(navigation image)
Home American Libraries | Canadian Libraries | Universal Library | Community Texts | Project Gutenberg | Children's Library | Biodiversity Heritage Library | Additional Collections
Search: Advanced Search
Anonymous User (login or join us)
Upload
See other formats

Full text of "Studies on Myxosporidia; a synopsis of genera and species of Myxosporidia; with 25 plates and 2 textfigures"

/ 



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 

AXTESSACE, M. 

1906. Ein Myxobolus im Kopfe von Gadus aeglefini L. Zool. Anz., 30:568-570; 4 fig. 
1906a. Weitere Mitteilungen iiber Myxobolus aeglefini. Zool. Anz., 31 :1 15-1 19; 5 fig. 
1906b. Eine neuer Myxobolus im Brachsen {Ahramis brama L.). Zool. Anz., 31:386- 
391; 5 fig. 

1908. Bemerkungen iiber MjTcosporidien heimischer Siisswasserfische. Zool. Anz., 

32:456-465; 7 fig. 

1909. Bemerkungen iiber Myxosporidien. 2k>oL Anz., 34:65-82; 6 fig. 

1909a. Bericht iiber eine Studienreise nach Bergen (Norway). Verb, naturw. Ver. 
Karkruhe, 21:37-73; 2 pi. 

1910. Biologische und morphologische Bemerkungen fiber Myxosporidien. Zool. 

Anz., 35:57-63; 3 fig. 
•f 1910a. Die Sporenbildung von Zschokkella und das System der Myxosporidien. Zool. 

Anz., 35:240-256; 5 fig. 
1910b. Cnidosporidienstudien. Zool. Anz., 35:767-777; 4 fig. 
1910c. Die Cnidosporidien (Myxosporidien, Actinomyxidien, Microsporidien). Eine 

monographische Studie. 261 pp.; 83 fig. 
1910d. Zwei neue Cnidosporidien aus cyprinoiden Fischen. Zool. Anz., 36:440-441; 

Ifig. 

1911. Untersuchungen iiber Eenneguya psoras pertnica Th^lohan. Verb, naturw. 

Ver. Karlsruhe, 24:1-25; 2 textfig., 2 pi. 
1911a. Unsere heutigen Kenntnisse iiber die geographische Verbreitung der Myxo- 
sporidien. Zool. Jarhb., Syst., 30:471-494. 

1912. Studien iiber die Myxosporidien der norwegischen Seefische und ihre Verbreitung. 

Zool. Jahrb., Syst., 34:1-50; 5 textfig., 5 pi. 
1912a. Bemerkungen iiber den Infektionsmodus der Seefische mit Myxosporidien. 
Zool. Anz., 39:617-623. 
'^^ 1912b. Die Sporenbildung der M>'xosporidien. Zool. Anz., 40:204-207 
1917. Bemerkungen iiber Myxosporidien. Zool. Anz., 49:145-157; 5 fig. 
AWZEINZEW, S. 
, 1907. Ueber Myxosporidien aus Gallenblase der Fische. 2^1. Anz., 31:831-834. 

1908. Studies on parasitic Protozoa. I, II, and III (Russian). Trav. soc. imp. nat. 
St.-Petersbourg, 28 (fasc. 2), 67 pp.; 3 pi. 
'^1909. Studien iiber parasitischen Protozoen. I. Die Sporenbildung bei Ceratotnyxa 
. drepanopsettae mihi. Arch. Protist., 14:74-112, 2 pi. 

r 1911. Studien iiber parasitischen Protozoen. VII. Ueber die Sporenbildimg bei 
Myxidium sp. aus der Gallenblase von Coitus scorpius. Arch. Protist., 23:199- 
204; 7 fig. 

1913. Myxobolus magnus nov. sp. Zool. Anz., 42:75-76; 2 fig. 

1913a. Ergebnisse der Untersuchungen iiber parasitische Protozoen der tropischen 
Region Afrikas. III. Zool. Anz., 42:151-156; 4 fig. 
Balbiani, G. 

1863. Sur I'organisation et la nature des psorospermies. C.R. acad. sci., 57:157-161. 

1883. Myxosporidies ou psorospermies des poissons. Joum. Microgr., 7:143-147, 

197-204, 270-281; 12 fig. 

1884. Lefons sur les Sporozoaires. IV. Les psorospermies des poissons ou Myxospori- 

dies, p. 420-449; 12 fig. 



439] STUDIES ON MYXOSPORIDIA—KUDO 201 

Berg, 

*1898. Communicat. Mus. Nac, Buenos Aires, p. 41, 
Borne, M. von dem, B. Benecke and E. Dallmer. 

*1886. Handbuch der Fischzucht und Fischerei. 701 pp., 581 fig. Berlin. 
BOSANQUET, W. C. 

1910. Brief notes on two M30cosporidian organisms {Pleistophora hippoglossoides n. sp. 
and Myxidiutn mackiei n. sp.). Zool. Anz., 35:434-438; 13 fig. 

BtJTSCHLI, O. 

1881. Beitrage zur Kenntnis der Fischsporospermien. Zeitschr. wiss. Zool., 35:629- 

651; 1 pi. 

