias: length 8 to 9n, breadth 6/x, polar
capsule 2m by 1/u. Those from Raja hatis: length 7 to 8//, breadth 5m,
polar capsule 2n by 1^. Length of polar filament 20 to 30m (absolute alco-
hol warmed up to 40** C).

CHLOROMYXUM CAUDATUM Thelohan
[Fig. 114]
1895 Chhromyxum caudatum Th61ohan 1895 : 346

Habitat: Gall-bladder of Molge cristata Laur.; Vicinity of Rennes.

Vegetative form: Body yellowish with lobose pseudopodia. Proto-
plasm finely granular.

Spore: Oval or spheroidal. Shell enlarged at the anterior part, having
a simple or bifurcated tail-like process, as in Henneguya, at the posterior
end. Dimensions: total length 18m, length 8m, breadth 6 to 7m, length of
tail 10m.

CHLOROMYXUM QUADRATUM Thelohan

[Figs. 115 to 117]

1891

Keiffer

1891

111

1893

Pfeiffer

1893

81

1895

Chlorotnyxum quadratum

Thaohan

1895

347

1912

CMoromyxum quadratum

Parisi

1912

289

1913

Chloromyxum quadratum

Awerinzew

1913a

:155

1913

Chhromyxum quadratum

Fennor

1913

199

327] STUDIES ON MYXOSPORJDIA—KUDO 89

Habitat: Muscle of Syngnathus acus L., Trachurus trachurus L.,
Nerophis aequorens L., Callionymus lyra L., Coris julis L., Ariodes polysta-
phylodon, kidney of Blennius gattorugine Brunn; Helder, RoscoflF, Concar-
neau, Marseille, Beira (Africa), Napoli (summer).

Vegetative form: Not described by any of these authors.

Spore : Quadrangular pyramid with curved edges and roundish angles.
Four polar capsules at the anterior end. Dimensions: length 6)u, breadth
S/i, length of polar filament 8 to 10^.

CHLOROMYXUM FLUVIATILE Thelohan
[Fig. 118]

1892 Chloromyxum fluviatile Thdlohan 1892 : 173-176

1895 Chloromyxum fluviatile Th61ohan 1895 : 346

Habitat: Gall-bladder of Leuciscus cephalus L.; Paris.

Vegetative form: Young trophozoites colorless; adults yellowish.
Form highly variable. Active change of the form of body. Clear diflFer-
entiation between ectoplasm and endoplasm. Ectoplasm usually recog-
nizable at one end of the body where lobose pseudopodia are formed.
Size reaches 25 to 30^. Polysporous.

Spore: Spherical, generally small. Sutural ridge fairly well marked.
Dimensions: 7 to 8/i in diameter.

CHLOROMYXUM MUCRONATUM Gurley

[Figs. 119 to 122]

1854
1879
1882
1883 ■

1893 Chloromyxum mucronatum

1894 Chloromyxum mucronatum

1908 Chloromyxum mucronatum

1909 Chloromyxum mucronatum

Habitat: Urinary-bladder and kidney of Lota lota L.; Bodensee, other
localities not mentioned.

Vegetative form: Spherical, elliptical or irregular. Size up to 75/ii
in diameter. Protoplasm containing irregularily scattered fat-like globules.

Spore: Sharp-contoured; subglobular, mucronate anteriorly. Dimen-
sions: length 8/z.

Lieberkiihn

1854;

: 352-353

Leuckart

1879 ;

:248

Butschli

1882 ;

: PI. 38 : 17

Balbiani

1883 ;

; 201, 203

Gurley

1893 :

:419

Gurley

1894;

: 264, 265

Auerbach

1908 ;

:456

Auerbach

1909a

:71

Balbiani

1866 : 600-602

Balbiani

1867 : 275,276, 335

Butsrhli

1882 : 590

Keiffer

1890 : 559

Henn^iuy et Th€Iohan

1892 : 587

Gurley

1893 : 411

Gurley

1894 : 281

Thaohan

1895 : 347

90 ILLINOIS BIOLOGICAL MONOGRAPHS [328

CHLOROMYXUM DIPLOXYS (Gurley) Thelohan
[Figs. 123 to 125]

1866
1867
1882
1890
1892

1893 Cystodiscus ? diploxys

1894 Cystodiscus ? diploxys

1895 Chloromyxum diploxys

Habitat: Abdominal cavity of Tortrix viridana L. (imago);

Vegetative form: Trophozoites form spherical cysts, 230 to 400ju
in diameter. Cyst membrane rather thick. Protoplasm containing
brownish granules, and fat-like globules (red with iodine).

Spore: Elliptic or slightly flattened. Sutural line straight, forming a
ridge. Two polar capsules at each end.

CHLOROMYXUM PROTEI Joseph

1905 Chloromyxum protei Joseph 1905 : 450-451
1907 Chloromyxum protei Joseph 1907 : 398-412

Habitat: Renal tubules of kidney of Proteus anguineus L.; Vienna.

Vegetative form: Generally rounded or sausage form. No clear
differentiation between ectoplasm and endoplasm. Movements slow.
Probable occurrence of plasmotomy by budding and division. Size:
40 to 45m by 28 to 40m.

Spore: Spherical. Shell finely striated parallel to the sutural line.
Four polar capsules each with an independent opening. Dimensions:
10 to 13m in diameter, polar capsules 4 to 6m long. The polar filament
appears to be rather short.

CHLOROMYXUM TRUTTAE Leger

[Fig.126]

1906 Chloromyxum truUae L6ger 1906 : 267-270
Habitat: Gall-bladder and gall-duct of Truttafario L.; Dauphine.
Vegetative form: Ameboid form. Elongated. Form resembles an

Amoeba Umax of about 40m in length. Roundish or irregularly contoured,
with small pseudopodia. Ovoidal or spherical, 25 to 40m in diameter
without any visible pseudopodia (resting state). Body colorless, clear and
hyaline. Very active movements which last for several hours after the
death of the host. Broad and obtuse pseudopodia well developed at the
anterior end of the body. Endoplasm alveolar, contains variable numbers
of nuclei, which are seen in vivo, refractive bodies and chromatic granules.
Monosporous(?) and polysporous.

Spore: Spherical. Four polar capsules of different size. Shell- valves
marked with parallel ridges. Dimensions: 8 to 9m in diameter.

329] STUDIES ON MYXOSPORIDIA—KUDO 91

CHLOROMYXUM CRISTATUM Leger

[Figs. 127 and 128]

1906 Chloromyxutn crishUum L^ger 1906 : 270-272

Habitat: Gall-bladder of Tinea vulgaris Cuv.; Grenoble.

Vegetative form: Ordinarily massive, with oval or round contours,
without noticeable pseudopodium. Ectoplasm hyaline. Endoplasm
granular and colorless. Average diameter of the adults about 20^. Mono-
sporous, rarely disporous.-

Spore: Spherical or subspherical. Ten marked ridges run antero-
posteriorly on each shell- valve, so that it presents a cog-wheel form in cross
section. Four polar capsules at the anterior end, one pair being smaller
than the other. Sporoplasm with two nuclei. Dimension: 10 to ll/u.

CHLOROMYXUM DUBIUM Auerbach
[Figs. 129 to 133]
1908 Chloromyxutn dubium Auerbach 1908 : 456-459

1910 Chloromyxutn dubium Auerbach 1910c : 177

Habitat: Gall-bladder of Lota vulgaris Cuv.; Bodensee (April to Sep-
tember).

Vegetative form: Spherical or rounded. Rarely irregular forms.
Protoplasm is dififerentiated distinctly into ectoplasm and endoplasm.
Ectoplasm very thin, forms pseudopodia which move slowly. Endoplasm
granular, contains fat globules. Majority of the trophozoites appear to
live floating in the bile, while some are attached to the epithelium of the
bladder. Disporous and polysporous.

Spore: Spherical, with four polar capsules. Each shell valve has
longitudinal ridges, variable in number (6 ridges are found on the drawing),
which run parallel to the sutural line. Four polar capsules of nearly same
size and convergent. Sporoplasm finely granular with two nuclei. Dimen-
sions: diameter 10.8/i, length of polar capsule 3.6/i.

CHLOROMYXUM sp. Awerinzew

[Fig. 134]

1908 Chloromyxum sp. Awerinzew 1908 : 43, 47, 48

Habitat: Gall-bladder of Raja radiata; Murman coast?.

Vegetative form: Form rounded. The protoplasm is distinctly differ-
entiated into ectoplasm and endoplasm. Ectoplasm hyaline and compara-
tively abundant in quantity compared with endoplasm, forms lobose
pseudopodia of active movements. Endoplasm vacuolated, contains
enclosures. Between the two layers, a thin layer of protoplasm, reticular in
structure and stained deeply with hematoxylin, is present.

Spore: No figure.

92 ILLINOIS BIOLOGICAL MONOGRAPHS [330

CHLOROMYXUM THYMALLI Lebzelter

1912 Chloromyxum thymalli Lebzelter 1912 : 295-296
Habitat: Gall-bladder of Thymallus thymallus L.; Vienna?
Vegetative form : Irregular form, ?)2) to 35ju long in average. Endoplasm

contains fat globules which stain brown with carmine. Trophozoites
attached to the epithelium. In average, 6 spores formed in each individual.
Intracellular stage in the epithelial cell is supposed. Polysporous.

Spore: Spherical. Shell structure similar to C. protei, but ridges are
more developed and exhibit somewhat wavy courses. Polar capsules of
equal size. Dimensions: 9 to 9.5ju in diameter, polar capsules 3^.

CHLOROMYXUM KOI Fujita
[Fig 135]

1913 Chloromyxum koi Fujita 1913 : 257-259

Habitat: Gall-bladder of Cyprinus carpio L.; Sapporo (Nippon).

Vegetative form : Spherical, with greatest diameter lip to SOju, contain-
ing 1 to 3 spores. Each spore is situated in a clear space surrounded by a
membranous envelope (sporoblast?), around which there is some finely
granular matter (endoplasm?).

Spore: Spherical, exhibiting a somewhat angular contour at the ante-
rior end. Shell thick and has well marked ridges on the surface, i.e., 4 to 5
circular ridges and on both sides of these ridges, two more ridges each bent
in a loop-like manner, so that the outline of spore in cross section, is very
much like of a toothed wheel with nearly equidistant teeth, 16 to 18 in
number. Four polar capsules, two slightly larger than the other two.
Dimensions: length 16)u, breadth lO/x, length of polar capsule 4/i, length of
polar filament 64/x.

CHLOROMYXUM MAGNUM Awerinzew

[Figs. 136 to 138]

1913 Chloromyxum magnum Awerinzew 1913 : 155-156

Habitat: Gall-bladder of Acanthias blainvillei;* Algoa Bay, East Lon-
don, Liideritzbucht (Africa).

Vegetative form: Ameboid. Body yellowish by the presence of large
yellowish granules in endoplasm. Often round or rosary form. Pseudo-
podia sometimes absent, so that the trophozoites move like Amoeba Umax
with a cluster of small, hairy pseudopodia at the posterior end. In larger
form, small round pseudopodia, composed of homogeneous ectoplasm, are
formed. Plasmotomj'^ by budding, was often observed. Usually polyspor-
ous, rarely monosporous.

Spore: Elongated spherical form. Four polar capsules at the narrow,
anterior end. Sporoplasm with two nuclei. Dimensions: length 40 to
48/1, breadth 30 to 38/x, length of polar capsules 12 to ISfi.

* Misprinted in Awerinzew's paper as blainvilei.

331] STUDIES ON MYXOSPORJDIA—KUDO 93

CHLOROMYXUM FUNDULI Hahn
[Figs. 139 and 140]

1915 Chloromyxum funduli Hahn 1915:205-206

Habitat: Muscle of Fundulus sp.; Woods Hole. In one fish.

Vegetative form: Hahn made observations on few fresh and stained
smears. According to him, it is clear that the staining was abnormal.
It is hard to quote this here as he used different terms without giving any
definition. The reader is advised to consult Hahn's paper.

Spore: Form slightly resembles that of Choloromyxum quadratum.
Posterior end rounded, the anterior portion narrow and truncated at the
tip; optical cross-section thru the posterior part of the polar capsules,
circular. Four polar capsules at the anterior end. Dimensions: height
(length) 6)u, breadth and thiclcness 7.5^ respectively.

Remarlis: As to the comparison of the present species with Chloro-
myxum clupeidae Hahn, see p. 94.

CHLOROMYXUM MISGURNI Kudo
[Figs. 141 to 146]

1916 Chloromyxum tnisgurni Kudo 1916 : 6-7

Habitat: Gall-bladder of Misgurnus anguillicaudatus Cantor; Tokio
(September).

Vegetative form: Round or irregular. Semicircular when viewed from
side. From the flat surface, many fine root-like, filiform pseudopodia are
extruded. No clear differentiation between ectoplasm and endoplasm.
Endoplasm alveolar. Trophozoites always found attached to the lining
epithelial cells. Size up to 50/x by 20/i. Polysporous (6 to 8 spores), rarely
disporous.

Spore: Spherical, slightly attenuated at the anterior end. Sutural line
straight and forms a ridge. Fine longitudinal striations run parallel to
the sutural line. Four polar capsules at the anterior end. Sporoplasm
finely granular, has two nuclei of equal size. Dimensions: length 8 to
9)u, breadth 6 to 7/x, thickness 5 to 6^, length of polar capsule 2 to 3ai,
of polar filament 28 to 35m (KOH).

Remarks : The host is often infected at the same time by Chloromyxum
fujitai, the trophozoites of which can be distinguished from the present
form by the structure and the floating habit in the bile. Spores in the
two species are decidedly different in form, structure and size.

CHLOROMYXUM FUJITAI Kudo
[Figs. 147 to 152]
1916 Chloromyxum fujitai Kudo 1916:7-9

Habitat: Gall-bladder of Misgurnus anguillicaudatus Cant.; Tokio,
(5% of the fish examined in September, found infected).

94 ILLINOIS BIOLOGICAL MONOGRAPHS [332

Vegetative form: Round or irregular. No clear differentiation of
protoplasm. Endoplasm highly vacuolated. Ectoplasm being hardly
distinguishable. Size up to 40/t in diameter. Trophozoites float in the
bile in almost all cases. Disporous and polysporous (up to 8 spores).

Spore: Spherical, often attenuated at the anterior end. Sutural line
not straight. Shell very thick, shows thick ridges running longitudinally
on the surface. In optical cross section, the spore presents an outline like
a cog-wheel with 20 to 22 ridges. The thickness of ridges varies regularly;
the thickest ones being located on two lines where a plane perpendicular
to sutural plane cuts the shell longitudinally, others decreasing in thick-
ness as they approach the sutural line. Four polar capsules at the anterior
end. Sporoplasm with two nuclei. Dimensions: length 10 to 12jLt, breadth
8 to 10/i, polar capsules 2 to 3n, length of polar filament 23 to SOn (KOH).

CHLOROMYXUM CLUPEIDAE Hahn
[Figs. 153 to 156 and 562 to 565]

1900 Sporozoa Tyzzer 1900 : 66-^

1901 Sporozoa Linton 1901 : 438
1910 Chloromyxum sp. Auerbach 1910 : 178
1917 Chloromyxum dupeidae Hahn 1917 : 15-19

Habitat: Body musculature of Clupea harengus, Pomolobus pseudohar-
engus, P. aestivalis, P. mediocris, Brevoortia tyrannus, Stenototnus chrysops,
Tautogolahrus adspersus; Woods Hole.

Tyzzer mentioned in his paper and also in a letter to the writer that
he collected the material in August of 1900 and that he found the infection
occurred only among young fish. Hahn also called attention on the latter
fact.

Vegetative form: Hahn's observations are as follows:

Clusters of spores ("pseudocysts") are spindle-shaped, especially when
young, usually l5ang between the bundles of muscle fibres. Color white or
creamy. Larger ones usually "in pocket just beneath the integument."
Schizogonic multiplication probably exists. Parasites hard to stain,
anilin dyes being unable to stain at all. Large form (probably composed of
many individuals) 890ju by 30jli.

Tyzzer described as follows: Cysts up tO" 1 to 2 mm. in length, lying
between the muscle fibres of the myotomes, surrounded at times by mem-
braneous connective tissue. The parasites also occur in diffused infiltration.

Linton found two cysts, 1.74mm. by 1.16mm. and 1.16mm. by 0.58mm.
and also diffused state between the fibrillae.

The writer's observation on slides prepared by Dr. Tyzzer* is as fol-
lows: Two cysts in sections; one almost spherical, 480/x by 430/x, sur-

*The writer had recently the opportunity of examining the slides of the parasites prepared
by Dr. Tyzzer, which occasion he appreciated very much. As a result of this, the writer
became convinced of the identity of forms observed by Tyzzer, Linton and Hahn, tho he
could not examine the latter authors' specimens.

333] STUDIES ON MYXOSPORJDIA—KUDO 95

rounded by several layers (about lO/i thick) of connective tissue of the
host, the other oval, 120^ by 110^. The staining sufficed to reveal only
indistinct structure of the parasites. The homogeneous ectoplasm sur-
rounds the entire surface of the body as a uniform, but very thin layer.
Endoplasm granular, filled with spores of remarkably identical stages of
development. Isolated spores, also, occur in the muscle bundle in the
state of diffused infiltration. Polysporous.

Spore: Hahn describes it as follows: Low conical pyramid with round
base; square with bulging sides. No indication of valves in the spore
shell. Dimensions: height (length) 5)u, breadth and thickness 7/*, polar
capsule 2/x by l^i.

Linton's form: squarish in outline with rounded corners, 7/i* in di-
ameter.

Tyzzer describes his form as follows: Quadrilateral in anterior end
view; oval in side view. The four corners are a little protuberant and are
directed slightly forward. Shape varies considerably in different species
of host. The corners of the spore from Stenotomus chrysops, are greatly
drawn out, exhibiting stellate form. Four polar capsules radiating from
the anterior extremity toward the four corners. Shell shows four furrows
radiating from the anterior extremity outwards to the side. Sporoplasm
occupies extracapsular cavity. Polar filaments are extruded under the
action of acetic acid. Dimension: breadth 7 to 7.5/t.

The writer's observations are as follows:

Spores in fixed and decolorized smears. In smear, most of the spores
are seen lying on the base exposing the anterior end view toward the
observer's eyes, a few lying with the sutural diameter parallel to the surface
of the slide. Form quadrilateral with corners more or less drawn out in
anterior end view; oval, with concave posterior side in front view (Figs.
562 to 564). Shell apparently thin but was not clearly separated from the
sporoplasm which is finely granular and fills the extracapsular cavity of
the spore. Four polar capsules of nearly same size and pyriform. Coiled
polar filament indistinct. When stained, the polar capsules stained deeply.
It is remarkable to see almost all of the spores exhibit four deeply stained
nuclei of capsulogenous cells, which in ordinary case disappear as the spore
matures. Dimensions: height (length) 4 to 4.75)u, breadth and thickness
5.4 to 6.5ju, polar capsule about 1.5/1 by 0.75)li.

Remarks: Thus the forms of Tyzzer, Linton and Hahn had better
be treated as one and the same species. As to the distinction of Chloro-
myxum funduli and the present species, the writer is unable to make it
clear as he could not examine the preparation of the former species and
especially as he observed some intermediate forms between these two
forms in Dr. Tyzzer's preparations of the present species.

96 ILUNOIS BIOLOGICAL MONOGRAPHS [334

CHLOROMYXUM GRANULOSUM Davis

[Figs. 157 and 158]
1917 CUoromyxum granulosum Davis 1917 : 237

Habitat: Urinary bladder of Tylosurus marianus; Beaufort (July,
August).

Vegetative form: Elongated when first placed on the slide, but soon
becomes contracted and motionless; progressing by very slow ameboid
movements. Ectoplasm usually undistinguishable, being noticed only in
a few individuals which had formed one or two short, lobose pseudopodia
of hyaline ectoplasm. Body colorless to light yellow. After being on the
slide for some time rounded trophozoites often became surrounded by a
distinct ectoplasmic layer. Entire body usually coarsely granular, the
granules var\dng greatly in size and shape; sometimes indistinctly vac-
uolated. Fat globules also present. Size of rounded trophozoites about
30/i. Disporous and polysporous.

Spore: Spherical, with four distinct ridges on the posterior half of
each valve converging toward the anterior end. Sutural ridge distinct.
Polar capsules pyriform and convergent. Dimensions: diameter 7/i,
polar capsules 2n.

Remarks: Trophozoites from some fish were" all colorless, while the
larger trophozoites from others were distinctly yellow.

CHLOROMYXUM TRIJUGUM nov. spec.
[Figs. 159 to 182]

Habitat: Gall-bladder of Lepomis megalotis Raf.; Stony Creek, and
Homer Park, HI. (November). The parasite was only found in this species,
Lepomis humilis and L. cyaneUus seined at the same time being free from
the infection. Six specimens, three from each of the above mentioned
locaUties, harboured abundantly both free spores and trophozoites of
various stages of development in the bile. The fish, from 6.5 to 10.5cm
long, were normal in external appearance and the bladders did not show any
particular abnormality, compared with those of other fish, as is usually
the case.

Vegetative form: Trophozoites float usually free in the bile, younger
forms are most frequently attached to the epithelium of the bladder. Form
extremely polymorphous, manifesting various shapes such as, almost circu-
lar, rounded, oval, elongated or irregular, which is chiefly due to the active
extrusion and retraction of the pseudopodia from the body surface. Body
is highly transparent and colorless in both the young and the adult. The
differentiation of protoplasm into ectoplasm and endoplasm, is distinctly
visible in vivo as well as in stained preparations, especially in larger forms
(Figs. 159 to 165). The endoplasm presents an alveolar structure without

335] STUDIES ON MYXOSPORIDIA—KUDO 97

any enclosure except the nuclei and various stages of spore formation
(Figs. 161, 165, 168 to 171), the alveolar network being smaller at the
periphery than in the center. The ectoplasm is a hyaline, transparent and
homogeneous layer, free from any course granulation in fresh conditions.
It shows, however, a very fine reticular structure in stained preparations.
The pseudopodia are of two kinds in form, always, formed of ectoplasm
alone : the filose and bristle-like form, sometimes branching and protruding
from the entire surface or from a localized part of the body, vary in length
from 0.5 to 4/* according to the size of the individual (Figs. 159, 161, 164).
This form developed, sometimes, into a thicker form with two to four
branched finer processes. The blunt, lobose pseudopodium formed at a
localized part of the body is well recognizable in larger individuals. Fre-
quently the filose and the lobose pseudopodia are formed on a trophozoite
at the same time. The movements of the blunt pseudopodia were striking
in some specimens. At the beginning of the observation, ten minues after
the bile was removed from the host, two club-shaped pseudopodia (Figs.
161 to 163) which were extruded from a trophozoite, the largest diameter of
which being 20/x, moved very actively in the semicircular area changing their
forms, showing maximum length of 20/x. In about thirty minutes, they
were retracted and from the same place, a short, oval-shaped pseudopodium
was seen to be extruded, which remained in the same position for some
time without great change of form (Fig. 164). In another case, a tropho-
zoite with a very broad and rounded pseudopodium extruded actively two
to three rounded smaller processes at its extremity (Figs. 165 to 167).
After fifteen minutes the pseudopodium was retracted, the ectoplasm
forming a uniformly thick layer around the endoplasm. The observations
were done at room temperature in hanging drop preparations, sealed with
vaseline and paraffin, by using comp. oc. 12 and apo. imm. ob. 2mm,,
which caused no mechanical pressure upon the parasites. The change of
form and especially that of pseud«podia, was clearly observed for one hour
and twenty minutes under the above mentioned conditions after the bile
was removed from the host. The trophozoites when kept for sixteen hours
at room temperature, underwent degeneration and disintegrated, setting
free the spores which were formed in them.

No active multiplication by plasmotomy, was observed in vivo. In
fixed preprations, however, forms that suggested the occurrence of the pro-
cess in the present myxosporidian, were recognized. As was stated before,
the pseudopodia are always formed of the ectoplasm and as each portion
of these dividing forms has many nuclei, the author is inclined to record
the presence of plasmotomy in the present form.

Size varies greatly. The monosporous form 10/x by 14/i, disporous
15)Lt by 25)Lt and polysporous 30)u by 50/i, the largest individual, developing
and containing more than 200 spores, was 300/x by 50/i.

98 ILLINOIS BIOLOGICAL MONOGRAPHS [336

Spore: Generally circular in front view; oval in side view. Shell com-
paratively thick, consequently the coiled polar filament is frequently indis-
tinct. Sutural ridge straight and distinct. Each valve has a thick straight,
sometimes slightly zigzag-form ridge that runs parallel to the sutural line,
so that in side view, three distinct ridges encircling the spore are recognized
(Figs. 177 and 180), From each of these two ridges, eight to twelve short
ridges are directed toward the center of each valve, which can distinctly
be observed on the spores stained with Heidenhain's iron hematoxylin
(Figs. 179 and 180). They can be seen as faint markings rising from the
margin directed toward the center of the spore, in front view of fresh spores.
Four pyriform polar- capsules of slightly different size open their foramina
independently at the anterior end of the spore (Figs. 178 and 181). The
sporoplasm, granular and finely reticular, shows almost always two nuclei
when stained. Dimensions in vivo: length and breadth 8 to 10/i, thickness
5 to In, polar capsules 3 to Sn by 2 to 3/x, length of polar filament 32 to
40/i (H2O2, KOH).

Remarks: In carefully made smear of the bile, a number of empty
spores which had been seen in fresh hanging drop preparations, and often
spores, in which the sporoplasm with two elongated nuclei seemed to leave
the shell (Fig. 182), were recognized. As this particular spore was found
close to a thicker mass of the wall of the gall-bladder in the smear, it can
hardly be thought that the mechanical pressure during the preparation
lead to the mission of the sporoplasm from the spore. It is possible, on the
other hand, to think that this is one of the cases of the germination of the
spore in the host in which they were developed, as was reported by the
author in Nosema bombycis Nageli (Kudo, 1916).

CHLOROMYXUM CATOSTOMI nov. spec.
[Figs. 560 and 561]

Habitat: Gall-bladder of Catostomus commersonii Lac; Salt Fork,
Urbana, 111. (October). Four fish, from 8 to 14cm.; apparently normal.

Vegetative form: Form usually rounded, with fiUform pseudopodia.
Majority attached to the epithelium, a few being free in the bile. Body
colorless. Protoplasm is not well differentiated. Endoplasm occupying
the entire body is of granular structure with vacuoles and refringent
spherules. Size: from 15 to 35)li. When kept for 16 hours in a refrigerator,
the trophozoites liberated the spores. The number of spores in each tro-
phozoite is usually 2 or 3, rarely 5 to 6. Active plasmotomic multiplication
observed when examined. Spores were comparatively small in number,
while the trophozoites were attached abundantly to the epithehum of the
gall-bladder. Disporous and polysporous.

Spore: Form approximately spherical in front view; oval in profile.
Shell with very fine striations which run parallel to the sutural ridge that
is fairly well marked. Rounded polar capsules almost of same size, have

3371 STUDIES ON MYXOSPORIDJA—KUDO 99

independent openings at the anterior end. Coiled polar filament indis-
tinct. Abnormal spores with five polar capsules are sometimes seen.
Dimensions of fixed spores: length 8j«, breadth 7/i, thickness 5 to 6fi, polar
capsules 2 to 2.5/x by l.S/i.

CHLOROMYXUM WARDI nov. spec.
[Figs. 632 to 642]

Habitat: In the gall-bladder of Oncorhynchus nerka: Klutina Lake,
Alaska (August). A single gall-bladder collected and preserved in formol
by Professor Ward, was found to harbor the present species. The study
was done on preserved material and on stained smear preparations.

Vegetative form: Young trophozoites (Fig. 632) show ameboid form,
and are mostly multinucleated. The protoplasm is not well differentiated
either in unstained material or in stained specimens. It is granulated
thruout the body, and is vacuolated at places. The smallest form mea-
sured was 18m in largest diameter. The shape of the body suggests its
possession of ameboid movements when alive, altho the writer could not
examine fresh specimens. Large trophozoites in which the spore forma-
tion had partly been completed are generally rounded with reticular proto-
plasm. Size varies to some extent. The trophozoite shown in figure 633
contains six mature spores and is 23ai in largest diameter. The largest
one found was 38 by 30ju, showing ten spores and nuclei. Each spore
appears to develop independently from a single sporoblast. Disporous
and polysporous.

Spore: Rounded pyramidal in front view (Figs. 640 and 641); circular
in transverse section (Fig. 638). The shell is thickened near the posterior
margin (Figs. 640 and 641). Sutural line is not straight, the ridge being
fairly distinct. The striations on the shell vary to a considerable extent
(Figs. 634 to 637 and 639). Four polar capsules at the anterior end,
mostly unequal in size and shape. The coiled polar filament is invisible
in formol material. Potassium hydrate solution does not cause its extru-
sion in the preserved spores. The sporoplasm is finely granular with two
nuclei. Dimensions of unstained preserved spores: diameter 7.5 to 9//,
polar capsule 3 by 2.5/*.

Remarks : The writer was able to study forty specimens of gall-bladder
of Alaskan fishes, chiefly of salmon, which have been collected by Professor
Henry B. Ward, during the summer of 1919, for which he wishes to express
his deepest appreciation. The examination of these specimens showed
that myxosporidia were found only in one of the gall-bladders, and that
specimen presented a fairly heavy infection of the present species.

Family SPHAEROSPORIDAE Davis
1917 Sphaerosporidae Davis 1917 : 219

The characters of the family are described on page 57.

100 ILLINOIS BIOLOGICAL MONOGRAPHS [338

Genus SPHAEROSPORA Thelohan
1892 Sphaerospora Th61ohan 1892 : 167

The characters of the genus are described on page 57.
Type species: Sphaerospora diver gens Thelohan.

SPHAEROSPORA DIVERGENS Thelohan
[Figs. 183 to 186]
1895 Sphaerospora divergens Th61ohan 1895 : 339-340

1912 Sphaerospora divergens Parisi 1912 : 289

1912 sphaerospora divergens Auerbach 1912 : 41-42

Habitat: Urinary tubules of kidney of Blennius phoUs L., Crenilabrus
melops L., C. pavo Cuv. et V, and urinary bladder of Hippoglossoides
Hmandoides; Concarneau, Roscoff, Napoli (July), Smalfjorden.

Vegetative form: Rounded discoidal or spherical or more or less
elongate. Ectoplasm transparent, without real pseudopodium. Move-
ments extremely slow. Endoplasm, granular, contains fat globules and
small yellowish granules. Size of sporulating individuals: 65/i by 55/*,
60)u by 25/x, 60ju by 20/i, etc. Polysporous (Thelohan); monosporous, and
disporous (Auerbach).

Spore: Spherical. Shell with fine striations. Two polar capsules
divergent; coiled polar filament visible in fresh state. Sporoplasm fills the
extracapsular cavity of the spore. Dimensions: lO/x in diameter, often
10/i by 12/x, the larger diameter coinciding with sutural plane, thickness
Bn (Auerbach), polar capsules about 4/li long, length of polar filament
20 to 25m.

SPHAEROSPORA ELEGANS Thelohan
[Figs. 187 and 188]

1890

Thdohan

1890

193-209

1892

Sphaerospora elegans

Thelohan

1892

167-175

1894

Chloromyxum (Sphaerospora)

elegans

Gurley

1894

266

1895

Sphaerospora elegans

Th61ohan

1895

338-339

1909

Sphaerospora elegans*

Auerbach

1909a

:71

1912

Sphaerospora elegans

Parisi

1912 :

289

Habitat: Renal tubules of kidney, connective tissue of ovary and
urinary bladder of Gasterosieus aculeatus L., G. pungitus L., Loia vulgaris
Cuv., Phoxinus laevis L.; Paris, Bretagne, Karlsruhe, Lake Garda.

Vegetative form: Rounded or sUghtly elongated, not exceeding 20 to
25/Lt in diameter. Protoplasm homogeneous, very finely granular, contains
numerous refractive globules, probably of fatty nature. Pseudopodia
lobose. Movements slow. Disporous.

*Misprinted as Sphaeromyxa elegans.

339] STUDIES ON MYXOSPORIDIA—KUDO 101

Spore: Spherical, somewhat attenuated at the anterior end. Sutural
ridge present, terminating in a small projection at each end of the spore.
Two polar capsules spherical. Coiled polar filament not visible in fresh
state. Dimensions: diameter lO^t in average, sutural diameter about ll^i.

SPHAEROSPORA ROSTRATA Thelohan
[Fig. 189]
1895 Sphaerospora rostrata TWlohan 1895 : 339

Habitat: Malpighian bodies of kidney of Mugil sp.; Roscoff, Le
Croisic, Le Vivier-sur-mer, Marseille, Banyuls.
Vegetative form: Not described.

Sphore: Subspherical. Shell shows deep longitudinal striations which
end in sharp spinous edges at the posterior end. Sutural ridge well
marked. Anterior part shows enlargement of quadrangular lamella, which
is spinous in side view. Dimensions: 10 to 12/i in diameter, sutural
diameter 1 to 2/i longer, length of polar filament 40/z.

Remarks: The parasites cause the degeneration of the Malpighian
bodies.

SPHAEROSPORA MASOVICA Cohn

[Figs. 190 to 192]

1902 Sphaerospora tnasovka Cohn 1902 : 628-632

Habitat: Gall-bladder of Abramis brama L.; Mauersee.

Vegetative form: Polymorphous, due to active movements. Trans-
parent and colorless, while in motion. Endoplasm highly granular, contains
yellowish enclosures. Ectoplasm hyaline, forms a narrow layer around the
body, occasionally developing into a blunt lobose pseudopodium. Pseudo-
podia of two kinds; lobose and filose, also intermediate forms. Filiform
pseudopodia are formed and retracted more slowly than the lobose.
Plasmotomy is of probable occurrence. Two spores are formed in each
pansporoblast. Size variable: lOfj. (with no spore), 18/i (with sporob lasts),
29^ (with 4 sporoblasts), 38/i (with 22 sporoblasts). Disporous(?), poly-
sporous.

Spore: Spherical. Sutural ridge well marked. Polar capsules and
sporoplasm are comparatively small, the former convergent. By warming
the spore, polar filament is extruded and at the same time two filaments
("starren Faden") are made visible at the anterior part of the sutural plane.
Sporoplasm with two nuclei, no vacuole being present. Dimensions:
diameter 8/i, length of polar filament 38/f, length of sutural filament 14/i.

Remarks: Cohn did not observe free spores in the gall-bladder. He,
however, saw many free spores, separated from each other, in the intestine,
concluding that the body and pansporoblast membrane of trophozoites,
are destroyed in the intestine, setting the spores free.

102 ILLINOIS BIOLOGICAL MONOGRAPHS [340

SPHAEROSPORA PLATESSAE Woodcock

[Figs. 193 and 194]
1904 Sphaerospora platessae Woodcock 1904 : 59H50

Habitat: Otic-capsule of Pleuronectes platessa L.; England.

Vegetative form: Cysts opaque masses about 1mm. in diameter.
The cartilage was greatly hypertrophied. Polysporous (presumably).

Spore: Spherical. Shell unornamented. Two polar capsules. Sporo-
plasm with several refractive granules, but without any vacuole. Dimen-
sions: diameter 8 to 9fi, length of polar filament about 70/i.

Remarks: Woodcock placed this species provisionally in the genus
as he could not examine any fresh material, but had studied smears only.

SPHAEROSPORA ANGULATA Fujita

[Figs. 195 to 197]

1912 Sphaerospora angulata Fujita 1912 : 261-262

Habitat: Kidney of Cyprinus carpio L., Carassius auratus L.; Sapporo
(Nippon).

Vegetative form: Only description: "The number of the spore in the
sporoblast is in this case always less than in the others, rarely exceeding
two."

Spore: Somewhat triangular, with convex sides, oval in sideview.
Slightly pointed at the mid-posterior margin of the spore. Shell very thin,
faintly marked with concentric striations. Two oblong polar capsules
are of unequal size. Dimensions: length 7 to 8ju, breadth 6 to 7^, thickness
5/x, length of largest polar capsule 3.8/i, length of polar filament twice as
long as that of the spore.

SPHAEROSPORA POLYMORPHA Davis

[Figs. 198 and 199]

1917 Sphaerospora polymorpha Davis 1917 : 231-232

Habitat: Urinary bladder of Opsanus tau; Beaufort (June, July).

Vegetative form: Elongate, but never very irregular in shape. Slowly
ameboid. Body colorless. Ectoplasm clearly seen in younger forms,
forming one to several large lobate pseudopodia, which in turn extrude
several short, conical pseudopodia. In larger forms, ectoplasm is, often,
recognizable only at ends of pseudopodia, which in such cases are composed
chiefly of endoplasm. Endoplasm granular, vacuolated in some smaller
forms, but in larger individuals vacuoles are indistinct or absent; small fat
globules abundant in large forms; numbers of rounded sporoblast cells
can be distinctly seen. Size of large trophozoites 35n by 50/x. Disporous
and polysporous (polysporous forms rarely contain many spores at the
same time).

341] STUDIES ON MYXOSPORJDIA— KUDO 103

Spore: Spherical, sometimes slightly compressed infero-superiorly.
Sutural ridge ; on each side are a number of concentric striations extending
around each valve parallel to sutural line. Polar capsules pyriform and
large. Coiled polar filaments indistinct. Sporoplasm finely granular.
Dimensions: diameter 7 to 10/x, (8/i in average), polar capsules 4 to S/t by
2 to 2.5^.

SPHAEROSPORA sp. Davis

1917 Sphaerospora sp. Davis 1917 : 213

Habitat: Urinary bladder of Lepisosteus platystomus; Gainesville, Fla.
Vegetative form: No description.
Spore: Not described.

SPHAEROSPORA sp. Southwell et Prashad

1918 Sphaerospora sp. Southwell and

Prashad 1918 : 347-348 .

Habitat: Under the scales of Barilius barna; from the vicinity of the
Ruby Mines, Burma (June).

Vegetative form : The cysts occurred in very large numbers, one under
each scale.

Spore: Authors' description: "The poor condition of the material did
not allow of a complete account of its structure, but the bicapsulate,
rounded structure of its spores places it undoubtedly in the genus Sphaero-
spora Thelohan."

SPHAEROSPORA CARASSII nov. spec.
[Figs. 200 to 204]

Habitat: Gill filament of Carassius carassius L.; Tokio (February).

Vegetative form: Trophozoites small ameboid in groups or in diffused
condition in the connective tissue of the gill filament. No cyst formation.
The number of trophozoites in groups is generally small. The largest
group found in sections was 96/i by 36/n, the macroscopical examination
always failing to trace the parasites. The trophozoites, 10 to 20/i long,
with poorly differentiated protoplasm and usually reticular endoplasm
without any particular enclosure (Fig. 200). Ameboid movements not
observed. Schizogonic multiplication rapid, each of the daughter indivi-
duals developing into two spores. Disporous. Other sporous characters
could not be determined.

Spore: Spherical in front and side views, tho form variable to some
extent (Figs. 201-203). Shell smooth. Sutural ridge fairly distinct. Two
polar capsules, broadly pyriform, of equal size and convergent, located at

104 ILLINOIS BIOLOGICAL MONOGRAPHS [342

the anterior end, one on each side of the sutural plane. Coiled polar
filament highly distinct (5 to 6 times) in vivo. Sporoplasm granular, shows
two nuclei when stained; no vacuole of any nature. Dimensions in vivo:
diameter 8 to 13/i, polar capsules 4 to 5^ by 2.5 to 3.5/x, length of polar
filament 35 to 40/x (KOH or pressure).

Remarks: No species of the genus, has ever been found in the branchiae.
The characters of the spore, however, compel the writer to place the
form in the present genus.

Genus SINUOLINEA Davis
1917 Sinuolinea Davis 1917 : 219

The characters of the genus are described on page 57.
Type species: Sinuolinea dimorpha Davis.

SINUOLINEA DIMORPHA Davis
[Figs. 205 to 213]

1916 Sphaerospora dimorpha Davis 1916 : 333-377

1917 Sinuolinea dimorpha Davis 1917 : 232-233

Habitat: Urinary bladder and ureter of Cynoscion regaJis; Beaufort.

Vegetative form: Disporous and polysporous trophozoites differ
distinctly from each other. Disporous trophozoites irregular, colorless,
transparent and show slow movements. When attached to the epithelium,
rounded with one to several pseudopodia. Differentiation of protoplasm
distinct. Occasionally endoplasm contains one or more erythrocytes.
Average diameter of full-grown form 25 to 30//.

Polysporous form: when attached to the bladder epithelium, the free end
is drawn out into a long, cylindrical process, covered with numerous short,
hairlike ectoplasmic processes. While not movable, these processes are
readily absorbed and reformed. When the trophozoite is detached from
the epithelium, the larger end gives rise to numerous conical or arborescent
pseudopodia, by means of which the trophozoite moves slowly. Endoplasm
extends into the proximal portion of large pseudopodia. It is granular
and vacuolated, contains numerous fat globules, refractive granules,
yellowish crystals (hematoidin?) and erythrocytes in various stages of
disintegration. Endoplasm also contains gemmules, each composed of
outer layer and finely granular central portion. Size varies greatly: up to
575m by 90m.

Spore: Spherical. Sutural ridge well marked. Polar capsules large
and spherical. Sporoplasm forms a rounded granular mass. Dimensions:
diameter 15m, diameter of polar capsules 4.5m, length of polar filament
27 to 35m.

3431 STUDIES ON MYXOSPORJDJA—KUDO 105

SINUOLINEA CAPSULARIS Davis
[Figs. 214 to 216]
1917 Sinuolinea capstdaris Davis 1917 : 233

Habitat: Urinary bladder of Paralichthys albiguttus, P. dentatus,
Spheroides maculatus; Beaufort (July, August).

Vegetative form : Rounded to irregular shape. Body colorless or light
yellow. Progressive movements slow. Pseudopodia large branched or
arborescent, formed entirely of ectoplasm. Ectoplasm transparent and
usually granular, merging gradually with the endoplasm. Endoplasm
contains numberous fat globules. In large trophozoites, gemmules are
observed. The gemmules are more finely granular and more transparent
than the surrounding protoplasm and are practically identical with the
small, free trophozoites. Trophozoites containing several gemmules are
usually rounded and motionless and appear to be more or less degenerate.
Disintegration of such trophozoites were actually observed. Sporulating
trophozoites were rare and were never seen to contain gemmules. Size up
to 40m in diameter. Disporous and polysporous(?).

Spore: Spherical, sometimes slightly elongated. Sutural plane much
twisted on its axis. Sutural ridge very distinct. Polar capsules and cap-
sulogenous cells large occupying more than one-half of the cavity of spore.
Coiled polar filament distinct, Sporoplasm granular contains numerous
fat globules. Dimensions: diameter 12 to 14/i, diameter of polar capsules
4.5iu, length of polar filament SOyit.

SINUOLINEA ARBORESCENS Davis
[Figs. 217 and 218]
1917 Sinuolinea arbor escens Davis 1917 : 233

Habitat: Urinary bladder of Siphostonia floridae; Beaufort
Vegetative form: Rounded or irregular. Body colorless or light yellow.
Actively ameboid, forming large arborescent pseudopodia of ectoplasm.
Ectoplasm well developed, hyaline and homogeneous. Endoplasm coarsely
granular, sometimes containing a few fat globules. Larger trophozoites
are less active and the ectoplasm less distinct. In sporulating trophozoites
the ectoplasm may entirely disappear, the entire trophozoite consisting of
a coarsely granular mass. Diameter of rounded sporulating trophozoites
75ju. Polysporous.

Spore: Rounded, in front view, slightly elongated in the anterior end
view. Polar capsules large. Sutural ridge prominent, makes a character-
istic S-shaped turn on the anterior end. Coiled polar filaments distinct.
Dimensions: length 15fx, breadth 12/i, diameter of polar capsules 5/i.

106 ILLINOIS BIOLOGICAL MONOGRAPHS [344

SINUOLINEA OPACITA Davis
[Fig. 219]
1917 Sinuolinea opacita Davis 1917 : 234

Habitat: Urinary bladder of Paralichihys albiguttus; Beaufort (August).

Vegetative form: Rounded or slightly irregular. Body colorless and
opaque. Movements slow. Pseudopodia short lobose. Ectoplasm not
distinct, except around ends of pseudopodia, where it forms a thin hyaline
layer. Endoplasm opaque, finely granular, with numerous greenish-yellow
fat globules varying greatly in size. Diameter of rounded sporulating
trophozoites 22/Lt, exceptionally large trophozoites 100/x. Disporous.

Spore: Nearly spherical, with flattened, lateral appendages extending
from the posterior side. Sutural plane slightly twisted on its axis. Sutural
ridge distinct. Polar capsules large. Coiled polar filament distinct.
Sporoplasm finely granular, containing several comparatively large fat
globules. Dimensions: diameter 12 to 13/x, diameter of polar capsules ^n.

SINUOLINEA BRACHIOPHORA Davis
[Fig. 220]
1917 Sinuolinea brachiophora Davis 1917 : 234

Habitat: Urinary bladder of Paralichthys albiguttus; Beaufort (August
only in one fish).