1882. Myxosporidia. Bronn'sKlass.Ordn., Protozoa, 1:590-603; 19 fig. 
C£PEDE, C. 

1905. Myxosporidies des poissons des Alpes frangaises. C. R. ass. fr. I'avanc. sc. 

sess., 33:905-913. 

1906. Myxosporidies des poissons des Alpes frangaises. Ann. I'univ. Grenoble, 18: 

57-68. 
1906a. Sur la pr6tendu6 inimunit6 des Cobitis k regard des infections myxosporidiennes. 

C.R. soc. biol., 60:15-16. 
1906b. Myxidiutn giardi CdpSde, et la pr6tendu6 immunity des Anguilles k r6gard des 

infections myxosporidiennes. C.R. soc. biol., 60:170-173; 4 fig. 

1907. A propos de le dehiscence des spores des Myxosporidies. C.R. soc. biol., 62 :13S- 

137. 

1908. La myxosporidiose des Anguilles dans les eaux douces, saumitres et salves du 

Boulonnais. Feuille jenn. natur. (4) 38:93-95; 6 fig. 
"' 1913. Existence de la plasmotomie hivemale chez Henneguya legeri C6pMe. Arch. 
Parasit., 16 :302-305 ; 26 fig. 
Claparede, a. de 

*1874. Notice sur les psorospermies des poissons. Lunel's Hist, natur. des poissons 
du basin du L6man. Geneve, p. 113-114. 
Cleland, J. B. and Johnston, T. H. 

1910. The Haematozoa of Australian Batrachians. Proc. Roy. Soc. N.S.W., 44: 
252-261. 

COHN, L. 

1895. Ueber die Myxosporidien von Esox lucius und Perca fluviaiilis. Inaugural- 

Dissert. Albertus-Univ. Konigsberg, 48 pp.; 2 pi. 

1896. Ueber die Myxosporidien von Esox lucius und Perca fluviatilis. Zool. Jahrb., 

Anat., 9:229-272; 2 pi. 
1902. Zur Kenntnis der Myxosporidien. Centralb. Bakt. Parasit., (I) Orig. 32:628- 
632; 2 fig. 
Creplin, J, C. H. 

1842. Beschreibung der Psorospermien des Kaulbarsches nebst einigen Bemerkungen 
uber die der Plotze und andere. Arch. Naturg., 8:61-66; 1 pi. 
CuNHA, A. M. DA and O. da Fonseca. 

1917. Sobre os myxosporidios dos peixes brazileiros. Brazil-Medico, 31:321. 
Danilewsky, B. 

1887. Recherches sur la parasitologic du sang. Arch. slav. biol., 3 :35. 
Davis, H. S. 

f~\9\6. The structure and development of a Myxosporidian parasite of the squeteague, 
Cynoscion regalis. Journ. Morph., 27:333-377; 7 textfig., 7 pi. 
J) y 1917. The Myxosporidia of the Beaufort Region. A systematic and biologic study. 
C^^ BuU. Bur. Fish., 35:201-243; 8 pi. 

•Original paper not seen. 



202 ILLINOIS BIOLOGICAL MONOGRAPHS [440 

DE Drothn de Bouvtlle, R. 

1908. Maladie des abc^s du Bameau (Mj^oboliasis tubCTOsa). Bull. soc. sc. Nancy 
 (3) 9:525-548; 9 textfig., 1 pi. 

DOFLEIN, F. 

1898. Studien zur Naturgeschichte der Protozoen. III. Ueber Myxosporidien. Zool. 

Jahrb., Anat., 11:281-350; 20 textfig., 7 pi. 

1899. Fortschritte auf dem Gebiete der Myxosporidiraikunde. Zusammenfassende 

Uebersicht. Zool. Centr., 6:361-379. 

1901. Die Protozoen als Parasiten und Krankheitserreger nach biologischen Gesichts- 

punkten dargestellt. Jena. 

1902. Das System der Protozoen. Arch. Protist., 1:169-192, 

1909. Die Lehrbuch der Protozoenkunde. 2 ed. 
1911. Die Lehrbuch der Protozoenkunde. 3 ed. 
1916. Die Lehrbuch der Protozoenkunde. 4 ed. 

Erdmann, Rh. 

1911. Zur Lebensgeschichte von Chlorotnyxum leydigi, eine miktosporen Myxosporidie. 

Teil I. Arch. Protist.,. 24:149-162; 4 textfig., 3 pi. 
^^1917. New facts and views concerning the occurrence of a sexual process in the Myxo- 
sporidian life cycle. Amer. Natur., 51:719-739. 
1917a. Chlorotnyxum leydigi und seine Beziehungen zu anderen Myxosporidien. Teil II. 
Arch. Protist., 37:276-326; 17 textfig. and 4 pi. 

EVERMANN, B. W. 

1893. A report upon investigations made in Texas in 1891. Bull. U. S. Fish. Comm., 
11:76. 
Fantham, H. B. and Annie Porter, 

1912. Some effects of the occurrence of Myxosporidia in the gall-bladder of fishes 

(Prelim. Communic). Ann. Trop. Med. Parasit., 6:467-481. 
Fermor, X. 

1913. Wissenschaftliche Ergebnisse einer Reise von S. Awerinzew in die Tropen Afrikas. 

Zool. Anz., 42:196-199. 

FiCKERT, K. 

1895. Ueber die Barbenseuche. Zeitschr. Fischer., 3:209-214. 

FlEBIGER, J. 

1909. Ueber Protozoen als Parasiten der Fische. Verb, zool.-bot. Ges. Wien. 59 : 32-48. 
Fletcher, A. W. 

1888. On a Myxosporidium infesting Australian frogs. Rep. Austr. Ass. Adv. Sc, 1 :337. 
Fritsch, a. J. 

*1902. Ueber Lebensweise, Nahrung und Parasiten der Fische der Elbe. Arch, naturw. 
Landesforsch. Prag, 11 (No. 3). 

FUHRMANN, O. 

1902. Sur les Myxosporidies des Coregones du lac de Neuchitel. Arch. sc. phys. nat. 

Geneve, 14:172-173, 16:331-332. 
1904. Ueber eine Krankheit der weiblichen Geschlechtsorgane des Hechtes. Allg. 

Fischer.-Zeit., 29:469-471. 
FujiTA, T. 

1912. Notes on new Sporozoan parasites of fishes. 2k)ol. Anz., 39:259-262; 3 fig. 

1913. On a new species of Chloromyxum from the gall-bladder of the caip. Annot. 

Zool. Japon., 8:257-259; 1 fig. 
GEORGfivrrcH, J. 

1914. Sur le cycle 6volutif chez les myxosporidies. C.R. acad. sci., 158 :190-192. 

*Original paper not seen. 



441] STUDIES ON MYXOSPORIDIA—KUDO 203 

-.1914a. fitude de cycle dvolutif chez les Myxosporidies. Arch, zool.exp., 54:387-409; 3pl. 

1916. Note sur les Myxosporidies recueillies k Roscofif. Bull. soc. zool. France, 

41:86-95. 
1916a. Note sur les Myxosporidies des poissons de la baie de Villefranche et de Monaco. 

Bull. I'inst. ocean., no. 322; 5 fig. 
1916b. Sur les diverses formes de Ceratomyxa herouardi Georg6vitch. C.R. acad. sci., 
163:717-719. 
'^ 1916c. Sur le cycle 6volutif de Ceratomyxa herouardi Georg6vitch. C.R. acad. sci., 163: 

983-985. 
• 1917. Recherches sur le d6veloppement de Ceratomyxa herouardi Georg6vitch. Arch, 
zool. exp., 56:375-399; 3 pi. 
1917a. Esquisses protistologiques. Bull. I'inst. oc6an, no. 328; 50 fig. 
1917b. Esquisses protistologiques. Bull. soc. zool. France, 42:99-107. 
>C 1917c. Sur le cycle 6volutif de Myxidium gadi Georg6vitch. C.R. acad. sci., 165 :797-799. 

1918. Sur Chloromyxum leydigi. Bull. soc. zool. France, 42 :182-189; 18 textfig. 
.^1919. fitudes sur le d^veloppement de Myxidium gadi Georg€vitch. Arch. zool. exper., 
58:251-289; 3 pi. and 4 textfig. 
Gluge, G. 

1841. Beobachtungen zahlreicher Balggeschwulste als epidemische Krankheit bei 
Fischen. Anat.-Mikr. Unters. allg. u. sp. Path. 2:202-204; 1 pi. 
Gurley, R. R. 

1893. On the classification of Myxosporidia, a group of protozoan parasites infesting 

fishes. Bull. U. S. Fish Comm., 11 :407-420. 

1894. The Myxosporidia, or psorosperms of fishes, and t\ie epidemic produced by then. 

Rep. U. S. Fish Comm., 26:65-304; 47 pi. 
Hahn, C. W. 

1915. Sporozoan parasites of certain fishes in the vicinity of Woods Hole, Mass. Bull. 
Bur. Fish., 33:193-214; 2 pi. 

1917. On the Sporozoan parasites of the fishes of Woods Hole and vicinity. I. Further 

observations on Myxobolus »»M5CM/t from Fundulus. Joum.Parasit., 3:91-104; 
3 fig. 
1917a. On the Sporozoan parasites of the fishes of Woods Hole and vicinity. II. Addi- 
tional observations upon Myxobolus musculi of Fundulus and a nearly related 
species, Myxobolus pleuronectidae oi Pleuronectes americanus. Joum.Parasit., 
3:150-162; 1 pi. 
1917b. On the Sporozoan parasites of the fishes of Woods Hole and vicinity. III. On 
the Chloromyxum clupeidae of Clupea harengus {Young), Pomolobuspseudo- 
harengus (Young), and P. aestivalis (Young). Joum. Parasit., 4:13-20. 
Hartmaxn, M. 

1909. Autogamie bei Protisten und ihre Bedeutung fur das Befruchtungsproblem. 
Arch. Protist., 14:286-292. 
Haswell, W. a. 