Vegetative form: Rounded to somewhat irregular. Body colorless.
Ectoplasm hyaline. Endoplasm granular, with numerous large fat glob-
ules. Disporous.

Spore: Nearly spherical, with a long lateral appendage from each
valve. These appendages are empty except at extreme distal end, which
contains a granular mass, probably the remains of the parietal cell. Sutural
plane slightly oblique to longitudinal axis. Sutural ridge distinct. Polar
capsules and capsulogenous cells large, occupying more than half of cavity
of spore. Sporoplasm finely granular. Dimensions: length exclusive of
appendages 9 to ll/x, length of appendages 18 to 22/i, breadth of spore 9/i,
diameter of polar capsules 3.5/*.

Remarks: Davis mentions that in many respects this species is very
similar to S. opacita, which occurs in the same host.

Suborder PLATYSPOREA nom. nov.

The definition of the suborder is recorded on page 57.

Family MYXIDIIDAE Thelohan

1892 Myxididees Thaohan 1892 : 173, 175

1893 Myxidiidae Gurley 1893 : 412

The characters of the family are described on page 57.

345]

STUDIES ON MYXOSPORIDIA—KUDO

107

Genus MYXIDIUM Biitschli
1882 Myxidium Butschli 1882 : PI. 38

The characters of the genus are described on page 58.
Type species: Myxidium lieberkiihni Butschli.

MYXIDIUM LIEBERKtJHNI Butschli
[Figs. 221 to 240]

1854

1879

1881

1882

Myxidium

1883

1891

Myxidium

1894

Myxidium

1895

Myxidium

1895

Myxidium

1898

Myxidium

1902

Myxidium

1902

Myxidium

1906

Myxidium

1909

Myxidium

1912

Myxidium

1912

Myxidium

1916

Myxidium

1916

Myxidium

lieberkuhni
lieberkuhni
lieberkuhni
lieberkuhni
lieberkiihni
lieberkuhni
lieberkuhni

lieberkuhni
lieberkuhni
lieberkuhni
lieberkuhni
lieberkuhni
lieberkuhni

Lieberkuhn

1854

5H5,349

Leuckart

1879

246

Butschli

1881

638-648

Butschli

1882

593-595

Balbiani

1883

201-202, 274-275

Pfeiffer

1891

20, 91, 105, 127

Gurley

1894

283-289

Thelohan

1895

340

Cohn

1895

5-36

Doflein

1898

229, 341

Prenant

1902a

: 200-217

Laveran and

Mesnil

1902

469-472

L^ger and Hesse 1906

720

Auerbach

1909a

:71

Schroder

1912

326-327

Parisi

1912

286

Mavor

1916a

:66-68

Mavor

1916b

: 373-378

Habitat: Urinary bladder of Esox lucius L., Lota lota L. (L. vulgaris) \
France, Canada (Georgian Bay), U. S. A., (Wisconsin, Lake Mendota),
Italy (Lago Maggiore, Lago di Como, Milano), Germany.

Vegetative form: Form variable with lobose or immovable filiform
pseudopodia. Clear differentiation of protoplasm. Cohn described third
layer of protoplasm (mesoplasm). Endoplasm yellowish in older tropho-
zoites, contains yellow globules, fat globules and hematoidin crystals.
Size varying with age up to a maximum length of 300/i by a breadth of
136/x (Butschli). Plasmotomous multiplication active. Cohn described
budding of larger forms, while Laveran et Mesnil observed only the division
of smaller forms. Each pansporoblast develops into two spores. Poly-
sporous.

Spore: Elongated fusiform. Shell with longitudinal striations. Polar
capsule at each end of the spore. The longer axis of polar capsules coincides
with that of spore. Dimensions: length 18 to 20/i, width 5 to 6At. Mavor's
measurement: polar capsules 5/i by 2.5 to Zn, length of polar filament
40 to 45/i.

106 ILLINOIS BIOLOGICAL MONOGRAPHS [346

MYXIDIUM INCURVATUM Thelohan
[Figs. 241 to 251]

1892

Myxidium ? inciirvatum

Th61ohan

1892a

: 1093-1094

1895

Myxidium incurvatum

ThflohaTi

1895

341

1912

Myxidium incurvatum

Parisi

1912

286-287

1912

Myxidium incurvatum

Auerbach

1912

4,39

1916

Myxidium incurvatum

Georg6vitch

1916

90-91

1917

Myxidium incurvatum

Davis

1917

234^235

Habitat: Gall-bladder of Xerophis aequoreus L., N. annulatus, N.
lumbriciformis, Blennius pholis L., Callionymus lyra L., Fundulus majalis,
Gambusia affinis, Hippocampus brevirostris, Mugil cephalus, Scorpaena
scrofa L., Syngnathus acus L., S. typhle; Roscofif, Concarneau, Marseille,
Banyuls, Napoli, Bergen, Monaco, Beaufort (July).

Vegetative form: Thelohan describes as follows: Trophozoites usually
small, sometimes reaching a considerable size. Pseudopodia lobose.
Protoplasm pale and finely granular with refractive globules. Disporous.
According to Parisi and Davis rarely monosporous. Georgevitch observed
apparently the polysporous form.

Parisi's form: ectoplasm hyaline, endoplasm granular. Monosporous
form 25/1 long.

Davis's form: lobose pseudopodia, occasionally being drawn out into a
long process. Many trophozoites often cling together closely. Diameter
of rounded disporous forms about 13 to 15m, that of monosporous forms
about 10 to ll/t.

Spore: Thelohan's description is as follows: Irregular fusiform. Long-
est axis curved into S-form, both ends sharply pointed and directed toward
opposite directions. Polar capsule opening on opposite side of the spore,
in some spores the axis of the polar capsules being parallel to each other.
Dimensions: length 8 to 9/x, breadth 4 to 5/iz, length of polar filament 10
to 15m.

Parisi gave the following dimensions: length 10 to 12/x, breadth 5 to
6/t, length of polar capsule 3m, length of polar filament 28m.

According to Georgevitch, young spores are not curved (Fig. 245).

Davis's form; Polar filaments when extruded in HCl remained tightly
coiled. Dimensions: length 8 to 9m, Avidth 5 to 6m, diameter of polar
capsule about 3>n.

Remarks: As are shown in figures, Davis's form seems to be some-
what different from the European forms.

347] STUDIES ON MYXOSPORIDIA—KUDO 109

MYXIDIUM SPHAERICUM Thelohan
[Fig. 252]
1895 Myxidiunt sphericum (corr.

sphaericum) Thelohan 1895 : 341-342

Habitat: Gall-bladder of Belone acus (Belone belone L.); Banyuls,
Le Vivier-sur-Mer.

Vegetative form : Trophozoites spherical or subspherical, not exceeding
20 to 22)Li in diameter with lobose pseudopodia formed from the entire
surface. Endoplasm granular, contains small refractive granules. Dis-
porous.

Spore: Form similar to M. incurvatum, but much greater. Coiled polar
filament distinctly visible in fresh spore. Dimensions: length 15 to 20/t,
width 7 to 8m, length of polar filament 60/i (KOH).

MYXIDIUM HISTOPHILUM Thelohan
[Fig. 253]
1895 Myxidiunt histophilum Th61ohan 1895 : 341

Habitat: Connective tissue of kidney and ovary of Leuciscus phoxinus
L. (Phoxinus laevis Ag.); France.

Vegetative form: Small mass.

Spore: Fusiform, being compressed at the middle part. Shell with
longitudinal striations. Length of the spore 15/i.

MYXIDIUM sp. Gurley

[Fig. 254]

1851

Leydig

1851

: 226, 234

1852

Leuckart

1852

:436

1894

Myxidiunt ? sp. incert.' Gurley

1894 :

:290

1899

Myxidiunt sp. Labb^

1899 :

:92

Habitat: Gall-duct of Raja hatis L.
Vegetative form: No description.
Spore: Not described. One figure.

MYXIDIUM DANILEWSKYI Laveran
[Figs. 255 to 257]

1887 DanUewsky 1887 : 35

1897 Myxidiunt danilewskyi Laveran 1897 : 725-726

1898 Myxidiunt danilewskyi Laveran 1898 : 27-30

Habitat: Kidney of Emys orbicularis L.; France.

Vegetative form: Form elongated, circular in cross-section, tapering
toward the ends. Body of greenish color, occupying the lumen of the renal
tubules of the kidney. Body bent along the cavity of the tubule. Endo-

no ILLINOIS BIOLOGICAL MONOGRAPHS [348

plasm granular, ectoplasm covering the entire surface of the body as a
thin layer. Each pansporoblast develops two spores. Polysporous.

Spore : Elongated fusiform, similar to M. lieherkuhni, but much smaller.
Polar capsule at each end, extrudes filament under the action of nitric
acid. Sporoplasm granular with one nucleus. Dimensions: length 12/*,
breadth 3 to 4/i.

MYXIDIUM GIGANTEUM Doflein

[Fig. 258]

1898 Myxidium giganteum Doflem 1898 : 285-286

Habitat: Gall-bladder of Raja asterias; Napoli.

Vegetative form: Rounded trophozoites. Lobose pseudopodia with
slow movement, show remarkable dimensions. Posterior portion forms
"Stemm-pseudopodien." Small form club-shaped. Endoplasm is of
yellowish color. Diameter of large form 500/x, of medium sized 200/i, small
individuals 70-90^t, quite young ones, polymorphous 8 to 40/x. Larger
individual up to 700/i by 180/i. Many trophozoites form a cyst-like motion-
less stage, in which many individuals seem to be covered with a common
gelatinous envelope. Each pansporoblast forms two spores. Polysporous.

Spore: Elongated. Fusiform in front view; in side view, one valve
arch-form, the other being flat. Transparent. Two polar capsules, one
at each end. Coiled polar filament is clearly seen in larger polar capsules.
Dimensions : length 28ju, breadth 8/t, polar capsules 8/x by 4/1.

MYXIDIUM BARBATULAE Cepede

1906 Myxidium barbatulae C6pSde 1906 : 67

1906 Myxidium barbatulae Cepede 1906a : 15-16

Habitat: Kidney of Cobitis barbatula L.; Isere.

Vegetative form: Trophozoites form cysts. Form and size vary
greatly. Average size: 400 to 500m in length and 200/x in breadth.

Spore: Irregular fusiform. Polar capsule at each end of the spore.
Shell longitudinally striated, number being variable. Dimensions: length
12 to 15/i, breadth about 6/x, polar capsules 5/i by 2.5 to Six.

MYXIDIUM GIARDI Cepede
[Figs. 259 to 261]
1906 Myxidium giardi C6pede 1906a : 16; 1906b : 170-173

1908 Myxidium giardi C6pSde 1908 : 93-95

1908 Myxidium giardi C6pede 1908a : 8

Habitat: Kidney of Anguilla vulgaris Flem.; Wimereux (August).

Vegetative form: Subspherical white cysts, 800 to 900/x in diameter,
surrounded by a thick (up to 30/x) membrane, composed of the connective
tissue of the host.

349] STUDIES ON MYXOSPORIDIA—KUDO 111

Spore: Irregular fusiform, greatly enlarged at the middle portion.
Plane of symmetry of the spore coincides with the sutural plane. Shell
thick with 9 to 11 longitudinal striations on each valve, which are more
clearly seen on spores stained with iron hematoxylin. Polar capsule at
each end. Coiled polar filament distinct. Sporoplasm finely granular
with two nuclei and refringent globules. Dimensions in vivo: length 9 to
lO^t, width 5 to 5.6/1, thickness 4.75 to 5/x, polar capsules 3. 5m by 2ju.

MYXIDIUM PFEIFFERI Auerbach

[Figs. 262 to 265]

1908 Myxidium pfeiferi Auerbach 1908 : 459-464

1910 Myxidium pfeiferi Auerbach 1910c : 171-172

Habitat: Gall-bladder of Tinea vulgaris Cuv.; Karlsruhe.

Vegetative form: Observations in sections. More or less flattened,
disc-form, often enrolled. The ectoplasm finely granular, without large
pseudopodia. It is not usually distinguishable from the endoplasm which
is highly alveolar and contains numerous nuclei, but no enclosures.

Spore: Form varies to some extent. Similar to Myxidium lieberkiihni;
slightly curved. Shell with fine longitudinal striations. Polar capsules
two, one at each end. Polar filament is extruded by adding one drop of
water to the smear of the spore, which had been dessicated for 24 hours.
Sporoplasm with one or two nuclei, in one case with four small nuclei,
which is thought to be an abnormal. Dimensions: length 13 to IS^i,
breadth 5.2 to 5.8/x, length of polar capsule 5.2 to 6/i, length of polar fila-
ment 45 to 54/i.

MYXIDIUM INFLATUM Auerbach
[Fig. 266]

1909

Myxidium inflatum

Auerbach

1909 : 72-74

1909

Myxidium inflatum

Auerbach

1909a : 31

1910

Myxidium inflatum

Auerbach

1910c : 172

1912

Myxidium inflatum

Auerbach

1912 : 39

Habitat: Gall-bladder of Cydopterus lumpus L.; Bergen (September).

Vegetative form: Extremely polymorphous. Rounded, spherical,
or much elongated. Ameboid movements very active. Differentiation of
protoplasm is sharp and clear, which is best observed in individuals in
motion; highly hyaline ectoplasm forms very long lobose pseudopodia,
into which granular endoplasm flows in slowly. Size variable. Rounded
large form 44 to 45/i in diameter. Fully grown spores are set free from the
mother trophozoite in comparatively short time. Spore formation similar
to that of Myxidium bergense. Disporous and polysporous (5 spores in
maximum).

Keysselitz

1908 :

289

? Myxidium sphaericum

Auerbach

1909 :

75-76

Myxidium bergense

Auerbach

1910 :

61

Myxidium bergense

Auerbach

1910c

:172

Myxidium bergense

Auerbach

1912 :

18-39

Myxidium bergense

Mavor

1915 :

30-31

jall-bladder of Gadus

virens L., G. aet

'lefinis

,G.l

112 JLUNOJS BIOLOGICAL MONOGRAPHS [350

Spore: Very broad compared with the length. The longitudinal axis
is curved in S shape. Polar capsule situated in opposite way at each end
of the spore. Dimensions: length 20.8 to 23.4pi, breadth 13 to 15.6/i,
polar capsules 7.8^, length of polar filament 90 to lOOjit (KOH).

MYXIDIUM BERGENSE Auerbach
[Fig. 267]

1908
1909
1910
1910
1912
1915

Habitat:

Pleuronectes platessa and Sehastes viviparus, Melanogrammus aeglefinis;
Norway (Bergen), Canada (St. Andrew, July to September).

Vegetative form: Rounded or elongated, as the result of formation of
various pseudopodia. Trophozoites partly free, partly attached to the
epithelium of the bladder. Size up to 54/x in diameter. Pseudopodia of
two kinds: lobose and long filose, sometimes slightly branched. Mavor
observed a cyst-like stage under certain conditions, which, he thinks, may
be due to some exceptional conditions of the parasite. Plasmogamy.
Monosporous, disporous and polysporous.

Spore: Fusiform. Main axis curved into S shape. Form, roughly
speaking, very much similar to that of M. sphaericum Thel. Dimensions:
length 16.2 to IQjli, breadth 7 to 9n, length of polar capsules 5An, length
of polar filament about three times as that of spore.

MYXIDIUM PROCERUM Auerbach
[Fig. 268]
1910 Myxidium procerum Auerbach 1910 : 61-62

1910 Myxidium procerum Auerbach 1910c : 172-173

1912 Myxidium procerum Auerbach 1912 : 4, 39

Habitat: Gall-bladder of Argentina situs As.; Bergen.

Vegetative form: Not observed.

Spore: Greatly elongated and narrow. Sporoplasm with one or two
nuclei. Dimensions: length 21.6 to 25.2/i, breadth 3.6 to 4/x, length of
polar capsule 7.2/1.

MYXIDIUM MACKIEI Bosanquet

[Figs. 269 to 271]

1910 Myxidium mackiei Bosanquet 1910 : 436-438

Habitat: Renal tubules of kidney of Trionyx ganzeticus; Bombay.
Observations on three slides.

351] STUDIES ON MYXOSPORIDJA—KUDO 113

Vegetative form: The largest trophozoite 160^ by 21 fi. No distinction
between ectoplasm and endoplasm could be drawn, except in a few indi-
viduals in which there was a cyst- wall. Spores are formed in pairs. Proto-
plasm with two kinds of nuclei, some vesicular, others smaller and com-
pact. Polysporous.

Spore: Fusiform with rather pointed ends. Shell finely striated. Two
comparatively small polar capsules, one at each end. Sporoplasm with
one or two nuclei, contains, often, two large vacuoles. Dimensions:
length 16/x (a few 11 ix), breadth 5^ (many broader than this).

Remarks: The discoverer, J. P. Mackie mentioned that the parasites
did not appear to excite any reaction in the tissue of the host, the animal's
health being unaffected.

MYXIDIUM MACROCAPSULARE Auerbach
[Figs. 272 and 273]

1910 Myxidium macrocapsulare Auerbach 1910 : 440-441

Habitat: Gall-bladder of Scardinius eryihrophihalmus L.; Karlsruhe.

Vegetative form: Not observed.

Spore : Elongated elliptical when viewed at right angles to sutural plane.
Shell somewhat thick with longitudinal striations parallel to the sutural
line. In side-view, both ends pointed in diagonally opposite directions.
Polar capsules are comparatively large, one at each end, opening at the
sharply pointed end. Dimensions: length 10 to 12/x, breadth 6/x, polar
capsules 3 to 4^t.

Remarks: No pathological change. Bile was clear.

MYXIDIUM sp. Awe'rinzew

[Figs. 274 to 276]

1908

Myxidium sp. Awerinzew

1908

: 33, 43, 45, 55

1909

Myxidium sp. Awerinzew

1909

: 76, 78, 80, 81

1911

Myxidium sp. Awerinzew

1911

: 199-204

Habitat: Gall-bladder of Cottus scorpius; Aleksandrowsk, North Sea.

Vegetative form: Trophozoites are small. The protoplasm is differ-
entiated into ectoplasm and endoplasm in some specimens. Very active
formation of filiform pseudopodia of various length. Degenerating troph-
ozoites, with one or two empty spaces are often noticed. Each spore
is formed independently from each other. Monosporous, disporous and
polysporous (with three spores).

Spore: Form similar to Myxidium incurvatum. Young spores not
curved. Dimensions: length 20 to 35^1, breadth 10 to 15/i.

114 ILLINOIS BIOLOGICAL MONOGRAPHS [352

MYXIDIUM DEPRESSUM Parisi

[Figs. 277 and 278]
1912 Myxidium depressum Parisi 1912 : 287

Habitat: Gall-bladder of Citharus Unguatula Gthr.; Napoli (August).

Vegetative form: Not observed.

Spore: Fusiform with greatly attenuated extremities in front view;
flattened and curved in S-form in profile. The axis of polar capsules
parallel to each other. Coiled polar filament visible in vivo. Sporoplasm
with two nuclei, occupies the extracapsular cavity of the spore. Dimen-
sions: length 12 to 14/x, breadth 5.5 to 6/x, thickness 2.5 to 3^1, polar cap-
sules 5.5 to 6/Li by 2.3^1, length of polar filament 30m.

MYXIDIUM OVIFORME Parisi
[Figs. 279 and 280]

1912 Myxidium oviforme Parisi 1912 : 287-288

1912 Myxidium oviforme Auerbach 1912 : 39

Habitat: Gall-bladder of Apogon rex mullorum Cuv., Coris julis Gthr.,
Gadus callarias L., Trutta solar L.; Napoli (August), Norwegian coast.

Vegetative form: Unobserved by Parisi. Auerbach's observation is as
follows :

Trophozoites, small ameboid, usually spherical. Size 10 to 12/i in
diameter. Monosporous (probably).

Spore: Oval with rounded extremities, slightly pointed at the foramina
of polar capsules. Shell with numerous fine striations running longitudin-
ally. Polar capsules being often invisible, opening a little above and below
of the hypothetical horizontal plane. Sporoplasm fills the extracapsular
cavity of the spore, leaving little space at the extremities of the polar
capsules. Dimensions: length 11/i, breadth 8 to 8.5/i, polar capsules
4.5)u by 2>ix, length of polar filament 30 to 35/i. Auerbach's measurements:
length 12 to 13/i, breadth 8 to 9/i, polar capsules about 4/t long.

MYXIDIUM ANGUILLAE Ishii

[Figs. 281 to 284]

1915 Myxidium anguiUae Ishii 1915 : 372-382

Habitat: Integument of the side of the body of Anguilla japonic a
Temm. et Sch.; Schizuoka, Nippon (October). Number of the cysts
visible to unaided eye, 10 and 9 on the left and the right side respec-
tively.

Vegetative form: Trophozoites form white and sharply contoured cysts.
Cysts, spherical or oval, surrounded by a membranous connective tissue
(about 2ix thick) of the host. Protoplasm is clearly differentiated into
ectoplasm and endoplasm. Diffuse infiltration also occurs. Size measured
along the skin, 1.2 to 2mm. in diameter; in sections 1.174mm. by 0.658mm.

353] STUDIES ON MYXOSPORIDIA—KUDO 115

Spore: Form similar to Myxidium pfeiferi Auerbach, but rather
straight fusiform, rarely slightly bent. In many spores the shell tapers
to a sharp point at each end. Shell striated longitudinally, 22 in all
(2 sutural ridges?). Two polar capsules, one at each end. Sporoplasm
usually with two nuclei. Dimensions: length 9.1ai, breadth 2,8^, length
of polar capsule 3.5/x.

MYXIDIUM sp. Mavor
[Figs. 285 and 286]

1915 Myxidium sp. Mavor 1915 : 32

Habitat: Gall-bladder of Pseudopleuronectes americanus ; New Bruns-
wick (Canada), of rare occurrence.

Vegetative form: Observations in smears are as follows: Spheroidal,
with numerous long pseudopodia on one side, which suggests the attach-
ment of the trophozoite to the bladder. Trophozoites without any spore.
Pansporoblasts spherical, 15 to 16/i in diameter.

Spore: Spindle shaped. The long axis being slighly bent in S-form.
Two pear shaped polar capsules, one at each end of spore. Coiled polar
filament visible in fresh state. Dimensions: length 14 to 15/x, breadth
6 to 7.5/i, polar capsules 4/i by 2.5/x, length of polar filament 90 to 95/i
(ammonia water).

MYXIDIUM GADI Georgevitch
[Figs. 287 to 290]

1916 Myxidium gadi Georg6vitch 1916 : 88-89

1917 Myxidium gadi Georg6vitch 1917c : 797-799
1919 Myxidium gadi Georgevitch 1919 : 251-289

Habitat: Gall-bladder of Gadus pollachius, Solea vulgaris Quens;
Roscoff (September).

Vegetative form: Highly polymorphous. Spherical or oval. Large
forms fill up the bladder. Ectoplasm hyaline and transparent, forming
one long or many short lobose pseudopodia. Endoplasm colorless and
finely granular, contains more or less large numbers of nucleus. Mono-
sporous, disporous and polysporous.

Spore: Fusiform with attenuated ends. Young spores more attenuated
than the fully grown forms. The main axis of the spore coincides with the
longitudinal axis of the polar capsules, with slight deviation. Two nuclei
of the sporoplasm, are always smaller than those of the shell-valves or
of polar capsules. Dimensions: length 6 (?) to 14jli, breadth 4 to 6ju.

MYXIDIUM GLUTINOSUM Davis
[Fig. 291]
1917 Myxidium glutinosum Davis 1917 : 235

Habitat: Gall-bladder of Cynoscion regalis; Beaufort.

116 ILLINOIS BIOLOGICAL MONOGRAPHS [354

Vegetative form : Elongated or irregular. Slowly ameboid, moving by-
means of a broad, lobose pseudopodium of hyaline ectoplasm. Body
colorless. Ectoplasm only distinct in pseudopodium. Endoplasm finely
granular. The mature spores while still within the mother trophozoites,
are surrounded by a clear, refractive gelatinous envelope. Diameter of
rounded sporulating trophozoites 20/i. Disporous.

Spore: Cylindrical, ends of valves rounded except at one side, where
the polar capsules open at the apex of a small, conical elevation. Spore
characterized by the presence of a transparent, homogeneous, gelatinous
envelope. Polar capsules pyriform, opening on each side nearly at right
angles to the longitudinal axis. Dimensions: length 10 to 11/x, breadth
6/i, length of polar capsules 3/x.

MYXIDIUM PHYLLIUM Davis

[Figs. 292 and 293]
1917 Myxidium phylllum Davis 1917 : 235

Habitat: Gall-bladder of Gamhusia affinis; Beaufort.

Vegetative form: Exceptionally large; flattened, leaflike, usually folded
on itself; motionless. Pseudopodia were not observed. Ectoplasm forming
a distinct transparent layer around entire body. After being on slide for
some time ectoplasm usually becomes covered with very numerous, short,
hairlike processes. Endoplasm finely granular, contains numerous fat
globules. Diameter up to 1.35mm. Polysporous.

Spore: Fusiform, slightly truncated at each end where polar capsules
open. Shell with numerous longitudinal striations. Sporoplasm finely
granular, with several small fat globules. Dimensions: length 1 1/x, breadth
8/x, diameter of polar capsules 3^.

MYXIDIUM STRIATUM Cunha et Fonseca
1917 Myxidium striatus Cunha et Fonseca 1917:321

Habitat: Gall-bladder of Menticirrhus americanus L., Bairdiella
ronchus Cuv. et Val. ; Brazil.

Vegetative form: More or less spherical. Body small and colorless.
Endoplasm granular. Ectoplasm visible when pseudopodia are formed.
Pseudopodia filiform, being projected radially. Size variable, 16ju in
diameter in average.

Spore: Elliptical. Shell with fine longitudinal striations which run
parallel to sutural line. Sutural plane oblique to the longitudinal axis of
the spore which is thickened at the extremities. Two ovoidal polar cap-
sules, one at each end. Dimensions: length 10 to 14^t, breadth 6 to 8/i,
length of polar capsules 4/i, length of polar filament 30/i.

3551 STUDIES ON MYXOSPORIDIA—KUDO 117

MYXIDIUM KAGAYAMAI nov. spec.
[Figs. 294 and 295]
1916 Myxidium sp. Kudo 1916 : 6

Habitat: Gall-bladder of Misgurnus anguillicaudatus Cant.; Tokio
(September), 2% of the fish examined infected.

Vegetative form: Not observed.

Spore: Fusiform; one valve being more convex than the other. Suture
line straight. Shell with fine longitudinal striations. Dimensions in fixed
preparations: length 15 to 18m, breadth 6 to Ijx, length of polar capsules
7 to 8m, length of polar filament 60 to 70^.

Remarks: Tho the vegetative form is still unobserved, the author is
compelled to consider the present form as a new species by careful
reexamination of the material and proposes the name in honor of Dr.
T. Kagayama, Tokio, Nippon.

MYXIDIUM AMERICANUM nov. spec.
[Figs. 622 to 627]

Habitat: In the lumen of urinary tubules of the kidney of Trionyx
spinifera; Crystal Lake, Urbana, 111. (July). A single host specimen
showed a light infection in the above mentioned organ. No intracellular
stage was detected.

Vegetative form: The young trophozoite in the lumen of the tubule
of the kidney is multinucleate, and more or less irregular in shape which
suggests the ameboid movements of the animal (Figs. 622, 623). The
older form with mature spores is rather spherical in form with a distinct
outline. The protoplasm is fairly well differentiated into ectoplasm and
endoplasm (Fig. 624). The size of the trophozoites varies from 12 to 25/1
in diameter. A pansporoblast produces two spores. Polysporous.

Spore: Spindle-form; with the two pointed extremities stretched in
opposite directions. Circular in cross-section. The shell is rather thin;
sutural line is straight. Fine longitudinal striations on the shell, eight
to ten in number on each valve. The polar capsules are nearly spherical,
coiled polar filament being visible in fresh material (three turns). The
polar filament is easily extruded from the fresh spores under the influence
of potassium hydrate solution. The direction of the extruded polar fila-
ment forms an angle of about 45° with the main axis of the spore and the
two filaments are parallel to each other. Preserved spores do not show
any filament extrusion under the influence of the said chemical. The
sporoplasm is finely granular, and shows, upon staining, two small nuclei
of ring-shape, as their peripheral layer takes stain more deeply than the
central portion. Average dimensions of fresh spores: length 15 to 16m,
breadth 5.5 to 6m, polar capsule 4m by 3.5m, length of polar filament
25 to 32m.

118 ILLINOIS BIOLOGICAL MONOGRAPHS [356

Remarks: Two species of the genus Myxidium were reported to occur
in chelonian hosts; i.e., M. danilewskyi (page 109) and M. mackiei (page
112). The former diflFers from the present form in having an elongated
vegetative form which is greenish in color, and in having spores of different
shape, dimensions and structure, not to speak of the difference of the
host. The latter resembles closely to the species under consideration in
dimensions of the spores, but differences in the trophozoite and in the
structure of the spore do not allow one to consider two forms as identical.
The species is therefore treated as new.

Genus SPHAEROMYXA Thelohan
1892 Sphaerotnyxa Th61ohan 1892a : 1091-1093

The characters of the genus are described on page 58.
Type species: Sphaerotnyxa balbianii Thelohan.

SPHAEROMYXA BALBIANII Thelohan
[Figs. 296 to 307]

1892 Sphaeromyxa balbianii Thelohan 1892a : 1091-1093

1895 Sphaeromyxa balbianii Th61ohan 1895 : 342

1912 Sphaeromyxa balbianii Parisi 1912 : 288

1916 Sphaeromyxa balbianii Georg^vitch 1916 : 92-93

1917 Sphaeromyxa balbianii Davis 1917 : 235-236

Habitat: Gall-bladder of Motella tricirrata Bl., M. maculata Risso.,
Cepola rubescens L., Clupea pilchardus, Siphostoma floridae, S. louisianae;
Roscoff (September), Concarneau, Marseille, Banyuls, Napoli (Septem-
ber), Beaufort (June to August).

Vegetative form: Flattened leaf -like or disc-form, reaching 3 to 4mm.
in diameter. Often forms spherical with opaque appearance. The proto-
plasm is distinctly differentiated into endoplasm and ectoplasm. Ecto-
plasm forms rounded lobes which exhibit slow movements and show a clear
radially striated structure in sections. Endoplasm reticular, contains
nuclei, young and mature spores and fat globules. Each pansporoblast
develops two spores. Polysporous.

Davis mentions that the largest form he observed was 900^ in diameter.

Georgevitch recognized a large number of small trophozoites which were
formed by repeated plasmotomous multiplication.

Spore: Fusiform, with truncate ends. Shell longitudinally striated.
One polar capsule at each end. Polar filament is wound around an imagi-
nary axis perpendicular to the longitudinal axis of the spore. When
extruded, the polar filament is seen as a short, conical and hollow thread-
like structure. Sporoplasm finely granular with two nuclei. Dimensions:
length 15m, width 5fx, length of polar filament 15/Lt.

357] STUDIES ON MYXOSPORIDIA—KUDO 119

Parisi gave the following dimensions: length 15 to 20ju, width 5 to 6/i,
polar capsule 7/i by 4.7/x, length of polar filament 25 to 30jii.

Davis' measurements are as follows: length 17 to 20/i, breadth 5 to 6/i,
length of polar filament 20/i.

Georgevitch observed young spores with both ends tapering into a
point. Later they assume the typical form with truncated ends. He did
not recognize the striations on the shell. He also mentions the occurrence
of abnormal spores, such as elliptical, spherical forms, etc.; or with only
one polar capsule.

SPHAEROMYXA IMMERSA (Lutz) Thelohan
[Figs. 308 to 311]

1889 Cystodiscus immersus Lutz 1889 : 84-88

1895 Sphaeromyxa immersa Thelohan 1895 : 343

1899 Cystodiscus immersus Liihe 1899 : 291-293

Habitat: Gall-bladder of Bufo marinus L. and Leptodactylus ocellatus
L.; Brazil.

Vegetative form: Leaf-like or disc form, visible thru the bladder wall.
Upper and lower sides slightly convex. Size up to 1.5 or 2mm. in diameter;
thickness being 1/20 to 1/10 of the diameter. Protoplasm is well differ-
entiated. Ectoplasm transparent and membranous, often contains a
large number of micrococcus-like bodies. No ameboid movements nor
change of form. Endoplasm highly vacuolar, contains fat globules.
Plasmotomic multiplication probably occurs. Spores always arranged in
pairs. Polysporous.

Spore: Oval with rounded extremities. Shell more or less thick, with
fine transverse striations. Spherical polar capsule at each end. Sutural
plane is oblique to the longitudinal axis of the spore. Sporoplasm trans-
parent. Dimensions: length 12 to 14/x, breadth 9 to 10/x, length of polar
filament 50 to 70/11 (4 to 5 times that of the spore) (KOH).

SPHAEROMYXA INCURVATA Doflein

[Figs. 312 to 314]

1898 Sphaeromyxa incurvaia Doflein 1898 : 286-287

Habitat : Gall-bladder of Blennius ocellatus ; Napoli.

Vegetative form: Trophozoites are found in large masses (Plasmodia ?),
in which they form a thin, hollow ball, 5-7mm. in diameter.

As the surface is greater than the inner surface of the bladder, some
parts of the body are folded up. Body bluish white and transparent.
Protoplasm highly vacuolar, contains numerous fat globules, nuclei and
spores. Polysporous.

Spore : Curved to one side along sutural plane and also in a plane at right
angles to it. Polar capsule at each end. Sporoplasm with two nuclei.

120 ILLINOIS BIOLOGICAL MONOGRAPHS [358

Polar filament is wound along the longer diameter of the capsule, and
relatively thick, but thinner than that of S. balbianii Thel. Dimensions:
length (along the inner side of the arch) 30 to 35/*, breadth 8/t, distance
between two polar capsules 12 to IS^u, polar capsules 12 to 15m by 4 to 5/i.

SPHAEROMYXA SABRAZESI Laveran et Mesnil
[Figs. 315 and 322]

1900

Sphaeromyxa sabrazesi

Laveran et

Mesnil

1900

: 380-382

1906

Sphaeromyxa sabrazesi

Schroder

1906 :

: 455-466

1907

Sphaeromyxa labrazesi*

Schroder

1907 :

: 359-381

1910

Sphaeromyxa sabrazesi

Schroder

1910;

:l-5

1912

Sphaeromyxa sabrazesi

Parisi

1912 ;

:288

1913

Sphaeromyxa sabrazesi

Jameson

1913 ;

;2

1916

Sphaeromyxa sabrazesi

Georg6vitch

1916;

: 91-92

1916

Sphaeromyxa sabrazesi

Georg^vitch

1916a

:3

Habitat: Gall-bladder of Hippocampus hrevirostris Cuv., H. guttulatus
Cuv.; Syngnathus acus, Motella tricirrata, Nerophis annulatus, Siphonos-
toma rondeletii; Arcachon, Rovigno, Napoli, Roscoff (September), Monaco,
Villef ranee, (March to June).

Vegetative form: Disc form. Diameter up to 2mm. Thickness var-
iable. Body whitish in color. Ectoplasm thin, transparent and homo-
geneous. Young trophozoites may probably have lobose pseudopodia.
Endoplasm highly vacuolated, contains nuclei of various sizes, pansporo-
blasts, spores and more or less refringent granules. Polysporous.

Schroder observed larger forms up to 5mm. Ectoplasm also was found
to project numerous fine short (Iju) hair-like processes from the surface.
Each pansporoblast develops into two spores.

Spore: Cylindrical, bent in arch form; with truncated ends. Large
cylindrical polar capsule at each end. Sporoplasm granular, contains one
nucleus. Polar filament short and conical, is extruded under the action of
nitric acid. Dimensions: length 28)u, width 4.3m, polar capsule 9 to 10m
by 3n, distance between the polar capsules 8m, length of polar filament 8m.

Schroder noticed the stained sporoplasm contained one or two nuclei.
He observed indistinctly marked longitudinal striations on the shell.
Dimensions: length 22 to 25m, breadth 3 to 4m, polar capsule 8m by 2 to 3m,
length of polar filament about 12m.

Georgevitch described the presence of a hyaline substance, containing
pale granules, in the spore cavity. Young spores were found to take the
form of Myxidium type. In mature spores, he always found two nuclei by
staining.

*Misprmted in Schroder's pap)er.

359] STUDIES ON MYXOSPORIDIA—KUDO 121

SPHAEROMYXA HELLANDI Auerbach
[Figs. 323 and 324]

1909 Sphaeromyxa hellandi Auerbach 1909 : 78-79

1910 Sphaeromyxa hellandi Auerbach 1910b : 772-774
1910 Sphaeromyxa hellandi Auerbach 1910c : 174^175

1912 Sphaeromyxa hellandi Auerbach 1912 : 4, 40

Habitat: Gall-bladder of Molva vulgaris Flem., Centronotus gunellus,
Brosmius brosme Asc; Bergen, Torghatten.

Vegetative form: Large and rounded disc form. Protoplasm is dis-
tinctly differentiated into ectoplasm and endoplasm. Thickness up to
160/i, folded in the bladder. Ectoplasm finely granular; in unstained speci-
mens, it is recognizable as 10 to 12jli thick layer, in which about 2n thick
radially striated outer layer and 8 to 10/x thick inner finely granular region
can be distinguished. In stained sections, the outer layer remains
unstained. Endoplasm highly alveolar, contains refractive granules of
different size which are not stained with Sudan III. Each pansporoblast
develops into two spores. Polysporous.

Spore: Arch form in front view. The degree of curvature varies
greatly. Sutural line curved in S-shape and well marked. Both ends more
or less truncated. Polar capsule at each end. Polar filament being wound
along the longest axis of the polar capsule and is extruded with KOH.
Sporoplasm rounded with one or two nuclei. Dimensions: length 20.8 to
26;u, breadth and thickness 5 An, length of polar capsule 10 to lO.Sju.

SPHAEROMYXA EXNERI Awerinzew
[Figs. 325 and 326]

1913 Sphaeromyxa exneri Awerinzew 1913a : 155

Habitat: Gall-bladder of Thysanophris japonicus; Lorenzo IVIarques
(Africa).

Vegetative form: Not observed.

Spore: Somewhat resembles that of S. hellandi Auer. in being bent
to one side with sutural line of S-form but differs in dimensions. Both
ends slightly tapering. Polar capsules two, one at each blunt end, in
which the polar filament is wound parallel to its longer axis. Sporoplasm
comparatively small and sharp-contoured, contains only one nucleus.
Dimensions: length 75 to 80/x, breadth 18 to 20/i, length of polar capsule
30 to 35m.

SPHAEROMYXA GASTEROSTEI Georgevitch
[Fig. 327]
1916 Sphaeromyxa gasterostei Georgevitch 1916 : 88

Habitat: Gall-bladder of Gasterosteus spinachia; Roscoff (September).
Vegetative form: Trophozoites form large plasmodia.

122 ILLINOIS BIOLOGICAL MONOGRAPHS (360

Spore: Large, elongated fusiform; ends less truncated than those of
the spore of Sphaeromyxa balbianii. As the spore becomes mature, the
ends assume more pointed shapes. Polar capsules two, one at each end.
Sporoplasm with two nuclei, fills the extracapsular cavity. Dimensions:
twice or three times larger than those of Sphaeromyxa balbianii Thelohan.

Genus ZSCHOKKELLA Auerbach
1910 Zschokkella Auerbach 1910 : 62-63

1910a : 240-256
1910c : 175

The characters of the genus are described on page 58.
Type species: Zschokkella hildae Auerbach.

ZSCHOKKELLA HILDAE Auerbach
[Figs. 328 to 331]
1910 Zschokkella hildae Auerbach 1910 : 62-63

1910 Zschokkella hildae Auerbach 1910a : 240-254

1912 Zschokkella hildae Auerbach 1912 : 40-41

Habitat: Urinary bladder of Phycis blennioides Br., Gadus aeglefinis,
G. callarias L., G. virens L.; Norway.

Vegetative form: Trophozoites float in the bile or attach themselves
to the epithelial layer of the bladder. Youngest ameboid form about
4.5 to 6/x. In floating form, pseudopodia more or less long, lobose, are
formed; while in the attached form those similar to the pseudopodia of
Myxidium bergense Auer. are developed. Plasmogamy occurs. Size
varies greatly according to the number of spores which are formed in each
individual. Monosporous (with or without the remnant of protoplasm),
disporous and polysporous (up to 4 spores).

Spore: Semicircular in front view, with slightly and equally attenuated
ends. At each end, large spherical polar capsule is situated which opens
not at the extremity, but on the flat surface. Shell bivalve and thick.
Sutural line S-form. Sporoplasm with two nuclei. Dimensions: length
16 to 28.8/i, breadth 13 to 18^i, polar capsules 5.6 to 7.2/i in diameter, length
of polar filament 72m (KOH).

ZSCHOKKELLA NOVA Klokacewa
[Figs. 332 and 333]
1914 Zschokkella nova Klokacewa 1914 : 184-186

Habitat: Gall-bladder of Carassius vulgaris; Russia?
Vegetative form: Not observed.

Spore: Outline irregular. Observations on fixed materials alone. Two
large round polar capsules open at the side near ends. Sporoplasm with
two nuclei. On some spores, striations that run parallel to the sutural line
were observed. Dimensions: length 9.5 to 11.5/i, breadth 6.5 to 7/t,
diameter of polar capsule 3 to 3.5m.

361] STUDIES ON MYXOSPORIDIA—KUDO 123

ZSCHOKKELLA ACHEILOGNATHI Kudo
[Figs. 334 to 338]

1916 Zschokkella acheUognathi Kudo 1916 : 3-5

Habitat: Gall-bladder and gall-duct of Acheilognathus lanceolatum
Temm. et Schl.; Tokio (May). Over 80% of the fish examined were
found to be infected.

Vegetative form: Disc-shape. In bile duct, large trophozoites are
folded up. Body colorless and transparent. Protoplasm is well defined
into two regions. Ectoplasm finely granular in vivo. In stained sections,
it shows two layers; thin outer layer (2/x thick) presents very fine striations,
while inner layer (6 to 8/i thick) is finely vacuolated without any enclosure.
Endoplasm is highly vacuolated. Lobose pseudopodia formed only in
younger individuals (15 to 30/i in diameter), in which ameboid movements
are not slow. Size: up to 720/i by 550ju, thickness 5 to 30/i. Polysporous.

Spore: Form resembles Zschokkella hildae, but slightly elongated.
Form varies to some extent. Some spores are of Myxidium type. Sutural
line curved. Longitudinally striated. Spherical polar capsule at each end
opening near the extremity. Dimensions: length 10 to 14/*, breadth 6 to 7/x,
diameter of polar capsule 3 to 5/x, length of polar filament 65 to 70/i (KOH).

ZSCHOKKELLA GLOBULOSA Davis
[Figs. 339 and 340]

1917 Zschokkella globulosa Davis 1917 : 236

Habitat: Urinary bladder of Spheroides maculalus; Beaufort (August).

Vegetative form: Rounded; slowly ameboid, forming short, lobose
pseudopodia. Body colorless and transparent. Ectoplasm not distinct.
Protoplasm granular, characterized by the presence of several large fat
globules. Sporulating trophozoites about 15 to 16/x in diameter. Mono-
sporous and disporous.

Spore: Semicircular. Sutural line twisted on its axis and oblique to
longitudinal axis; sutural ridge distinct. Polar capsules opening on flat
surface. Sporoplasm finely granular and very transparent. Dimensions:
length 11/x, breadth 7/t, diameter of polar capsules 3/i.

Family MYXOSOMATIDAE Poche
1913 Myxosomatidae Poche 1913 : 230

The characters of the family are described on page 58.

Genus MYXOSOMA Thelohan

1892 Myxosoma Th61ohan 1892 : 175

The characters of the genus are described on page 58.
Type species: Myxosoma dujardini Thelohan.

124 ILLINOIS BIOLOGICAL MONOGRAPHS [362

MYXOSOMA DUJARDINI Thelohan

[Figs.

341 to 343]

1841

MiiUer

1841

:486-487

1845

Dujardin

1845

:644

1892

Myxosoma dujardini

Thdohan

1892

:175

1895

Myxosoma dujardini

Thflohan

1895

-.343-344

1905

Myxosoma dujardini

Nufer

1905

: 77, 79, 186

1910

Myxosoma dujardini

Wegener

1910

: 72-73

1916

f Myxosoma dujardini

Kudo

1916

:3

Habitat: Branchial lamellae of Scardinius erythrophthalmus L., Perca
fiuviatilis, Leuciscus rutilus L. and Cyprinus carpio L.; France, Frisches
Haff, Kurisches HafF (February, April, May), Tokio (May), Switzerland.

Vegetative form: White cysts being branched, rounded, spherical or
irregular; 1 to 1.5mm. in diameter.

Wegener's form 1 to 1.7mm. long.

Spore : Ovoidal, flattened, with attenuated anterior end which is slightly
bent laterally. Two pyriform polar capsules at the anterior end. Sporo-
plasm without any iodinophilous vacuole. Dimensions: length 12 to
13n, breadth 7 to 8/i.

Wegener's form: polar capsules 6jj, by 3/*.

Kudo's form: polar capsules 6 to Ifx by 2/i, length of polar filament TO/*.