1890. A remarkable flat worm in the golden frog. Proc. Linn. Soc. N.S.W., 5:661-666. 
Heckel, J. and R. Kner. 

*1858. Die Siisswasserfische der oestereichischen Monarche mit Rucksicht auf die 
angrenzenden Lander. Leipzig. 388 pp., 204 fig. 
HOFER, B. 

1893. Ueber die Drehkrankheit der Regenbogenforelle. Allg. Fishereiz., 18:7-8. 

1895. Ueber Fischkrankheiten. Zeitschr. Fisch., 3:179-209; 23 fig. 

1896. Die sogenannte Pockenkrankheit der Karpfen. Allg. Fischereiz., 2 1 :2, 186, etc. 
1896a. Die Infektion der Fische mit Myxosporidien. Allg. Fischereiz., 21 :38-39. 
1904. Handbuch der Fischkrankheiten. Stuttgart. 359 pp., 222 fig. 

*Original paper not seen. 



204 ILLINOIS BIOLOGICAL MONOGRAPHS [442 

HUBER, J. C. 

1888. Ueber Piesbergens Fischsporospermien. Centralbl. Bakt. u. Parasit., 3:663-664. 
ISHH, S. 

1915. Myxosporidiosis of Nipponese eel (Nipponese). Jour. Zool., 27:372-382; 1 pi. 

1915a. Lentospora-disease of the eel (Nipponese). Jour. Zool. 27 :471-474; 4 fig. 
Jameson, P. 

1913. A note on some Myxosporidia collected at Monaco. Bull. I'inst. oc6an., no. 273. 
JOHKSTON, T. H. 

1909. Exhibit of Myxobolus. Proc. Roy. Soc. N.S.W., 43:29. 
Johnston, T. H. and M. J. Bancroft. 

1918. A parasite, Myxoholus hylae sp. nov., of the reproductive organs of the golden 

swamp frog, Hyla aurea. Australian Zoologist, 1 :171-175; 5 textfig.; 1 pi. 

1919. Some new sporozoan parasites of Queensland freshwater fish. Jour. Proc. Roy. 

Soc. N.S.W., -52:520-528; 5 pi. 
Johnstone, J. and H. M. Woodcock. 

1907. On a Myxosporidian infection of Gadus esmarkii, with a note on the identification 
of the parasite. Trans. Biol. Soc. Liverpool, 21:204-208; 1 pi. 
Joseph, H. 

1905. Chloromyxum protei n. sp. Zool. Anz., 29:450-451. 

1907. Chloromyxum protei n. sp., ein in der Niere des Grottenohns parasitierendes Myxo- 

sporidium. Arch. Protist., 8:398-412; 1 textfig., 2 pi. 
Keysselitz, G. 

1908. Ueber durch Sporozoen (Mjocosporidien) hervorgerufene pathologische Verander- 

ungen. Verb. Ges. deutsch. Natur. u. Arzte, Vers. 79:452-453. 
1908a. Die Entwicklung von Myxoholus pfeifferi. Teils I, II, Arch. Protist., 11 :252-308; 
14 textfig., 4 pi. 
Klokacewa, S. 

1914. Ueber die Myxosporidien der Karausche. Zool. Anz., 44:182-186; 2 fig. 

KOLESNIKOFF, N. F. 

*1886. O psorospermiakh (mikrosporidiakh) v muskulature rib. Vet. Vestnik, Kharkoff, 
5:242-248; 1 pi. 
Kudo, R. 

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- 

dia found in some fresh-water fishes of Japan, with the description of three new 
species. Joum. Parasit., 3:3-9; 4 fig. 

1917. Contributions to the study of parasitic protozoa. II. Myxobolus toyamai nov. 

spec., a new Myxosporidian parasite in Cyprinus car pioL. Joum. Parasit., 
3:163-170; 2 pi. 

1918. Experiments on the extrusion of polar filaments of Cnidosporidian spores. Joum. 

Parasit., 4:141-147; 1 pi. 
1918a. Contributions to the study of parasitic protozoa. IV. Note on some Myxospori- 
dia from certain fish in the vicinity of Woods Hole. Joum. Parasit., 5:11-16; 
4 textfig., 2 pi. 
Labb£, a. 

1899. Sporozoa. Das Tierreich. 5 Lief. 180 pp. 
Laveran, a. 

1897. Sur une Myxosporidie des reins de la tortue. C.R. soc. biol. 49:725-726. 

1898. Surle Myxidiumdanilewskyi. C.R. soc. biol., 50:27-30; 9 fig. 
Laveran, A. and F. Mesnil. 

1900. Sur une Myxosporidie des voies biliares de I'Hippocampe (Sphaeromyxa sahrasesi 

nov. spec). C.R. soc. biol., 52:380-382; 4 fig. 
♦Original paper not seen. 



443] STUDIES ON MYXOSPORIDIA—KUDO 205 

1902. Sur la multiplication endogene des Myxosporidies. C.R., socbiol., 54:469-472; 

5 fig. 
Lebzelter, v. 
1^^ 1912. Ueber Protozoen aus der Gallenblase von Thymallus thymallus L. Zool. Anz., 
*^ 40:295-297. 

L£ger, L. 

^y- 1906. Myxosporidies nouvelles parasites des poissons. I. Sur una nouvelle maladie 
myxosporidienne de la truite indigne. II. Sur une nouvelle Myxosporidie de la 
tanche commune. Ann. I'univ. Grenoble, 18:267-272; 3 fig. 
L£ger, L. and E. Hesse. 

1906. Sur la structure de la parol sporale des Myxosporidies. C.R. acad. sci., 142: 
720-722; 8 fig. 
Leuckart, R. 

1852. Parasitismus und Parasiten. Arch, physiol. Heilkde, 1 1 :434-436. 
1879. Die Parasiten des Menschen. 336 pp.; 130 fig. 
Leydig, F. 

1851. Ueber Psorospermien und Gregarinen. Arch. Anat., Phys. u. wiss. Med., 18: 
221-234; 1 pi. 

LlEBERKtJHN, N. 

1854. Notice sur les psorospermies. Bull. ac. roy. Belg., 21:21-23. 

1855. Les psorospermies des poissons. M6m, cour. et m6m. sav. etrang. acad. 
roy. Belg., 26:36-38; 2 pi. 

Linton, E. 

1891. On certain wart-like excrescences, occurring on the short minnow, Cyprinodon 

variegatus, due to psorosperms. Bull. U.S. Fish Comm., 9:99-102; 1 pi. 
1891a. Notice of the occurrence of protozoan parasites (psorosperms) on Cyrinoid 

fishes in Ohio. Bull. U. S. Fish Comm., 9:359-361 ; 1 pi. 
1901. Parasites of fishes of the Woods Hole region. Bull. U. S. Fish Conim.,19:407- 

492;34pL 

LO GlXJDICE, 

1912. Studi sui Cnidosporidi. Pavia, 91 pp. ; 1 pi. 
LXJDWIG, H. 

*1888. Ueber die Myxosporidienkrankheit der Barben in der Mosel. Jahresb. rhein. 
Fisch. Ver., Bonn, p. 27-36. 
LtjHE, M. 

1896. Die Infektion der Fische mit Myxosporidien. Die Natur, 45:284-285; 5 fig. 
1899. Cystodiscus immersus. Verb, deutsch. zool. Ges., 9:291-293; 1 fig. 
LuTz, A. 

1889. Ueber eine Myxosporidium aus der Gallenblase brazilianischer Batrachier (Cys- 
todiscus immersus). Centralbl. Bakt. u. Parasit., 5:84r-88; 10 fig. 
Mavor, J. W. 

1915. Studies on the Sporozoa of the fishes of the St. Andrew's Region. Ann. Rep. 
Dept. Marine Fishery, 47, no. 39b:25-38; 6 textfig.; 1 pi. 

1916. On the life-history of Ceratomyxa acadiensis, a new species of Myxosporidia from 
the eastern coast of Canada. Proc. Amer. Acad. Arts and Sci., 51 :55 1-574; 3 
textfig., 3 pi. 

1916a. Studies on the protozoan parasites of the fishes of the Georgian Bay. Trans. 

Roy. Soc. Canada., (3) 10:63-74; 6 fig. 
1916b. On the occurance of a parasite of the pike in Europe, Myxidium lieberkiihni 

Butschli, in the pike on the American continent and its significance. Biol. Bull., 

31:373-378; 1 pi. 
*Original paper not seen. 



206 ILLINOIS BIOLOGICAL MONOGRAPHS [444 

Mayor, J. W., and W. Strasser. 

1916. On a new Myxosporidian, Henneguya wisconsinensis n. sp., from the urinary blad- 

der of the yellow perch, Perca fluvescens. Trans. Wise. Acad. Sci., Arts and 
Letters, 18:676-682; 3 fig. 
Mazzarelli, G. 

1905. Sulla pseudodifterite degli Agoni. Atti soc. ital. sc. nat. mUs. civ. stor, nat. 

Milano, 44:71-72. 

1906. Le condizioni della pesca nella provincia di Milano. Riv. mens, pesca, Milano, 

8:301-325. 
♦1908. Di alcune malattie di pesci e gamberi osservate in Lombardia. Atti terzo con- 
greso naz. di pesca, Milano, p. 272. 
MiGNIN, T. P. 

1885, Sur le r6le pathog6nique de certaines psorospermies. Bull. soc. zool. France, 
10:351-352. 
Mercier, L. 

1906. Phenomenes de sexualite chez Myxobolus pfeiferi. C.R. soc. biol., 110:427-428. 
1906a. Contribution h. I'etude du d^veloppement des spores chez Myxobolus pfeifferi. 
C. R. soc. biol., 110:763-764. 

1908. Notes sur les Myxosporidies. Arch. zool. exp., 8JLIII-LXII; 5 figs. 

1909. Contribution a I'^tude de la sexualit6 chez les Myxosporidies et chez les Micro- 

sporidies. M6m. class, sc. I'acad. roy. Belg., 2:3-30; 1 pi. 

MiLEWSKI, A. 