MYXOSOMA (?) LOBATUM Nemeczek

[Fig. 348]

1911 Myxosoma (?) lobalum Nemeczek 1911 : 160-162

Habitat: Branchiae of Leuciscus leuciscus L. and Aspius rapax Ag.;
Austria.

Vegetative form: Cysts spherical, oval or elongated; of white color.
Size from 0.5 to 3mm. by 0.5 to 1mm. Those in Aspius rapax, oval to
spindle-shape, 1 to 3mm. long and 1 to 1.5mm. wide.

Spore: Ovoidal; anterior end narrowly pointed and straight; posterior
end rounded, with lobose appendix (about 6/x long). A transverse fold
on the shell behind the polar capsules in fresh as well as preserved spores.
No iodinophilous vacuole. Dimensions: length 12. 6/x, breadth 8.2/t,
length of polar capsule 4.2^, length of polar filament 80 to 90/x. Spores
found in Aspius rapax, had slight difference in dimensions, the structure,
however, being similar to the above.

Remarks: Nemeczek doubts if this form is really Myxosoma because
of the following: 1) different shape of the cysts compared with that of
the type species as described by Thelohan; 2) spores observed might
develop later into other forms like Henneguya.

363] STUDIES ON MYXOSPORJDIA— KUDO 125

MYXOSOMA FUNDULI Kudo
[Figs. 344 to 347]
1918 Myxosoma fundidi Kudo 1918 : 12-14

Habitat: Branchiae of Fundulus majalis Wal. and F. heteroclitus L;
Woods Hole (August, September).

Vegetative form: Cysts. Spherical and small; 150/* in average diame-
ter. Largest form observed 360/x by 264)u. Spores, young and mature,
were found in the cysts. Polysporous.

Spore: Pyriform. Shell uniformly thick with 7 to 10 folds on sutural
edge at the posterior portion. Sutural ridge, fairly well marked. Two
polar capsules pyriform and of equal size at the anterior end. Sporoplasm
finely granular with two nuclei but without any iodinophilous vacuole.
Dimensions: length 14/*, breadth 8/i, thickness 6/i, polar capsule 8/i by 2yLi,
length of polar filament 38 to 42/i (perhydrol, KOH).

Remarks: The writer could not find any evidence of an iodinophilous
vacuole by treatment with various iodine mixtures, which is the most
important characteristic of the genus. Hahn's form {Myxobolus funduli,
p. 151) should be distinguished from the present form.

Genus LENTOSPORA Plehn
1905 Lentospora Plehn 1905 : 150

The characters of the genus are described on page 58.
Type species: Lentospora cerebralis (Hofer) Plehn.

LENTOSPORA CEREBRALIS (Hofer) Plehn

[Figs. 349 to 354]

1903

Myxobolus cerebralis

Hofer

1903 :

8

1904

Myxobolus chondrophagus

Hofer

1904:

:53

1905

Lentospora cerebralis

Plehn

1905 ;

; 145-166

1909

Lentospora cerebralis

Plehn

1909 :

:38

1910

Lentospora cerebralis

Auerbach

1910c

:176

Habitat: Cartilage and perichondrium of Trulta iridea Gibb., Salmo
fontinalis Mitch., Trutta salar L.; Germany (Karlsruhe and other localities).

Vegetative form: Ameboid form. Size varies greatly. Small ameboid
form probably grows up into large individual which has often fifty or more
ringform nuclei and breaks up into numerous small forms by division. No
sporous character is observed except a figure of a disporous form.

Spore: Circular in front view; lenticular in side view, with more or less
extensive variation in length and breadth. Shell smooth. Sutural ridge
distinctly thickened. Two polar capsules pyriform and convergent, are
usually of same size. Extruded polar filaments cross each other. Sporo-
plasm with two ring-form nuclei but without any iodinophilous vacuole.

126 ILLINOIS BIOLOGICAL MONOGRAPHS [364

Dimensions: diameter 6 to 10/*, length of polar capsule 2/5 that of the
spore, length of polar filament 40 to SOfi (limewater, 1% KOH).

Remarks: Plehn noticed that the present form causes the chronic
form of "Drehkrankheit" among young fish in German waters. She was
unable to extrude the polar filament with mineral (?) acids. Auerbach,
however, could extrude the filament by means of acids.

LENTOSPORA MULTIPLICATA Reuss

[Fig. 355]
1906 Lentospora multiplicata Reuss 1906 : 203

Habitat: Muscle of Idus melanotus Heck. ; Volga?, Russia.

Vegetative form: Not described.

Spore: Oval. Sutural edge broad with many folds. No iodinophilous
vacuole. Dimensions: length 12^, breadth 9.5m, thickness 6/i, polar
capsules 4/t by 2.25/*.

LENTOSPORA ENCEPHALINA Mulsow

1911 Lentospora encephalina Mulsow 1911 : 483-485

Habitat: In the blood vessel of the brain, especially of the mid-brain of
Cyprinus carpio L.; Munich (spring). Blood vessels are the only seat of
infection. In most cases many individuals lie parallel to one another. The
infection occurs frequently and heavily. The effect, however, is undeter-
mined.

Vegetative form: Trophozoite elongated, worm-like and circular in
cross section. The body is covered with a pellicula. The protoplasm is
distinctly differentiated into homogeneous ectoplasm layer and inner endo-
plasm. In the latter are found numerous granules, small nuclei and spores.

Spore : Almost circular in front view ; profile ? No iodinophilous vacuole
is found. The polar filament is easily extruded by means of a highly diluted
KOH solution. Diameter: 5 to 5.5^.

LENTOSPORA ASYMMETRICA Parisi
[Figs. 356 to 359]

1912 Lentospora asymmetrica Parisi 1912 : 292-293

Habitat: Connective tissue of kidney of Crenilabrus pavo C. et V.;
Napoli (September).

Vegetative form: One trophozoite found; a small, rounded form with
thin and hyaline ectoplasm which could be distinguished from the endo-
plasm with coarse yellowish globules, containing two spores. Disporous?

Spore: Oval from the front; flattened and fusiform in profile. Sutural
edge with many triangular folds, which are more clearly seen in material

365] STUDIES ON MYXOSPORIDIA—KUDO 127

preserved in formalin than in fresh condition. Two polar capsules of same
size, are situated asymmetrically, opening at the side near the anterior
end. Sporoplasm granular and with two nuclei, but without any iodino-
philous vacuole. The polar filament not being extruded by ordinary
reagents, probably because the spores were not full-grown. Dimensions:
length 10 to ll)u, breadth 6.5 to 7/i, length of polar capsules Six.

LENTOSPORA ACUTA (Fujita) Kudo

[Figs. 360 to 362]

1912 Sphaerospora acuta Fujita 1912 : 260-261

Habitat: Epithelium of branchial lamellae of Carassius auratus L.;
Sapporo, Nippon.

Vegetative form: Fujita's description is simply as follows: Sporoblast
contains about two spores.

Spore: Spherical in front view, with slightly pointed anterior end;
spindle shaped in side-views. Shell thin and smooth. Two convergent
polar capsules are of different sizes, occupying about 5/8 in space of the
spore. No vacuole could be made out in sporoplasm. Dimensions:
length 8 to lO^t, breadth 7 to Sn, thickness 5 to 6/x, polar capsules 5^ by Afi.

Remarks: This species, recorded incompletely by Fujita as Sphaero-
spora, shows characters of the genus Lentospora in spore form so that it
is provisionally placed here.

LENTOSPORA DERMATOBIA Ishii
[Figs. 594 to 596]
1916 Lentospora dermatobia Ishii 1916 : 472-474

Habitat: In the integument of Anguilla japonica Temm. et Schl.;
Shizuoka, Nippon. From the same specimen which harboured Myxidium
a«gwj^/ae, see page 114. The number of cysts reaches probably "several
hundreds."

Vegetative form: Cysts, beneath the epidermis, usually subcircular,
more or less irregularly triangular or quadrilateral under the magnifier,
with the largest diameter of from 142 to 267/*. The epidermis is slightly
lifted up by the cyst. No chromatophore on the surface of the cyst. The
cysts separated from each other, are found mostly in the central region of
the body, head and fins being free from cysts. In cross-section, cysts
exhibit oval or lenticular shape with the longest diameter, which is twice
as long as the depth, placed parallel to the surface of the skin. No partic-
ular pathological change was noticed.

Spore: Circular in front view; broad fusiform or lenticular in side view.
Sutural ridge fairly distinct. Sutural edge comparatively broad, especially

128 ILLINOIS BIOLOGICAL MONOGRAPHS [366

at the posterior margin, where a few folds (three are figured) are seen. Two
oval polar capsules convergent and of equal size. Sporoplasm is sharply-
contoured, no iodinophilous vacuole being recognized. Dimensions in
preserved material (?): diameter 6.3 to 7n, thickness 4.2 to 4.9/u, length
of polar capsule 2.8 to 3.5/x.

Family MYXOBOLIDAE Thelohan

1892 Myxobolidies Thdlohan 1892 : 173, 176

1893 Myxoholidae Gurley 1893 : 412, 413
1895 Myxobolidies Thdohan 1895 : 347

The characters of the family are described on page 58.

Genus MYXOBOLUS Biitschli
1882 Myxobolus ButschU 1882 : PI. 38 : 6-10

The characters of the genus are described on page 58.
Tjrpe species: Myxobolus miilleri Biitschli.

MYXOBOLUS MULLERI BiitschH

[Figs.

397 to 4031

1881

Batsrhli

1881

630

1882

Myxobolus miilleri

Butschli

1882

595

1895

Myxobolus miilleri

Thdlohan

1895

349

1905

Myxobolus miiUeri

Nufer

1905'

77, 79, 186

1906

Myxobolus miiUeri

C6p^de

1906

64-65

1906

Myxobolus miiUeri

Schroder

1906

195

1908

Myxobolus miilleri

Auerbach

1908

:456

1909

Myxobolus miiUeri

Auerbach

1909a

:54, 71

1910

Myxobolus miiUeri

Wegener

1910

:81

1911

Myxobolus miiUeri

Nemeczek

1911

160

1912

Myxobolus miiUeri

Parisi

1912

:293

Habitat: Air bladder and branchiae of Leuciscus cephalus L.; kidney
and ovary of L. phoxinus L.; eye of Crenilabrus melops L. and Alburnus
lucidus; branchiae of As pro as per L., Barbus vulgaris Flem., Leuciscus ruii-
lus L., Squalius cephalus L., S. agassizii Heckel, Lota vulgaris L., Phoxinus
loevis Ag. ; pseudobranchiae of Cottus gobio L. ; intestine of Mugil auratus
Risso.; France, Germany [Karlsruhe, AUe (October), Pregel, Frisches
Haff], Switzerland (Neuchatel Lake), Italy (Napoli, September).

Vegetative form: White cysts in the connective tissue. Form elongated
oval, 2 to 3mm. in diameter. No clear differentiation of protoplasm is
observed even in young forms. In sections, some cysts show radiate
striations in the thick granule-free ectoplasm. Endoplasm filled with nuclei.

Cepede writes as follows : Cysts in branchiae, subspherical or elliptical,
1.5mm. by 0.5mm.

Wegener's form: Cysts small and rounded, 0.2 to 0.3mm. in diameter.

Spore: Ordinarily spherical or subspherical. Two polar capsules

367] STUDIES ON MYXOSPORJDIA—KUDO 129

with a small triangular intercapsular appendix. Polar capsules pyriform
and of same size. Sutural edge exhibits folds (7 to 9).

Thelohan's dimensions: length 10 to 12)Lt, breadth 9 to ll/x, length of
polar capsule 5/i.

Cepede gave the following dimensions in vivo: length lO/x, breadth
9/x, thickness 6/t, length of polar capsule 5/ix.

Wegener's form. Usually oval, often almost spherical. Length 10 to
11/x, breadth 8 to 9/i, diameter of spherical form 9^, polar capsule 4 to
5/i by 2 to 2>n.

MYXOBOLUS PIRIFORMIS Thelohan

[Figs. 363 to 364]

1852

Remak

1852 :

:144

1883

Balbiani

1883

: 197-198

1884

Balbiani

1884:

:125

1891

Pfeiffer

1891

:132

1892

Myxobolus piriformis

Th61ohan

1892 :

:177

1893

Myxobolus piriformis

Gurley

1893 :

:414

1894

Myxobolus piriformis

Gurley

1894 ;

:211

1895

Myxobolus piriformis

Th61ohan

1895

:348

1905

Myxobolus piriformis

Nufer

1905

: 77, 186

1910

Myxobolus piriformis

Plehn

1910

: 22-27

1910

Myxobolus piriformis

Wegener

1910

:73

Habitat: Branchiae, spleen, kidney of Tinea tinea L., Cohitis fos silts L.
and subcutaneous connective tissue, spleen, liver, connective tissue of the
intestine of Leuciscus sp.; France, Germany (Pregel), Switzerland.

Vegetative form: Small, long thread-like cysts. Color white. Poly-
sporous.

Wegener's form: average size, length 1mm., breadth 0.09 to 0.1mm

Spore: Elongated oval; flattened. Anterior end highly attenuated
and slightly bent to one side. One pyriform polar capsule at this end.
Dimensions: length 16 to 18/i, breadth 7 to 8/i, length of polar filament
30/x.

Wegener gives the following dimensions: length 18/x, breadth 7. 5/i,
polar capsule 7. 5/i by 3. 5/i.

MYXOBOLUS UNICAPSULATUS Gurley
[Figs. 365 to 366]

1841 Miiller 1841 : 487

1893 Myxobolus unicapsulatus Gurley 1893 : 414

1894 Myxobolus unicapsulatus Gurley 1894 : 210-211

Habitat: In the skin of Labeo niloticus For.; Nile.

Vegetative form: Cysts very small pustules in the skin of the head.

130 ILLINOIS BIOLOGICAL MONOGRAPHS (368

Spore: Form similar to Myxosoma dujardini. A single polar capsule
at the anterior end, obliquely directed. Dimensions: length 0.0051'",
breadth 0.0034'".

MYXOBOLUS FUHRMANNI Auerbach
[Fig. 367]

1909 Myxobolus fuhrmanni Auerbach 1909 : 65-68

1910 Myxobolus fuhrmanni Auerbach 1910c : 178-179

Habitat: Connective tissue under the mucous membrane of the mouth
of Leuciscus rutilus L.; Neuchatel Lake.

Vegetative form: Cysts, of pea-size, surrounded by several membra-
nous layers of connective tissue with a few nuclei. Finely granular ecto-
plasm forms outer layer. Endoplasm is dense and contains faintly stained
nuclei. Pansporoblasts and spores are found in the central portion of the
cyst. Polysporous.

Spore: Elongated pyriform, with attenuated anterior and rounded
posterior ends. Majority with a single polar capsule; spores with two
polar capsules were also observed. Shell thick, at the posterior end. 4 to
6 notch-like markings on the posterior part of the shell. Sutural ridge
thickened and fairly well marked. Coiled polar filament visible in pre-
served material. The opening of the polar capsule is either at the anterior
end or near it. Sporoplasm with two nuclei of unequal size and a com-
paratively large iodinophilous vacuole, stained brown with iodine alcohol.
Dimensions: length 18 to 20fx, breadth about 8fx, thickness 6fi, length of
polar capsule 9 to lO^t.

MYXOBOLUS OCULI-LEUCISCI Trojan
[Fig. 368]

1909 Myxoholus oculi-leucisci Trojan 1909 : 679-682

Habitat: Vitreous body of the eye of Leuciscus rutilus L.; Prague
(May?).

Vegetative form: Two cysts, spherical and subspherical, 100 to 180/i in
diameter. Ectoplasm finely granular. Outer portion of endoplasm with
small nuclei, then larger nuclei each surrounded by protoplasm, while the
central portion contains spores. Polysporous.

Spore: Elongated oval, flattened dorso-ventrally. Posterior margin
rounded. At the anterior end, a single polar capsule with distinctly visible
coiled polar filament. Shell smooth without any markings. Sporoplasm
with one nucleus, usually elongated oval (2.8/i in diameter) and one
vacuole, occupies more than half of the space of the spore. Dimen-
sions: length 9 to lOjit, breadth 4.5 to 5.5/i, thickness 3n, polar capsule
5ju by 2fi.

369) STUDIES ON MYXOSPORIDIA—KUDO 131

MYXOBOLUS TOYAMAI Kudo
[Figs. 369 to 370]

1915 Myxoholus toyamai Kudo 1915 : 517-523
1917 Myxoholus toyamai Kudo 1917 : 163-170

Habitat: — Connective tissue of branchial lamellae of Cyprinus carpio
L.; Tokio (July).

Vegetative form : Cysts, ovoidal or in shape of calabash. Small form 67
by 50/1, shows clear differentiation of protoplasm. Ectoplasm radially
striated, often, differentiates fine processes (2 to 3/i long). Endoplasm
coarsely granular, contains nuclei from 1 to 4/i in diameter. Size up to 190/x
in greatest diameter in sections. Two spores are formed in each pansporo-
blast. Polysporous.

Spore: Pyriform, with attenuated anterior and rounded posterior ends.
No bilateral symmetry. Lateral sides are curved. Calabash shaped
spores often occur. Shell without any marking, thickened at the anterior
end. Sutural ridge shows sometimes a short {l.Six long) tail-like process at
the posterior tip. A single pyriform polar capsule at the anterior end; in
stained preparations, a small, oblong mass of protoplasm is seen between
the polar capsule and the shell. Coiled polar filament distinct. Sporo-
plasm with two nuclei of usually same size and a relatively large iodino-
philous vacuole, 3)U in diameter. Dimensions: length \Sn, breadth 7 to 8ju,
thickness 5 to 6/i, polar capsule 7 to 8/* by 3 to 4/i, length of polar filament
40 to 45m (pressure, perhydrol, KOH).

MYXOBOLUS NOTATUS Mavor
[Figs. 371 to 372]

1916 Myxoholus notatus Mavor 1916a : 70-71

Habitat: Connective tissue of the voluntary muscles on the sides or
tail of Pimephales notatus Raf.; Georgian Bay, Canada (Summer).

Vegetative form: Cysts as large as 3mm. in diameter, are surrounded
by a layer of columnar epithelial cells (origin and significance?) and a dense
layer of connective tissue. Protoplasm is not clearly differentiated, tho
the cyst is surrounded by an area devoid of nuclei. In the outer region of
endoplasm, numerous nuclei each with a caryosome, are recognized. In
the course of spore formation, two nuclei for polar capsules appear at first,
one of which degenerates later. Polysporous.

Spore: Pyriform, with a posterior extension forming a process, 5/i in
length and as broad as the spore. A single polar capsule at the anterior
end. An iodinophilous vacuole in the sporoplasm. Dimensions: length
17 to 18/c, breadth 7.5 to 8/t, polar capsule 7/t by 4/i, length of polar filament
95/i.

132 ILLINOIS BIOLOGICAL MONOGRAPHS [370

MYXOBOLUS sp. Kudo
1918 Myxobolus sp. Kudo 1918 : 15

Habitat: Spleen of Perca flavescens; West Falmouth, Mass. (August).
Isolated spores were noticed in one fish, in smears and section preparations.

Vegetative form: Not observed.

Spore: Ovoidal, attenuated at the anterior end. Shell uniformly thick.
A single polar capsule opens at the anterior tip. Sporoplasm contains an
iodinophilous vacuole and two nuclei of equal size (2)u). Dimensions:
length 18 to 20/1, breadth 8/x, polar capsule 7 to 9/* by 3 to 6/x.

MYXOBOLUS ROHITAE Southwell et Prashad

[Figs. 373 to 374]

1918 Myxobolus rohitae Southwell et Prashad 1918 : 344-347

Habitat: Branchiae of Labeo rohita; Turag river, Mirpur, Dacca
district, Bengal (June). Type specimens of Indian Museum P48/1.
Infection was heavy. In one case SZ cysts were found on one surface of
a single gill.

Vegetative form: Cysts in the gill-filaments. The cysts preserved in
alcohol are of a creamy-yellow color, oval to cylindrical in form, lying with
the long axis parallel to the gill- filaments. Cysts attached to the gill-
filaments with the flattened surface. Size: length 3.1 to 3.8mm.; breadth
0.8 to 1.2mm. Cyst- wall striated vertically, covered with an epithelium,
two to three layers thick. In the central portion and at the periphery,
mature spores and pansporoblasts as well as immature spores were found
respectively. Polysporous.

Spore: Elongated pyriform, rounded at the posterior end and acutely
pointed anteriorly. Sutural ridge slightly raised. One polar capsule
present, being of conspicuous size. Coiled polar filament is distinctly
observed in the polar capsule. An iodinophilous vacuole, 3.6/x in diameter,
in the sporoplasm. "Lying just posterior to it is the nucleus of the spore.
A few granules of chromatin were also seen lying scattered in the proto-
plasm." Dimensions: length 30 to 32/i, breadth 7 to 8/x, length of the polar
capsule 22 to 23/*, that of polar filament 92 to 97/i.

MYXOBOLUS SENI Southwell et Prashad

[Figs. 375 to 376]

1918 Myxobolus seni Southwell et Prashad 1918 : 347

Habitat: On the median and caudal fins of Labeo rohita; Mirpur,
Dacca (January). Type specimens in Indian Museum numbered P 53/1.

Vegetative form: Trophozoites form cysts which are elongated ellip-
soidal. Size from 4.7mm. to 5.4mm. in length, 2.9mm. to 3.7mm. in

371]

STUDIES ON MYXOSPORJDIA—KUDO

133

breadth. Color of the cyst whitish with black scattered granules on the
surface.

Spore: Oval, much wider behind than in front and pointed at the
anterior end. Sutural ridge is slightly thickened. A single polar capsule,
showing much coiled polar filament. lodinophilous vacuole 2.3m in diame-
ter. Dimensions: length 13.2 to 13.6/i, breadth 10.1 to 10.3)li, length of
polar capsule 4/i, length of polar filament 43^1 (in one case) .

MYXOBOLUS MISGURNI nov. spec.

[Figs. 377 to 378]
1916 Myxobolus fuhrmanni Kudo 1916 : S

Habitat: GaW-hladder oi Misgurnus anguillicaudatus ; Tokio (Septem-
ber). About 50% of the fish examined showed a few isolated spores
floating in the bile.

Vegetative form: Unobserved.

Spore: Form elongated pyriform, with attenuated anterior and rounded
posterior ends. Shell uniformly thick. Over sutural edge, shell exhibits
many (up to 12) triangular markings. Sutural ridge distinct. A single
pyriform polar capsule at the anterior end. Sporoplasm contains an
iodinophilous vacuole and two nuclei. Coiled polar filament distinct in
vivo. Dimensions of fresh spores: length 14 to 15.5/x, breadth 6 to 7.3^,
thickness 5 to 6fi, polar capsule 6.3/i by 2 to 3n, length of polar filament up
to lOOju.

Remarks : The writer reported this species as identical with Myxobolus
fuhrmanni Auerbach. By repeated reexamination and comparison with
Auerbach's description, however, he came to the conclusion that the
present form should be treated as a new species, on account of the difference
of the host and the characters of the spore.

MYXOBOLUS PFEIFFERI Thelohan
[Figs. 379 to 385]

1890

Myxosporidian

Pfeiffer

1890;

: 30-37

1891

Myxosporidian

Pfeiflfer

1891 ;

: 100, 105-110, 130

1893

Myxosporidian

PfeiflFer

1893;

: 118-130

1895

Myxobolus pfeifferi

Th61ohan

1895

:350

1898

Myxobolus pfeijfferi

Doflein

1898

: 306, 320, etc.

1906

Myxobolus pfeijfferi

C6pede

1906

:59

1906

Myxobolus pfdferi

Stazzi

1906

:14hl9

1906

Myxobolus pfdferi

Mercier

1906
1906a

: 427^28;
: 763-764

1908

Myxobolus pfeiferi

Keysselitz

1908

: 253-273,
286-306

1909

Myxobolus pfeiferi

Mercier

1909

:.5-30

134 ILLINOIS BIOLOGICAL MONOGRAPHS [372

Habitat: Muscle and connective tissue of kidney, spleen, intestine,
ovary, etc., of Barbus barbus L., and branchiae of B. fluviatilis Ag. and
B. plebejus Val.; Drac (June), Neckar, Prag, Milano. The cause of well
known ''Boil disease" (Beulenkrankheit) or Myxoboliasis tuberosa (Hofer)
of the barbels in European waters. Among many observers Keysselitz
made a thoro study of the parasite. His observations are as follows: The
disease occurs among the fish at any stage of growth. About 8% of the
fish, 7 to 15cm, long, caught in May and June between Conz and Trier
were infected with the parasites. The heaviest infection, however, occurs
among fish up to 40cm. in length; fish 50cm, long or larger show the tumors
caused by the parasites, rather rarely. Most of the fish die as the result
of the infection between the early part of April and the end of October.
The highest mortality is reached in the hottest months, i.e., July and
August. The temperature greatly affects the growth of the parasites.
Fish kept in the aquarium at a temperature of 25° C. or higher demon-
strate the growth of the boil in size daily. The boils are not noticed
during the winter and spring, they are formed from the early part of April
to the middle of October.

Vegetative form: The parasites develop tumors of conspicuous size.

Keysseltiz's observations are as follows: The tumor varies in size from
millet-grains to hen's eggs. Form spherical, oval or elongated. The
number of cysts on a single fish, is usually 3 to 4; often one, in some fish,
however, 23 were recognized on one fish. Usually tumors separated from
each other, rarely many forming one tumor. In one fish, 27cm. in length,
a tumor of 7cm. long, 4cm. broad and 3cm. thick, was observed in July.
The seat of infection is: the muscle of the body, muscle of pectoral and
anal fins, often in peritoneum and rarely in intestine. As the result of
breaking up of the cyst membrane, spores are also found in the testis, liver
and kidney.

The tumor is composed of many vegetative forms, rounded, oval,
elongated, variously branched or flattened. Size reaches to 1.5mm. in
diameter. Protoplasm is usually differentiated into ectoplasm and endo-
plasm. The surface is not often smooth, but shows irregular outline.
Ectoplasm is seen often as a very thin, uniformly hyaline, indistinctly
granular or radially striated layer, giving the network-like appearance to
the surface of the body. Endoplasm, stained more deeply around the
peripheral part than other portion, shows a coarsely alveolar structure in
the central region. It contains vegetative nuclei, developmental stages of
propagative nuclei, granules, fat-like, often leucocytes and red blood
'corpuscles. The leucocytes, uninuclear or multinuclear, were seen at the
periphery, apparently in the course of degeneration. Red blood corpuscles
were found, in section, inside of the apparently intact parasite. Each
pansporoblast develops into two spores. Polysporous.

373] STUDIES ON MYXOSPORIDAJ—KUDO 135

Cepede observed one cyst, about 2mm. in diameter, in the connective
tissue of the third gill arch.

Spore: Thelohan described as follows: Ovoidal. Sutural edge shows
folds. A small triangular intercapsular appendix. Dimensions: length
12/x, breadth 10/i. Cepede's form showed exactly the same dimensions.

Keysselitz gave the characters of the spore as follows:

Flattened oval. Shell smooth. A small intercapsular appendix.
Sutural edge having a number of small flat enlargement, size and number
being variable. Two convergent narrow canals (foramina) penetrate the
shell at the anterior end. Two polar capsules, pyriform and of equal or
nearly equal size, are located at the anterior half. Coiled polar filament
distinct, coiled 7 to 8 times. No distinct connection between polar
capsule and the filament. Sporoplasm fills the posterior half of the spore,
extending into intercapsular cavity. It is finely reticular, exhibits one or
two rounded or oval vesicular nuclei and an iodinophilous vacuole. Fat-
like substance is often seen around the polar capsules. Spores kept in water
for four months remain intact in large numbers. Dimensions: length 12
to 12.5/x, breadth 10 to lO.S/it, length of polar capsule, 5.5 to 6/x, length
of polar filament 28 to 34)u.

MYXOBOLUS INAEQUALIS Gurley
[Fig. 411]

1841 MUUer 1841 : 487-488

1893 Myxobolus inaequalis Gurley 1893 : 414

1894 Myxobolus inaequalis Gurley 1894 : 212

Habitat: In the skin of the head of Piramutana blochi Cuv. et Vil. and
Synodontis schall Bl. Schn.; Guiana, Surinam.

Vegetative form: Very small pustules in the skin of the head.

Spore: Ovoidal. Two polar capsules of unequal size at the anterior
end. Dimensions: length 0.0052'", breadth 0.0033'".

MYXOBOLUS DISPAR Thelohan

[Fig. 386]

1895

Myxobolus dispar

Thelohan

1895

:348

1904

Myxobolus dispar

Hofer

1904 ;

:50

1910

Myxobolus dispar

Wegener

1910

: 73-74

1911

Myxobolus dispar

Nemeczek

1911

:145

Habitat: Branchiae of Carassius carassius L., branchiae and epithelium
of intestine of Cyprinus carpio L., also muscle and spleen of Scardinius
erythrophthalmus L. and in the skin and the connective tissue of Alhurnus
lucidus Heck.; France, Austria, Konigsburg (March, July, September).

Vegetative form: Not described by Thelohan.

Wegener's description is as follows: Cysts: white in color; spindle shape
with pointed ends. Cysts in Carassius carassius L. smaller and oval.

136 ILLINOIS BIOLOGICAL MONOGRAPHS [374

Size 3.5mm. by 0.8mm. Cysts are surrounded by thick layers (7 to 8/li) of
the connective tissue of the host. Ectoplasm seems to be undifferentiated.
Endoplasm granular, contains a larger number of spores. Polysporous.

Spore: Thelohan's diagnosis is as follows:

Ellipsoidal or slightly oval. Shell with 3 to 5 folds along sutural edge.
Polar capsules of unequal size, mth a small intercapsular body. The
vacuole is difficult to stain with iodine. Dimensions: length 10 po
12/i, breadth Sju, polar capsule Tfx by 5/x. •

Wegener's form is as follows: length 11 to 12/i, breadth 7.5 to 8/x,
larger polar capsule 6 to 7n by 3.5jLt, smaller one 4yn by 2.5 to 3/i. The
sporoplasm is shifted toward the smaller polar capsule.

MYXOBOLUS ELLIPSOIDES Thelohan

[Figs.

387 to 389]

1852

Remak

1852

: 144-146

1892

Myxobolus ellipsoides

Th61ohan

1892

:177

1895

Myxobolus ellipsoides

Th61ohan

1895

: 350-351

1898

Myxobolus ellipsoides

Doflein

1898 ;

: 324, etc.

1905

Myxobolus ellipsoides

Nufer

1905

: 77, 79, 186.

1910

Myxobolus ellipsoides

Wegener

1910

:74r-75

1912

Myxobolus ellipsoides

Lo Giudice

1912

:l-79

Habitat: Connective tissue of air bladder, branchiae, kidney, spleen,
liver and cornea of Tinea tinea L., branchiae oi Ahramis brama L., Alburnus
lucidus Heck., Leueiseus rutilus L., Squalius eephalus L., Ahramis vimpa
Cuv., Blieea bjorkna L., Idus melanotus; France, Vierwaldstatter See,
Prague, Masurische See, Italy.

Vegetative form: Thelohan does not describe.

According to Wegener, white cysts, elongated oval; 2mm. by 0.5mm.
in size. Polysporous.

Spore: Thelohan described as follows: Flattened elliptical, rather
elongated. Sutural edge broad without any folds. Shell with no marking.
Form of the spore somewhat variable. Two polar capsules of equal size,
capsulogeneous nuclei present even when fully grown. Abnormal spores
are of frequent occurrence. Dimensions: length 12 to lAfi, breadth 9 to
ll/i, length of polar capsule 4/x.

Wegener's form: length 14 to 15/i, breadth 10 to ll/t, polar capsule
4 to 5n by 3fi. Shell comparatively thick. One spore had a tail 5/i long.

MYXOBOLUS EXIGUUS Thelohan

[Figs.

390 to 395]

1891

Myxosporidium mugilis ?

Perugia

1891

:23

1895

Myxobolus exiguus

Thaohan

1895

: 349-350

1906

Myxobolus exiguus

Schroder

1906

:195

1910

Myxobolus exiguus

Wegener

1910

:75

1912

Myxobolus exiguus

Parisi

1912

: 294-295

375] . STUDIES ON MYXOSPORJDIA— KUDO 137

Habitat: Branchiae of Ahramis brama L. and Chondrostoma nasus L.,
wall of stomach, pyloric coecum and intestine, branchiae, spleen, kidney of
Mugil chelo Cuv., M. capita Cuv. and M. auratus Riss.; Le Vivier-sur-mer,
Banyuls, Marseille, Heidelberg, Pregel, Frisches Haff, Kurisches HafiF,
Geneva, Napoli.

Vegetative form: No description by Thelohan.

Wegener writes as follows:

Cysts of variable size. Color white. Usually small and narrow, 0.5
to 0.7mm. long and 0.2mm. wide. Frequently large round cysts of 1.2 to
1.5mm. in diameter, filling the lamella. Cysts are surrounded by 10 to
1 l/i thick membrane composed of the connective tissue of the host. Ecto-
plasm, 5/x thick, is faintly stained by hematoxylin. Outer region of endo-
plasm, alveolar and densely loaded with nuclei, while in the central portion
with mature spores in granular ground-mass.

Parisi's observations are as follows:

Cysts in the intestinal wall of Mugil auratus, large; reaching a length
of 3mm.

Spore: Thelohan's description is as follows:

Flattened ovoidal, with more or less attenuated anterior end. Sutural
edge shows fairly noticeable folds. A small triangular intercapsular appen-
dix. Vacuole in the sporoplasm is usually hard to stain with iodine.
Dimensions: length 8 to 9^i, breadth 6 to In, length of polar filament
15/i (KOH).

Wegener observed as follows: Rounded with slightly pointed anterior
end. Length 8 to 9. 5/x, breadth 6 to 7.5ju, polar capsule 4.5/i by 2 to 3/i.
Shell exhibits small folds around the sporoplasm. An intercapsular triangu-
lar body indistinctly visible.

Parisi gave the following dimensions: length 8 to 8.5ju, breadth 6 to 7/i,
thickness 5.5m, polar capsule 3 to An by 1.5 to 2ju, length of polar filament
30ybt (alkaline). Folds usually 6 in number. Coiled polar filament visible

m vivo.

MYXOBOLUS OVIFORMIS Thelohan

[Fig. 396]

1854

Lieberkahn 1854

: 21-22

1892

Myxobolus oviformis

Th61ohan 1892

:177

1895

Myxobolus oviformis

Th6Iohan 1895

:351

1905

Myocobolus oviformis

Nufer 1905 ;

: 77, 186

1906

Myxobolus oviformis

CdpMe 1906

:60

1910

Myxobolus oviformis

Wegener 1910

: 76-78

Habitat: Fin (subcutaneous tissue), spleen, kidney and liver of Gobio
gobio L.; branchiae of Alburnus lucidus Heck., Cyprinus carpio L., Blicca
bjorkna L., Abramis brama L. and A. vimba L. ; France (Isere), Frisches
HafiF (especially spring months), Switzerland.

138 ILLINOIS BIOLOGICAL MONOGRAPHS [376

Vegetative form: Thelohan gave no description.

Wegener's observations are as follows:

Cysts, white, 0.75 to 1.7mm. by 0.4 to 0.7mm. In sections, cysts are
shown to be surrounded by a thick (10 to 20/*, average 16/*) layer of connec-
tive tissue. Ectoplasm a thin (6 to 8/i thick) layer, exhibits a transverse
striation. The striation is often absent at places in ripe cysts. Endoplasm
finely granular. In young cysts, it is, however, reticulated, with nuclei of
1.5/i in diameter.

Spore: Thelohan described as follows: Flattened ovoidal with pointed
anterior end. Shell smooth. No folds. Polar capsule comparatively large.
Dimensions: length 10 to 12/i, breadth 9/x, polar capsule 6/i.

Cepede observed numerous spores in the liver and kidney of Gobio
gohio. Dimensions in vivo: length 10 to 12/i, breadth 9/*, length of polar
capsule 6/1. Polar capsules of equal size. Coiled polar filament distinct.

Wegener's form: length 10.5 to ll/z, breadth 7.5 to 8/i, polar capsule 5
to 6/i by Zn.

Remarks: Wegener recognized another form, which seems to be of very
rare occurrence and which can not be distinguished distinctly from the
above described form. Cysts at the end of the branchial lamellae. Size
1.7 to 2mm. in largest length. Spore resembles more closely the figure
given by Thelohan for Myxobolus oviformis than the above mentioned form
which he observed. A small intercapsular appendix (rounded) indistinct.
Sporoplasm comparatively small. Length 12.5 to 13.5/*, breadth 9/*,
polar capsule 7.5/i by 3/*.

MYXOBOLUS LINTONI

Gurley

[Figs. 404 to 408]

1891
1893

1894

Linton

Myxobolus lintoni Gurley
Myxobolus lintoni Gurley

1891
1893
1894

: 99-102

:414

:238

Habitat: Superficial musculature and subcutaneous tissue of Cyprino-
don variegatus; Woods Hole (August).

Vegetative form: Cysts, not closed, but fungoid masses of an irregular
shape, varying in size from 4mm. by 2.5mm. to 10mm. by 4mm., projecting
as much as 3mm. above general surface of skin. The skin of the host overly-
ing these tumors, is more or less cracked and broken, the scales being
scattered.

Spore: Elliptical in the front view; lenticular in side view. Shell
thick. Sutural ridge marked. Two polar capsules, convergent, at the
anterior end. Spores kept in alcohol, extruded polar filaments under the
action of iodine water and sulphuric acid. Sporoplasm with a large iodino-
philous vacuole. Dimensions: length 13.9/i, breadth ll/x, thickness 8/x.

377) STUDIES ON MYXOSPORIDIA—KUDO 139

MYXOBOLUS GLOBOSUS Gurley
[Figs. 409 and 410]

1893 Myxoholus globosus Gurley 1893 : 415

1894 Myxobolus globosus Gixrley 1894 : 241

Habitat: Branchial lamellae of Erimyzon sucetta ohlongus Lac. (Catos-
tomus tuberculatus Le Sueur); Kinston (N.C.), Columbia, (S.C., March),
tributaries of Fox River.

Vegetative form: Cysts, whitish, elongated elliptical or rod-shaped,
surrounded by very thin membrane? Size up to 0.5mm. in max. length.
PolysporoUs.

Spore: Globose, subcircular in outline. Shell thin and very transparent.
Sutural ridge very wide, being one third of the thickness of the spore.
Polar capsules two, of equal size, divergent. Vacuole present, but not
clearly contoured. Dimensions: length 7 to 8jw, breadth 6 to 7/t, thickness

MYXOBOLUS OBLONGUS Gurley

[Figs. 412 to 416]

1841 Miiller 1841 : 487-490

1893 Myxobolus oblongus Gurley 1893 : 414

1894 Myxobolus oblongus Gurley 1894 : 234^38

Habitat: Beneath the skin, chiefly of the head of Erimyzon sucetta
ohlongus Lac. {Catostomus tuberculatus Le Sueur); Kinston, tributaries
of Fox River.

Vegetative form: Cysts, round or elliptic, not over 1mm. in diameter,
covered by resistant membrane. Color whitish. Polysporous.

Spore: Spatular, approaching roundish-oblong. Shell thin and trans-
parent. Sutural ridge wide. Two polar capsules, pyriform, of equal size.
Sporoplasm extending forward along the upper surface. Vacuole could not
be detected. Dimensions: length 14 to 17/i, breadth 8.5/x, thickness 5
to 6/i.

MYXOBOLUS TRANSOVALIS Gurley
[Figs. 417 and 418]

1893 Myxobolus transovalis Gurley 1893 : 415

1894 Myxobolus transovalis Gurley 1894 : 242

Habitat: Under scales on external surface of Phoxinus (Clinostomus)
funduloides Girard; 4 Mile Run, Carlius, Va., tributary of Potomac River
(June), No fish of the same species caught from the same locality on
August 29 of the same year was found infected.

Vegetative form: It is not certain whether cysts exist or not. Spores
in mass, appear to be held together by a small gelatinous or mucoid mass

140 ILLINOIS BIOLOGICAL MONOGRAPHS [378

which has no attachment to the subjacent connective tissue. It forms
a thin discoidal mass situated in the center of the concave surface of the
scale. The color of the mass slightly more yellowish than the surrounding
tissue, when coagulated. It is exceedingly diflScult to detect its presence
in Vhe fresh state.

Spore: Elliptical, with the largest diameter passing thru two polar
capsules. Shell thin. Sutural edge narrow. Two polar capsules of equal
size convergent. Polar filament is extruded under the action of glycerine
and sulphuric acid. The vacuole in the sporoplasm is difficult to
detect. Sporoplasm also contains two nuclei, rarely one, 1 to 1.5/* in
diameter. Dimensions: length 6 to 7/i, breadth Sju. ^

MYXOBOLUS OBESUS Gurley

[Figs.

419 and 420]

1883

Balbiani

1883

:203

1893

Myxobolus obesus

Gurley

1893

:415

1894

Myxobolus ? obesus

Gurley

1894

.239

1899

Myxobolus obesus

Labb^

1899

:100

1906

Myxobolus obesus

C^pMe

1906

:60HS1

Habitat: On Alburnus alburnus L.; branchiae and kidney of A. lucidus
Heck. (A . jnirandella Bl.); Lac du Bourget.

Vegetative form: Balbiani gave no observation.

Cepede observed as follows: Cysts, ovoidal, more or less elongated
or variable in form, not exceeding 800ju in length. In kidney, numerous cysts
were of subspherical, ovoidal or rarely irregularly elongated form. Sub-
spherical cysts 500 to 600/x in average diameter. Polysporous.

Spore: Cepede describes as follows: Subcircular or ovoidal in front
view; lenticular in side view. Sutural edge exhibits variable numbers (4
to 5) of fold-like markings on the shell. Polar capsules pyriform and of
equal size. Coiled polar filament distinct. A small triangular intercapsular
appendix. Sporoplasm with a subspherical and clearly outlined vacuole
and two nuclei. Dimensions in vivo: length 11.5 to 12/li, breadth 7.5 to 8/i,
thickness 5^. Those of fixed and stained spores: length 11.25 to 11.50/*,
breadth 7.25 to 7.50/*, length of polar capsule 5/t.

Remarks: Cepede mentions that Alburnus alburnus L. mentioned by
Gurley is "without doubt" identical with A. lucidus Heckel.

MYXOBOLUS CYCLOIDES Gurley

[Fig. 421]

1841

Miiller

1841

: 481, 486

1893

Myxobolus cycloides

Gurley

1893

:41S

1894

Myxobolus cycloides

Guriey

1894

:239

1906

Myxobolus cycloides

C6pede

1906

: 61-63

1910

Myxobolux cycloides

Wegener

1910

: 79-80

379) STUDIES ON MYXOSPORIDIA—KUDO 141

Habitat: Opercle, pseudobranchiae and kidney of Leuciscus ruHlus;
branchiae of Scardinius erythrophthalmus, Blicca bjorkna L., Gohio gobio
L., Abramis vintba L., A. brama L., Rhodeus amarus Bl., Alburnus alburnus
L., Lota lota L.; France (Isere), Germany (Pregel, Frisches and Kurisches
Haff, Masurische See, January, May).

Vegetative form: Wegener observed cysts as follows. A type: 1 to
2mm. by 0.4 to 0.7mm. Form exactly like that of Myxobolus oviformis.
B type: small and round, present in groups. C type: small 0.5mm. by
0.2mm.

Spore : Gurley gave the following short diagnosis from the observations
of J. Miiller: subcircular-ovate or broadly rounded elliptic, length 12/*.

Cepede distinguishes three different types of spores as follows : Lenticu-
lar in side view; subcircular (13.5/x by 13/i), oval (14.7/i by 11.4ju) and
ovoidal (16/i by 11^) in front view. Two polar capsules of equal size (6^
by 4jLi), closely set or separated (3n apart) from each other. Coiled polar
filament distinct. A small triangular intercapsular appendix. Sporoplasm
refractive and finely granular. Sutural edge exhibits folds of variable
number at the posterior portion. Dimensions of fixed and mounted
spores: length 10.5 to 12/x, breadth 7.5 to S/x.

Wegener, without noticing Cepede's paper, also mentions three different
types chiefly distinguished by the spore as follows:

A type (common form), in the branchiae of Lota lota, Abramis brama,
A. vimba, Blicca bjorkna, Leuciscus rutilus, Alburnus alburnus and scardi-
nius erythrophthalmus . Cysts mentioned above.

Spore. Rounded or oval; flattened. A tail, 15/i long, was noticed
twice. A triangular intercapsular appendix. Sutural edge usually having
folds. Polar capsules often differ in form and size in different cysts, tho
they are constant in one and the same cyst, causing the variability in size
of sporoplasm. Dimensions: length 11 to 12.5/i, breadth 8 to 9ju, polar
capsule 4.5 to 6^ by 3 to 3.7/x, in many cysts7.5/x by 4jli.

B type. In the fifth gillarch of Gobio gobio L. Cysts mentioned above.

Spore. Elongated oval. A triangular intercapsular appendix. Indis-
tinct folds on sutural edge. Dimensions: length 12.5 to 13. 5m, breadth 8
to lOju, polar capsule 5 to 6^ by 3 to 4ju.