1917. Myxoboliasis tuberosa, die Barbenseuche oder Beulenkrankheit der Barbe. 

Wochenschr. Aquar.-Terrar.-Kde., 14:14-17. 
MiNCHIN, E. A. 

1912. An Introduction to the study of the protozoa. 517 pp.; 194 textfig. 
Mingazzini, p. 

1890. Sullo svilui^o dei Myxosporidi. Boll. soc. nat. Napoli, 4:160-164; 1 pi. 
Miyairi, K. 

1909. Guide to the study of parasitic protozoa (Nipponese). 170 pp.; 8 pi. 
MOROFF, T. 

1906. Ueber die Gelbsucht der Bachforelle. Osterr. Fischereiz. 3 :285. 

MiJLLER, J. 

1841, Ueber eine eigenthiimliche krankhafte parasitische Bildung mit specifisch organi-" 

sierten Samenkorperchen. Arch. Anat. Phys. u. wiss. Med., 5:477-488; 1 pi, 
1841a. Krankhafter Hautsusschlag mit specfisch organisierten Korperchen (Psorosper- 
mien). Ber. preuss. Akad. Wiss. Berlin, p. 212-221, 246-250, 232 pp., 4 pi. 
MuLSOw, K. 

1911. Ein neuer Gehimparasit des Karpfens. Allg. Fisch. Zeit., 36:483-485; 3 fig. 
Nemeczek, a. 

1911. Beitrage zur Kenntnis der Myxo- und Microsporidien der Fische. Arch. Protist., 
22:143-169; 19 textfig., 2 pi. 
Ntifer, W. 

1905. Die Fische des Vierwaldstattersees und ihre Parasiten. Inaug.-Diss, Basel, 
230 pp.; 21 fig. 
*1905a. Die Parasiten der Fische des Vierwaldstattersees. Festschr. 50-jahr. Besteh. 
naturf. Geselisch., Luzem, pp. 65-232; 4 pi. 
Ohlmacher, A.P. 

1893. Myxosporidia in the common toad, with preliminary observations on two chro- 
mophile substances in their spores. Jour. Amer. Med. Assoc, 20:561-567; 1 pi. 
Parisi, B. 

1910. Sphaerosporacaudatan.sp. Zool. Anz., 36:253-254; 3 figs. 
*Original paper not seen. 



445] STUDIES ON MYXOSPORIDIA— KUDO 207 

1912. Primo contributo alia distribusione geografica dei missosporidi in Italia. Atti 

soc. ital. sc. nat., 50:283-290; 11 fig. 

1913. Svi)laL Sphaerospora caudataVa.ri'&i. Atti soc. ital. sc. nat. 51:5-12; 1 pi. 
Perugia, A. 

*1890. SuUe Myxosporidie dei pesci marini. Boll, scient. Pavia, 12:134-139. 
*1891. Sulle Myxosporidie dei p>esci marini. Boll, scient. Pavia, 13:22-25; 1 pi. 
Pfeiffer, L. 

1890. Die Protozoen als Krankheitserreger. 1 ed. 100 pp.; 34 fig. 

1891. Die Protozoen als Krankheitserreger. 2 ed. pp. 216; 91 fig. 

1893. Die Parasitismus des Epithelcarcinoms sowie der Sarco-, Micro- und Myxo- 

sporidien in Muskelgewebe. Centralbl. Bakt. u. Parasit., 14:118-130. 
1895. Die Protozoen als Krankheitserreger. Nachtrage. Jena. pp. 122; 52 fig. 

PlESBERGEN, F. 

1886. Eine neue Form von Fischpsorospermien aus Perca fivviatUis. Jahresb. Ver. 
vater. Naturk. Wurttemberg, 42:73. 
Plehn, M. 

1904. Woher kommt die Drehkrankheit der Salmoniden? Allg. Fischereiz., 29 :151-153. 
1904a. Weiteres uber die Drehkrankheit. Allg. Fischereiz., 29:183-184. 

1905. Ueber die Drehkrankheit der Salmoniden (Lentospora cerebralis [Hofer] Plehn). 

Arch. Protist., 5:145-166; 7 textfig., 1 pi. 

1906. Die Drehkrankheit der Salmoniden. Allg. Fischereiz., 31 :465-470. 

1906a. Nochmals iiber die Drehkrankheit der Salmoniden. AUg. Fischereiz., 31:516- 

519. 
1910. Die pathogene Bedeutung der Myxoboliden fiir die Fische. Sitz. Ges. Morph. u. 
Physiol. Munchen, 26:20-27; 3 fig. 
POCHE, F. 

1913. Das System der Protozoen. Arch. Protist., 30:125-321; 1 fig. 
Prenant, a. 

1902. Striation et ciliation de la partie adherente du Myxidium Ueberkuhni Biltschli. 

C.R. soc. biol, 54:844-846. 
1902a. Notes cytologiques. VII. Contribution a I'etude de la ciliation. Striation et 
ciliation de la partie adherente du Myxidium Ueberkuhni. Arch. anat. micr., 
Paris, 5:212; 7 textfig. 
Railliet, a. 