C type. In the branchiae of Rhodeus amarus Bl. and Alburnus alburnus
L. (April and May). Cysts mentioned above.

Spore. Rounded. Distinct intercapsular appendix. Folds distinct
on sutural edge. Dimensions: length 12 to 15/x, breadth 9 to lOju, polar
capsule 5 to 7/x by 3 to 4/i.

MYXOBOLUS SPHAERALIS Gurley

1874 Clapar^de 1874 : 113-114

1893 Myxobolus sphaeralis Gurley 1893 : 415

1894 Myxobolus sphaeralis Gurley 1894 : 240

142 ILLINOIS BIOLOGICAL MONOGRAPHS [380

Habitat: Mucosa of branchiae of Coregonus lavaretus L. (C. fera);
Lake Geneva.

Vegetative form: Cysts, 0.25 to 0.33mm. in diameter. Polysporous.
Spore: Spherical, 9/z in diameter.

MYXOBOLUS ANURUS Cohn
[Figs. 422 and 423]

1895

Myxobolus anurus

Cohn

1895

: 42-43

1896

Myxobolus anurtts

Cohn

1896

:266

1899

Henneguya psorospermica

anura

Labb£

1899

:102

1910

Myxobolus anurus

Wegener

1910

:76

1911

Henneguya psorospermica

anura

Nemeczek

1911

:146

Habitat: Branchiae of Esox lucius L.; Konigsberg (March, December),
Frisches Haff, Pregel, Masurische See, Lotzen, (September, October).

Vegetative form: Cysts small rounded and of white color. Cohn
measures length 0.6mm., breadth 0.34mm. Wegener's form: length 0.3
to 0.5mm. and breadth 0.2 to 0.3mm.

Spore: Cohn's descriptions are as follows: More or less oval. Dimen-
sions: length 12 to 15/i, breadth 4 to 6.8/*, polar capsule 5.5 to 7/x by 2.1 to
2.5/i, length of polar filament 32 to 38m.

Wegener's form : Elongated and narrow, often with a tail. Dimensions :
length 15/i (maximum up to 18^), breadth 6 to 7/*, polar capsule 8n by 3n.

Remarks: Tho Labbe classified this as a subspecies of Henneguya pso-
rospermica Thelohan, Wegener's observation gives stronger basis for placing
this form in the genus Myxobolus.

MYXOBOLUS sp. Gurley
[Fig. 424]

1882 B&tschli 1882 : 590

1894 Myxobolus sp. incert. Gurley 1894 : 214

1899 Myxobolus sp. Labb6 1899 : 100

Habitat: Nais lacustris L. {N. prohoscidea); Locality?
Vegetative form: Cysts, 8mm. by 4.25mm. Polysporous.
Spore: Oval or circular; tailed or untailed. These spores of diflrerent
form occur, often, without order in the same cyst.

MYXOBOLUS sp. Gurley
[Fig. 425]
1894 Myxobolus sp. incert. Gurley 1894 : 239

Habitat: Body cavity of Carassius carassius L.; Leipsic.
Vegetative form: Not observed.

381] STUDIES ON MYXOSPORIDIA—KUDO 143

Spore: Broadly elliptic; shell bivalve; valves equally convex. Sutural
ridge. Two equal polar capsules. Sporoplasm with a vacuole. Dimen-
sions: length 14/i, breadth lO/i, thickness Six.

Remarks: This species seems to be very similar to M. carassii Kloka-
6ewa (page 150).

MYXOBOLUS sp. Gurley
[Figs. 426 to 429]

1841 MiiUer 1841 : 480

1894 Myxobolus sp. Gurley 1894 : 240-241

1899 Myxobolus sp. Labb6 1899 : 100

Habitat: Skin of opercle, in the branchiae, on the head or on the fin of
Lucioperca lucioperca L.; Germany, Don.

Vegetative form: Cysts 1.09 to 2.18mm. in diameter. Color whitish.
Polysporous.

Spore: Rounded. Thickness equal to haK the breadth. Sutural ridge.
Two polar capsules, of equal size, converging.

MYXOBOLUS CYPRINI Doflein
[Figs. 430 to 432]

1896 Hofer 1896 : 2, 38-39

1898 Myxobolus cyprini Doflein 1898 : 288, 320, 325

1904 Myxobolus cyprini Hofer 1904 : 66-67

1909 Myxobolus cyprini Doflein 1909 : 780-783

1916 Myxobolus cyprini Doflein 1916 : 1026-1027

Habitat: Suppurative connective tissue and epithelium of kidney,
liver and spleen of Cyprinus carpio L., rarely Tinea vulgaris Cuv. and
Abramis brama L.; Germany, Austria. According to Hofer the parasites
cause so-called "small pox of carp" among carp in German waters.

Vegetative form: Small ameboid. Form irregular. The youngest
form with a single or many nuclei, is found in the epithelium of the kidney.
Multiplication by multiple division, the nuclei undergoing amitotic
division. Endoplasm contains homogeneous, yellow and refractive bodies.
Also found in the state of diffuse infiltration. Spores are found in the
parenchym of the kidney.

Spore: Oval. Shell thickened (1.5)li wide) along the sutural edge.
Two converging polar capsules cross each other, in front view, at the
anterior tip. Sporoplasm with an iodinophilous vacuole. Dimensions:
length 21m, breadth ISjj., length of polar capsule 6^i. Doflein (1916:1027)
gives the following dimensions: length 10 to 16/x, breadth 8 to 9/i.

Hofer gives the following dimensions: length 10 to \2n (up to 16/i),
breadth 8 to \\n, polar capsule 5 to 6m by Zn, sutural edge LSju.

144 ILUNOIS BIOLOGICAL MONOGRAPHS [382

MYXOBOLUS NEUROBIUS Schuberg et Schroder
[Figs. 433 to 436]
1905 Myxobolus neurobius Schuberg and Schroder 1905 : 49-56

Habitat: Nervous tissue of TruUa fario L.; Gutach (May?).

Vegetative form: Cysts, usually elongated, often spherical. Elongated
form 0.9mm. by 0.02mm. The seat of the cysts is between the medullary
sheath and sheath of Schwann. Neither medullary sheath nor axis-cylinder
was infected. Cyst-membrane could not be made out. Cysts contained
only full-grown spores without any younger stage. Polysporous.

Spore: Broad oval in front view; spindle shaped in side view. Anterior
end attenuated, posterior end rounded. Shell somewhat thick. Sutural
ridge is not particularly marked. Edge without any fold. No intercapsu-
lar appendix. Sporoplasm, with a large and spherical iodinophilous
vacuole and a single nucleus, occupies less than one half of the inner space
of the spore. Two polar capsules, pyriform, fuse into one at the anterior
end. Coiled (8 to 10 times) polar filament distinct. Dimensions: length
10 to 12/i, breadth 8ju, thickness 6^, polar capsule 6 to Tju by 2jLt.

MYXOBOLUS AEGLEFINI Auerbach
[Figs. 437 to 441]
1906 Myxobolus aeglefini Auerbach 1906 : 568-570

1906 Myxobolus aeglefini Auerbach 1906a : 115-119

1907 Myxobolus esmarkii Johnstone and 1907 : 204-208

Woodcock

1909 Myxobolus aeglefini Auerbach 1909 : 76-78

1910 Myxobolus aeglefini Auerbach 1910c : 181-182

1911 Myxobolus aeglefini Nemeczek 1911 : 162

Habitat: Cartilage and bone of cranium and eye of Gadus aeglefinis
G. callarias, G. merlangus L., G. morrhua L., G. esmarkii and Molva vulgaris
Flem.; Norway, Morecambe (March).

Vegetative form: Cysts in cartilage and bone of cranium and in carti-
lagineous layer of the sclerotic of the eye. Protoplasm is distinctly
differentiated. Ectoplasm somewhat vacuolated; endoplasm granular
with numerous small nuclei. Polysporous.

Johnstone's observations are as follows: Round the peripheral part
of the cornea, and covered loosely by conjunctiva are a number of milk-
white rounded or oval bodies, from about 1 to 3mm. in diameter. Several
of these fused to form elongated mass which lie along the curvature of the
periphery of the eye. These cysts also invade the lateral and posterior
parts of the bulbus ocuU. In sections, the cysts lie within the thickness
of cartilaginous layer of the sclerotic. This latter is enlarged into thick
layer (2mm.) by the presence of the cysts.

Nemeczek mentions irregular cysts of 1.5mm. in diameter.

383] STUDIES ON MYXOSPORJDJA—KUDO 145

Spore: Elliptical in front view. Two polar capsules convergent. No
intercapsular appendix. Sutural edge rather thick with a number of folds
on the posterior margin. Sporoplasm with two nuclei and an iodinophilous
vacuole. Dimensions: length 10.8 to 11.7/i, breadth 9.9 to 10.4ju, thick-
ness 7.2 to 9n, length of polar capsule 4.5 to 5/x.

Woodcock's form has a spore with the following characters:
Slightly ovoid. Sporoplasm always contains a large and well defined
vacuole and two nuclei. Dimensions: length lO/x, breadth 8m, length of
polar capsule 3.25 to 3.5ju.

MYXOBOLUS GIGAS Auerbach
[Figs. 442 to 445]

1906 Myxoholus gigas Auerbach 1906 : 386-391

1910 Myxobolus gigas Auerbach 1910c : 182

1912 Myxoholus gigas Parisi 1912 : 293-294

Habitat: Subcutaneous connective tissue of the operculum of Abramis
hrama L.; Karlsruhe, Pavia. Parisi observed cysts on the side, on the
caudal fin (5 cysts on rays), on other fins, branchiae and in the internal
organs of the fish.

Vegetative form: Cysts, spherical or ovoidal. No cyst membrane
composed of the connective tissue of the host. Protoplasm is indistinctly
differentiated. Ectoplasm thin and radially striated, which gradually
turns into endoplasm. Endoplasm finely granular, contains numerous
nuclei (2.5 to 2.7/i in diameter). Size of greatest form 360ju by 290 to 300/i.
According to Parisi size up to 1.5mm.

Spore : Elliptical when viewed from the front. Sutural edge somewhat
narrow, having a number of folds at the posterior portion. Sporoplasm
with an iodinophilous vacuole and two nuclei. Dimensions: length 16.9 to
21.6m, breadth 13 to 16. 2^ thickness 9ju, length of polar capsules 7.8/i,
length of polar filament 90m (sulphuric acid) .

Parisi gives 150m for the length of polar filament.

MYXOBOLUS VOLGENSIS Reuss

[Figs. 446 to 448]

1906 Myxoholus volgensis Reuss 1906 : 200-201

Habitat: Branchiae, cornea and dorsal fin of Lucioperca volgensis
Pall; Volga.

Vegetative form: Cysts, spherical, 0.3 to 1mm. in diameter. Poly-
sporous.

Spore: Broad elliptic or rounded. Sutural edge has at least 3 folds.
Sporoplasm with an iodinophilous vacuole. Dimensions: length 8.25 to
9.5m, breadth 7.25 to 8.25m, thickness 4.5 to 5.5m, polar capsule Sp, by Ipi.

146 ILLINOIS BIOLOGICAL MONOGRAPHS [384

MYXOBOLUS SCARDINII Reuss
[Fig. 449]
1906 Myxobolus scardinii Reuss 1906 : 201

Habitat: Branchiae of Scardinius erythrophthalmus L. ; Volga.
Vegetative form: Cysts, elongated oval. Smaller cysts rounded oval,
0.8mm. by 0.5mm., the larger forms elongated, 1.2mm. by 0.5mm. Poly-
sporous.

Spore: Broad elliptical. Sutural edge narrow, having folds. A larger
triangular intercapsular process. An iodinophilous vacuole in sporoplasm.
Dimensions: length 11 to i2fi, breadth 9 to 9.5ju, thickness 4.5 to 5^,
polar capsules 5n by 2.5/i.

MYXOBOLUS PHYSOPHILUS Reuss
[Figs. 450 and 451]
1906 Myxobolus physophilus Reuss 1906 : 201-202

Habitat: Surface of air bladder of Scardinius erythrophthalmus L.;
Volga.

Vegetative form: Cysts, rounded, 1.5mm. in diameter. Polysporous.

Spore: Oval, with attenuated anterior end. Sutural edge narrow

and smooth. Polar capsules rather large. An iodinophilous vacuole in

sporoplasm. Dimensions: length 12 to 13/i, breadth 8.25 to 9/i, thickness

6.5 to 7/i, polar capsules 6/i by 2.5/*.

MYXOBOLUS MACROCAPSULARIS Reuss
[Fig. 452]
1906 Myxobolus macrocapsularis Reuss 1906 : 202

Habitat: Branchiae of Blicca bjorkna L.; Volga.

Vegetative form: Cysts, Elongated oval. Size: 1mm. by 0.5mm.
Polysporous.

Spore: Oval with greatly attenuated anterior portion. Sutural edge
broad and without any fold. Polar capsules rather large. An iodinophil-
ous vacuole in sporoplasm. Dimensions: length 11 to 13;*, breadth 8.25 to
9.25/*, thickness 5.5/*, polar capsules 6/* by 2.5 to 3/i.

MYXOBOLUS SANDRAE Reuss

[Fig. 453]

1906 Myxobolus sandrae Reuss 1906 : 202-203

Habitat: Muscle of Luciop&rca sandra Cuv.; Volga.

Vegetative form: Cysts. Rounded, 0.5mm. in diameter. Polyspor-
ous.

Spore: Oval. Sutural edge broad with many distinct folds. An
iodinophilous vacuole in sporoplasm. Dimensions: length 9.25 to 10/*,
breadth 7.25 to 8.25/*, thickness 4 to 5/*, polar capsules 3.5/i by 2/*.

3851 STUDIES ON MYXOSPORIDIA— KUDO 147

MYXOBOLUS BRAMAE Reuss

[Fig. 454]

1906 Myxobolus bramae Reuss 1906 : 203-204

Habitat: Branchiae of Abramis brama L.; Volga.

Vegetative form: Cysts. Oval, O.Smm. long, 0.25mm. broad. Poly-
sporous.

Spore: Oval to nearly spherical. Sutural edge narrow and with indis-
tinct folds. Two polar capsules, with a small triangular intercapsular
process. An iodinophilous vacuole. Dimensions: length 11 to 12/*,
breadth 9.25 to 10/x, thickness 4.5 to 5.5/i, polar capsules 4 to 5/i by 2.25/*.

MYXOBOLUS CYPRINICOLA Reuss
. [Fig. 456]
1906 Myxobolus cyprinicola Reuss 1906 : 204

Habitat: Branchiae of Cyprinus carpio L.; Volga.

Vegetative form: Cysts, oval, 0.5mm. by 0.3mm. Polysporous.

Spore: Elongated oval. Sutural edge narrow with many indistinct
folds. An iodinophilous vacuole. Dimensions: length 9.25 to 10/t, breadth
7 to 7.25/x, thickness 5 to 5.5/*, polar capsules 4.5/* by 2.5 to 3/*.

MYXOBOLUS BALLERI Reuss

[Fig. 455]
1906 Myxobolus balleri Reuss 1906 : 204-205

Habitat: Branchiae of Abramis ballerus L.; Volga.

Vegetative form: Cysts. Elongated, 1.5mm. by 0.5mm. Polysporous.

Spore: Oval, slightly pointed at the anterior end. A triangular
intercapsular appendix. Sutural edge smooth. An iodinophilous vacuole.
Dimensions: length 11 to 12/*, breadth 9.25 to 10/*, thickness 5.5 to 6.5/*,
polar capsules 5.5/t by 2.75/*.

MYXOBOLUS SQUAMAE Keysselitz
[Figs. 457 to 459]
1908 Myxobolus squamae Keysselitz 1908 : 273-274

Habitat: Inner surface of the scales oi Barbus fluviatilis Agass.; Mosel
and Neckar.

Vegetative form: Form variable; rounded, oval, elongated or rarely
branched. The outline of the body is not smooth but irregular with numer-
ous small tooth-like projections with which the body comes in contact
with the surrounding substance. The parasites seem to be able to dissolve
the substance composing the scale. Length 50 to 800/*. In one scale, one
or many, up to 8, individuals were found. All showed only advanced stages
of spore formation. The parasites are surrounded by a variously developed
envelope of connective tissue. Polysporous.

148 ILLINOIS BIOLOGICAL MONOGRAPHS (386

Spore: Elongated oval. Two polar capsules, with 7 to 8 times coiled
polar filament. A triangular intercapsular projection. Sporoplasm with
an iodinophilous vacuole. Dimensions: length 10 to 10.5)li, breadth 8 to
8.5m, length of polar capsule 4.5^.

MYXOBOLUS CORDIS KeysseUtz

[Figs. 460 and 461]

1908 Myxobolus cordis Keysselitz 1908 : 279-282

Habitat: Muscle of ventricle, rarely that of bulbus arteriosus of Barbus
fluviatilis Ag., spores found in kidney, liver and spleen in the condition of
somewhat scattered infiltration; Germany (Mosel and Neckar).

Vegetative form: Elongated, oval, sausage or club form. The body
whitish, later yellowish. Size from 0.25 up to 4mm., usually 1 to 1.5mm. in
length. Propagative stage and cysts observed. One end of the body is
held more or less deeply in the muscle and is covered by cellular envelope
as in Myxobolus tnusculi, while remaining larger portion of the body is sus-
pended freely inside of the ventricle, covered with a thin layer probably of
endocardiac cells. Fish 30 to 45cm. long harboured 40 to 60 parasites.
No movements. Ultimately the cysts are formed with differentiated
protoplasm. Polysporous.

Spore: Oval. Shell very thin at the anterior end. At the posterior
end, cell-like appendage, 2 to 3n wide which is probably formed by both
valves, is present. Two pyriform polar capsules at the anterior end, which
show the polar filament coiled 7 to 8 times. Sporoplasm with a
comparatively large and oval iodinophilous vacuole and two nuclei,
rarely one (syncaryon). Dimensions: length 12/i, breadth 10/x, length of
polar capsule 4.5)u.

MYXOBOLUS MUSCULI Keysselitz

[Figs. 462 to 464]

1908 Myxobolus tnusculi Keysselitz 1908 : 282-286

Habitat: Muscle of the main body, rarely that of fins and operculum,
B.nd kidney oi Barbus Jluviatilis AgSLSS. of various size (youngest fish found
infected, 2 months old), spores in liver, spleen, kidney and ovary (not the
ovum) in diffuse infiltration; Mosel and Neckar.

Vegetative form: Elongated. Body whitish opaque, with differentiated
protoplasm. Smallest individual observed, 24/i. Large form 2mm. in
length. Many trophozoites are found closely situated, forming a large
mass of parasites that reached dimensions of 4mm. by 2mm. The
surrounding envelope, varying in thickness, composed of cells with
elongated nuclei as those of perimysium. Young cysts surrounded by thin
layer of ectoplasm. Polysporous.

3871 STUDIES ON MYXOSPORJDIA—KUDO 149

Spore: Oval. Two polar capsules usually unequal. Shell as in M.
cordis with a small peg closer to the anterior end, polar filament coiled 4 to
5 times, visible in the capsule. Sporoplasm with rarely one (syncaryon),
but usually two nuclei and an iodinophilous vacuole. A posterior process
as is seen in the spores of M. cordis, but much smaller, was occasionally
observed. Dimensions: length 11/i, breadth Sn, polar capsules 6jLt and
4/i long.

MYXOBOLUS sp. Miyairi

1909 Myxobolus sp. Miyairi 1909 : 126
Habitat: Branchiae of loach {Misgurnus anguillicaudatus Cant.?);

Fukuoka? (Nippon).

Vegetative form: Cysts were not observed.
Spore: No description.

MYXOBOLUS sp. Wegener
[Fig. 465]

1910 Myxobolus sp. Wegener 1910 : 78
Habitat: Branchiae (gill-arch) of Percafluviatilis L.; Germany (Frisches

Haff, March). Only one case.

Vegetative form: Cysts on a gill-arch, white and round, with a diameter
of 1.1mm. Polysporous.

Spore: Form and size very variable. Rounded or elliptical, pointed at
the anterior end. Sutural edge showing folds at the posterior portion.
Dimensions: length 8 to 10)u (in round form) and Wjx (in elliptical form),
breadth 8 to 9/i, polar capsules 4 to 5ju by 2 to 3/x, length of polar filament
40m.

MYXOBOLUS PERMAGNUS Wegener
[Fig. 466]

1910 Myxobolus permagnus Wegener 1910 : 78-79
Habitat: Branchiae and operculum of P^rca /wwa^^Vu L., air bladder

of Scardinius erythrophthalmus L.; Konigsberg (May), Pregel (March).

Vegetative form: Cysts rounded in form and white in color, resemble
to those of M. gigas. No clear ectoplasm layer, nor typical protoplasmic
structure. Polysporous.

Spore: Oval, sharply pointed at the anterior end. Sutural edge with
5 to 6 distinct folds at the posterior portion. Polar filament visible in' the
polar capsules. Dimensions: length 17 to 18/x, breadth 10 to 13/:, polar
capsules 7 to 8/x by 3.5 to 4ju.

MYXOBOLUS ROTUNDUS Nemeczek
[Fig. 467]

1911 Myxobolus rotundus Nemeczek 1911:156-157
Habitat: Branchiae of Abramis brama L.; Austria.

ISO ILLINOIS BIOLOGICAL MONOGRAPHS [388

Vegetative form: Cysts, ovoidal or spindle form, 1 to 3mm. long and
1 to 1.5mm. wide. Body white. An extraordinary large number of spores
were found in the cysts. Polysporous.

Spore: Round or slightly oval, when viewed from the front. Greatly
flattened in side view. Polar capsules convergent, with no intercapsular
body. Shell smooth. Sutural edge narrow, without folds. Dimensions:
length lO/i, breadth 9,8/i, thickness 3/x, polar capsules 3.8 to 5/x long, length
of polar filament 40jtt.

MYXOBOLUS MINUTUS Nemeczek
[Fig. 468]

1911 Myxobolus mintUus Nemeczek 1911 : 160
Habitat: Branchiae of Leuciscus sp.; Austria.

Vegetative form: Cysts spherical, oval or elongated with white color.
Size: 0.5 to 3mm. by 0.5 to 1mm. Polysporous.

Spore: Rounded oval, similar to that of Myxobolus rotundus. Shell
smooth. Sutural edge narrow without folds. Sporoplasm with an iodino-
philous vacuole. No intercapsular appendix. Dimensions: length 6m,
breadth 4.2 to 5ju, polar capsule ^n by 2/i, length of polar filament 50 to 60,
often 70/Lt.

MYXOBOLUS sp. Lebzelter

1912 Myxobolus sp. Lebzelter 1912 : 296-297
Habitat: Gall-bladder of Thymallus thymallus L.
Vegetative form: Not observed.

Spore: Sutural ridge distinct. Dimensions, length 5^, breadth 3^.

MYXOBOLUS MAGNUS Awerinzew
[Figs. 469 and 470]

1913 Myxobolus magnus Awerinzew 1913 : 75-76
Habitat: Eye of Acerina cernua L.; Petrograd.

Vegetative form: Trophozoites form white spots in the tissue of iris,
with many spores (300 to 400). Each pansporoblast forms in most cases
two, sometimes 3 or 5 spores! Polysporous.

Spores: Large, elongated roundish, slightly flattened. Sutural edge
somewhat thick, forming a wide ridge, with 4 to 5 folds at the posterior
portion. Polar capsules do not cross each other. Sporoplasm with an
iodinophilous vacuole and two nuclei. Dimensions: length 38 to 45m,
breadth 32 to 38m, thickness 28 to 35m, length of polar capsules 15 to 17m,
diameter of the vacuole 12 to 16m.

MYXOBOLUS CARASSII Klokacewa
[Figs. 471 to 473]

1914 Myxobolus carassii Klokacewa 1914 : 182-184
Habitat: Body cavity, liver and intestine of Carassius vulgaris L.;

Petrograd?

389] STUDIES ON MYXOSPORJDJA—KUDO 151

Vegetative form: Cysts spherical. Those in liver and intestine yellowish,
surrounded by an envelope composed of fibrous connective tissue. Second-
ary cysts are formed. Polysporous.

Spore: Oval, in front view. Two ovoidal polar capsules convergent
at the slightly attenuated anterior end. Coiled polar filament visible.
Sporoplasm with an iodinophilous vacuole and two nuclei. Sutural edge
shows folds in some cases. Dimensions: length 13 to 17ju, breadth 8 to
10/i, thickness 5 to In, polar capsules 6 to Ifx long.

Remarks: Compare with Myxobolus sp. Gurley on page 142.

MYXOBOLUS sp. Southwell
1915 Myxobolus sp. Southwell 1915 : 312-313

Habitat: Subcutaneous intermuscular tissue of Rasbora (Cyprinus)
daniconius Day; from a stream near Katwan, Mirzapore (U.P.), India.

Vegetative form: 6 cysts found on four fish. The seat is immediately
below the scales, in the epidermis. Color milky white. Soft, flattened and
roughly oval in shape. Greatest length found, 1.1mm, No pigment was
present on the cyst.

Spore : Two equal capsules, with a very short tail-like process. Sporo-
plasm with vacuole; iodine treatment could not be carried out.
Dimensions: length 13^, breadth I3fi, polar capsule 4)u by 4/i(?).

Remarks: Dimensions, especially that of polar capsule seem to
be misprinted. Southwell gave one figure of a fish with a cyst near the
dorsal fin. He thinks that "it is quite possible that our parasites may
belong to Myxobolus cyprini." The incomplete observation without any
figure, leads the writer to leave the form also as Myxobolus sp. Southwell.

MYXOBOLUS FUNDULI Kudo

[Figs. 474 to 476]

1915 Myxobolus musculi Hahn 1915 : 201-205

1917 Myxobolus musculi Hahn 1917 : 91-104

Habitat: Branchiae and muscle of Fundulus heteroclitus, F. majalis;
Woods Hole. Hahn claims that he succeeded in causing experimental
infection in F. diaphanus and Cyprinodon variegatus by inoculation.

Vegetative form: Hahn uses quite a number of different terms from
those that are ordinarily used in describing Myxosporidia, without giving
any definitions. Naturally it is hard to put what he wrote in several pages
in the following lines. Granular vegetative forms produce a great many
pansporoblasts, each with a single spore. "Trophoplasm" is difficult to
stain. Size: 74 by 33n, 24 by I9fi. Cysts within and between the muscle
fibers, containing several hundred spores.

Spore: Hahn's descriptions may be summarized as follows: Dimen-
sions: length 14.3^, breadth 6.7/x, thickness 6.7/t to 2/3 of width, polar

152 ILLINOIS BIOLOGICAL MONOGRAPHS [390

capsule, 6.5/x by 2n, polar filament 3 to 4 times the length of the spore
(42.9 to 57.2)Li). Polar filament coiled 10 to 14 times. Shell thin, almost
invisible. The spores found in the gill: length 12 to 13.4ju, breadth 6/i to
10.4ai. a vacuole is present in the sporoplasm.

Remarks: Examination of Hahn's first paper suggested that he was
dealing with the present form as a new species tho he did not mention
at all Keysselitz who gave the name Myxoholus musculi Keysselitz to
the parasite of Barbus fiuviatilis from German riverg. I was informed by
Hahn that he gave the name, Myxoholus musculi, without knowing the
fact that it was preoccupied by Keysselitz (1908) (see page 148) and that
tho he became aware of it later, he can not determine differences by which
the two forms can be distinguished. A comparison of the descriptions
of Keysselitz and Hahn, however, shows that these two forms differ in
several respects. Hence the latter form is recorded here as a distinct
species under the new name.

It is interesting to note that very similar forms, one without an iodino-
philous vacuole at any stage of spore-formation (Myxosoma funduli Kudo,
see page 125) and the other with a vacuole, occur in the same hosts
in the same locality. As mentioned above, the reader is requested to refer
to Hahn's original paper for further data.

. MYXOBOLUS PLEURONECTIDAE Hahn

[Fig. 477]

1917 Myxoholus pleuronectidae Hahn 1917 : 160-161

Habitat: Subcutaneous muscular tissue of Pseudopleuronectes ameri-
canus; Woods Hole.

Vegetative form : Similar to that of Myxoholus f unduli.

Spore: Hahn writes as follows: Shape and appearance resembles
Myxoholus pfeifferi. Dimensions: length 14.5/x, breadth 11.9ju, polar
capsules 6n by 3.7 fx.

MYXOBOLUS CAPSULATUS Davis

[Fig. 478]
1917 Myxoholus capsulatus Davis 1917 : 237

Habitat: Visceral connective tissues of Cyprinodon variegatus; Beaufort.

Vegetative form: Irregular form. In the state of diffuse infiltration.
Polysporous.

Spore: Pyriform, flattened. Polar capsules large and pyriform, filling
almost entire cavity of the spore. Sporoplasm relatively small. lodino-
philous vacuole visible in living spore. Dimensions: length 16)li, breadth
10 to 1 Iju, polar capsules 1 1/x by 4/i, length of polar filament 84/i.

3911 STUDIES ON MYXOSPORJDJA—KUDO 153

MYXOBOLUS NODULARIS Southwell et Prashad

[Figs. 479 and 480]

1918 Myxobolus nodularis Southwell and Prashad 1918 : 347

Habitat: In the muscles of Rashora daniconius occurring in two fish
on the sides, and in another as a globular cyst near the anus; Mirpur,
Dacca (June). Type specimens, numbered P 52/1,

Vegetative form: Cysts rounded or slightly elongated, varying in length
3.5 to 3.8mm. and 2.3 to 2.8mm. in width. Creamy yellow in color, in one
case appearing blackish owing to the large number of black granules
scattered in its surface.

Spore: Ovoidal. Sutural ridge very wide (about 1/5 thickness of the
spore). Two polar capsules of equal size, which show coiled polar filaments
clearly. Dimensions: length 9/i, breadth 7.2/i, length of polar capsule
3.4;ti, that of polar filament 18.3/i.

MYXOBOLUS HYLAE Johnston et Bancroft
[Figs. 591 to 593]

1888

Fletcher

1888

:337

1890

Haswell

1890

:661

1909

Myxobolus sp.

Johnston

1909

:29

1910

Myxobolus sp.

Cleland and

Johnston

1910

:25

1918

Myxobolus hylae

Johnston and

Bancroft

1918 ;

: 171-175

Habitat: In the testes, vasa efferentia and oviducts of Hyla aurea;
Sidney, Australia (April, other months not mentioned). Fletcher observed
the parasites also in the urinary bladder of both sexes, which fact was not
confirmed by Johnston and Bancroft on account of the scarcity of the
material. The latter authors could not infect Hyla caerulea by feeding
infected testes or the cysts, giving the conclusion that the parasite is
specific to H. aurea. The male is more often attacked by the parasite
than the female. The infected animal appeared sickly and emaciated.
As to the infection in kidneys, they write as follows: In one male specimen
both testes and both kidneys were affected, and the upper parts of the
ureters adjacent to the kidneys were swollen and milky in appearance.
In another, in addition to the testes, the adjacent kidney and mesentery
were attacked. No spores have yet been detected by them in sections of
the kidney tubules.

Vegetative form: Johnston and Bancroft describe as follows:
Cysts: in male, either imbedded in the tissue or may project freely into
the coelom of the testes; in female, lying between the layers of the wall,
being projected into the lumen of the oviduct. Size from those of micro-
scopic dimensions up to 2 to 3mm. in diameter. In sections, the protoplasm

154 ILLINOIS BIOLOGICAL MONOGRAPHS [392

is differentiated into two regions. The outer layer (ectoplasm) surrounds
the body as a thin, light-staining region, while the endoplasm being den-
ser and of more or less granular structure filled with spores especially in
the central portion.

Spore: Johnston and Bancroft describe as follows:

Form somewhat variable, caused by the reduction in length. Oval,
egg-shaped or nearly circular in front view. Sutural ridge slightly thick-
ened. Two pyriform polar capsules are located at the anterior end.
Sporoplasm with an iodinophilous vacuole (2ju in diameter), shows usually
two distinct nuclei, rarely but one. Dimensions: length variable, diameter
of circular form 7 to 8/x, breadth 8 to lOju, thickness about 6/x, thickness of
shell 1/x, polar capsules 4 to 5/i by 2/i, length of polar filament 90 to 98ju
(acids or alkalies). ,

MYXOBOLUS AUREATUS Ward
[Figs. 643 to 649]
1919 Myxoholus aureatus Ward 1919 : 49

Habitat: Between the ectodermal layers of the fin membrane of
Notropis anogenus', Put-in-Bay, Lake Erie (August). Out of thirty fish,
two to three cm. in length, seven were found to be infected. The infected
fish were not inferior in size or vigor to others of the same species. The
most heavily infected one was the most vigorous of all. The number of
cysts, in the individual fish, varied from one to forty, being confined in the
fin. The cysts are always separated from each other, tho in a few instances
they were apparently connected.

Vegetative form: The parasite forms cysts between the ectodermal
layers of the fin membrane. The cyst is a smooth margined ellipsoid,
measuring from 1 to 1.6 mm. in layer diameter and from 0.8 to 1.2 mm.
along its transverse axis. The opaque cyst is of a clear orange yellowish
color. This gilt color is contained in the cyst wall, fading away in alcohol
and formol. The chromatophores of the skin of the host are distinctly
more abundant on the cyst than in other parts of the skin, and the older
the cyst the more abundant the chromatophores. The wall of the cyst is
noticeably tough and thick. In section, the protoplasm shows a poor
differentiation into ectoplasm and endoplasm. The former granular and
reticular, covers the entire surface as a thin layer, while the latter is highly
vacuolated, containing only mature spores. Polysporous.

Spore: Ovoid; slightly pointed anterior and rounded posterior ends in
front view; slightly compressed in lateral view. Sutural ridge distinct.
The shell is of moderate thickness, and bears a flange at the posterior
half in some spores. Two pyriform polar capsules, frequently of slightly
different dimensions, are at the anterior part of the spore. No intercapsu-
lar appendix is present. When the spore is allowed to stand for 24 hours

393] STUDIES ON MYXOSPORIDIA—KUDO ISS

or more in water, the polar filaments are extruded. The binucleated
finely granular sporoplasm shows an iodinophilous vacuole. Dimensions:
length 12.4 to 13.5/i, breadth 6.5 to 7.5/*, thickness 5/i, length of polar
capsule 6 to 7/i (rarely 7.5/x), length of polar filament about 20 to 26/i,
diameter of iodinophilous vacuole about 2n.

MYXOBOLUS MIYAIRII nov. spec.
[Fig. 481]
1909 Myxoholus sp. Miyairi 1909 : 130, 131-132

Habitat: Intestinal wall of Parasilurus asotus L.; Fukuoka ? (Nippon)

Vegetative form: Cysts. Size rather small up to 0.5mm. Full-grown
spores as well as those in developmental stages fill the central portion of
cysts, while numerous nuclei are chiefly found along the periphery of end-
plasm.

Spore: Elongated elliptic. Two polar capsules of nearly same size.
Sporoplasm with a comparatively large iodinophilous vacuole. Dimen-
sions: length 13 to 14.5)u, breadth 6 to 7/x, length of polar capsules 4.5iLt,
length of polar filament 30 to 35/i.

Remarks. As the descriptions show the form and structure are dis-
tinguishable from other species, the writer establishes the present species.

MYXOBOLUS KOI nov. spec.
[Figs. 482 to 485]

Habitat: In the connective tissue of the gill filament of CypHnus carpio
L.; Tokio (April). One fish was found infected in a slight degree.

Vegetative form: Cysts small and spherical; white in color. Size up to
230m in largest diameter. The seat similar to Myxoholus toyamai. The
structure of the cysts, observed in section preparations, is also similar to
the above mentioned unicapsular Myxoholus.

Spore: Oval with attenuated anterior and rounded posterior ends in
front view; elongated pyriform in side view. Shell comparatively thin. No
marking on shell. Sutural ridge fairly well marked. No intercapsular
appendix. Two polar capsules are pyriform, large, and of usually equal
form and size. Coiled polar filament distinct in vivo. Sporoplasm rather
small, finely granular, shows two nuclei in almost all spores. An iodino-
philous vacuole is deeply stained by Lugol's solution. Dimensions: 14 to
16m, breadth 8 to 9fx, thickness 5 to 6m, polar capsule 8 to 9m by 2.5 to 3m,
length of polar filament 72m in average (KOH).

MYXOBOLUS ORBICULATUS nov. spec.
[Figs. 566 to 576]

Habitat: Muscle of myotomes of Notropis gilherti J. et M.; Stony Creek,
111. (November). The fish was kept alive in an aquarium from November

156 ILLINOIS BIOLOGICAL MONOGRAPHS [394

11, 1918, until March 10, 1919, when it was killed, being then nearly
dead. The material was examined on March 15. A few isolated spores
occurred in the muscle of Notropis blennius (Homer Park, 111., November).

Vegetative form: In and between the muscle bundles of the myotomes.
Size variable. Color opaque white under the dissecting microscope.
Smallest rounded ameboid forms with a single or numerous nuclei, in the
muscle bundle, have the size of from 10/t to 30n in greatest diameter
(Figs. 573 to 575). The largest form observed was 400/x by 120ju. Young
forms without any diflferentiated protoplasm, shows indistinct granular
and reticular structure with deeply staining spherical or ring-form chro-
matinic granules. The number of the nuclei increases with the growth of
the body. Larger form (Fig. 576), spindle shape, circular in cross-section,
lies with its long axis parallel to the muscle fibres. The protoplasm
vacuolated, contained mostly mature spores. Spores were also found in
the state of diffuse infiltration. Polysporous.

Spore: Form somewhat variable. Typical form almost circular,
slightly pointed at the anterior end (Fig. 566) in front view; spindle shaped
in profile (Figs. 569 and 570). Sutural ridge marked. Shell uniformly
thick, usually exhibiting four triangular folds on the surface along the
posterior margin (Figs. 566, 568 and 571). No intercapsular appendix.
Two pyriform polar capsules are, as a rule, of the same size and form.
Frequent occurrence of the inequality of the polar capsules together with
abnormalities in the form of the spore, were noticed especially among
comparatively young spores. The granular sporoplasm, shows two spher-
ical nuclei when stained. The iodinophilous vacuole, spherical and 2fi in
average diameter, is deeply stained with Lugol's solution. Dimensions
of unstained preserved spores: length and breadth 9 to lO/x, thickness
6.5 to 7/x, polar capsule 6 to 7.5/x by 2.5 to 3n.

MYXOBOLUS DISCREPANS nov. spec.
[Figs. 597 to 601]

Habitat: Branchial lamellae of Carpiodes diformis; Salt Fork, Urbana,
U.S.A. (May). One fish caught, died (soon after the capture) two hours
before being fixed. Length 8.5cm.

Vegetative form: The parasites formed numerous cysts on the branchial
lamellae. Cysts slightly yellowish white and mostly rounded or elongated
along the lamella, occur in groups, often occupying the entire lamella.
Infection was fairly heavy. Every gill arch harbored ten to twenty cysts
mostly on the outer surface. Size of the cyst varies, small rounded one
500/x in diameter up to elongated forms 2mm. by 0.5mm., the majority
being from 0.5 to 1mm. in diameter. The cyst is surrounded by a thin
connective tissue layer of the host. The protoplasm shows little differen-

395) STUDIES ON MYXOSPORIDIA—KUDO 157

tiation. The ectoplasm is a rather narrow zone around the entire body
and the endoplasm is filled with various nuclei, several stages of developing
pansporoblasts, and mature spores. Each pansporoblast produces two
spores. Polysporous.

Spore: Approximately circular with broad anterior and more or less
narrower posterior end in front view; broadly fusiform in profile. Shell
uniformly thin with 5 to 6 markings on the posterior margin. Two polar
capsules broadly oval and convergent, fill the anterior half of the spore.
A small triangular intercapsular appendix presents. Coiled polar filament
is fairly visible in vivo. The spores from the cysts which were fixed with
alcohol-acetic and preserved in 95 per cent alcohol, showed the extrusion
of the polar filament under the influence of potassium hydrate solution
(35 per cent) even after a considerable length of time as is shown in the
following:

Material fixed on May 29. .

June 2; Extrusion took place in almost all spores.

June 10; Extrusion took place in almost all spores.

June 26; Extrusion took place in almost all spores.

July 28; Extrusion took place in almost all spores.

August 29; Extrusion took place in numerous spores.

September 29; Extrusion took place in about 70 per cent of the spores, some filaments
being rather short, and not fully extended.

October 20; Extrusion took place in about 50 per cent of the spores, most filaments being
short, and not fully extended.

Sporoplasm coarsely granular shows clearly two ring-form nuclei in
fresh preparations. Dimensions of preserved spores: length 11.4 to
13.5/1, breadth 9.5 to llju, thickness 8.5 to 9.5ju, polar capsule 5.5 to 6n
by 3.5 to 4/x, length of polar filament 50 to 55/i.

Remarks: The present species differs from the hitherto known species,
Myxobolu» lintoni (page 138) and Myxobolus orbiculatus (page 155) which
are the nearest to the present form, differ from Myxobolus discrepans in
the host, organ of infection, vegetative form and form and structure of
the spore.

MYXOBOLUS MESENTERICUS nov. spec. ,
[Figs. 628 to 631]

Habitat: In the mesentery, liver, spleen and wall of stomach, pyloric
coecum, intestine, and gall-bladder of Lepotnis cyanellus; Crystal Lake,
Urbana, 111. (June and July). Out of thirty-six host fish, 10 cm. in average
length, seven were found to be infected. In every case, except one, the
mesentery was the main seat of infection, harboring conspicuous cysts.
The number of cysts found in the host body varied from three to seven.
The infected fish did not exhibit any recognizable pathological changes.
Other species of fish caught at the same time, were free from the infection.

158 ILLINOIS BIOLOGICAL MONOGRAPHS [396

Vegetative form: The cysts are mostly spherical in form, and are
covered by a tough resistant envelope composed of the connective tissue
of the host. They are uniformly white in color, and have the variable
dimensions of from 0.5 to 1.5mm. in diameter. In section, the protoplasm
shows a coarsely reticulated structure without distinct differentiation.
In all cysts of various sizes fully mature spores were only observed. The
spore formation could not be worked out. Polysporous.

Spore: Broadly oval with a slightly truncated anterior end in front
view (Fig. 628), lenticular in side or end view (Fig. 629). No intercapsular
appendix is seen. The shell is rather thick, and shows about eight folds
on the sutural edge, two of which located laterally being more conspicuous
than others. The sutural ridge is rather fine. Two convergent polar
capsules equal in size occupy the anterior half of the spore. The coiled
polar filament becomes more distinctly visible with the addition of Lugol's
solution, altho it is faintly observable in fresh state. Fresh spores ex-
truded their polar filaments under the action of potassium hydrate solu-
tion. In some spores, the extruded filaments cross each other near the
foramina. The preserved spores showed no extrusion of the filament
as in the last species. The sporoplasm is extremely finely granu-
lated. The iodinophilous vacuole is comparatively large. When stained,
the spore shows two nuclei in the sporoplasm. Dimensions of fresh
material: length 10 to 11.5jli, breadth 8.5 to 9.5m, thickness 6.5ai, polar
capsule 4.75m by 1.5 to 2n, length of polar filament 32 to 40^. Average
dimensions of unstained preserved spores: length 9.5/*, breadth 8m, polar
capsule 4.75 m by 2m.

Remarks : The habitat and the structure of the spores, lead the writer
to record the species as a new species.

Genus HENNEGUYA Thelohan
1892 Henneguya Thelohan 1892 : 167, 176

1895 Henneguya Th61ohan 1895 : 352

The characters of the genus are described on page 59.
Type species : Henneguya psorospermica Thelohan.

HENNEGUYA PSOROSPERMICA Thelohan
[Figs. 486, 487 and 496]

1895

Henneguya psorospermica

Thdlohan

1895

:ZS3

1896

Myxobolus psorospermica s.

str. Cohn

1896

261

1899

Henneguya psorospermica

typica

Labb6

1899

101

1905

Henneguya psorospermica

Nufer

1905

: 77, 185

1910

Henneguya psorospermica

Wegener

1910

: 81-82

1911

Henneguya psorospermica

typica

Auerbach

1911

: 5, etc.

Habitat: Branchiae of Esox lucius L. and Ferca fluviatilis; France,

397] STUDIES ON MYXOSPORIDIA—KUDO 159

Frisches and Kurisches Hafif, Pregel, Masurische Seen (all the year round,
but rarer in Winter) Switzerland.

Vegetative form: Thelohan's observations on the structure of the
cyst, are as follows: The surface of the cyst is covered by a la5''er, homo-
geneous, refringent and deeply stained, with which the cyst comes in
direct contact with the surrounding epithelial cells of the host. Inside
of this layer, there is a "pseudoectoplasmic" zone, in which the protoplasm
is dense at places, forming radiate irregular striations, enclosing numerous
irregular masses which are composed of apparently the same substance that
forms the external layer. Toward the central portion of the cyst, there are
masses of spores (Fig. 496).

Cohn's descriptions are as follows: The purely white cyst is elliptical;
length 1.15mm. and breadth 0.85mm. The seat is under the epidermis.
It is surrounded by the host tissue with small, elongated and scattered
nuclei. The outer layer of the cyst is a thin membraneous protoplasm.

Wegener writes as follows: The white cysts are round or elliptical,
usually on the upper end of the branchial lamella. Size of larger cysts,
1.5 to 2mm. long and 1.1 to 1.5mm. wide.

Spore: Elongated; anterior part fusiform and anterior end blunt.
Polar capsules elongated and parallel to each other. Coiled polar filament
visible in fresh conditions. Shell unstriated. Dimensions: total length
40)u in average, largest breadth 7/x, length of polar capsule 7 to 8/x.

Cohn's form is described by him as follows: Spore narrow with blunt
anterior end. Sporoplasm with 6 horns (no figure to explain this expres-
sion!). When kept in water, sporoplasm takes round form and becomes
highly refractive. Dimensions: length 29 to 38/x, length between the tip
and the posterior margin of the cavity (15 to 20/x) 18;u, breadth 9 to 10/x,
polar capsule (8 to ll)u) 9/i by 2tx, length of "starren Faden" 14/n, length of
tail 14 to 18m.

Wegener's form is as follows: total length 35 to 38/i, breadth 7 to 8/*,
length of the spore cavity 15^, length of tail 15 to 20/x, polar capsule 8)li by
2 to 3)u.

HENNEGUYA TEXTA (Cohn) Labbe

1895 Myxoholiis textus Cohn 1895 : 38-39

1899 Henneguya psorospermicatexta Labb6 1899 : 101

1910 Henneguya texta Wegener 1910 : 82-83

Habitat: Branchiae of Perca fluviatilis L.; Pregel, Frisches and
Kurisches Haff (all the year round).

Vegetative form : Cohn observed as follows : Cyst distinctly elliptical.
Length 0.75mm., breadth 0.375mm. The cysts surrounded by a thick layer
of the host tissue. In the peripheral portion of the cyst, the protoplasm
exhibits a network-like structure which forms a fibrous structure further
inside, crossing the cyst at right angles to the long axis of the cyst.

160 ILLINOIS BIOLOGICAL MONOGRAPHS (398

Wegener writes as follows: The white cysts are elongated, 1.2 to 1.8mm.
long and 0.5 to 0.7mm. wide.

Spore: Cohn mentions dimensions exactly the same as those of
Henneguya psorospertnica and can not distinguish the two species by the
spore.

Wegener gives the following dimensions: length 30 to 40/i, breadth
7 to 8m, length of the cavity of spore 15 to 18/i, length of tail 15 to 25^,
polar capsule 8/t by 2 to 3jn.

HENNEGUYA MINUTA (Cohn) Labbe

[Figs. 488 and 489]

1895 Myxobolus minutus Cohn 1895 : 39-40

1899 Henneguya psorospertnica

minuia Labb€ 1899 : 102

Habitat: Branchiae of Percafluviatilis L.; Frisches Haflf, Lesina.

Vegetative form: Cohn's description is as follows: Cysts oval and
small, difficult to distinguish them from those of Henneguya psorospermica.
Size, 130/1 by 115/x. The parasite was met only once. But the number of
the cysts was far greater than that of Henneguya psorospertnica, often 5 to 6
on one lamella, reaching up to 200 cysts on a single gillarch.

Spore: Cohn gives the following dimensions: total length (28 to 45)u)
about 36/i, length from the tip to the end of cavity (20 to 28m) about 26m,
breadth 10 to 11m, thickness 8m, polar capsule 11 to 14m by 2 to Zn, length
of polar filament 42 to 45m, length of tail (8 to 17m) 12m. Cohn gives a
figure (Fig. 489) of a spore with two vacuoles(?).

HENNEGUYA OVIPERDA (Cohn) Labbe
[Figs. 490 and 491]
1892 Weltner 1892 : 28-36

1895 Myxobolus oviperdus Cohn 1895 : 40-41

1899 Henneguya psorospertnica ovi-

perda Labbfi 1899 : 102

1904 Hentieguya ^psorospertnica ovi-

perda Fuhnnann 1904:469-471

19 1 1 Henneguya psorospertnica ovi-

perda Auerbach 1911 : 5-22

1911 Henneguya psorospertnica ovi-

perda Nemeczek 1911 : 146

Habitat: Ovary of Esox lucius L.; Switzerland, Berlin, Frisches Haff
(all the year round), Upsala (May), Austria (December).

Vegetative form: Cohn writes as follows: No real cyst exists. The
parasite occupies the ovum.

Auerbach, however, mentions the presence of cysts in the connective
tissue and follicle epithelium of the ovary. Dimensions, 1mm. up to 5 or
6mm. in diameter.

399\ STUDIES ON MYXOSPORJDIA—KUDO 161

Spore: Cohn states the form and dimensions are very much similar
to those of H. psorospermica.

HENNEGUYA LOBOSA (Cohn) Labbe
[Figs.' 492 and 493]

1895 Myxobolus lobosus Cohn 1895 : 42
1899 Henneguya psorospermica lo-

bosa Labb6 1899 : 102

1910 Henneguyai?) lobosa Wegener 1910 : 83

1911 Henneguyai?) lobosa Auerbach 1911 : 22-25

Habitat: Branchiae of Esox lucius L.; Frisches Hafif, Pregel, Karlsruhe.

Vegetative form : Cysts irregular in shape, size up to 2.5mm.

Wegener noticed that the cyst resembles that of Myxosoma dujardini
with the dimensions of 2.2 to 2.8mm. by 1 to 1.1mm.

Spore: Cohn gives the dimensions as follows: total length 30 to 40/i,
length from the tip to the posterior margin of cavity 11.5 to 15/i, breadth
5 to 6.5m, polar capsules 6.5 to Sfi by 2 to 2.5/x, length of tail 22 to ISfi.

Wegener's form: oval; length 35 to 40^1, breadth Sfi, polar capsule 6 to
7n by 2.5 to 3n, length of the cavity of spore 13 to 15^, length of tail 20 to
25/i, the iodinophilous vacuole could not be detected.

Auerbach gave the following dimensions: total length 30/*, breadth
4 to 6n, length of polar capsule 6n, length of polar filament 48 to 54/u.

Remarks: Wegener and Auerbach did not observe the iodinophilous
vacuole.

HENNEGUYA PERI-INTESTINALIS Cepede

1906 Henneguya psorospermica peri-

intestinalis Cfipfide 1906 : 67

1907 Henneguya psorospermica peri-

intesiinalis C^de 1907 : 137

1912 Henneguya psorospermica peri-

intestinalis Parisi 1912 : 295

Habitat: Intestine of Esox lucius L.; Lac du Bourget, Pavia. (June).
Vegetative form: Cysts.

Spore: Cepede mentions that it resembles that of Henneguya psoro-
spermica.

HENNEGUYA MEDIA Thelohan
[Figs. 494 and 495]
1890 Thelohan 1890 : 198-200

1892 Henneguya media Thdohan 1892 : 177

1894 Myxobolus medius Gurley 1894 : 248

1895 Henneguya media Thglohan 1895:353
1898 Henneguya media Doflein 1898 : 342

Habitat: Renal tubules of kidney and ovary of Gasterosieus aculeatus
and G. pungitius L.; France. Mixed infection with Sphaerospora elegans.

162 ILLINOIS BIOLOGICAL MONOGRAPHS [400

Vegetative form: Rounded or elongated. In larger individuals, clear
differentiation of protoplasm. Monosporous (?) and polysporous.

Spore: Fusiform. Shell striated. A vacuole in sporoplasm. Dimen-
sions: length 20 to 24ju, breadth 5 to 6/t, polar capsules 4 to 5/x. Tail short.

HENNEGUYA BREVIS Thelohan
1854 Lieberkuhn 1854 : 357

1892 Henneguya brevis Th61ohan 1892 : 177
1895 Henneguya brevis TWlohan 1895:354

Habitat: Similar to H. media Thelohan.
Vegetative form: Undescribed.

Spore: Fusiform with short tail. Dimensions: length 14 to ISju,
breadth 5 to 6/1, polar capsules 1.4 to 5jU, tail 4 to 5)u long.

HENNEGUYA SCHIZURA (Gurley) Labb6

[Figs. 497 to 499]

1841 MuUer 1841 : 477-478

1893 Myxobolus schizurus Gurley 1893 : 417

1894 Myxobolus schizurus Gurley 1894:255
1899 Henneguya schizura Labb6 1899 : 102-103

Habitat: In cellular tissue of the eye muscles, in that of the sclerotic,
and in that between the sclerotic and choroid of Esox lucius L.; Germany,
U. S. A.

Vegetative form: Cysts white; membrane delicate; 0.44 to 1.09mm. in
diameter.

Spore: Oval. Dimensions: length 12/x, breadth 6/i, thickness one-
half the breadth, tail 3 to 4 times length of the body.

HENNEGUYA CREPLINI (Gurley) Labbe

[Figs.

500 to 503]

1842

Creplin

1842

: 61-63

1894

Myxobolus creplini

Gurley

1894

: 248-249

1899

Henneguya creplini

Labb€

1899

:103

1910

Henneguya creplini

Wegener

1910;

:84

Habitat: Branchiae of Acerina cernua L.; Pregel (March), Frisches
and Kurisches Haff.

Vegetative form: Wegener describes as follows: Cysts, usually elon-
gated oval and are located at the end of branchial lamella. Color white.
Size 1 to 1.1mm. by 0.5mm. During winter, the cyst has only pansporo-
blasts, but no fully grown spores.

Spore: Creplin writes as follows: Elongated elliptical. Length 1/120'",
breadth 1/360'", tail about as long as or a little longer than the body.

Wegener's form: elongated spindle shape; length 20At, breadth 8 to
9iJL, polar capsule 8/i by 2 to 3/x (parallel to each other).

4011 STUDIES ON MYXOSPORJDJA—KUDO 163

Remarks: Wegener thinks that the present species and Henneguya
acerinae Schroder, are one and the same species, and that the differences
between the dimensions are due to the miscalculation of measurement in
lines given by Creplin on the part of Gurley and Labbe.

HENNEGUYA LINEARIS (Gurley) Labbe
[Fig. 504]

1841 MuUer 1841 : 489

1893 Myxobolus linearis (part) Gurley 1893 : 417

1894 Myxobolus linearis Gurley 1894 : 255
1899 Henneguya linearis Labb6 1899 : 103

Habitat: Membrane lining branchial cavity of Pimelodus sebae Cuv.
et Val., branchiae of Platystoma fasciatum L.; South American rivers.
Vegetative form: Not described.
Spore: Very narrow. Length 3 to 4 times breadth.

HENNEGUYA GURLEYI Kudo

[Fig. 505]

1893 Myxobolus linearis ipaxt) Gurley 1893:417

1894 Myxobolus ci. linearis Gurley 1894:253-254
1899 Henneguya linearis v&T. Labb6 1899 : 103

Habitat: Base of spines of the second dorsal fin of Ameiurus melas
Raf.; Iowa (Storm Lake) (August).

Vegetative form: Spherical cysts, 1mm. in diameter.

Spore: Lanceolate. Dimensions: length of the body 19/i, width
5 to 6/i, thickness about 3)it.

Remarks: The species is most probably different from Henneguya
linearis judging from the difference in the form and structure of spores,
the seat of infection, and host species. Hence, it is recorded here as an
independent species.

HENNEGUYA STRONGYLURA (Gurley) Labb6
[Fig. 506]
1841 MuUer 1841 : 480

1894 Myxobolus strongylurus Gurley 1894:249

1899 Henneguya strongylura Labb6 1899:103

Habitat: Skin of cephalic region of Synodontis schall Bl. Schn.; Nile.
Vegetative form: Cysts over 2.18mm. in diameter.
Spore: Dimensions: length of the body 9/i, breadth 5.4/1. Tail always
undivided. Two polar capsules of equal size.

IW ILUNOJS BIOLOGICAL MONOGRAPHS (182

HENNEGUYA MONURA (Gurley) Labbe
[Fig. 507]

1880 Ryder 1880 : 211-212

1893 Myxobolus monurus Guriey 1893 : 416

1894 Myxobolus monurus Guiley 1894 : 249-250
1899 Eentuguya monura Labb6 1899 : 103

Habitat: Subcutaneous intermuscular tissue of Aphredoderus sayanus
Gill.; New Jersey (Woodbury).

Vegetative form: Cysts, lenticular, large, white, opaque and numerous
(20). Membrane thin.

Spore: Lenticular or slightly obovate. Tail 2 to 3 times longer than
the body.

HENNEGUYA KOLESNIKOVI (Gurley) Labbe
[Fig. 508]

1886 Kolesnikov 1886 : 242-248

1894 Myxobolus koksnikovi Gurley 1894 : 256-257 •

1898 Myxobolus bicaudatus (part) Zschokke 1898 : 602-604, 646-

655, 699-703

1899 Hmneguya koUsnikovi Labh€ 1899 : 103-104

Habitat: Interstitial connective tissue of the thoracic and intercostal
muscles of Coregonus lavaretus L.; Russia.

Vegetative form: Cysts numerous (80), spherical or oval; length 10
to 30mm., breadth 7 to 20mm.

Spore: Oval with a pointed anterior end. Tail three times longer than
the body.

Remarks: Zschokke thinks the present species is identical with Henne-
guya zschokkei. But the evidence is not clear enough to bring one to
agree with him due to the incomplete description of the present species.

HENNEGUYA MACRURA (Gurley) Thelohan
[Figs. 509 to 512]

1893 Evennann 1893 : 76

1894 Myxobolus macrurus Gurley 1894 : 250-253

1895 Henneguya macrura Thdohan 18^5 : 354

Habitat: Subcutaneous connective tissue of head of Hybognaihus
nuchalis Ag.; Neches River, Texas (November, temperature of water
9*'.4C.) Of frequent occurrence.

Vegetative form: Cysts, elongated 6mm. by 2mm. or less.

Spore: Rounded oblong. Dimensions: length 10 to 11m, breadth
6 to 8/x, thickness 4^. Shell- valves unequally convex. Tail 30 to 40^.

403] STUDIES ON MYXOSPORIDJA—KUDO 165

HENNEGUYA ZSCHOKKEI (Gurley) Doflein
[Fig. 513]

1884

Zschokke

1884 :

: 234-235

1894

Myxoholus (?) zschokkei

Gurley

1894 :

244

1898

Myxobolus bicaudatus (part]

1 Zschokke

1898

: 602-607, 646-
655, 699-703

1898

Myxoholus bicaudatus

Zschokke

1898a

: 213-214

1901

Henneguya zschokkei

Doflein

1901 ;

:202

1904

Henneguya zschokkei

Hofer

1904;

:56

1905

Henneguya zschokkei

Nufer

1905

: 77, 185

Habitat: Subcutaneous and superficial intermuscular tissue of Core-
gonusfera, C. schinzii Fatio, C. hiemalis Jur. and muscular tissue and bran-
chia of C. wartmanni nobilis and C. exiguus albellus; Neuch^teler See, Zurich
See, Genfer-see, Thuner-see, Vierwaldstatter-see.

Vegetative form: Zscholclce writes as follows: Cysts rounded or oval
surrounded by a compact membrane with many nuclei. The largest
32mm. by 16mm. Protoplasm granular. Polysporous.

Spore : Rounded oval in front view ; broad elliptical in side view. Anter-
ior end rounded^ posterior end tapering, forming tail. Sutural ridge
distinct. Tail is either bifurcated along the entire length or a single form,
no intermediate form being observed. Dimensions: total length 5Sfx,
length of the body 10)u, breadth 7/i, length of tail 4 to 5 times the length
of the spore-body, length of polar filament 6 to 10 times that of the body
of the spore.

Remarks: Zschokke thinks that Henneguya kolesnikovi, H. zschokkei
and H. sp. Gurley are one and the same species, for which he proposed
the name Myxobolus bicaudatus.

HENNEGUYA sp. (Gurley) Labbe
[Fig. 514]
1886 Benecke 1886 : 211

1894 Myxoholus sp. inc. Gurley 1894 : 244

1899 Henneguya sp. Labbe 1899 : 104

1904 Henneguya sp. Hofer 1904 : 51

Habitat: Integument (?) of Leuciscus rutilus L. The parasites formed
boil-like enlargement in the skin.
Vegetative form: Not described.
Spote: Not described.

HENNEGUYA sp.

(Gurley)

Labbe

1874

Claparede

1874

:114

1894

Myxoholus sp. inc.

Gurley

1894;

;253

1898

Myxobolus bicaudatus (part)

Zschokke

1898

: 602-607, 646-
655, 699-703

1899

Henneguya sp.

Labbe

1899;

:104

Habitat: Branchial-arches of Coregonus /era; Genfer-see.

166 ILLINOIS BIOLOGICAL MONOGRAPHS [404

Vegetative form: One cyst, 1mm. in diameter.
Spore: Tail short. Zschokke quotes: length 8 to lO/i.
Remarks: According to Zschokke, this species is identical with H.
zschokkei.

HENNEGUYA TENUIS Vaney et Conte
[Fig. 515]
1901 Hmneguya tenuis Vaney and Conte 1901 : 103-106

Habitat: Connective tissue of alimentary tract of Acerina cernua L.;
Lyon (February).

Vegetative form: Numerous cysts particularly in the pyloric coecum.
Usually spherical. Size: 30 to 150/* in diameter.

Spore: Oval and small. Tail short. Two polar capsules at the anterior
end. Sporoplasm with a nucleus, rod-shaped, with somewhat enlarged
ends which is located at right angles to the longitudinal axis. lodinophilous
vacuole could not be traced. Dimensions: length 4/i, breadth 2/i.

HENNEGUYA NUSSLINI Schuberg et Schroder

[Figs. 516 and 517]

1905 Hettneguya nUsdini Schubetg and Schroder 1905 : 56-59

Habitat: Subcutaneous connective tissue at the base of dorsal fin of
Truitafario L.; Gutach.

Vegetative form: Trophozoites form cysts (2 cysts found). Cysts
lenticular, 1.5 to 2mm., surrounded by many concentric layers of fibrous
connective tissue. Cysts containing only mature spores.

Spore: Broad oval form, flattened. Anterior end rounded. Tail at
the posterior end. Shell somewhat thick, often shows sutural ridge. Tail
filaments two. A "dark part" which in side-view is of triangular form,
runs into the tail. Sporoplasm, occupying the posterior half of the spore,
projects a narrow portion between the polar capsules beyond the middle of
the capsules. Sporoplasm, uniformly granular, contains an iodinophilous
vacuole and one, sometimes two nuclei connected by nuclear bridge.
Polar capsules, pyriform, opening independently. Coiled polar filament
observable, coiled 6 to 7 times. Dimensions: length excluding tail 12^,
length with tail 32/x, breadth 8 to 9/x, polar capsules Sn by 3ai, length of
polar filament 4 to 5 times longer than that of spore excluding tail (48 to
60m).

HENNEGUYA L£GERI Cepede
[Figs. 518 to 523]

1905 Hameguya Ugen Cepede 1905 : 905-913

1906 Heitneguya Ugeri Cepede 1906 : 66
1913 Hermeguya Ugen Cepede 1913 : 302-305

Habitat: Urinar\' bladder of Cohitis barbaittla L.; Isere (January).

405] STUDIES ON MYXOSPORIDIA—KUDO 167

Vegetative form: Young trophozoites subcircular, irregularly elliptical
or elongated with distinct differentiation of protoplasm into ectoplasm
and endoplasm. Plasmotomic multiplication takes place during winter
months, when no spore is formed.

Spore: Oval with short tail, mostly bifurcated at the free end. The
anterior end is more rounded, occasionally acuminated. Two polar capsules
of equal size. Coiled polar filament distinct in vivo. Sporoplasm granular,
contains two nuclei and a vacuole. The spore often shrinks in fresh con-
ditions, probably owing to the poorly developed thin valves. Dimensions
of spores mounted in balsam: length variable. Examples: Total length
22.5m, tail 8.5/i; total length 19.5pt, tail 8/i; length of main part 8.5/^, breadth
(comparatively constant) 6/*.

HENNEGUYA ACERINAE Schroder
[Figs. 525 and 526]

1906 Henneguya acerinae Schroder 1906 ; 186-196

1910 Henneguya creplini Wegener 1910 : 84

1911 Henneguya acerinae Nemeczek 1911 : 155

Habitat: Branchiae of Acerina cernua L., Aspro zingel Cuv., Lucioperca
lucioperca L. and L. sandra Cuv. (?) ; Heidelberg (Necker), Apatin, Komitat
Baco-Bodrog, Hungary (May).

Vegetative form: Schroder describes as follows: Rounded or spherical
cysts in the connective tissue of branchial lamella. Full-grown cysts up to
300/1 in diameter. Protoplasm is differentiated into ectoplasm and endo-
plasm. Ectoplasm shows fine radial striations. Endoplasm granular,
contains many nuclei, especially lying in the middle portion. Well devel-
oped cyst, containing only spores, is surrounded by a membrane. On the
surface of the ectoplasm, numerous edge-like elevations, branched and
joining together, were recognized. Polysporous.

Nemeczek observed the largest cyst, spherical and 600// in diameter. .

Spore: Pyriform in front view; flattened. The anterior end is more or
less blunt. Shell uniformly thin. Sutural edge slightly enlarged. Sporo-
plasm finely granular, contains an iodinophilous vacuole and two nuclei.
Polar capsules approximated closely, each having an independent opening.
Dimensions: length 20 to 22)li, breadth 8 to 9/i, thickness 6 to 7/x, length of
tail 50 to 60m, polar capsules lO/x by 2 to 3)u, length of polar filament 80 to
90/ii (water and nitric acid).

Nemeczek's form is as follows:

The tail is bifurcated along its entire length. In one case (May, 1909),
however, all the spores had no bifurcated tail, while the polar capsules were
of unequal size. Dimensions in fresh state: total length 37.6 to 41. 8/i,
length, excluding tail 12.6 to 16.8m, breadth 4.5/x, length of polar capsule
6.3 to 8.4jLi, length of polar filament 67/x, length of tail 25m.

168 ILLINOIS BIOLOGICAL MONOGRAPHS [406

Nemeczek observed two more different (?) forms. One form found in
Lucioperca sandra, tho the size differs from the dimensions given by
Schroder, is thought to be identical with the present species. Another
form in the branchiae of As pro zingel, which is also to be one and the same
species with the present species has the following dimensions : total length
35)u, length of spore excluding tail 15)u, breadth Sn, length of polar capsule
6fx, length of tail 20/x.

HENNEGUYA GIGANTEA Nemeczek
[Figs. 527 to 535]

1911 Henneguya giganlea Nemeczek 1911:146-154

1914 Henneguya gigantea Georg6vitch 1914 : 387-409

Habitat: Branchiae of Lucioperca sandra Cuv.; Apatin, Komitat
Bacs-Bodrog, Hungary, Petrograd. Nemeczek mentions that the infection
takes place only among young fish.

Vegetative form: Cysts numerous and of conspicuous size in the free
end of branchial lamella. In average, each gill-arch has about 100 cysts
which are of creamy color. Young cysts 400 to 450/i in diameter. They
gradually begin to increase the size, from autumn until toward the end of
spring, during which period, the contents remaining in the stages of pan-
sporoblasts formation. Older cysts rounded spindle shape with the length
of 4 to 7mm. and the breadth of 2 to 3mm. The connective tissue and
epithelial cell layers form the cyst membrane. The connective tissue
either simply surrounds the parasite or branches in the surface of the para-
site, increasing in thickness and forming more or less enclosed chambers
of the parasite. The membrane of the cyst which contains mature spores
is usually very thin. Throughout the growth of the cyst, "chromatoid
body" is seen in the endoplasm, which appears first as a filiform struc-
ture,, stained deeply with nuclear stain. Later they gather together and
form a compact body, situated excentrically. Fine branches from it
become directed toward the surface of the body, anastomosing each other
so that a network is formed on the surface of the cyst. The latter develops
small ovoidal or columnal bodies (1.2/u long and about In wide), which
are arranged radially and densely. The number and quantity of these bod-
ies increase in proportion to the number of propagative nuclei and they
begin to disappear, first in the central portion, then in the periphery,
so that in fully grown cysts (in summer months) these chromatoidal bodies
are more rudimentary. Differentiated protoplasm is only recognized in
young individuals, in which case ectoplasm is homogeneous and endoplasm
reticular. Polysporous.

Spore: Nemeczek gives the following accounts.

407] STUDIES ON MYXOSPORJDIA—KUDO 169

Spindle shape, with truncate anterior end and very long thread like
tail at the posterior end. The tail seems split into two at about the middle
part of its length. Gentian violet stains the tail so intensively that its
entire length could easily be made out. Dimensions: total length 87.5 to
IIO.Sm, length of the body 10.5^, breadth 5/i, length of tail 77 to lOO/ii,
length of polar capsule 5/i, length of polar filament 70/i (pressure or dessica-
tion followed by immersion in water).

Georgevitch's form: length excluding tail 15jli, breadth 6/i, length of
tail 75^1, length of polar capsule 6/*, length of polar filament 75/*, diameter of
the iodinophilous vacuole 4/i.

Remarks: Nemeczek mentions that from October on, cysts had no
spores, only containing propagative cells. The velocity of the development
of spores depends upon the temerature of water.

Georgevitch worked out the spore formation of the species and observed
that the binucleated sporeplasm emerged from the posterior end of the
spore.

HENNEGUYA (?) sp. Nemeczek
[Figs. 536 to 539],

1911 Henneguya sp. Nemeczek 1911 : 1S7-1S9

Habitat: Branchiae of Abramis bratna; Komorn, Komitat Komorn,
Hungary (March).

Vegetative form: Cysts in the branchiae.

Spore: Besides normal spores of Myxobolus rotundus (page 149), spores
of Henneguya type in small number were found. The anterior part of
these spores resembles that of the species mentioned above, while the
breadth is much smaller (8/i) than the latter. Majority of spores have a
thread like tail, 10 to 15/i long, which was often bifurcated. An iodinophil-
ous vacuole was fairly marked.

Remarks : It is placed here as a species of Henneguya by reason of the
bifurcate tail.

HENNEGUYA GASTEROSTEI Parisi
[Figs. 540 to 543]

1912 Henneguya gasterostei Parisi 1912 : 296-297

Habitat: Kidney of Gasterosteus aculeatus L.; Lago di Garda (Feb-
ruary).

Vegetative form: Rounded or oval, usually with two, but rarely with
four spores. Ectoplasm thin and hyaline. Endoplasm contains numerous
granules, most probably of fatty nature and decreasing in number as
spores grow. Free full-grown spores were seen abundantly in the connec-
tive tissue of renal tubules, glomeruli, etc. Disporous and polysporous.

170 ILLINOIS BIOLOGICAL MONOGRAPHS [¥»

Spore: Oval with slightly attenuated anterior end; posterior end
tapering into tails, which end in one point or bifurcated; asymmetrical in
shape, one valve is more curved than the other. This asymmetry of the
sheU-valves in profile enables the present species to be distinguished from
other species. Shell striated longitudinally. Two polar capsules pyriform
and well developed, reaching to the middle of the spore. Sporoplasm with
a round iodinophilous vacuole. Dimensions: total length 38 to 48;*,
length of the cavity of the spore IS^t, breadth 6 to 7.5/i, polar capsules 7.5
to 9jtt by 3 to 3.5/i, length of polar filament 50/*.

HENNEGUYA NEAPOLITANA Parisi

[Figs. 544 and 545]

1912 Eenneguya neapolUana Parisi 1912 : 297-298

Habitat: Connective tissue of the renal tubule of kidney of Box sal pa
C. et v.; Napoli (August).

Vegetative form: Small cyst (40 to 50/t in diameter) surrounded by
thin membrane, containing a number of spores, numerous pigment granules
and coarse yellowish globules.

Spore: Oval, slightly flattened. Anterior end rounded when seen from
the front, but attenuated in profile. Shell tapering into a long fine tail
posteriorly. The fine distal portion of the tail wraps around the thicker
part. Two polar capsules, pyriform, occupying the anterior half of the
cavity of the spore, cross each other when seen from the front. Sporoplasm
finely granular with two nuclei, the iodinophilous vacuole being hardly
visible. Dimensions: total length 50 to 60ix, length of the cavity of spore
8.5 to 9.5/4, breadth 8.5 to 9.5/i, internal breadth 6.3 to 7/i, thickness 8/*,
polar capsules 4.7 to 5.5/* by 3/t. «

HENNEGUYA WISCONSINENSIS Mavor et Strasser

[Figs. 558 and 559]

1916 Eenneguya wisconsinensis Mavor et Strasser 1916 : 676-682

Habitat: Urinary bladder of Perca flavescens; Lake Mendota, Wiscon-
sin (April).

Vegetative form: Trophozoites are usually elongated and have the
general form and shape of a limax ameba. It may reach a size of 300/i by
70/i. Clear differentiation of ectoplasm and endoplasm. Pseudopodia
lobose. Two spores are formed in each pansporoblast. Polysporous.

Spore: Ovoid, bilaterally symmetrical, and have a bifurcated caudal
process. Two polar capsules at anterior end. Coiled polar filament visible
in vivo (5 windings). Dimensions: length excluding tail 11.5/i, breadth
7/i, tail 9.6;t, polar capsules 3.5ft by 2.5/i, length of filament 33/i.

4091 STUDIES ON MYXOSPORIDIA— KUDO 171

HENNEGUYA BRACHYURA Ward
[Figs. 650 to 653]
1919 Henneguya brachyura Ward 1919 : 57

Habitat: In the cartilageous fin ray of the caudal fin of Notropis
anogenus] Put-in-Bay, Lake Erie (August). The species was found
encysted in the same fish which was heavily infected by Myxobolus aureatus.

Vegetative form: Cysts rounded with slightly irregular contour im-
bedded in the fin ray. The size varies from 160^ in diameter up to 360/* by
240/x. No particular cyst membrane could be recognized. The differen-
tiation of the protoplasm into ectoplasm and endoplasm is distinct. The
ectoplasm covering the entire surface of the parasite as a layer 4 to 6/it
thick, shows structure of a very finely granular nature. The endoplasm
coarsely alveolar, is filled with mature spores in the central portion, while
numerous nuclei and young spores in various developmental stages are
present at the peripheral portion. Polysporous.

Spore: Rounded oval in front view; spindle shape with symmetrically
built valves in profile. Shell rather thick. Sutural ridge fairly well
marked; sutural edge exhibiting a variable number of folds (8 to 10).
Two pyriform polar capsules are usually of the same size and form. The
tail is a single process, usually more or less bent or irregularly curved,
very rarely being straight. In general, it is sinuous with two or three
shallow curves and is rather short, tapering gradually to a point. In
young spores which are less deeply stained by any stain, various develop-
mental stages of the tail are reasily recognized. Giemsa solution stains
the shell proper in clear blue, while the tail takes on a beautiful pink
color, a distinct difference in affinity for dyes between the material in the
tail and the shell. It seems probable that the tail of this type is entirely
different in its development from that of the ordinary bifurcated type.
Dimensions in section: length 10 to ll.5fi, breadth 8 to 8.75)u, thickness
4 to 5fi, polar capsules 3 to 4/x by 2n, length of the tail up to 17/*.

HENNEGUYA SALMINICOLA Ward
[Figs. 654 to 656]
1914 IHenneguya zschokkei Zschokke and Heite 1914 : 200-201

1919 Henneguya salminicola Ward 1919 : 59

Habitat: In the sub-dermal tissue of Onchorhynchus keta and 0. kisutch
(Zschokke and Heitz, Kamtschatka) and in the connective tissue in body
muscles of Oncorhynchus keta, Stickeen River, Alaska (Ward, September).
The last named author undertook a careful examination of a part of the
infected tissue preserved in formol. The species forms conspicuous C)^ts
in the muscle from the sub-peritoneal to the sub-dermal connective tissue,
tho all are sub-peritoneal in position.

172 ILUNOIS BIOLOGICAL MONOGRAPHS [410

Vegetative form: Ward describes as follows: The whitish opaque cysts
are pjniform, and fairly uniform in size (3 to 6mm. in diameter). The
cyst is covered by numerous layers of connective tissue which form a
tough membrane around the parasite. The cyst contains young spores
in various stages of development, which showed that two spores are formed
in one pansporoblast, and mature spores thickly massed together in the
central area. Polysporous.

Spore: Oval with rounded anterior and more or less attenuated poster-
ior ends; elliptical in profile with attenuated anterior end. Shell smooth.
Sutural edge exhibits folds variable in number (usually 6 to 7). Tail
double, composed of two fine and equal halves which are the prolongation
of the shell valves. The processes usually run roughly parallel to each
other. Two pyriform polar capsules are of slightly difiFerent dimensions.
Coiled polar filament is indistinct in preserved unstained specimens.
Sporoplasm finely granular, shows a large iodinophilous vacuole. Dimen-
sions of stained and mounted spores: total length 47m (42.75 to 52.44/i),
length of the main part Un (11.97 to 14.25>x), breadth 8^ (7.12 to 8.43m),
thickness 4.78At, length of tail 35/* (30.78 to 38. 19m), polar capsule 3.70 to
4.55m by 1.59 to 2.85m.

Remarks: Zschokke and Heitz (1914) observed a species from Kam-
tschatka, which they thought to be identical with Henneguya zschokkei
(page 165). The writer is inclined to think that the species is identical
with the species just described from Alaska.

HENNEGUYA MIYAIRII nov. spec.
[Fig. 524]
1909 Henneguya sp. * Miyairi 1909 : 127-129 <

Habitat: Subcutaneous tissue of head of Carassius auratus L.; Fukuoka
(Nippon).

Vegetative form: Trophozoites form cysts and are also found in the
condition of diffuse infiltration around the cysts. Cyst-membrane fibrous
and thin. Ectoplasm and endoplasm fairly well differentiated, though
the border line is not sharply marked. At the periphery of endoplasm,
pansporoblasts with 7 to 12 nuclei are present (Two spores are formed in
each pansporoblast?). Polysporous.

Spore: Oval, with broadly rounded anterior and slightly elongated
posterior ends, the latter ending in long and fine tails. Two polar capsules
at the anterior portion, are pyriform, small and convergent. Sporoplasm
with an iodinophilous vacuole. Dimensions: length 12m, breadth 8m,
length of the tails 10 to 30m, length of polar filaments 23 to 40m.

Remarks: As the description gives the details by which the species
can be distinguished from other species, the writer estabUshes it on an
independent basis.

4111 STUDIES ON MYXOSPORIDIA—KUDO 173

HENNEGUYA MICTOSPORA nov. spec.
[Figs. 546 to 557]

Habitat: Urinary bladder of Lepomis cyanellus Raf., L. humilis Gir.
and Micropterus salmoides Lac; Stony Creek, 111. (November).

In one out of three (6.5 to 8cm. long) of the first, in one out of two
(7 and 9.5cm. long)" of the second and in one of the third species, examined
in the middle of November, was found the present form. None showed a
heavy infection, a number of scattered trophozoites and spores being
observed. The host did not show any pathological change.

Vegetative form : Polymorphous. Generally rounded or elongated oval.
In small monosporous and disporous forms, the tail of the spores developed
inside, is extruded from the body, so that these trophozoites show long
processes (Figs. 546, 553, 555). Pseudopodia lobose, and extruded from the
entire surface of the body (Fig. 547), tho sometimes they are well formed
at one end of the body. Protoplasm is differentiated distinctly into ecto-
plasm and endoplasm. Ectoplasm is homogeneous and hyaline, forming
the outer layer. Endoplasm is of reticular structure. The body is colorless,
often yellowish, when the endoplasm is loaded with numerous yellowish
coarse granules. The size varies from 6 or 7/i up to 60)Lt. In a rounded form
of 38/i in longest diameter, five pansporoblasts, each developing two spores
and many nuclei were observed. In another oval form of 45^ by 60/* in
size, numerous nuclei were stained, showing that no development of
pansporoblast has yet taken place. Disporous, polysporous and mono-
sporous, tho of rare occurrence.

Spore: Broad spindle shape with attenuated anterior end. Shell rather
thin. Each valve has 6 to 8 longitudinal striations on the surface. A long
tail composed of two halves, is developed at the posterior end. Two pyri-
form polar capsules with distinctly visible coiled polar filament opens at
the anterior tip. Sporoplasm, finely granular, contains an iodinophilous
vacuole which is made distinctly visible by treating with Lugol's solution.
When stained two typical nuclei are recognized in the sporoplasm. Dimen-
sions of the fresh spores: length excluding tail 13.5 to 15/t, breadth 8 to
9/x, thickness 6 to 7.5/i, length of tail 30 to 35/1, often up to 40/i, polar
capsule 5 to 6/* by 3/i, length of polar filament 40/i.

Genu

s HOFERELLUS Berg

1898

Hoferdlus

Berg 1898 : 41

1898

Hoferia

Doflein 1898 : 288-289

The characters of the genus are described on page 59.
Type and only species: Hoferellus cyprini Doflein.

1T4 ILLINOIS BIOLOGICAL MONOGRAPHS [412

HOFERELLUS CYPRINI Doflein
[Figs. 577 to 581]

1898 Hojeriacy print Doflein 1898:289-290

1908 Hoferellus cyprini Mercier 1908 : LIII-LIV

1910 Hoferellus cyprini Plehn 1910 : 20-22

Habitat: In lumen and epithelial cells of renal tubules of kidney of
Cyprinus carpio L.; France and Germany.

Vegetative form: Young trophozoites live in epithelium. Adults free
in the urinary tubules. Form rounded or oval. No clear differentiation of
protoplasm. Pseudopodium unobserved. Endoplasm contains numerous
granules and many nuclei. Each pansporoblast forms two spores. Smaller
individuals 20 to 30/* in diameter. Polysporous.

Spore: Pyramidal with two short tail-like processes at the posterior
end, which are formed from the shell-valves like those of Henneguya.
Between these two processes, rarely small protoplasmic pointed processes
occur. Each shell-valve has 9 to 10 longitudinal striations on it. Two
polar capsules at the anterior part, show clearly the coiled polar filaments.
Sporoplasm has two neclei and an io<iinophilous vacuole. Dimensions:
total length 10 to \2n, breadth 8/i, tail-process 2/i long, polar capsule 3/x long.

MYXOSPORIDIA GENERA ET SPECIES INCERTAE

Gen. et spec, incert. Leydig
1851 Leydig 1851 : 222

1894 Gen. et spec. incerU Gurley 1894 : 186

Habitat: Cysts in the root of tongue of Chondrostoma nasus L.; Ger-
many.

Gen. et spec, incert. Leydig
1851 Leydig 1851 : 222

1894 Gen. et spec, incert. Gurley 1894 : 186

Habitat: Heart (auriculo- ventricular valve) of Leuciscus rutilus L.

Gen. et spec, incert. Leydig

1851 Leydig 1851 : 223

1894 Gen. et spec, incert. Gurley 1894 : 186

Gen. et spec, incert. Heckel et Kner
1851 Heckel and Kner 1851 : 12

1894 Gen. et spec, incert. Gurley 1894 : 186-187

Habitat: Branchiae of Lucioperca lucioperca L.; Austria.

Gen. et spec, incert. Borne
1886 Borne 1886 : 211

1894 Gen. et spec incert Gurley 1894 : 187

Habitat: Scomber scombrus L.

4131 STUDIES ON MYXOSPORIDIA— KUDO 175

Genus incert. MERLUCII Perugia
[Figs. 582 and 583]
1891 Myxosporidium merlucii Perugia 1891 : 22-24

1894 Myxobolus ? merlucii Gurley 1894 : 242-243

1899 Myxobolus merlucii Labb6 1899 : 100

Habitat: Gall-bladder of Merlucius merlucius L.; Italy.

Vegetative form: Various form. No differentiation of protoplasm.
Disporous (?).

Spore: Oval, with two polar capsules.

Remarks: The species was placed in the genus Myxobolus by previous
authors. The figures given by Perugia show that the spores are at least
dimorphous. From the habitat and the disporous characters, one should
place it rather in one of the genera of the Family Ceratomyxidae.

Genus incert. CONGRI Perugia
[Figs. 584 and 585]
1891 Myxosporidium congri Perugia 1891 : 24-25

1894 Genus incert. congri Gurley 1894 : 182

1912 Myxobolus congri Parisi 1912 : 284

Habitat: Gall-bladder of Conger conger L.; Genova.
Vegetative form : Floating in the bladder. Form variable. Movements
incessant, slow and ameboid.
Spore: Not described.

Gen. et spec, incert. Linton

[Fig. 590]

1891

Linton

1891 : 359-361

1894

Gen. et spec, incert. Gurley

1894 : 182-183

Habitat: Subcutaneous tissue of Notropis megahps Raf.; Ohio (Black
River; September, October).

Vegetative form: Cysts. Globular, discrete or aggregated into clusters,
white, with minute patches of black pigment from host; size varying from
2.5mm. (single cyst) to 7mm. by 5mm. (clusters) ; cyst-membrane composed
of connective tissue.

Spore: Top-shaped, somewhat flattened; with pointed anterior and
broadly rounded posterior end. Shell thick, with elevated sutural ridge.
Polar capsules could not be detected. Protoplasm finely granular. Dimen-
sions: length 17/i, breadth 10/t, thickness 6;u.

Remarks : The cysts and figures of spores given by Linton suggest that
it is most probably a unicapsular Myxobolus. As Linton could not detect
(?) the polar capsule, tho his figures faintly show the said structure, it is
placed in this group.

176 ILLINOIS BIOLOGICAL MONOGRAPHS [414

Gen. et spec, incert. Mingazzini
1892 Mingazzini 1892 : 398

1899 Labbe 1899 : 113

Habitat: Ovarian egg of Lacerta sp.

Vegetative form: Ameboid with hyaline pseudopodia and granular
protoplasm.

Spore: Not observed.

Gen. et spec, incert. Nufer
1905 Myxobolus sp. Nufer 1905 : 71, 77, 79, 85, 186

Habitat: In the connective tissue of branchia of Chondrostoma nasus;
Lake of Lucerne. A single cyst in a single host fish.

Vegetative form: Cyst white, and of 1mm. in diameter.

Spore: With two polar capsules at one pole and the sporoplasm.
^ Dimensions or any other characters are not given.

Remarks: Altho Nufer placed the form in the genus Myxobolus, this
must be brought into the present group in view of the fact that the iodin-
ophilous vacuole was not detected, and that the observation is too incom-
plete to place it to any one of the genera.

Gen. et spec, incert. Mavor

[Figs. 586 and 587]

1915

Mavor

1915 : 27-28, 32-33

1916

Mavor

1916 : 553-554

Habitat: Gall-bladder of Urophycis chuss; St. Andrews (July to Sep-
tember).

Vegetative form: Mavor writes as follows:

Attached, usually in large numbers, to the epithelium of the bladder,
occurs a spherical or ellipsoidal trophozoite which in stained preparations
is found to contain numerous nuclei. Very often clusters of Ceratomyxa
acadiensis are found adhering to the free surface of myxosporidium. In
fresh preparations the appearance is that of budding from a parent organ-
ism. An examination of sections has shown a sharp division between the
myxosporidium and Ceratomyxa acadiensis.

Spore: Not found.

Remarks: Mavor supposed that the form under discussion probably
was some species of Myxidium or Chloromyxum.

Gen. et spec, incert. Mavor
[Figs. 588 and 589]
1916 Mavor 1916a : 68-69

Habitat: Urinary bladder of Stizostedion vitreum Mitch.; Georgian
Bay (Canada).

4151 STUDIES ON MYXOSPORJDJA— KUDO 177

Vegetative form: Free forms vary greatly in shape, being rounded,
elongated or branched. The largest individual 200/*, Ectoplasm layer
clearly visible, sometimes projecting many bristle-like short processes.
Endoplasm contains greenish granules. Trophozoites also attached to
the epithelium by means of deeply stainable portion of the body.

Spore: Not observed.

Remarks: Mavor mentions resemblance of the present form to Myxi-
dium lieberkuhni Biitschli in many respects.

178 ILLINOIS BIOLOGICAL MONOGRAPHS [416

KEYS TO THE GENERA AND SPECIES OF MYXOSPORIDIA

No key to the genera and species of Myxosporidia has been published
up to the present time. This is due of course to the difficulties which
accompany such an attempt. These difficulties lie chiefly in the incom-
pleteness of the observations and descriptions of the majority of the species
of Myxosporidia.

The writer has attempted in the following pages to carry out this task.
The key is by no means complete, as is unavoidable in the present state of
knowledge concerning this particular group of the Protozoa.

Altho the spore is the fundamental factor used in constructing this key,
it was necessary to refer also to some other secondary characters such as
vegetative form and habitat.

Some authors are inclined to think that the difference in host species
gives an ample basis on which to record the parasite as a new species. In
some cases the parasite is specific in a certain host species while in other
cases a number of different host species are infected by one and the same
parasite. Therefore one can not lay much emphasis upon a difference of
hosts in fixing the identification of a Myxosporidian.