1886. Maladie des Barbeaux causae par des psorospermies. Bull. M6m. soc. centr. 

med. v6t. Paris, 4:134-137. 
1890. La maladie des Barbeaux de la Mama. Bull. soc. centrale d'aquic. Paris, 
2:117-120. 
Remak, R. 

1852. Ueber runde Blutgerinsel und iiber pigmentgukelhaltige Zellen. Arch. Anat., 
Phys. u. wiss. Med., 144-146; 1 pi. 
Reuss, H. 

1906. Neue Myxosporidien von Susswasserfischen. Bull. I'acad. imp. sc. St.-Peters- 
bourg, 25:199-205; 1 pi. 
Ryder, J. A. 

1880. The psorosperms found in Aphredoderus sayanus. Amer. Nat., 14:211-212; 2 fiigs. 
Sandeman, G. 

1894. On the multiple tumors in Plaice and Flounders. Ann. Rep. Scott. Fish. 

Board, 11:391-392; p. 391-392. 
*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 



PLATE I 



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). 



ILLINOIS BIOLOGICAL MONOGRAPHS 



VOLUME V 




KUDO 



9 ^<:=f==:^ ■" "~<=i=^ 14 
STUDIES ON MYXOSPORIDIA 



PLATE I 



4S1J STUDIES ON MYXOSPORIDIA—KUDO 213 



PLATE II 



214 ILLINOIS BIOLOGICAL MONOGRAPHS [452 



• EXPLANATION OF PLATE 

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. 



ILLINOIS BIOLOGICAL MONOGRAPHS 



VOLUME V 




KUDO 



STUDIES ON MYXOSPORIDIA 



PLATE II 



453] STUDIES ON MYXOSPORIDIA— KUDO 215 



PLATE III 



216 ILLINOIS BIOLOGICAL MONOGRAPHS [454 



EXPLANATION OF PLATE 

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. 



ILLINOIS BIOLOGICAL MONOGRAPHS 



VOLUME V 




KUDO 



STUDIES ON MYXOSPORIDIA 



PLATE III 



4SS] STUDIES ON MYXOSPORIDIA—KUDO 217 



PLATE IV 



218 ILLINOIS BIOLOGICAL MONOGRAPHS [456 



EXPLANATION OF PLATE 

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. 



ILLINOIS BIOLOGICAL MONOGRAPHS 



VOLUME V 




KUDO 



STUDIES ON MYXOSPORIDIA 



PLATE IV 



457} STUDIES ON MYXOSPORIDIA—KUDO 219 



PLATE V 



2ip ILUNOIS BIOLOGICAL MONOGRAPHS [458 



EXPLANATION OF PLATE 

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. 



ILLINOIS BIOLOGICAL MONOGRAPHS 



VOLUME V 




KUDO 



STUDIES ON MYXOSPORIDIA 



PLATE V 



459] STUDIES ON MYXOSPORIDIA—KUDO 221 



PLATE VI 



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). 



ILLINOIS BIOLOGICAL MONOGRAPHS 



VOLUME V 




KUDO 



STUDIES ON MYXOSPORIDIA 



PLATE VI 



461] STUDIES ON MYXOSPORJDJA— KUDO 223 



PLATE VII 



224 ILUNOJS BIOLOGICAL MONOGRAPHS (462 



EXPLANATION OF PLATE 

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. 



ILLINOIS BIOLOGICAL MONOGRAPHS 



VOLUME V 




KUDO 



STUDIES ON MYXOSPORIDIA PLATE VII 



Hi] STUDIES ON MYXOSPORJDIA—KUDO 225 



PLATE VIII 



226 ILLINOIS BIOLOGICAL MONOGRAPHS [464 



EXPLANATION OF PLATE 

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. 



ILLINOIS BIOLOGICAL MONOGRAPHS 



VOLUME V 




KUDO 



172 N;^W 173 

STUDIES ON MYXOSPORIDIA 



PLATE VIII 



465] STUDIES ON MYXOSPORIDIA— KUDO 227 



PLATE IX 



328 ILLINOIS BIOLOGICAL MONOGRAPHS [466 



EXPLANATION OF PLATE 

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). 



ILLINOIS BIOLOGICAL MONOGRAPHS 



VOLUME V 




KUDO 



STUDIES ON MYXOSPORIDIA 



467] STUDIES ON MYXOSPORJDIA—KUDO 229 



PLATE X 



230 ILUNOIS BIOLOGICAL MONOGRAPHS [46S 



EXPLANATION OF PLATE 

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). 



ILLINOIS BIOLOGICAL MONOGRAPHS 



VOLUME V 




KUDO 



223 (3>'"''^*^ 221 

STUDIES ON MYXOSPORIDIA 



I 



469] STUDIES ON MYXOSPORIDIA— KUDO 231 



PLATE XI 



232 ILLINOIS BIOLOGICAL MONOGRAPHS [470 



EXPLANATION OF PLATE 

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). 



ILLINOIS BIOLOGICAL MONOGRAPHS 



VOLUME V 




249 ^^^.c=e^ 250 ^ ^ 251 

STUDIES ON MYXOSPORIDIA 



PLATE XI 



471] STUDIES ON MYXOSPOBJDJA—KUDO 233 



PLATE XII 



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. 