KEY TO THE GENERA OF MYXOSPORIDIA

1(6) Spore approximately spherical

Suborder Sphaerosporea Kudo 1919 2

2(3) Spore with four polar capsules

Family Chloromyxidae Thdohan 1890

Genus Chloromyxum Mingazzini 1890 (183)

3( 2) Spore with two polar capsules •

Family Sphaerosporidae Davis 1917 4

4( 5) Sutural line of spore straight

Genus Sphaerospora Thdohan 1892 (185)

5( 4) Sutural line of spore sinuous

Genus Sinuolinea Davis 1917 (186)

6( 1) Spore not spherical 7

7(16) Largest diameter of spore at right angles to sutural line; two polar capsules, one
on each side of sutural line

Suborder Eurysporea Kudo 1919

Family Ceratomyxidae Doflein 1899 / 8

8(11) Shell-valves prolonged laterally 9

9(10) Shell-valves hemispherical or rounded

Genus Leptotheca Th^lohan 1895 (179)

10( 9) Shell-valves conical; free end tapering to a more or less pointed end

Genus Ceratomyxa Th61ohan 1892 (180)

11(8) Shell- valves rather elongated; circular in cross-section 12

12(13) Spore rounded oblong; shell longitudinally striated; polar capsules pyriform, with
or without long and fine posterior filaments

Genus Mitraspora Fujita 1912 emend. Kudo 1919 (183)

Numbers enclosed in parentheses refer to the page of the article on which is found the description of the species
named.

4171

STUDIES ON MYXOSPORIDIA—KUDO

179

13(12
14(15:

15(14;

16( 7

17(22

18(21
19(20;

20(19;

21(18;

22(17
23(26;

24(25

25(24;

26(23;

27(30;
28(29;

29(28;
30(27

Spore angular, not rounded 14

Spore pyramidal in front view; with its base at anterior end; with or without dis-
tinct anterior processes; shell smooth

Genus Myxoproteus Doflein 1898 (183)

Spore isosceles triangular in front view; anterior end attenuated; polar capsules
spherical and large; shell with fine network-like ridges; with posterior fringe-
like processes

Genus Wardia Kudo 1919 (183)

Largest diameter of spore coincides with or at an acute angle to sutural plane;
one or two polar capsules which ^e in sutural plane

Suborder Platysporea Kudo 1919 17

Spore fusiform; two polar capsules, one at each end of spore

Family MYXiDiiDAETh61ohan 1892 18

Spore more or less regularly fusiform; shell-valves symmetrical 19

Polar filament fine and long

Genus Myxidium Butschli 1882 (186)

Polar filament thick and short

Genus Sphaeromyxa Th^lohan 1892 (188)

Spore semi-circular in front view; polar filament fine

Genus Zschokkella Auerbach 1910 (188)

Spore not fusiform; with one or two polar capsules at anterior extremity 23

Sporoplasm without iodinophilous vacuole

Family Myxosomatidae Poche 1913 24

Spore elongated ovoid in front view; anterior end mostly pointed

Genus Myxosoma Th61ohan 1892 (189)

Spore more or less rounded in front view

Genus Lentospora Plehn 1905 (189)

Sporoplasm always with iodinophilous vacuole

Family Myxobolidae Thelohan 1892 27

Spore with posterior process; shell sometimes striated 28

Process more or less long, projecting posteriad along median line of spore; process
either single or double; shell sometimes striated

Genus Henneguya Thelohan 1892 (J^^)

Process short projecting posteriad from sides; shell longitudinally striated

Genus Hoferellus Berg 1898 (173)

Spore without posterior process; shell unstriated; one or two polar capsules

Genus Myxobolus Butschli 1882 (189)

11. KEY TO THE SPECIES
• Genus LEPTOTHECA Th61ohan 1895

1(14) Spore: sutural diameter always more than half of greatest breadth 2

2( 7) Average sutural diameter less than 10^ 3

3( 4) Posterior margin of spore concave in front view; sutural diameter 8 to 9n, breadth

12 to 14m. Trophozoite usually with a long process

Leptotheca longipes Auerbach 1910 (63)

4( 3) Posterior margin of spore not concave 5

5( 6) Posterior margin of spore flattened; polar capsules p3ndform; sutural diameter 6

to 7m, breadth 11 to 12^. Trophozoite with actively motile long filiform

pseudopodia at rounded end

Leptotheca agilis Thelohan 1892 (60)

Numbers enclosed in parentheses refer to the page of the article on which is found the description of the species
named.

180 ILLINOIS BIOLOGICAL MONOGRAPHS [418

6( 5) Spore regularly ovoidal ; polar capsules short pynf onn, opening on opposite sides ;
sutural diameter 9/u, breadth \6y.. Trophozoite pyriform without any recog-
nizable pseudopodium

Leptotheca fusiformis Davis 1917 (63)

7( 2) Average sutural diameter of spore equal to or larger than 10/* 8

8(13) Shell-valves symmetrically built; sutural ridge straight 9

9(10) Posterior margin of spore concave in front view; sutural diameter lO/iz, breadth
18 to 20/*; each spore formed independently

Leptotheca informis Auerbach 1910 (63)

10( 9) Spore regularly ovoidal 11

11(12) Trophozoite extremely polymorphous. Spore: sutural diameter 10 to 12/«,
breadth 18 to 20/1

Leptotheca polymorpha Th61ohan 1895 (61)

12(11) Typical form of trophozoite elongated; anterior end depressed surrounded by
short often branched pseudopodia. Spore: sutural diameter 12 to 15/i,
breadth 18 to 20/t

Leptotheca elongata Thdohan 1895 (60)

13( 8) Shell-valves asymmetrically built; sutural ridge sinuous; sutural diameter 9 to
lO/i, breadth 16 to 18/*. Trophozoite rounded; movements slow

Leptotheca lobosa Davis 1917 (64)

14( 1) Sutural diameter equal to or less than half of greatest breadth 15

15(18) Average sutural diameter smaller than 10/« 16

16(17) Spore arch-shaped in front view; polar capsules pyriform; sutural diameter 3 to
4/*, breadth 8 to lO/i

Leptotheca parva Thelohan 1895 (61)

17(16) Spore cylindrical; sutural diameter 4.5/t, breadth 9/»

Leptotheca glomerosa Davis 1917 (65)

18(15) Average sutural diameter greater than 10/* 19

19(20) Posterior margin of spore slightly concave in front view ; anterior end attenuated ;
polar capsules pyriform; sutural diameter 13/*, breadth 26/t. Trophozoite
rounded; with active amoeboid movements

Leptotheca macrospora Auerbach 1909 (62)

20(19) Posterior margin of spore more or less flattened; anterior end smoothly rounded;
polar capsules rounded; sutural diameter 11/*, breadth 22 /t. Trophozoite
elongated; pseudopodia often anastomose

Leptotheca scissura Davis 1917 (64)

Incompletely described species

Leptotheca renicola Thelohan 1895 (61)

Leptotheca hepseti Thelohan 1895 (62)

Leptotheca perlata Gurley 1894 (62)

Leptotheca sp. Awerinzew 1908 (62)

Genus CERATOMYXA Thelohan 1892

1(52) Spore constant in form and size 2

2(21) Sutural diameter equal to or less than one-eighth of total breadth 3

3(10) Sutural diameter not less than one-tenth of total breadth 4

4( 9) Pseudopodia of vegetative form located at rounded end 5

5( 6) Pseudopodia long filiform; with slow whiplash-like movements toward pointed
extremity. Spore: sutural diameter 12/t, breadth 118/*

Ceratomyxa flagelUfera Davis 1917 (77)

Numbers enclosed in parentheses refer to the page of the article on which is found the description of the species
named. ^

419J STUDIES ON MYXOSPORIDIA—KUDO 181

6( 5) Pseudopodia short lobose 7

7( 8) Extremities of spore attenuated; spore large; sutural diameter 10 to 12;*, breadth
90 to 100^

Ceratomyxa spkaerulosa Thdlohan 1892 (66)

8( 7) Extremities of spore rounded; spore small; sutural diameter 4/«, breadth 34 to
39m

Ceratomyxa slreptospora Davis 1917 (79)

9(4) Pseudopodia unlocalized ; from main part of sporulating trophozoite are branched
out from one to six long prolongations. Spore: sutural diameter 5 to 7/«,
breadth 50^

Ceratomyxa a-ppendiculata Thelohan 1892 (67)

10 ( 3) Sutural diameter equal to or less than one- tenth of total breadth 11

11(14) Shell-valve terminating in a fine thread-like process at distal end 12

12(13) Sutural diameter 7 to 8/u, breadth 40 to 50^, lateral process 250 to 300/i

Ceratomyxa acadiensis Mavor 1915 (71)

13(12) Sutural diameter Sfi, breadth 10 to 12/i, length of lateral process 20ju

Ceratomyxa linospora Doflein 1898 (69)

14(11) Shell-valves not terminating in thread-like processes 15

15(20) Shell-valve drawn out into a delicate process 16

16(17) Lateral process ribbon-like; sutural diameter 6ft, breadth 140 to 150m

Ceratomyxa taenia Davis 1917 (74)

17(16) Lateral process not ribbon-like, but circular in cross-section 18

18(19) Posterior margin of main part of spore flattened; sutural diameter 12m, breadth
115 to 140m; trophozoite disporous

Ceratomyxa sphairophora Davis 1917 (73)

19(18) Posterior margin of main part of spore rounded; sutural diameter 7m, breadth
80m. Trophozoite monosporous or disporous

Ceratomyxa spinosa Davis 1917 (80)

20(15) Shell-valve tapering gradually to attenuated point; asjoimietrical; sutural diame-
ter 9m, breadth 115m

Ceratomyxa attenuata Davis 1917 (75)

21( 2) Sutural diameter more than one-eighth of total breadth 22

22(45) Sutural diameter equal to or more than one-fifth of breadth 23

23(34) Shell-valves symmetrically built 24

24(29) Shell-valves attenuated at distal end 25

25(26) Pseudopodia peculiar network-like foim. Spore: sutural diameter 12 to 20m,
breadth 50 to 80m

Ceratomyxa ramosa Awerinzew 1907 (69)

26(25) Pseudopodia never unite together 27

27(28) Shell-valves curved greatly posteriad; polar capsules rounded; sutural diameter
8 to 9m, breadth 16(?)m. Trophozoite elongated pyriform.

Ceratomyxa recurvata Davis 1917 (75)

28(27) Shell-valves not curved; two thickenings on posterior margin equidistant from
sutural line; polar capsules pyrifonn; sutural diameter 40 to 45m, thickness 25
to 30m, breadth 124 to 140m

Ceratomyxa tylosuri Awerinzew 1913 (70)

29(24) Shell-valve rounded at distal end 30

30(31) Spore arch-shaped; sutural diameter and thickness 12 to 15m, breadth 50 to 60m.
Trophozoite large amoeboid.

Ceratomyxa spari Awerinzew 1913 (70)

31(30) Spore straight 32

Numbers eiiclose(} in parentheses refer to the page of the article on which is found the description of the species
named.

182 ILLINOIS BIOLOGICAL MONOGRAPHS [420

32(33) Shell-valves shorter (sutural diameter: breadth = 1:1.6)

Ceratomyxa coris Georg6vitch 1916 (72)

33 (32) Shell-valves longer (sutural diameter : breadth = 1 :2 . 6)

Ceratomyxa herouardi Georgevitch 1916 (72)

34(23) Shell-valves as3Tmnetrically built 35

35(40) Spore arch-shaped in front view 36

36(37) Sutural diameter equal to one-fifth of breadth; sutural diameter lO/t, breadth
50m

Ceratomyxa globulifera Thelohan 1895 (67)

37(36) Sutural diameter more than one-fifth of total breadth 38

38(39) Spore: breadth shorter; sutural diameter 14m, breadth 17m

Ceratomyxa abbreviata Davis 1917 (76)

39(38) Spore: breadth longer; sutural diameter 12 to 15m, breadth 45 to 50m

Ceratomyxa reticularis Thelohan 1895 (68)

40(35) Spore straight 41

41(42) Sutural diameter 11m, breadth 27m. Trophozoite alwajrs roimded

Ceratomyxa amorpha Davis 1917 (78)

42(41) Sutural diameter 6m 43

43(44) Trophozoite with active pseudopodia. Spore: sutural diameter 6m, breadth

22 to 24m

Ceratomyxa undtdala Davis 1917 (79)

44(43) Trophozoite with inactive pseudopodia. Spore: sutural diameter 6m, breadth
31m

Ceratomyxa inaequalis Doflein 1898 (68)

45(22) Sutural diameter less than one-fifth of total breadth 46

46(49) Breadth of Sf)ore equal to or greater than 50m 47

47(48) Shell-valve tapering gradually toward distal end; sutural diameter 8m, breadth
50 to 56m. Trophozoite usually elongated pyriform

Ceratomyxa mesospora Davis 1917 (73)

48(47) Shell-valve rounded at distal end; sutural diameter 6 to 7m, breadth 50m. Tro-
phozoite usually rounded or irregular form; size small

Ceratomyxa aggregate Davis 1917 (79)

49(46) Breadth of spore smaller than 30m SO

50(51) Trophozoite ordinarily spherical, diameter not exceeding 16 to 20m; protoplasm
extremely pale looking. Spore : sutural diameter 5m, breadth 25 to 30m.

Ceratomyxa pallida Thelohan 1895 (67)

51(50) Trophozoite pyriform with a long posterior process; movements by wavelike
motion of ectoplasm; also active backward movements of pseudopodia. Spore
asymmetrically built; sutural diameter 5m, breadth 24 to 28m

Ceratomyxa agglomerata Davis 1917 (77)

52( 1) Spore variable in size and form 53

53(54) Variation in number of shell-valves conspicuous; sutural diameter 5m, breadth
25m

Ceratomyxa truncata Thelohan 1895 (67)

54(53) Variable in size and form of spore, but not in niunber of shell- valve 55

55(60) Trophozoite more or less definite in shape 56

56(59) Trophozoite usually pyriform 57

57(58) Trophozoite disporous. Spore: sutural diameter 7 to 9m, breadth 15 to 38m

Ceratomyxa lunata Davis 1917 (76)

58(57) Trophozoite monosporous or disporous. Spore: sutural diameter 5 to 6m,
breadth 18 to 25m

Ceratomyxa monospora Davis 1917 (78)

Niunbers enclosed in parentheses refer to the page of the article on which is found the description of the species
named.

4211 STUDIES ON MYXOSPORIDIA—KUDO 183

59(56) Trophozoite always rounded, never pyrifonn. Spore: sutural diameter 6/«,
breadth \(>n

Ceratomyxa navicularia Davis 1917 (80)

60(55) Trophozoite polymorphous 61

61(62) Shell-valves symmetrically built; sutural diameter 5 to 8^, breadth 20 to 31/a

Ceratomyxa arcuata Thelohan 1892 (65)

62(61) Shell- valves often asymmetrically built; sutural diameter 8 to 14^, breadth SO to
80/i

Ceratomyxa drepanopsettae Awerinzew 1907 (70)

Incompletely described species

Ceratomyxa sp. (?) Awerinzew 1913 (71)

Ceratomyoca sp. (?) Awerinzew 1913 (71)

Ceratomyxa sp. Georg6vitch 1916 (72)

Genus MYXOPROTEUS Doflein 1898
1( 2) Spore with two long (5ju) processes extending anteriad from sides; sutural diameter
9 IX, breadth 12 m

Myxoproteus cornuttis Davis 1917 (82)

2( 1) Spore without long process 3

3( 4) Spore with two small spinous processes at anterior end; sutural diameter
25 m, breadth 18 to 20/*

Myxoproteus ambiguus Thelohan 1895 (81)

4( 3) Spore without any process; posterior end slightly pointed; sutural diameter 12^,
breadth 10 to 11m

Myxoproteus cordiformis Davis 1917 (81)

Genus WARDIA Kudo 1919
1 Spore isosceles triangular form; shell with network-like striations which end in

fringe-like processes at posterior margin; sutural diameter 9 to 10m, breadth 10
to 12m, diameter of polar capsule 4m.

Wardia ovinocua Kudo 1919 (82)

Doubtfully placed species

Wardia ohlmacheri (Gurley 1894) (83)

Genus MITRASPORA Fujita 1912 emend. Kudo 1919

1( 4) Spore with posterior filaments 2

2( 3) Posterior filaments short (5 to 6m long); length 10m, breadth 8 to 9m, thickness
6 to 8m, polar capsule 4m by 1.5 to 2m

Mitraspora cyprini Fujita 1912 ! (84)

3( 2) Poisterior filaments of spore long (up to 28m) ; length 10 to 11m> polar capsule 4 to
4.5m long

Mitraspora caudata Parisi 1910 (85)

4( 1) Spore without posterior filament; anterior end slightly attenuated; posterior end
truncate; length 15 to 17m, breadth 5 to 6m, thickness 4.5 to 5.5m, polar cap-
sule 7.5m by 2m

Mitraspora elongala Kudo 1919 (85)

Genus CHLOROMYXUM Mingazzini 1890

1( 4) Spore with posterior appendage 2

2( 3) Posterior appendage fine and numerous; length 6 to 9ii., breadth 5 to 6m, polar
capsule 2 to 3 m by 1 to 2 m

Chloromyxum leydigiMrngSiZzmi 1890 * (87)

Numbers enclosed in parentheses refei to the page of the article on which is found the description of the species
named.

184 ILLINOIS BIOLOGICAL MONOGRAPHS [422

3( 2) Posterior appendage single or bifurcated; length 8^, breadth 6 to 7^, appendage
10/t long

Chloromyxum caudatum Th61ohan 1895 (88)

4( 1) Sp)ore without posterior appendage 5

5(34) Spore circular or subcircular in front view; parasitic in body cavity of host 6

6(29) Shell-valves marked with striations or ridges 7

7(24) Main part of striations or ridges parallel to sutural line 8

8(11) Shell-valves partially marked 9

9(10) Ridges on each shell- valve variable in number (six found in original drawing)
ruiming closely to sutural line; diameter lO.Sfi

Chloromyxum duhium Auerbach 1908 (91)

10( 9) Each shell-valve with one ridge from which eight to twelve short ones are directed
toward centre of valve; oval in profile; length and breadth 8 to lO/z, thickness
5 to 7m

Chloromyxum trijugum Kudo 1919 (96)

11( 8) Entire shell-valve marked 12

12(19) Shell-valve marked with fine striations 13

13(16) Spore oval in lateral view 14

14(15) Trophozoite larger; size up to SOn by 20/*; polysporous (up to eight spores) rarely
disporous. Spore: length 8 to 9/x, breadth 6 to 7/x, thickness 5 to 6ft

Chloromyxum misgurni Kudo 1916 (93)

15(14) Trophozoite smaller; size up to 35m in diameter; polysporous (up to six spores)
or disporous. Spore: length 8m, breadth 7m, thickness 5 to 6m

Chloromyxum catostomi Kudo 1919 (98)

16(13) Spore circular in lateral view 17

17(18) Trophozoite rounded; 40 to 45m by 28 to 40m. Spore: diameter 10 to 13m, polar
capsule 4 to 6m long

Chloromyxum protei Joseph 1905 (90)

18(17) Trophozoite irregular form; 33 to 35m in average length. Spore: striations
thicker and somewhat wavy; diameter 9 to 9 . 5m, polar capsule 3n long

Chloromyxum thymalli Lebzelter 1912 (9Z)

19(12) Shell-valves marked with ridges 20

20(21) Trophozoite small (average diameter of adults about 20m) 5 monosporous, rarely
disporous. Spore: shell- valves with ridges marked antero-posteriad; diameter
10 to 11m

Chloromyxum crislatum Leger 1906 (91)

21(20) Trophozoite large, diameter reaching 40 to 50m 22

22(23) Ridges on shell-valves united into a line at each end and unequal in thickness;
spore small; length 10 to 12m, breadth 8 to 10m

Chloromyxum fujitai Kudo 1916 (93)

23(22) Shell-valve with two circular and two small ridges; spore large; length 16m,
breadth 10m

Chloromyxum koi Fujita 1913 (92)

24( 7) Striations or ridges not parallel to sutural line 25

25(26) Striations irregular; posterior margin thickened at sides; diameter 7 . 5 to 9m

Chloromyxum wardi Kudo 1919 (99)

26(25) Striations parallel to each other < 27

27(28) Striations forming acute angles with sutural line; diameter 8 to 9m

Chloromyxum truttae L^ger 1906 (90)

28(27) Four ridges on posterior half of shell- valve converging toward anterior end;
diameter 7 m

Chloromyxum granulosum Davis 1917 (96)

Niunber senclosed in parentheses refer to the page of the article aa which is found the description of the spedes
named.

423] STUDJES ON MYXOSPORIDIA—KUDO 185

29( 6) Shell-valves without marking, beside sutural ridge 30

30(31) Anterior end of spore rounded; diameter 7 to S/z; one or two short spinous thick-
enings at posterior margin

Chloromyxum fluviatile Thdlohan 1892 (89)

31(30) Anterior end of spore mucronate or truncate 32

32(33) Anterior end of spore mucronate; length 8m

CMorotnyxum mucronatum Gurley 1893 (89)

33(32) Anterior end of spore truncate; spore large; length 40 to 48 /x, breadth 30 to 38^

Chloromyxum magnum Awerinzew 1913 (92)

34 ( 5) Spore rounded quadrangular in end view; conical in front view; parasitic in mus-
cular tissue of fish 35

35(36) Length of spore larger than breadth; length 6/i, breadth 5m

Chloromyxum quadratum Thelohan 1895 (88)

36(35) Length (sutural diameter) of spore smaller than breadth 37

37(38) Spore variable in form; anterior end narrower or broader than posterior end;
length 4 to 4.75m, breadth 5.4 to 6.5m

Chloromyxum clupeidae Hahn 1917 (94)

38(37) Anterior end of spore drawn out; almost circular in end view; length 6m, breadth
7.5m

Chloromyxum funduli Hahn 1915 (93)

Incompletely described species

Chloromyxum diploxyi Gurley 1893 (90)

Chloromyxum sp. Awerinzew 1908 (91)

Genus SPHAEROSPORA Thflohan 1892

1(8) Shell-valve of spore without marking except sutural ridge 2

2( 7) Vegetative form amoeboid 3

3( 4) Movements of vegetative form active. Spore: sutural ridge fairly well marked;
a pair of short filaments become visible at anterior end on warming; di-
ameter 8m

Sphaerospora masovica Cohn 1902 (101)

4( 3) Vegetative form without active movements 5

5( 6) Spore: sutural ridge not prominent; polar capsule pyriform; diameter 8 to 13Mf
polar capsule 4 to 5m by 2 . 5 to 3 . 5m

Sphaerospora carassii Kudo 1919 (103)

6( 5) Spore: sutural ridge prominent; polar capsule spherical; slightly attenuated at
anterior end; diameter 10 to 11m

Sphaerospora elegans Thelohan 1892 (100)

7( 2) Vegetative form produces cyst in tissue. Spore: diameter 8 to 9m

Sphaerospora platessae Woodcock 1904 (102)

8( 1) Shell-valves striated 9

9(10) Polar capsules divergent; diameter of spore 10 to 12m, thickness 8m

Sphaerospora diver gens Thdlohan 1895 . . ; (100)

10( 9) Polar capsules not divergent 11

11(14) Striation marked antero-posteriad 12

12(13) Spore with a quadrangular lamella at anterior margin; striations ending in small
spines at posterior margin; length 12 to 14m, breadth 10 to 12m

Sphaerospora rostrata Th61ohan 1895 (101)

13(12) Spore smooth-contoured; polar capsules parallel to each other; diameter 7 to 10m

Sphaerospora polymorpha Davis 1917 (102)

Numbers enclosed in parentheses refer to the page of the article on which is found the description of the species
named.

186 ILLINOIS BIOLOGICAL MONOGRAPHS [424

14(11) Faint concentric striations; pointed at sides and middle part of posterior margin;
polar capsules unequal in size; length 7 to 8m, breadth 6 to 7ai, thickness Sn

Splioerospora angulata Fujita 1912 (102)

Incompletely described species

Sphaerospora sp. Davis 1917 (103)

Sphaerospora sp. Southwell et Prashad 1918 (103)

Genus SINUOLINEA Davis 1917

1( 4) Spore with two processes 2

2( 3) Processes lateral and long (20m); spore: 9 to lift long, 9^ broad, process 18 to
22ft long

Sinnolinea brachiophora Davis 1917 (106)

3( 2) Processes posteriad from sides and short; diameter 12 to 13^. Trophozoite
opaque

Sinuolinea opacita Davis 1917 (106)

4( 1) Spore without process 5

S( 6) Trophozoite with active amoeboid movements. Spore: sutiural ridge S-shaped
at anterior part; length ISm, breadth 12m, thickness 8m

Sintiolinea arborescens Davis 1917 (105)

6( 5) Trophozoite with slow amoeboid movements ■ 7

7( 8) Sutural plane much twisted on its axis; capsulogenous cells large occupying more
than half of sporal cavity; polar capsules opening on opposite sides; diameter
12 to 14m

Sinuolinea capsularis Davis 1917 (lOS)

8( 7) Sutural plane not much twisted; diameter ISm

Sinuolinea ditnorpha Davis 1916 (104)

Genus MYXIDIIIM ButschU 1882

1(16) Breadth of spore equal to or more than half of length 2

2( 7) Shell-valves imstriated 3

3( 6) Sutural plane curved into anS 4

4( 5) Spore small; length 8 to 12m, breadth 4 to 6m

Myxidium incurvatum Thelohan 1892 (108)

5( 4) Spore large; much broader; length 20.8 to 23.4m, breadth 13 to 15.6m

Myxidium inflatum Auerbach 1909 (HI)

6( 3) Sutural plane straight; spore cylindrical; surroimded by a gelatinous envelope;

length 10 to 11m, breadth 6m

Myxidium glutinosum Davis 1917 (115)

7( 2) Shell-valves striated 8

8( 9) Sutural line curved into an S; form oval; circular in cross-section; openings of

polar capsules pointed; length 11 to 13m, breadth 8 to 9m

Myxidium oviforme Parisi 1912 (114)

9( 8) Sutural line straight 10

10(13) Sutural line coincides with longitudinal axis of spore 11

11(12) Sutural ridge distinct; extremities mucronate; length 9 to 10m, breadth 5 to 5 . 6m,

thickness 4 . 75 to 5 m. Vegetative form produces cysts, 800 to 900m in diameter

Myxidium giardi Cep^de 1906 (HO)

12(11) Sutural ridge faintly marked; extremities gradually drawn out; length 11m,

breadth 8m. Trophozoite large and leaf-like; diameter up to 1 .35 mm.

Myxidium phyllium Davis 1917 (116)

Numbers enclosed in parentheses refer to the page of the article on which is found the description of the species
named.

425] STUDIES ON MYXOSPORIDIA— KUDO ' 187

13(10) Sutural line forming an acute angle with longitudinal axis of spore 14

14(15) Shell thickened at extremities; polar capsules ovdidal; length 10 to 14m, breadth
6 to 8iu, length of polar capsule 4^

Myxidium striatum Cunha et Fonseca 1917 (116)

15(14) Shell uniformly thick; polar capsules rounded pyriform; length 10 to 12^, breadth
6n, length of polar capsule 3 to 4/i

Myxidium macrocapsulare Auerbach 1910 (113)

16( 1) Breadth of spore less than half of length 17

17(34) Breadth more than one-third of length 18

18(25) Shell-valves unstriated 19

19(22) Extremities of spore ported 20

20(21) Spore: extremities sharply pointed; greatly curved; narrow; length 12 to 14/x,
breadth 5 . 5 to 6ft, thickness 2.5 to 3 ft

Myxidium depressum Parisi 1912 (114)

21(20) Spore: extremities not so sharply pointed; not greatly curved; broader; length
16.2 to 19m, breadth 7 to 9m

Myxidium bergense Auerbach 1909 (112)

22(19) Extremities of spore not pointed 23

23(24) Spore larger; length 15 to 20m, breadth 7 to 8m

Myxidium sphaericum Th^lohan 1895 (109)

24(23) Spore smaller; length 6(?) to 14m, breadth 4 to 6ft. Trophozoite mictosporous

Myxidium gadi Georg^vitch 1916 (115)

25(18) Shell-valves striated 26

26(33) Spore definite in shape 27

27(28) Spore constricted in middle part of length; length 15m

Myxidium histophilum Th^lojian 1895 (109)

28(27) Spore regularly fusiform 29

29(30) Vegetative form produces cyst. Spore: length 12 to 15m, breadth 6m

Myxidium harhatulae C6pede 1906 (HO)

30(29) Vegetative form does not produce cyst 31

31(32) Sutural line slightly curved in S-form; length 15 to 16m, breadth and thickness

5.5 to 6m

Myxidium americanum Kudo 1919 (117)

32(31) Sutural line not curved in S-shape, but bent to one side; length 15 to 18m, breadth
and thickness 6 to 7 m

Myxidium kagayamai Kudo 1919 (117)

33(26) Spore variable in form; straight and constricted; one side concave, the other con-
vex; arch-shaped, etc.; length 13 to 18m, breadth 5.2 to 5.8m

Myxidium pfeifferi Auerbach 1908 (HI)

34(17) Breadth of spore equal to or less than one-third of length 35

35(40) Shell-valves unstriated 36

36(37) Spore greatly elongated (breadth: length = 1:6. 2); length 2 1.6 to 25. 2 m, breadth

3.6 to 4m

Myxidium procerum Auerbach 1910 (112)

37(36) Spore less elongated (breadth: length = l:3 or 1:3.4) 38

38(39) Spore large; valves asymmetrical; length 28m, breadth 8m

Myxidium giganleum DoiBein 1898 (IK^)

39(38) Spore small; valves symmetrical; length 12m, breadth 3 to 4m

Myxidium danilewskyi Laveran 1897 (109)

40(35) Shell-valves striated 41

41(44) Spore definite in shape 42

Numbers enclosed in parentheses refer to the page of the article on which is found the description of the species
named.

188 ILLINOIS BIOLOGICAL MONOGRAPHS [426

42(43) Length of polar capsule more than one-fourth of that of spore; spore 18 to 20^
long, 5 to 6m broad

Myxidium lieberkuhni Biitschli 1882 (107)

43(42) Length of polar capsule less than one-seventh of that of spore; length 16 to 17/x,
breadth 5^

Myxidium mackiei Bosanquet 1910 (112)

44(41) Spore variable in shape; S-form, straight fusiform, etc.; length 9.1/u, breadth
2 . 8fx. Vegetative form produces cyst

Myxidium anguillae Ishii 1915 (1^4)

Incompletely described species

Myxidium sp. Gurley 1894 (109)

Myxidium sp. Awerinzew 1908 (H^)

Myxidium sp. Mavor 1915 (115)

Genus SPHAEROMYXA Th61ohan 1892

1( 6) Spore straight, not arch-shaped 2

2( 5) Shell-valves symmetrical 3

3(4) Ends of spore truncate ; stria tions longitudinal ; length 1 5 to 20//, breadth 5 to 6/* .

Sphaeromyxa balbianii Thelohan 1892 (H^)

4( 3) Ends of spore rounded; sutural plane forming some angles with longitudinal axis
of spore; striations transverse; length 12 to 14/x, breadth 9 to 10m

Sphaeromyxa immersa Lutz 1889 (119)

5( 2) Shell-valves asymmetrical; unstriated; ends less truncate; dimensions about
twice or three times larger than those of Sphaeromyxa balbianii

Sphaeromyxa gaskrostei Georg^vitch (121)

6( 1) Spore arch-shaped, not straight. . ^ 7

7( 8) Shell-valves indistinctly striated; ends truncate; length 22 to 28m, breadth 3 to
4.3m

Sphaeromyxa sabrazesi Laveran et Mesnil 1900 (120)

8( 7) Shell-valves unstriated 9

9(10) Spore extremely large; length 75 to 80m, breadth 18 to 20m; ends slightly tapering

Sphaeromyxa exneri Awerinzew 1913 (121)

10( 9) Spore less than 35m in length 11

11(12) Extremities rounded; length 30 to 35m, breadth 8m

Sphaeromyxa incurvata Doflein 1898 (119)

12(11) Extremities truncate; sutural ridge often twisted in S-form; length 20.8 to 26m,
breadth and thickness 5 . 4m

Sphaeromyxa hellattdi Auerbach 1909 (121)

Genus ZSCHOKKELLA Auerbach 1910

1( 4) Shell-valves unstriated 2

2( 3) Openings of polar capsules on flattened side; spore large; length 16 to 28.8m,
breadth 13 to 18m

Zschokkella hildae Auerbach 1910 (122)

3( 2) Openings o^ polar capsules at pointed ends; spore small; length 11m, breadth 7m

Zschokkella globulosa Davis 1917 (123)

4( 1) Shell-valves striated 5

5( 6) Openings of polar capsules at pointed ends; polar capsules spherical; spore larger;
length 10 to 14m, breadth 6 to 7m

Zschokkella acheUognathi Kudo 1916 (123)

Numbers enclosed in parentheses refer to the page of the article on which b found the description of the species
named.

427] STUDIES ON MYXOSPORIDIA— KUDO 189

6( 5) Openings of polar capsules on side; polar capsules rounded pyriform; spore
smaller; length 9.5 to 11. 5^, breadth 6.5 to 7^

Zschokkella nova Klokacewa 1914 (122)

Genus MYXOSOMA Th^lohan 1892
1( 2) Spore: shell thickened at anterior end; length 12 to \d>ix, breadth 7 to 8/*, polar
capsule 6 to 7/t by 2 to 3/i. Cysts polymorphous

Myxosoma dujardini Thelohan 1892 (124)

2( 1) Spore: shell of uniform thickness and with seven to ten folds on suturaledge;
length 14^*, breadth 8m, thickness 6^, polar capsule 8/* by 2>i. Cysts spherical
up to 360 jLi in largest diameter

Myxosoma fundtdi Kudo 1918 (125)

Ambiguous form

Myxosoma lobatutn Nemeczek 1911 (124)

1(8:

2( 3

3( 2
4(5:

5(4
6(7

7(6:
8(1

9(io:

10( 9

1(18
2( 9
3( 6

4(5

5(4:

6( 3

7( 8:

Genus LENTOSPORA Plehn 1905

Spore circular in front view 2

Vegetative form produces cysts. Spore: length and breadth 6.3 to Tn, thick-
ness 4.2 to4.9/t

Lentospora dermatobia Ishii 1916 (1^)

Vegetative form does not produce cysts or cysts unobserved 4

Spore small; trophozoites found in the blood vessel of the brain. Spore 5 to 5.5/«
in diameter.

Lentospora encephalina Mulsow 1911 (126)

Spore large, greater than 7 . S^i in average diameter 6

Spore slightly pointed at anterior end; length 8 to 10/t, breadth 7 to 8iu, thickness
5 to6/n

Lentospora acuta Fujita 1912 (127)

Anterior end of spore roimded; diameter 6 to lO/i

Lentospora cerebralis Plehn 1905 (125)

Spore oval in front view 9

Spore symmetrically built; length 12/a, breadth 9.5/i, thickness 6>x

Lentospora multiplicata Reuss 1906 (126)

Spore asymmetrically built; length 10 to ll/x, breadth 6.5 to 7/z

Lentospora asymmetrica Parisi 1912 (126)

Genus MYXOBOLUS Butschli 1882

Spore with one polar capsule 2

Breadth of spore equal to or more than half of length ; 3

Breadth of spore equal to half of length 4

Spore larger; often calabash-shaped; anterior end drawn out into a rounded tip;
shell thickened at tip; length 15^i, breadth 7 to 8/z, thickness 5 to 6/i

Myxobolus toyamai Kudo 1915 (131)

Spore smaller; anterior end pointed; shell of uniform thickness; length 9 to 10|«,
breadth 4.5 to 5.5|i, thickness Six

Myxobolus oculi-leucisci Trojan 1909 (130)

Breadth of spore more than two-thirds of length 7

Polar capsule small and oblique in position

Myxobolus unicapsulatus Gurley 1893 (12i^)

8( 7) Polar capsule long and median in position; spore broader; length 13 . 2 to 13 . 6/t,
breadth 10.1 to 10.3^1

Myxobolus sent Southwell et Prashad 1918 (132)

Numbers enclosed in parentheses refer to the page of the article on which is found the description of the species
named.

190 ILLINOIS BIOLOGICAL MONOGRAPHS [428

9( 2) Breadth of spore less than half of length 10

10(17) Breadth of spore more than two-fifths of length 11

11(16) Spore without any process 12

12(15) Shell of uniform thickness 13

13(14) Spore bent to one side; shell thickened at slightly rounded anterior tip; sutural
edge without marking; length 16 to 18/z, breadth 7 to 8/tt, polar capsule 5
to 7 m long

Myxobolm piriformis Th61ohan 1892 (129)

14(13) Spore straight; shell thickened at posterior part; sutural edge with four to six
markings; length 18 to 20)Lt, breadth 8^, thickness 6)u, polar capsule 9 to 10^
long

Myxobolus fuhrmanni Auerbach 1909 (130)

15(12) Shell of uniform thickness; valves symmetrical; sutural edge with markings up to
12 in number; spore 14 to 15 . 5ju long, 6 to 7 . 3/x broad, 5 to 6m thick, polar cap-
sule 6.3m by 2 to 3m

Myxobolus misgurni Kudo 1919 (133)

16(11) Spore with a posterior process, 5m in length and as broad as spore; length 17 to
18m, breadth 7 .5 to 8m, polar capsule 7m by 4m

Myxobolus notatus Mavor 1916 (131)

17(10) Breadth of spore about one-fourth of length; spore large; polar capsule extremely
large; length 30 to 32m, breadth 7 to 8m, length of polar capsule 22 to 23m

Myx<jibolus rohitae Southwell et Prashad 1918 (132)

18( 1) Spore with two polar capsules 19

19(24) Form of spore variable 20

20(23) Spore with an intercapsular appendix at anterior end 21

21(22) Spore oval; length 10 to 12m, breadth 8 to 9m, thickness 6m, polar capsule 5m by 2
to 3m

Myxobolus midleri BUtschli 1882 (128)

22(21) Spore pyriform or elongated oval; length 11 to 16m, breadth 8 to 13m, polar cap-
sule 6m by 4m

Myxobolus cydoides Gurley 1893 (140)

23(20) Spore without intercapsular appendix; circular form 7 to 8m, breadth 8 to 10m,
thickness 6m, polar capsule 4 to 5m by 2m

Myxo6o/Mj Ay/a€ Johnston et Bancroft 1918 (1S3)

24(19) Form of spore definite 25

25(28) Polar capsules in each spore regularly of considerably different size 26

26(27) Spore with an intercapsular appendix; anterior end roimded; sutural edge with
folds (3 to 5); length 10 to 12m, breadth 8m

Myxobolus dispar Th^lohan 1895 (135)

27(26) Spore without intercapsular appendix; anterior end pointed; no fold on sutural
edge

Myxobolus inaequalis Gurley 1893 (135)

28(25) Polar capsules approximately of equal form and size 29

29(30) Sutural diameter smaller than breadth; length 6 to 7m, breadth 8/i

Myxobohis transovalis Gurley 1893 ' (139)

30(29) Length equal to or more than breadth of spore 31

31(102) Length longer than breadth 32

32(37) Breadth of spore less than half of length 33

33(34) Extremities of spore equally pointed; length 13 to 14.5m, breadth 6 to 7m, polar
capsule 4.5m long

Myxobolus miyairii Kudo 1919 (155)

34(33) Anterior end of spore attenuated; posterior end rounded 35

Numbers enclosed in parentheses refer to the page of the article on which is found the description of the species
named.

429] STUDIES ON MYXOSPORIDIA—KUDO 191

35(36) Shell thickened at posterior margin; spore 12 to ISy. long, 4 to 6.8^ broad, polar
capsule 5.5 to7juby2.1 to2.5/i

Myxobolus anurus Cohn 1895 (142)

36(35) Shell of uniform thickness; spore 14 . 3n long, 6.7|x broad, polar capsule 6.5/* by 2/t

Myxobolus funduli Kudo 1919 (ISl)

37(32) Breadth of spore greater than half of length 38

38(41) Length of spore greater than 20 ft 39

39(40) Spore large; subcircular and anterior end flattened in front view; sutural edge
with markings; length 38 to 45 ju, breadth 32 to 38^1, thickness 28 to35A(, polar
capsule 15 to 17/i long

Myxobolus magnus Awerinzew 1913 (ISO)

40(39) Spore small; extremities equally rounded; length 21/*, breadth 15/i, polar capsule
6n long

Myxobolus cyprini Doflein 1898 (143)

41(38) Length of spore less than 20/t 42

42(43) Spore with a wide (2 to Zn) membraneous posterior process; length 12/x, breadth
10/i, length of polar capsule 4.5/t

Myxobolus cordis Keysselitz 1908 (148)

43(42) Normal spore without appendage 44

44(49) Breadth of sutural ridge one-third of thickness of spore 45

45(46) Length of spore smaller than 10/t; subcircular in front view;*length 7 to 8/*,
breadth 6 to 7/x, thickness 5/x

Myxobolus globosus Gurley 1893 (139)

46(45) Length of spore greater than 10/* 47

47(48) Spore large; elliptical in front view; with an intercapsular appendix, sutural edge
with markings; length 16.9 to 21.6/*, breadth 13 to 16.2/*, thickness 9/*

Myxobolus gigas Auerbach 1906 (145)

48(47) Spore small; subcircular in front view; markings on entire sutural edge;
length 10.8 to 11.7/*, breadth 9.9 to 10.4/*, thickness 7.2 to 9/*

Myxobolus aeglefini Auerbach 1906 (144)

49(44) Sutural ridge narrower 50

50(69) Spore with an intercapsular appendix 51

51(68) Intercapsular appendix triangular 52

52(57) Anterior end of spore attenuated in front view 53

53(54) Sutural edge without marking; length 11 to 12/*, breadth 9.25 to 10.5/i

Myxobolus balleri Reuss 1906 (147)

54(53) Sutural edge with markings 55

55(56) Spore small; length 8 to 9 . 5/*, breadth 6 to 7 . 5/*, thickness 5.5/*

Myxobolus exiguus Thelohan 1895 (1^)

56(55) Spore large; often subcircular; length 1 1 . 5/t to 12/*, breadth 7 . 5 to 8/*, thickness
5/x

Myxobolus obesus Gurley 1893 (140)

57(52) Anterior end of spore broadly rounded in front view > 58

58(59) Posterior portion of spore narrower; polar capsule rather large; length 11.4 to
13.5/*, breadth 9.5 to U/t, thickness 8.5 to 9.5/*, polar capsule 5.5 to 6/* long

Myxobolus discrepans Kudo 1919 (156)

59(58) Extremities of spore approximately equal 60

60(61) Sutural edge without markings; spore 10 to 10.5/* long, 8 to 8.5/* broad, polar
capsule 4.5/* long

Myxobolus squamae Keysselitz 1908 (147)

61(60) Sutural edge with markings 62

62(65) Markings distinct 63

Numbers enclosed in parentheses refer to the page of the article on which is found the description of the species
named.

192 ILLINOIS BIOLOGICAL MONOGRAPHS [430

63(64) Marking variable in number along posterior margin of spore; spore more elon-
gated; length 12 to 12.5a(, breadth 10 to IO.Sai, polar capsule 5.5 to 6^ long

Myxobolus pfeiferi Thdohan 1895 (133)

64(63) Sutural edge with four markings around posterior margin; spore rather short;
length 1 1 to 12m, breadth 9 to 9.5At, thickness 4.5 to 5m, polar capsule 5m by 2.5m

Myxobolus scardinii Reuss 1906 (14^)

65(62) Markings indistinct 66

66(67) Markinp about five at posterior margin; spore larger and shorter; length 11 to
12m, breadth 9.25 to 10m, thickness 4.5 to 5 . 5m, polar capsule 4 to 5m by 2 . 25m

Myxobolus bramae Reuss 1906 (J'i?)

67(66) Markings many along entire sutural edge except anterior tip; spore smaller,
longer and thicker; length 9 . 25 to 10m, breadth 7 to 7 . 25m, thickness 5 to 5 . 5m,
polar capsule 4. 5m by 2 . 5 to 3m

Myxobolus cyprinicola Reuss 1906 (l^)

68(51) Intercapsular appendix rounded; sutural edge smooth; length 11m, breadth 8m,
polar capsule 4 to 6m long

Myxobolus musculi Keysselitz 1908 (i48)

69(50) Spore without intercapsular appendix 70

70(75) Length of spore less than IOm 71

71(72) Spore very much flattened and small; length 6m, breadth 4 . 2 to 5m, polar capsule
3n by 2m

Myxobolus minutus Nemeczek 1911 (ISO)

72(71) Thickness of spore about half of length 73

73(74) Shell thick; length 9 . 25 to IOm, breadth 7 . 25 to 8 . 25m, thickness 4 to 5m

Myxobolus sandrae Reuss 1906 (146)

74(73) Shell thin; spore 8.25 to 9.5m long, 7.25 to 8.25m broad, 4.5 to 5.5m thick

Myxobolus volgensis Reuss 1906 (145)

75(70) Length of spore greater than 10m 76

76(85) Extremities of spore approximately equal 77

77(80) Spore elongated (breadth: length = l:1.8 or 1:1.4) 78

78(79) Spore larger; length 14 to 17m, breadth 8.5m, thickness 5 to 6m

Myxobolus oblongus Gurley 1893 (139)

79(78) Spore smaller; length 12 to 15 m, breadth 9 to 11m

Myxobolus ellipsoides Thdohan 1892 (136)

80(77) Spore shorter (breadth: length = l:1.3, 1:1.2 or 1:1.1) 81

81(82) Sutural edge with markings; slightly truncate at anterior end in front view; spore
9.5 to 11.5m long, 8.5 to 9. 5m broad, 6.5m thick, polar capsule 4. 75m by 1.5
to 2m

Myxobolus mesenlericus Kudo 1919 (157)

82(81) Sutural edge without markings 83

83(84) Polar capsule larger; spore 13 .9m long, 11m broad, 8m thick

Myxobolus lintoni Guriey 1893 (138)

84(83) Polar capsules smaller; spore 14 . 5 m long, 1 1 . 9m broad, polar capsule 6m by 3 . 7m

Myxobolus pleuronectidae Hahn 1917 (152)

85(76) Anterior end of spore more attenuated than posterior 86

86(89) Sutural edge with markings 87

87(88) Markings five or six in number; spore 17 to 18m long, 10 to 13m broad, polar cap-
sule 7 to 8m

Myxobolus permagnus Wegener 1910 (149)

88(87) Markings sometimes present; spore 13 to 17m lo°gj 8 to 10m broad, 5 to 7m thick,
polar capsule 6 to 7 m long

Myxobolus carassii Klokacewa 1914 (150)

Numben enclosed in parentheses refer to the page of the article on which is found the description of the species
named.

431] STUDIES ON MYXOSPORIDIA—KUDO 193

89(86) Sutural edge without markings 90

90(99) Anterior end of spore highly attenuated 91

91(96) Length of polar capsule equal to or less than half of that of spore 92

92(93) Spore: anterior end pointed; length 12.4 to 13.5m, breadth 6.5 to 7.5/tt, thick-
ness 5jix, polar capsule 6 to 7/i long. Cysts of bright golden color

Myxobolus aureatus Ward 1919 (JS4)

93(22) Anterior tip of spore not pointed 94

94(95) Spore greater in thickness (6 . 5 to 7)u) , length 12 to 13 ft, breadth 8 . 25 to 9/*, polar
capsule 6/i by 2.Sm; anterior end more rounded

Myxobolus physophilus Reuss 1906 (1^)

95(94) Spore smaller in thickness (5.5/ti), length 11 to 13^, breadth 8.25 to9.25A», polar
capsule 6/i by 2 . 5 to 3m; anterior end less rounded

Myxobolus macrocapsularis Reuss 1906 (146)

96(91) Length of polar capsule greater than half of that of spore 97

97(98) Length of polar capsule greater than two-thirds of that of spore; spore 16^ long,
10 to 11m broad, polar capsule 11m by 4m

Myxobolus capsulalus Davis 1917 (i52)

98(97) Length of polar capsule less than two-thirds of that of spore; spore 14 to 16m long,
8 to 9m broad, 5 to 6m thick, polar capsule 8 to 9m by 2 . 5 to 3 m

Myxobolus koi Kudo 1919 (155)

99(90) Anterior end of spore rounded 100

100(101) Cysts: size up to 1.7 mm. by 0.7 mm.; parasitic in various tissues of host.
Spore 10 to 12m long, 9m broad

Myxobolus oviformis Th6lohan 1892 (137)

101(100) Cysts: 0.9 mm. by 0.02 mm.; parasitic in nervous system. Spore 10 to 12m
long, 8m broad, 6m thick, polar capsule 6 to 7m by 2m

Myxobolus neurobius Schuberg et Schroder 1905 (144)

102(31) Spore almost circular in front view 103

103(104) Anterior end somewhat attenuated; sutural edge with four markings; spore 9
to 10m long and broad, 6.5 to 7m thick, polar capsule 6 to 7.5m by 2.5 to 3m

Myxobolus orbicuiatus Kudo 1919 (1S5)

104(103) Spore regularly circular in front view lOS

105(106) Cysts large, up to 3 mm. by 1 .5 mm. Spore 10m long, 9.8m broad, 3m thick,
polar capsule 3.8 to 5m long

Myxobolus roiundus Nemeczek 1911 (149)

106(105) Cysts small, up to 0.33 mm. in diameter. Spore 9m in diameter

Myxobolus sphaeralis Gurley 1893 (141)

Incompletely described species

Myxobolus sp. Guriey 1894 (142)

Myxobolus sp. Guriey 1894 , (142)

Myxobolus sp. Guriey 1894 (143)

Myxobolus sp. Miyairi 1909 (149)

Myxobolus sp. Wegener 1^10 (149)

Myxobolus sp. Lebzelter 1912 (150)

Myxobolus sp. Southwell 1915 (1^1)

Myxobolus sp. Kudo 1918 • (132)

Genus HENNEGUYA Th61ohan 1892

1(10) Parasitic in urinary bladder or urinary tubule of kidney of host 2

2( 7) Shell-valves striated 3

Ntunbers enclosed ir pa rentheses refer to the page of the article on which is found the description of the species
named.

194 ILLINOIS BIOLOGICAL MONOGRAPHS [432

3( 6) Length of tail equal to two-thirds of length of main part of spore* 4

4( 5) Shell-valves asymmetrical; spore smaller; total length 38 to 48/*, length of main
part 15/i, breadth 6 to 7.5;^, polar capsule 7.5 to 9m by 3 to 3.5m. Tropho-
zoite disporous and polysporous

Henneguya gasterostei Parisi 1912 (169)

S( 4) Shell-valves symmetrical; spore broader; length of main part 13.5 to 15ai,
breadth 8 to 9^, thickness 6 to 7 . 5m, polar capsule 5 to 6m by 3ft, length of tail
30 to 40m. Trophozoite mictosporous

Henneguya mictospora Kudo 1919 (173)

6( 3) Length of tail equal to half of length of main part of spore and single(?) ; length
20 to 24m, breadth 5 to 6m, length of polar capsule 4 to 5m

Henneguya media Thelohan 1892 (161)

7( 2) Shell-valves unstriated 8

8( 9) Spore elongated; polar capsule longer; posterior portion of main part broad; tail
wider and bifurcated to same direction; total length 19.5 to 22.5m, length of
main part 8.5m, breadth 6m, length of tail 8 to 8.5m

Henneguya Ugeri Cepdde 1905 (166)

9( 8) Spore oval; polar capsule shorter; posterior part of main portion narrow; tail
narrower and bifurcated to opposite directions; length of main part 11.5m,
breadth 7m, length of tail 9.6m, polar capsule 3.5m by 2.5m

Henneguya wisconsinensis Mavor et Strasser 1916 (170)

10( 1) Parasitic in tissue of host 11

11(28) Tail always appears as a single process 12

12(13) Spore small; length 4m, breadth 2m

Henneguya tenuis Vaney et Conte 1901 (166)

13(12) Spore longer and larger, at least 27m long 14

14(15) Sutural edge with eight to ten markings; tail rather long; length of main part 10
to 11 .5m, breadth 8 to 8 . 75m, thickness 4 to 5m, polar capsule 3 to 4/* by 2m, tail
up to 17m long

Henneguya brackyura Ward 1919 (171)

15(14) Sutural edge without markings 16

16(19) Total length of spore greater than 40m 17

17(18) Anterior end rounded; polar capsule large; sheU-valves asymmetrical; tail long;
main part 10 to 11m long, 6 to 8m broad, 4m thick, tail 30 to 40m long. Cysts
elongated and large up to 6 mm. by 2 mm.

Henneguya macrura Gurley 1894 (164)

18(17) Anterior end attenuated; polar capsule smaller; shell-valves symmetrical; tail
shorter; length 20m, breadth 8 to 9m, polar capsule 8m by 2 to 3m. Cysts
elongated oval and smaller, 1 . 1 mm. by 0.5 mm.

Henneguya creplini Gurley 1894 (162)

19(16) Total length of spore less than 40m 20

20(21) Tail about one- third of main part; total length 38m, niain part 26m long, 10 to 11m
broad, 8m thick, polar capsule 11 to 14m by 2 to 3m. Cysts oval, numerous and
smaU (130m by 115m)

Henneguya minuta Cohn 1895 (160)

21(20) Tail about three-sevenths of main part; total length 29 to 40m, main part 15 to

20m long, 7 to 8m broad, polar capsule 8m by 2m 22

22(25) Cysts large 23

*Length of main part of spore denotes in all possible cases the distance between the outer
anterior tip and the posterior margin of sporal cavity; and consequently that between the
latter and the distal end of the tail is the length of the taiL

Numbers enclosed in parentheses refer to the page of the article on which is found the description of the q>ecies

named.

433] STUDIES ON MY XOSPORIDI A— KUDO 195

23(24) Cysts spherical up to 6 mm. in diameter; parasitic in ovary

Henneguya oviperda (Cohn 1895) (^60)

24(23) Cysts elongated up to 2 mm, by 1 .5 mm.; parasitic in branchia

Henneguya psorospermica Thelohan 1895 (^58)

25(22) Cysts rather small or size not observed 26

26(27) Elongated cysts 0.75 mm. by 0.375 mm.; parasitic in branchia

Henneguya texta (Cohn 1895) (159)

27(26) Parasitic in intestinal wall

Henneguya peri-intestinalis Cep^de 1906 (161)

28(11) Tail composed of two processes 29

29(30) Total length reaches 87.5 to 110. 5^; length of main part 10.5 to 15/x, breadth
5n, length of polar capsule 5/i, length of tail 75 to lOO/x

Henneguya gigantea Nemeczek 1911 (168)

30(29) Total length of spore less than S2/j. 31

31(34) Sutural edge with markings 32

32(33) Tail longer and spore larger; anterior end more rounded; total length 47 /i. Cysts
spherical and large, up to 6 mm.

Henneguya salminicola Ward 1919 (171)

33(32) Tail shorter and spore smaller; anterior end slightly more attenuated; total
length 32 ju. Cysts lenticular, up to 2 mm. in length

Henneguya niisslini Schuberg et Schroder 1905 (166)

34(31) Sutural edge without markings 35

35(38) Distal end of tail thread-like 36

36(37) Tail 40 to 50^ in length; total length 50 to 60^, main part 8 . 5 to 9 . 5/x long, 8 . 5 to
9.5m broad, polar capsule 4.7 to 5. 5m by ?>n. Cysts small, up to 50^ in
diameter

Henneguya neapolitana Parisi 1912 (170)

37(36) Tail 10 to 30/* in length; main part 12^ long, 8/x broad

Henneguya miyairii Kudo 1919 (172)

38(35) Distal end of tail tapers to a point and not thread-like 39

39(40) Cysts irregular in shape; size up to 2.5 mm. Spore: total length 30 to 40/x,
main part 11.5 to 15^ long, 5 to 6.5^ broad, polar capsule 6.5 to 8/x by 2 to
2.5m, tail 22 to 28m

Henneguya lobosa (Cohn 1895) (161)

40(39) Cysts spherical or oval 41

41(42) Anterior end of spore rounded; tail either single or double processes; total length
55m, length of main part 10m, breadth 7m, length of tail 40 to 50m. Cysts
spherical or oval up to 32 mm. by 16 mm.

Henneguya zschokkei (Gurley 1893) (165)

42(41) Anterior end attenuated; spore large; main part 20 to 22m long, breadth 8 to
9m, 6 to 7m thick, polar capsule 10m by 2 to ifi, tail 50 to 60m long

Henneguya acerinae Schroder 1906 (167)

Incompletely described species

Henneguya schizura (Gurley 1893) (162)

Henneguya linearis (Gurley 1893) (163)

Henneguya gurleyi Kudo 1893 (163)

Henneguya monura (Gurley 1893) (164)

Henneguya strongylura (Gurley 1894) (163)

Henneguya kolesnikovi (Gurley 1894) (164)

Henneguya brevis Thelohan 1895 (162)

Henneguya sp. (Gurley 1894) (165)

Henneguya sp. (Gurley 1894) (165)

Henneguya sp. Nemeczek 1911 (169)

Numbers enclosed in parentheses refer to the page of the article on which is found the description of the species
named.

196 ILLINOIS BIOLOGICAL MONOGRAPHS [434

SUMMARY

1) All species of Myxosporidia which have been observed in various
parts of the world, reaching 237 in number, are recorded with figures.

2) A new classification of Myxosporidia is proposed after discussion
of those of previous authors.

3) A complete list of the specific names of the hosts that harbor Myxo-
sporidia, is given together with the names of the organ of infection and
the localities from which recorded.

4) By study of the geographical distribution of Myxosporidia, it
is shown that few species are common both to American and European
waters or Asiatic and European waters, while the majority of Myxosporidia
are localized in definite and limited regions.

5) The study of the organal distribution of Myxosporidia in the host,
shows that the gall-bladder is the organ most frequently invaded by the
parasite. The kidney, branchia and urinary bladder have less chances of
being attacked.

6) One new genus, Wardia, is established.

7) Nine new species; Wardia ovinocua, Mitraspora elongaia, Chloromy'
xum trijugum, Chloromyxum catostomi, Myxidium americanum, Myxoholus
orhiculatus, Myxoholus discrepans, Myxoholus mesentericus and Eenneguya
mictospora, are described from fresh-water fish collected in the vicinity
of Urbana, HI.

8) Six new species; Sphaerospora carassii, Myxidium kagayamai,
Myxoholus tnisgurni, Myxoholus miyairii, Myxoholus koi and Eenneguya
miyairii, are recorded from fresh-water fish of Nippon.

9) One new species; Chloromyxum wardi, is described from Alaska.
This is the second species of Myxosporidia from that part of North America.

10) Keys to the genera and species of known Myxosporidia are in-
cluded.

435] STUDIES ON MYXOSPORIDIA—KUDO 197

APPENDIX: NEW MYXOSPORIDIA FROM AUSTRALIA

The following six species described by Johnston and Bancroft did not
reach the wtiter until the page proof was read. For this reason they could
not be put in the text and are recorded here separately,

MYXIDIUM THERAPON Johnston et Bancroft
1919 Myxidium therapon Johnston and Bancroft 1919 : 520-521

Habitat: In the gall bladder of Therapon carho and Th. hillii; Thomson
River at Longreach, Australia.

The parasite occurred in one specimen of the former host fish and in
nine out of thirteen specimens of the latter. No visible effect of the infec-
tion on the part of the host fish was recognized.

Vegetative form: Body pale yellowish to green in color. Form(?).
Size varies from 3 to 12mm. in diameter. The protoplasm is differentiated
into a clear narrow ectoplasm, about 10/i in width, and a coarsely grained
endoplasm. No movements could be seen on slides; but undulations were
observed to travel round the margin of the trophozoite. Polysporous.

Spore: Spindle-shaped with slightly pointed extremities. Polar cap-
sules are more or less rounded. Shell with faintly marked longitudinal
stria tion. The sporoplasm is binucleated. Average dimensions: length
9 to 10m, breadth 4/x, polar capsules 2 to 3/i by 1 to 2/x.

MYXOSOMA OGILBYI Johnston et Bancroft
1919 Myxosoma ogilbyi Johnston and Bancroft 1919 : 521-522

Habitat: In the fibrous tissue of the gill arch of Plectroplites amhiguus;
Thomson River at Longreach, Australia. Three out of nine host specimens
examined showed the infection.

Vegetative form: The parasite forms white cysts usually close to the
bases of the gill filaments. Cysts are small and rounded, being less than
1mm. in diameter. The authors simply mention that sections revealed the
structure usually present in a Myxosporidian cyst.

Spore : Oval with pointed anterior end. The inner margin of the shell
is indented posteriad. The sporoplasm contains a single nucleus, but not
any iodinophilous vacuole. Average dimensions: length 11 to 13ju, breadth
6 to 2>ix, thickness 5/x, polar capsules 5 to 6/i by 2n.

MYXOBOLUS PLECTROPLITES Johnston et Bancroft

1919 Myxobolus plectroplites Johnston and Bancroft 1919 : 522-523

Habitat: In the kidney and gall-bladder of Plectroplites amhiguus;
Thomson River at Longreach, Australia. The parasite was observed in

198 ILLINOIS BIOLOGICAL MONOGRAPHS [436

four out of nine host fish examined; in two cases in the kidney only, in one
case only in the gall-bladder, and in one instance in both gall-bladder and
kidney. Cysts were found in the kidney, while only spores were recognized
in the gall-bladder.

Vegetative form: The cysts which could only be detected in sections,
lie in the connective tissues of kidney. They are of minute size, ranging
somewhat widely from 36m in diameter to 144 by 100/i. According to the
authors no definite structure could be found.

Spore: Rounded oval. It bears quite a close resemblance to that of
Myxobolus hylae (page 153), which is slightly longer, and which has a longer
polar filament than the present form. The vacuole, however, is apparently
not iodinophilous(?). Average dimensions: length 10 to 12ju, breadth 7 to
8/x, polar capsules 5 by 2^, length of polar filament 30 to 40^.

HENNEGUYA AUSTRALIS Johnston et Bancroft
1919 Henneguya australis Johnston and Bancroft 1919 : 523-524

Habitat: In the branchiae of Plectroplites amhiguus; Thomson River
at Longreach, Australia. The parasite was detected in four out of nine host
fish examined. The infection was extremely light in all cases.

Vegetative form: The parasites form cysts. They lie embedded in the
spongy mass of the gill filament, and in many cases occupy a relatively
large area of the section. Cysts showed two layers in section; the outer-
most clear ectoplasm and inner endoplasm with developing spores, the
central portion of which being filled with mature spores. The spores appear
to lie in a definite manner, the long axis of the spore commonly being at
right angles to the boundary of the cyst, the anterior end of the spore
pointing outwards.

Spore : Elongated ovoidal. Anterior end pointed, posterior end drawn
out into a tail. The tail appears single when the spore is removed from the
cyst but separates soon afterward into two halves which usually diverge
widely. Two polar capsules parallel to each other are quite frequently of
different length. The sporoplasm contains two nuclei and a small vacuole
(iodinophilous?). Average dimensions: length 11 to IS/x, breadth 3 to 5/i,
thickness 3 to 4/z, polar capsules 5 to 6^ by 1 to 2jli, length of tail about 20/Lt.

HENNEGUYA GRACILIS Johnston et Bancroft
1919 Henneguya gracilis Johnston and Bancroft 1919 : 524-526

Habitat: In the gill filament of Therapon hillii; Thomson River at
Longreach, Australia. Out of thirteen specimens examined, eight were
infected. Heavy infection was recognized only in one case.

Vegetative form: The cyst is of definite, narrow, pear-shaped form, and
lie transversely, i.e., at right angles to the long axis of the gill filament.

437] STUDIES ON MYXOSPORIDIA—KUDO 199

Spore: The spore resembles Eenneguya australis, but is slightly
smaller, while the tail is longer in proportion. The spores are arranged
with long axis parallel to that of the cyst. Average dimensions: length
10 to 14/i, breadth 2 . 5 to 3^. thickness 3ju, polar capsules 5 to 6/i by 1 to
2/1, length of tail about 20 to 26)u.

HENNEGUYA sp. Johnston et Bancroft
1919 Henneguya sp, Johnston and Bancroft 1919 : 526

Habitat: In the branchiae of Nematalosa elongata; Thomson River at
Longreach, Australia.

The authors state that they observed a number of spores of a Henne-
guya in the scrapings of the gill of one of four host fish.

Vegetative form: Undescribed.

Spore: Undescribed.

200 ILLINOIS BIOLOGICAL MONOGRAPHS {438

BIBLIOGRAPHY

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Balbiani, G.

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439] STUDIES ON MYXOSPORIDIA—KUDO 201

Berg,

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441] STUDIES ON MYXOSPORIDIA—KUDO 203

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1893. Ueber die Drehkrankheit der Regenbogenforelle. Allg. Fishereiz., 18:7-8.

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*Original paper not seen.

204 ILLINOIS BIOLOGICAL MONOGRAPHS [442

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1888. Ueber Piesbergens Fischsporospermien. Centralbl. Bakt. u. Parasit., 3:663-664.
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1909. Exhibit of Myxobolus. Proc. Roy. Soc. N.S.W., 43:29.
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1918. A parasite, Myxoholus hylae sp. nov., of the reproductive organs of the golden

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1905. Chloromyxum protei n. sp. Zool. Anz., 29:450-451.

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1914. Ueber die Myxosporidien der Karausche. Zool. Anz., 44:182-186; 2 fig.

KOLESNIKOFF, N. F.

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1915. On a new Myxosporidian from a carp (Nipponese). Joum. Zool., 27 :5l7-523; 2 pi.

1916. Contributions to the study of parasitic protozoa. Ill, Notes on some Mjrxospori-

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1917. Contributions to the study of parasitic protozoa. II. Myxobolus toyamai nov.

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1918. Experiments on the extrusion of polar filaments of Cnidosporidian spores. Joum.

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1899. Sporozoa. Das Tierreich. 5 Lief. 180 pp.
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1900. Sur une Myxosporidie des voies biliares de I'Hippocampe (Sphaeromyxa sahrasesi

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♦Original paper not seen.

443] STUDIES ON MYXOSPORIDIA—KUDO 205

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1852. Parasitismus und Parasiten. Arch, physiol. Heilkde, 1 1 :434-436.
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1891. On certain wart-like excrescences, occurring on the short minnow, Cyprinodon

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1889. Ueber eine Myxosporidium aus der Gallenblase brazilianischer Batrachier (Cys-
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1915. Studies on the Sporozoa of the fishes of the St. Andrew's Region. Ann. Rep.
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1916. On the life-history of Ceratomyxa acadiensis, a new species of Myxosporidia from
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*Original paper not seen.

206 ILLINOIS BIOLOGICAL MONOGRAPHS [444

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MiLEWSKI, A.

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1912. An Introduction to the study of the protozoa. 517 pp.; 194 textfig.
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1909. Guide to the study of parasitic protozoa (Nipponese). 170 pp.; 8 pi.
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1906. Ueber die Gelbsucht der Bachforelle. Osterr. Fischereiz. 3 :285.

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1841, Ueber eine eigenthiimliche krankhafte parasitische Bildung mit specifisch organi-"

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1911. Ein neuer Gehimparasit des Karpfens. Allg. Fisch. Zeit., 36:483-485; 3 fig.
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445] STUDIES ON MYXOSPORIDIA— KUDO 207

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1890. Die Protozoen als Krankheitserreger. 1 ed. 100 pp.; 34 fig.

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*Original paper not seen.

208 ILLINOIS BIOLOGICAL MONOGRAPHS [446

Schroder, O.

1906. Eine neue Myxosporidienart aus den Kiemen von Acerina cernua {Henneguya
acerinae n. sp.). Arch. Protist., 7:186-196; 1 pi.
-t4907. Beitrage zur Entwicklungsgeschichte der Myxosporidien. Sphaeromyxa sabrazesi
Laveran et Mesnil. Arch. Protist., 9:359-381; 3 textfig., 2 pi.

1910. Ueber die Anlage der Sporocyste (Pansporoblast) bei Sphaeromyxa sabrazesi

Laveran et Mesnil. Arch. Protist, 19:1-5; 10 fig.

1912. Myxosporidien. Prowazek's Handbuch Path. Protoz., 1^324-336.
ScHUBERG, A. and O. Schroder.

1905. Myxosporidien aus dem Nervensystem und der Haut der Bachforelle (Myxoholus
neurobius n. sp. and Henneguya nUssUni n. sp.). Arch. Protist., 6:45-60; 1 pi.

SOUTUWKLL, T.

1915. Notes from the Bengal Fisheries Laboratory, Indian Museum. On some Indian
Parasites of Fish with a note on Carcinoma in Trout. Rec. Ind. Mus., 11 :311-
330; 2 pi.
Sotn^WEix, T. and B. Prashad.

1918. On some Indian Myxosporidia. Rec. Ind. Mus., 15 :344r-348; 1 pi.
Stazzi, p.

m 1906. Psorospermosi o myxoboliasi de BarbL Riv. mens. pesca.,Milano, 8:14-19; 3 fig.
Sttrbeck, G.

1900. Eine neue Krankheit beim Bachsaibling (Salvelinus fontinalis). Allg. Fischereiz.,
25:367-368.

1911. Eine grosse Sporencyste von Henneguya zschokkei. Schweiz. Fisch.-Zeitg., 19:

163-165; 1 fig.

1913. Ueber eine eigenartige Form des Auftretens von Henneguya zschokkei Gurley.

Schweiz. Fisch.-Zeitg., 21:30-31.

1914. BeitragzurFischpathologie. lu. II. Schweiz. Fisch.-Zeitg., 22:296-299.

TsfeLOHAN, p.

1889. Sur la constitution des spores des Myxosporidies. C.R. acad. sci., 109 :919-922.
-x 1890. Recherches sur le developpement des spores chez les Myxosporidies. C.R. soc.
biol., 2:602-604.
1890a. Nouvelles recherches sur les spores des Myxo^xjridies (structure et developpe-
ment). C.R. acad. sci., 111:692-695.
1890b. Contribution k I'fitude des Mj^xosporidies. Microgr., 2:193-213; 1 pi. Annal.
1891. Sur deux Sporozoaires nouveaux, parasite des muscles des poissons. C.R. soc.
biol., 3:27-29.
fl fl «v^ 1892. Observation sur les myxosporidies et 6ssai de classification de ces organismes.
f'^^ -^ Bull. soc. philom., 4:165-178.

1892a. Myxosporidies de la vesicule biliaire des poissons. C.R. acad. scL, 115561-964,
1091-1094.

1893. Alteration du tissue muscvdaire duhs k la pr&ence de Myxosporidies et de microbes

C.R. soc. biol., 5:267-270.

1894. Sur les aflSnitfe r6ciproques des Myxosporidies. C.R. acad. sci., 118:428-430.

1895. Recherches sur les Myxosporidies. Bull. sc. France et Belg., 26:100-394; 6 fig.,

3 pi.
TiojAN, E.

1909. Ein MjTJobolus im Auge von Leuciscus rutilus. Zool. Anz., 34:679-682; 3 figs.
Tyzzer, E. E.

1900. Tumors and Sporozoa in fishes. Joum. Boston Soc. Med. Sc, 5:62-68; 1 pi.
Vaney, C. and A. Conte,

1901. Sur deux nouveaux Sporozoaires dndospores parasites de VAcerina cernua Cuv.

Aim. Soc. Linn. Lyon, 47: 103-106; 4 fig.

447] STUDIES ON MYXOSPORJDIA—KUDO 209

Ward, H. B.

1919. Notes on North American Mjrxosporidia. Journ. Parasit., 6:49-64, 1 pi,
Wasieiewsky, von

1896. Sporozoenkunde. Jena. 162 pp.; Ill textfig.
Wegener, G.

1910. Die Ektoparasiten der Fische Ostpreussens. Inaugur.-Dissert. Konigsberg, p.
72^0; 4 figs.
Weltnes, W.

1892. Ueber Myxosporidien Sporen in den Eiem von Esox lucius. Sitz. Ges. nat. Fr.

Berlin, p. 7-41; 16 fig.
Whinery, J. B.

1893. Some additional notes on a M3Tcosporidian infection in the common toad. N. Y.

Med. Journ., 18:660H562; 1 fig.
Woodcock, H. M.

1904. On M3Tcosporidia in flat-fish. Trans. Biol. Soc. Liverpool, 18:46-62; 1 pi.
1907. On a Myxosporidian infection of Gadus estnarkii. With Johnstone.

ZSCHOKKE, F.

1884. Psorospermies de Coregonus fera. Arch. biol. gand, Leipzig et Paris, 5:234-235;

Ipl.
1898. Die Myxosporidien der Gattimg Coregonus. Centralbl. Bakt. u. Parasit., Abt. L

Orig., 23:602-607, 646-655, 699-703; 4 fig.
1898a. Die M3TJOsporidien in der Muskulature der Gattung Coregonus. Zool. Anz.,

21:213-214.
1898b. Myxobolus bicaudatus n. sp., ein Parasit der Coregoniden des Vierwaldstattersees.

Mitt. Naturf. Ges., Luzem, p. 205-217.
1900. Myxobolus psorospermicus Th61ohan im Vierwaldstattersee. Mitt. Naturf. Ges.

Luzem, p. 441-442.
ZscHOKKE, F, and A. Heitz.

1914. Entoparasiten aus Salmoniden von Kamtschatka. Revue Suisse zool., 22:195-

256.

210 ILLINOIS BIOLOGICAL MONOGRAPHS [448

GENERAL EXPLANATION OF FIGURES

For the type species of each genus, both the vegetative form and the
spore are illustrated. For the other species, except those which are new,
figures of the spore are given, unless the vegetative forms are different
from those of the type species or the species were reported in papers which
seem to be of less universal distribution.

The original drawings were made with the Abbe drawing apparatus.
The combinations used were Zeiss apochromatic objectives 16, 8, 3, and
homogeneous oil immersion 2 mm. with compensation oculars 2, 4, 6, 8,
12 and 18. All the other drawings were copied from the original figures of
the respective observers, an exact citation of which is given in each case,
and were also made with the same drawing apparatus on the same scale
except that a few figures were enlarged among those that were taken
from other authors.

Magnifications were also calculated and given for those quoted figures,
for which the respective authors failed to mention the scale at which the
drawings were made.

44^ STUDIES ON MYXOSPORIDIA— KUDO 211

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212 ILUNOIS BIOLOGICAL MONOGRAPHS [450

EXPLANATION OF PLATE
Figs. 1 to 5. Leptotkeca agUis.

Fig. 1. A typical trophozoite in vivo. After Thfilohan (1895, Fig. 29); X750. *
Fig. 2. A young form. After Th^lohan (1895, Fig. 31). X750.
Fig. 3. A trophozoite in motion. After Doflein (1898, Fig. 5).
Fig. 4. A trophozoite in contracted condition. After Doflein (1898, Fig. 7).
Fig. 5. A fresh spore. After Th61ohan (1895, Fig. 30). X1500.
Fig. 6. A fresh mature spore of Leptotkeca dongata. After Thdohan (1895, Fig. 38).

X1500.
Fig. 7. A fresh mature spore of Z^/><o//(«ca ^orra. After Thdohan (1895, Fig, 25). X1500.
Fig. 8. A fresh spore of Leptotkeca perlata. After Balbiani (1884, Fig. 40).
Fig. 9. A spon oi Leptotkeca macros pora. After Auerbach (1909, Fig. 2a). X1350.
Fig. 10. A spore of Leptotkeca informis, preserved in formol. After Auerbach (1910b, Fig.

la). X about 2000.
Fig. 11. A spore of Leptotkeca longipes, preserved in formol. After Auerbach (1910b, Fig.

Id). X about 2200.
Fig. 12. AiresiispoitoiLeptotkecafusiformis. After Davis (1917, Fig. 1). X1500.
Fig. 13. Airesh spore ol Leptotkeca scissura. After Davis (1917, Fig. 8). X1500.
Fig. 14. A fresh spore of Leptotkeca lohosa. After Davis (1917, Fig. 11). X1500.
Fig. 15. A fresh spore of Leptotkeca glomerosa. After Davis (1917, Fig. 13). X1500.
Fig. 16, A trophozoite of Leptotkeca sp. After Awerinzew (1908, PI. 2, Fig, 14). 1/12

and comp, oc, 12.
Fig. 17. Another trophozoite of the same. After Awerinzew (1908, PL 2, Fig. 17). 1/12

and comp. oc, 12,
Figs. 18 to 20. Ceratomyxa arcuata. After Th^lohan (1895).
Figs. 18 and 19. Typical young form. After Thflohan (1895, Figs. 16 and 17).
Fig. 20. A trophozoite with two spores. After Thelohan (1895, Fig. 18).

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Fig. 2L A sporulating trophozoite of Ceratomyxa arcuata. After Parisi (1912, Fig. 6a).
Fig. 22. A spore of Ceratomyxa arcuata treated with KOH. After Th6lohan (1895, Fig. 19).

X1500.
Figs. 23 and 24. Ceratomyxa sphaendosa. After Thdlohan.
Fig. 23. A part of the tropzoite (1895, Fig. 2). X750.
Fig. 24. A fresh spore (1895, Fig. 3). X750.

Fig. 25. A fresh spore of Ceratomyxa globulifera. After Th^lohan (1895, Fig. 43). X 1500.
Fig. 26. An adult trophozoite of Ceratomyxa appendiculata. After Thfilohan (1895, Fig. 4) .

X about 400.
Fig. 27. A spore of Ceratomyxa truncata. After Th^lohan (1895, Fig. 51). X1500.
Fig. 28. A spoK oi Ceratomyxa reticularis. After Th61ohan (1895, Fig. 27). X1500.
Fig. 29. A siporeoi Ceratomyxa inaequclis. After Doflein (1898, Fig. 10). X1250.
Figs. 30 and 31. Ceratomyxa linospora. After Doflem (1898). X about 1900.
Fig. 30. A spore (1898, Fig. 39).
Fig. 31. A trophozoite with spores (1898, Fig. 43).
Figs. 32 and 33. Ceratomyxa ramosa. After Awerinzew (1908).
Fig. 32. A trophozoite (1908, PI. 2, Fig. 20). Zeiss obj. D and comp. oc. 4.
Fig. 33. A spore (1908, PI. 2, Fig. 19). Zeiss obj. E and comp. oc. 4.

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Figs. 34 to 39. Ceratomyxa drepanopsettae. Awerinzew (1908).

Fig. 34 and 35. Trophozoites (1908, PL 2, Figs. 7 and 9). Obj. D and oc. 4.

Fig. 36. The part of a trophozoite attached to the epithelium of the gall-bladder of the host

1908, PL 2, Fig. 10). Obj. E and oc. 4.
Figs. 37 to 39. Spores (1908, PL 1, Figs. 2, 3 and 1). Obj. D and oc. 4.
Figs. 40 and 41. Two different views of the spore of Ceratomyxa tylosuri. After Awerinzew

(1913a, Fig. 1). X about 350. 4

Figs. 42 and 43. Ceratomyxa{?) spari. After Awerinzew (1913a, Fig. 2). *

Fig. 42. A trophozoite.
Fig. 43. A spore. X about 345.
Figs. 44 to 47. Spores of Ceratomyxa acadiensis. After Mavor (1916). Figs. 44 and 45

(1916, Fig. B). X270. Fig. 46 (1916, Fig. A) X1800. Fig. 47 (1916, Fig. 40) X2950.
Fig. 48. A spore of Ceratomyxa coris. After Georg6vitch (1916a, Fig. 1).
Fig. 49. A spore of Ceratomyxa herouardi. After Georg6vitch (1917, Fig. 1).
Fig. 50. A spore of Ceratomyxa mesospora. After Davis (1917, Fig. 15). X1500.
Fig. 51. A spore of Ceratomyxa sphairopkora. After Davis (1917, Fig. 23). X950.
Figs. 52 and 53. Spores oi Ceratomyxa taenia. After Davis (1917, Figs. 26 and 25). X700.

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218 ILLINOIS BIOLOGICAL MONOGRAPHS [456

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Fig. 54. A spore of Ceratomyxa aUenuata. After Davis (1917, Fig. 28). X950.

Figs. 55 and 56. Spores of Ceratomyxa recurvata. After Davis (1917, Figs. 32 and 33).

X1500.
Figs. 57 to 60. Spores of Ceratomyxa lunata. After Davis (1917, Figs. 34 to 37). X 1500.
Fig. 61. A spore oi Ceratomyxa abbreviata. After Davis (1917, Fig. 41). X1500.
Fig. 62. A spore of Ceratomyxa flagdlifera. After Davis (1917, Fig. 42). X1500.
Fig. 63. A spore oi Ceratomyxa agglomerata. After Davis (1917, Fig. 45). X1500.
Fig. 64. A spore oi Ceratomyxa amorpha. After Davis (1917, Fig. 47). X1500.
Figs. 65 to 67. Spores of Ceratomyxa monospora. After Davis (1917, Figs. 57, 56 and 55).

X1500.
Figs. 68 and 69. Spores of Ceratomyxa streptospora. After Davis (1917, Figs. 59 and 60).

X1500.
Fig. 70. A spore oi Ceratomyxa aggregata. After Davis (1917, Fig. 63). X1400,
Fig. 71. A spore of Ceratomyxa undulata. After Davis (1917, Fig, 66), X1500.
Figs. 72 and 73. Spores of Ceratomyxa navicularia. After Davis (1917, Figs. 69 and 68).

X1500.
Fig. 74. A spore oi Ceratomyxa spinosa. After Davis (1917, Fig. 72). X1500.

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Figs. 75 to 80. Myxoproteus anibiguus. After Doflem (1898).

Figs, 75 and 76. Trophozoites of typical forms (1898, Figs. 12 and 52).

Figs. 77 and 78. Young trophozoites produced by budding (1898, Figs. 55 and 56).

Figs. 79 and 80. Spores (1898, Figs. 54 and 64). X about 800 and 1080.

Figs. 81 to 83. Spores of Myxoproteus cordiformis. After Davis (1917, Figs. 79, 80 and 78).

X1500.
Fig. 84. ks^itoi Myxoproteus cornutus. After Davis (1917, Fig. 85). X1400.
Figs. 85 to 95. Wardia ovitwcua. Original.
Fig. 85. A portion of the cross-section thru an infected ovary of Lepomis humilis, showing

the parasite in one ovum and the hypertrophied nurse cells and several connective tissue

layers. X 160.
Fig. 86. A portion of the cross-section of a cyst. X640.
Figs. 87 to 89. Three diflEerent views of fresh spore. X2000.
Figs. 90 and 91. Stained spores. X1700.
Fig. 92. A spore mechanically pressed and stained with Giemsa's mixature, showing the

escaping polar capsules without extruding polar filament, and the sp>oroplasm. X1700.
Figs. 93 to 95. Front and lateral views of the shell valves, exhibiting the network-like fine

ridges on the surface and the posterior processes. X1700.
Figs. 96 and 97. Spores of Wardia ohlmacheri. After Ohlmacher (1893, Figs. 4a and 2).

2mm. and oc. 4.

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222 ILLINOIS BIOLOGICAL MONOGRAPHS (460

EXPLANATION OF PLATE
Figs. 98 to 104. Mitraspora cyprini.
Figs. 98 and 99. Trophozoites from the ureter of Cyprinus carpio in vivo. Original. X

about 1500.
Figs. 100 and 101. Different views of fresh spores. Original. X about 2000.
Figs. 102 to 104. Spores. After Fujita (1912, Figs, la to Ic).
Figs. 105 to 107. Mitraspora caudaia. After Parisi (1910 and 1913).
Fig. 105. A trophozoite (1910, Fig. 1).
Figs. 106 and 107. Front and lateral views of the spore (1913, Fig. 20 and 1910, Fig. 2).

X about 1600.
Figs. 108 to 113. Chloromyxum leydigi.

Figs. 108 and 109. Trophozoites. After Th^lohan (1895, Figs. 7 and 6). X750.
Figs. 110 and 111. Trophozoites in division. After Doflein (1898, Figs. 57 and 58).
Figs. 112 and 113. Different views of spores. After Th61ohan (1895, Figs. 10 and 9).

X1500.
Fig. 114. A spore of Chloromyxum catidatum. After Thdohan (1895, Fig. 36). X1500.
Figs. 115 and 116. Different views of the spore of Chloromyxum quadratum. After Th61ohan

(1895, Figs. 100a and 100b). X1500.
Fig. 117. A spore of Chloromyxum quadratum treated with nitric acid. After Th6Iohan

(1895, Fig. 100c). X1500.
Fig. 118. A spoie oi Chloromyxum fluviaiile. After Th61ohan (1892, Fig. 2). X1500.
Figs. 119 and 120. Different views of the spore of Chloromyxum mucronatum. After Lieber-

kiihn from Guriey (1894, PI. 39, Fig. 5). X about 1750.
Figs. 121 and 122. The same after Balbiani (1884, Fig. 41). X about 1200.
Figs. 123 and 125. Spores of Chloromyxum diploxys. After Balbiani from Guriey (1894

PI. 42, Figs. 13a, 13b and 13c).

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224 ILUNOJS BIOLOGICAL MONOGRAPHS (462

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Fig. 126, Trophozoite of Chloromyxum truUae. After L^ger (1906, Fig. 4). XlOOO.

Figs. 127 and 128. Trophozoites of Chloromyxum cristatum. After L6ger (1906, Fig. 7).
XlOOO.

Figs. 129 to 133. Chloromyxum dubium. After Auerbach (1908).

Figs. 129 and 130. Trophozoites (1908, Figs. 2 and 1).

Figs. 131 and 132. Spores (1908, Figs. 3 and 4). X about 2250.

Fig. 133. A stained young spore (1908, Fig. 5).

Fig. 134. Trophozoite of Chloromyxum sp. After Awerinzew (1908, PI. 2: Fig. 12). Objec-
tive D and ocular 4.

Fig. 135. Chloromyxum koi. After Fujita (1913). X about 800.

Figs. 136 to 138. Chloromyxum magnum. After Awerinzew (1913a, Fig. 4).

Figs. 136 and 137. Trophozoites.

Fig. 138. A spore. X about 320.

Figs. 139 and 140. Spores oi Chloromyxum funduli. After Hahn (1915, Figs. 34 and 33).
X2000.

Figs. 141 to 146. Chloromyxum misgurni. After Kudo (1916). X1750.

Figs. 141 and 142. Trophozoites (1916, Figs. 3f and 3g)

Figs. 143 to 145. Different views of fresh spore (1916, Figs. 3a, 3c and 3b).

Fig. 146. A spore treated with potassium hydrate (1916, Fig. 3e).

Figs. 147 to 152. Chloromyxum fujitai. After Kudo (1916). X1750.

Fig. 147. A trophozoite (1916, Fig. 4a),

Fig. 148. A fresh spore (1916, Fig. 4e).

Fig. 149. A spore stained with Giemsa's mixture (1916, Fig. 4g).

Figs. 150 to 152. Two surface views and an optical cross-section of a stained spore, showing
the ridges on the shell valves (1916, Figs. 4b, 4d and 4c),

Figs. 153 to 156. Spores of Chloromyxum clupeidae.

Figs. 153 to 155. Fresh spares. After Linton (1901, Fig. 3). "Variously magnified."

Fig. 156. A spore. After Hahn (1917b, Fig. 10). X1650.

Figs. 157 and 158. Two views of Chloromyxum granulosum. After Davis (1917, Figs. 137 and
138). X1500.

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Figs. 159 to 182. Chloromyxum trijugum. Original.

Figs. 159 to 174. Trophozoites of various form and age.

Figs. 159 and 160. Trophozoites of medium size. X640.

Fig. 161, A trophozoite, ten minutes after it was removed from the host. X1700.

Figs. 162 and 163. The movements of pseudopodia of the same specimen in ten minutes.
X1700.

Fig. 164. The same specimen after thirty minutes. X1700.

Figs. 165 to 167. A trophozoite, showing the change of pseudopodia in five and ten minutes.
X1700.

Figs. 168 to 172. Small trophozoite with different nimibers of the nuclei. Fig. 172 is prob-
ably a disporous form. X2350.

Fig. 173. A monosporous trophozoite with a young spore. X2350.

Fig. 174. A young spore. X2350.

Figs. 175 to 177. Different views of fresh spores. X1700.

Fig. 178. A Giemsa stained spore. X1700.

Figs 179 and 180. Side views of the valves showing the ridges by Giemsa staining. X1700.

Fig. 181. A spore treated with potassium hydrate solution, and stained with Giemsa solu-
tion. X1700.

Fig. 182, A spore from which the sporoplasm is leaving the shelL From the Giemsa smear
of the infected bUe. X 1700.

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Fig. 183 to 186. Sphaerospora divergens.

Fig. 183. Trophozoite. After TWlohan (1895, Fig. 12). X7S0.

Figs. 184 and 185. Two views of spore. After Auerbach (1912, PI. 5, Fig. 4). X about

1500.
Fig. 186. A spore treated with nitric acid. After Thflohan (1895, Fig. 13). X1500.
Figs. 187 and 88. Spores of Sphaerospora elegans. After Th61ohan (1890b, Fig. 1). X

about 1000.
Fig. 189. A spore of Sphaerospora rostrata. After Thdiohan (1895, Fig. 93). Xabout 1635.
Figs. 190 to 192. Spores of Sphaerospora masovica. After Cohn (1902, Fig. 3). X 1000.
Fig. 192. Spore with extruded polar filaments and "starren Ffiden" by warming.
Figs. 193 and 194. Spores of Sphaerospora platessae. After Woodcock (1904, Fig. 7d).

X900.
Figs. 195 to 197. Spores of Sphaerospora angulata. After Fujita (1912, Fig. 3). X about

2800.
Figs. 198 and 199. Spores of Sphaerospora polymorpha. After Davis (1917, Figs. 91 and 92)

X1500.
Figs. 200 to 204. Sphaerospora carassii. OriginaL
Fig. 200. A trophozoite. X2250.
Figs. 201 to 203. Different views of spores. XI 800.
Fig. 204. A young spore. X2250.

Figs. 205 to 209. Sinuolinea dimorpha. After Davis (1916).
Fig. 205. A fresh trophozoite (1916, Fig. 1). X1400.
Figs. 206 and 207. Trophozoites with erythrocytes in different stages of disintegratioa

(1916, Figs. 2 and 57). X640.
Fig. 208. A stained disporous trophozoite (1916, Fig. 41).
Fig. 209. A stained genmiule (1916, Fig, 72).

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230 ILUNOIS BIOLOGICAL MONOGRAPHS [46S

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Figs. 210 to 213. Sinuolinea dimorpha. After Davis (1916 and 1917).

Fig. 210. A living trophozoite (1916, Fig. 56). X640.