ILLINOIS BIOLOGICAL MONOGRAPHS 



VOLUME V 




KUDO 



288 \ 1/ 289 \l/ 290 

STUDIES ON MYXOSPORIDIA 



PLATE XII 



473] STUDIES ON MY XOSPORIDI A— KUDO 235 



PLATE XIII 



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. 



ILLINOIS BIOLOGICAL MONOGRAPHS 



VOLUME V 




KUDO 



STUDIES ON MYXOSPORIDIA 



i 

1 



^H STUDIES ON MYXOSPORIDIA— KUDO 237 



PLATE XIV 



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. 



ILLINOIS BIOLOGICAL MONOGRAPHS 



VOLUME V 




339 340 341 ' ^''^ 343 

STUDIES ON MYXOSPORIDIA 



PLATE XIV 



4771 STUDIES ON MYXOSPORJDIA—KUDO 239 



PLATE XV 



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. 



ILLINOIS BIOLOGICAL MONOGRAPHS 



VOLUME V 




372 



KUDO 



STUDIES ON MYXOSPORIDIA PLATE XV 



4791 STUDIES ON MYXOSPORJDJA—KUDO 241 



PLATE XVI 



242 ILLINOIS BIOLOGICAL MONOGRAPHS i^» 



EXPLANATION OF PLATES 

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). 



ILLINOIS BIOLOGICAL MONOGRAPHS 



VOLUME V 




KUDO 



STUDIES ON MYXOSPORIDIA 



PLATE XVI 



4»l] STUDIES ON MYXOSPORIDIA— KUDO 243 



PLATE XVII 



2M ILLINOIS BIOLOGICAL MONOGRAPHS [482 



EXPLANATION OF PLATE 

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. 



ILLINOIS BIOLOGICAL MONOGRAPHS 



VOLUME V 




437 ^^^^444 v^^^445 

STUDIES ON MYXOSPORIDIA PLATE XVII 



4831 STUDIES ON MYXOSFORIDIA—KUDO 3^ 



PLATE XVIII 



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. 



ILLINOIS BIOLOGICAL MONOGRAPHS 



VOLUME V 




466 ^^^ 468 ^^==^4671 

KUDO STUDIES ON MYXOSPORIDIA PLATE XVIII 



485] STUDIES ON MYXOSPORIDIA—KUDO 247 



PLATE XIX 



248 ILLINOIS BIOLOGICAL MONOGRAPHS [186 



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). 



ILLINOIS BIOLOGICAL MONOGRAPHS 



VOLUME V 




KUDO 



502 / 503 5041 

STUDIES ON MYXOSPORJDIA 



PLATE XIX 



487] STUDIES ON MYXOSPORIDIA—KUDO 249 



PLATE XX 



250 ILLINOIS BIOLOGICAL MONOGRAPHS [488 



EXPLANATION OF PLATE 

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). 



ILLINOIS BIOLOGICAL MONOGRAPHS 

1^ 



VOLUME V 




 521 

STUDIES ON MYXOSPORIDIA 



PLATE XX 



STUDIES OW MTZOSPOEIDI A— KUDO 2H 



PLATE XXI 



252 ILLINOIS BIOLOGICAL MONOGRAPHS [490 



EXPLANATION OF PLATE 

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. 



ILLINOIS BIOLOGICAL MONOGRAPHS 



VOLUME V 




638 



KUDO 



539 



549 

STUDIES ON MYXOSPORIDIA 



544 

PLATE XXI 




491] STUDIES ON MYXOSPORIDIA— KUDO 2S3 



PLATE XXII 



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. 



ILLINOIS BIOLOGICAL MONOGRAPHS 



VOLUME V 




KUDO 



STUDIES ON MYXOSPORIDIA PLATE XXII 



493] STUDIES ON MYXOSPORIDIA—KUDO 255 



PLATE XXIII 



2M ILLINOIS BIOLOGICAL MONOGRAPHS [494 



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. 



ILLINOIS BIOLOGICAL MONOGRAPHS 



VOLUME V 



^^•'<)B.<3..?i-J' 




KUDO 



STUDIES ON MYXOSPORIDIA 



590 
PLATE XXIII 



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. 



ILLINOIS BIOLOGICAL MONOGRAPHS 



VOLUME V 



.w^- i%b^ K<M^;. K% 







KUDO 



STUDIES ON MYXOSPORIDIA 



627 

PLATE XXIV 



497] STUDIES ON MYXOSPORIDIA—KUDO 259 



PLATE XXV 



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. 



ILLINOIS BIOLOGICAL MONOGRAPHS 



VOLUME V 




649 
KUDO 



650 



661 652 

STUDIES ON MYXOSPORIDIA 



PIATE XXV 



499] 



STUDIES ON MYXOSPORIDIA—KUDO 



261 



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] 



STUDIES ON MYXOSPORIDIA—KUDO 



263 



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 



503] 



STUDIES ON MYXOSPORIDIA—KUDO 



265 



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