Fig. 211. Alivingtrophozoitefromwhichagenunuleis just escaping (1916, Fig. 60). X640L

Figs. 212 and 213. Spores (1917, Figs. 99 and 100). X1400.

Figs. 214 to 216. Spores of Sinuolinea capsularis. After Davis (1917, Figs. 105 to 107).

X1500.
Figs. 217 and 218. Spores of Sinuolinea arborescens. After Davis (1917, Figs. 109 to 110).

X1500.
Fig. 219. Spore oi Sinuolinea opacita. After Davis (1917, Fig. 112). X1500,
Fig. 220. Spore ot Sinuolinea brackiophora. After Davis (1917, Fig. 113). XlSOO.
Figs. 221 to 224. Myxidium lieberkukni. After Butschli (1881 and 1882).
Fig. 221. A trophozoite (1882, PI. ^, Fig. 12). X about 60.
Fig. 222. A trophozoite (1882, PI. 38, Fig. 13). Xl60.
Figs. 223 and 224. Trophozoites (1881, Figs. 27 and 26).

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Figs. 225 to 240. Myxidium lieherkUhni.

Fig. 225. A trophozoite. After LieberkOhn from Gurley (1894, PI. 43, Fig. la). X330.

Figs. 226 and 227. Trophozoites. After Btitschli (1881, Figs. 25 and 31).

Figs. 228 to 230. Stamed trophozoites. After Thflohan (1895, Figs. 44, 46 and 45). X7S0.

Fig. 231. A trophozoite forming daughter individuals by budding. After Cohn (1895

Fig. 5).
Fig. 232. Four figures showing the separation of a bud. After Cohn (1895, Figs. 6a, 6b,

6d and 6e).
Fig. 233. 1^ A cross-section of a trophozoite, showing the ectoplasm, mesoplasm and endo-

plasm. After Cohn (1895, Fig. 2).
Fig. 234. A trophozoite. After Laveran and Mesnil (1902, Fig. 3). X500.
Fig. 235. A part of a trophozoite. After Laveran and Mesnil (1902, Fig. 3), X800.
Figs. 236 and 237. Young trophozoites undergoing division. After Laveran and Mesnil

(1902, Figs. 4 and 5). X 1000.
Fig. 238. An isolated spore. After BUtschli (1881, Fig. 32). X about 2500.
Figs. 239 and 240, Fresh and stained spores. After Th6Iohan (1895, Figs. 47 and 48).

X1500.
Figs. 241 to 251. Myxidium incurvatum.

Fig. 241. A monosporous trophozoite. After Parisi (1912, Fig. 1).
Fig. 242. A spore. After Th61ohan (1895, Fig. 54). X1500.
Figs. 243 and 244. Different views of spore. After Parisi (1912, Fig. 1).
Fig. 245, A young spore. After Georgdvitch (1916, Fig. 11).
Figs. 246 to 248. Spores. After Georg6vitch (1916, Figs. 10, 9 and 8).
Figs. 249 to 251. Spores. After Davis (1917, Figs. 119 to 121). X1500.
Fig.*252. Trophozoite of Myxidium sphaericum. After Th61ohan (1895, Fig. 28). XlSOO.
Fig.*253. A spore of Myxidium histophUum. After Th6Iohan (1895, Fig. 49). X 1500.
Fig,^254. A spore of Myxidium sp. After Leydig from Gurley (1894, PI. 47, Fig. 6).

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234 ILLINOIS BIOLOGICAL MONOGRAPHS [472

EXPLANATION OF PLATE

Figs. 255 to 257. Myxddium danilewskyi. After Laveran (1898).

Figs. 255 and 256. Longitudinal and transverse sections thru renal tubules, showing the

trophozoites (1898, Figs. 1 and 2). Fig. 255. X350.
Fig. 257. Spores (1898, Figs. 4, 5 and 6). X800.

Fig. 258. Spores of Myxidium giganteum. After Doflein (1898, Fig. 48). X about 500.
Figs. 259 to 261. Spores of Myxidium giardi. After CepSde (1906a, Figs. 1, 3 and 2).

X2000.
Figs. 262 to 265. Spwres of Myxidium pfeifferi. After Auerbach (1908, Figs. 6 and 7).

X about 2000, except Fig. 265.
Fig. 266. A spore of Myxidium inflatum. After Auerbach (1909, Fig. 3a). X about 1500.
Fig. 267. A spore of Myxidium bergense. After Auerbach (1910a, Fig. 57). X about 1820.
Fig. 268. A spore of Myxidium procerum. After Auerbach (1910a, Fig. 58). X about 2000.
Figs. 269 to 271. Spores of Myxidium mackiei. After Bosanquet (1910, Fig. 12). X

about 1250.
Figs. 272 and 273. Spores of Myxidium macrocapsidare. After Auerbach (1910d, Figs, la

and lb). X3000.
Figs. 274 to 276. Myxidium sp. After Awerinzew (1908 and 1911).
Fig. 274. A monosporous trophozoite (1911, Fig. C).
Fig. 275. A disprous trophozoite (1908, PI. 2, Fig. 6). Obj. E and oc. 4.
Fig. 276. A spore (1908, PI. la. Fig. 17). X about 1000.
Figs. 277 and 278. Spores of Myxidium depressum. After Parisi (1912, Figs. 2a and 2b).

X about 1600.
Figs. 279 and 280. Spores of Myxidium oviforme. After Parisi (1912, Fig. 3). X about

1600.
Figs. 281 to 284. Spores of Myxidium anguiUae. After Ishii (1915, Fig. 3a). X1450,
Figs. 285 and 286. Myxidium sp. After Mavor (1915).
Fig. 285. A spore treated with ammonia water (1915, Fig. 3a). X660.
Fig. 286. A spore (1915, Fig. 3b). X1320.

Figs. 287 to 290. Spores of Myxidium gadi. After Georg6vitch (1916, Figs. 1, 4, 3 and 2).
Fig. 228. A spore from Solea vulgaris.
Figs. 289 and 290. Young spores.

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236 ILLINOIS BIOLOGICAL MONOGRAPHS [474

EXPLANATION OF PLATE

Fig. 291. A spore of Myxidium glutinosum. After Davis (1917, Fig. 124). X1400.

Figs. 292 and 293. Spores of Myxidium phyllium. After Davis (1917, Figs. 126 and 127).

X2000.
Figs. 294 and 295. Spores of Myxidium kagayatnai. After Kudo (1916, Fig. 2). X1750

and X 1000 respectively.
Figs. 296 to 307. Sphaeromyxa balbianii.
Fig. 296. A trophozoite. After Thdohan (1895, Fig. 57). X3.
Fig. 297. A trophozoite. After Davis (1917, Fig. 128). X640.
Fig. 298. A trophozoite in plasmotomy. After Georg6vitch (1916, Fig. 15).
Figs. 299 and 300. Spores. After Th61ohan (1895, Figs. 58 and 59). X1500.
Fig. 301. An end of a spore. After Thflohan (1895, Fig. 60).
Fig. 302. A spore treated with nitric acid. After Thflohan (1895, Fig. 61).
Fig. 303. A spore. After Parisi (1912, Fig. 4). X about 1750.
Fig. 304. A spore. After Davis (1917, Fig. 130). X2100.
Figs. 305 to 307. Mature and young spores (Figs. 306 and 307). After Georg6vitch (1916,

Figs. 17, 20 and 19).
Figs. 308 to 311. Sphaeromyxa immersa. After Lutz (1889: 301).
Fig. 308. An infected gall-bladder of Bufo aqua (1889, Fig. 1). XL
Fig, 309. Spores (1889, Figs. 4, 5 and 6).
Fig. 310. A spore (1889, Fig. 10). X600.
Fig. 311. A spore with extruded polar filaments (1889, Fig. 7).
Figs. 312 to 314. Spores of Sphaeromyxa incurvata. After Doflein (1898, Fig. 49). X

about 1000.
Figs. 315 and 316. Sphaeromyxa sabrazesi. After Schroder (1907).
Fig. 315. A trophozoite (1907, Fig. 1). X15.
Fig. 316. A cross section thru a trophozoite (1907, Fig. 3). XlSOO.

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238 JLUNOIS BIOLOGICAL MONOGRAPHS [476

EXPLANATION OF PLATE

Figs. 317 to 322. Spores of Sphaeromyxa sabrazesi.

Figs. 317 to 319. After Laveran and Mesnil (1900, Figs. 1, 3 and 4). XlSOO.

Fig. 318. A spore treated with nitric acid (1900, Fig. 3).

Figs. 320 and 321. Spores. After Schroder (1907, Figs. 39 and 41). X1500.

Fig. 322. A polar capsule. After Schroder (1907, Fig. 45).

Figs. 323 and 324. Spores of Sphaeromyxa hellandi. After Auerbach (1909, Fig. 5). X

about 1500.
Figs. 325 and 326. Spores of Sphaeromyxa exneri. After Awerinzew (1913a, Fig. 3). X

about 365.
Fig. 327. A spore of Sphaeromyxa gasterostei. After Georg6vitch (1916, Fig. 22).
Figs. 328 to 331. Zschokkella hildae. After Auerbach (1910a and 1912).
Fig. 328. A monosporous trophozoite (1912, PL 5, Fig. 2).
Figs. 329 to 331. Spores (1910a, Fig. 62).
Figs. 332 and 333. Spores of Zschokkella ruma. After Klokacewa (1914, Fig. 2). X about

2500.
Figs. 334 to 338. Spores of Zschokkella acheilognathi. After Kudo (1916).
Figs. 334 to 336. Different views of fresh spores (1916, Figs. 3d, 3e and 3f). X2250.
Fig. 337. A young spore. Original. X2785.
Fig. 338. A stained spore (1916, Fig. 3h). X2800.
Figs. 339 and 340. Spores of Zschokkella globulosa. After Davis (1917, Figs. 135 and 134).

X1500.
Figs. 341 to 343. Spores of Myxosoma dujardini. After Th6Iohan (1895, Figs. 90, 91, and

89). X1500.
Figs. 344 to 347. Spores of Myxosoma funduli. After Kudo (1918a, Fig. A). X1500.

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210 ILUNOJS BIOLOGICAL MONOGRAPHS [478

EXPLANATION OF PLATE

Fig. 348. A spore of Myxosomai?) lobatum. After Nemeczek (1911, Fig. 18). XlOOO.

Fig. 349 to 354. Lentospora cerebralis. After Plehn (1905). X1200.

Fig. 349. Various young ameboid forms; stained (1905, Textfig. 5),

Fig. 350. A stained larger form (1905, Textfig. 5).

Fig. 351. A trophozoite with two spores (1905, Textfig. 4).

Fig. 352. Various spores (1905, Textfig. 2).

Fig. 353. A stained spore (1905, Textfig. 3).

Fig. 354. A spore with extruded polar filaments (1905, Textfig. 2).

Fig. 355. A spore oi Lentospora multiplicata. After Reuss (1906, Fig. 8). X1500.

Figs. 356 to 359. Spores of Lentospora asymmetrica. After Parisi (1912, Fig. 7). X about

1500.
Figs. 360 to 362. Spores of Lentospora acuta. After Fujita (1912, Fig, 2). X2000.
Figs. 363 and 364. Spores of Myxobolus piriformis. After Thdohan (1895, Figs. 117 and

116). X1500.
Figs. 365 and 366. Spores of Myxobolus unicapsulaius. After Miiller (1841, Fig. 5).
Fig. 367. A spore of Myxobolus fukrmanni. After Auerbach (1909, Fig. lb). X1840.
Fig. 368. A spore of Myxobolus oculi4eucisci. After Trojan (1909, Fig. 3). X2000,
Figs. 369 and 370. Spores of Myxobolus toyatnai. After Kudo (Original and 1917, Fig. 40).

X25O0.
Figs. 371 and 372. Spores of Myxobolus notatus. After Mavor (1916, Figs. 6C and 6B).

X2600.
Figs. 373 and 374. Spores of Myxobolus rohitae. After Southwell and Prashad (1918,

Figs. 26 and 27). X about 1720 and 700 respectively.
Figs. 375 and 376. Spores of Myxobolus sent. After Southwell and Prashad (1918, Figs.

29 and 30). X about 1700.
Figs. 377 and 378. Sports oi Myxobolus misgurni. Original. X1500.

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Figs. 379 to 385. Myxobolus pfeifferi.

Figs. 379 and 380. Parts of section thru cyst. After Keysselitz (1908a, PI. 15, Figs. 1 and 2) .

Fig. 381. A spore. After TMlohan (1895, Fig. 77). X1500.

Fig. 382. An optical section of spore. After Keysselitz (1908a, Fig. A).

Fig. 383. A spore treated with Lugol's solution. After Keysselitz (1908a, PI. 14: Fig. 92).

Fig. 384. A spore with extruded filaments. After Keysselitz (1908a, Fig. C).

Fig. 385. A stained young spore. After Keysselitz (1908a, PI. 14, Fig. 81).

Fig. 386. A spore of Myxobolus dispar. After Thdlohan (1895, Fig. 86). X about 1500.

Figs. 387 to 389. Spores of Myxobolus ellipsoides. Fig. 389. An abnormal spore with six

polar capsules. After Thelohan (1895, Figs. 112, 113 and 115).
Fig. 390. A part of the infected intestine of Mugil auratus, showing cysts of Myxobolus

exiguus. After Parisi (1912, Fig. 9e). X about 3.
Fig. 391. A spore of Myxobolus exiguus. After Thelohan (1895, Fig. 98). X 1500.
Figs. 392 to 395. Spores of Myxobolus exiguus. After Parisi (1912, Fig. 9). X2500.
Fig. 396. A spore of Myxobolus oviformis. After Th61ohan (1895, Fig. 81). X1500.
Figs. 397 to 398. Spores of Myxobolus miiUeri. After Thdohan (1895, Figs. 96 and 97).

X1500.
Figs. 399 and 400. Spores of Myxobolus millleri. After Biitschli (1881, Figs. 1 and 2).
Fig. 401. A spore of Myxobolus millleri in cone, sulphuric acid. After Biitschli (1881, Fig. 6).
Figs. 402 and 403. Abnormal spores of Myxobolus millleri. After Stitschli (1881, Figs. 9

and 8).
Figs. 404 and 405. Spores of Myxobolus lintoni. After Linton (1891, Figs. 3 and 5).
Fig. 406. Diagram of the cross section of a spore of Myxobolus lintoni. After Linton (1891,

Fig. 8).
Fig. 407. A spore of Myxobolus linloni with extruded polar filaments. After Linton (1891,

Fig. 10).
Fig. 408. A stained spore of Myxobolus lintoni. After Gurley (1894, PI. 26, Fig. 7). X

about 2000. •

Figs. 409 and 410. Spores of Myxobolus globosus. After Gurley (1894, PI. 26, Fig. 7).

X about 2900.
Fig. 411. Sp>ores of Myxobolus inaequalis. After Miiller (1841, Fig. 6).

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Figs. 412 to 416. Spores of Myxobolus oblongus. Figs. 412 to 414. After Gurley (1894,
PI. 26, Fig. 6). X2300. Figs. 415 and 416. After MiiUer (1841, Fig. 9).

Figs. 417 and 418. Spores of Myxobolus transovdis. After Gurley (1894, PI. 29, Fig. 1).

Figs. 419 and 420. Spores of Myxobolus obesus. After Balbiani (1884, Fig. 39).

Fig. 421. Spores of Myxobolus cycloides. After Miiller (1841, Fig. 3).

Figs. 422 and 423. Spor^ oi Myxobolus anurus. After Cohn (1895, Fig. 25). X1500.

Fig. 424. Spores of Myxobolus sp. X700. After Butschli (1882, PL 36, Fig. 23).

Fig. 425. Myxobolus sp. After Gurley (1894, PL 28: Fig. 4a). X about 1500.

Figs. 426 to 429. Spores of Myxobolus sp. After Miiller (1841, Fig. 4).

Fig. 430. A vegetative form of Myxobolus cyprini. After Doflein (1898, Fig. 112).

Figs. 431 and 432. Spores of Myxobolus cyprini. After Doflein (1898, Figs. 113 to 115).

Figs. 433 to 436. Myxobolus neurobius. After Schuberg and Schroder (1905).

Figs. 433 and 434. Longitudinal and transverse sections thru infected nerve fibres (1905,
Figs. 2 and 4a). X520.

Figs. 435 and 436. Sp>ores (1905, Figs. 5 and 6). Comp. oc. 12 and imm. obj. 2mm.

Figs. 437 to 441. Myxobolus aeglefini. After Auerbach (1906a).

Fig. 437. A cyst in the sclerotic cartilage of the eye of Gadus aeglefini (1906a, Fig. 2).

Figs. 438 and 439. Spores (1906a, Figs. 5a and 3d). Xabout 1320.

Figs. 440 and 441. Abnormal spores (1906a, Figs. 5b and 5c). Xabout 1320.

Figs. 442 to 445. Myxobolus gigas. After Auerbach (1906b).

Fig. 442. A part of the section of a cyst (1906b, Fig. 1).

Figs. 443 to 445. Spores (1906b, Figs. 3a, 3c, 5 and 3b). X about 850.

Figs. 446 and 447. Spores of Myxobolus volgensis. After Reuss (1906, Fig. 1). X2000.

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246 ILLINOIS BIOLOGICAL MONOGRAPHS [484

EXPLANATION OF PLATE

Fig. 448. The branchia of Lucioperca volgensis with cysts of Myxobolus volgensis. After

Reuss (1906, Fig. 2). X2.25.
Fig. 449. A spore of Myxobolus scardinii. After Reuss (1906, Fig. 3). X1500.
Fig. 450. Air bladder of Scardinius erythrophtkalmus with the cysts of Myxobolus pkyso-

philus. After Reuss (1906, Fig. 5). X2.
Fig. 451. A spore of Myxobolus physophilus. After Reuss (1906, Fig. 4). X1500.
Fig. 452. A sport of Myxobolus macrocapsular is. After Reuss (1906, Fig. 6). X1500.
Fig. 453. A spore of Myxobolus sandrae. After Reuss (1906, Fig. 7). X2000.
Fig. 454. A spoK ol Myxobolus bramae. After Reuss (1906, Fig. 9). X1500.
Fig. 455. A spore ot Myxobolus balleri. After Reuss (1906, Fig. 10). X1500.
Fig. 456. A spore of Myxobolus cyprinicola. After Reuss (1906, Fig. 11). X1500.
Figs. 457-459. Myxobolus squamae. After Keysselitz (1908a).
Fig. 457. A part of the infected scale (1908a, Fig. G.)
Fig. 458. A spore treated with Lugol's solution (1908a, PI. 14, Fig. 94).
Fig. 459. A stained spore (1908a, PL 14, Fig. 96).

Figs, 460 and 461'. Spores of Myxobolus cordis. After Keysselitz (1908a).
Fig. 460. A spore treated with Lugol's solution (1908a, PI. 16, Fig. 16).
Fig. 461. A strained spore (1908a, Fig. B on page 281).
Figs. 462 to 464. Spores of Myxobolus tnusculi. After Keysselitz (1908a).
Fig. 462. A spore treated with Lugol's solution (1908a, PI. 15, Fig, 13).
Fig. 463 and 464. Stained spores (1908a, Figs. D and E on page 286).
Fig. 465. Sports oi Myxobolus sp. After Wegener (1910, Fig. 44). X1050.
Fig. 466. A sport oi Myxobolus per magnus. After Wegener (1910, Fig. 45). X1050.
Fig. 467. Spores of Myxobolus rotundus. After Nemeczek (1911, Figs. 10 and 11). XlOOO.
Fig. 468. Spores of Myxobolus minutus. After Nemeczek (1911, Figs. 16 and 17). XlOOO.
Figs. 469 and 470. Spores of Myxobolus magnus. After Awerinzew (1913, 76). X about

340.
Figs. 471 to 473. Spores of Myxobolus carassii. After Klokacewa (1914, Fig. 1). X about

2400.

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EXPLANATION OF PLATE

Figs. 474 to 476, Myxobolus funduli. After Hahn (1915 and 1917).

Fig. 474. A cyst from the gill filament (1917, Fig. 1).

Fig. 475. A stained spore (1915, Fig. 28). X2000.

Fig. 476. Diagram of the cross section of a spore (1915, Fig. 30).

Fig. 477. A spore of Myxobolus pleuronectidae. After Hahn (1917a, Fig. 2). X157S.

Fig. 478. A spore of Myxobolus capsulatus. After Davis (1917, Fig. 139). X1500.

Figs. 479 and 480. Spores of Myxobolus nodularis. After Southwell and Prashad (1918,
PI. 11, Figs. 34 and 35). X about 1450.

Fig. 481. A sf)ore of Myxobolus miyairii. After Miyairi (1909, Fig. 14).

Figs. 482 to 485. Spores of Myxobolus koi. Original. X 1300.

Figs. 482 to 484. Different views.

Fig. 485. A spore stained with Giemsa's mixture.

Figs. 486 and 487. Henneguya psorospermica. After Th€lohan (1895).

Fig. 486. A cross section thru branchial lamella of Esox lucius with a cyst (1895, Fig. 82).

Fig. 487. Two spores (1895, Figs. 83 and 84). X about 1000.

Figs. 488 and 489. Henneguya minuta. After Cohn (1895).

Fig. 488. A longitudinal section of an infected branchial lamella (1895, Fig. 29.)

Fig. 489. Two spores. One with two vacuoIes(?) (1895, Fig. 30). X about 450.

Fig. 490 and 491. Spores of Henneguya oviperda. After Cohn (1895, Fig. 31).

Fig. 491. A spore with extruded "starren Faden" and polar filaments.

Figs. 492 and 493. Henneguya lobosa. After Cohn.

Fig. 492. An external view of the parasite on the gill (1895, Fig. 18).

Fig. 493. Two spores and one unseparated young spores (1895, Fig. 21).

Figs. 494 and 495. Henneguya media. After TMlohan (1890b).

Fig. 494. A sporoblast in the ovary of Gasterosteus, with one spore (1890b, Fig. 18).

Fig. 495. Spores (1890b, Fig. 1).

Fig. 496. The peripheral portion of a section of a cyst of Henneguya psorospermica, show-
ing the characteristic structure. After Th^lohan (1895 : 237).

Figs. 497 to 499. Spores of Henneguya schizura. After Muller (1841, Fig. 1).

Figs. 500 to 503. Spores of Henneguya creplini. After Creplin (1842, Figs. B, E, A and C).

Fig. 504. Spores of Henneguya linearis. After Miiller (1841, Fig. 10).

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Fig. 505. Spores of Henneguya Gurleyi. After Gurley (1894, PI. ii, Figs. 8c, 6 and 7).

X about 3100.
Fig. 506. Spores of Henneguya strongylura. After Muller (1841, Fig. 2).
Fig. 507. Spores of Henneguya tnonura. After Ryder (1880, Figs, Ic and 2d).
Fig. 508. Sp>ores of Henneguya kolesnikovi. After Kolesnikov from Gurley (1894, PL 35,

Fig. 7).
Figs. 509 to 512, Henneguya macrura. After Gurley (1894). X about 2100.
Figs. 509 and 510. Spores (1894, PI. 32, Fig. 5, PI. 33, Fig. 1).

Fig. 511. A spore treated with iodine, showing the "beading of the tail" (1894, PI. 33, Fig. 3).
Fig. 512. A tail separated from the main part by iodine (1894, PI. ii, Fig. 4).
Fig. 513. Spores of Henneguya zschokkei. After Zschokke (1898, Figs. 2 and 1).
Fig. 514. Spores of Henneguya sp. After Benecke from Gurley (1894, PI. 29, Fig. 8).
Fig. 515. Spore of Henneguya tenuis. After Vaney and Conte (1901, Fig. 2).
Figs. 516 and 517. Spores of Henneguya nusslini. After Schuberg and SchrSder (1905,

Figs. 13 and 14), Comp. oc. 12 and obj. 2mm.
Figs. 518 to 523. Henneguya legeri. After C6pMe (1913).
Figs. 518 to 521. Trophozoites (1913, Figs. 2, 23, 15 and 25). X900.
Fig. 519. A trophozoite in division (1913, Fig. 23), X900.
Fig. 521. A trophozoite stained with iron hematoxylin (1913, Fig. 25). X900.
Fig. 522. An elongated spore (1913, Fig. 26). X900,
Fig, 523, An ovoidal spore (1913, Fig. 24), X450.
Fig. 524. A spore of Henneguya miyairii. After Miyairi (1909, Fig, 11).

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252 ILLINOIS BIOLOGICAL MONOGRAPHS [490

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Figs. 525 and 526. Spores of Eenneguya acerinae. After SchrSder (1906, Figs. 5 and 6).

X1650.
Fi^. 527 to 535. Eenneguya giganUa. After Nemeczek (1911). XlOOO.
Fig. 527. A mature spore with extruded polar filaments (1911, Fig. 1).
Figs. 528 to 535. Stages in development of spores (1911, Figs. 2 to 9).
Figs. 536 to 539. Spores of Henneguya(?) sp. After Nemeczek (1911, Figs. 12 to 15).

XlOOO.
Figs. 540 to 543. Eenneguya gasterostet. After Parisi (1912). X about 1500.
Fig. 540. A disporous trophozoite (1912, Fig. lOd).
Figs. 541 and 542. Two spores (1912, Figs. lOf and lOe).
Fig. 543. A young sp>ore (1912, Fig. 10c).
Figs. 544 and 545. Spores of Eenneguya neapolilana. After Parisi (1912, Fig. 11). X

about 1500.
Figs. 546 to 549. Various trophozoites of Eenneguya mictospora. OriginaL
Fig. 546. A monosporous trophozoite with a yoimg spore. X950.
Figs. 547 and 549. Trophozoites in vivo. X650.
Fig. 548. A stained binucleated yoimg trophozoite. X1700.

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254 ILLINOIS BIOLOGICAL MONOGRAPHS [492

EXPLANATION OF PLATE

Figs. 550 to 557. Henneguya mictospora. Original.

Fig. 550. A stained trophozoite. X1700.

Figs. 551 to 553. Three different stages of development of disporous trophozoites. Giemsa.

X1700.
Figs. 554 and 555. Two stages of monosporous trophozoites. Giemsa. X1700.
Figs. 556 and 557. Different views of mature spores in vivo. X2000.
Figs. 558 and 559. Henneguya wisconsinensis. After Mavor and Strasser (1916).
Fig, 558. A trophozoite in vivo (1916, Fig. la). X570.
Fig. 559. A fresh spore (1916, Fig. 3d). X4000.

Figs 560 and 561. Trophozoites oi Chloromyieum catostomi. Original. X1500.
Figs. 562 to 565. Chlorotnyxum clupeidae. Original drawn from Dr. Tyzzer's smears which

were restained. X2360.
Fig. 562. Anterior end view of two spores in preserved and decolorized smears.
Fig. 563. The same views of three spores restained with Giemsa mixture.
Fig. 564. Front view of a preserved and decolorized spore.
Fig. 565. The same views of two spores restained with Giemsa's mixture.
Figs. 566 to 572. Spores of Myxobolus orbiculatus. Original.

Figs. 566, 569 and 570. Different views of normal spores in preserved specimen. X 1500.
Figs. 567 and 568. Abnormal spores. X1500.
Fig. 571. A spore stained with Lugol's solution. X1500.
Kg. 572. A spore stained with Giemsa's mixture. X2360.

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EXPLANATION OF PLATE

Figs. 573 to 576. Myxobolus orbiculatus. Original

Fig. 573. Young vegetative forms in the muscle fibres. From a section stained with Heid-

enhain's iron hematoxylin. X900.
Figs. 574 and 575. Two vegetative forms under higher magnifications. X1500.
Fig. 576. A cross section thru the infected muscle fibres, stained with Heidenhain's iron

hematoxylin. X900.
Figs. 577 to 581. Hoferellus cyprini. After Doflein (1898).
Figs. 577 and 578. Two vegetative forms (1898, Figs. 106 and 105).
Figs. 579 to 581. Spores (1898, Figs. 108 and 107).
Figs. 582 and 583. Genus incert. merlucii. After Perugia from Gurley (1894, PI. 29,

Figs. 2 and 7).
Figs. 584 and 585. Genus incert. congri. After Perugia from Gurley (1894, PL 6, Figs. 7

and 3).
Figs. 586 and 587. Genus et species incertae. After Mavor (1915).
Fig. 586. A trophozoite with attached Ceratomyxa acadiensis (1915, Fig. 8). X830.
Fig. 587. A trophozoite (1915, Fig. 6).
Figs. 588 and 589. Trophozoites of genus et species incertae. After Mavor (1916a, Figs.

3d and 3b). X660.
Fig. 586. A trophozoite with attached Ceratomyxa acadiensis (1915, Fig. 8). X830.
Fig. 587. A trophozoite (1915, Fig. 6).
Figs. 588 and 589. Trophozoites of genus et species incertae. After Mavor (1916a, Figs.

3d and 3b). X660.
Fig. 590. SjK)res of genus et species incertae. After Linton (1891a, Fig. 2).
Figs. 591 to 593. Myocobolus hylae. After Johnston and Bancroft (1918).
Fig. 591. A transverse section of a heavily infected testis of Hyla aurea, (1918, Rg. 1).

X about 11.
Fig. 592. Different views of normal spores, stained (1918, Fig. 3). X about 800.
Fig. 593. Abnormal spores (1918, Fig. 4). X about 800.
Figs. 594 to 596. Lentospora dermaiobia. After Ishii (1915).
Fig. 594. A part of the infected skin of the host (1915, Fig. 2). X140.
Figs. 595 and 596. Different views of spore (1915, Figs. 4 and 3). X1450.
Figs. 597 to 601. Myxobolus discrepans. Original.
Fig. 597. An infected branchial lamella showing the cysts of various size and form. X

about 4.
Figs. 598 to 601. Unstained preserved spores, showing different views. X about 1500.

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4951 STUDIES ON MYXOSPORIDIA—KUDO 257

PLATE XXIV

258 ILLINOIS BIOLOGICAL MONOGRAPHS [496

EXPLANATION OF PLATE

Figs. 602 to 621. MUraspora elongate. Original. X 1500, except Figs. 620 and 621, X2500.
Giemsa staining, unless otherwise stated.

Fig. 602. A trophozoite showing various nuclei and sporoblasts at different stages of devel-
opment.

Fig. 603. A trophozoite with a mature and a young spore. Iron hematoxylin.

Fig. 604. A trophozoite with two mature spores.

Figs. 605 to 609. Formation and development of sporoblasts.

Figs. 610 to 613. Developing spores. Fig. 612; Delafield's hematoxylin.

Fig. 614. A surface view of a preserved spore.

Fig. 615. An optical section of a preserved spore.

Fig. 616. Front view of a stained spore.

Fig. 617. Lateral view of a spore stained with Heidenhain's iron hematoxylin.

Fig. 618. A slightly elongated spore.

Fig. 619. An abnormal spore.

Fig. 620. A longitudinal section thru a polar capsule.

Fig. 621. An oblique view of a polar capsule, showing the spirally coiled polar filament.

Figs. 622 to 627. Myxidium americanum. Original. X1500.

Figs. 622 and 623. Two young trophozoites. Giemsa.

Fig. 624. A sporulating trophozoite in imstained preserved state.

Fig. 625. A spore in preserved state.

Fig. 626. A fresh spore treated with potassium hydrate solution.

Fig. 627. A spore stained with Giemsa's mixture.

Figs. 628 to 631. Myxobolus mesentericus. Original. X1500.

Figs. 628 and 629. Front and lateral views of unstained preserved spores.

Fig. 630. A Giemsa stained spore.

Fig. 631. A spore with extruded polar filament (potassium hydrate), stained with Giemsa's
mixture.

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200 ILUNOIS BIOLOGICAL MONOGRAPHS [498

EXPLANATION OF PLATE

Figs. 632 to 642. CUoromyxum wardi. OriginaL

Fig. 632. A 3roung trophozoite. Smear and Giemsa, XlSOO.

Fig. 633. A sporulating trophozoite. Giemsa. X 1500.

Figs. 634 to 637. Surface views and optical sections of four unstained spores, showing the

sutural ridge and the fine striations on the shelL X 1500.
Fig. 638. A polar view of unstained spore, X2360.
Figs. 640. Surface view and optical section of a single spore. X2360.
Fig. 641. A front view of an unstained spore. X2360.
Fig. 642. A Giemsa stained spore. X1500.
Figs. 643 to 649. Myxoholus aureatus. After Ward (1919).
Fig. 643. A fresh spore (1919, Fig. Aa).

Figs. 644 and 645. Fresh spores kept for 24 hours or more in water (1919, Fig. Aa).
Fig. 646. A preserved unstained spore (1919, Fig. Ba).

Fig. 647. A spore stained with Delafield's hematoxylin (1919, Fig. Bd). X1500.
Fig. 648. A spore with an extruded polar filament from section preparation stained with

Giemsa's mixture (1919, Fig. Bg). X1500.
Fig. 649. A yoimg spore, stained with Giemsa's mixture (1919, Fig. Bi). X1500.
Figs. 650 to 653. Henneguya brachyura. After Ward (1919). X1500.
Fig. 650. A young spore stained with Giemsa's mixture (1919, Fig. Ce).
Figs. 651 and 652. Front and lateral views of spore (1919: F^. Cb and Cc).
Fig. 653. A detached taU (1919, Fig. Cf).

Figs. 654 to 656. Henneguya salminicola. After Ward (1919). X1500.
Fig. 654. A young spore stained with Giemsa's mixture (1919, Fig. Fc).
Figs. 655 and 656. Front and lateral views of unstained preserved spores.

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INDEX

African Myxosporidia, 14

Air bladder, Mj^osporidia parasitic in, 39

Alaskan Myxosporidia, 16

Alimentary canal, Myxosporidia parasitic in,

39,41
Amphibian hosts of Myxosporidia, 36
Annelida as host of Mjrxosporidian, 25
Anterior end of spore, 47, 50

process of spore, 47, 50, 51
Asiatic Myxosporidia, 13
Australian Myxosporidian, 14
Austrian Myxosporidia, 24
Blood vessel, Mj^tosporidia parasitic in, 39
Body cavity, Myxosporidia parasitic in, 37,

43,45
Bone, Myxosporidia parasitic in, 39
Branchia, Myxosporidia parasitic in, 38
Breadth of spore, 50
Burma, Myxosporidia of, 14
Canadian M5Tcosporidia, 16
Capsulogenous cell of spore, 50
Cartilage, Myxosporidia parasitic in, 39
Ceratomyxa, 9, 56, 65, 178, 180
Ceratomyxa abbreviata, 76

acadiensis, 71

agglomerata, 77

aggregata, 79

amorpha, 78

appendiculata, 67

arcuata, 65

attenuaia, 75

coris, 72

drepanopsettae, 70

flagellifera, 77

globurifera, 67

herouardl, 72

inaequdis, 68

Unospora, 69

lunata, 76

mesospora, 73

tnonospora, 78

navicularia, 80

pallida, 67

ramosa, 69

recurvata, 75

reticularis, 68

spari, 70

sp. (?) Awerinzew, 71

sp. (?) Awerinzew, 71
sp. Georg^vitch, 72
sphaerulosa, 66
sphairophora, 73
spinosa, 80
streptospora, 79
taenia, 74
truncata, 67
tylosuri, ^0
undtdata, 79
Ceratom)^dae, 9, 56, 60, 178
Chloromyxidae, 10, 57, 87, 178
Chloromyxum, 10, 57, 87, 178, 183
CMoromyxum catostomi, 98
cattdatutn, 88
dupeidae, 94
cristatum, 91
diploxys, 90
dubium, 91
fluviatile, 89
fujitai, 93
funduli, 93
granulosutn, 96
yfeoi, 92
leydigi, 87
magnum, 92
misgurni, 93
tnucronaium, 89
^ro<e«, 90
quadratum, 88
sp. Awerinzew, 91
thymalli, 92
trijugum, 96
truttae, 90
wardi, 99
Classification of Myxosporidia after
Auerbach, 52
Davis, 54
Doflein, 52
Kudo, 56
Parisi, 54
Poche, 54
Thelohan, 52
Connettive tissue, Myxosporidia parasitic in,

37
Cyst, 50
Disporea, 52, 54
Disporous, 50

262

ILUNOIS BIOLOGICAL MONOGRAPHS

[500

England, Myxosporidia of, 22
European, M)aosporidia, 17
Eurysporea, 9, 56, 60
Eye, Myxosporidia parasitic in, 38
Fish as hosts of Myxosporidia, 25
fresh water, 44
marine, 44
Foramen of polar capsule, 47, 48, 50
France, M3^osporidia of, 18
Front view of spore, 47, 50
Gall bladder, Myxosporidia parasitic in, 39
Gemmule, 50
German MjTsosporidia, 20
Heart, Myxosporidia parasitic in, 39
Henneguya, 11, 59, 158, 179, 193
Henneguya acerinae, 167

brachyura, 171

brevis, 162

creplini, 162

gasterostei, 169

gigantea, 168

gurleyi, 163

kolesnikovi, 164

legeri, 166

linearis, 163

lobosa, 161

macrura, 164

media, 161

mictospora, 173

minuta, 160

miyairii, 172

monura, 164

neapolitana, 170

nilsslini, 166

oviperda, 160

peri-intestinalis, 161

psoros per mica, 158

salminicola, 171

schizura, 162

sp. (Gurley) Labb6, 165

sp. (Gurley) Labb6, 165

sp. (?) Nemeczek, 169

strongylura, 163

tenuis, 166

texta, 159

wisconsinensis, 170

zschokkei, 165
HofereUus, 12, 59, 173, 179
Hoferettus cyprini, 174
Indian Myxosporidia, 14
Insect as host of Myxosporidia, 25
Int^ument, Myxosporidia parasitic in, 37

Intercapsular appendix of spore, 47
Intestine, Mj^osporidia parasitic in, 41
lodinophilous vacuole, 47, 50
Italian Myxosporidia, 17
Kamtschatka, Myxosporidian of, 14
Klidney, Myxosporidia parasitic in, 42
Lateral process of spore, 50
Length of spore, 50
Lentospora, 11, 58, 125, 179, 189
Lentospora acuta, 127

asymmetrica, 126

cerebralis, 125

dermatobia, 127

encephalina, 126

multiplicata, 126
Leptotheca, 9, 56, 60, 178, 179
Leptotheca agUis, 60

dongaia, 60

fusiformis, 63

glomerosa, 65

hepseti, 62

informis, 63

lobosa, 64

longipes, 63

macrospora, 62

parva, 61

perlata, 62

polymorpha, 61

renicola, 61

scissura, 64

sp. Awerinzew, 62
Liver, Myxosporidia parasitic in, 39
Longitudinal striation on spore, 50
Mesentery, Myxosporidia parasitic in, 43
Mesoplasm, 50
Mictosporea, 52, 54
Mictosporous, 50
Mitraspora, 9, 56, 84, 178, 183
Mitraspora caudata, 85
cyprini, 84
elongata, 85
Monaco, Myxosporidia of, 17
Monosporous, 50

Muscle, Myxosporidia parasitic in, 38
Myxidiidae, 10, 57, 106, 179
Myxidium, 10, 58, 107, 179, 186
Myxidium americanum, 117

anguiUae, 114

barbatulae, 110

bergense, 112

danilewskyi, 109

depressum, 114

• 501]

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gadi, 115
giardi, 110
giganteum, 110
glutinosum, 115
histophilum, 109
incurvatum, 108
infiatum, 111
kagayamai, 117
lieberkuhni, 107
mackiet, 112
macrocapstdare, 113
oviforme, 114
pfeifferi, 111
phyllium, 116
sp. Awerinzew, 113
sp. Gurley, 109
sp. Mavor, 115
sphaericum, 109
striatum, 116
MyxoboUdae, 11, 58, 128, 179
Myxobolus, 11, 58, 128, 179, 189
Myxobolus aeglefini, 144
anurus, 142
aureatus, 154
6a//er>, 147
bramae, 147
capsulatus, 152
carassii, ISO
cordis, 148
cycloides, 140
cyprini, 143
cyprinicola, 147
discrepans, 156
dispar, 135
eUipsoides, 136
exiguus, 136
fuhrmanni, 130
fundtdi, 151
gtg<w, 145
globosus, 139
Ay/oe, 153
inaequalis, 135
^o», 155
lintoni, 138
tnacrocapsularis, 146
magnus, 150
mesentericus, 157
minutus, 150
tnisgumi, 133
miyairii, 155
/»tf//eft, 128
musculi, 148

neurobins, 144
nodularis, 153
notatus, 131
obesus, 140
oblongus, 139
oculi-leucisci, 130
orbiculatus, 155
oviformis, 137
Permagnus, 149
pfeifferi, 133
physophilus, 146
piriformis, 129
pleuronectidae, 152
rohitae, 132
rotundus, 149
sandrae, 146
scardinii, 146
■yewj, 132
sp. Gurley, 142
sp. Gurley, 142
sp. Gurley, 143
sp. Kudo, 132
sp. Lebzelter, 150
sp. Miyairi, 149
sp. Southwell, 151
sp. Wegener, 149
sphaeralis, 141
squamae, 147
toyamai, 131
transovalis, 139
unicapsulatus, 129
volgensis, 145
Myxoproteus, 9, 56, 81, 179, 183
Myxoproteus ambiguus, 81
cordiformis, 81
cornutus, 82
Myxosoma, 11, 58, 123, 179, 189
Myxosoma dujardini, 124
funduli, 125
? lobatum, 124
Myxosomatidae, 11, 58, 123, 179
M3^osporidia in air bladder, 39

alimentary canal, 39, 41

Amphibia, 36

Annelida, 25

blood vessel, 39

body cavity, 37, 43, 45

bone, 39

branchia, 38

cartilage, 39

connective tissue, 37

eye, 38

264

ILUNOIS BIOLOGICAL MONOGRAPHS

[502

gall bladder, 39
heart, 39
Insecta, 25
integument, 37
intestine, 41
kidney, 41
liver, 39
mesentery, 41
muscle, 38
nervous tissue, 39
organs of host, 37
ovary, 41
Pisces, 25
pyloric coecum, 39
Reptilia, 36
spleen, 41
stomach, 39
testis, 41
tissue, 37, 43, 45
unknown seat, 41
urinary bladder, 41
of Africa, 14
Asia, 13

Burma, 14

India, 14

Japan, 13

Kamtschatka, 14

Nippon, 13
Australia, 14
Europ>e, 17

Austria, 24

England, 22

France, 18

Germany, 20

Italy, 17

Monaco, 17

Netherland, 22

Norway, 22

Russia, 24

Serbia, 24

Switzerland, 23
North America, 15

Alaska, 16

Canada, 16

United States, 15
South America, 16
unknown genera and species,

12, 174
Nervous tissue, Myxosporidia parasitic in, 39
Netherland, Myxosporidian of, 22
New species listed, 196

Nipjwn, Myxosporidia of, 13

North American Myxospwridia, 15

Norwegian Myxosporidia, 22

Organs of host infected by Myxosporidia, list

of, 37
Ovary, Myxosporidia parasitic in, 41
Pansporoblast, 50
Plasmogamy, 51
Plasmotomy, 51
Platysporea, 10, 57, 106, 179
Polar capsule, 48, 51
filament, 48, 51
Polysporea, 52, 54
Polysporous, 51
Posterior filament, 47, 51
process, 47, 51
Pyloric coecum, Myxosporidia parasitic in, 39
Reptilian hosts of Myxosporidia, 36
Ridge of spore, 51
Russian Myxosp)oridia, 24
Serbian Myxosporidian, 24
Shell of spore, 47, 51

-valve, 47, 51
Sinuolinea, 10, 57, 104, 178, 186
Siniwlinea arborescens, 105
brachiophora, 106
capsularis, 105
dimorpha, 104
opacita, 106
South American Mj^osporidia, 16
Species, scheme of description of, 7
Sphaeromyxa, 10, 58, 118, 179, 188
Sphaeromyxa balbianii, 118

exneri, 121

gaskrostei, 121

heUandi, 121

immersa, 119

incurvata, 119

sabrazesi, 120
Sphaerospora, 10, 57, 100, 178, 185
Sphaerospora angulata, 102

carassii, 103

divergens, 100

elegans, 100

masovica, 101

plaiessae, 102

polymorpha, 102

rostrata, 101

sp. Davis, 103

sp. Southwell et Prashad, 103
Sphaerosporea, 10, 57, 86, 178

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Sphaerosporidae, 10, 57, 99, 178
Spleen, Myxosporidia found in, 41
Spore, description of, 47
Sporoplasm, 51

Stomach, Myxosporidia parasitic in, 39
Sutural diameter, 51

edge, 51

line, 51

plane, 51

ridge, 51
Switzerland, Myxosporidia of, 23
Tail of spore, 51

Testis, Myxosporidia parasitic in, 41
Thickness of spore, 51

Tissue, Myxosporidia parasitic in, 37, 43, 45

Trophozoite, 51

United States, Myxosporidia of, 15

Urinary bladder, Myxosporidia parasitic in,

41
Vegetative form, 51
Wardia, 9, 56, 82
Wardia ohlmackeri, 83

ovinocua, 82
Zschokkella, 10, 58, 122
Zschokkella acheiiognalhi, 123

globulosa, 123

hildae, 122

nova, 122

Kfttaral HlAtory Eiinrey
i- Library